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Journal of Biology Advance Access published 23 November 2018 Journal of Crustacean Biology The Crustacean Society Journal of Crustacean Biology 39(1) 36–39, 2019. doi:10.1093/jcbiol/ruy092

An additional representative of Lecythocaridae (: Brachyura: ) from the Upper

Jurassic of southern Germany Downloaded from https://academic.oup.com/jcb/article/39/1/36/5202037 by guest on 29 September 2021

Günter Schweigert Staatliches Museum für Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany

Correspondence: Günter Schweigert; email: [email protected] (Received 14 September 2018; accepted 23 October 2018)

ABSTRACT A new species of the previously monotypic brachyuran genus Prolecythocaris Schweigert & Robins, 2016, P. rieberi n. sp., is described from southern Germany. The holotype was col- lected from an Upper massive siliceous sponge-microbial limestone of early late age. Relationships within Lecythocaridae are briefly discussed. Key Words: diversification, , palaeoecology

INTRODUCTION & Feldmann, 2009a), Czech Republic (Moericke, 1889; Patrulius, 1959), Poland (Patrulius, 1966; Schweitzer & Feldmann, 2009a), In modern marine benthic environments, brachyurans are well and Romania (Patrulius, 1959, 1966; Franţescu, 2010). represented by approximately 7,250 species (Davie et al., 2015), The aim of this study is to report a second species of the previ- with several families entering freshwater (Yeo et al., 2008) and even ously monotypic genus Prolecythocaris. The abbreviation SMNS is terrestrial habitats (Bliss & Mantel, 1968). The most primitive ones, used for Staatliches Museum für Naturkunde Stuttgart, Germany. belonging to a clade of brachyurans taxonomically summarized in the section Dromiacea De Haan, 1833, were exclusively marine. Their earliest known representatives occurred in shallow marine, GEOLOGICAL SETTING mostly non-reefal environments of the Early and Middle Jurassic (Krobicki & Zatoń, 2008, 2016). Soon after, in the Late Jurassic, The described specimen was collected from massive limestones siliceous sponge-microbial magnafacies became widespread along of a sponge-microbial reef complex exposed in the upper part the northern margin of the Tethys Ocean (e.g., Leinfelder et al., of the street (“Messelbergsteige”) connecting the small town of 1994, 1996). This probably triggered a rapid diversification and Donzdorf and Schnittlingen village just before reaching the top biogeographical expansion of dromiacean (Wehner 1988; of the Swabian Alb (Fig. 1). These sponge reefs are summarized as Müller et al., 2000; Klompmaker et al., 2013). The Upper Jurassic belonging to the Oberjura-Massenkalk Formation (Mönnig et al., of southern Germany, which is part of this Tethyan reef belt, is 2018). There are no further data available about additional a classical area for brachyuran studies (von Meyer, 1860). Most from this locality because the traffic along the street makes any of the occurring species were originally lumped in the single sampling very difficult. A few meters up this section, however, bed- genus Prosopon von Meyer, 1835. During the last decades, numer- ded limestones of the Untere-Felsenkalke Formation are exposed, ous publications have been dedicated to taxonomic revisions of which are interfingering with the massive sponge-microbial lime- the original type material (Feldmann et al., 2006; Schweitzer et al., stones and hence are of the same age. Ammonite remains from 2007; Schweitzer & Feldmann, 2008a, b, 2009a, b, c; Schweigert there including fragments of Aulacostephanus sp. (Horst Kuschel, & Koppka, 2011). A few additional taxa have been reported from personal communication, 2018), which point to a position of the this area more recently (Schweigert & Koppka, 2011; Starzyk, finding horizon within the late Kimmeridgian Acanthicum Zone. 2015; Schweigert & Robins, 2016). Within their revision of homol- The sponge-microbial limestones of the Swabian and Franconian odromioid crabs, Schweitzer & Feldmann (2009a) established the Alb developed on a deep-shelf ramp position along the northern monotypic family Lecythocaridae Schweitzer & Feldmann, 2009 margin of the Tethys Ocean, with estimated maximum water (cf Schweitzer & Feldmann, 2009a). Prolecythocaris Schweigert & depths of ca. 150–200 m (e.g., Leinfelder et al., 1994, 1996; Schmid Robins, 2016 was added in 2016 from the early Kimmeridgian of et al., 2005). In the entire Swabian Upper Jurassic, patch reefs with southern Germany (Franconia). Lecythocarids have been reported hermatypic corals never occur in the stratigraphic position of the from southwestern Germany (von Meyer, 1860; Quenstedt, 1867; exposed section. Coral patch reefs locally appeared in this area in Beurlen, 1925; Wehner, 1988; Müller et al., 2000; Schweigert & the latest Kimmeridgian Ulmense Subzone of the Beckeri Zone, Robins, 2016; Schweigert & Kuschel, 2018), Austria (Schweitzer immediately after a sudden shallowing, probably caused by a

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Figure 1. Locality of the material described. Arrow indicates location of the Messelbergsteige near Donzdorf, southwestern Germany. tectonically induced uplift of the sea bottom (Schweigert & Franz, 2004). Type horizon: Oberjura-Massenkalk Formation (Upper Jurassic, late Kimmeridgian, Acanthicum Zone).

SYSTEMATIC PALAEONTOLOGY Studied material: Holotype only. The classification follows Schweitzer et al. (2010). Occurrence: At present only known from the type locality. Order Decapoda Latreille, 1802 Infraorder Brachyura Latreille, 1802 Diagnosis: A species of Prolecythocaris with significantly inflated subhepatic areas, otherwise matching the original diagnosis of Section Dromiacea De Haan, 1833 the genus. Superfamily Patrulius, 1959 Description: The holotype SMNS 70451 (Fig. 2) is three-dimension- Family Lecythocaridae Schweitzer & Feldmann, 2009 ally preserved with its calcified cuticle. Surface of carapace coarsely pustulose in all areas, with pustules of irregular size. Outline of Genus Prolecythocaris Schweigert & Robins, 2016 carapace rounded-subtrapezoidal, slightly wider than long. Total Type species: Prolecythocaris hauckei Schweigert & Robins, 2016, by length of carapace including the rostrum 3.6 mm; maximum width original designation. 4.0 mm, maximum height (in centre of mesogastric area) 2.0 mm. Hepatic regions large, triangular, weakly separated from adjacent Species included: Prolecythocaris hauckei Schweigert & Robins, 2016; regions. Epigastric region grades into forwardly directed blunt, trap- Prolecythocaris rieberi n. sp. (herein). ezoidal rostrum. Orbital margin straight, perpendicular to longitu- dinal axis. Orbits very large, rounded, laterally bordered by inflated, coarsely tuberculated subhepatic region. Cervical groove well devel- Prolecythocaris rieberi n. sp. oped; branchiocardiac groove indistinct, hidden by shallow epi-, (Fig. 2A–D) mesobranchial areas. Urogastric region extremely narrow, hardly separated from adjacent regions. Epibranchial lobes well developed, Etymology: Named after Artur Rieber (1894–1994), who collected laterally separated from epigastric areas by weak transverse furrows. the sole specimen during a hiking tour in 1949, and his son Volkmar Cardiac region triangular, moderately elevated. Branchial regions Rieber (Horb am Neckar, Germany), who donated the specimen. prominent, slightly inflated. Marginal spines lacking. Posterior mar- gin of carapace concave in the intestinal area, forming a smooth Holotype: Specimen SMNS 70451 (Fig. 2). rim. Sternal region, appendages not preserved.

Type locality: Messelbergsteige (steep road connecting Donzdorf Remarks: Prolecythocaris rieberi n. sp. differs from P. hauckei Schweigert and Schnittlingen) near Donzdorf, eastern Swabian Alb, Germany. & Robins, 2016 in its inflated subhepatic region, a much coarser

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Figure 2. Prolecythocaris rieberi n. sp., holotype, SMNS 70451, Upper Jurassic, Oberjura-Massenkalk Formation, late Kimmeridgian, Acanthicum Zone, Donzdorf, southwestern Germany. Dorsal (A, B), frontal (C), lateral (D) views; B–D whitened with ammonium chloride prior to photography. Scale bar = 2 mm. This figure is available in color at Journal of Crustacean Biology online. ornamentation, and the lack of any lateral spines. Since only one Prolecythocaris and the oldest specimen of Lecythocaris paradoxa (von specimen of P. rieberi n. sp. is available, it is impossible to state Meyer, 1858) are recorded from southern Germany, this area whether this species was smaller than P. hauckei or just a juvenile. could have been the originating area for Lecythocaridae.

DISCUSSION ACKNOWLEDGEMENTS Prolecythocaris rieberi n. sp. from the lower Kimmeridgian of I thank Volkmar Rieber (Horb, Germany), who kindly donated Swabia represents the second species assigned to this genus. the studied specimen and numerous other fossils from his and It is stratigraphically younger than the type species P. hauckei his father’s collections. Cristina Robins (University of California, Schweigert & Robins, 2016, which comes from the early Berkeley, USA) checked an early draft of the manuscript. Carrie Kimmeridgian Platynota or Hypselocyclum zones. Both the E. Schweitzer (Kent State University, OH, USA), an anonymous stratigraphically oldest record of the more heavily ornamented one, and the editors are heartily thanked for their valuable sugges- Lecythocaris paradoxa (von Meyer, 1858) (see Schweigert & Robins, tions and criticism that helped to improve the quality of this paper. 2016: fig. 2D) andP. rieberi n. sp. come from a coeval bed of the same area. This co-occurrence supports the previously supposed origin of Lecythocaridae at the norther Tethyan margin and the REFERENCES phyletic relationship between the two genera as sister taxa shar- Beurlen, K. 1925. Über Brachyuren- und Anomurenreste des ing a common ancestor (Schweigert & Robins, 2016). Both taxa Schwäbischen Jura. Neues Jahrbuch für Mineralogie, Geologie und inhabited relatively deep environments within sponge-microbial Paläontologie, Beilage-Bände, 52: 464–532. mounds. Younger Lecythocaris spp. are predominantly found in Bliss, D.E. & Mantel, L H. 1968. Adaptations of to land: a later Kimmeridgian and coral reefs not only in south- summary and analysis of new findings. American Zoologist, 8: 673–685. ern Germany but elsewhere in Europe, and thus representatives Davie, P.J.F., Guinot, D. & Ng, P.K.L. 2015. Systematics and classifica- of this taxon are considered a typical component of coral reef tion of Brachyura. In: Treatise on zoology – anatomy, , biology. The Crustacea (P. Castro, P.J.F. Davie, D. Guinot, F.R. Schram & J.C. von communities (Schweitzer et al., 2018). The very few isolated and Vaupel Klein, eds.), Vol. 9C-II, pp. 1049–1130. Brill, Leiden, The extremely rare occurrences of Prolecythocaris spp. either point to Netherlands. an adaptation to very special environmental conditions, or could Feldmann, R.M., Lazăr, I. & Schweitzer, C.E. 2006. New crabs (Decapoda: result from collection biases due to their small size and a lithol- Brachyura: Prosopidae) from Jurassic (Oxfordian) sponge bioherms of ogy unfavorable for sampling macrofossils. Since both taxa of Dobrogea, Romania. Bulletin of the Mizunami Museum, 33: 1–20.

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