© Author(s) 2020. CC Atribution 4.0 License New early Paleocene (Danian) paguroids from deep-water coral/bryozoan mounds at Faxe, eastern Denmark Novi zgodnjepaleocenski (danijski) raki samotarji iz globokovodnih koralno- briozojskih kop nahajališča Faxe na vzhodnem Danskem Sten L. JAKOBSEN1, René H.B. FRAAIJE2, John W.M. JAGT3 & Barry W.M. VAN BAKEL2,4 1Geomuseum Faxe, Østsjællands Museum, Rådhusvej 2, 4640 Faxe, Denmark; e-mail: SLJ@oesm.dk 2Oertijdmuseum Boxtel, Bosscheweg 80, 5283 WB Boxtel, the Netherlands; e-mail: info@oertijdmuseum.nl 3Natuurhistorisch Museum Maastricht, de Bosquetplein 6-7, 6211 KJ Maastricht, the Netherlands; e-mail: john.jagt@maastricht.nl 4Oertijdmuseum Boxtel, Bosscheweg 80, 5283 WB Boxtel, the Netherlands, and Naturalis Biodiversity Center, Leiden, the Netherlands; e-mail: barryvanbakel@gmail.com Prejeto / Received 9. 11. 2019; Sprejeto / Accepted 2. 4. 2020; Objavljeno na spletu / Published online 22. 4. 2020 Key words: Anomura, Paguroidea, Paleogene, cheliped morphology, Denmark Ključne besede: Anomura, Paguroidea, paleogen, morfologija škarnikov, Danska Abstract During recent decades, decapod crustacean faunas from middle Danian (lower Paleocene) strata at Faxe (Sjælland, Denmark) have been studied in detail. However, paguroid anomurans have not yet been described formally. Two new species of hermit crab have lately been recognised in the collections of the Geomuseum Faxe. Percentages of total paguroid assemblages and feeding behaviour derived from the morphology of its chelae indicate that one of these, Dardanus faxensis sp. nov., as a generalist, was better adapted to inhabit the deep-water reefal environment of the Faxe carbonates than the more specialised, suspension-feeding Paguristes frigoscopulus sp. nov. Izvleček V zadnjih desetletjih so bile podrobno proučene združbe rakov deseteronožcev iz srednjedanijskih (spodnjepaleocenskih) plasti v Faxu (Sjælland, Danska), vendar raki samotarji do sedaj še niso bili opisani. Pri pregledu zbirk muzeja Geomuseum Faxe sta bili prepoznani dve novi vrsti rakov samotarjev, ki ju predstavljamo v tem prispevku. Zastopanost ostankov v celotni združbi in način prehranjevanja, ki ga kaže funkcionalna morfologija škarij, pričajo, da je bil eden od teh, to je vrsta Dardanus faxensis sp. nov., kot generalist bolje prilagojen za bivanje v globokomorskem grebenskem okolju od bolj specializiranega suspenziofaga Paguristes frigoscopulus sp. nov. GEOLOGIJA 63/1, 47-56, Ljubljana 2020 https://doi.org/10.5474/geologija.2020.005 Introduction In general, records of paguroid remains from Danian-aged rocks are scarce and hermit crab assemblages of this age remain largely under- studied. Vega et al. (2007) recorded a right and an incomplete left chela of a diogenid from the Rancho Nuevo Formation at Coahuila (Mexico), while Armstrong et al. (2009) described a right paguroid palm, without systematic placement, from the Midway Group of Texas, USA. In addi- tion, Cope et al. (2005) noted Paguristes johnsoni Rathbun, 1935, on the basis of a single, well-pre- served propodus and an additional fragment of a carpus that were closely comparable to the type material illustrated and described by Rathbun (1935) from the Sucarnoochee Formation of Al- abama (USA). Robin et al. (2017) mentioned an indeterminate species of Paguristes Dana, 1851 from a private- ly held collection of Danian decapod crustaceans from Vigny, France. It would appear that this locality was very rich in paguroids of earliest Paleocene age (Wallaard et al., 2020), but unfor- tunately this material is not currently accessible for scientific assessment and publication. A brief visit to the collections of the Geomu- seum Faxe (September 2019) has now resulted in the recognition of fourteen hermit crab speci- 48 Sten L. JAKOBSEN, René H.B. FRAAIJE, John W.M. JAGT & Barry W.M. VAN BAKEL mens along with the material from the collections at the Natural History Museum (Copenhagen), all partially complete chelae. The two new paguroid taxa erected to accommodate these forms from the middle Danian of Faxe present a valuable ad- dition to the hermit crab faunas from this time interval. Geological setting A large, cool-water, subphotic coral/bryozoan mound complex at Faxe developed in the Danish Basin during the early Danian, shortly after the Cretaceous/Paleogene boundary event (e.g., Bjer- ager et al., 2018; Schrøder & Surlyk, 2019). During the middle Danian, a low-diversity scleractinian coral fauna initiated the formation of extensive cold-water coral mound complexes that interca- lated with bryozoan mounds (e.g., Lauridsen et al., 2012). The coral mounds started to grow be- low the photic zone over the easternmost part of the Ringkøbing-Fyn High, only 2 million years after the mass extinction at the K/Pg boundary (Lauridsen et al., 2012). The middle Danian limestones exposed at Faxe quarry (Fig. 1) document the extraordi- nary preservation of a 63-myr-old, cold-water coral mound ecosystem. Bryozoan mounds dom- inate the stratigraphically lower parts and are overlain by interfingering coral and bryozoan mounds. The Faxe quarry exposes a large mound complex that includes individual coral mound bodies. The mound complex is dominated by the frame-building scleractinian coral species, Den- drophyllia candelabrum (Hennig, 1899), which is associated with common Faksephyllia faxoensis (Lyell, 1837) and minor numbers of Oculina becki (Brünnich Nielsen, 1922). A highly diverse fauna is found in middle Danian strata at Faxe quarry, including serpulid annelids, arthropods, brachi- opods, bryozoans, coelenterates, echinoderms, sponges and molluscs (Lauridsen et al., 2012, ta- ble 1). The quarry at Faxe comprises the best-stud- ied Danian decapod crustacean fauna worldwide (Jakobsen & Collins, 1997). Studies dealing with such taxa, in chronological order, are the follow- Fig. 1. Position of Faxe qu- arry and the main structure of the Danish area during the middle Danian (compi- lation by Dr Erik Thomsen, Geological Institute, Aarhus University, Denmark). 49New early Paleocene (Danian) paguroids from deep-water coral/bryozoan mounds at Faxe, eastern Denmark ing: von Schlotheim (1820), Reuss (1859), von Fis- cher-Benzon (1866), Segerberg (1900), Woodward (1901), Rasmussen (1973), Förster (1975), Jagt et al. (1993), Collins & Jakobsen (1994), Jakobsen & Collins (1997), Fraaije (2003), Jakobsen (2003), Jakobsen & Feldmann (2004), Collins (2010), Robin et al. (2015) and Klompmaker et al. (2015, 2016). However, in none of these are any paguroid remains recorded, so that our present note is the first formal description of middle Danian hermit crab taxa from Faxe. Material and methods The taxonomic descriptions are based on ma- terial stored at the Geological Museum (now Natural History Museum of Denmark) and and Geomuseum Faxe. Our material comprises four- teen specimens belonging to two genera, repre- sented by two new species. The material includ- ed in the present study was collected mainly by Curator Søren Bo Andersen and the late ama- teur palaeontologist, Alice Rasmussen, of Faxe. In 2012, the latter was awarded the prestigious Mary Anning Prize of the Palaeontological As- sociation (London) in recognition of her out- standing work on fossils from the type locality of the Danian Stage at Faxe. The present mate- rial was collected in various parts of the quarry over many years. The chelae are preserved mainly in the form of internal and external moulds in the coral lime- stone and therefore our descriptions are in part based on casts. However, that of the holotype of Paguristes frigoscopulus relied on the internal mould (‘steinkern’). Preparation and photogra- phy of the material followed the procedures out- lined in Jakobsen & Feldmann (2004). Institutional abbreviations. MGUH, Natural History Museum of Denmark (Geological Muse- um), Copenhagen, Denmark; OESM, Geomuseum Faxe, Faxe, Denmark. Systematic palaeontology Order Decapoda Latreille, 1802 Infraorder Anomura H. Milne Edwards, 1832 Superfamily Paguroidea Latreille, 1802 Family Annuntidiogenidae Fraaije, 2014 Genus Dardanus Paul’son, 1875 Dardanus faxensis sp. nov. (Figs. 2-5) 2005 Eremitkrebs 20 mm**; Damholt & Rasmus- sen, p. 22, unnumbered figures. 2005 Klo fra eremitkrebs 20 mm**; Damholt & Rasmussen, p. 38, unnumbered figure. 2010 Eremitkrebs 20 mm**; Damholt et al., p. 23, unnumbered figures. 2010 Klo fra eremitkrebs 20 mm***; Damholt et al., p. 42, unnumbered figure. Diagnosis: Palm transversely oval; inner and outer sides longitudinally and transversely con- vex; outer side of palm covered with forward- ly directed, largely spinose tubercles; proximal part of outer side of palm and distal part of inner side irregularly covered with capsulated setae arranged in a curved row; fixed finger elongated, with a horseshoe-shaped, spoon-like tip. Etymology: In reference to the type locality. Type material: The holotype, OESM 6811, is a near-complete right palm with fixed finger, measuring 10 mm in length and 6 mm in great- est width; both external and internal moulds are present (Fig. 2A-G). Paratypes are MGUH 33401, MGUH 33402, OESM 581, OESM 10178 OESM 10179 and OESM 10180, all representing incom- plete right palms; MGUH 33403, MGUH 33404, MGUH 33405, MGUH 33406 and MGUH 33408 are incomplete left palms. Locality and stratigraphy: The coral lime- stone assigned to the Faxe Formation has been known for centuries due to the extensive quar- rying. The middle Danian levels exposed at Faxe quarry represents an extraordinary preservation of a 63-myr-old, cold-water coral mound com- plex. This coral limestone complex was formally described as a new formation by Lauridsen et al. (2012). The type locality of the Faxe Formation is a Danish GeoSite and is accessible to all mem- bers of the public. The formation was dated as middle Danian (calcareous nannoplankton zones D5-6 of Perch-Nielsen, 1979 and NNTp3-NNT- p4a of Varol, 1998). Description: Cross section of palm transverse- ly oval; inner and outer sides longitudinally and transversely convex, curving slightly inwards; dorsal edge straight, ventral edge concave at cen- tral part; outer side of palm covered with forward- ly directed, largely spinose tubercles, largest tu- bercles irregularly arranged along dorsal edge and outer distal half and proximal half of fixed finger, proximal part of outer side of palm irregularly covered with multiple capsulated setae arranged in circles or rows; inner side of palm smooth with on distalmost part few curved rows of multiple setae (Fig. 3B); fixed finger elongated, curved in- wardly, with horseshoe-shaped, spoon-like tip, which is covered on outer side with a row of very large pits; the cutting edge is covered with about three variably sized, molar-like calcareous teeth. 50 Sten L. JAKOBSEN, René H.B. FRAAIJE, John W.M. JAGT & Barry W.M. VAN BAKEL Remarks: Normally, setal pits are situated around the bases of tubercles and/or on the ad- jacent integument, but usually not on the tuber- cles themselves. Multiple capsulated pits have previously been recorded for both fossil and ex- tant species of paguroids (see Fraaije et al., 2011, 2015a; Hyžný et al., 2016). During the Paleogene, the genus Dardanus appears to have been high- ly successful and widely distributed (Fraaije & Polkowsky, 2016). In the case of Dardanus faxen- sis sp. nov. all tubercles on the outer side of the palm have large setal pits (Fig. 2A); this morpho- logical trait differentiates it immediately from all congeners. For a list of all known fossil spe- cies referred to the genus, reference is made to Fraaije et al. (2011) and to Fraaije & Polkowsky (2016). The shape of the chelae is indicative of a deposit feeder that was also capable of browsing, scavenging and suspension feeding. This type of paguroid fed by scraping material from cor- als or other solid surfaces using the tips of the spoon-like chelae (compare Schembri, 1982). One specimen, MGUH 33406, is totally covered with epibionts, mainly subcircular colonies of bryozo- ans (Fig. 4). Fig. 2. Dardanus faxensis sp. nov., holotype (OESM 6811). The top three - silicone rubber casts of external mould; the bottom four internal mould. Scale bar in millimetres. 51New early Paleocene (Danian) paguroids from deep-water coral/bryozoan mounds at Faxe, eastern Denmark Fig. 3A. Dardanus faxensis sp. nov., silicone rubber cast of external mould of incomplete right palm, outer side (OESM 10178), showing setal pits and tubercles; B. Silicone rubber cast of external mould of incomplete right palm (MGUH 33401, GMF 2004- 1802), showing rows of multiple setal pits. Scale bar in millimetres. Fig. 4. Dardanus faxensis sp. nov., MGUH 33405, incomplete left palm covered with epibionts. Scale bar in millimetres. 52 Sten L. JAKOBSEN, René H.B. FRAAIJE, John W.M. JAGT & Barry W.M. VAN BAKEL Three specimens of D. faxensis sp. nov. bear remains of serpulid/spirorbid annelids on the outer palm surface at the base of the fixed finger (Fig. 5). This appears to have been a symbiotic relationship, rather than post-mortem serpulid attachment to moulted anomurans, as described by Jakobsen & Feldmann (2004). Family Diogenidae Ortmann, 1892 Genus Paguristes Dana, 1851 Paguristes frigoscopulus sp. nov. (Fig. 6A-E) Diagnosis: Right cheliped transversely oval; outer, dorsal and ventral sides convex, inner side almost straight; outer side covered with regular, dense cover of fine, forwardly directed tubercles, fixed finger short, stout and triangular. Etymology: From Latin, frigus, meaning cold, and scopulus, meaning reef. Type material: The holotype, OESM 6705, is a near-complete right palm with fixed finger, meas- uring 8.5 mm in length and 5.5 mm in greatest width. The paratype, MGUH 33407, is an incom- plete right palm, still embedded in coral-limestone. Fig. 5. Dardanus faxensis sp. nov., specimens OESM 6811, OESM 10180 and OESM 10179, respectively, with remains of serpu- lid/spirorbid annelids (arrows) on the outer palm surface near the base of the fixed finger. Scale in millimetres. Fig. 6A-E. Paguristes frigoscopulus sp. nov., A-D. Holotype (OESM 6705), a near-complete right palm in internal mould pre- servation; E. Paratype (MGUH 33407), an incomplete right palm, still embedded in coral limestone. Scale bar in millimetres. 53New early Paleocene (Danian) paguroids from deep-water coral/bryozoan mounds at Faxe, eastern Denmark Locality and stratigraphy: The same as for Dardanus faxensis sp. nov. (see above). Description: Cross section of right palm trans- versely oval; outer, dorsal and ventral sides con- vex, inner side almost straight; outer side with regular, dense cover of fine, forwardly directed tubercles; fixed finger short, stout and triangu- lar; occlusal margin straight without teeth, sur- rounded by few large pores; interdigital margin oblique; length of fixed finger about one quarter of total length. Remarks: Fraaije et al. (2015b) listed all ex- tinct species of Paguristes known at that time. With a convex dorsal and ventral edge and a short, convex triangular fixed finger, P. frigo- scopulus sp. nov. is morphologically closest to the late Albian P. liwinskii Fraaije, van Bakel, Jagt & Machalski, 2015 from east-central Poland and P. santamartaensis Feldmann, Tshudy & Thom- son, 1993, from the Campanian of Antarctica. It differs in having a less dense, more forwardly di- rected, tuberculose ornamentation. Conclusions In total, fourteen specimens were available for the present study. Of these, about 86 % (12 specimens) are attributed to D. faxensis sp. nov. and only c. 14 % (2 specimens) to P. frigoscopu- lus sp. nov. Unfortunately, there are no detailed studies yet that relate cheliped morphology to feeding strategy. Schembri (1982) is one of the few studies that documented a close relationship between cheliped morphology and types of feed- ing. If we extrapolate his data to Dardanus fax- ensis sp. nov., that species may be considered to have been a generalist and better adapted to the Danian deep-water reefal environment of Faxe than P. frigoscopulus sp. nov., which probably was mainly a filter feeder. Acknowledgements We are greatly indebted to Mr Søren Bo Andersen (Geological Institute, University of Aarhus) for do- nation of some specimens from Faxe. We also thank the late Mrs Alice Rasmussen for having transfer- red very important material (both holotypes) to the Geomuseum Faxe. 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