© Author(s) 2020. CC Atribution 4.0 License Paguroid anomurans from the upper Tithonian–lower Berriasian of Štramberk, Moravia (Czech Republic) Zgornjetitonijski–spodnjeberiasijski raki samotarji iz Štramberka, Moravska (Češka) René H.B. FRAAIJE1, Barry W.M. VAN BAKEL1, John W.M. JAGT2 & Petr SKUPIEN3 1Oertijdmuseum, Bosscheweg 80, 5283 WB Boxtel, the Netherlands; e-mail: info@oertijdmuseum.nl 2Natuurhistorisch Museum Maastricht, de Bosquetplein 6-7, 6211 KJ Maastricht, the Netherlands; e-mail: john.jagt@maastricht.nl 3Department of Geological Engineering, VŠB-Technical University of Ostrava, 17. listopadu 15, Poruba, 708 33 Ostrava, Czech Republic; e-mail: petr.skupien@vsb.cz Prejeto / Received 10. 8. 2019; Sprejeto / Accepted 26. 11. 2019; Objavljeno na spletu / Published online 30. 1. 2020 Key words: Paguroidea, upper Tithonian - lower Berriasian, Štramberk Limestone, Moravia, Czech Republic Ključne besede: Paguroidea, zgornji titonij - spodnji beriasij, Štramberk, Moravska, Češka republika Abstract Subsequent to a preliminary report on a handful of paguroid remains from the Tithonian (uppermost Jurassic) to lower Berriasian (Lower Cretaceous) Štramberk Limestone in Moravia (eastern Czech Republic), published in 2013, several field campaigns were organised by our research team during the summers of 2012–2015 and 2018. These resulted in the recovery of additional paguroid shields (or, anterior carapaces) that form the basis of the present study. The currently available material documents a diverse paguroid fauna. In fact, it ranks amongst the most diverse fossil paguroid assemblages known, following faunas from the upper Kimmeridgian of Nusplingen (southern Germany) and the Tithonian of Ernstbrunn (northeast Austria). New representatives of five families and five genera are described, named and illustrated, as follows: Annuntidiogenes sagittula sp. nov. (Diogenidae), Protopagurus cerebellum sp. nov. and Protopagurus duopupae sp. nov. (Paguridae), Mesoparapylocheles janetjacksonae sp. nov. (Parapylochelidae), Masticacheles septemgradu sp. nov. (Pilgrimchelidae) and Ammopylocheles romankijoki sp. nov. (Pylochelidae). Izvleček Po prvih poročilih o ostankih rakov samotarjev (Paguroidea) titonijske (zgornje jurske) do spodnje beriasijske (spodnje kredne) starosti iz apnencev v kamnolomu Štramberk na Moravskem (vzhodna Češka), ki so bila objavljena leta 2013, smo med leti 2012-2015 in poleti 2018 opravili več dodatnih terenskih raziskav. Pri raziskavah smo odkrili številne nove ščite (sprednje dele oklepa) rakov samotarjev, ki so predstavljeni v pričujočem članku. Opisan fosilni material predstavlja raznovrstno favno rakov samotarjev, ki se uvršča med najbolj raznolike znane fosilne paguroidne združbe, primerljive z zgornjo kimeridžijsko združbo regije Nusplingen (južna Nemčija) in titonijsko paguroidno združbo nahajališča Ernstbrunn (severovzhodna Avstrija). Opisani in predstavljeni so novi predstavniki petih družin in petih rodov rakov samotarjev: Annuntidiogenes sagittula sp. nov. (Diogenidae), Protopagurus cerebellum sp. nov. in Protopagurus duopupae sp. nov. (Paguridae), Mesoparapylocheles janetjacksonae sp. nov. (Parapylochelidae), Masticacheles septemgradu sp. nov. (Pilgrimchelidae) in Ammopylocheles romankijoki sp. nov. (Pylochelidae). GEOLOGIJA 63/1, 9-18, Ljubljana 2020 https://doi.org/10.5474/geologija.2020.001 10 René H.B. FRAAIJE, Barry W.M. VAN BAKEL, John W.M. JAGT & Petr SKUPIEN Introduction The Štramberk Limestone, exposed along sev- eral exploitation levels at Kotouč quarry in the immediate vicinity of the town of Štramberk (Moravia, Czech Republic), comprises variably sized carbonate megablocks, breccias and con- glomerates that represent deposition on a car- bonate platform along the northern Tethyan mar- gin in the area of the present-day Outer Western Carpathians during the latest Jurassic and ear- liest Cretaceous (Vašíček et al., 2018; Vaňková et al., 2019). In recent years, numerous macrofossils have been collected from this quarry thanks to an agreement between the VSB-Technical Uni- versity of Ostrava and the management of Kotouč quarry. From about 1910 onwards, the quarry at Kotouč Hill has been the main source of mac- ro- and microfossils that have been described in numerous palaeontological studies (see Vašíček & Skupien, 2004, 2005, 2019 for references). The reefal limestone facies at the quarry varies wide- ly, ranging from very coarse-grained to gravelly layers or lenses, formed by e.g. molluscan shells and corals, to very fine-grained micritic lime- stones and (most commonly) fine-grained biode- trital limestones (e.g., Houša & Vašíček, 2005). Fieldwork carried out by our research team in the Upper Jurassic–Lower Cretaceous reefal limestones at Kotouč quarry during the summers of 2012–2015 and 2018, has provided a highly di- verse decapod crustacean fauna comprising re- mains of isopods, macrurans, anomurans, and brachyurans. Paguroid material collected during the first campaign was recorded in a preliminary paper; this included a handful of shields (or, por- tions of anterior carapaces) and a single sixth abdominal tergite (Fraaije et al., 2013). In 2015, Gašparič et al. described the galatheoid Gala- theites zitteli (Moericke, 1889) from the infill of a test of a nucleolitid echinoid, collected in June 2014. With at least 18 species, in eight families, the Tithonian (Late Jurassic) paguroid fauna from Ernstbrunn (Austria) is by far the most diverse fossil assemblage recorded to date. In second place, with 13 species in six families, follows that from Nusplingen (upper Kimmeridgian, southern Germany), just ahead of the assemblage described below, which comprises 11 species in seven fam- ilies. Herein, we follow Vašíček & Skupien (2019) and Vaňková et al. (2019) for the stratigraphical placement of the material studied. Seven species are here recorded from the upper Tithonian, and four from the lower Berriasian part of the section exposed at Kotouč. Material and methods Specimens, usually only partially exposed, were mechanically prepared under a LOMO MBS-10 stereomicroscope, using needles and pneumatic airscribes of the types Hardy Winkler HW-1 and HW-70/3. Accidentally chipped pieces were glued back by Starbond super fast thin CA glue. For photography, specimens were first dyed with black water colour, and when dry, coated with ammonium chloride, in order to gain max- imum contrast of fine details. Specimens were photographed using a Canon digital SLR in ap- erture priority, Zeiss Luminar 100 mm and 63 mm macrolenses on a Nikon PB-4 bellows unit. A copy stand was used, and a Euromex coldlight source for illumination. Post-processing was done in GIMP 10.0; levels and curves were ad- justed for white balance and contrast, the sharp- ness slightly enhanced with an ‘unsharp mask’. Systematic palaeontology We here adopt the carapace-based classifica- tion and descriptive terminology of extinct pa- guroids proposed by Fraaije (2014) and Fraaije et al. (2019). All material is contained in the collec- tions of the Oertijdmuseum, Boxtel (the Nether- lands; abbreviation: MAB). Order Decapoda Latreille, 1802 Infraorder Anomura H. Milne Edwards, 1832 Superfamily Paguroidea Latreille, 1802 Family Annuntidiogenidae Fraaije, 2014 Genus Annuntidiogenes Fraaije, Van Bakel, Jagt & Artal, 2008 Type species: Annuntidiogenes ruizdegaonai Fraaije, Van Bakel, Jagt & Artal, 2008, by origi- nal designation. Included species: Annuntidiogenes sagittula sp. nov., An. elongatus Fraaije, Robins, Van Ba- kel, Jagt & Bachmayer, 2019, An. hoelderi Fraa- ije, Robins, Van Bakel, Jagt & Bachmayer, 2019, An. jurassicus Fraaije, 2014, An. massetispino- sus Fraaije, Van Bakel & Jagt, 2017, An. sunucio- rum Fraaije, Van Bakel, Jagt & Artal, 2008, An. ruizdegaonai and An. worfi Fraaije, Van Bakel, Jagt, Klompmaker & Artal, 2009. Annuntidiogenes sagittula sp. nov. (Pl. 1.1) Diagnosis: Shield longer than wide, divided into distinct regions by grooves; long triangu- lar rostrum extending beyond postocular and 11Paguroid anomurans from the upper Tithonian–lower Berriasian of Štramberk, Moravia (Czech Republic) postantennal spines; convex postrostral ridge; scabrous ornament on anterior gastric region; long and wide central gastric groove forming ar- row-shaped figure in conjunction with rostrum; elongated, bipartite massetic region; pronounced triangular anterior branchial area; posterior in- tragastric grooves parallel to cervical groove. Derivation of name: Latin sagittula, meaning small arrow, in reference to the typical arrow- head shape of the central anterior groove in con- junction with the triangular rostrum. Type material: The holotype, and sole speci- men known to date (MAB k.3631), is a near-com- plete shield; as preserved, maximum carapace length measures 4.0 mm, maximum shield width is 3.0 mm. Type locality and type level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 5; lower Berriasian (see Vašíček & Skupien, 2019, p. 39, fig. 3, locality 10; Vaňková et al., 2019, section B, layer B22). Description: Shield longer than wide (L/W ra- tio 1.14), divided into distinct regions by grooves; long, spinose and ridged triangular rostrum extending beyond postocular and postanten- nal spines; very shallow convex orbital and an- tennal cavities; convex postrostral ridge; long and wide central gastric groove, forming ar- rowhead-shaped configuration in conjunction with triangular rostrum; scabrous ornament on anterior gastric region, central gastric groove not extending posteriorly; elongated, bipartite massetic region; small, elongated keraial region; pronounced triangular anterior branchial area; posterior intragastric grooves parallel to cervi- cal groove. Remarks: Intragastric grooves (also known as Y-linea in extant paguroids) are parallel to the cervical groove; this is a unique feature shared by representatives of the family Annuntidioge- nidae. Annuntidiogenes sagittula sp. nov. can be distinguished from all congeners known to date by the unique combination of a very wide central anterior gastric groove, a bipartite mas- setic region and a triangular, ridged rostrum; in conjunction with the gastric groove, this forms a typical arrowhead shape. Family Diogenidae Ortmann, 1892 Genus Bachmayerus Fraaije, Van Bakel, Jagt & Skupien, 2013 Type species: Bachmayerus cavus Fraaije, Van Bakel, Jagt & Skupien, 2008, by original designation. Bachmayerus cavus Fraaije, Van Bakel, Jagt & Skupien, 2013 (Pl. 1.2) Type locality and type level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 7; upper Tithonian (see Vašíček & Skupien, 2019, p. 39, fig. 3). Type material: The holotype, and sole speci- men known to date (MAB k.3631), is a near-com- plete shield; as preserved, maximum carapace length measures 4.0 mm, maximum shield width is 3.0 mm. Remarks: For a detailed description, reference is made to Fraaije et al. (2013). Genus Eopaguropsis Van Bakel, Fraaije, Jagt & Artal, 2008 Type species: Eopaguropsis loercheri Van Ba- kel, Fraaije, Jagt & Artal, 2008, by original de- signation. Eopaguropsis cf. nidiaquilae Fraaije, Krzemiński, Van Bakel, Krzemińska and Jagt, 2012 (Pl. 1.3) Locality and level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 6; upper Titho- nian (see Vašíček & Skupien, 2019, p. 39, fig. 3, locality 3). Material: The specimen (MAB k.3759), is an incomplete shield; as preserved, maximum car- apace length measures 7.0 mm, maximum shield width is 5.0 mm. Remarks: For a detailed description, reference is made to Fraaije et al. (2012c). Family Gastrodoridae Van Bakel, Fraaije, Jagt & Artal, 2008 Genus Gastrodorus von Meyer, 1864 Type species: Gastrodorus neuhausensis von Meyer, 1864, by monotypy. Included species: Gastrodorus bzo- wiensis Krzemińska, Krzemiński, Fraaije, Van Bakel & Jagt, 2015, G. cretahispani- cus Klompmaker, Artal, Fraaije & Jagt, 2011, G. kotoucensis Fraaije, Van Bakel, Jagt & Skupien, 2013 and G. neuhausensis von Meyer, 1864. 12 René H.B. FRAAIJE, Barry W.M. VAN BAKEL, John W.M. JAGT & Petr SKUPIEN Gastrodorus kotoucensis Fraaije, Van Bakel, Jagt & Skupien, 2013 (Pl. 1.4) Type locality and type level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 7; upper Tithonian (see Vašíček & Skupien, 2019, p. 39, fig. 3). Remarks: For a detailed description, reference is made to Fraaije et al. (2013). Family Paguridae Latreille, 1802 Genus Protopagurus Fraaije, Robins, Van Bakel, Jagt & Bachmayer, 2019 Type species: Protopagurus janoscheki Fraa- ije, Robins, Van Bakel, Jagt & Bachmayer, 2019, by original designation. Included species: Protopagurus janoscheki, Protopagurus cerebellum sp. nov. and Protopa- gurus duopupae sp. nov. Remarks: To date, we are unaware of any rep- resentative of the family Paguridae from Oxford- ian and Kimmeridgian strata, in spite of intensive fieldwork in southern Germany and southern Po- land over several years. The oldest known pagu- rid has recently been described from the middle to lower upper Tithonian of Ernstbrunn (Austria; see Fraaije et al., 2019). The new taxa from the lower Berriasian of Moravia appear to substanti- ate the notion that this group rose to dominance during the latest Jurassic (and up to the present day) and ousted the more ancient groups of sym- metrical hermit crabs. Protopagurus duopupae sp. nov. (Pl. 1.5) Diagnosis: Well-areolated shield, slightly longer than wide; large, elongated massetic re- gion, anteriorly covered with scale-like ornamen- tation; shallow central gastric groove centrally indenting convex postfrontal ridge; anterior part of gastric region covered with scale-like orna- mentation; thin, elongated anterior branchial area; well-delineated, reniform keraial region; shield irregularly covered with (setal) pores. Derivation of name: Latin duo and pupa (-e), a noun in apposition, or two puppets, in allusion to the morphology of the gastric region. Type material: The holotype, and sole speci- men known to date (MAB k.3626), is an incom- plete shield of a maximum carapace length, as preserved, of 12.0 mm; the maximum shield width is 13.0 mm. Type locality and type level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 5; lower Berriasian (see Vašíček & Skupien, 2019, p. 39, fig. 3, locality 10; Vaňková et al., 2019, section B, layer B22). Description: Well-areolated shield, slightly longer than wide; large, elongated massetic re- gion, anteriorly covered with scale-like ornamen- tation, posteriorly covered with broad, shallow, pitted furrow slightly curving from anteriormost keraial region to mid-massetic edge; shallow central gastric groove centrally indenting convex postfrontal ridge; anterior part of gastric region covered with scale-like ornamentation; posteri- orly a row of large pits is forming subtransverse furrow; thin, elongated anterior branchial area; well-delineated, reniform keraial region; shield irregularly covered with (setal) pores. Frontal area and posteriormost part of shield not pre- served. Protopagurus cerebellum sp. nov. (Pl. 2.1) Diagnosis: Well-areolated shield, slightly longer than wide; convex orbital cavity, ending in triangular postocular projection; large, elon- gated massetic region, with large pores; central gastric groove centrally indenting convex post- frontal ridge; anterior part of gastric region cov- ered with brain-like ornamentation; thin, elon- gated anterior branchial area; small, reniform keraial region; shield irregularly covered with (setal) pores. Derivation of name: Latin cerebellum, or brains (noun used in apposition), referring to the brain-like ornament of the anterior gastric re- gion. Type material: The holotype, and sole speci- men known to date (MAB k.3628), is a near-com- plete shield of a maximum carapace length, as preserved, of 3.0 mm; the maximum shield width is 3.0 mm. Type locality and type level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 5; lower Berriasian (see Vašíček & Skupien, 2019, p. 39, fig. 3, locality 10; Vaňková et al., 2019, section B, layer B22). Description: Well-areolated shield, longer than wide; rostrum not preserved; convex orbit- al cavity with smooth rim, ending in triangular postocular projection; large, elongated massetic region, irregularly covered with large pores, also on anterior lateral edge; central gastric groove centrally indenting convex postfrontal ridge; an- 13Paguroid anomurans from the upper Tithonian–lower Berriasian of Štramberk, Moravia (Czech Republic) 1 - Annuntidiogenes sagittula sp. nov.; 2 - Bachmayerus cavus Fraaije, Van Bakel, Jagt & Skupien, 2013; 3 - Eopaguropsis cf. nidiaquilae Fraaije, Krzemiński, Van Bakel, Krzemińska and Jagt, 2012, original (left), composite (right); 4 - Gastrodorus kotoucensis Fraaije, Van Bakel, Jagt & Skupien, 2013; 5 - Protopagurus duopupae sp. nov., original (left), composite (right); all scale bars 2 mm. PLATE 1 14 René H.B. FRAAIJE, Barry W.M. VAN BAKEL, John W.M. JAGT & Petr SKUPIEN terior part of gastric region covered with brain- like ornamentation, posteriorly ending convexly; thin, elongated anterior branchial area; small, reniform keraial region; shield irregularly cov- ered with (setal) pores. Family Parapylochelidae Fraaije, Klompmaker & Artal, 2012a Genus Housacheles Fraaije, Van Bakel, Jagt & Skupien, 2013 Type species: Housacheles timidus Fraai- je, Van Bakel, Jagt & Skupien, 2013, by original designation. Housacheles timidus Fraaije, Van Bakel, Jagt & Skupien, 2013 (Pl. 2.2) Type locality and type level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 5 (see Vašíček & Skupien, 2019, p. 39, fig. 3). Remarks: For a detailed description, reference is made to Fraaije et al. (2013). Genus Mesoparapylocheles Fraaije, Klompmaker & Artal, 2012a Type species: Mesoparapylocheles michael- jacksoni Fraaije, Klompmaker & Artal, 2012a, by original diagnosis. Included species: Mesoparapylocheles jaegeri Fraaije, 2014, M. michaeljacksoni, M. schweigerti Fraaije, 2014, M. strouhali Fraaije, Robins, Van Bakel, Jagt & Bachmayer, 2019 and M. zapfei Fraaije, Robins, Van Bakel, Jagt & Bachmayer, 2019. Mesoparapylocheles janetjacksonae sp. nov. (Pl. 2.3) Diagnosis: Shield well calcified, longer than wide, well areolated; globose massetic region; prominent triangular rostrum; triangular posto- cular spines. Gastric region of arrowhead shape, pointing posteriorly. Distinct and complete U-shaped branchiocardiac groove, parallel to V-shaped cervical groove. Derivation of name: Named after Janet (Dami- ta Jo) Jackson, well-known American singer, songwriter, actress, dancer and sister of the late Michael Jackson after whom the first member of this genus was named. Type material: The holotype, and sole spec- imen known to date (MAB k.3623a, b), is a near-complete carapace of a maximum carapace length, as preserved, of 5.0 mm; the maximum shield width is 3.5 mm. Type locality and type level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 7; upper Tithonian (see Vašíček & Skupien, 2019, p. 39, fig. 3). Description: Well-calcified, smooth, areolat- ed shield, subcylindrical transversely, slight- ly convex longitudinally; pronounced, slightly downarched triangular rostrum, base wider than long, slender spinose tip; ocular-frontal area ex- ceeding half of total maximum width; orbital cavity subcircular, bounded by distinct triangu- lar postocular spines; thin, central gastric groove centrally indenting convex postfrontal ridge; gastric region of arrowhead shape, pointing pos- teriorly with a pair of gastric pits close to keraial region; elongated keraial region with straight lateral margin; prominent, reniform, globose massetic region; cardiac region anteriorly not delineated; elongated mesobranchial region with deep incision centrally running parallel to cer- vical groove; distinct U-shaped branchiocardi- ac groove, parallel to deep, V-shaped cervical groove. Remarks: Mesoparapylocheles janetjacksonae sp. nov. differs from all other Jurassic paguroids in the combination of an elongated keraial region with a straight, rather than convex, lateral mar- gin; a narrower reniform, rather than broader, trapezoidal, massetic region, as well as a very convex postfrontal ridge. The new species dif- fers from the mid-Cretaceous M. michaeljack- soni in having elongated keraial and massetic regions (rather than globose ones) and a cardiac region that is not posteriorly delineated as in M. michaeljacksoni. Family Pilgrimchelidae Fraaije, 2014 Genus Masticacheles Fraaije, Krzemiński, Van Bakel, Krzemińska & Jagt, 2014 Type species: Masticacheles longirostris Fraaije, Krzemiński, Van Bakel, Krzemińska & Jagt, 2014, by original diagnosis. Included species: Masticacheles longirostris and Masticacheles minimus Fraaije, 2014 and Masticacheles septemgradu sp. nov. Masticacheles septemgradu sp. nov. (Pl. 2.4) Diagnosis: Shield well calcified, well areo- lated, with distinct regions; convex orbital cav- 15Paguroid anomurans from the upper Tithonian–lower Berriasian of Štramberk, Moravia (Czech Republic) 1 - Protopagurus cerebellum sp. nov., original (right), composite (left); 2 - Housacheles timidus Fraaije, Van Bakel, Jagt & Skupien, 2013; 3 - Mesoparapylocheles janetjacksonae sp. nov.; 4 - Masticacheles septemgradu sp. nov.; 5 - Ammopylocheles mclaughlinae Van Bakel, Fraaije, Jagt & Artal, 2008, original (left), composite (right); 6 - Ammopylocheles romankijoki n. sp.; all scale bars 2 mm. PLATE 2 16 René H.B. FRAAIJE, Barry W.M. VAN BAKEL, John W.M. JAGT & Petr SKUPIEN ity with sharp postocular projection, convex post-rostral ridge centrally indented by long cen- tral groove; anterior part of gastric region cren- ulated; large, elongated massetic region; crescent keraial region; narrow anterior branchial area. Derivation of name: Named after ‘Level 7’ at Kotouč quarry (see e.g., Vašíček & Skupien, 2019, fig. 3); Latin septem and gradu, noun used in ap- position. Type material: The holotype, and sole speci- men known to date (MAB k.3757), is an incom- plete shield of a maximum carapace length, as preserved, of 2.5 mm; the maximum shield width is 2.5 mm. Type locality and type level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 7; upper Tithonian (see Vašíček & Skupien, 2019, p. 39, fig. 3). Description: Well-calcified and clearly areo- lated shield, convex transversely, slightly convex longitudinally; convex orbital cavity bordered with sharp postocular projection; ocular-frontal area equalling about 60 per cent of total maxi- mum width; convex post-rostral ridge centrally indented by long central groove; anterior part of gastric region crenulated; prominent, globose and elongated massetic region; crescent keraial region laterally convex with its anterior tip cen- trally/forwardly directed; relatively narrow an- terior branchial area; rostrum and posterior part of carapace not preserved. Remarks: Until now, the family Pilgrimchel- idae appeared to be confined to the Jurassic, to be replaced subsequently by, for instance, annun- tidiogenids. Masticacheles septemgradu sp. nov. can be differentiated from congeners by the typ- ical crescentic morphology of the keraial region, with its anterior tip directed centrally/forwardly rather than laterally/forwardly, as well as a nar- rower anterior branchial area. Family Pylochelidae Bate, 1888 Subfamily Trizochelinae Forest, 1987 Genus Ammopylocheles Van Bakel, Fraaije, Jagt & Artal, 2008 Type species: Ammopylocheles mclaughlinae Van Bakel, Fraaije, Jagt & Artal, 2008, by origi- nal designation. Included species: Ammopylocheles mclaugh- linae, Am. petersi Fraaije, 2014, Am. robertbore- ki Fraaije, Krzemiński, Van Bakel, Krzemińska & Jagt, 2012b and Am. romankijoki sp. nov. Ammopylocheles mclaughlinae Van Bakel, Fraaije, Jagt & Artal, 2008 (Pl. 2.5) Locality and level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 8; upper Tithoni- an (see Vašíček & Skupien, 2019, p. 39, fig. 3). Material: The specimen (MAB k.3760) is an incomplete shield with part of the posterior car- apace; as preserved, maximum carapace length measures 7.0 mm, maximum shield width is 5.5 mm. Remarks: For a detailed description, reference is made to Van Bakel et al. (2008). Members of the genus Ammopylocheles range from the mid- dle Oxfordian (Fraaije et al., 2012b) to the lower Berriasian (the present study). Ammopylocheles mclaughlinae is by far the commonest element in Kimmeridgian deposits at Nusplingen (Fraaije, 2014) and at Geisingen (Van Bakel et al., 2008) in southern Germany, but it is rather uncommon to rare at Ernstbrunn (Austria). The same appears to hold true for Štramberk. Ammopylocheles romankijoki n. sp. (Pl. 2.6) Diagnosis: Typical smooth carapace of py- lochelid; carapace longer than broad, shield of equal width and length; broad rostrum and con- vex, rimmed orbital cavity; pronounced post- frontal ridge, centrally indented by deep, short central gastric groove; elongated massetic region; reniform keraial region, distinct V-shaped cervi- cal groove. Derivation of name: Named after Roman Ki- jok (Poland), who collected the specimen and kindly donated it to the Oertijdmuseum, Boxtel. Type material: The holotype, and sole specimen known to date (MAB k.3758), is a near-complete shield with part of the posterior carapace, meas- uring 10.0 mm in maximum total length and 7.0 mm in width. Type locality and type level: Kotouč quarry (Štramberk, Moravia, Czech Republic), level 5; lower Berriasian (see Vašíček & Skupien, 2019, p. 39, fig. 3, locality 10; Vaňková et al., 2019, section B, layer B22). Description: Carapace longer than broad, shield as wide as long, strongly convex in trans- verse section, slightly convex in longitudinal sec- tion; broad rostrum posteriorly extending into pronounced central ridge, effacing towards cen- tral gastric groove; broad and convex, rimmed orbital cavity; postantennal projections obtuse; 17Paguroid anomurans from the upper Tithonian–lower Berriasian of Štramberk, Moravia (Czech Republic) transverse, convex, post-rostral ridge, with few large pores, medially subdivided by a short, deep, central gastric groove; elongated, more or less oval massetic region; subrounded keraial re- gion not well delineated, about one third size of massetic region; deep V-shaped cervical groove, posterior part of carapace less well calcified (partially preserved), smooth with irregularly distributed large (setal) pores. Remarks: This new species, of early Berria- sian age, is the youngest member of the genus. It differs from its middle Oxfordian congener A. robertboreki in having a larger, wider rostrum, a shorter central gastric groove and a more clear- ly V-shaped cervical groove. Ammopylocheles romankijoki sp. nov. differs from A. mclaughli- nae in having a much larger massetic region, a wider and more pronounced rostrum and a more angular V-shaped cervical groove. The new spe- cies differs from Am. petersi in having a much smaller and more subrounded keraial region, in lacking ornament on the anterior and posteri- or gastric regions and in having a more angular V-shaped cervical groove. Acknowledgements We wish to thank Dr Ewa Krzemińska (Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland) for bringing the specimen collected by Roman Kijok to our atten- tion, Roman himself for kindly donating it to the Oertijdmuseum, Boxtel, and Yvonne Coole for col- lecting and donating the new species of Masticacheles. We are very grateful to the management of Kotouč qu- arry for access to all levels in their quarry over re- cent years and the journal reviewers, Rok Gašparič (Ljubljana) and Guenter Schweigert (Stuttgart), for pertinent comments on an earlier version of the typescript. References Bate, C. S. 1888: Report on the Crustacea Macrura collected by the H.M.S. Challenger during the years 1873–1876. In: Report on the scienti- fic results of the voyage of H.M.S. Challenger during the years 1873-76, Zoology 24: 1–942. Forest, J. 1987: Les Pylochelidae ou «Pagures symétriques» (Crustacea Coenobitoidea). In: Crosnier, A. (ed.): Résultats des Campagnes MUSORSTOM, Volume 3. Mémoires du Muséum national d’Histoire naturelle, Série A137: 1–254. Fraaije, R. H. B. 2014: Diverse Late Jurassic ano- muran assemblages from the Swabian Alb and evolutionary history of paguroids based on carapace morphology. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 273: 121–145. Fraaije, R. H. B., Klompmaker, A. A. & Artal, P. 2012a: New species, genera and a fa- mily of hermit crabs (Crustacea, Anomura, Paguroidea) from a mid-Cretaceous reef of Navarra, northern Spain. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 263: 85–92. Fraaije, R. H. B., Krzemiński, W., Van Bakel, B. W. M., Krzemińska, E. & Jagt, J. W. M. 2012b: The earliest record of pylochelid hermit crabs from the Late Jurassic of southern Poland, with notes on paguroid carapace terminology. Acta Palaeontologica Polonica, 57: 647–654. Fraaije, R. H. B., Krzemiński, W., Van Bakel, B. W. M., Krzemińska, E. & Jagt, J. W. M. 2012c: The earliest record of a diogenid hermit crab from the Late Jurassic of the southern Polish Uplands, with notes on paguroid carapace terminology. Acta Palaeontologica Polonica, 57: 655–660. Fraaije, R. H. B., Krzemiński, W., Van Bakel, B. W. M., Krzemińska, E. & Jagt, J. W. M. 2014: New Late Jurassic symmetrical hermit crabs from the southern Polish Uplands and early paguroid diversifcation. Acta Palaeontologica Polonica, 59: 681–688. https://doi.org/10.4202/ app.2012.0022 Fraaije, R. H. B., Van Bakel, B. W. M. & Jagt, J. W. M. 2017: A new paguroid from the type Maastrichtian (Upper Cretaceous, the Netherlands) and erection of a new family. Bulletin de la Société géologique de France, 188/3. https://doi.org/10.1051/bsgf/2017185. Fraaije, R. H. B., Van Bakel, B. W. M., Jagt, J. W. M. & Artal, P. 2008: New decapod crustaceans (Anomura, Brachyura) from mid-Cretaceo- us reefal deposits at Monte Orobe (Navarra, northern Spain), and comments on related type-Maastrichtian material. In: Steurbaut, E., Jagt, J. W. M. & Jagt-Yazykova, E. A. (eds.): Annie V. Dhondt Memorial Volume. Bulletin de l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 78: 193–208. Fraaije, R. H. B., Van Bakel, B. W. M., Jagt, J. W. M., Klompmaker, A. A. & Artal, P. 2009: A new hermit crab (Crustacea, Anomura, Paguroidea) from the mid-Cretaceous of Navarra, northern Spain. Boletín de la Sociedad Geológica Mexicana, 61/2: 211-214. 18 René H.B. FRAAIJE, Barry W.M. VAN BAKEL, John W.M. JAGT & Petr SKUPIEN Fraaije, R. H. B., Van Bakel, B. W. M., Jagt, J. W. M. & Skupien, P. 2013: First record of paguroid anomurans (Crustacea) from the Tithonian-lower Berriasian of Štramberk, Moravia (Czech Republic). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 269/3: 251–259. https://doi. org/10.1127/0077-7749/2013/0348. Fraaije, R. H. B., Robins, C., Van Bakel, B. W. M., Jagt, J. W. M. & Bachmayer, F. 2019: Paguroid anomurans from the Tithonian Ernstbrunn Limestone, Austria – the most diverse extin- ct paguroid assemblage on record. Annalen des Naturhistorischen Museums in Wien, 121: 257–289. Gašparič, R., Fraaije, R. H. B., Van Bakel, B. W. M., Jagt, J. W. M. & Skupien, P. 2015: Mesozoic–Cenozoic crustaceans preserved within echinoid tests and bivalve shells. Bulletin of Geosciences, 90/3: 601–611. Houša, V. & Vašíček, Z. 2005: Ammonoidea of the Lower Cretaceous deposits (Late Berriasian, Valanginian, Early Hauterivian) from Štramberk, Czech Republic. GeoLines, 18 (for 2004): 7–57. Krzemińska, E., Krzemiński, W., Fraaije, R. H. B., Van Bakel, B. W. M. & Jagt, J. W. M. 2015: Allometric ontogenetic changes in two Late Jurassic gastrodorid hermit crabs (Crustacea, Decapoda, Anomura) from central Europe. Journal of Systematic Palaeontology, 14/2: 139-148. https://doi.org/10.1080/14772019.201 5.1018968 Latreille, P. A. 1802–1803: Histoire naturelle, générale et particulière, des Crustacés et des Insectes 3. F. Dufart, Paris: 1–467. Meyer, H. von. 1864: Briefliche Mittheilungen an Prof. H. B. Geinitz. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, 1864: 206–211. Milne Edwards, H. 1832: Extrait d’une lettre sur les caractères des Crustacés Anomures, adressée à M. Audouin. Annales des Sciences naturelles, Zoologie et Biologie Animale, 26: 255–256. Moericke, W. 1889: Die Crustaceen der Stramberger Schichten. Palaeontologische Mitteilungen aus dem Museum des königlich Bayerischen Staates, 3: 43–72. Ortmann, A. E. 1892: Die Decapoden-Krebse des Strassburger Museums, mit besonderer Berücksichtigung der von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln ge- sammelten und zur Zeit im Strassburger Museum aufbewahrten Formen, IV. Theil. Die Abtheilungen Galatheidea und Paguridea. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 6: 241–325. Van Bakel, B. W. M., Fraaije, R. H. B., Jagt, J. W. M. & Artal, P. 2008: An unexpected diversity of Late Jurassic hermit crabs (Crustacea, Decapoda, Anomura) in Central Europe. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 250: 137–156. Vaňková, L., Elbra, T., Pruner, P., Vašíček, Z., Skupien, P., Reháková, D., Schnabl, P., Košťák, M., Švábenická, L., Svobodová, A., Bubík, M., Mazuch, M., Čížková, K., & Kdýr, Š. 2019: Integrated stratigraphy and palaeoenvironment of the Berriasian pe- ri-reefal limestones at Štramberk (Outer Western Carpathians, Czech Republic). Palaeogeography, Palaeoclimatology, Palaeoecology, 532: 109256. https://doi. org/10.1016/j.palaeo.2019.109256 Vašíček, Z. & Skupien, P. 2004: Historie geolo- gických a paleontologických výzkumů svrch- nojurských a spodnokřídových sedimentů na Štramberku [The Štramberk fossil site (uppermost Jurassic/Lower Cretaceous, ou- ter Western Carpathians) – two centuries of the geological and paleontological resear- ch]. Sborník vědeckých prací Vysoké školy báňské, Technické univerzity Ostrava Řada hornicko-geologická, 50: 83–102. Vašíček, Z. & Skupien, P. 2005: Doplňky k histo- rii geologických a paleontologických výzku- mů na Štramberku [Supplements to history of geological and paleontological research of Štramberk territory]. Sborník vědeckých prací Vysoké školy báňské Technické univer- zity Ostrava, Řada hornicko-geologická, 51: 1–6. Vašíček, Z. & Skupien, P. 2019: New Tithonian and Lower Berriasian aptychi of Štramberk Limestone from the Kotouč Quarry (Outer Western Carpathians, Czech Republic). Paläontologische Zeitschrift, 93: 37–48. Vašíček, Z., Skupien, P. & Jagt, J. W. M. 2018: Current knowledge of ammonite assemblages from the Štramberk Limestone (Tithonian- Lower Berriasian) at Kotouč Quarry, Outer Western Carpathians (Czech Republic). Cretaceous Research, 90: 185–203.