Atti Soc. it. Sci. nat. Museo civ. Stor. nat. Milano, 150 (II): 229-238, Giugno 2009 Alessandro Garassino* **, Antonio De Angeli" & Giovanni Pasini" In situ hermit crab (Crustacea, Anomura, Paguroidea) from thè Early Eocene (Ypresian) of NE Italy Abstract - Hermit crabs from thè Eocene of Italy have heen reported by several authors who described only fragmentary and complete chelae of chelipeds. The studied specimen from thè Early Eocene (Ypresian) of Monte Baldo (Verona, NE Italy) has been ascribed to Paguristes Dana, 1851, with P. baldoensis n. sp. (Diogenidae Ortmann, 1892). It represents thè first report of hermit crab in situ from Italy. Key words: Crustacea, Anomura, Paguroidea, Early Eocene, Italy. Riassunto - Paguro in situ (Anomura, Paguroidea) dell’Eocene inferiore (Ypresiano) dell’Italia nordorientale. Paguridi dell'Eocene d’Italia sono stati segnalati da alcuni autori che hanno descritto solo chele complete o incomplete dei chelipedi. L’esemplare studiato dell'Eocene inferiore (Ypresiano) del Monte Baldo (Verona, NE Italia) è stato ascritto a Paguristes Dana, 1851, con P. baldoensis n. sp. (Diogenidae Ortmann, 1892). Si tratta della prima segnalazione di un paguride in situ per l’Italia. Parole chiave: Crustacea, Anomura, Paguroidea, Eocene inferiore, Italia. Introduction As reported by Glaessner (1969), fossil hermit crabs are usually known only for thè strongly calcified parts, like chelipeds and ambulatory legs, while thè shield is rarely preserved. Shafer (1972) and later Walker (1992) pointed out that “as a rule, hermit crabs are not preserved within thè gastropod shed”. In fact, thè fossil record of hermit crabs preserved in situ is exceptionally rare, limited to date to about ten specimens formerly recognized, that ranges from thè Early Jurassic to thè Pliocene, presumably due to a previous selective sampling in fìeld prospections or, according to van Bakel et al. (2008), for merely inattention rather than of ready paucity. * Museo di Storia Naturale di Milano, Corso Venezia 55, 20121 Milano, Italy, e-mail: [email protected] ** Associazione Amici del Museo “G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy, e-mail: [email protected] ”* Museo Civico dei Fossili di Besano, Via Prestini 5, 21050 Besano (Varese), Italy, e-mail: [email protected] 230 .ALESSANDRO GARASSINO, ANTONIO DE ANGELI & GIOVANNI PASINI Jagt et al. (2006) reported thè earliest hermit crab in situ referred to Palaeopa- gurus Van Straelen, 1925, including a probable new species, from thè upper Plien- sbachian (Early Jurassic) of southern Germany, preserved in a fragmentary body chamber of thè amaltheid ammonite Pleuroceras solare. van Bakel et al. (2008) reported Ammopylocheles mclaughlìnae van Bakel, Fraaije, Jagt & Aitai, 2008, preserved within thè body chamber of an empty peris- phinctid ammonite shell from thè Kimmeridgian (Early Jurassic) of southern Ger¬ many. Fraaije (2003) reported Palaeopagurus vandenengeli Fraaije, 2003, preserved within thè shell of thè ammonite Simbirskìtes gottscheì from thè middle Hauteri- vian (Fower Cretaceous) of United Kingdom. Breton & Collins (2007) reported a single specimen with chelipeds and ambula- tory legs, poorly preserved and very compressed, of an indeterminate IPagurus sp. within a ?natacid gastropod from thè Cenomanian (Fate Cretaceous) of Fe Mans (France). Feldmann et al. (1993), described Paguristes santamartaensis Feldmann, Tshudy & Thomson, 1993, by two chelipeds associated with an extemal mould of thè gastropod Taioma from thè ?late Santonian- middle Campanian (Fate Creta¬ ceous) of Antarctica. Fraaije et al. (2008) reported Anmmtidìogenes sunuciorum Fraaije, van Bakel, Jagt & Artal, 2008, preserved within a volutoid gastropod from thè Maastrichtian (Fate Cretaceous) of The Netherlands. Collins & Jakobsen (2003) described Pagurusl langei Collins & Jakobsen, 2003, within thè body chamber of Volutocorbis sp. from thè Eocene (Ypresian/ Futetian) of Fillebaelt Clay Formation of Jutland (Denmark). Jagt et al. (2006) reported chejipeds and ambulatory legs of an indeterminate paguroid (“Pagurus damesii” Noetling, 1885, nomea dubium) within a inter¬ nai mould of a ?volutid gastropod from thè Middle Eocene of eastem Prussia (Russia). Karasawa (2002) reported remains of pereiopods 1-3 of Pagurus (s.l.) sp. pre¬ served within Turritella ìnflralirata from thè Early Fate Oligocene of Japan. Hyden & Forest (1980) reported Pagurus clifdenensis Hyden & Forest, 1980, within a damaged shell of thè gastropod Struthiolaria subspinosa from thè Early Miocene of New Zeland. Finally, Dunbar & Nyborg (2003) reported thè discovery of three specimens of hermit crabs, ascribed to Isocheles sp., associated with their host gastropod shell from thè Pliocene of thè San Diego Formation of California (United States). Geological setting The mountain chain of Monte Baldo, between Verona and Trento provinces (NE Italy), expands from N to S. It is included between thè eastem bank of Garda Fake and thè right orographic bank of Adige River. The geological section of thè eastem side of thè mountain chain includes some sedimentary series from thè Fate Jurassic to thè Fate Eocene (Fabiani, 1915; Fuciani, 1987, 1989). A continuous series of marly limestones and Eocene marls of marine origin crops out dose to thè village of Ferrara del Baldo (Verona) (Fig. 1). The bottom of thè series has been referred to thè Early Eocene, including different invertebrates, such as brachyurans belonging to Harpactocarcinus and echinoderms belonging to Pentacrinus (Fabiani, 1915). INSITI' HERMIT CRAB FROM THE EARLY EOCENE (YPRESIAN) OF NE ITALY 231 Fig. 1 - Geographical map with fossiliferous locality (*) mappa geografica con ubicazione della località fossilifera (*). The studied specimen was discovered in this level and therefore ascribed to thè Early Eocene (Ypresian), also for thè presence of one fragmentary specimen of Pentacrinns cfr. P. diaboli, typical of thè Early Eocene (Fabiani, 1915; Siliotti, 1971), associated with thè studied specimen within thè body chamber of thè coeval cerithid Pseudovertagus. Previous reports of hermit crabs from thè Eocene of Italy As reported bv De Angeli & Garassino (2006) and Beschin et al. (2007), thè tossii record of Paguroidea from thè Eocene of Italy includes only Diogenidae Ortmann. 1892. Diogenidae includes Calcinus Dana, 1851, Ciliopagums Forest. 1995, Darda- mis Paulson. 1875, Diogenes Dana, 1851, Eocalcinus Via Boada, 1959, Paguri- stes Dana. 1851, and Petrochirus Stimpson, 1859. Only fragmentary and complete chelae of chelipeds are known from thè Eocene of Italy to date for this family, as reported by Beschin et al. (1994, 2002, 2005, 2006, 2007), De Angeli (1995), and Vicariotto (1997). So thè studied specimen represents thè first report of hermit crab in situ from Italy, enlarging also thè very poor knowledge about thè life-style ways among thè tossii paguroids. Material The studied specimen is preserved in situ in an inner mould of gastropod. pro- bably belonging to Pseudovertagus (Striovertagus) striatus (Brughiere, 1792), a cerithid usually common from thè late Ypresian to middle Lutetian of Veneto (Qua- giotto, pers. comm., 2008). Even though we suppose that thè complete shield is 232 ALESSANDRO GARASSINO, ANTONIO DE ANGELI & GIOVANNI PASINI preserved within thè body chamber of thè second tum of thè shell, only thè chelae and thè right carpus of chelipeds were cleaned after thè preparation by air gravers due to thè hard nature of thè carbonatic mould of thè gastropod shell. The studied specimen has been ascribed to thè infraorder Anomura MacLeay, 1838, including Paguristes baldoensis n. sp. (Diogenidae Ortmann, 1892). The specimen is housed in thè Palaeontological Codections of thè Museo di Storia Naturale, Milano (MSNM). Measurements are given in millimetres (mm). Systematic Palaeontology Order Decapoda Latreille, 1802 Infraorder Anomura MacLeay, 1838 Superfamily Paguroidea Latreille, 1802 Family Diogenidae Ortmann, 1892 Included fossil genera: Annuntidiogenes Fraaije, van Bakel, Jagt & Aitai, 2008 (late Albian - Spain; Maastrichtian - The Netherlands); Calcinus Dana, 1851 (Lutetian - Italy; Pliocene - Italy); Ciìiopagurus Forest, 1995 (Oligocene - Bel- gium; Badenian - Hungary); ?Cìibanarius Dana, 1852 (?Late Eocene - Egypt); Dardanus Paulson, 1875 (Eocene - United States and Egypt; Oligocene - Asia; Lutetian - Italy; Miocene - Europe; Pliocene - Italy and Fiji; Pleistocene - United States); Diogenes Dana, 1851 (Ypresian - Italy); Eocaìcinus Via Boada, 1959 (Eocene - Spain and Italy); Eopaguropsis van Bakel, Fraaije, Jagt & Artal, 2008 (late Oxfordian - Germany); Paguristes Dana, 1851 (?late Santonian-middle Cam- panian - Antarctic; Campanian, Maastrichtian - United States; Maastrichtian - The Netherlands; Paleocene - United States; Lutetian and Priabonian - Italy; Miocene - United States); Petrochirus Stimpson, 1859 (Late Cretaceous - United States; Lutetian and Priabonian - Italy; Oligocene - W India; Miocene - Panama, N Ame¬ rica, and N Africa; Pliocene - N America); Striadiogenes Garassino, De Angeli & Pasini, 2009 (Cenomanian - Lebanon) (updated after Glaessner, 1969). Genus Paguristes Dana, 1851 Type species: Paguristes hirtus Stimpson, 1858. Included Italian fossil species: P. extentus Beschin, Busulini, De Angeli & Tes- sier, 2007 (Early Eocene - Ypresian); P. prealpinus Beschin, De Angeli, Checchi & Zarantonello, 2005 (Middle Eocene - Lutetian); P ìineatuberculatus Beschin, De Angeli, Checchi & Mietto, 2006 (Late Eocene - Priabonian). Paguristes baldoensis n. sp. Figs. 2-4 Diagnosis: chelae subequal, subrectangular in outline; left chela sbghtly longer and larger than right chela; right chela elongate with upper and lower margins almost rectilinear; coirugated upper margin of paini; lower margin of palm straight and smooth; movable and fìxed fìngers elongate and strong; corrugated upper margin of IXSITI' HERMIT CRAB FROM THE EARLY EOCENE (YPRESIAN i OF NE ITALY 233 movable fìnger; interdigital margin strongly inclined; articulation carpo-propoidale straight; omamentation of chela with strong tubercles not aligned. Etymology: thè trivial name alludes to Monte Baldo where thè studied speci¬ men was discovered. Holotvpe: MSNM Ì27220. Geological age: Early Eocene (Ypresian). Type locality: Ferrara di Monte Baldo (Verona). Fig. 2 - Paguristes baldoensis n. sp., MSNM Ì27220. holotvpe olotipo (x 2) 234 ALESSANDRO GARASSINO, ANTONIO DE ANGELI & GIOVANNI PASINI Occurrence: one specimen in situ that preserves only thè chelipeds. Length of carpus of right chela: 4 Length of right chela: 8 Length of palm of right chela: 5 Width of palm of right chela: 4 Length ofleft chela: 10 Length of palm ofleft chela: 5 Width of palm ofleft chela: 5 Length of fìxed fìnger of right chela: 4 Length of movable fìnger of right chela: 6 Maximum length of shell: 60 Maximum width of shell: 20 Description. Left and right chelae subequal in size, subrectangular in outline, strongly broader proximally; left chela slightly longer and larger than right chela; thè upper margin is slightly curved and corrugated; thè lower margin is straight and smooth, continuous with thè stout fìxed fìnger and smooth from thè tip of thè fìxed fìnger to thè carpai margin; thè palm is subquadrate in outline; thè interdigital margin is strongly inclined, smooth, with two slight depressions divided by a mediai slight bulge; thè fìxed finger is almost as long as thè palm with thè tip shallowly spooned; thè movable finger is stout, elongate, with corrugated upper margin and lateral pits; thè outer surface of thè left and right chelae is gently convex with strong tubercles not aligned and uniformly distributed; thè articulation carpo-propoidale is straight; thè carpus is subquadrate in outline with thè carpus-merus articulation less wide than thè carpo-propodiale articulation; upper margin of carpus smooth and gently inflated laterally. Discussion. As reported by many authors, thè systematics of Recent Paguroi¬ dea uses essentially morphological dorsal and ventral characters of thè body to distinguish genera and species of each family. Since, thè studied specimen does not preserve dorsal or ventral characters useful for its identification, only thè shape of thè chelae of chelipeds and thè omamentation of chelipeds can be used for its taxonomic ascription. As reported by McLaughlin (2003) thè superfamily Paguroidea includes at present thè following Recent families, Coenobitidae Dana, 1851, Pylochelidae Bate, 1888, Diogenidae Ortmann, 1892, Lithodidae Samouelle, 1819, Pylojac- quesidae McLaughlin & Lemaitre, 2001, Paguridae Latreille, 1802, and Parapagu- ridae Smith, 1882. Comparing thè generai morphology of thè chelipeds of thè studied specimen with thè diagnostic characters of these families (McLaughlin & Lemaitre, 2001; Poore, 2004), it is assigned to thè Diogenidae and to Paguristes Dana, 1851, for thè chelae subequal, thè palm subquadrate having a corrugated upper margin, and thè interdigital margin strongly inclined. In thè fossil record from thè Eocene of Veneto, Paguristes is represented with three species, P. prealpiniis Beschin, De Angeli, Checchi & Zarantonello, 2005, P. ìineatubercuìatus Beschin, De Angeli, Checchi & Mietto, 2006, and P. extentus Beschin, Busulini, De Angeli & Tessier, 2007. A morphological resemblance exists between thè studied specimen and P extentus for thè elongate outline of thè chela and thè interdigital margin strongly inclined and smooth. However, two characters distinguish thè new species from P extentus, thè stout fìxed fìnger almost as long as thè palm with thè tip shallowly spooned and thè corrugated upper margin of INSITI1HERMIT CRAB FROM THE EARLY EOCENE (YPRESIAN) OF NE ITALY 235 thè paini. The elongate chelae subequal with corrugateci upper margin, thè elon- gate movable fìnger with corrugateci upper margin, thè interdigital margin strongly inclined, smooth, with two slight depressions divided by a mediai slight bulge, and thè omamentation of paini with strong tubercles not aligned distinguish P bal- doensis n. sp. from P. prealpinus and P. lineatuberculatus, both having a fìxed fìnger with distai tip directed downwards, interdigital margin slightly inclined and straight, and omamentation of paini with small tubercles. Fig. 3 - Paguristes baldoensis n. sp., MSNM Ì27220. A) detail of thè chelae / dettaglio delle chele. B) reconstruction of thè chelae / ricostruzione delle chele, (x 6). 236 ALESSANDRO GARASSINO, ANTONIO DE ANGELI & GIOVANNI PASINI Taphonomic and palaeoecological notes The studied specimen shows both chelipeds exposed, ranged subparallel in lateral view and in life position. The originai shell, probably belonging to Pseudo- vertagus (Striovertagus) striatus, was dissolved during thè diagenesis process and only thè generai form of thè helicoidal spire, laterally compressed, and thè outline development are visible, as internai mould. This kind of preservation allows to recognize thè presence of thè chelae into thè shell. Recent species of Pseudovergatus live on sandy bottoms and soft sediments in subtropical intertidal-sublitoral and brackish waters near thè costai line. A similar environment is supposed for thè Eocene in Veneto area, intercalated with locai important volcanic events. Moreover, some Recent small sized hermit crabs living in thè Mediterranean Sea, such as Clibanarius erytropus, have a preference to inhabit thè empty shells of several Cerithiidae in addition to a few other gastropods (Grecchi & Balestrazzi, 2002). Other extra-Mediterranean examples are thè “hermitted” shells (sensi/ Walker, 1992) of Recent Cerithium stercusmuscarum from thè Cortez Sea (Mexico) (Walker, 1992). As reported by Tunberg et al. (1994), also thè small shells of Cerithium muscarum, in thè estuarine subtropical environment of thè Indiana River Lagoon on thè Atlantic coast of Florida (United States) are observed as an important habitat resource for Pagurus maclaughinae. Another example is Diogenespugilator Roux, 1828, inhabiting thè shells of Ceritium vulgatum along thè Coastal environment of thè Crimea Peninsula in thè Black Sea. Acknowledgements We wish to thank R. H. B. Fraaije, Oertijdmuseum De Groene Poort, Boxtei, The Netherlands, for careful review and criticism, E. 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