In Situ Hermit Crab (Anomura, Paguroidea) from the Oligocene Pysht Formation, Washington, USA

N. Jb. Geol. Paläont. Abh. 295/1 (2020), 17–22 Article E Stuttgart, January 2020

In situ hermit crab (Anomura, Paguroidea) from the Oligocene Pysht Formation, Washington, USA

Giovanni Pasini, Alessandro Garassino, Torrey Nyborg, Stephan G. Dunbar, and René H.B. Fraaije With 2 figures

Abstract: An unusual fossil in situ paguroid hermit crab is here reported from the Oligocene Pysht Formation, Washington (USA). Paguroid specimens preserved within their host gastropod shells are rare in the fossil record. Only a few reports of in situ paguroid hermit crabs preserved within their host gastropod shells have been reported from the Cenozoic.

Key words: Crustacea, Decapoda, Anomura, Paguroidea, Oligocene, Pysht Formation, USA.

1. Introduction 2. Geological setting

We report on a paguroid crab specimen preserved in The Pysht Formation is one of three including the Hoko situ within a shell of Priscofusus stewarti Tegland, River, Makah, and Pysht formations, which are part of 1933 (Gastropoda, Fusinidae). This is the first record the Twin River Group, an Eocene to Oligocene deposit of a hermit crab from the Oligocene Pysht Formation, exposed in a northward-dipping homoclinal sequence Washington (USA). Although the preservation of the along the northwestern coast of the Olympic Peninsula studied specimen does not allow a precise systematic of Washington (Snavely et al. 1978; Prothero et al. assignment, this report is important due to the scarce 2009). The Twin River Group unconformably overlies samples of fossil hermit crabs preserved in situ within pillow basalts and breccias of the Crescent Volcanics their host gastropod shell. So far, relatively complete in (MacLeod & Snavely 1973; Snavely & MacLeod situ Cenozoic paguroids have been described from the 1977; Snavely et al. 1978). The Pysht Formation was Eocene of Austria (Fraaije & Polkowsky 2016), Italy named after the Pysht River, which flows into the (Garassino et al. 2009) and Russia (Jagt et al. 2006), Strait of Juan de Fuca at Pillar Point (Snavely et al. from the Oligocene of Japan (Karasawa 2002), and 1978). The Pysht Formation is exposed for 18 km from from the Miocene of New Zealand (Hyden & Forest Pillar Point State Park east to approximately 3.5 km 1980) and The Netherlands (Jagt et al. 2006). west of Low Point along the Strait of Juan de Fuca near the northwest tip of the Olympic Peninsula in Clallam County, Washington as shore-cliff exposures and wave-cut terraces (Gower 1960; Addicott 1976; Snavely et al. 1978) (Fig. 1). The Pysht Formation consists of gray to olive-gray massive mudstone, san-

eschweizerbart_xxx 18 G. Pasini et al.

Fig. 1. Generalized locality map. X marks the area where the paguroid hermit crab specimen described herein was collected.

dy siltstone, with conglomerate and sandstone inter- Goedert 1996; Schweitzer & Feldmann 1999; beds (Snavely et al. 1978). Thickness ranges from East 2006), and marine tetrapods (Olson 1980; 1,100 to 1,400 m (Snavely et al. 1978) to up to 2000 m Barnes et al. 1994; Dyke et al. 2011; Mayr & (Prothero et al. 2001). Based on magnetostratigra- Goedert 2016; Peredo & Uhen 2016; Vélez- phy, the lower Pysht Formation has been correlated Juarbe 2017) have been previously described from with Chrons C11r–C8r (26.5–30.5 Ma) and the upper- the Pysht Formation. most Pysht Formation with Chron C6Cr and C6Cn3n (23.7–24.7 Ma), making the Pysht Formation late early Oligocene to latest Oligocene in age (Prothero et al. 2001). Foraminiferal biostratigraphy place the Pysht 3. Material Formation in the Oligocene Zemorrian Stage (Nes- bitt et al. 2010). Foraminifera indicate an outer slope One small specimen including pereiopods, dorsally com- to bathyal depositional environment in cold water pressed, crushed in dorsal view and preserved in situ with- (Goedert et al. 2003; Peckmann et al. 2003). in a Priscofusus stewarti gastropod shell. The specimen is housed at The Burke Museum of Natural History and Cul- The Pysht Formation is highly fossiliferous, with ture, Invertebrate Paleontology Collections, University of many fossils contained within calcareous concre- Washington, Seattle, Washington, USA. tions. Benthonic foraminifera (Rau 1964), mol- Abbreviations: UWBM IP – The Burke Museum of Natural luscs (Durham 1944; Addicott 1976; Squires History and Culture, Invertebrate Paleontology Collections, 1989; Squires & Goedert 1994; Kiel & Goedert University of Washington, Seattle, Washington, USA. P1– 2007), crustaceans (Rathbun 1926; Berglund & P3 first to third pereiopods.

eschweizerbart_xxx In situ hermit crab (Anomura, Paguroidea) from the Oligocene Pysht Formation 19

Fig. 2. Paguroid hermit crab specimen preserved in a Priscofusus stewarti gastropod shell, UWBM IP 110567. C: carapace, P1–P3: first to third pereiopods. Scale bar equals 1 cm.

4. Systematic paleontology Locality: The sole specimen was collected from wave-cut terraces of the Pysht Formation near the mouth of Deep Creek at UWBM locality no. B9416 (Fig. 1). Order Decapoda Latreille, 1802 Infraoder Anomura MacLeay, 1838 Geological age: Oligocene. Superfamily Paguroidea Latreille, 1802 Measurements: Carapace shield not measurable; Prisco- Family, genus and species indet. fusus stewarti gastropod shell – total length: 13 mm; max- Fig. 2 imum width: 7 mm (at the last coil). Description: Carapace shield very crushed, poorly pre- Examined material: UWBM IP 110567, one crushed and served, partially covered dorsally by the inhabited shell; poorly preserved specimen including pereiopods preserved it seems longer than wide. ?Right P1 cheliped with sub- in situ within a Priscofusus stewarti gastropod shell. triangular carpus; sub-rectangular P1 palm dorsal and ventral margins smooth, slightly convex; triangular index rounded distally; dactylus slender, index incurved downward; oc-

eschweizerbart_xxx 20 G. Pasini et al. clusal margin of P1 dactylus straight, with alternate small Dunbar & Nyborg (2003) found that if buried rapid- rounded teeth; stout triangular P1 index, with some posterior ly with dry sand, hermit crabs did not abandon their rounded teeth bigger than those of dactylus; smooth outer shells and died where buried. In contrast, when buried surface of P1 palm and carpus; ?left cheliped covered by right cheliped; left and right P2–P3 equal, exceeding che- slowly, P. samuelis dug up through the sediment, often liped; P2–P3 elongate, smooth, with a transverse median de- carrying the shell with them. In additional experiments pression along length of merus and carpus; elongate, curved with the same species, but with combined sand and dactylus. P4–P5 not observables. Cephalic appendages not seawater, Shives & Dunbar (2010) found the orien- preserved. tation of the aperture during burial was a significant Discussion: The systematics of extant Paguroidea makes factor in whether hermit crabs abandoned the shell or use of morphological characters such as antennules, pleo- not, with hermit crabs buried aperture up much more pods, maxillipeds or pereiopods, and several internal and likely to abandon. Those authors suggested that when ventral characters to distinguish genera and species of each family whereas the dorsal carapace or chelipeds proxy the aperture was down, shell architecture may cause characters (sensu Schweitzer 2003), mainly preserved as sediment to flow over the shell and settle around the fossils, are mostly used on fossil specimens (Fraaije et al. organism allowing the crab to push upward against the 2017). Though the P1 cheliped is preserved, it does not al- sediment overburden. In contrast, when the aperture low us to recognise its homo- or heterochely. Therefore, the faces upward, sediment can fill the internal volume of lack of above-mentioned diagnostic characters in the studied specimen hampers its systematic assignment both to extant the shell. This may cause the hermit crab to extend and fossil families and genera described to date among the outward, allowing more sediment to infiltrate the shell Paguroidea. The only possible remark is that the apparent and eventually inhibiting the hermit crab from retreat- absence in the studied specimen of any kind of ornamen- ing inward. This may result in the hermit either suc- tation on the preserved P1–P3 and the P1 probably being cumbing to anoxia while partially extended from the isochelous could suggest that it does not belong to the Dio- genidae and Paguridae, which usually have more or less shell, or abandoning the shell altogether in an effort to ornamentated pereiopods and P1 heterochelous (Zariquiey escape the burial. In either case, the hermit crab may Alvarez 1968; McLaughlin 2003). then be deposited partially or fully disarticulated from The studied specimen is preserved in situ within a small the shell. It is therefore not surprising that other au- gastropod shell, slightly deformed due to the different thors, such as Walker (1992), found fossil pagurized hardness of the mineralized shell, assigned to Priscofusus gastropod shells lacking the associated pagurid. stewarti due to its similarities to described species of Prisci- fusus from the overlying Clallam Formation (Addicott 1976). This is the first report of a pagurid hermit crab from the Oligocene Pysht Formation of Washington. Acknowledgements

We wish to thank John Cornish of Port Angelas, Washing- 5. Taphonomy ton, who collected and kindly donated the sole specimen for research. Joachim Haug of the Ludwig-Maximilians- University of Munich Munich, Bavaria, Germany, took Twenty-five paguroid specimens, many preserving the color image of the specimen used in this paper. Austin both chelipeds and in some cases the walking legs, Hendy and Shawn Wiedrick, Natural History Museum of were reported from the underlying Late Eocene Hoko Los Angelas County, helped with the identification of the River Formation (Schweitzer & Feldmann 2001). gastropod. We also thank Guenter Schweigert, State Mu- Preservation of a paguroid associated within its host seum of Natural History Stuttgart, Germany, and Carrie Schweitzer, Kent State Univeristy, Ohio, USA, for their gastropod shell is scarce in the fossil record (see Jagt kind reviews. et al. 2006 and references therein). However, empty gastropod shells with signs of hermit crab inhabitation are quite common in the fossil record (e.g., Walker 1992; Feldmann et al. 2019). References Dunbar & Nyborg (2003) speculated that the paucity of fossil hermit crabs associated with gastropod Addicott, W.O. (1976): Molluscan paleontology of the shells is due to hermit crabs abandoning their host lower Miocene Clallam Formation, northwestern Wash- gastropod shells and climbing upward through the ington. – U.S. Geological Survey, Professional Paper, sediment during times of stress, such as burial. In pre- 976 : 44 pp. doi: 10.3133/pp976 liminary experiments with shelled Pagurus samuelis, Barnes, L.G., Kimura, M., Furusawa, H. & Sawamura, H. (1994): Classification and distribution of Oligocene Ae-

eschweizerbart_xxx In situ hermit crab (Anomura, Paguroidea) from the Oligocene Pysht Formation 21

tiocetidae (Mammalia; Cetacea; Mysticeti) from western new records. – Revista Mexicana de Ciencias Geológicas, North America and Japan. – Island Arc, 3 (4): 392–431. 23 (3): 364–369. doi: 10.1111/j.1440-1738.1994.tb00122.x Karasawa, H. (2002): Fossil uncinidean and anomalan De- Berglund, R.E. & Goedert, J.L. (1996): A new crab capoda (Crustacea) in the Kitakyushu Museum and In- (Brachyura: Cancridae) from Lower Miocene rocks of stitute of Natural History. – Bulletin of the Kitakyushu the northwestern Olympic Peninsula, Washington. – Museum of Natural History, 21: 13–16. Journal of Paleontology, 70 (5): 830–835. doi: 10.1017/ Kiel, S. & Goedert, J.L. (2007): New mollusks associated S0022336000023866 with biogenic substrates in Cenozoic deep-water sedi- Dunbar, S.G. & Nyborg, T. (2003): Three specimens of her- ments of Washington State. – Acta Palaeontologica Po- mit crabs found associated with their gastropod shell from lonica, 52 (1): 41–52. the Pliocene San Diego Formation, California and reas- Latreille, P.A. (1802): Histoire naturelle, générale et par- sessment of the paucity of fossil hermit crabs associated ticulière, des crustacés et des insects, 3: 468 pp.; Paris with gastropod shells in the fossil record. – Geological (Dufart). Society of America, Abstract with Program, 35: 56. MacLeay, W.S. (1838): On the brachyurous decapod Crusta- Durham, J.W. (1944): Megafaunal zones of the Oligocene cea brought from the Cape by Dr. Smith. – In: Smith, A. of northwestern Washington. – University of California (ed): Illustrations of the Annulosa of South Africa; being Publications, Bulletin of the Department of Geological a portion of the objects of natural history chiefly collect- Sciences, 27 (5): 101–212. ed during an expedition into the interior of South Africa, Dyke, G.J., Wang, X. & Habib, M.B. (2011): Fossil Plo- under the direction of Dr. Andrew Smith, in the years topterid Seabirds from the Eo-Oligocene of the Olympic 1834, 1835, and 1836; fitted out by “The Cape of Good Peninsula (Washington State, USA): Descriptions and Hope Association for Exploring Central Africa”: 53–71 Functional Morphology. – PLoS One, 6 (10): e25672. pp.; London (Smith, Elder, and Co.). doi: 10.1371/journal.pone.0025672 PMID: 22065992 MacLeod, N.S. & Snavely Jr., P.D. (1973): Volcanic and East, E.H. (2006): Reconstruction of the Fossil Mud Shrimp intrusive rocks of the central part of the Oregon Coast Callianopsis clallamensis. – Journal of Crustacean Biol- Range. – In: Geologic Field Trips in Northern Oregon ogy, 26 (2): 168–175. doi: 10.1651/C-2562.1 and Southern Washington. – Oregon Department of Ge- Feldmann, R.M., Schweitzer, C.E. & Phillips, G.E. ology and Mineral Industries Bulletin, 77: 47–74. (2019): Oligocene pagurized gastropods from the River Mayr, G. & Goedert, J.L. (2016): New late Eocene and Bend Formation, North Carolina, USA. – Bulletin of the Oligocene remains of the flightless, penguin-like plo- Mizunami Fossil Museum, 45: 7–13. topterids (Aves, Plotopteridae) from western Washing- Fraaije, R.H.B. & Polkowsky, S. (2016): Dardanus col- ton State, U.S.A. – Journal of Vertebrate Paleontology, osseus, a new paguroid from the Eocene of Austria pre- 36 (4): e1163573. doi: 10.1080/02724634.2016.1163573 served in its host gastropod shell. – Neues Jahrbuch für McLaughlin, P.A. (2003): Illustrated keys to families and Geologie und Paläontologie, Abhandlungen, 279 (1): genera of the superfamily Paguoridea (Crustacea: De- 57–62. doi: 10.1127/njgpa/2016/0540 capoda: Anomura), with diagnoses of genera of Pagu- Fraaije, R.H.B., van Bakel, B.W.M. & Jagt, J.W.M. ridae. – Memoirs of Museum Victoria, 60 (1): 111–144. (2017): A new paguroid from the type Maastrichtian doi: 10.24199/j.mmv.2003.60.16 (Upper Cretaceous, the Netherlands) and erection of Nesbitt, E.A., Martin, R.A., Carroll, N.P. & Grieff, J. a new family. – Bulletin de la Société géologique de (2010): Reassessment of the Zemorrian foraminiferal France, 188: 155–158. doi: 10.1051/bsgf/2017185 Stage and Juanian molluscan Stage north of the Olympic Garassino, A., De Angeli, A. & Pasini, G. (2009): In situ Mountains, Washington State and Vancouver Island. – hermit crab (Crustacea, Anomura, Paguroidea) from the Newsletters on Stratigraphy, 43 (3): 275–291. doi: Early Eocene (Ypresian) of NE Italy. – Atti della Società 10.1127/0078-0421/2010/0043-0275 italiana di Scienze naturali e del Museo civico di Storia Olson, S.L. (1980): A new genus of penguin-like pelecan- naturale in Milano, 150 (2): 229–238. iform bird from the Oligocene of Washington (Pele- Goedert, J.L., Thiel, V., Schmale, O., Rau, W.W., Mi- caniformes: Plotopteridae). – Contributions in Science, chaelis, W. & Peckmann, J. (2003): The Late Eocene Natural History Museum in Los Angeles County, 330: ‘Whiskey Creek’ Methane-Seep Deposit (Western 51–57. Washington State) Part I: Geology, Palaeontology, and Peckmann, J., Goedert, J.L., Heinrichs, T., Hoefs, J. & Molecular Geobiology. – Facies, 48 (1): 223–240. doi: Reitner, J. (2003): The Late Eocene ‘Whiskey Creek’ 10.1007/BF02667541 Methane-Seep Deposit (Western Washington State). Gower, H.D. (1960): Geological map of the Pyshy quad- Part II: Petrology, Stable isotopes, and Biogeochemis- rangle, Washington. – Geological Survey, Geologic try. – Facies, 48 (1): 241–253. doi: 10.1007/BF02667542 Quadrante 129. doi: 10.3133/gq129 Peredo, C.M. & Uhen, M.D. (2016): A new basal chaeo- Hyden, F.M. & Forest, J. (1980): An in situ hermit crab mysticete (Mammalia: Cetacea) from the Late Oligo- from the Early Miocene of southern New Zealand. – Pa- cene Pysht Formation of Washington, USA. – Papers in laeontology, 23 (2): 471–474. Palaeontology, 2 (4): 533–554. doi: 10.1002/spp2.1051 Jagt, J.W.M., van Bakel, B.W.M., Fraaije, R.H.B. & Neu- Prothero, D.R., Bitboul, C.Z., Moore, G.W. & Moore, E.J. mann, C. (2006): In situ fossil hermit crabs (Paguroidea) (2001): Magnetic stratigraphy of the early and middle from northwest Europe and Russia. Preliminary data on Miocene Astoria Formation, Newport Embayment, Ore- gon. – In: Prothero, D.R. (ed.): Magnetic Stratigraphy

eschweizerbart_xxx 22 G. Pasini et al.

of the Pacific Coast Ceonozoic: Pacific Section. – Society shells. – Journal of Paleontology, 66: 535–558. doi: for Sedimentary Geology, Book 91: 272–283. 10.1017/S0022336000024410 Prothero, D.R., Draus, E. & Burns, C. (2009): Magne- Zariquey Alvarez, R. (1968): Crustáceos Decápodos Ibéri- tostratigraphy and Tectonic Rotation of the Eocene- cos. – Investigación Pesquera, 32: 499 pp. Oligocene Makah and Hoko River Formations, Northwest Washington, USA. – International Journal of Geophys- ics, 2009: ID 930612. doi: 10.1155/2009/930612 Rathbun, M.J. (1926): The fossil stalk-eyed Crusta- Manuscript received: November 14th, 2019. cea of the Pacific slope of North America. – United Revised version accepted by the Stuttgart editor: Novem- States National Museum Bulletin, 138: 1–155. doi: ber 27th, 2019. 10.5479/si.03629236.138.i Rau, W.W. (1964): Foraminifera from the northern Olympic Peninsula, Washington. – U.S. Geological Survey, Pro- fessional Papers, 374-G: 33 pp. Schweitzer, C.E. (2003): Utility of proxy characters for classification of fossils: an example from the fossil Xan- thoidea (Crustacea: Decapoda: Brachyura). – Journal of Addresses of the authors: Paleontology, 77 (6): 1107–1128. doi: 10.1666/0022-3360 (2003)077<1107:UOPCFC>2.0.CO;2 Giovanni Pasini, Via Alessandro Volta 16, 22070 Ap- Schweitzer, C.E. & Feldmann, R.M. (1999): Fossil de- piano Gentile (Como), Italy; capod crustaceans from the Late Oligocene to Early Miocene Pysht Formation and Late Eocene Quimper e-mail: [email protected] Sandstone, Olympic Peninsula, Washington. – Annals of Alessandro Garassino, Torrey Nyborg (corre- Carnegie Museum, 68 (4): 215–273. Schweitzer, C.E. & Feldmann, R.M. (2001): New Cre- sponding author), Department of Earth and Biological taceous and Tertiary decapod crustaceans from western Sciences, Loma Linda University, Loma Linda, CA North America. – Bulletin of the Mizunami Fossil Mu- 92350, USA; seum, 28: 173–210. e-mails: [email protected]; Shives, J.A. & Dunbar, S.G. (2010): Behavioral responses [email protected] to burial in the hermit crab, Pagarus samuelis: Impli- cations for the fossil record. – Journal of Experimental Stephen G. Dunbar, Marine Research Group, De- Marine Biology and Ecology, 388 (1–2): 33–38. doi: partment of Earth and Biological Sciences, Loma Lin- 10.1016/j.jembe.2010.03.008 da University, Loma Linda, CA 92350, USA; e-mail: Snavely Jr., P.D. & MacLeod, N.S. (1977): Evolution of the Eocene continental margin of western Oregon and [email protected] Washington. – Geological Society of America, Abstract René H.B. Fraaije, Oertijdmuseum, Bosscheweg 80, with Programs, 9 (7): 1183. Snavely Jr., P.D., Niem, A.R. & Pearl, J.E. (1978): Twin 5283 WB Boxtel, The Netherlands; River Group (Upper Eocene to Lower Miocene) –defined e-mail: [email protected] to include the Hoko River, Makah, and Pysht Forma- tions, Clallam County, Washington. – In: Sohl, N.F. & Wright, W.B. (eds.): Changes in stratigraphic nomen- clature by the U.S. Geological Survey, 1977. – U.S. Geo- logical Survey Bulletins, 1457-A: A111–A120. Squires, R.L. (1989): Pteropods (Mollusca: Gastropo- da) from Tertiary Formations of Washington and Ore- gon. – Journal of Paleontology, 63 (4): 443–448. doi: 10.1017/S0022336000019673 Squires, R.L. & Goedert, J.L. (1994): A new species of the volutid gastropod Fulgoraria (Musashia) from the Oli- gocene of Washington. – The Veliger, 37 (4): 400–409. Tegland, N.M. (1933): The fauna of the type Blakeley upper Oligocene of Washington. – University of California publications. Bulletin of the Department of Geology, 23 (3): 81–174. Vélez-Juarbe, J. (2017): A new stem odontocete from the late Oligocene Pysht Formation in Washington State, U.S.A. – Journal of Vertebrate Paleontology, 37 (5): e1366916. doi: 10.1080/02724634.2017.1366916 Walker, S.E. (1992): Criteria for recognizing marine hermit crabs in the fossil record using gastropod

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