Cainozoic Research, 2(l-2)(2002), pp. 109-116,October 2003

obesus a new of hermit Ciliopagurus , species Oligocene

from northwest Belgium

³ van Bakel John+W.M. ² & René+H.B. Barry+W.M. ¹, Jagt Fraaije

'Schepenhoek 235, NL-5403 GB Uden, the Netherlands:e-mail: [email protected]

2 Natuurhistorisch Museum Maastricht, de Bosquetplein 6, NL-6211 KJ Maastricht, the Netherlands:

e-mail: john.jagt@maastricht. nl

2 Oertijdmuseum de Groene Poort, Bosscheweg 80, NL-5283 WB Boxtel, the Netherlands: e-mail: [email protected]

Received 21 October 2002; revised version accepted 22 March 2003

The basal portion of the Belsele-Waas Clay Member (Boom Clay Formation, Rupelian, Oligocene) as exposed at the Scheerders van

Kerchove (SVK) clay pit, southwest ofSint-Niklaas (province of Oost-Vlaanderen,NW Belgium), has recently yielded materialofa diogenid

the with annulations and a stridulatory apparatus on the inner surface of cheliped. This is here described as a new taxon,

Ciliopagurus obesus n. sp., documentingone of the rare instances where a paguroid maybe assigned with certainty to an extant .

Previous records of fossil hermit (Paguroidea)are listed.

KEY WORDS:: Paguroidea, , Oligocene, Belgium, new species.

Introduction Of note is the fact that van Straelen (1921, p. 122) listed

but a single Cainozoic pagurid, Pagurus sp., from the ‘Le-

Although decapod remains abound in many dien’ (= Lede Formation, Lutetian, ) ofthe Brussels

Cainozoic strata in Belgium, in particular those of Eocene area, based on previous records by Galeotti (1837) and Le

and Oligocene age, there are but few modem papers de- Hon (1862). From much younger, Pliocene, strata in the

these notable Collins Smith scribing faunas; exceptions are & Antwerp area, van Bakel et al. (2000) listedPagurus sp.; this

(1993) and Verheyden (2002). From personal observations, is closely related to or conspecific with P. bernhardus Linne,

we know that both public (see also Feldmann & Dhondt, 1758 (see van Bakel et al, in press). The new diogenid

material 1991) and private collections contain abundant of described in the present note thus constitutes not only the

often exquisitely preserved specimens. In addition to the new thirdhermit crab on record from Cainozoic strata in Belgium,

diogenid hermit crab described in the present paper, a new but also the first that is formally named. It is to be expected

cancrid from the Lower/Middle Pliocene of Oelegem (van that there will be additionalrecords of Cainozoic pagurids

Bakel and and faunules from rich et al, 2003) anomuran brachyuran Belgium, once the collections of molluscs, includ-

from the Lillo Formation (Piacenzian, Pliocene) at Kallo ing bulk samples, in particular from Pliocene strata in the

(van Bakel et al, in press) will shortly be published. Other Antwerp area, have been meticulously screened. There is

and from the indirect evidence in the form of broken and/or abraded projects (e.g., linuparids brachyurans Ypresian or

(Eocene) of Marke, Oligocene stomatopods from Sint- encrusted/bored gastropod shells (compare Ehrenberg, 1931;

Niklaas) are underway. Walker, 1988, 1989, 1992), which are known from various

The most complete, albeit outdated, listing ofCainozoic localities and stratigraphic levels.

decapod from Belgium is still that by van

Straelen strata of he (1921). From Oligocene age, recorded

a single nephropid lobster (Homarus perceyi van Beneden, Fossil paguroid records crab 1872), a geryonid (Coeloma rupeliense Stainier, 1887)

well Portunus nodosus Hermit crabs as as van Beneden, 1883. According (Paguroidea) are obligatory occupants ofempty

to Verheyden (2002), the last-named taxon is a nomen gastropod and other molluscan shells, which explains why

nudum based the Stainier later ; on same material, (1887) their often dissociated remains are fairly common in shell-

erected its valid Coeloma These rich name, rupeliense. two deposits, such as coquinas. With the exception of a from the species Boom Clay Formation, also illustrated by handful ofrecords documenting in situ preservation offossil

have been within Geys & Marquet (1983, pi. 57, figs 1,2), recently pagurids molluscan shells (see e.g., Hyden & Forest, revised by Verheyden (2002). 1980; Feldmann & Keyes, 1992; Hu & Tao, 1996; - 110-

McLaughlin & Forest, 1997; Jagt et al, 2000; Karasawa, Angeli, 1995, 2000; Vicariotto, 1997);

2002; Fraaije, 2003), most occurrences comprise isolated Spain (Via, 1959; Sole & Via, 1989);

major claws (chelipeds). On account of their characteristic Hungary (Muller, 1975, 1979, 1984; Muller & Collins,

such claws and morphology, are easily recognised as pagurid 1991);

distinguished from associated anomuran and brachyuran Poland(Forster, 1979; Muller, 1996);

chelipeds. Kyushu, Boso Peninsula and Ainoshima Island, Japan

For the most recent discussion of fossil hermit crabs, (Karasawa & Inoue, 1992; Kato & Karasawa, 1998; reference is madehere to Schweitzer & Feldmann(2001) and Karasawa, 1993, 1997,2002);

Schweitzer et al. (2002). These authors notedthat the Pagu- Taiwan(Hu & Tao, 1996); roidea known in the fossil reord in Zealand as currently are urgent New (Hyden & Forest, 1980; McLaughlin &

need ofrevision; most fossil taxa have typically, and often Forest, 1997). routinely, beenreferred to the Diogenidae or Paguridae, thus

the level. Characters the masking true diversity at family impor- In present note we describe a new species of diogenid tant at the generic level include morphological features of fromRupelian (Oligocene) strata exposed near Sint-Niklaas,

dorsal carapace, abdomen, mouthparts, and appendages which we assign to the genus Ciliopagurus Forest, 1995a.

than the which (other chelipeds), are rarely preserved in Forest (1995a) subdivided the widely distributed diogenid

. genus Trizopagurus Forest, 1952 into the genera Trizopagu-

For most fossil hermit crabs, only (a) major claw(s) is rus Strigopagurus, and Ciliopagurus as based on the proper, ,

(are) available; however, based on claw shape, size and shape ofthe cephalothoracic shield, the ornament ofthoracic

ornament, a rough generic placement has generally proved appendages, the organisation of pleopods and the develop- possible. This is the case for pagurids which have ment of stridulatory structures on chelipeds. In a subsequent

beenrecorded from; paper (Forest, 1995b), he reassigned a species from

Hungary to Ciliopagurus, making reference to its stridulatory

and features. Alabama, Arkansas, Texas, Mississippi New Jersey The new species described here shows well-

(USA; see Rathbun, 1935; Stenzel, 1945; Roberts, developed stridulatory structures as well, and extends the

1962; Whetstone & Collins, 1982; Bishop, 1983a,b, range of the genus Ciliopagurus downwards for another 15

1986a, b); Ma.

Mexico (Vega et al, 1995);

Germany (Merlin, 1941);

northeast Belgium and the southeastNetherlands(Collins Systematic description

et al, 1995; Jagt et al, 2000, work under way);

France (de Tribolet, 1875); Abbreviation- Type specimens are deposited in the collec-

Spain (Sole & Via, 1989); tions of the Oertijdmuseum de Groene Poort, Boxtel (the

Antarctica (Aguirre-Urreta & Olivero, 1992; Olivero & Netherlands; abbreviation MAB).

Aguirre-Urreta, 1994).

Superfamily Paguroidea Latreille, 1802

These have been assigned to the genera Paguristes Dana, Family Diogenidae Ortmann, 1892

1851, Pagurus Fabricius, 1775 (s. lat), Palaeopagurus van Genus Ciliopagurus Forest, 1995a, p. 43

Straelen, 1925,Petrochirus Stimpson, 1858;Bishop (1983a)

and erected and — Cancer Bishop (1986b) new genera, Roemerus Type species strigatus Herbst, 1804, by original Parapaguristes, respectively. designation.

Cainozoic pagurids have mostly been assigned to the

Calcinus genera Anapagurus Henderson, 1888, Dana, 1851, Ciliopagurus obesus n. sp. Dardanus Paul’son, 1875, Diacanthurus McLaughlin &

Forest, 1997, Diogenes Dana, 1851, Eocalcinus Via, 1959, Figures 1,2

Pagurus (s. lat.), Paguristes, Petrochirus and Pylopagurus

A. Milne Edwards & Bouvier, 1891, often with a query. Such Diagnosis - Heterochelous diogenid, with identical shape

records include: and ornament of claws, stridulatory structures less pro-

nouncedin minorclaw; ornament oftransverse, more or less

Chiapas, Mexico (Vega et al, 2001); evenly distributed, entire striae, beaded in some larger

Jamaica (Morris, 1993; Collins et al, 1996; Collins & specimens.

Portell, 1998);

North and South and California — Carolina, Washington Derivation ofname Latinobesus, meaning thick, bulky, in

(USA; Rathbun, 1926; Blow & Manning, 1996; reference to the shape and size of the major claw.

Schweitzer & Feldmann, 2001);

Netherlands Material— the (Holthuis, 1949); Holotype, a leftpropodus, is MAB k.2365 (leg. Austria (Glaessner, 1928; Bachmayer, 1951); T. Lambrechts) (Figures 1/1-3; 2/1); paratype, a right cheli-

northern Italy (Beschin et al, 1994, 2000, 2002; de ped, is MAB k.2366 (leg. FH. Mollen)(Figure2/2, 3). -Ill -

1. MAB Member Figure Ciliopagurus obesus n. sp., k.2365 (holotype), basal portion of the Belsele-Waas Clay (Boom Clay Formation ;

SVK 1 - Rupelian, Oligocene) ofthe clay pit near Sint-Niklaas (Belgium); detail ofstridulatory apparatus (scale bar equals 1 mm); 2,

3 - inner and dorsal aspects, respectively, of left propodus (scale bar equals 2 mm) (SEM). - 112-

MAB and MAB basal of the Figure 2. Ciliopagurus obesus n. sp., k.2365 (holotype) k.2366 (paratype), portion Belsele-Waas Clay Member

(Boom Clay Formation ; Rupelian, Oligocene) ofthe SVK clay pit near Sint-Niklaas (Belgium); 1 - detail ofstridulatory apparatus

(holotype; scale bar equals 200 pm); 2, 3 - inner and dorsal aspects, respectively, of right chela (paratype; scale bar equals 1 mm) (SEM). - 113 -

and - The material & described Geography stratigraphy present comes Beurlen, 1929) was (see Muller, 1984), later from the basal of the Belsele-Waas Member portion Clay reassigned to Ciliopagurus (Forest, 1995b). The same strata

(Boom Clay Formation), as exposed at the Scheerders van yielded Dardanus hungaricus (Lorenthey in Lorenthey & Kerchove’s Fabrieken Sint- Verenigde (SVK) clay pit near Beurlen, 1929), a species subsequently also recorded from Niklaas (province of Oost-Vlaanderen, NW Belgium), and the Miocene of northern Spain (Muller, 1993). The oldest

collected Frederik H. Theo Lambrechts and record of was by Mollen, a striated paguroid, as yet specifically unnamed, is

Luc Anthonis. On calcareous nannofossil evidence that of Beschin al. (Steur- et (2000, p. 9, pi. 1, fig. 3, as Dardanus also this unit baut, 1986, 1992; see Laga et al., 2002), was sp.), who illustrateda specimen from the Eocene of northern dated as Early to MiddleRupelian (Oligocene) or zone NP23 Italy (see also de Angeli & Beschin, 2001).

More detaileddata be found in al. have (sensu Martini, 1971). may a Vega et (2001) recently noted that some fossil

et al. to which reference is the which paper by Vandenberghe (2002), species assigned to genusDardanus, amongst are made. D. hungaricus and D. arrosor (Herbst, 1794), are in needof

re-evaluation. The absence of apparent stridulatory structures

- First are Description chelipeds heterochelous, with the left and the fact that cheliped ornament differs, means that these claw larger than the right one. Shape and ornament of both two species cannot be assigned to Ciliopagurus.

claws identical, but stridulatory less in The taxon which most apparatus pronounced closely resembles C. obesus n. sp. the minor claw. Manus suboval with oblique upper margin, is C. substriatiformis, whose cheliped surface is also covered

surface in and with ciliated outer markedly convex longitudinal transverse striae, although these are more closely spaced sections. cross Lower propodus margin rounded, elevated and show more distinct tubercles (see Muller, 1984, pi. 17, towards of the fixed into distally tip finger which it curves figs 1-4). In the Flungarian species, striae extend straight towards the continuously. lower cheliped margin, whereas in C. obesus n.

Fixed about thirdof finger short, one palmar length, stout sp. they are curved towards the fixed finger. The outer

and face offixed like with surface in the is triangular; opposable finger spoon new species more convex than that in C.

of rounded. There is sharp cutting edges; tip finger an emar- substriatiformis, with a much broader, more intumed fixed

in the situated The gination cutting edge, proximally. finger. stridulatory ridges are longer and fewer in

The curved is the number than in the weakly dactylus broadly flattened, new species, and not as distinctly

opposable face armed with beaded ribs and cusped viewed grouped. from above. The cutting edge is straight with a weak notch

situated proximally, opposite the emargination on the fixed

finger. Acknowledgements

Interdigital margin slanted, slightly concave with an

indistinct indentationfor the articular boss of the dactylus. We thank P. Verhaert (director of SVK Verenigde The changeover of the interdigital margin with the upper Fabrieken, Sint-Niklaas) for allowing access to their clay pit, margin ofthe fixed is obscure viewed from above. The finger F.H. Mollen, T. Lambrechts, L. Anthonis and E. Wille for carpal articulationmargin is strongly curved, thickened. bringing material to our attention and for donating it, Dr J. The outer propodus surface is ornamented with trans- Forest (Museum national d’FIistoire naturelle, Paris) for less verse, more or evenly distributed, entire striae, which are supplying items ofliterature, Dr J. Herman (Institut royal des beaded in faintly larger specimens. Striae are slightly sinu- Sciences naturelles de Belgique, Brussels) for preparing

ous, running parallel to the carpal articulation margin, and photomicrographs, as well as Professor R.M. Feldmann curving towards the fixed finger, near the lower margin. Both (Kent State University, Kent, Ohio) and Dr P. Muller(Geo- and lower notched upper propodus margins are distinctly by Institute of for logical Hungary, Budapest) commenting on the the striae, upper more than the lower. earlier an typescript. The inner surface is convex and bluntly noded in trans-

section and in verse cross convex longitudinal section, with both fingers intumed. The lower halfofthe innersurface is References the smooth, upper half striatedand scaled.

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