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Tf^A^(J€, W^ COLLINS-EVOLUTION of ANOMALOCARIS f ^v^ Tf^A^(j€, W^ COLLINS-EVOLUTION OF ANOMALOCARIS . 1911a. Middle Cambrian Merostomata. Cambrian geology and Middle Cambrian, Bxu^ess Shale, British Columbia. Philosophical paleontology II. Smithsonian Miscellaneous Collections, 57:17-40. Transactions of the Royal Society of London, B271:1-43. 1911b. Middle Cambrian Holothurians and Medusae. Cambrian , AND D. E. G. BRIGGS. 1982. A new conundrum from the Middle geology and paleontology II. Smithsonian Miscellaneous Collections, Cambrian Burgess Shale. Proceedings of the Third North American 57:41-68. Paleontological Convention, Montreal, 2:573-575. 1911c. Middle Cambrian Annelids. Cambrian geology and pa­ , AND . 1985. The largest Cambrian animal, Anomalocaris, leontology II. Smithsonian Miscellaneous Collections, 57:109-144. Burgess Shale, British Coliunbia. Philosophical Transactions of the . 1912. Middle Cambrian Branchiopoda, Malacostraca, Trilobita Royal Society of London, B309:569-609. and Merostomata. Cambrian geology and paleontology II. Smithson­ WOODWARD, H. 1902. The Canadian Rockies. Part I: On a collection ian Miscellaneous Collections, 57:145-228. of Middle Cambrian fossils obtained by Edward Whymper, Esq., WmxEAVES, J. F. 1892. Description of a new genus and species of F.R.G.S., from Mount Stephen, British Columbia. Geological Mag­ Phyllocarid Crustacea from the Middle Cambrian of Moimt Stephen, azine, 4(9):502-505, 529-544. B.C. Canadian Record of Science, 5:205-208. WHrmNGTON, H. B. 1975. The enigmatic animal Opabinia regalis, ACX:EPTED 8 AUGUST 1995 /. Paleont., 70(2), 1996, pp. 293-296 Copyright © 1996, The Paleontological Society O022-3360/96/OO70-0293$03.0O A NEW TETHYAN MIGRANT: CRETACHLORODIUS ENCIENSISN, GEN., N. SP. (CRUSTACEA, DECAPODA), FROM THE MAASTRICHTIAKT^YPE AREA RENEH. Geo Centrum Brabant, St. Lambertuswi oxtel, The Netherlands ABSTRACT—A new genus and species of crab is described from the late Maastrichtian Meerssen Member of the Maastricht Formation at the type area of the Maastrichtian Stage, SE Netherlands. This new taxon could well be the ancestor of the xanthid Prochlorodius- Chlorodiella group and its occurrence provides additional evidence of a late Cretaceous radiation among the decapod crustaceans. INTRODUCTION are described by Collins et al. (in press). The new decapod data inuNG DEPOSITION of the upper part of the Maastricht For­ provide indications for an immigration event at the base of the D mation, a number of typically Tethyan organisms (e.g., Meerssen Chalk (Figiu-e 1). rudistid biviilves) first appeared in the Maastrichtian type area. TAXONOMY Among them, benthic foraminifera (Hofker, 1966; Villain, 1977) and ostracods (Bless, 1989; Deroo, 1966) are the best docu­ Order DECAPODA Latreille, 1803 mented groups. A major immigration of Tethyan thermophilic Infraorder BRACHYURA Latreille, 1803 foraminifera and ostracods is recorded between the top of the Section BRACHYRHYNCHIA Borradaile, 1907 Emael Member and the base of the Nekum Chalk (Liebau, 1978; Superfamily XANTHOIDEA Dana, 1851 Bless, 1989; Bless et al., 1993) (Figure 1). This interval is char­ Family XANTHIDAE Dana, 1851 acterized by a change in paleobathymetry from a quiet water, Genus CRETACHLORODIUS n. gen. sublittoral setting (>20 meters of depth) to a phytal platform Diagnosis. —Carapace sub-hexagonal, twice as broad as long, with high-energy conditions (depth fluctuating between 2 and with a pronounced postfrontal ridge, otherwise smooth; orbi- 20 meters) (Liebau, 1978; Sprechmann, 1981). Parts of the Nek­ tofrontal margin about half the carapace width, long orbital um and Meerssen Members are characterized by bryozoan-sea- margins with distinct fissures; anterolateral margin with tuber- grass associations (Voigt, 1981). The occurrence of encrusting culation and indefinite lobation terminating in forwardly di­ red algae, corals and rudistid bivalves indicates that water tem­ rected spines at the lateral angles. perature was in excess of 20°C. Etymology.—Indicating a Cretaceous member of the Pro- In 1987, Fraaye and Collins described a new calappid crab, chlorodius-Chlorodiella group. Prehepatus werneri, whose occurrence seemed to correspond Type species. — Cretachlorodius enciensis n. sp. with an increasing Tethyan influence in the Maastrichtian type area. The genus Prehepatus Rathbun, 1935, was hitherto known CRETACHLORODIUS ENCIENSIS new species only by five species, all from North America. That paper trig­ Figures 2.1-2.2 gered a revival of decapod studies in the type Maastrichtian Diagnosis. —As for genus. area. Almost all species decribed previously (Binkhorst, 1857, Material. —The holotype is the only specimen known; it is an 1861; Bosquet, 1854;Forir, 1887a, 1887b, 1889;Noethng, 1881; almost complete carapace (width 25 mm, length 12 mm) found Pelseneer, 1886) suffer from a lack of detailed stratigraphic data. in a fine-grained bioclastic limestone, about three meters above A new xanthid crab recently has been described by Jagt et al. the base of the late Maastrichtian Meerssen Member (Maastricht (1991) and ten new (two anomuran and eight brachyuran) taxa Formation) in the quarry ENCI. 2 km southwest from Maas- 294 JOURNAL OF PALEONTOLOGY, V. 70, NO. 2, 1996 MAASTRICHTIAN TYPE AREA FIGURE 7—Composite lithostratigraphic log of the Maastrichtian type area. Five key sections, all quarries, including quarry ENCI, are in­ dicated under exposures. A = immigration event of thermophilic benthic foraminifera and ostracoda, B = immigration event of Teth- yan decapod crustaceans, 1 = range of Xanthosia semiornata, 2 = occurrence of Cretachlorodius enciensis, 3 = range of Prehepatus wer- neri. tricht, The Netherlands. The specimen is deposited in the col­ FIGURE 2.1-2--Cretachlorodius enciensis holotype, collection Geo Cen­ lections of Geo Centrum Brabant, Boxtel, The Netherlands (MAB trum Brabant (MAB k.0014), view of carapace outline (1) and details k0014). of orbitofrontal area, showing fine granulation, orbital fissures and Description.—CsLvapace is sub-hexagonal in outline, twice as p)ostfrontal ridge (2). Bar scale is 1 cm. broad as long, widest about midlength. A pronounced, slightly lobate ridge extends from anterolateral angle transversely across entire carapace; ridge just posterior orbital furrows; interupted superficial resemblance in outline and ornamentation often masks medially by broad, shallow emargination. Carapace is smooth, a major phylogenetic gap. except for some granulation along anterolateral margin, kidney­ The dorsal postfrontal ridge, broad orbitoft-ontal margin with like cardiac furrows, and two gastric pits. a median depression, and the anterolateral spinose margin of Orbitofrontal margin about half carapace width, front me­ the new genus resemble the xanthids Prochlorodius spp. and dially broadly emarginate and sulcate. Two fissures in finely Coral licarcinus spp., recorded from Eocene rocks of Hungary ridged, subcircular orbital margins, orbits directed obliquely (Miiller and Collins, 1991) (Figure 3); however, the former genus away from long axis. A short, distinct ridge midway between has a greater carapace length and the latter has more than one inner orbital angle and inner fissure runs parallel with front. frontal ridge; both have more equal-sized anterolateral spines, Anterolateral margin granulated, posteriorly tending to tu- a significantly broader orbitofrontal margin and lack fissures in berculation and indefinite lobation. At lateral angles, postfrontal the orbital margins. Miocene species of Chlorodiella Rathbun, ridge terminates in prominent, forwardly directed spine with incipient spine behind. Posterolateral margin smooth and si­ nusoidal; distinct, narrow ridge bounding concave, posterior margin. liiocene ChLorodieUa / Etymology.—The specific name alludes to its occurrence in / the ENCI quarry. Eocene N^ptocarcinus CoraUicarcinus Prochlorodius Remarks. —Cretachlorodius enciensis n. sp. exhibits a com­ bination of morphological characters which suggests placement yaastrichtian Xa'ni?u)sia semiornata Cretachlorodvus within the Xanthidae. The interfamilial relationships of this \ large family are complex, which holds particularly true for ex­ Cenomanlan Xanttu>sia buchii tinct genera for which further studies are required (Glaessner, I 1969, p. R515). In prehminary studies of several Recent genera of the Xan­ FIGURE i—Possible evolutionary relationships of Cretachlorodius and thidae, Guinot (1967, 1969) pointed out that, for example, a related genera. FRAA YE-MAASTRICHTIAN DECAPOD 295 1897, and extant species ofLeptodius (A. Milne Edwards, 1863), Cretachlorodius enciensis have been foimd whereas its closest Cataleptodius Guinot, 1967 and Gaudichaudia Rathbun, 1930 congener, Xanthosia semiornata, is known from a dozen more differ in having distinct furrows, especially in the mesogastric or less complete carapaces and many fragments. An interpre­ region, and a multiple-lobed anterolateral margin. tation of Cretachlorodius enciensis being a drifted, Tethyan jel- Cretachlorodius enciensis n. sp. differs from the most closely lyfish-hitchhicker could explain the single, atypical occurrence. resembling portunid, Neptocarcinus spp. in having a frontal ridge and lacking a distinct, lobed anterolateral margin. This resem­ CONCLUSIONS blance could be only superficial (Miiller and Collins, 1991) but Cretachlorodius enciensis seems to have an important evo­ more material and fUrther studies are needed to clear this matter. lutionary position and is a late, but not last (Fraaye, in prep.) The genus most closely approaching Cretachlorodius
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