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Harpactocarcinus from the Eocene of Istria, Croatia, and the Paleoecology of the Zanthopsidae Via, 1959 (Crustacea: Decapoda: Brachyura)

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Harpactocarcinus from the Eocene of Istria, Croatia, and the Paleoecology of the Zanthopsidae Via, 1959 (Crustacea: Decapoda: Brachyura) J. Paleont., 79(4), 2005, pp. 663±669 Copyright q 2005, The Paleontological Society 0022-3360/05/0079-663$03.00 HARPACTOCARCINUS FROM THE EOCENE OF ISTRIA, CROATIA, AND THE PALEOECOLOGY OF THE ZANTHOPSIDAE VIA, 1959 (CRUSTACEA: DECAPODA: BRACHYURA) CARRIE E. SCHWEITZER,1 VLASTA C OSOVIC ,2 AND RODNEY M. FELDMANN3 1Department of Geology, Kent State University Stark Campus, Canton, Ohio 44720, ,[email protected]., 2Department of Geology and Paleontology, Faculty of Science, University of Zagreb, Croatia, ,[email protected]., and 3Department of Geology, Kent State University, Kent, Ohio 44242, ,[email protected]. ABSTRACTÐHarpactocarcinus punctulatus istriensis Bachmayer and Nosan, 1959 is elevated to species level. Analysis of the larger foraminiferans associated with specimens of H. istriensis suggests a habitat preference for off-shore, clear, shelf environments below fair-weather wave base and an age of early to middle Lutetian (Eocene). A review of the paleoenvironmental indicators for nearly all species within the genera referred to the Zanthopsidae Via, 1959 suggests that all exhibit similar habitat preferences. Description of the paleoenvironmental preference for an entire extinct decapod family has not before been possible. INTRODUCTION The ``marls with crabs'' have been assigned a lower middle OSSIL DECAPOD crustaceans have been reported from various Eocene, speci®cally middle Lutetian, age in the FloricÏicÂi region localities in Istria (Croatia and Slovenia) on several occa- (JuracÏicÂ, 1979), from which we collected specimens of Harpac- F  sions; however, these reports have been limited to nineteenth- tocarcinus (Drobne et al., 1979; Pavlovec et al., 1991; Cosovic century and early twentieth-century descriptions (Bittner, 1883, and Drobne, 1998). Subsequently, an early Lutetian age was as- 1884, 1893; Stache, 1889; Toniolo, 1909; Vogl, 1912), anecdotal signed to the ``marls with crabs'' in the regions of Buzet and RocÏ reports, and short notes (Salopek, 1954; Bachmayer and Nosan, (Pavlovec and PavsÏicÏ, 1986) at localities from which we also 1959; Tarlao, 2000; MikuzÏ, 2002) (Table 1). Toniolo (1909) de- collected specimens of Harpactocarcinus. Bergant et al. (2003) scribed numerous localities in Croatia from which decapods were assigned a middle Lutetian age for the sediments in the Pazin collected, giving them Italian names that are no longer used (Table Basin, which encompasses central to northeastern Istria. The Gra- 2). The region would be expected to contain a robust decapod cÏisÏcÏe, Slovenia, sediments, from which we collected additional fauna, due to the presence of a well-developed carbonate ramp in specimens of Harpactocarcinus, belong to the Brkini ¯ysch com- Ï the area during the middle Eocene (C osovic et al., 2004) and the plex (Savrin Littoral Basin) and are of early Eocene (Ypresian) robust Eocene decapod record in nearby Hungary (MuÈller and age according to the Geological Map, Sheet Trieste 1:100,000 Collins, 1991) and Spain (Via, 1959, 1969). However, the deca- (PlenicÏar et al., 1973), or are Lutetian (Pavlovec and PavsÏicÏ, 1986; pod record in Istria is surprisingly sparse (Table 1). All of the PavsÏicÏ and Peckmann, 1996). Our analyses of sediments collected fossil decapods previously reported from localities in Istria are with specimens of Harpactocarcinus con®rm the age differences brachyurans, and the overwhelming majority of the occurrences between the GracÏisÏcÏe, Slovenia, and FloricÏicÂi, Croatia, samples are of species of Harpactocarcinus A. Milne Edwards, 1862 and and those from Buzet-RocÏ, Croatia. The depth of deposition of Harpactoxanthopsis Via, 1959, both members of the extinct fam- these ``marls with crabs'' is estimated to have been from 60 to ily Zanthopsidae Via, 1959. Specimens of Lophoranina sp. are 100 m in the lower ``marls with crabs,'' deepening to up to 1,000 also common (Bittner, 1893). Harpactocarcinus sp. and Harpac- m in the upper part of the ``marls with crabs'' (JuracÏicÂ, 1979). toxanthopsis sp. are so common that the layers in which they are The GracÏisÏcÏe packstones contain a rich foraminiferal assem- found are known in the literature as the ``marls with crabs.'' The blage, composed of larger and planktonic foraminifera along with ¯ysch deposits overlying the ``marls with crabs'' contain trace authigenic glauconite (foraminiferan identi®cations after Toumar- fossils referrable to Ophiomorpha Lundgren, 1891 and Thalassi- kine and Luterbacher, 1985; Less, 1987). Larger foraminifera as- noides Ehrenberg, 1944 (Tunis and Uchman, 1996), but no deca- semblages with ¯at nummulitids, discocyclinids such as Disco- pod fossils have yet been reported from the ¯ysch sediments. cyclina dispansa (Sowerby, 1840) and D. archiaci (Schlumberger, Limestones of the underlying carbonate ramp sediments are pri- 1903), and operculinids are characteristic of low-energy, lower marily packstones containing numerous foraminiferans but they photic conditions. The planktonic foraminifera Morozovella leh- do not contain decapod fossils. neri (Cushman and Jarvis, 1929) and ``Globigerinatheka'' senni (Beckmann, 1953) are typical of the P-11 Biozone (sensu Berg- gren et al., 1995) of Lutetian age. These sediments were deposited DEPOSITIONAL SETTING in a quiet, open-marine, outer ramp setting with a signi®cant pe- The unit from which the specimens of Harpactocarcinus de- lagic in¯uence, but that was shallow enough to support larger scribed herein were collected has variously been called the ``marls foraminifera in the lower photic zone. The presence of glauconite with crabs'' or the ``transition beds'' (JuracÏicÂ, 1979; Tunis and indicates deeper water and low sedimentation rates, with less ox- Uchman, 1996; Bergant et al., 2003). These beds lie between the ygenated conditions (Odin and Matter, 1981). foraminiferal (Nummulites±Orthophragmina) limestones, the plat- The Buzet sample is composed of packstones in which larger formal sediments, and the ¯ysch deposits, the basin sediments, of foraminifera outnumber the planktonic ones, and glauconite the Istrian Peninsula. Most of the specimens of Harpactocarcinus grains are present in signi®cant quantity. Planktonic foraminifera and Harpactoxanthopsis recovered from the ``marls with crabs'' are present which are typical of the biostratigraphic zone P-11 (s. are corpses, because they retain the ventral portion of the carapace Berggren et al., 1995) of middle Lutetian age, including Globi- as well as poorly preserved appendages; thus, we can infer that gerinatheka mexicana (Cushman, 1925), Turborotalia frontosa the crabs were living in the described environments, and were not (Subbotina, 1953), Turborotalia possagnoensis (Toumarkine and transported into the environment as buoyant molts of the cara- Bolli, 1970), and Subbotina inaequispira (Subbotina, 1953). Larg- pace. er foraminifera are well preserved and discocyclinids dominate, 663 664 JOURNAL OF PALEONTOLOGY, V. 79, NO. 4, 2005 TABLE 1ÐFossil decapods previously reported from Istria and localities in Croatia other than Istria. Decapod taxon Localities References Achelous krambergeri Bittner, 1893 Croatia Bittner, 1893 Calappa sp. Weber, 1795 Istria Toniolo, 1909 Harpactoxanthopsis quadrilobatus (Desmarest, 1822) Istria, Croatia Bittner, 1883; Toniolo, 1909; Vogl, 1912 (Drvenik); Tar- lao, 2000; MikuzÏ, 2002 Harpactocarcinus punctulatus (Desmarest, 1822) Istria Bittner, 1883; Toniolo, 1909 Harpactoxanthopsis souverbiei (A. Milne Edwards, 1862) Istria Toniolo, 1909 Lophoranina marestiana (KoÈnig, 1825) Istria Bittner, 1883 Mioplax socialis Bittner, 1884 Croatia Bittner, 1884 Neptunus radobajanus Bittner, 1884 Croatia Bittner, 1884 Neptunus stenaspis Bittner, 1884 Croatia Bittner, 1884 including Discocyclina radians (d'Archiac, 1850). Their abun- SYSTEMATIC PALEONTOLOGY dance implies an open marine setting with winter water temper- Order DECAPODA Latreille, 1802 8 atures greater than 16 C (Hollaus and Hottinger, 1998). The depth Infraorder BRACHYURA Latreille, 1802 is recognized as that of the lower photic zone, the reproductive Section HETEROTREMATA Guinot, 1977 depth for planktonic foraminifera. During Eocene time, when the Superfamily XANTHOIDEA MacLeay, 1838 studied area was located around 358N paleolatitude (Butterlin et Family ZANTHOPSIDAE Via, 1959 al., 1993a, 1993b), the estimated depth of the euphotic zone was Included genera.Harpactocarcinus A. Milne Edwards, 1862; about 100 m in clear water with no terrestrial input. This is sug- Harpactoxanthopsis Via, 1959; Martinetta Blow and Manning, gested by the presence of glauconite grains. Oligotrophic condi- 1997; Neozanthopsis Schweitzer, 2003; Zanthopsis McCoy, 1849. tions are inferred because of the presence of larger foraminifera. Discussion.Schweitzer (2003) raised Via's (1959) Zanthop- The FloricÏicÂi packstones contain rich planktonic foraminiferal sinae to family status and reevaluated all species previously re- associations. Fragments of discocyclinids are scattered throughout ferred to Zanthopsis and Harpactocarcinus. Therein are diagnoses the micritic matrix. The foraminifera are typical of the middle for the family and the various genera it contains. Lutetian (P-12 Biozone, s. Berggren et al., 1995), including Mo- rozovella lehneri, Dentoglobigerina yeguaensis (Weinzierl and Genus HARPACTOCARCINUS A. Milne Edwards, 1862 Applin, 1929), and Turborotalia frontosa. The foraminifera col- Harpactocarcinus A. MILNE EDWARDS, 1862, p. 64, pls. 3±6, 8±10. lected by JuracÏic (1979), at the site of our collection, allow
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