EOCENE ISOPODS OF PESCIARA DI BOLCA (ITALY) REVISITED

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1 JOURNAL OF CRUSTACEAN BIOLOGY, 35(4), , 2015 EOCENE ISOPODS OF PESCIARA DI BOLCA (ITALY) REVISITED Ronald Vonk 1,2,, Leonardo Latella 3, and Roberto Zorzin 3 1 Naturalis Biodiversity Center, Department of Marine Zoology, Darwinweg 2, 2333 CR Leiden, The Netherlands 2 Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, Science Park 904, 1098 XH Amsterdam, The Netherlands 3 Museo Civico di Storia Naturale of Verona, Lungadige Porta Vittoria, 9, I Verona, Italy ABSTRACT The scarcity of the isopod material of the fossil sites from the Pesciara di Bolca (Verona Province, Northern Italy) did not leave much room up to now for conclusions concerning the systematic relationship of this fauna. In this study, we reinvestigate fossils of the cymothoidan Palaega acuticauda Secretan, 1975 and the sphaeromatidean Heterosphaeroma veronensis Secretan, We argue that both species do not differ substantially in dorsal morphology from recent genera, and we place them among more likely congeners: Cirolana acuticauda n. comb., and Dynamenella veronensis n. comb. A detailed redescription of both species is made on the basis of additional material that came from Pesciara di Bolca and Monte Postale marine fish and terrestrial plant bearing layers. Two environmental scenarios are discussed: one in which the palaeoenvironment is thought to be mainly lacustrine, another in which discoveries of more fossil sphaeromatideans and other cymothoidans in Italy suggest seawater affinities, coral reef environments, or estuarine conditions. KEY WORDS: Cymothoida, Sphaeromatidea, Ypresian, lacustrine, lagoon, coral reef environment DOI: / X INTRODUCTION The Lower Eocene isopod fossils from the Pesciara di Bolca (Verona Province, Northern Italy) were described by Secretan as part of her very well-documented and complete work on Les Crustacés du Monte Bolca (Secretan, 1975). There was very little isopod material at her disposal at that time. She placed one specimen in the popular fossil genus Palaega Woodward, 1870 and four other specimens in Heterosphaeroma Munier-Chalmas, 1872, an undiagnosed genus with one representative from lacustrine deposits of late Paleocene age of the Champagne region in France (Munier-Chalmas, 1872; van Straelen, 1928). With additional material incorporated at a later date into the palaeontological collections of the Museo Civico di Storia Naturale of Verona from Pesciara di Bolca and Monte Postale, it is now possible to reinvestigate the systematic affinities of these fossils. New material combined with the old specimens allows us to extend the species descriptions for two of the three species mentioned by Secretan from the Veneto region, Northern Italy. It also requires a new combination of names that place them in the cirolanid genus Cirolana Leach, 1818, and in the sphaeromatid genus Dynamenella Hansen, 1905, respectively. A third poorly preserved specimen that Secretan studied (1975: 318) was designated by her as Sphaeroma sp.; we have no more insights to add to that specimen. In addition, recent palaeo-environmental insights into the biostratigraphy of the study area have revealed a more de- tailed picture of local circumstances in a coastal area (Trevisani et al., 2005; Papazzoni et al., 2006, 2014; Schwark et al., 2009). Benthic crustaceans such as the isopods studied here form a minor part of the fossil assemblage that mainly consist of the famous Pesciara di Bolca fishes. The environmental requirements of the isopods as they are perceived today by studying related taxa can help elucidate microhabitat conditions at the time of burial. SYSTEMATICS Isopoda Latreille, 1817 Suborder Scutocoxifera Dreyer and Wägele, 2002 Infraorder Cymothoida Wägele, 1989 Cirolanidae Dana, 1852 Cirolana, Leach, 1818 Cirolana acuticauda (Secretan, 1975) n. comb. (Figs. 1-2) 1975 Palaega acuticauda Secretan, p Material Examined. Holotype, specimen no 13, Museo Civico di Storia Naturale, Verona, consisting of a negative (Fig. 1A, B) and a positive part (Fig. 2A, B), with some features present on one side only. Body length 21 mm, width 9 mm. Additional material: 2 more specimens, IG positive (Fig. 1C) and VR negative (Fig. 1D) and positive. IG body length 19 mm, width 8 mm. VR body length 18 mm, width 8 mm. Diagnosis. Small cirolanid, mildly flattened dorsoventrally; cephalon not completely embedded, pereiopods 6 and Corresponding author; ronald.vonk@naturalis.nl The Crustacean Society, Published by Brill NV, Leiden DOI: / X

2 VONK ET AL.: EOCENE ISOPODS FROM ITALY 541 Fig. 1. Cirolana acuticauda (Secretan, 1975), no. 13 MCSNV, holotype. A, negative part, numbers 1-5 outline pleonite segments; Ex, exopod of uropod; En, endopod of uropod; Pl3, pointed extension of pleonite 3. B, idem, anterior part with Antl, antennule; Fr. lam, frontal lamina; Antn, antenna; Da, dactylus of pereiopod 3 or 4; Pr, propodus; B, basis. C, IG 91160, positive part, pleonites 1-5; Ex, exopod of uropod; En, endopod of uropod. D, VR 27746, Ceph, cephalon, arrows indicate posterior margin; Pn 1, pereionite; Pn 2, pereionite 2. Scale bars = 2 mm. 7 slender; pleotelson slightly longer than the 5 pleonites combined, posterior margin rounded but ending in an ogival arch ( acuticauda ); uropodal endopods slightly protruding past posterior margin of pleotelson; entire body covered with small fine grained tubercles, resembling a puckered surface.

3 542 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 35, NO. 4, 2015 Fig. 2. Cirolana acuticauda (Secretan, 1975), no. 13 MCSNV, holotype. A, positive part; P6, pereiopod 6; P7, pereiopod 7; Pt, pleotelson; Ex, exopod of uropod; En, endopod of uropod. B, outline of divides between body segements. C, Dynamenella veronensis (Secretan, 1975) nov. comb. IG , negative part, numbers 1-7 mark the pereionites; Pl 1-3, pleonites; Ex, exopod of uropod; En, endopod of uropod; Pt, pleotelson. D, IG , positive part, Pn, pereionite; Pl 1-3, pleonites; Ex, exopod of uropod; En, endopod of uropod. Scale bars = 2 mm. Revised Description. Cephalon (Figs. 1D; 2A) not embedded in pereionite 1, but well fitted and forming a rounded frontal section. Frontal lamina slightly protruding (Fig. 1B). Eyes present, probably on the place of hollows near the frontal margin of cephalon, facets not preserved (Fig. 2A).

4 VONK ET AL.: EOCENE ISOPODS FROM ITALY 543 Antennule represented by one fragment in the counterpart of the holotype (Fig. 1B). Antenna with broad peduncular segment and a tapering flagellum with a visible length of 3 mm (Fig. 1B). Pereionite 1 axially wider than the following 6, lateral margin not in any place clearly demarcated, coxal plates not visible. Pereionites 2-7 irregular in axial width, possibly caused by telescopic displacement, distal margins without spines or knobs. Coxal plates observable on left side of pereionite 5, 6, and 7 (Fig. 2A). Pereiopods: details of limb morphology not observable. Pereiopods 6 and 7 slender. Pleon with five distinct pleonites (Fig. 1A, C), the fifth axially longer than previous four. Four pleonites similar in length; pleurae thin and pointed (Fig. 1A). Pleotelson about as wide as long, tapering distally, rounded triangular; posterior margin rounded, ending in an ogival form (Fig. 2B). Uropods pronounced; exopod narrower than endopod, tapering distally without crenulations or indented margins; endopods more rounded, slightly longer than pleotelson, also no ornamented margins. Remarks. As Secretan (1975) stated herself, the assignment of the Monte Bolca fossils to Palaega was a matter of tradition, but she expressed some unease about this. She hinted at reuniting the genera Aega and Palaega (p. 320); but she did not carry that through, still placing her new species into Palaega. Earlier investigations in this matter were carried out by Malzahn (1968), who compared his Lower Cretaceous Palaega stemmerbergensis with the common Recent species, Aega psora L., 1758, and he found the genera descriptions overlapping. Van Straelen (1930), Polz (2006), and Hansen and Hansen (2010) actually described fossils in Aega, thinning the distinction between the genera. Also, Feldmann and Rust (2006) proposed removing P. acuticauda from the genus, together with 11 other species of Palaega that did not fit the generic diagnosis of Woodward (1870). Etter (2014) gave incertae sedis status to several species of Palaega (among them P. acuticauda) within Cymothoida. Redefining Palaega has resulted in a more precise delimitation of the genus (Feldmann and Goolaerts, 2005; Hyžný et al., 2013; Etter, 2014). Following the key in Hyžný et al. (2013), the specimens of P. acuticauda studied here would better fit within Cirolana. We adhere to this ranking for the moment until more material discloses the anatomy of the anterior pereiopods in detail, as well as structures of the head, both frontal and lateral views. The eyes were interpreted by Secretan (1975: 318) as large and positioned dorsally on the second segment (pereionite 1). We did notice a bulge on the spot where the suggested large dextral eye is supposed to have been, but we think smaller eyes, both visible as depressions positioned frontally near the anterior margin of the cephalon, are more likely. This conclusion actually conforms to what occurs in many modern species of Cirolanidae and Aegidae; but it also is not unlike the situation in Brunnaega tomhurleyi Wilson et al., 2011, a Lower Cretaceous scavenging cirolanid from Australia (Wilson et al., 2011) with a similar axially long first pereionite as in Cirolana acuticauda. In cirolanids, all pereiopods are ambulatory (Roman and Dalens, 1999; Feldmann, 2009). In the case of C. acuticauda, the pereiopods cannot be divided into ambulatory and prehensile types because the pereiopods are not well preserved. We did discern parts of a pereiopod 3 or 4 with either a basis and a propodus and dactylus, or a carpus and propodus (Fig. 1B) laying on its side and dorso-ventrally flattened. Pereiopod 6 and 7 are also present in the holotype on one side, containing all limb segments, somewhat dislocated and looking rather thin (Fig. 2A, B). The broad merus, carpus or propodus parts, in the equally uncertain pereiopod 3 or 4 does not hold enough information to ascribe a clear clinging or walking function to the limb. As for the pleonites, Secretan mentions 3, but we can distinguish 5 that are also visible in the holotype (Fig. 1A). Cirolana holds seven fossil representatives, including C. acuticauda. The latter species differs from its congeners in having a slightly acute pleotelson apex. The pleotelson of C. enigma Wieder and Feldmann, 1992 has a rounded posterior margin; C. fabianii de Angeli and Rossi, 2006 has spines on each side; C. makihiki Feldmann et al., 2008 has a small pleotelson; and the pleotelson of C. garassinoi Feldmann, 2009 has indents on its margin. The pleotelson of C. feldmanni Hyžný et al., 2013 does have a slightly acute posterior margin but lacks a pointed tip. Furthermore, C. feldmanni has uropodal exopods that are clearly more narrow than those of C. acuticauda. Lastly, the pleotelson of C. cottreaui (Feldmann and Charbonnier, 2011) is broad and of quadrate form. Infraorder Sphaeromatidea Wägele, 1989 Sphaeromatoidea Latreille, 1825 Sphaeromatidae Latreille, 1825 Dynamenella Hansen, 1905 Dynamenella veronensis n. comb. (Figs. 2-3) 1868 Sphaeroma A. Milne-Edwards, p Heterosphaeroma Munier-Chalmas, p Heterosphaeroma van Straelen, p. 59, 40, 58, 68 (Fig. 9) Heterosphaeroma veronensis Secretan, p Material Examined. Additional material, in addition to the 4 specimens of Secretan (1975), were depositied in the Verona museum in 1988 from Monte Postale excavations (IG 91180, IG 91162, IG 91161, IG , IG ) and from Pesciara di Bolca excavations (IG , IG ). Diagnosis. Cephalon large and conical, eyes large and positioned dorsolaterally with articulated facets. First pereionite variable in axial length. Uropodal endopod large and broad but not extending the apex of pleotelson. Pleotelson with heightened anterior part, divided in three strong bulges, the middle being the highest; posterior part flat, without indented apex. Body largely covered with granules, pleon and pleotelson with larger pustules. Revised Description. Small sphaeromatid, length cm; pleon with 3 pleonites, one completely developed; pleotelson with three bulges, the middle one higher, and a

5 544 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 35, NO. 4, 2015 Fig. 3. Dynamenella veronensis (Secretan, 1975) nov. comb. A, IG 91180, pleotelson with 3 bulges, lateral view. B, IG , Pn, pereionite; Pln, pleonites; Pt, pleotelson. C, IG , head region, the cephalon posteriorly crushed and parts of exocuticle drifted loose but demarcation with first peronite is clear, Pn, pereionite. Depression on both sides of the pereion axial ridge might be an artefact caused by crushing. D, reconstruction. Scale bars = 2 mm. flat posterior part without rear margin indent; uropods not protruding past the end of the pleotelson, exopods folded beneath endopods. Entire body covered with tubercles, the strongest ones on the pleotelson. Cephalon large and conical with large eyes positioned on the posterolateral margins (Figs. 2C, D; 3C). No traces of antennae or frontal lamina. Pereionites 1-7 (Figs. 2C; 3C) all similar in outline, increasing in axial length towards pleotelson. Coxal plates and pereiopods not visible. Pereionites of seemingly reduced width because a larger part of the lateral sides of the body is still obscured by matrix. Three pleonites (Fig. 2C, D), with only the third one completely distinct; pleotelson triangular, with three bulges (Fig. 3A), the middle one rising higher than the flanking ones, followed by a steep depression and a flat triangular posterior part without indentation, posterior margin rounded. Uropods large (Fig. 2C, D), exopod narrower than endopod, tapering distally without crenulations on margins; endopods not extending to the posterior margin of the pleotelson. Remarks. Heterosphaeroma as a genus was first mentioned in a letter to the French Geological Society on 5 February 1872 by Charles Munier-Chalmas (Munier-Chalmas, 1872), which suggested a generic diagnosis had been given at some other place in the literature. However, this description was never found or referred to. Munier-Chalmas proposed the name Heterosphaeroma as an alternative assignment for Sphaeroma priscum (A. Milne-Edwards, 1868), but he did not provide a diagnosis for the new genus (van Straelen, 1928: 59; Secretan, 1975). The single specimen of H. priscum from Sézanne in France, which consists of a part and counterpart, differs significantly from the specimens from Pesciara di Bolca in having a broad body shape, supposedly uniramous uropods, eyes placed more dorsal than lateral, and a triangular elevation joining the pleon and pleotelson. This fossil was dated Thanetian (Upper Paleocene) and therefore is older than a Late Ypresian age (Lower Eocene) suggested for the Bolca isopod beds. We place the specimens of our fossil species D. veronensis into the living genus Dynamenella, although they are widely

6 VONK ET AL.: EOCENE ISOPODS FROM ITALY 545 separated in time. In doing this, we admittedly are using Dynamenella as a form genus, as was done for Cirolana in the case of fossil cirolanoideans (Hyžný et al., 2013; Jones et al., 2014). We cannot add for the present any more evidence as to whether these forms are related or not. The recent discovery of a sphaeromatid isopod described as Dynamenella miettoi de Angeli and Lovato, 2011, from the Middle Eocene near Vicenza, shows a striking resemblance with the Bolca specimens; it is of marine origin, considering the associated organisms such as mollusks, echinoids, corals, and shellfish (de Angeli and Lovato, 2011). Dynamenella veronensis differs from D. miettoi in not having pronounced, blunt spines on the dorsal margin of the pereionite 7 and pleon. It also lacks the forked indentation on the apex of the pleotelson. Dynamenella is for the most part a modern genus with some 52 species (which have been for the greater part now split up into three genera), several of which live in brackish estuarine conditions (Holdich and Harrison, 1983; Schotte, 2013). They typically have a granulated cuticle and a bulging anterior part of the pleotelson, trilobed, with often ridges or mounts in various forms followed by a flat posterior part ending in a more or less indented apex. DISCUSSION Palaeoenvironment The suborder Scutocoxifera is present in the fossil record from the Triassic to Recent, with fossils allocated to the families Cirolanidae, Serolidae Dana, 1852, Sphaeromatidae, Aegidae White, 1850, Urdidae Kunth, 1887, Schweglerellidae Brandt et al., 1999, Archaeoniscidae Haack, 1918, Chaetillidae Dana, 1849, Idoteidae Samouelle, 1819, Trichoniscidae Sars, 1899, Armadillidiidae Brandt, 1833, Porcellionidae Brandt and Ratenberg, 1831, Oniscidae Latreille, 1802, and some fossils unassigned to a family. Often isopod fossil remains consist of moult stages; entire animals, such as the corpses in this study, are rare. Secretan (1975) commented on the possibility of parasitism as the main mode of living for the isopods found in the Bolca layers because they are so scarce. Parasites are less probable to be buried in sediment as they cling to their hosts and do not frequently walk on stretches of bottom. For the sphaeromatids she concludes that all forms found until the Miocene come from lacustrine environments and that the Pesciara di Bolca isopods might have lived in waters of low salinity. Recent palaeoenvironmental reconstructions confirm Secretan s hypothesis and suggests that the Pesciara di Bolca in the late Ypresian age (between 49 and 50 Ma) was a basin or a subtropical lagoon, close to land, and in some proximity to rivers and wetlands (Petit et al., 2014). Pieces of amber in the lowest fish bearing level contained bryophytes and pteridophytes as well as conifer pollen from Araucariaceae, and specimens of angiosperms (Trevisani et al., 2005). The fish bearing levels, named L1 to L5 (Papazzoni et al., 2006) contain in addition to marine reef fishes also brackish genera such as the perch Cyclopoma (Trevisani et al., 2005) and the atherinid genus Atherina, of which the Recent species inhabit seas and estuaries (Woodward, 1901), confirming the estuarine features of the environment. Arguing against the lacustrine hypothesis are the discoveries, after 1975, of fossil sphaeromatideans and other scutocoxiferans in Italy that are all reported from marine deposits (de Angeli and Rossi, 2006; de Angeli and Lovato, 2009; Passini and Garassino, 2012a, b). In the case of Dynamenella miettoi de Angeli and Lovato, 2011, also of marine origin, fossils of terrestrial plants were found amongst marine invertebrates suggesting transport in times of strong geological activity (de Angeli and Lovato, 2011). These authors report the isopod preserved in a volcanic matrix, part of a volcanic breccia, demonstrating eruptive activity in the late Middle Eocene (Beccaro and de Angeli, 2001). Nonvolcanic events of comparable impact could have played a role in the preservation of the isopods in this study. Rapid burial after major floods by either marine incursions, or river outflows with ensuing mixture of marine and terrestrial organisms is not uncommon in coastal deposits (Alonso et al., 2000; Peñalver et al., 2007; Vonk and Schram, 2007; Najarro et al., 2009; Forey et al., 2010; Dick et al., 2014; Sánchez- Gárcia et al., 2015). It remains uncertain whether the Bolca isopods lived in true marine and shallow reef like conditions, or inhabited a supra-tidal and fluvial-terrestrial habitat. High frequency sea level fluctuations can result in alternating marine and non-marine sediments in a given section. What is needed in future excavations is microstratigraphic, bed-by-bed sampling that records for every layer the entire fossil assemblage. Until such a database exists, conclusions about the microenvironment are rather speculative. ACKNOWLEDGEMENTS We gratefully acknowledge the help of Anna Vaccari and Roberta Salmaso (Museo Civico di Storia Naturale of Verona) in enabling the work in the collections. We thank Viviana Frisone, Claudio Beschin, and Antonio de Angeli (Museum G. Zannato, Montecchio Maggiore) for showing us holotype material. Alessandro Garassino (Natural History Museum of Milan) and Cesare Papazzoni (University of Modena and Reggio Emilia) are thanked for sending literature. 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Roghi Early Eocene amber from the Pesciara di Bolca (Lessini Mountains, Northern Italy). Palaeogeography, Palaeoclimatology, Palaeoecology 223: van Straelen, V Contribution a l étude des Isopodes Méso- et Cénozoïques. Mémoires de l Académie royale des sciences, des lettres et des beaux-arts de Belgique 4, Série 2, 9(5), 68 pp Présentation d un Isopode de l Yprésien du Jutland. Bulletin de la Sociéte Belge de Géologie, de Paléontologie et d Hydrologie 39(1929): 86. von Malzahn, E Über einen neuen Isopoden aus dem Hauterive Nordwestdeutschlands. Jahreshefte des Geologischen Landesamtes in Baden-Württemberg Bd. 10: Vonk, R., and F. R. Schram Three new tanaid species (Crustacea, Peracarida, Tanaidacea) from the Lower Cretaceous Álava amber in northern Spain. Journal of Paleontology 81: Wieder, R. W., and R. M. Feldmann Mesozoic and Cenozoic fossil isopods of North America. Journal of Paleontology 66: Wilson, D. F., J. R. Paterson, and B. P. Kear Fossil isopods associated with a fish skeleton from the Lower Cretaceous of Queensland, Australia direct evidence of a scavenging lifestyle in Mesozoic Cymothoida. Palaeontology 54: Woodward, A. S Catalogue of the Fossil Fishes in the British Museum (Natural History). Vol. I, Part IV. British Museum (Natural History), London, 638 pp. RECEIVED: 24 November ACCEPTED: 11 March AVAILABLE ONLINE: 30 March 2015.