Geo.Cro.2-3-61-KB.pdf 177 � Carles Martín-Closas1, Jean-Paul Colin2, Jean-Paul Billon-Bruyat3, Abder El-Albani4, Jean-Michel Mazin5 and Johann Schnyder6 AB STRA CT A charophyte assemblage including Latochara tenuicostata (PECK 1937) and Mesochara gr. voluta (PECK 1937) was found in the Lower Tithonian bone-bed 148 of the stratigraphic section at the Pointe de Chassiron, Oléron Is- land (Charente-Maritime, Southwestern France). This bed, which is well known for its rich and diverse fauna of tetrapods, was deposited in a margino-littoral environment of Purbeckian facies in the Late Jurassic Charente pal- aeogulf. The occurrence of Latochara at Oléron Island is the most southern location known in the distribution of this genus during the Late Jurassic. This suggests that the boundary between the charophyte Boreal and Tethyan prov- inces, defi ned on the basis of this distribution, is not solely determined by latitudinal (i.e. thermal) factors. Ecologi- cal factors such as salinity and competition with other charophyte families are hypothesized as alternative explana- tions. Keywor ds: Charophytes, ostracods, vertebrates, Tithonian, France, bioprovinces 1 Departament d’Estratigrafi a, Paleontologia i Geociències Marines, Facultat de Geologia, Universitat de Barcelona, Carrer de Martí i Franqu’s, 08028 Barcelona, Catalonia, Spain; (cmartinclosas@ub.edu) 2 3 Impasse des Biroulayres, 33610 Cestas, France 3 Section d’archéologie et paléontologie, Offi ce de la Culture, République et Canton du Jura, Hôtel des Halles, 2900 Porrentruy, Switzerland 4 UMR 6531 Hydrasa, UFR SFA Bâtiment Sciences Naturelles, Université de Poitiers, 40 avenue du Recteur Pineau, 86022 Poitiers Cedex, France 5 UMR 5125 PEPS CNRS, France; Université Lyon 1, Campus de la Doua, Bt. Géode, 69622 Villeurbanne Cedex, France 6 UPMC-Université Paris 06, CNRS, UMR Tectonique, case 117, 4, pl. Jussieu, 75005 Paris, France Palaeobiogeography of the genus Latochara (fossil Charophyta) in the Upper Jurassic of Southern Europe. New data from Oléron Island (France) Geologia Croatica 61/2–3 177–184 3 Figs. Zagreb 2008 Geologia CroaticaGeologia Croatica � 1. INTRODUCTION Palaeobiogeography of Late Jurassic and Early Cretaceous charophytes is a relatively new and poorly understood subject in charophyte research. The pioneer studies of SIRNA (1968) and PALAMAREV (1971) on Early Cretaceous clavatora- ceans, showed for the fi rst time that large palaeogeographic regions shared a similar charophyte fl ora, which was useful not only for characterising them biogeographically, but also for large-scale biostratigraphic correlation. Late Jurassic cha - rophyte biogeography of the Northern Hemisphere, (Europe and North America), was characterized by SCHUDACK (1996) and SCHUDACK et al. (1998). These studies were signifi cant in showing that fl oras from North America and Northern Eu- rope were much more like each other than like fl oras of the Tethyan domain. Thus, without calling them bioprovinces, SCHUDACK et al. (1998) characterized two different regions on the basis of Late Jurassic charophyte assemblages. A North- ern or Boreal region was determined with characean-poro- characean fl oras in opposition to a Southern or Tethyan region, dominated at that time by clavatoracean fl oras. The genus Geologia Croatica Geologia Croatica 61/2–3 178 m thick, is Late Kimmeridgian (biozone Autissiodorensis) to Early Tithonian (biozone Gigas) in age (HANTZPERGUE, 1989; HANTZPERGUE et al., 2004). A sequence boundary associated with a drop in sea-level is found at the top of the previous limestone-dominated suc- cession and is marked by a conglomerate 50 cm thick, show- ing at the top, an accumulation of large woody remains (Fig. 1). Upon this boundary bed, a Purbeckian succession, 56 m thick, developed with non-marine lutites dominating in the lower and upper members, whilst marine carbonates are char- acteristic of the central member. Bioclasts accumulated in par- ticular horizons. The top of the section includes one gypsum layer. Sedimentological evidence and the occurrence of layers with marine fossils, such as benthic foraminifera, marine os- tracods, echinoderm debris and brachiopod-rich horizons, al- ternating with other layers rich in charophytes and non-marine ostracods, indicate that this succession was formed in a shal- low marine, littoral context infl uenced by freshwater discharg- es (BOUSQUET, 1967; SCHNYDER, 2003). The age of the Fi gu re 1: The studied locality and stratigraphic section at the Pointe de Chassiron (Oléron Island, Southwestern France) with the charophyte bearing bed (Bed 148) indicated. Latochara was considered as a key taxon of the Northern Re - gion. Furthermore, the boundary between both regions was set at the 20°C isotherm of mean winter temperature. Later studies show ed that even in the Northern part of the Iberian pa laeoisland, Tithonian charophyte assemblages lacked this genus and were instead dominated by clavatoracean genera such as Globator and Clavator (HERNÁNDEZ et al., 1999; MARTÍN -CLO- SAS, 2000). This suggested that the 20°C iso therm could be a real Late Jurassic bioprovincial boundary. Now, the occurrence of the marker of the northern province, (the genus Latochara), south of the boundary, raises the question of whether factors other than thermal boundaries should be taken into account if we are to understand charophyte palaeobiogeography. 2. GEOLOGICAL AND PALAEONTOLOGICAL SETTING 2.1. GEOLOGICAL BACKGROUND The beginning of the uppermost Jurassic is generally repre- sented in Western Europe by a major drop in relative sea lev- el. Thus, the epicontinental marine deposits of the lowermost Upper Jurassic are covered by lagoonal to non-marine facies, including evaporites that develop well into the Lower Creta- ceous (JACQUIN et al., 1998; RUSCIADELLI, 1999). These facies are generally referred to as “Purbeckian”, through com- parison with reference-sections in England. In France, the Charente Basin is one such Purbeckian basin that is rich in evaporites. The palaeogeography of this basin shows a narrow gulf opening to the West according to DELFAUD & GOTTIS (1966) and HANTZPERGUE & LAFAURIE (1994). This hy- pothesis has been challenged by a recent palaeogeographic reconstruction by THIERRY (2000), showing a connection southward and eastward between the Aquitanian Basin and the Mediterranean Tethys. The outcrops are located in the northern part of Oléron Island, along the cliffs of the Pointe de Chassiron, near the Chassiron lighthouse (Fig. 1). This sec- tion represents the most complete stratigraphic record of the Purbeckian basin of the Charentes. In particular, it is an excel- lent outcrop of the boundary between the underlying Upper Jurassic marine rocks and the Purbeckian non-marine facies. The stratigraphy and biostratigraphy of the section were studi- ed in detail by LANTZ (1958), DONZE (1960), MALZ (1958, 1966) and BOUSQUET (1967), who provided a de tail ed sed- imentological study and BOURGUEIL et al. (1976). More recently, HANTZPERGUE (1989) and HANTZPERGUE et al. (2004) reported new biostratigraphic data and SCHNY- DER (2003) proposed a new palaeoclimatic interpretation based on palynological and mineralogical evidence. 2.2. DESCRIPTION OF THE STRATIGRAPHIC SECTION AT POINTE DE CHASSIRON The base of the section consists of grey-beige limestones (mud- stone to bioclastic packstone), showing poorly stratifi ed or ir- regular, nodular bedding, alternating with marls (Fig. 1). In sedimentological and palaeontological terms, the facies cor- respond to a shallow, open platform of normal salinity, with ammonites, echinoderms and dinofl agellates (BOUSQUET, 1967; SCHNYDER, 2003). This succession which is up to 24 Geologia CroaticaCarles Martín-Closas et al.: Palaeobiogeography of the genus Latochara (fossil Charophyta) in the Upper Jurassic of Southern Europe... 179 Purbeckian succession is not easy to establish in the absence of ammonites. However, an Early to Late Tithonian age was suggested on the basis of ostracods, brachiopods (HANTZ- PERGUE et al., 2004), calcareous nannofossils and dinofl ag- ellates (SCHNYDER, 2003). Bed 148, which provided the fossils that are the subject of this study, is a marl with small carbonaceous nodules, lo cat- ed in the lower part of the Purbeckian succession, upon a gre- enish marly-silty layer (Fig. 1). It shows an accumulation of woody remains at the base, charophyte gyrogonites and ver- tebrate remains. The member of the section containing this layer was attributed to the Early to Middle Tithonian. This member is formed by dark blue marly claystones interlayered with siltstones and silty, fi nely bedded limestone. These rocks are thought to have been deposited in a littoral or estuarine bay with abundant freshwater discharges. The lack of fauna indicating normal marine salinity such as echinoderms is of particular interest and, in contrast, the presence of rare ostra- cods of the “Cypris” type, indicates brackish to freshwater conditions, together with charophytes (BOUSQUET, 1967; SCHNYDER, 2003). 2.3. THE FAUNA ASSOCIATED WITH CHAROPHYTES Two main groups of fossils were found associated with the charophytes studied: ostracods and vertebrates. Ostracods are represented by about 20 species. The euryaline form Fabanel- la boloniensis ornata (STEGHAUS, 1953) and the limnic ge- nus Mantelliana cf. perlata WIENHOLZ, 1968 are dominant. The remaining fauna consists mainly of non-marine species: Cetacella armata MARTIN, 1958, Darwinula oblonga (RO- EMER, 1839), Alicenula leguminella (FORBES in LYELL 1855) The rio synoecum aff. forbesii (JONES, 1885), Therio- synoecum cf. albertense (PINTO & SANGUINETTI, 1962), Cypridea spp., Eo paracypris sp., Stenestroemia sp., Damonella pygmaea (AN DERSON, 1941), Damonella ellipsoidea (WOL- BURG, 1962), “Bisulcocypris” dilatata ANDERSON, 1971, and Rhi nocypris jurassica (MARTIN, 1940). Marine genera are rare and represented by a few specimens of Dicrorygma, Asciocythere? and Cytheropterina. The vertebrate fauna from the Pointe de Chassiron con- sists of chondrichthyans, osteichthyans and tetrapods. Tetra- pods have been reported in 14 beds from the Purbeckian suc- cession (beds B104 to B203a from BOUSQUET, 1967). They include remains from a lissamphibian, turtles, a lepidosaurian, crocodilians, pterosaurs and dinosaurs (LAPPARENT & OUL- MI, 1964; RIEPPEL, 1980; BILLON-BRUYAT et al., 2001; BIL LON -BRUYAT, 2003). A detailed list and a description of these tetrapods is given in BILLON-BRUYAT (2003). Bed 148 (B148 from BOUSQUET, 1967) was termed a bone-bed and is considered the horizon with the most diverse vertebrate fauna in the section. A preliminary study of chon- drichthyans and osteichthyans from this bed was carried out by VULLO (2001), who reported a number of families living in marine or margino-littoral environments. A recent isotopic geo- chemical study showed that Semionotidae remains from bed B148 are of marine origin (BILLON-BRUYAT et al., 2005). Tetrapod remains from the same bed include a lissamphib- ian (Urodela incertae sedis), turtles (Plesiochelyidae), croco- dilians (Teleosauridae, Atoposauridae, Goniopholididae, Ber - nissartiidae), pterosaurs (Pterodactyloidea indet, Pterosauria indet.) and dinosaurs (Theropoda indet.). Most of these taxa were identifi ed on the basis of microremains, mainly isolated teeth. Only the plesiochelyids, teleosaurids, goniopholidids and a theropod are represented by larger remains. The strati- graphic distribution of these tetrapod families is consistent with a latest Jurassic age. They were known from the Jurassic and Cretaceous with the exception of plesiochelyids, which disappeared in the Tithonian, and teleosaurids, which are only represented in the Cretaceous by a single questionable speci- men (CORNÉE & BUFFETAUT, 1979; BENTON, 1993; BARDET, 1995). A margino-littoral depositional environment is consistent with the composition of the tetrapod fauna. Ple- siochelyids and teleosaurids in particular are considered as co ast- al marine reptiles (BUFFETAUT, 1982; BROIN, 1994; BAR- DET, 1995). In addition, BILLON-BRUYAT et al. (2005) reported that samples from these families in bed B148 indi- cate a marine isotopic signal. Their remains were abundant and include some connected bones. All the other tetrapod fam- ilies were non-marine, either freshwater/terrestrial (lissam- phibians), freshwater/brackish (goniopholidid, bernissartiid crocodilians), terrestrial (atoposaurid crocodilians, theropod dinosaurs) and aerial elements (pterosaurs). The occurrence of these continental families in the margino-littoral deposi- tional environment clearly refl ects a freshwater infl ow. 3. SYSTEMATIC PALAEONTOLOGY Division Charophyta MIGULA, 1897 Class Charophyceae SMITH, 1938 Order Charales LINDLEY, 1836 Family Nitellaceae von LEONHARDI, 1864 emend. MARTÍN-CLOSAS & SCHUDACK, 1991 Genus Latochara MÄDLER 1955 emend. FEIST in FEIST & CUBAYNES, 1984 Latochara latitruncata (PECK, 1937) MÄDLER, 1955 (Figs. 2.1–2.4) 1937 Aclistochara latitruncata PECK, n. sp. – PECK, p. 89, pl. 14, fi gs 1–4 1955 Latochara latitruncata (PECK) nov. comb. – MÄ- DLER, p. 271 1957 Latochara latitruncata (PECK) – PECK, p. 32–33, pl. 5, fi gs. 7, 21–33 1957 Latochara concinna PECK, n. sp. – PECK, p. 34, pl. 5, fi gs. 5–6, 8–9 1990 Latochara latitruncata (PECK 1937) MÄDLER 1955 – SCHUDACK, p. 219, pl. 2, fi gs. 1–4 1993 Latochara latitruncata (PECK 1937) MÄDLER 1955 – SCHUDACK, p. 56, pl. 4, fi gs. 1–4 Material: About 200 gyrogonites from sample 148b. Description: Medium-sized gyrogonites, in shape sub - spherical to ellipsoidal. Length ranges 351–432 mm (average 404 mm), including the neck. Width ranges 270–351 mm (av- erage 312 mm). Isopolarity index 123–136 (average ISI 129). Geologia Croatica Geologia Croatica 61/2–3 180 Spiral cells are fl at or slightly concave with carinated sutures. The number of convolutions visible in frontal view ranges from 10 to 13. Apex generally pointed. Apical neck generally short and pyramidal but in isolated specimens may be almost prismatic, similar to the neck of Stellatochara. When ap- proaching the neck, spiral cells become fi rst wider and then narrow suddenly up to the summit. In this area the spiral cells are strongly concave. A periapical depression is present in some specimens. The apical pore is small and hardly visible. Base rounded or truncated. Basal plate not observed, but prob- ably multipartite, as in other species of the genus. Discussion: The population studied belongs to medium- sized Jurassic and Lower Cretaceous Latochara species. These are L. latitruncata, L. collina PECK, 1957, L. concinna PECK, 1957, L. tenuicostata (PECK, 1937) and L. maedleri SCHU- DACK, 1996. Other species are smaller, i.e. about 300 mm average length (L. mensinki SCHUDACK, 1990 and L. bel- latula PECK, 1957) or larger, i.e. more than 500 mm average length (L. sphaerica PECK, 1957 and L. durand-delgai FEIST 1984) and will be not be considered for comparison. The population studied is similar in size and shape to L. latitruncata (PECK, 1937) from the Tithonian and Kim mer- idgian of the Morrison Formation (USA), with the difference that this population is more ellipsoidal and has more numer- ous and thinner spiral cells. The specimen shown by PECK (1957, pl. 5, fi g. 22) is very close to this population in its numb- er of convolutions and general shape. Apart from the Morrison Formation, this species was reported from the Lower Kimme- ridgian of the Saxony Basin (NW Germany), by SCHUDACK (1993) and from the Tithonian of the Dnieper-Donets basin (Ukraine) by SHAĎKIN (1967). The German material shown by SCHUDACK (1990, 1993) differs from the population stu- died here in the stronger calcifi cation of spiral cells. The numb- er of convolutions is somewhat smaller but overlaps with the range of the French population. Latochara collina PECK 1957 is very close to the previ- ous species and PECK (1957) suggested that they may be dis- tinct only at the sub-species level. The only noticeable differ- ence between the two species is the shape of the intercellular sutures which is wavy in L. collina. This may give a knotty aspect to spiral cells. This characteristic is not present in this material. SCHUDACK (1993) points to the larger width (sub- lat shape) and the truncated summit of gyrogonites to distin- guish this species; however these characters are also present in Latochara latitruncata, as clearly indicated in its species description. Latochara concinna PECK 1957 is considered in the orig- inal description as being very similar to L. latitruncata and SCHUDACK (1993) concludes that both species are synony- mous, a point of view to which we adhere. Latochara tenuicostata PECK 1957 from the Aptian of Wyoming is somewhat smaller than the material here, but is similar in the number of convolutions. However, the apical pore of this species is large and rose-shaped, unlike the apical pores of other Latochara. This pore is reminiscent of the api- cal pores of Clavatoraceae. Also the overall aspect of the gy- rogonite, including its pointed base, is similar to internal casts of the fructifi cation of genus Atopochara. PECK (1957) no- ticed the presence of clavatoracean remains associated with Latochara tenuicostata. In consequence, it appears that this species should not be ascribed to the genus Latochara but rather to an internal cast of a clavatoracean oosporangium. Latochara maedleri SCHUDACK, 1996 from the Berria- si an of NW Germany is distinct from other species in its ex- tremely truncated apex, which results in a very wide gyrogonite. This form does not occur in the material described here. Fi gu re 2: Early Tithonian charophytes from bed 148 at the Pointe de Chassiron. 1, 2: Latochara latitruncata. Lateral view of two gyrogonites showing globulosity and pointed apex. 3: Lateral view of gyrogonite showing an elongated shape and an apical neck. 4: Lateral view of a well- calcifi ed gyrogonite showing a conical apex. 5: Mesochara gr. voluta. Lateral view of gyrogonite showing drop shape and prominent sutures. Geologia CroaticaCarles Martín-Closas et al.: Palaeobiogeography of the genus Latochara (fossil Charophyta) in the Upper Jurassic of Southern Europe... 181 Family Characeae (R. ex C. AGARDH, 1824) emend. MARTÍN-CLOSAS and SCHUDACK, 1991 Genus Mesochara GRAMBAST, 1962 Mesochara gr. voluta (PECK, 1937) emend. PECK, 1957 nov. comb. GRAMBAST, 1965 (Fig. 2.5) 1937 Chara voluta PECK, n. sp. – PECK, p. 85–86, pl. 14, fi gs. 16–19 1957 Praechara voluta (PECK) – PECK, p. 39, pl. 7, fi gs. 7, 22–27 1965 Mesochara voluta (PECK) nov. comb. – GRAM- BAST, p. 581 Material: Six gyrogonites from sample 148b. Description: The gyrogonite is small, 270–350 mm in high and 240–300 mm wide. Isopolarity index ranges between 100 and 120. The gyrogonite is normally drop-shaped with a fl at apex and a pointed base. Some specimens also show a point- ed apex. The maximum width is above the middle point. In pro fi le, 7–8 convolutions are visible, separated by prominent ridges. In the equatorial zone, spiral cells are between 25–60 mm high. Apex closed by simple junction of spiral cells. Base shows a pentagonal pore with a maximum diameter of about 50 mm. Basal plate not observed. Discussion: The group voluta has a wide chronostratigra- phic range, from the Kimmeridgian to the Barremian (SCHU- DACK, 1993; MARTÍN-CLOSAS, 2000). 4. PALAEOBIOGEOGRAPHY Whilst Latochara from the Lower Jurassic is known from south ern France, (FEIST and CUBAYNES, 1984), in the Up per Jurassic and Lower Cretaceous, the genus was exclusively known from the Boreal domain, i.e. regions north of the Teth- yan Sea. PECK (1957) and SCHUDACK et al. (1998) report- ed the genus in the Kimmeridgian and Berriasian of the Mor- rison Formation (Western United States), FEIST et al. (1995) in the Upper Tithonian–Lower Berriasian from the Purbeck facies in England, SCHUDACK (1993) from the Kimmerid- gian, Tithonian and Berriasian in the North-western German basins, and SHAĎKIN (1967), from the Tithonian of the Dnie- per-Donetz basin (Fig. 3). However, the genus is completely un known in the Tithonian–Berriasian basins of Central Teth- ys, i.e. in the Iberian Peninsula, Sardinian High, Provence and Northern Africa, where charophyte associations were gener- ally dominated by clavatoraceans during that period (SCHU- DACK et al., 1998; MARTÍN-CLOSAS, 2003). In consequ- ence, the Latochara genus was considered by SCHUDACK (1996) as one of the most characteristic genera of the North- ern charophyte biogeographic province during the Kimmer- idgian–Tithonian and Berriasian. He hypothesized that a ther- mal barrier at 20°C of mean winter temperature would account for the biogeographic boundary (Fig. 3). The occurrence of Latochara tenuicostata at Oléron Is- land, Southwestern France, during the Late Jurassic, allows us to shift the known biogeographic distribution of this genus 570 km southwards, to touch the northern margins of the Me- sogea, in Central Tethys. The new distribution suggests that the biogeography of the genus was not just controlled by temperature as proposed by SCHUDACK (1996). Other factors, in addition to temper- ature, may have hindered the expansion of the genus into the central Tethyan islands (Iberian Peninsula, Sardinian high and Morocco Meseta). The next Upper Jurassic-Lower Cretaceous charophyte locality south of Oléron Island occurs in the Lower Berriasian lacustrine limestone of the Aguilar de Campoo Fm, Cantabrian Chain, where charophyte assemblages are domi- nated by Globator maillardii praecursor and Clavator reidii (HERNÁNDEZ et al., 1999). This formation is attributed to deposition in large, permanent, alkaline and freshwater lakes without any marine infl uence. Also the other localities from Iberia (SCHUDACK, 1993), the Sardinian High (COLIN et al., 1985) and the Morocco Meseta (MOJON, 2002) corre- spond to lacustrine limestone belonging to freshwater alkaline lakes, whilst localities dominated by Latochara, such as the new occurrence described here, correspond to brackish or lit- toral marshes. In conclusion, palaeoecology also appears to play a signifi cant role in the distribution of Latochara in ad- dition to climate. It is known that Late Jurassic and Early Cre- taceous charophyte assemblages dominated by porocharac- eans, (the family to which Latochara belongs), are cha racteristic of brackish environments (MARTÍN-CLOSAS & GRAM- BAST-FESSARD, 1986; MOJON, 1989). Palaeoenvironmen- tal factors, such as water alkalinity or salinity, may be an im- portant factor in the palaeogeographic distribution of Meso zoic charophytes. Also, the conjoint or disjoint disposition of ga- metangia has to be taken into account when drawing the bio- geography of fossil species. PROCTOR (1980) showed that monoecious species of extant Chara were cosmopolitan whilst dioecious ones were restricted to single land masses. This was found to be signifi cant in at least one Early Cretaceous spe- cies, Atopochara trivolvis according to MARTÍN-CLOSAS & WANG (2008). Fi gu re 3: Global distribution of Tithonian Latochara with its southern- most location on Oléron Island, beyond the winter isotherm of 20°C (modifi ed from SCHUDACK et al., 1998). Legend: 1–5 localities in the Mor- rison Fm (United States), 1. Montana, 2. South Dakota, 3. Central and Eastern Wyoming, 4. East-Central Utah, 5. Colorado, 6. Purbeck basin, S. England, 7. Saxony basin, N. Germany, 8. Oléron Island, Charente basin, France. Geologia Croatica Geologia Croatica 61/2–3 182 5. CONCLUSION Bone bed 148 of the Pointe de Chassiron section on Oléron Island, (Charente-Maritime, France) contains a Tithonian as- semblage of Latochara tenuicostata and Mesochara gr. volu- ta that represents the southernmost occurrence of the genus Latochara in the Latest Jurassic-Earliest Cretaceous of Eu- rope. This genus, which was considered by SCHUDACK et al. (1998) to be characteristic of a Boreal charophyte region, occurs beyond the southern boundary (as set by these authors at the 20°C isotherm of winter temperature) of this Boreal Re- gion. Since all the Latest Jurassic to Earliest Cretaceous lo- calities without Latochara, occur south of Oléron Island and were deposited in freshwater alkaline lakes we suggest that palaeonvironmental factors as well as thermal boundaries were involved in the biogeographic distribution of the genus. ACKNOWLEDGEMENT This study is a contribution to the project CGL2005-00404 of the Spanish Ministry of Science and Technology. 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