2019 | 72/3 | 163–172 | 2 Figs. | 3 Pls. | www.geologia-croatica.hr Journal of the Croatian Geological Survey and the Croatian Geological Society 1. INTRODUCTION Cordaitalean remains from the Bohemian part of the Intrasudetic Basin have been known to palaeobotanists since the 19th century [e.g. Cordaites platynervis (GÖPPERT, 1864) GRAND’EURY, 1877]. Since then, tens of Cordaites species have been described based on leaf morphology and venation. However, it has been shown that venation can be strongly influenced by both tapho­ nomic and preservation conditions (CROOKALL, 1970). More­ over, many species have been described based on incomplete spec­ imens, without their base, or apex, and such specimens are difficult to compare with other specimens only by the means of venation. In reality, the situation is more complicated, because both leaf morphology and venation have some natural variability caused by the edaphic and climatic conditions in which the plant grew. The most stable feature seems to be the cuticle. Cordaitalean cuticles have been studied by FLORIN (1931), and more systematically by BARTHEL (1962, 1964, 1976), LEDRAN (1960, 1966), and RABITZ (1966). BARTHEL (1962, 1964, 1976) found that one species [e.g. Cordaites principalis (GERMAR) GEINITZ] can have different cuticles, that he named Type 1 to Type 8. BAR­ THEL (1976) was aware of the problem of the correct determina­ tion of Cordaites species and came to the conclusion that it is not possible to distinguish three long­leaved Cordaites species – C. borassifolius (STERNBERG) UNGER, C. palmaeformis (GÖP­ PERT) WEISS and C. principalis (GERMAR) GEINITZ, and he suggested calling them “Cordaites sp.” This interpretation helps to avoid problems in the determination of Cordaites species, but it does not enable recognition of the diversity of this group and/or the specification of the number of Cordaites species per locality. BARTHEL (2009, 2016) did not change his mind and classified all samples as Cordaites sp. From the quantity of Cordaites names, only Cordaites boras- sifolius, C. palmaeformis and C. principalis dominate in the Revision of the genus Cordaites UNGER from the Permian of the Intrasudetic Basin (Broumov Formation, Olivětín Member, Czech Republic) Zbyněk Šimůnek Czech Geological Survey, Klárov 3/131, 118 21 Praha 1, Czech Republic; (zbynek.simunek@geology.cz) doi: 10.4154/gc.2019.16 Abstract Cordaitalean remains are not very common in the Permian of the Intrasudetic Basin (Czech Re- public and Poland) and have usually been classified simply as Cordaites sp. Based on leaf mor- phology, venation and cuticles (where possible), five cordaitalean species are recognised: Cor- daites cf. beinertianus, C. cf. affinis, C. cf. roesslerianus, C. cf. foliolatus and Cordaites sp. A. The determination of Cordaites leaves is difficult because of the variability and similarities be- tween several species. The biggest leaf was determined as Cordaites cf. beinertianus. The cu- ticles of Cordaites cf. affinis resemble the cuticles of C. palmaeformis and C. radvanicensis, however, the latter is known only in a fragmentary state. Cordaites beinertianus and C. palmae- formis are very similar and are considered as conspecific by some authors. Cordaites affinis has generally smaller leaves. Cordaites cf. roesslerianus is comparable with C. principalis with which it is sometimes synonymised. C. cf. foliolatus is represented by a much narrower leaf than is typical for cordaitalean leaves from France. Cordaites sp. A is a relatively small leaf-form and differs from all other forms previously described. pa laeobotanical literature. As shown above, these species are under­ stood by different authors in different ways, and only cuticles from the type material will help to solve the problem of the accurate determination of these species. Cuticles of Cordaites borassifolius were described as amphistomatic with dispersed stomata on the adaxial cuticle, and stomatiferous and non­stomatiferous bands on the abaxial cuticle (ŠIMŮNEK et al., 2009). Studying cuticles of Cordaites palmaeformis was difficult due to poor preservation, but eventually the stomata were shown to be arranged in stomatal rows on both cuticles (ŠIMŮNEK, 2015). A problem remains with Cordaites principalis, because its holotype is preserved as an impression in fine-grained sandstone that does not allow cu­ ticular study (ŠIMŮNEK, 2015). Some cordaitalean remains were discovered during the study of flora from the Olivětín Member in the Intrasudetic Basin. Be­ cause some of these remains were large and some contain coal matter suitable for cuticular study, description of these specimens by the author was possible. 2. MATERIAL AND METHODS The specimens are stored in the collections of the National Mu­ seum in Prague and the Museum of Eastern Bohemia in Hradec Králové (No. 74574 and 74575). All specimens came from the Otovice locality (National Museum), and Otovice, Černý potok (Black Creek) limestone (Museum of Eastern Bohemia). All were from the Permian (latest Asselian) Broumov Formation, Olivětín Member (OPLUŠTIL et al. 2016, Table 1). Three Cordaites speci­ mens were macerated and slides No. 698/1­16, 699/1 and 700/1­4 prepared. The method used involved maceration in Schulze’s reagent, as described previously by KERP (1990), KRINGS & KERP (1997) and KERP & KRINGS (1999). Coalified fragments of Article history: Manuscript received June 06, 2019 Revised manuscript accepted September 10, 2019 Available online October 31, 2019 Keywords: Cordaites, Permian flora, Intrasudetic Basin G eo lo gi a C ro at ic a Geologia Croatica 72/3164 leaves were separated from the rock by needle or by hydrofluoric acid (HF) digestion. These fragments were bleached in Schulze’s reagent (40% HNO3 with crystals of KClO3) for 3–7 hours. After the treatment in Schulze’s reagent, cuticles were washed in water, subsequently immersed in 2% or 10% potassium hydroxide (KOH) and finally rinsed in distilled water. Some cuticles were stained with safranin, Bismarck brown, malachite green or neu­ tral red (KRINGS, 2000) for 1 to 2 hours to accentuate the anti­ clinal walls and stomata. Before embedding in glycerine­gelatine slides, the cuticles were dehydrated in pure glycerine. 3. GEOLOGICAL SETTING The Intrasudetic Basin, which is situated along the eastern mar­ gin of a suite of Late Palaeozoic Bohemian continental basins (Fig. 1), is the most important among the Sudetic basins. Struc­ turally, the Intra­Sudetic Basin is a complicated NW–SE­elon­ gated syncline with up to 5 km of infill that occupies an approxi- mately 1800 km2 area, of which about one third is located in the territory of the Czech Republic (formerly Czechoslovakia). De­ po sition spans a wide stratigraphic interval from the middle Visean to the Triassic and includes several hiatuses. Continental depo sits unconformably overlie Saxo­Thuringian crystalline complexes of Pre­Cambrian to Early Palaeozoic age (TÁSLER et al., 1979, BOSSOWSKI, 1995). Visean sediments in Poland consist of a few kilometres of fluvial to alluvial-fan strata with interca­ lated marine to deltaic deposits. They are followed in Poland by Serpukhovian (early Namurian) fluvial and coal-rich deposits of the Wałbrzych Formation. In Bashkirian times, the depositional center enlarged further to the south along a tectonically active western basin margin. This resulted in deposition of the Late Bashkirian–Early Moscovian Žacleř Formation (Yeadonian–Bol­ sovian), with about 60–80 coal seams in three members. Locally important coal seams of Kasimovian – Gzhelian (Cantabrian to Saberian) age occur only in the Czech part of the basin in the Odolov Formation (Svatoňovice and Jivka members) (Table 1). The Stephanian/Permian Chvaleč Formation is mostly com­ posed of coal-barren fluvial red beds and contains only one thin coal seam in the Vernéřovice Member. Lacustrine horizons a few tens of metres thick, with bituminous limestones and shales are rarely intercalated in red beds (Bečkov Horizon). The overlying Broumov Formation contains the Nowa Ruda, Olivětín and Mar­ tínkovice members that correspond mostly to the Asselian (OPLUŠTIL et al. 2016). The Nowa Ruda Member is formed mostly by thick complexes of rhyolite tuffs and tuffites, The Olivětín Member contains fossiliferous “Walchia” shales several tens of metres thick and thin bituminous shales and Ruprechtice and Otovice limestones. The Martínkovice Member also contains limestone horizons, but only animal fossils have been discovered so far. The Upper Permian Trutnov and Bohuslavice formations are composed of red and brown mudstone and sandstone depo­ sits; and the Middle Triassic Bohdašín Formation is represented mostly by white sandstones. 4. SYSTEMATICS Order Cordaitanthales MEYEN, 1984 Family Cordaitanthaceae MEYEN, 1984 Genus Cordaites UNGER, 1850 Type species Cordaites borassifolius (STERNBERG) UNGER, 1850 As mentioned in the Introduction, there are difficulties with the systematics of Cordaites leaves. More than 40 cordaitalean spe­ cies have been described based on leaf adpressions from Europe, Table 1. Stratigraphic table of the youngest part of the Intrasudetic Basin. Adopted according to OPLUŠTIL et al. (2016). Remark: Time intervals of gaps between younger units are not known. Figure 1. Map of the Czech Republic with Permo-Carboniferous basins and the studied locality. G eologia C roatica Šimůnek Zbyněk: Revision of the genus Cordaites UNGER from the Permian of the Intrasudetic Basin (Broumov Formation, Olivětín Member, Czech 165 defined on the shape, dimensions and venation type. Even with complete, or nearly complete leaves, there can still be difficulties in classifying specimens to particular species, because the vena­ tion is often poorly preserved due to taphonomic processes (CROOKALL, 1970). About 15 species of Cordaites UNGER and Poa-Cordaites GRAND’EURY ex BRONGNIART have been described from the Stephanian and Permian, or reported from these strata. These species were compared with specimens found in the Otovice lo­ calities but the latter are problematic because their cuticles are poorly preserved and so cannot be used in species determination. In both museums, seven fragments, probably belonging to four or five species, were selected for study. CROOKALL (1970) only described five species from Great Britain. The last compre­ hensive paper on cordaitaleans was published by LEDRAN (1966) who described 22 species from France. She also tried to describe the cuticles but they were usually poorly preserved and only in a few cases did they add significant information to sup­ plement the morphological description. BARTHEL (1962, 1964) distinguished in Cordaites principalis several types of cuticles and considered Cordaites borassifolius, C. palmaeformis and C. principalis as indistinguishable from each other (BARTHEL, 1976, 2009, 2016). Specimens from the Otovice locality demonstrate the vari­ ability in leaf shape, dimension and venation. Here, comparison of these specimens will be made with other specimens described from the uppermost Stephanian and Permian. Cordaites cf. beinertianus (GÖPPERT, 1841) GRAND’EURY, 1877 (Fig. 2a, h, i; Pl. 1, Figs. a, b) 1841 Noeggerathia beinertiana GÖPPERT, Vol. 5­6, p.108, pl. 12, fig. 3. 1877 Cordaites beinertianus GÖPPERT; GRAND’EURY, p. 215. 1914 Dorycordaites beinertianus (GÖPPERT) GRAND’EURY; Bureau, p. 304, Atlas, 1913, pl. 66, fig. 1. Description: The leaf is lanceolate with a bluntly pointed apex. The fragment is 565 mm long and 97 mm wide. The venation is very fine and dense, 45–60 veins per cm in different parts of the leaf (Fig. 2a, h, i; Pl. 1, Figs. a, b). Remarks: The complete leaf may have been more than 600 mm long, and because its right margin is enrolled (Fig. 2a), it could be 110–120 mm wide. The holotype of Cordaites beiner- tianus (GÖPPERT, 1842) represents the basal half of a leaf. It is wedge­shaped, widening from 3 mm (base) to 55 mm (in the mid­ dle) along its length of 170 mm. The veins are very thin, and it seems from the picture that there are about 30 veins per cm. As the apex is not preserved, determination by means of venation only is uncertain. The type locality of Cordaites beinertianus is the Charlottenbrunn (nowadays Jedlina Zdrój), of Poland and is probably Bolsovian in age. This combination was used for the first time by GRAND’EURY (1877) in comparison with Cord- aites affinis, but C. beinertianus was not formally transferred to the genus Cordaites. GRAND’EURY (1877) used this name without description, only with reference to (GÖPPERT, 1842). C. beinertianus was also reported from the Upper Stephanian (BUREAU, 1914), though it is not known that these young speci­ mens are of the same species as the holotype. This specimen is also similar to Cordaites palmaeformis, however the lectotype represents several leaf fragments 160–180 mm long and 12–20 mm wide (ŠIMŮNEK, 2015). Other speci­ mens figured by GÖPPERT (1852) also do not have a preserved apex, which is an important diagnostic feature, but the leaves are wider, up to 50 mm wide. GÖPPERT (1852) did not specify the venation density; he only noted that the venation was fine and dense. Only subsequent authors more precisely defined the vena­ tion density: GEINITZ (1855) – 3–4; WEISS (1871) – 3–5 and ZEILLER (1886­1888) – 8–12 veins per mm. CROOKALL (1970) mentioned 35–50 veins per mm. This confusion may be because each specimen came from a different country and stra­ tigraphy. The lectotype of Cordaites palmaeformis came from the Langsettian of Wałbrzych in the Intrasudetic Basin (ŠIMŮNEK, 2015), however the other syntype (GÖPPERT, 1852, pl. 16, fig. 2) is from the Upper Silesian Basin. The leaves of Cordaites palmaeformis are amphistomatic (ŠIMŮNEK, 2015), with stomata on both cuticles arranged in rows. One cuti­ cle (probably adaxial) has 5–6 rows per mm, whilst the second cuticle (probably abaxial) has 12–16 rows per mm. These densi­ ties correspond to the densities of fine veins in the adpressions, and also correspond to the specimen from Otovice. However, there is a large stratigraphic distance between these specimens, and cuticles from this specimen from Otovice are not known. GÖPPERT (1852) compared Cordaites palmaeformis with C. beinertianus. Figure 2. Shapes and venations of Cordaites leaves: a. – Cordaites cf. beinertian- us, b. and c. –Cordaites cf. affinis, d. and e. – Cordaites sp. A; f. – Cordaites cf. roess- lerianus, g. – Cordaites cf. foliolatus; Scale bars: a = 5 cm; b–g = 2 cm. Venations of leaves: h, i – Cordaites cf. beinertianus (fig. a), j – Cordaites cf. affinis (fig. b), k – Cordaites sp. a (fig. d), l – Cordaites cf. roesslerianus (fig. f ). (Left scale bar is only for a; right upper scale bar is for b–g and left lower scale is for h–l). G eo lo gi a C ro at ic a Geologia Croatica 72/3166 Cordaites cf. affinis GRAND’EURY, 1877 (Fig. 2b, c, j; Pl. 1, Fig. c–e; Pl. 2) 1877 Cordaites affinis GRAND`EURY; GRAND`EURY: p. 215–216, pl. XVIII, figs 6. 7. 1896 Cordaites (Dorycordaites) affinis GRAND`EURY; RENAULT: p. 344, pl. LXXXVI, fig. 17. Material: Two specimens. National Museum and Museum of East Bohemia (No. 74574). Cuticle slides 698/1–16 have been prepared from specimen 74574. Description: Two incomplete leaves. The larger (Pl. 1, Fig. c) is 215 mm long and 36.5 mm wide. Maximum leaf width is below its middle. The second specimen (Pl. 1, fig. e) is a leaf fragment 165.5 mm long and 20.5 mm wide in its upper part. This leaf was probably thin and lacerated in its upper part. The venation is the same in both specimens, formed by very thin veins with density of 55 veins per cm. Cuticles are very poorly preserved with details only visible by staining with safranin, malachite green or neutral red. It is dif­ ficult to distinguish adaxial and abaxial cuticles as both probably have stomatal rows and non­stomatiferous bands. Cuticles on Pl. 2, Figs a, b come from the same leaf fragment and were stained in malachite green. Pl. 2, Fig. a probably represents an adaxial cuticle. Dark bands 120–170 mm wide are distinguishable from light bands around 60 mm wide. Pl. 2 Fig. b probably belongs to an abaxial cuticle. The lighter bands on the adaxial cuticle prob­ ably correspond to stomatal rows on the abaxial cuticle and darker bands to non-stomatiferous bands (costal fields). Abaxial cuticle (Pl. 2, Figs b–h). The cells are differentiated into 90–160 mm wide non­stomatiferous bands and 50–60 mm wide stomatal rows. In contrast to the adaxial cuticles, the stomatal rows are darker than the non­stomatiferous bands. The cells of the non­stomatiferous bands are oblong with straight or slightly bent walls, 15–22 mm wide and 45–80 mm long. The structure of the stomatal rows is difficult to observe. Usually, there are 2 guard cells, 2 polar and 2 lateral subsidiary cells per stomatal complex. The stomatal complexes were probably collapsed and it is impos­ sible to distinguish cell walls from folds on the periclinal walls. In a few cases, guard cells have been preserved. They have a crescent shape, are 28–40 mm long and 12–14 mm wide (Pl. 2, Figs. e, f). Remarks: GRAND`EURY (1877) described similar speci­ mens with fine venation as Cordaites affinis. They are narrow, lanceolate leaves with sharp apices, about 200–300 mm long and 18–30 mm wide. The widest part of the leaf is also approximately in the middle of the leaf. The dimensions of the leaves described here fit to this description, only they (Pl. 1, fig. 3) are a little wider – 36.5 mm. The vein density is comparable with data from LE­ DRAN (1966): 50–60 per mm. Leaves of Cordaites affinis start gradually to widen from their base, the leaf on Pl. 1, fig. c widens more and thus resembles more closely Cordaites palmaeformis that is compared closely with previous species. Cuticles prepared from the specimen on Pl. 1, fig. c resemble the cuticles of Cordaites palmaeformis (ŠIMŮNEK, 2015) with stomatal rows on both the abaxial and adaxial cuticles. Cuticles from the type material of Cordaites palmaeformis were also poorly preserved, however, the structure of the stomatal complexes of adaxial and abaxial cuticles has eventually been resolved. Corda- ites cf. affinis from Otovice and Cordaites palmaeformis from Walbrzych are both amphistomatic leaves with stomata arranged in stomatal rows on both cuticles. However, it is still not possible to demonstrate that the specimen on Pl. 1, fig. c is Cordaites affinis, or if it belongs to Cordaites palmaeformis, because cells of the sto­ matal rows are poorly preserved and a comparison is not possible, therefore these specimens are named as Cordaites cf. affinis. LEDRAN (1960, 1966) studied cuticles from cordaitaleans from France which were also poorly preserved. Her drawings of the cuticles show that the stomatal complexes are drawn as cy­ clocytic complexes with 4 or 5 equal subsidiary cells around the stomatal pore. Such a stomatal type is not known from cordaita­ leans. Abaxial cuticles of Cordaites angulosostriatus also have stomatal rows (LEDRAN, 1960, 1966). Its reconstruction by RABITZ (1966) is more realistic. The adaxial cuticle of Corda- ites angulosostriatus has dispersed stomatal complexes. BARTHEL (1964) published Cordaites sp. type 4 from the Asturian of Zwickau that also has dark stomatal rows on abaxial cuticles and dispersed stomata on adaxial cuticles. Cordaites regu- laris described by LEDRAN (1966) has stomatal rows on the abaxial cuticle and dispersed stomata on the adaxial cuticle. Cu­ ticles from the other nine cordaitalean species described by LE­ DRAN (1966) have either stomatiferous bands or irregular sto­ matal rows on the abaxial cuticles. Several cordaitalean species have been described that have stomatal rows on the abaxial and adaxial cuticles: e.g. Cordaites principalis type 2 ZODROW et al. (2000) from the Bolsovian of Bay St. George, Newfoundland, Canada (see also ŠIMŮNEK, 2007). Similar cuticles are also known from the Bolsovian of the Czech Republic – Cordaites rerichensis (ŠIMŮNEK, 2007) and from the Asturian of the Heřmanova Huť locality – Cordaites pil- sensis (ŠIMŮNEK, 2007, 2008) and from the Blažkovice locality – Cordaites blazkovicensis ŠIMŮNEK, 2007; however, these spe­ cies have a “Cordaites principalis­type” venation (ŠIMŮNEK, 2007, 2008). The Stephanian species Cordaites risutensis ŠIMŮNEK, 2007 is similar to Cordaites cf. affinis having fine venation with 50–60 veins per cm, however the stomatal densities within the stomatal rows are not so high and individual stomatal complexes are separated by ordinary epidermal cells. The last comparable species is Cordaites radvanicensis ŠIMŮNEK, 2007 from the Stephanian of the Intrasudetic Basin, Czech Republic. The type material consists of leaves less than 30 mm wide with a venation consisting of 30 thick veins per mm and 2–3 thin sclerotic bundles between each of the two thick veins. However, the “thick” veins are not as thick as those in Cordaites principalis. When we count all the veins, there are 90 veins per cm. A comparison of the cuticles is interesting: it was demonstrated in Cordaites radvani- censis that on the adaxial cuticle there are dark stomatiferous bands 120–160 mm wide, separated by light non­stomatiferous bands 60–80 mm wide. The dark stomatal rows of the abaxial cu­ ticles are 50–70 mm wide and are separated by 50–80 mm wide non–stomatiferous bands. The data from the abaxial cuticle do not fully compare with Cordaites cf. affinis, but dark bands on the adaxial and abaxial cuticles are common for both species. It is pos­ sible that the specimen from Otovice, Černý potok, belongs to Cordaites radvanicensis ŠIMŮNEK, 2007. The guard cells of Cordaites cf. affinis are 28–40 mm long and the guard cells of Cordaites radvanicensis are 25–38 mm long (ŠIMŮNEK, 2007). Cordaites cf. roesslerianus GEINITZ, 1862 (Fig. 2f, l; Pl. 3, Figs. d–f) 1862 Cordaites roesslerianus n. sp.; GEINITZ: p. 149, pl. 35, fig. 5. Material: Museum of East Bohemia, Hradec Králové, specimen No. 74575. Description: The leaf fragment is 71.5 mm long and 15 mm wide; the margins are non­parallel, but the base and apex not preserved. G eologia C roatica Šimůnek Zbyněk: Revision of the genus Cordaites UNGER from the Permian of the Intrasudetic Basin (Broumov Formation, Olivětín Member, Czech 167 The venation differs from all previously mentioned species. The leaf relief is three­dimensional with prominent ribs and grooves that make prominent veins in oblique light; vein density 20 per cm of the leaf (Pl. 3, Fig. e). Thin veins were not observed. Remarks: The incomplete nature of the specimen renders the correct determination problematical. Cordaites roesslerianus GEINITZ are also preserved in fragments up to 120 mm long and 15–20 mm wide is similar. According to GEINITZ (1862), the vein density is 20–22 veins per cm. Cuticle macerations of Cordaites cf. roesslerianus were unsuccessful, with only the dark and light bands being discernible (Pl. 3, Fig. f). Further macera­ tion resulted in destruction of the cuticles. The dark bands that probably reflect vascular bundles are 370–500 mm wide and the light bands are only 170–300 mm wide. No sclerotic (thin) bun­ dles were observed. The other comparable species is Cordaites principalis (GERMAR) GEINITZ. Its holotype is preserved as a bunch of 32 leaves, 350–410 mm long and 15–36 mm wide. The leaf mar­ gins are slightly bent, as in the currently described specimen. The vein density of Cordaites principalis is 20–25 veins per cm (ŠIMŮNEK, 2015), which is also in concordance with the pre­ sently described specimen. However, the ridges in Cordaites principalis are more prominent and the places between the ridges are flatter. Perhaps C. roesslerianus is conspecific with C. prin- cipalis, however it is difficult to prove because the base and apex of C. roesslerianus is not preserved. LIPPS (1927) described Cordaites principalis from the Lower Permian of the Intrasudetic Basin (Unislaw Ślaski locality). It has 18–24 veins per cm an 3–4 sclerotic bundles between 2 veins. Cordaites cf. foliolatus GRAND’EURY (Fig. 2g; Pl. 3, Fig. g) 1877 Cordaites foliolatus GRAND`EURY; GRAND`EURY: p. 219, pl. XXI, fig. 3 Material: A single specimen from the Otovice locality is stored in the National Museum, Prague. Description: The leaf is small, narrow, 69.5 mm long and 13 mm wide. The base is 9 mm wide. The apex is blunt. The venation is not prominent, fine, about 40 veins per cm. Remarks: This specimen to some extent resembles Cordaites foliolatus GRAND`EURY. Proportionally, the specimen from the Otovice is much more slender than the GRAND’EURY type specimens of Cordaites foliolatus, which have a narrower base (5–7 mm) and wider leaves (10–25 mm). Length/width ratio of this specimen is 5.4, whereas in GRAND’EURY’s specimens it is around 4, and the apex is blunt to rounded, and the venation is prominent with 30 veins per cm and 4–6 sclerotic bundles be­ tween each pair of veins. It is also possible that the described specimen represents a juvenile leaf of some long-leaved form, therefore it is described here as Cordaites cf. foliolatus GRAND’EURY. Cordaites sp. A (Fig. 2d, e; Pl. 3, Figs. a-c) Material: Two fragments, stored in the National Museum, Prague. One represents the basal part and the other an apical part. They probably come from different leaves but may belong to one species. Description: The fragment from the basal part of a leaf is 51 mm long and only 12 mm wide near the base. The leaf widens rela­ tively progressively from the base. The second fragment repre­ sents a terminal part of a leaf. It is widest in the middle – about 28 mm and suddenly narrowed to the bluntly pointed apex. The venation is fine, but not dense, about 25 veins per cm of the leaf. Remarks: These specimens are difficult to compare with European Cordaites species. The estimated length of this leaf is 120–130 mm. The small­leaved forms described by GRAND’EURY (1877) and LEDRAN (1966) are usually wider and have a blunt or rounded apex. MEYEN (1966) described some similar forms as Rufloria MEYEN. Some species have a large variability, mostly also with rounded apices, but there were other forms with nearly bluntly pointed apices. However, the venation of Rufloria is quite different from Cordaites species. The shape of the leaf is to some extant similar to Cordaites laevis (GRAND’EURY, 1877, pl. 25, fig. 2), which represents a drawing of a twig with leaves and fructifications. Some leaves have blunt apices but others have bluntly pointed apices. Cordaites laevis is maybe a nomen nudum, because GRAND’EURY (1877) wrote only that the twig is with Cordaites laevis leaves and Antholithes fructifications. There was no description of the holotype, but the drawing of GRAND’EURY (1877) was illustrated many times in subsequent publications and also on the internet. 5. DISCUSSION It is difficult to classify Euramerican cordaitaleans only by mac­ romorphological characteristics because of the variability of the leaves; e.g. IGNATIEV & MEYEN (1989, fig. 6) showed great variability of Rufloria synensis. Similar leaves in the Eurameri­ can Realm could be classified to several species, if we only take into account leaf shape. However, when studying cordaitaleans from one locality, we do not have such variability, e.g. in the case of Cordaites borassifolius (ŠIMŮNEK et al, 2009): the length of complete leaves is about 400–700 mm and width 35–90 cm, but the leaf shape remains practically the same. Unfortunately, the localities near Otovice yielded specimens, where cuticular study was mostly not possible, so we are reliant on morphological features of cordaitalean leaves only. Still, it is not clear, if four or five Cordaites species are present in the Oto­ vice localities. In this paper, five species are described, but it can­ not be excluded that the specimens described here as Cordaites cf. beinertianus and C. cf. affinis do not belong to one species. The other species have different types of venation and Cordaites cf. foliolatus is small with a somewhat different leaf shape. CROOKALL (1970) and BARTHEL (1976) realised the dif­ ficulty in the determination of Cordaites species. CROOKALL (1970) used the names of species, but BARTHEL (1976) consi­ dered all species (e.g. Cordaites borassifolius, C. principalis and C. palmaeformis) as “one” species, even if diversity of the cuti­ cles was much higher, and BARTHEL (1976) and in subsequent papers (e.g. BARTHEL, 2009, 2016) preferred using “Cordaites sp.”. RAGOT (1966) studied many French species of Stephanian and Permian age, but Cordaites cuticles were poorly preserved and did not contribute to the identification of individual species. Due to their variability in leaves and venation all species in this paper are determined with some uncertainty as “cf.”, even if it is evident that more Cordaites species also occur in the Eurameri­ can Permian (Asselian). 6. FLORAL ASSEMBLAGE WITH Cordaites The flora of the Olivětín Member has been neglected in the palaeo­ botanical literature. Except for museum collections, there are un­ published works by RIEGER (1966) and BORS (1988). A taxo­ nomic list compiled from these works and museum collections consists of: Sigillaria brardii BRONGNIART, Calamites gigas G eo lo gi a C ro at ic a Geologia Croatica 72/3168 BRONGNIART, Palaeostachya thuringiaca (WEISS) BAR­ THEL, Cyathocarpus cyatheus (BRONGNIART) MOSBRUG­ GER, “Pecopteris” thuringiaca NĚMEJC, Autunia conferta (STERNBERG) KERP, A. naumannii (GUTBIER) KERP, Arnhardtia scheibei (GOTHAN) HAUBOLD et KERP, Spheno- pteris germanica WEISS, Odontopteris lingulata (GÖPPERT) SCHIMPER, O. subcrenulata ROST, Neurodontopteris auriculata (BRONGNIART) POTONIÉ, Barthelopteris germarii (GIEBEL) ZODROW & CLEAL, Taeniopteris coriacea GÖPPERT, T. cf. multinervia WEISS, Ernestiodendron filiciforme (SCHLOTHEIM ex STERNBERG) FLORIN, Culmitzschia angustifolia (FLO RIN) CLEMENT­WESTERHOF, C. parvifolia (FLORIN) CLEMENT ­ ­WESTERHOF, C. intermedia (FLORIN) CLEMENT­WES­ TERHOF, C. speciosa (FLORIN) CLEMENT­WESTERHOF, Hermitia goeppertiana (FLORIN) CLEMENT­WESTERHOF, H. hirmeri (FLORIN) CLEMENT­WESTERHOF, Otovicia hypnoides (BRONGNIART) KERP et al., Walchia piniformis SCHLOTHEIM ex STERNBERG and Gomphostrobus bifidus GEINITZ. This assemblage, which has been collected for more than 100 years, consists mainly of mesophilous to xerophilus elements. The hygrophilous species are practically limited to Sigillaria brardii and pecopteroids, which occur only very rarely. Walchian conifers dominate in the “walchia shales” and Autunia conferta is the most common species in the limestone facies. The other species occur partly in limestone and partly in the “walchia shales”. 7. CONCLUSION Cordaitaleans are not very common in the Permian of the Intra­ sudetic Basin. Seven specimens studied from the museum collec­ tions represent four or five species. The first specimen has been classified as Cordaites cf. bei- ne rtianus. The type material of Cordaites beinertianus has a simi­ lar leaf shape, but the veins are sparser than in C. cf. beinertianus. The leaves of C. cf. beinertianus also resemble Cordaites palma- eformis, which has slightly narrower leaves but a comparable vein density. A further two specimens were classified as Cordaites cf. af- finis. They have a comparable shape and venation with the type material of Cordaites affinis GRAND’EURY. However, the apex is not preserved, so a comparison of this feature is not possible. One specimen, from which the cuticles have been studied, is wider than typical C. affinis and could also be compared with C. beinertianus. The cuticles of Cordaites cf. affinis are poorly pre­ served, but stomatal rows and non­stomatiferous bands are dis­ tinguishable on both cuticles. The same cuticular type also occurs in Cordaites palmaeformis, and also C. radvanicensis, which is known from the Stephanian of the Intrasudetic Basin, but unfor­ tunately only as fragments. The second specimen of Cordaites cf. affinis is narrow and compares well with the variability of Cordaites affinis. On the other hand, Cordaites palmaeformis can also have narrow leaves and all specimens mentioned here could represent the variability of one species similar to Cordaites pal- maeformis or C. beinertianus. The next specimen has been classified as Cordaites cf. roes- slerianus. The determination is difficult because the base and apex are not preserved, however, the venation is very typical with wide and sparse veins. Such a venation is known from Cordaites roesslerianus and C. principalis. Another specimen has been classified as Cordaites cf. folio- latus. It resembles the type specimens of that species, however, it has somewhat narrower leaves. It is the smallest cordaitalean leaf from the Otovice locality and could simply represent a juve­ nile leaf of one of the other species. The final species is represented by two fragments: a basal and an apical part. 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(2000): New cuticular mor­ photypes of „Cordaites principalis (GERMAR) GEINITZ” from the Canadian Carboniferous Maritimes Basin.– Canadian Journal of Botany, 78/2, 135–148. G eo lo gi a C ro at ic a Geologia Croatica 72/3170 Plate 1 a – Cordaites cf. beinertianus (GÖPPERT) GRAND’EURY, coll. National Museum, Prague, no. E 7601, scale bar = 5 cm. b – Venation of the leaf from Fig. a., scale bar = 2 mm c – A leaf of Cordaites cf. affinis GRAND’EURY, coll. Museum of Eastern Bohemia, HRADEC KRÁLOVÉ, no. 74574, scale bar = 2 cm d – Venation of the leaf from Fig. c., scale bar = 2 mm e – A leaf of Cordaites cf. affinis GRAND’EURY, coll. National Museum, Prague, no. E 7601, scale bar = 2 cm f – Venation of the leaf from Fig. e., scale bar = 2 mm G eologia C roatica Šimůnek Zbyněk: Revision of the genus Cordaites UNGER from the Permian of the Intrasudetic Basin (Broumov Formation, Olivětín Member, Czech 171 Plate 2 Cuticles of Cordaites cf. affinis GRAND’EURY from the leaf figured on Pl. 1, Fig. c. a – b Adaxial and abaxial cuticle from the same leaf fragment. Slide 698/13, scale bar = 200 mm. c – h Abaxial cuticles. c. Detail from Fig. b. with two stomatal rows and non-stomatiferous bands. Scale bar = 100 mm. d. Abaxial cuticle with many stomatal rows. Slide 698/15, scale bar = 200 mm. e. Close up of Fig. d. with guard cells (arrow). Scale bar = 100 mm. f. Close up of two guard cells from Fig. e. Scale bar = 50 mm. g – h Two stomatal rows and a non-stomatiferous band, Slides. g. 698/7; h. 698/14, scale bar = 50 mm. G eo lo gi a C ro at ic a Geologia Croatica 72/3172 Plate 3 a – c Cordaites sp. A; a. Terminal part of the leaf, b. Basal part of the leaf, coll. National Museum, Prague, no. E 7602 and E 7603, (a and b – scale bar = 1 cm), c. Close up of the venation, scale bar = 2 mm. d – f Cordaites cf. roesslerianus GEINITZ. d. Fragment of the leaf, coll. Museum of Eastern Bohemia, HRADEC KRÁLOVÉ, no. 74575, scale bar = 1 cm, e. Close up of the venation, scale bar = 2 mm, f. Macerated leaf fragment with veins (vascular bundles) and tissue between them. Slide 700/3, scale bar = 500 mm. g Cordaites cf. foliolatus GRAND’EURY, coll. National Museum, Prague, no. E 7599, scale bar = 1 cm.