Ab strA ct Upper Westphalian coals are usually rich in organic-matter of different plant taxa, including gymnosperms. The as- semblage from Kalinovo (Eastern Ukraine) is exceptional in being dominated by Sphenophyllum, both cuticles and conducting tissues. Sphenophyllum cuticles are easily distinguished by their parallel oriented cells with sinuous an- ticlinal walls, and paracytic stomata on the abaxial cuticle. Tracheids with multiseriate bordered pits that occur in Sphenophyllum can be also found in the Calamitaceae and some pteridosperms. However, rectangular shaped remains of parenchyma cell strips along the radial wall of tracheids are only known in Sphenophyllum. Keywords: Sphenophyllum, dispersed cuticles, Carboniferous – Westphalian, Donets Basin Dispersed cuticles and conducting tissue of Sphenophyllum BRONGNIART from the Westphalian D of Kalinovo, Donets Basin, Ukraine  Zbyněk Šimůnek1 and Jan Bureš2 1 Czech Geological Survey, Klárov 131/3, 118 21 Praha 1, Czech Republic; (zbynek.simunek@geology.cz) 2 West Bohemian Muzeum Plzeň, Kopeckého sady 2, 30100 Plzeň, Czech Republic; (rallus@seznam.cz) doi: 10.4154/gc.2015.01 lar conditions in a peat-swamp, however, they come from different geographical regions and stratigraphical horizons. Sphenophyllum priveticense is known from the lower Bols- ovian (middle Westphalian) of Central Bohemia, whereas the Sphenophyllum sp. studied in this paper occurs in the up- permost Asturian (uppermost Westphalian) of the Kalinovo locality, Donets Basin, Ukraine. 2. GeoloGicAl settinG The Carboniferous sequence in the research area transgres- sively overlies marine and younger terrestrial Devonian de- posits. Carboniferous and Permian deposits in the southeast- ern part of the Donets Basin achieve a thickness of 12 km. Coal-bearing cyclothems consist of basinal marine to shal- low marine claystones and limestones with marine fauna; and terrestrial sandstones, siltstones, claystones with roots, and coal (HAvlEnA, 1965). Lithostratigraphic units are marked with letters and their subunits by letters with num- bers; limestones with capital letters and coal seams above the limestone with corresponding small letters (Fig. 1). Geologia CroaticaGeologia Croatica 1. introduction The genus Sphenophyllum BROnGnIART belongs to the sphenopsids and is characterized by linear, spatulate or fan- shaped leaves that extend up to several cm in length. The leaves are arranged along the axis in nodal whorls of six or nine. The name Sphenophyllum is used for both impression- compression and petrified Permo-Carboniferous samples. Although more than 100 compression-impression and petrified Sphenophyllum species have been described in the world, cuticles are known only from 15 of those species. The cuticles known so far, have been prepared by maceration of coalified Sphenophyllum leaves; studied directly on fossils by means of optical techniques (BARTHEl, 1997); or stud- ied in sections of coal balls (GOOD, 1973). This paper brings unique information on sphenophyllalean cuticles and con- ducting tissue obtained directly from the coal. Unfortunately, taxonomic affiliation was not possible because the leaf out- line is not preserved. Similar cuticles have only been previ- ously observed in Sphenophyllum priveticense lIBERTÍn et al., 2014. We presume that both species lived under simi- Geologia croatica 68/1 1–9 3 Figs. 1 tab. 3 Pls. Zagreb 2015 Geologia croatica 68/1Geologia Croatica 2 sandstones (AIzEnvERG et al. 1975). Mudstones with roots and coal seam no. n3 (0.3 m thick) are located above the sandstone. A sample from this coal seam from the Kalinovo locality, which is situated about 55 km W. from Luhansk in the Donets Basin (Fig. 2) was macerated for dispersed cuti- cles (sample position: N = 48° 35.355’; E = 38° 31,956’; H = 123 m). Above the coal seam, there are mudstones and claystones of lacustrine origin. Stigmaria occurs in the rooted horizon, while Linopteris obliqua (BUnBURY) zEIllER, Neuropteris ovata HOFFMAnn, Laveineopteris rarinervis (BUnBURY) ClEAl et al., Alethopteris sp., Annularia sphenophylloides (zEnKER) GUTBIER and Pecopteris sp. have been determined above the coal seam (AIzEnvERG et al. 1975). This flora is of Asturian (Westphalian D) age and is the youngest Westphalian flora found in the Donets Basin. 3. MAteriAl And Methods In order to obtain cuticles authors applied a new method. A 2.5 g sample of coal was macerated in Schulze’s reagent: 35 ml concentrated (65%) nitric acid (HNO3) and 1.5 g of po- tassium chlorate (KClO3) for 2 days and 18 hours. The black residue was fully washed under running water in a sieve and then treated with 10% potassium hydroxide (KOH) for up to one hour. During this process, the “coal matter” was com- pletely dissolved and only the cuticles and vascular tissue remained. The cuticles were stained in Safranin, Bismarck brown, Malachite green or neutral red, and mounted in Glycerine Jelly, or were attached to an SEM stub for obser- vation under a scanning electron microscope (SEM). About 30 foliar fragments of cuticles and a similar number of conducting tissue fragments were mounted on six slides. About ten cuticular fragments are longer than 1 mm (up to 1.3 mm), and represent both the adaxial and abaxial leaf surfaces. Some fragments of conducting tissue were up to 3 (4) mm long. The dispersed cuticles are stored in slides no. 595/1–6. Figure 1: Stratigraphy of the Pennsylvanian of the Donets Basin. The coal sample was taken from the upper part of the section. Adapted from PrivAlov et al. (2005). limestone n3 in the Isayevskaya Suite (Formation) is overlain by a 7 m thick sequence of unfossiliferous bluish- grey claystones and siltstones and about 30 m of fine-grained Figure 2: Map of Ukraine with Kalinovo locality highlighted. Šimůnek and bureš: Dispersed cuticles and conducting tissue of Sphenophyllum BroNGNiArT from the Westphalian D of Kalinovo, Donets Basin, Ukraine Geologia Croatica 3 4. systeMAtics Systematics performed according to ClEAl & THOMAS (1995) class equisetopsida (‘horsetails’) order bowmanitales (‘sphenophylls’) Meyen, 1978 Family Bowmanitaceae Meyen, 1987 Genus Sphenophyllum bronGniArt, 1828 Sphenophyllum sp. (Fig. 3, Pls. 1, 2, 3) Description: Adaxial cuticle: The cells are elongate, more or less fusi- form or elongate tetragonal in shape with coarsely sinuous anticlinal walls that affect the original cell shape. The cells are oriented parallel to the veins, and are 150–300 μm long and 25–50 μm wide. The costal (vein) areas are usually prominent with original black vascular tissue (Pl. 1, fig. 2), which is about 100 μm wide. When this tissue is not pre- served (Pl. 1, fig. 1), the costal area is about 50 μm wide and shows narrow cells only 15–20 μm wide. Abaxial cuticle: The cells have essentially the same shape and dimensions as the cells of the adaxial cuticle. The difference is in the presence of stomata (Pl. 1, figs. 3–4). The stomata tend to be concentrated in “bands” along the veins and they are oriented parallel to the veins. The guard cell pairs are fusiform or elliptical, 26–32 μm long and 10–14 μm wide (Pl. 1, figs. 4, 6, Pl. 2, fig. 4). Each stoma has two lateral subsidiary cells (paracytic stomatal type), one of which is shorter – 78–90 μm long; and the longer is 176–192 μm long. Both cells have approximately the same width – 20–35 μm. The cuticles in scanning electron microscope have very fine striated periclinal walls in outer view (Pl. 2, fig. 1) and the anticlinal walls are prominent in the inner view (Pl. 2, figs. 2–4B). Using the same approach as described by POOlE & KüRSCHnER (1999) stomatal density and stomatal index were counted from several cuticle fragments; the area of each fragment was 0.1–0.15 mm2. The stomatal density of Sphe- nophyllum sp. is 83–90 stomata per mm2 and the stomatal index is 20.5–23. Conducting tissue: Fourteen tracheids from slides 595/1, 2, 3 and 5 were studied. Fragments of tracheids have walls perforated mainly by multiseriate bordered pits (Pl. 3, figs. 1, 3), which locally become in reticulate cell wall thickening (Pl. 3, fig. 4). Bordered pits are free, and according to the number of their rows and tight configuration they belong to the alternate pitting surface type . The number of bordered pit rows on the tracheid wall is (2)–4–6. Bordered pits have a circular to elliptical shape and approximately the same di- mension. The tracheids are 60–100–(125) mm wide. The thickening on some tracheids is interrupted by the walls of parenchyma cells that form a rectangular outline on the tra- cheid (Pl. 3, fig. 2). Due to the fragmentary size of the pre- served tracheids, it was not possible to determine their orig- inal total length. 5. reMArKs Fragments of the xylem – tracheids were found in association with the Sphenophyllum cuticles. According to TAYlOR et al. (2009, p. 206), annular or helical thickenings of tracheids are most often found in the earliest matured primary xylem. Secondary xylem is made up predominantly of pitted tra- cheids, although some plant groups also have secondary xy- lem tracheids. Bordered pits in tracheal elements are accord- ing to NěMEjC (1963, p.76) very rare in many pteridophylls (Pteridophyta). It is common in equisetopsids (Equisetopsida) and in many pteridosperms and gymnosperms. 6. Sphenophyllum cuticles And their coMPArison Most species of Sphenophyllum BROnGnIART have irre- gu larly isodiametric to elongate epidermal cells that usually posses sinuous anticlinal walls. Stomatal complexes occur on the abaxial surface of the leaf and consist of two guard cells with polar and circumpolar thickenings, and two para- cytic lateral subsidiary cells. Subsidiary cells may have the same shape and size as normal epidermal cells, or they can differ. One subsidiary cell is usually distinctly larger than the other (TAYlOR et al. 2009). These characteristics fit very well with the cuticles de- scribed above. Individual species differ in the distribution of the stomata, the orientation and size of the guard cells, and the shape, arrangement and size of the intercostal cells (Ta- ble 1). BARTHEl (1997) distinguished two groups among the Sphenophyllum species. Group 1 consists of Sphenophyl- lum cuneifolium, S. emarginatum, S. majus ABBOT (=S. geinitzii STORCH), S. thonii, S. speciosum. S. longifolium and S. saxonicum. This group is hypostomatic, with sinuous anticlinal walls and larger cells. Group 2 consists of Sphe- nophyllum oblongifolium and the anatomically preserved species S. quadrifidum REnAUlT and S. reedae GOOD. In this group, the epidermis has rectangular cells with straight anticlinal walls. Figure 3: A paracytic stoma of Spheno- phyllum sp. from Kalinovo locality. Note the uneven lateral subsidiary cells, ls = lat- eral subsidiary cells, g = guard cells. Scale bar = 50 μm. Geologia croatica 68/1Geologia Croatica 4 Plate 1 Sphenophyllum sp., locality Kalinovo, Donets Basin, coal seam n3, Upper Asturian (Moscovian, Pennsylvanian) 1 and 2 – Adaxial cuticle, slide 595/3, scale bar = 100 μm; 2 – Note black costal area. 3–6 – Abaxial cuticle; Fig. 3 – Cuticle with black costal area and holes where guard cells, have fallen out, slide 595/3, scale bar = 100 μm; 4 – Detail of a stomatal complex with guard cells, scale bar = 20 μm; 5 – Cuticle with 5 stomatal complexes, slide 595/5, scale bar = 50 μm; 6 – Close up to two stomatal complexes from fig. 5 scale bar = 20 μm Šimůnek and bureš: Dispersed cuticles and conducting tissue of Sphenophyllum BroNGNiArT from the Westphalian D of Kalinovo, Donets Basin, Ukraine Geologia Croatica 5 Plate 2 Sphenophyllum sp., loc. Kalinovo, Donets Basin, coal seam n3, Upper Asturian (Moscovian, Pennsylvanian) cuticles in scanning electron micro- scope. SEM stub no. 84. 1 – Adaxial or abaxial cuticle in outer view. Note the periclinal walls with very fine longitudinal striations and sinuous anticlinal walls, scale bar = 50 μm. 2 – Adaxial or abaxial cuticle in inner view. Note prominent sinuous anticlinal walls, scale bar = 50 μm. 3 – Abaxial cuticle with a stoma (S) in inner view, scale bar = 50 μm. 4 – Close up of the stomatal complex from fig. 3, scale bar = 20 μm. Geologia croatica 68/1Geologia Croatica 6 Plate 3 Typical structure of secondary xylem tracheids in the studied specimens resembling the tracheid structure of Sphenophyllum sp. 1 – Secondary xylem tracheids with multiseriate bordered pits, the shape of the bordered pits is circular – elliptical, slide 595/3. Scale bar = 100 µm. 2 – view of the tracheid walls in places where strips of parenchyma cells were attached. p – tracheid surface (? wall) in the place where parenchyma cells were attached. c – Contiguous tracheid wall is demarcated and perforated. The tracheid has reticulate cell wall thickening. Slide 595/3, scale bar = 100 µm. 3 – A close up of tracheidal wall with pentastichous pitting of a circular shape. SEM stub 84. Scale bar = 20 µm. 4 – A close up of a tracheidal wall, a – circular bordered pits, b – elliptical bordered pits, c – reticulate cell wall thickening, the tracheid wall is partly sag- ging. SEM stub 84, scale bar = 20 µm. Šimůnek and bureš: Dispersed cuticles and conducting tissue of Sphenophyllum BroNGNiArT from the Westphalian D of Kalinovo, Donets Basin, Ukraine Geologia Croatica 7 The Sphenophyllum sp. described in the present paper belongs to Group 1. The elongate cells with coarsely sinu- ous anticlinal walls occur in many species: e.g. Sphenophyl- lum emarginatum, S. cuneifolium, S. thonii, S. majus ABBOT (non BROnn) (=S. geinitzii), S. kobatake and S. zwicka- wense. However, the cell size in the present species (150–300 μm) is rather large compared to all the species mentioned above, where it is usually only up to 200 μm. The exception is the newly described Sphenophyllum priveticense lIB- ERTÍn, BEK et DRáBKOvá, 2014, in which the cells are 125–350 μm long and so more comparable to our samples of Sphenophyllum sp. However, they differ in other charac- teristics (see Tab. 1) and it is unlikely that they belong to the same species, especially as Sphenophyllum priveticense is of early Bolsovian age whereas the present Sphenophyllum sp. is of latest Asturian age. Sphenophyllum guard cells are usually reniform and the outline of the pair is elliptical. Their size differs according to species, however some overlap exists. Our Sphenophyl- lum sp. has medium sized guard cells among the spheno- phylls, 28–32 μm in length. The smallest ones are known from permineralised S. multirame (only 15 μm long) and the largest ones are known in four species S. emarginatum (31– 57 μm), S. apiciseratum (34–54 μm), S. koboense (27–54 μm) and S. priveticense (45 μm). An important feature is also the shape of the subsidiary cells. Many species have two lateral subsidiary cells per stoma (usually one of them is larger than the other); they are of the same shape as normal epidermal cells in Sphenophyllum sp., S. emarginatum, S. thonii, S. spe- ciosum, S. longifolium, S. apiciseratum, S. koboense and S. priveticense. According to lIBERTIn et al. (2014), the sto- matal complex of S. priveticense is anomocytic, but it seems that all those mentioned above have paracytic stomata. In other species, the subsidiary cells differ significantly from the ordinary epidermal cells: Sphenophyllum cuneifolium, S. thonii, S. geinitzii (S. majus ABBOT), S. saxonicum, S. mul- tirame and S. reedae. Some species have trichomes and papillae on the adaxial surface of the leaves: Sphenophyllum majus, S. speciosum, S. saarensis, S. trichomatosum STUR and probably also S. sewardii BATEnBURG (lIBERTÍn et al. 2014). No trichomes or emergences have been observed on cuticles from Sphenophyllum sp. from Kalinovo. 7. coMPArison oF conductinG eleMents The characteristic pitting of the tracheid walls in the studied samples corresponds to the description of secondary xylem anatomy of the Sphenophyllum plant as mentioned in table 1: Epidermal structures of Sphenophyllum species. Species (C-compression; P-petrification) Distribution of stomata orientation of guard cells Guard cell size [mm] Subsidiary cells no./specialised in shape and size Shape and arrangement of intercostal cells intercostal cell size [mm] Shape of anticlinal walls references S. emarginatum BroNGNiArT [C] random usually parallel to veins 31-57x5-18 2/no long, irregular 79-179x26-50 coarsely sinuous BATENBUrG (1977), (1981) S. cuneifolium (STErNBErG) ZEillEr [C] random parallel to veins 25x15 2/few long, irregular 80-200x30-60 coarsely sinuous BArTHEl (1997) S. thonii MAHr [C] random to stripped parallel to veins 30x20 2/yes long, irregular 100-200x20-30 coarsely sinuous MEYEN (1970) S. majus ABBoT non BroNN (=S. geinitzii STorCH) [C] random random 30x20 5-6/few long, irregular 40-60-10-15 coarsely sinuous ABBoT (1958) S. speciosum (roYlE) ZEillEr [C] random random 25x15 2/no long, irregular 100-200x30-70 sinuous PANT & MEHrA (1963) S. longifolium (GErMAr) GUTBiEr[C] random parallel to veins 20-28x12-15 2/no longitudinal rows 140-200x40-55 finely sinuous BArTHEl (1997) S. saxonicum rEMY & rEMY [C] random parallel to veins 25x15 2/few longitudinal rows 100-200x35-40 finely sinuous BArTHEl (1997) S. oblongifolium (GErMAr & KAUlFUSS) UNGEr [C] ? ? ? ? rectangular in rows 100-180x15-25 straight BArTHEl (1997) S. multirame DArrAH (P) in 2 sunken rows or furrows random 15x10 2-4/yes long, irregular 100-150x15-25 finely sinuous GooD (1973) S. reedae GooD (P) random parallel to veins 25x18 3-5/few rectangular in rows 70-150x20-30 straight GooD (1973) S. apiciseratum YAo et al. [C] random random 34-54x14-24 2/no irregular 65-147x 41-112 coarsely sinuous YAo et al. (2000) S. koboense KoBATAKE [C] random (near veins) random 27-54x10-24 2/no long, irregular 81-189x20-41 finely to coarsely sinuous YAo et al. (2000) S. zwickauvense STorCH [C] ? ? ? ? longitudinal rows 60-177x14-52 sinuous BATENBUrG (1981) S. priveticense liBErTÍN, BEK & DrÁBKovÁ [C] in rows parallel to veins 45x15 2/few longitudinal rows 125-350x50-60 sinuous liBErTÍN et al. (2014) S. sp. [dispersed] near veins parallel to veins 26-32x10-14 2/no longitudinal rows 150-300x25-50 coarsely sinuous present study only well preserved compression [C] species, and petrifaction (P) species whose foliar grooss morphology is known, have been included. other compression species are known only by fragments without stomata (see BATEN BUrG, 1981). Geologia croatica 68/1Geologia Croatica 8 BOUREAU (1964, p. 98, 99), who figured Sphenophyllum with multiseriate bordered pits. He also figured the distribu- tion of parenchyma cell strips along the radial walls of a tra- cheid. It has been shown that strips of parenchyma cells cre- ated a rectangular contour on the radial wall of the tracheids. These contours are the only remnants of parenchyma cells. TAYlOR et al. (2009) also described tracheids of secondary xylem with circular–eliptical bordered pits on the lateral walls in Sphenophyllum plurifoliatum WIllIAMSOn et SCOTT. Similarly, BUREš et al. (2013) described tracheids with circular–elliptical bordered pits in Sphenophyllum cf. myriophyllum CRéPIn. BATEnBURG (1982) described xylem with scalariform thickenings and elements with uni- or multiseriate bordered pits in Sphenophylllum speciosum zEIllER (PAnT & MEHRA). RÖSSlER & nOll (2007 p. 174) described a similar structure in the secondary xylem of Calamitea striata COTTA (Calamitaceae). Cell walls show simple to bifurcate scalariform to reticulate thickening with elongate oval pits in radial and tangential sections. Mul- tiseriated bordered pits of tracheids are common in pterido- sperms as noted by AnDREWS (1940), NěMEjC (1968), and TAYlOR et al. (2009). 8. conclusion Cuticles and conducting tissue of Sphenophyllum BROnG- nIART have for the first time been identified in the dispersed cuticle spectrum. Many Sphenophyllum species have the same cuticular pattern, so it is difficult to identify Spheno- phyllum species only using cuticles, as noted by BARTHEl (1997). nevertheless, as it has paracytic stomata and elon- gate cells with sinuous anticlinal walls, the Sphenophyllum from Kalinovo clearly belongs to group 1 (sensu BARTHEl 1997) together with Sphenophyllum cuneifolium, S. emar- ginatum, S. thonii, S. speciosum. S. longifolium, S. saxoni- cum and S. priveticense. Although the Kalinovo species has medium-sized guard cells that fit with several species, to- gether with Sphenophyllum priveticense lIBERTÍn, BEK et DRáBKOvá, 2014, it has the longest cells that have been reported so far in a Sphenophyllum species, 300 – 350 μm long, in contrast to usually no more than 200 μm long seen in other species. Despite the similarity of their cuticles, the Sphenophyllum sp. from Kalinovo and Sphenophyllum priv- eticense cannot be regarded as conspecific as the former is stratigraphically much younger. Sphenophyllum sp. was discovered in Kalinovo coal, and S. priveticense was found in a tuff that buried the peat-swamp “in situ” and also contained many peat-forming floral ele- ments. It is likely, therefore, that both Sphenophyllum sp. and S. priveticense lived in similar habitats. lIBERTÍn et al. (2014) supposed that Sphenophyllum priveticense lived in peat-swamps with a high water table and also in slightly drier habitats with a transition to an Omphalophloios-phase. Maybe the larger cell dimension is caused by living in such an environment. It is not possible to systematically classify the studied tracheid fragments of the genus Sphenophyllum sp. based on tracheid thickening. A similar structure of tracheids is also common in the other plant groups from the Pennsylvanian (Calamitaceae, pteridosperms). Any palaeoecological inter- pretation based on the tracheids is not possible, as their width depends on vegetation conditions, and we are not sure if all the studied tracheids in fact belong to this Sphenophyllum. AcKnoWledGeMent This study was part of the International Geoscience Pro- gramme (IGCP Project 575) and the GACR project P210/10/0232. The authors are obliged to V. I. POlETAEv, n. BOYARInA and A. SCHEGOlEv (Kiev, Ukraine) for guiding in the field and to C.j. 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