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Research paper

New sphenophyllaleans from the Pennsylvanian of the Czech Republic

Milan Libertín a, Jiří Bek b,⁎, Jana Drábková c

a National Museum, Václavské nám. 64, 118 21 Prague 1, Czech Republicb Department of Palaeobiology and Palaeoecology, Institute of Geology v.v.i., Academy of Sciences, Rozvojová 269, 165 00 Prague 6, Czech Republicc Czech Geological Survey, Klárov 3, Prague 1, Czech Republic

a b s t r a c ta r t i c l e i n f o

Article history:Received 21 May 2013Received in revised form 26 August 2013Accepted 27 September 2013Available online 16 October 2013

Keywords:SphenophyllumBowmanitesin situ sporemonolete sporeCarboniferous

Two new species of sphenophyllalean cones, Bowmanites myriophyllus sp. nov. and Bowmanites priveticensissp. nov., from the Radnice Basin of the Czech Republic are proposed. The most characteristic feature is thesporangiophore having lanceolate expansions bearing sporangia. Strobili were studied morphologically,including cuticle analysis and examination of in situ spores. Both new species yielded monolete spores of theLaevigatosporites/Latosporites-type. A review is given of the dispersed and in situ Laevigatosporites/Latosporitesspores of sphenopsid origin and their parent plants are compared. A new species of the genus SphenophyllumBrongniart, Sphenophyllum priveticense sp. nov. is proposed and the diagnosis of Sphenophyllum myriophyllumCrépin is emended. Some B. myriophyllus cones are born on vegetative axes with leaves of the Sphenophyllummyriophyllum-type and some B. priveticensis cones are connected with stems and leaves of S. priveticense.Epidermal structures of the leaves (hook-like structures at the end of lobes of leaves) of both Sphenophyllumspecies could enable plants to climb. The stratigraphical interval of all four species is the Whetstone volcanichorizon, directly overlying the Lower Radnice Coal, Radnice Member, Kladno Formation, Lower Bolsovian,Pennsylvanian.

© 2013 Elsevier B.V. All rights reserved.

1. Introduction

Sphenophyllalean plants are ecologically andmainly stratigraphicallysignificant fossils (Němejc, 1953). The classification of sphenophyllshas largely been based on their vegetative organs (stems, leaves);a large number of “leafy” taxa were described (Abbott, 1958; Boureau,1964; Storch, 1981), but several of them are probably synonymous dueto prominent heterophylly and the fragmentary nature of specimens.Sphenophyllalean reproductive organs (strobili) are rare in the fossilrecord, but several of them are considered good biostratigraphicmarkers(Libertín et al., 2005, 2008). The earliest sphenophylls are reported fromDevonian strata but they were most abundant in the Carboniferous,especially in the Westphalian. Most sphenophylls became extinct attheWestphalian–Stephanian boundary and their last known occurrenceis from strata of the Permian age (Boureau, 1964).

Almost all sphenophyllalean cones have been assigned to the genusBowmanitesBinney, despite differentmodes of preservation (petrifactionsand compressions). Themain criteria for their classification have been themorphology and/or anatomy of the cones; the significance of their in situspores was often ignored. This contrasts with the modern concept of theclassification of fossil reproductive organs, where in situ spores are anequally important part of diagnoses and the descriptions of their parent

plants (e.g., Thomas, 1970; Bek and Opluštil, 2004, 2006). The authorsfollow this concept, and are working (e.g., Bek, 1998; Libertín andBek, 2003, 2006) toward a new approach for the classification ofCarboniferous sphenophyllalean fertile organs based on the combinationofmorphological and/or anatomical features of the cones and their in situspores. This new classification for Carboniferous sphenophyllaleancones will be closer to the natural systematics for sphenophyllsand preliminarily consists of eight morpho-genera (e.g., Libertínand Bek, 2006). Each of these new morpho-genera possesses adifferent characterization of the cones and different in situ spores.The authors are convinced that this concept reflects a more naturaldivision of these plants that can help with the general interpretationof the whole group of sphenophylls.

The genus Sphenophyllum Brongniart was erected by Brongniart(1822) to formally accommodate fossil plants formerly namedSphenophyllites Brongniart. Binney (1871) proposed the nameBowmanites for a compressed sphenophyllalean cone attached toa leafy axis. Sphenophyllostachys Seward was proposed for conesof Sphenophyllum by Seward (1898). Sternberg (1821) describedsphenophyllalean plants from the Carboniferous of the CzechRepublic, first as Rotularia Sternberg (Sternberg, 1821) and thenas Volkmannia Sternberg (Sternberg, 1833). The name Rotularia isrejected in favour of Sphenophyllum and Volkmannia is a genusthat is no longer used, but has been used to accommodate varioussphenopsid strobili that are now assigned to other morpho-generasuch as CalamostachysWeiss and PalaeostachyaWeiss, and even sterile

Review of Palaeobotany and Palynology 200 (2014) 196–210

⁎ Corresponding author.E-mail addresses: milan_libertin@nm.cz (M. Libertín), mrbean@gli.cas.cz (J. Bek),

jana.drabkova@geology.cz (J. Drábková).

0034-6667/$ – see front matter © 2013 Elsevier B.V. All rights reserved.http://dx.doi.org/10.1016/j.revpalbo.2013.09.008

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axes. Bek (1998) and Bek and Opluštil (1998) were the first to describein situ spores from sphenophyllalean plants in the Czech Republic.

There have been two approaches to the subdivision and classificationof sphenophyllalean cones. The first divides them into three groups(Jugati, Conferti and Simplices) based on the number and positionof sporangia on each sporangiophore (Hoskins and Cross, 1943).The second approach was suggested by Remy (1955), who proposedfour new sphenophyllmorpho-genera,KoinostachysRemy,AspidiostachysRemy, Lilpopia Conert and Schaarschmidt and Anastachys Remy, basedon similar criteria. Remy (1955) especially focused on the numberof sporangia, but he also included some non-sphenophyllalean taxa(e.g., Anastachys). Both of these divisions combined parent plantsproducing monolete, trilete and trilete operculate spores together intoone group and/or genus.

This paper focuses on the study of two species of Sphenophyllumwhich bear monolete-producing sphenophyllalean cones of the genusBowmanites from the Pennsylvanian of the Czech Republic. This researchcombines both palaeobotanical (mainly cuticular analysis) andpalynological methods of investigation. Cones were studied underbinocular and scanning electron microscopes (SEM) and their insitu spores are described.

2. Material and methods

Specimens of Bowmanites myriophyllus sp. nov., Bowmanitespriveticensis sp. nov., and Sphenophyllum myriophyllum Crépinare from Radnice (Pokrok Mine/Ovčín), Radnice Basin, Whetstonevolcanic horizon, Radnice Member, Kladno Formation, Lower Bolsovian,Pennsylvanian. Specimens of B. myriophyllus: E6324, E6415a,b andE6497 (holotype) are stored in the National Museum, Prague, CzechRepublic. Specimens of B. priveticensis sp. nov.: F11355 and F11356(holotype) are stored in the West Bohemian Museum, Pilsen and thespecimen E6305 is in the National Museum, Prague, Czech Republic.Specimens of S. myriophyllum: E54 (lectotype), E3001, E3009, E3011,E5417 and E5634 are stored in the National Museum, Prague. Specimensof Sphenophyllum priveticense are from Přívětice (Pokrok Mine/Ovčín),specimens F11355-6 are stored in the West Bohemian Museum, Pilsen;E5414, E5416, E6498, E6499 and E6306 (holotype) are from Přívětice(Pokrok Mine/Ovčín) and are stored in the National Museum, Prague,Czech Republic.

A Nikon Eclipse 80i microscope was used for the study ofthe specimens. Samples for cuticular analysis were macerated inhydrofluoric acid (70%) for 12 h and then in a Schulze's solution for16 h. Samples were afterwards neutralised in KOH (5%) for 8–10min.Cuticles were observed using a Nikon microscope. Isolated sporangiaand leaves were studied with Hitachi SEM. Spores were recoveredby dissolving small portions (separated from the cone specimenswith a mounted needle) of cones in nitric acid for 24–48 h and KOHfor 1–2 h. Most spores were mounted in glycerine jelly for directmicroscopic examination. Some spores were sputter-coated with goldfor examination with a Cameca SX100 SEM. Descriptive terms forthe spores follow the latest edition of the Glossary of Pollen andSpore Terminology (Punt et al., 2007). Spores are classified accordingto the system of dispersed spores suggested by Potonié and Remp

(1954), Potonié and Kremp (1955), Dettmann (1963), and Smithand Butterworth (1967). In situ spores were compared directlywith the original diagnoses (type specimens), descriptions, andillustrations of dispersed spore species. Species determinationsare based only on the original diagnoses, and not on the interpretationsof subsequent authors.

There was no possibility for the authors to physically studyall holotypes and specimens of sphenophyllalean cones producingmonolete spores. Species described herein are compared accordingto the descriptions by first authors. Therefore, the taxonomic classesare identical to the original works of those authors (summarised inTable 1). Map showing the geographical position of the localities ofBowmanites myriophyllus, B. priveticensis, Sphenophyllum priveticense andS. myriophyllum is in Fig. 1.

Table 1Selected measurements of Laevigatosporites/Latosporites-producing sphenopsids.

B. nindelii B. simonii S. oblongifolium S. laciniatum B. bifurcatus S. kettneri S. zwickaviense Leeites oblongifolis

Size of sporangia (mm) 0.5–0.7 0.5–1 0.8–1.2 0.4–0.5 0.6–1 0.5–0.8 0.5–1 1–1.3Number of sporangiaper sporagiophore

? ? 6 ? 4 4 2? 4?

Cone length (mm) 40 40 ➢ 50 ➢ 25 ➢ 40 10–16 55 50Cone width (mm) 10–12 5–8 10 6 3 5–8 6–9 5Shape of bracts Linear Lanceolate Peltate Cuneiform spread

to 2 lanceolate lobesLanceolate Cuneiform spread

to 3 lanceolate lobesCuneiform spreadto 4 lanceolate lobes

Claw-like

Number of bracts ? ? 6 6 ? 6 ? 6?

Fig. 1.Map showing the geographical position of the localities of Bowmanitesmyriophyllussp. nov., B. priveticensis sp. nov., Sphenophyllum priveticense sp. nov. and S. myriophyllum(Crépin) emend.

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3. Systematic palaeontology

Order sphenophyllalesFamily sphenophyllaceaeGenus Bowmanites Binney, 1871 emend. Hoskins and Cross, 1943Type species B. cambrensis Binney, 1871

Bowmanites myriophyllus sp. nov.Plate I, 1–2; Plate II, 5, 7–9; Plate IIIHolotype: E6497 (Pl. I, 1), National Museum, Prague, Czech Republic.Etymology: Following the name of the parent plant, Sphenophyllummyriophyllum which produced these cones.Type locality: Radnice (Pokrok Mine/Ovčín), Radnice Basin, CzechRepublic.Type horizon: Base of theWhetstone volcanic horizon, directly overlyingthe Lower Radnice Coal, Radnice Member, Kladno Formation, LowerBolsovian, Pennsylvanian.Material: Specimens E6324, E6415a,b and E6497 (holotype), NationalMuseum, Prague.Diagnosis: Cylindrical strobili with sterile apices. Sporangiophoreswith scutelliform margins between axis and sterile bracts arranged inwhorls. Five pyriform sporangia at scutelliform apex of sporangiophore.Cuneiform sterile bracts divided into two or four narrow thread-likelobes. Margins of sterile bracts serrate. Teeth directed towards apexof leaves. Lobes of leaves with terminal hook-like apex. Twelve sterilebracts in whorls equal to the number of ribs on the axis. Monoletespores circular to oval. Exine laevigate. Laesurae simple, one-third tothree-quarters of spore diameter in length.Description: Cylindrical strobili are more than 60 mm long, and10–12 mm wide without sterile bracts. Strobili are always connectedterminally with sterile stems (Plate I, 1). Internodes in cones are6–8mm long (Plate I, 1; Plate III, 1). Stems are 2–3mmwide on average.Sporangiophores are 1–3mm long with scutelliform margins (Fig. 2. A;Plate II, 8, 9). Pyriform sporangia are 0.5–1.1 mm wide (Plate II, 5,7–9). The cuticles of sporangia are milled. Sterile bracts in strobili areidentical to leaves on sterile stems. Cuneiform leaves are divided intotwo to four very narrow lobes (Plate I, 3, 7; Plate II, 1, 6) with themedialincision always more than two-thirds of the whole diameter. The depthof distal incisions is about five-sixths of the whole length of leaves. Thelength of leaves is 10–40 mm and the width of lobes is 0.3–0.5 mm.Only one vein per leaf that divides into the leaf lobes. Margins of leafylobes are serrate with hook-like apices. Sporangia and sporangiophoresabscised and therefore some cones are not compact (Plate I, 1–2). Somesporangiophores possess sessile sporangia (Plate II, 5, 7–9). Monoletelaevigate thin-walled spores (Plate III, 2–5). Amb is oval to circular,margin smooth. Diameter is 35 (49) 78 μm. Exine is about 1 μm thick,seldom folded.Remarks: Spores with a circular amb correspond to the spore genusLatosporites Potonié & Kremp, and spores with an oval amb belongto the genus Laevigatosporites Ibrahim. Spores with an oval ambare more common (about 80–85%) than those with a circular amb(15–20%).

Bowmanites priveticensis sp. nov.Plate IV, 1, 5–7; Plate V, 8–11Holotype: Specimen F11356 (Pl. IV, 1),West BohemianMuseum, Pilsen,Czech Republic.

Etymology: After Přívětice, the type locality.Type locality: Přívětice (PokrokMine/Ovčín), Radnice Basin, CzechRepublic.Type horizon: Base of theWhetstone volcanic horizon, directly overlyingthe Lower Radnice Coal, Radnice Member, Kladno Formation, LowerBolsovian, Pennsylvanian.Material: Specimens F11355 and F11356 (holotype) stored in the WestBohemian Museum, Pilsen; specimen E6305 stored in the NationalMuseum, Prague, Czech Republic.Diagnosis: Cylindrical strobili with sterile apices. Sporangiophoresbetween axis and sterile bracts arranged in whorls. Sporangiophoresrelatively long with scutelliform apices. Four pyriform sporangia onsporangiophore. Six sterile bracts per verticil with cuneiform expansionson two lanceolate leafy lobes. The leafy lobes are terminated by two fork-like teeth. Monolete spores circular to oval. Exine laevigate. Laesuraeone-third to three-quarters of spore diameter in length.Description: Cylindrical strobili are not always compact and possess asterile apex (Plate IV, 6). Strobili are connected terminally with distalstems (Plate IV, 1, 5) and 50–100mm long, and 10–12mmwidewithoutsterile bracts. Internodes are 6–8mm long. Stems are 2–3mm wideon average. Sporangiophores are 3 mm long. Pyriform sporangia are1–2 mm wide on average (Plate IV, 7). Cuneiform sterile bractsin cones are divided into two lanceolate lobes (Plate IV, 6). Lobemarginsare smooth and lack a hook-like apex. Sterile bracts near the apexpossess a broader and slightly convex frontal margin characterised byround teeth without prolonged vein (Plate IV, 6). Monolete laevigatethin-walled spores. Amb is oval to circular, margin smooth. Diameteris 30 (45) 76 μm. Exine is about 1 μm thick, seldom folded.Remarks: Spores with a circular amb correspond to the spore genusLatosporites, and those with an oval amb to the genus Laevigatosporites(Plate V, 8–11). Spores with an oval amb are generally more common(about 80–85%) than those with a circular amb (15–20%).

Genus Sphenophyllum Brongniart, 1828Type species S. emarginatum Brongniart, 1828

Sphenophyllum myriophyllum Crépin emend.Plate I, 3–7; Plate II, 4, 6, 8–9; Fig. 4

Synonymy: 1823Volkmannia gracilis— Sternberg p. 53; taf. 15, Fig. 1, nonFigs. 2, 3.

1880 Sphenophyllum myriophyllum sp. nov. — Crépin, p. 25–28.1966 Sphenophyllum myriophyllum Crépin — Storch, p. 275.1968 Sphenophyllum myriophyllum Crépin — W. Remy, p. 125–130;

pl. 23, 24.1980 Sphenophyllum myriophyllum Crépin — Storch, p. 198–200.

Lectotype: E54 (No. 125, Pl. I, 5–6) selected by Storch (1981); Sternberg,1833, p. 53; pl. 15, Fig. 1.Material: Specimens E54, E3001, E3009, E3011, E5417 and E5634.Emended diagnosis: Stems monopodially branched with prominentexpansion in nodes. Cuneiform leaves divided into four narrow linearlobes. Leaves are divided by central incision close to the base and bytwo marginal incisions to two-thirds of the length of leaves. Marginsof leafy lobes serrate. Teeth of serrate margin directed towards tipof leaf lobe. Leaf lobes with hook-like tips. Each tooth comprises of asingle cell. Epidermal cells rectangular without undulate anticlinalwalls.

Plate I. 1. Bowmanites myriophyllus sp. nov. (E6497 holotype), Pokrok Mine (Ovčín). General view. 2. Bowmanites myriophyllus sp. nov. (E5415a), Pokrok Mine (Ovčín). Cone with sterileapex. 3. Sphenophyllummyriophyllum Crépin (E5417), PokrokMine (Ovčín). Detail of the apex. 4. Sphenophyllummyriophyllum Crépin (E3011), PragoMine. General view of the distal part.5. Sphenophyllum myriophyllum Crépin (E54 lectotype), Radnice locality. General view. Notice vegetative axes with leaves arranged in verticils. 6. Sphenophyllum myriophyllum Crépin(E54 lectotype), Radnice locality. Monopodial branching. 7. Detail of cuneiform leaves of Sphenophyllum myriophyllum Crépin (E3001), Prago Mine. Leaves are spread into twos or threesnarrow small leaves arranged in verticils.

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Plate II. 1. Sphenophyllum myriophyllum Crépin (E5417), Pokrok Mine (Ovčín). Detail of cuneiform leaves spread to four narrow leaves arranged in verticils. Scale bar 10 mm.2. Sphenophyllummyriophyllum Crépin (E5634), PragoMine. Notice that leaves donot fuse at their basis. Scale bar 10mm. 3. Sphenophyllummyriophyllum Crépin (E3009),Merklín locality.General view of leaves arranged in verticils. Scale bar 10mm. 4. Sphenophyllum myriophyllum Crépin (E6496), Prago Mine. Detail of narrow lobes with hook-like apex and teeth on themargin. Scale bar 1 mm. 5. Sphenophyllum myriophyllum Crépin (E6496), Prago Mine. Detail of connection of pyriform sporangium with sporangiophore. SEM. Scale bar 500 μm.6. Sphenophyllum myriophyllum Crépin (E6496), Prago Mine. Detail of the lower part of cuneiform leaves spread to four narrow liny. SEM. Scale bar 4 mm. 7. Bowmanites myriophyllussp. nov. (E6496), Prago Mine. Detail of five pyriform sporangia at the scutelliform apex of sporangiophore. SEM. Scale bar 1 mm. 8. Sphenophyllum myriophyllum (Crépin) emend.(E6496), Prago Mine. General view on sporangium at the scutelliform apex of sporangiophore. SEM. Scale bar 500 μm. 9. Sphenophyllum myriophyllum (Crépin) emend. (E6496), PragoMine. Detail of scutelliform apex of sporangiophore. Detail of fig. 8. SEM. Scale bar 500 μm.

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Description: Stems are monopodially branched with prominentexpansions in nodes (Plate I, 5–6). Longitudinal ribs (usually runningover nodes) are part of the periderm of the stem. Ribs may sometimesalternate (Plate I, 6). The number of ribs is equal to the number of leavesin whorls. Internodes are 1–20mm long and 1–10mmwide. Cuneiformleaves are 10–40mm long and leaf lobes are 0.3–0.5mm wide (Plate I,3–4, 7). Cuneiform leaves are divided into two to four narrow lobes.

Twelve to eighteen leaves are arranged in a whorl (Plate I, 7; Plate II,1, 3). Leaves are not fused at the base (Plate II, 1–2).Epidermal structures: It was not possible to prepare upper and lowercutinised surface of leaves due to poor preservation. Epidermal structureare probably covered by inorganic crystal structures. Epidermal cells areirregularly rectangular 30–40 μm wide and 50–60 μm long. Anticlinalwalls are straight or slightly undulated.

Plate III. Bowmanites myriophyllus sp. nov. (E6324), Prago Mine. 1. Part of relatively mature specimen. Scale bar 10 mm. 2.-5. In situ spores of the Laevigatosporites–Latosporites-type.Oval specimens are of the Laevigatosporites-type and circular ones belong to the Latosporites-type. 2.–4. All × 500. SEM. Scale bar 50 μm.

Fig. 2. Reconstruction of sporangiophore with sporangia. A. Bowmanites myriophyllus sp. nov. B. Bowmanites priveticensis sp. nov.

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Plate IV. Scale bars 10 mm. 1. Bowmanites priveticensis sp. nov. (F11356holotype), PokrokMine(Ovčín). General viewon theaxis. 2. Sphenophyllumpriveticense sp. nov. (E6306), PokrokMine(Ovčín). Detail showing cuneiform leaves. 3. Sphenophyllum priveticense sp. nov. (E5414), Doubrava locality. General view of distal leafy stem. 4. Sphenophyllum priveticense sp. nov. (E6306),Pokrok Mine (Ovčín). General view of major leafy stem. 5. Association of Corynepteris angustissima (Sternberg) Němejc and B. priveticensis sp. nov. (F11355), Ovčín locality. 6. Bowmanitespriveticensis sp. nov. (E6305), Pokrok Mine (Ovčín). Sterile apex. 7. Bowmanites priveticensis sp. nov. (E6305), Pokrok Mine (Ovčín). Detail of sporangiophore with four pyriform sporangia.

Plate V.1. Sphenophyllumpriveticense sp. nov. (E6498), PokrokMine (Ovčín). Cuneiform leavesdividedby twoadditional incisions into four lobes. Scale bar 10mm.2. Sphenophyllumpriveticensesp. nov. (E65416), PokrokMine (Ovčín). Compact cuneiform leaves ondistal axis. Scale bar 10mm.3. Sphenophyxllumpriveticense sp. nov. (E6416), PokrokMine (Ovčín). Very narrow lanceolateleaves with a hook-like apex. Scale bar 10mm. 4. Sphenophyllum priveticense sp. nov. (E6499), Pokrok Mine (Ovčín). Detail of anomocytic stomata arranged in rows parallel to veins. Scale bar100 μm. 5. Sphenophyllum priveticense sp. nov. (E6499), Pokrok Mine (Ovčín). Epidermal cells of rounded rectangle shape and undulate anticlinal walls. Scale bar 100 μm. 6. Sphenophyllumpriveticense sp. nov. (E6499), Pokrok Mine (Ovčín). Detail of hook-like apex. Scale bar 500 μm 7. Sphenophyllum priveticense sp. nov. (E6499), Pokrok Mine (Ovčín). Detail of distal teeth-likeexpansions of leaf blade. Scale bar 1 μm. 8.–11. Bowmanites priveticensis sp. nov. (E6305), Pokrok Mine (Ovčín). In situ spores of the Laevigatosporites–Latosporites-type. Oval specimens are ofthe Laevigatosporites-type and circular ones belong to the Latosporites-type. Notice the length of laesurae, variable amb of spores and occasional folds of exine. All × 500.

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Sphenophyllum priveticense sp. nov.Plate IV, 2–4; Plate V, 1–7; Plate VIHolotype: Specimen E6306, National Museum, Prague.Etymology: After Přívětice, the type locality.Type locality: Přívětice (Pokrok Mine/Ovčín), Radnice Basin, CzechRepublic.Type horizon: Base of theWhetstone volcanic horizon, directly overlyingthe Lower Radnice Coal, Radnice Member, Kladno Formation, LowerBolsovian, Pennsylvanian.Material: F11355, F11356 Přívětice (PokrokMine/Ovčín),West BohemianMuseum, Pilsen; E5414, E5416, E6498, E6499, E6306 Přívětice (PokrokMine/Ovčín), National Museum, Prague, Czech Republic.Diagnosis: Sterile stems monopodially branched. Lanceolate leavesdivided into two to four lobes. Leaves divided from one-third to two-thirds of the length of leaves. Two leaf lobes can be divided into foursmaller by two narrow incisions. Frontal margin of leaves straight or

slightly convex. Convex round teeth terminated by prolongated vein.Hook-like teeth on frontalmargin of leaves on themain stems. Epidermalcells roundly rectangular in shape and anticlinal walls undulated.Anomocytic stomata arranged in rows along veins. Multicellular teethon leaf margins oriented towards frontal margin.Description: Sterile stems are monopodially branched and wider atnodes (Plate IV, 1). Longitudinal ribs extend across nodes (Plate IV, 2).The number of ribs is equal to the number of leaves in whorls (PlateIV, 2, 4). Internodes are 10–15mm long and 2–3mm wide (Plate IV, 2,4). Lanceolate leaves are divided into two to four lobes (Plate V, 1–3,7). Heterophylly is prominent (Plate V, 1–3). Leaves can be of threetypes and often grade into each other (Fig. 5). Entire leaves can bedivided by a single incision reaching one-third of the length of theleaf; resulting two lobes can be further divided by two incisions intofour additional lobes. The central incision can reach about two-thirdsof the length of the leaf (Plate V, 2). Leaves can be lanceolate andvery narrow. The frontal margin of leaves is slightly convex (Plate V,6). Rounded teeth have a prolongated vein at their apex. Leavesare 9–20mm long and up to 5–12mm wide. A single vein enters thebase of the leaf (Plate IV, 2). Teeth at the end of leaves on main stems

Fig. 3. Reconstruction of three morphotypes of leaves of Sphenophyllum priveticense sp.nov. and comparing with leaves of S. majus Bronn and S. zwickaviense Storch. A. Leaffrom distal axis of Sphenophyllum priveticense sp. nov. (E6498). Cuneiform leaves dividedby two additional incisions into four lobes with teeth with prolongated vein (a). B.Sphenophyllum priveticense sp. nov. (E6499). Compact cuneiform leaves on distal axiswith a hook-like apex of teeth (a). C. Sphenophyllum priveticense sp. nov. (E65416).Very narrow lanceolate leaves with a hook-like apex of teeth (a). D. Reconstructionof leaf of Sphenophyllum majus Bronn (from Storch, 1980). E. Part of leaf ofSphenophyllum zwickaviense Storch (from Batenburg, 1981).

Fig. 4. The reconstruction of Sphenophyllum myriophyllum Crépin showing the position ofdifferent types of leaves on the axis.

Plate VI. 1. Sphenophyllum priveticense sp. nov. (E5414), Doubrava Mine (Marie). Part of cuneiform leaves divided into four lobes. SEM. Scale bar 5 mm. 2. Sphenophyllum priveticense sp.nov. (E5414), DoubravaMine (Marie). Detail of hook-like apex. SEM. Scale bar 5 mm. 3. Sphenophyllum priveticense sp. nov. (E6499), DoubravaMine (Marie). Detail of trichome onmarginof the leaf. Scale bar 100 μm. 4. Sphenophyllum priveticense sp. nov. (E6499), Doubrava Mine (Marie). Detail of papillae on hook-like tip of the leaf. Scale bar 50 μm. 5. Sphenophyllumpriveticense sp. nov. (E6499), Doubrava Mine (Marie). Detail of papillae on the margin of the leaf. Scale bar 100 μm. 6. Sphenophyllum priveticense sp. nov. (E6499), Doubrava Mine(Marie). Detail of guard cells of anomocytic stoma. Scale bar 100 μm. 7. Sphenophyllum priveticense sp. nov. (E5414), Doubrava Mine (Marie). Detail of anomocytic stomata and undulateanticlinal walls of epidermal cells. SEM. Scale bar 200 μm 8. Sphenophyllum priveticense sp. nov. (E6499), Doubrava Mine (Marie). Detail of papillae on distal margin of the leaf. Scale bar100 μm. 9. Sphenophyllum priveticense sp. nov. (E6499), DoubravaMine (Marie). Surface of leaves are covered by small coniwhich probably represent trichome base. Scale bar 100 μm. 10.Sphenophyllum priveticense sp. nov. (E6499), Doubrava Mine (Marie). Detail of papillae on the margin of the leaf. Scale bar 500 μm.

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possess hook-like structures (Plate V, 7). The apex of lanceolate narrowleaves also have hook-like structures (Plate V, 3, 6; Plate VI, 1, 2, 4;Fig. 3. A–C). Teeth-like expansions are 0.5–1mm long on lateralmarginsof leaves. The length of teeth-like expansions is shorter close to frontalmargins (Plate VI, 4, 5, 8, 10). Lateral margins of leaves, including

teeth-like expansions and hook-like teeth on frontal margins of leavesare covered by small coni which probably represent trichome base(Plate VI, 9). Very few trichomes are preserved probably due tothe maceration process. Connected trichome was observed only once(Plate VI, 3).Epidermal structures: Upper and lower leaf surfaces are only slightlycutinised compared to strongly cutinised anticlinal walls. The shapeof epidermal cells on abaxial and adaxial surfaces of leaves differ onlyslightly. Cells on abaxial surface are irregularly rectangular, 50–60 μmwide and 125–350 μm long. Cells are narrow and prolongated aboveveins. Anticlinal walls are sinusoid and undulated (Plate V, 4, 5, 7).Irregular small circular structures probably represented trichomebases. Adaxial surface was probably covered by trichomes (Plate VI, 3,9). Intercostal cells are less prolongated on abaxial surface, but close toveins, their shape is more rectangular and prolongated (Plate V, 5).Anticlinal walls are straight or slightly sinusoid mainly close to stoma.Their shape is sinusoid and undulated above veins and more similar toadaxial surface (Plate V, 4; Plate VI, 9). Anomocytic stomata are onlyon abaxial surface and arranged in rows parallel to veins (Plate V, 4;Plate VI, 6–7). Guard-cells are parallel to leafy venation (Plate V, 5;Plate VI, 6, 7). Guard-cells of stoma are 15 μm wide and 45 μm long.Adaxial surface of leaves was probably covered by trichomes (Plate VI,9). Only trichome basis is preserved probably due to the maceration(Plate VI, 3). Teeth on leaf margins consist of several cells (Plate V, 7),and are oriented toward the apex.

4. Spores

4.1. Dispersed Laevigatosporites and Latosporites spores

We know of only few genera of Carboniferous laevigate monoletespores, which are usually referred to Laevigatosporites, Latosporites,Leioaletes Staplin, Renisporites Winslow or maybe TuberculatosporitesImgrund. All of these genera are similar, and some of them may besynonymous.

The genus Leioaletes was defined for alete circular to oval laevigatespores. However, the absence of an aperture is very questionable, andit is probable that spores previously assigned to this genus in fact maybelong to Laevigatosporites and/or Latosporites. In addition, the erectionof the genus Renisporites may be questionable and these spores alsoprobably belong to Laevigatosporites and/or Latosporites (Alpern andDoubinger, 1973).

Laevigatosporites/Latosporites spores belong to the simplestmorphological spore types and are defined for monolete thin-walledspores that usually possess a laevigate sculpture. This type of spore

Fig. 5. The reconstruction of Sphenophyllum priveticense sp. nov. showing the position ofdifferent types of leaves on the axis.

Table 2Sphenopsid plants producing Carboniferous Laevigatosporites/Latosporites spores.

Parent plants Diameter (μm) Stratigraphy References

Bowmanites bifurcatus 34–46 Asturian Andrews and Mamay (1951)Bowmanites bifurcatus 34–46 Asturian Courvoisier and Phillips (1975)Bowmanites nindelii 55–70 Permian Remy (1960)Bowmanites simonii 35–55 Permian Remy (1961)Sphenophyllum cuneifolium 37–61 Bolsovian Storch (1980)Sphenophyllum myriophyllum 30–70 Bolsovian Storch (1980)Sphenophyllum myriophyllum 32–70 Bolsovian Remy (1968)Sphenophyllum myriophyllum 45–75 Bolsovian Bek and Opluštil (1998)Sphenophyllum zwickaviense 21–64 Asturian Storch (1966 1980)Sphenophyllum kettneri 25–59 Asturian Storch (1980)Sphenophyllum oblongifolium type A 47–70 Permian Barthel (1976)Sphenophyllum oblongifolium type B 50–60 Permian Barthel (1976)Sphenophyllum oblongifolium 35–71 Stephanian Lugardon and Brousmiche–Delcambre (1994)Sphenophyllum angustifolium 40–75 Permian Barthel (1976)Sphenophyllum cuneifolium 37–61 Bolsovian Asturian Storch (1980)Sphenophyllum laciniatum 18–45 Asturian Storch (1966 1980)Tetraphyllostrobus broganensis 27–53 Cantabrian Gao and Zodrow (1990)Lilpopia crockensis 48–105 Permian Remy (1961)Leeites oblongifolius 15–30 Cantabrian Zodrow and Gao (1991)

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is referred to turma Monoletes, suprasubturma Acavatomonoletes,subturma Azonomonoletes, and infraturma Laevigatomonoletes (Smithand Butterworth, 1967). Specimens of both genera are very similar anddiffer only in the amb. The genus Laevigatosporites was proposed formostly laevigate, oval monolete spores (Ibrahim, 1933), whereas sporesof the genus Latosporites are characterised by a circular amb (Kosanke,1950). All other morphological features including the type of laesurae,sculpture and the thickness of exine are the same. The separation

of both genera based only on amb shape indicates that such puremorphological criteria need not represent natural spore taxa.

Over 40 dispersed Laevigatosporites species and over 20 dispersedLatosporites species have been described from Carboniferous sporeassemblages (Bek, 1998). Both genera and related laevigate monoletespores are not only recorded in the Carboniferous, but they have alsobeen reported in stratigraphically older (Devonian) as well as younger(Permian and Cenozoic) strata (Balme, 1995). Table 4 shows the

Table 4Selected Carboniferous species of Laevigatosporites Ibrahim (upper part) and Latosporites Potonié and Kremp (lower part) and their size ranges. Red vertical lines indicate usual size range(40–70 μm) of Laevigatosporites/Latosporites spores. At least thirteen species of Laevigatosporites Ibrahim and five species of Latosporites Potonié and Kremp fit into this size range.

Laevigatosporites majorLaevigatosporites maximusLaevigatosporites longusLaevigatosporites plicatusLaevigatosporites dunkardensisLaevigatosporites contactusLaevigatosporites vulgarisLaevigatosporites colliensisLaevigatosporites minorLaevigatosporites desmoinesensisLaevigatosporites ovalisLaevigatosporites vulgaris f. maiorLaevigatosporites cranmorensisLaevigatosporites striatusLaevigatosporites densusLaevigatosporites haardtiiLaevigatosporites mediusLaevigatosporites pennovalisLaevigatosporites minimusLaevigatosporites vulgaris f. minorLaevigatosporites scissusLaevigatosporites minimus sensu D & JLaevigatosporites perminutus

Latosporites robustusLatosporites infragranulosusLatosporites puntatusLatosorites planorbisLatosporites singularisLatosorites latusLatosporites ficoidesLatosporites falkenbergensisLatosporites saarensisLatosporites globosusLatosporites minutus

10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 µm

Table 3Epidermal structures of compression Sphenophyllum.

Species Distributionof stomata

Orientationof guard-cells

Average size ofguard-cells (μm)

Subsidiary cellsno./specializedin shape in size

Shape and arrangementof itercostal cells

Intercostalcells of size (μm)

Shape ofanticlinalwalls

References

S. priveticense In rows Parallel to veins 45 × 15 2/few Longitudinal rows 125–300/50–60 Sinuous Present studyS. myriophyllum ? ? ? ? ? 35–40/10–15? straight? Present studyS. majus Random Random 30 × 20 5–6/few Long, irregular 40–60/10–15 Coarsely sinuous Abbott (1958)S. zwickaviense ? ? ? ? Longitudinal rows 68–177/14–52 Sinuous, usually

somewhat inclinedS. emarginatum Random Usually parallel

to veins40 × 10 2/no Long, irregular 80–100/25–50 Coarsely sinuous

S. cuneifolium Randomto striped

Parallel to veins 25 × 15 2/few Long, irregular 80–200/30–60 Coarsely sinuous Barthel (1997)

S. thonii Random Parallel to veins 30 × 20 2/yes Long, irregular 100–200/20–30 Coarsely sinuous Meyen (1970)S. saarensis ? ? ? ? ? 70–140/20–40 finely sinuous Remy (1962)S. longifolium Random Parallel to veins 25 × 15 2/no Longitudinal rows 100–180/25–50 Finely sinuous Barthel (1997)S. oblongifolium ? ? ? ? Rectangular in rows 100–180/15–25 Straight Barthel (1997)S. saxonicum Random Parallel to veins 25 × 15 2/few Longitudinal rows 100–200/35–40 ?S. angustifolium ? ? ? ? Longitudinal rows 35/8 Finely sinuous Abbott (1958)S. speciosum Random Parallel to veins 25 × 15 2/no Long, irregular 100–200/20–30 Straight Pant and

Mehra (1963)

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size ranges of a selection of the most abundant Laevigatosporites andLatosporites species and their most frequent size range (red verticallines). It is clear that monolete, laevigate thin-walled spores with themost frequent size range from 40 to 70 μm can be correlated with atleast thirteen species of Laevigatosporites and five of Latosporites. Infact, monolete spores larger than 35 μm with an oval, subcircular andoval amb always occur together on a slide from one sporangium.Therefore, it is evident that they all belong to the same natural sporespecies although they are referred to two different dispersed genera.

4.2. In situ Laevigatosporites and Latosporites spores

Miospores of the Laevigatosporites–Latosporites type were describedfrom several compressed and permineralised fertile organs (Balme,1995; Bek, 1998). The smallest specimens (≤30–35μm)were producedby several marattialeans (Laveine, 1969, 1970; Balme, 1995), whereasmiospores of intermediate and larger (≥35μm)diameterwere producedby sphenopsids, mainly sphenophyllalean plants (Table 2). Monoletesof both origins (i.e., marattialean and sphenophyllalean) differ only bytheir diameters.

Sphenopsid plants mentioned in Table 2 are not the only producersof monolete Carboniferous spores, and other sources include thespecies Sentisporites goodii Riggs and Rothwell, Bowmanites fertilis(Scott) Hoskins and Cross and Peltastrobus reedae Baxter. All specimensof these species are permineralised and yielded (Baxter, 1950; Potonié,1962; Courvoisier and Phillips, 1975; Riggs and Rothwell, 1985)monolete perisporate spores with a costate/reticulate sculpture ofthe Columinisporites-type. These spores differ from those of theLaevigatosporites/Latosporites-types by having a strongly costate/reticulate perispore, i.e., outer exine layer.

5. Discussion

Sternberg (1833) illustrated a specimen of the Sphenophyllummyriophyllum-type and classified it as Volkmannia gracilis Sternberg.The species S. myriophyllum was established by Crépin (1880). Storch(1966) emended S. myriophyllum and later erected (Storch, 1980) alectotype (No. E54 in the National Museum, Prague, Czech Republic)that was originally illustrated by Sternberg (1833). The first descriptionof strobili assigned to S. myriophyllumwas published by Remy (1968).

Cones of the new species B. myriophyllus were born on vegetativestems of Sphenophyllum myriophyllum (Plate I, 1). Identical taxonomicfeatures were observed (cuticle analysis) on leaves of B. myriophyllus(including holotype) and leaves preserved on sterile stems ofS. myriophyllum including lectotype. We believe this proves that thistype of strobili belonged to the species S. myriophyllum. Both, leaves ofthe S. myriophyllum and cones of B. myriophyllus occur at the samelocality (Pokrok mine/Ovčín, Radnice Basin, Czech Republic) withinidentical stratigraphic level (i.e., the base of the Whetstone volcanichorizon, directly overlying the Lower Radnice Coal, Radnice Member).

The new species Sphenophyllum priveticensis is similar toSphenophyllum majus Bronn and S. zwickaviense Storch describedfrom the same stratigraphical level (Storch, 1966, 1980).

Division of distal edge of the leaf by central incision is characteristicfor these three species of sphenophylls. Distal leafy edge is divided intotwo lobes which are often divided into four minor lobes. It is the reasonwhy S. priveticense, S. majus and S. zwickaviensemay be confused of eachother. Reconstruction of three morphotypes of leaves of S. priveticenseand comparing with leaves of S. majus and S. zwickaviense is in Fig. 3.

S. majus differs mainly by its dimensions, with leaves are 15–20mmlong and 5–10mm wide with convex distal edges. Convexly roundedteeth on the distal leafy edge are very typical for this species. Venationis regularly divided three times within the lower, middle and upperthird of the leaf.

S. zwickaviense has leaves arranged in six whorls, leaves are up to10mm long and 5–10mm wide and generally is smaller than S. majus.

Lateral leafy margin is straight or slightly convex. Central incisionwhich divided the leaf into two lobes reaches one-third to a half ofthe length of the leaf. Lobes are often subdivided into four minor lobeswithin three-fifth to a third of the length of the leaf. Ten to twelveconcave teeth of triangular shape 0.5–1.5mm long occur on the distalmargin. Axis is covered by 1mm long trichomes.

S. priveticense has wedge-shaped leaves that are 9–15mm long and5–12 mm broad and divided into two to four lobes. Central incisiondivided leaves into two lobes within their one to two-thirds. Theselobes can be subdivided into four minor secondary lobes. Slightlyconvex frontal leafy margin is ended by convexly rounded teeth withhook-like prolongated vein. Two or three teeth occur on the lateralmargin of the lobe, i.e., eight to fourteen teeth are on the whole distalmargin. S. priveticense is characteristic by its epidermal structures.The comparison with other compression species of Sphenophyllum isgiven in Table 3 (modified from Barthel, 1997).

The genus Sphenophyllum has typically relatively slightly cutinisedupper and lower surfaces of leaves compared to the strongly cutinisedanticlinal walls. The shape of intercostal epidermal cells is irregularlypolygonal to rectangular, but cells are narrow and prolongated aboveveins. Anticlinal walls are mostly sinusoid and undulated, sometimesthey can be straight. Slightly undulated to straight anticlinal wallsoccur in S. oblongifolium (Germar and Kaulfuss) Unger (Barthel, 1997).Typical only for S. priveticense is different cell shapes on abaxial andadaxial surfaces, i.e., intercostal cells on adaxial surface are irregularlyrectangular with sinusoid and undulated anticlinal walls and lesselongated irregular cells on abaxial surface with straight anticlinalwalls. Intercostal cells of S. priveticense, especially on adaxial surfaceare the biggest within the whole genus. They are 50–60 μm wideand 125–350 μm long. The genus Sphenophyllum is characterised byanomocytic stomata that are only on the adaxial surface and theirstomatal fissure is usually parallel to the venation of the leaf or canbe occasional, as described on S. speciosum Pat and Merha. Stomatahave never been observed on Sphenophyllum oblongifolium (Batenburg,1981; Barthel, 1997). Hook-like teeth on the frontal margin of leavesare known (Batenburg, 1981; Barthel, 1997) only in S. priveticense,Sphenophyllum cuneifolium and Sphenophyllum emarginatum Brongniart.Teeth-like expansions on the lateral margins of leaves are known also onsterile bracts of S. myriophyllum (in the present study), S. angustifoliumGermar, S. emarginatum, S. cuneifolium and S. oblongifolium (Abbott,1958; Barthel, 1976, 1997; Batenburg, 1981). Trichomes and papillaeon the adaxial surface of leaves are described from S. majus, S. speciosum,Sphenophyllum saarensis, Sphenophyllum trichomatosum Stur and pro-bably occur on Sphenophyllum sewardii Batenburg. Lateral marginsof leaves, including hook-like teeth on frontal margins, and the abaxialsurface were probably covered by trichome bases. The axes of someSphenophyllum species were also covered by trichomes or emergences(S. zwickaviense, S. emarginatum, S. taylorii Bek et al., and S. trichomatosum,see Stur, 1887; Batenburg, 1981; Bek et al., 2009). Axes of S. priveticenseare smooth without trichomes. Strobili that were referred by Storch(1980) to S. zwickaviense are different from those of B. myriophyllum.Both types of strobili yielded identical laevigate monolete in situspores, but the number of sporangia per sporangiophore is different.In addition, the shape of the sterile bracts of S. zwickaviense is differentand sporangiophores do not possess the typical cuneiform expansion(see Table 1).

Bowmanites myriophyllus differs from B. priveticensis, especiallyby the shape of the sterile bracts and the number of sporangia on thescutelliform sporangiophore (Fig. 2). This type of sporangiophore isalso observed on the strobili of Sphenophyllum kettneri Storch, wherefour sporangia occur. However, the shape of the sporangia of S. kettneriidiffers from that of B. priveticensis, as does the strobili length andthe shape of the sterile bracts. The main difference between the strobiliof Sphenophyllum oblongifolium and Bowmanites bifurcatusAndrews andMamay is the dichotomous division of the sporangiophore, which doesnot possess a scutelliform expansion. S. oblongifolium has six sporangia on

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each sporangiophore while B. bifurcatus possesses only four. The shape ofthe sterile bracts and sterile apex of Bowmanites nindeliiRemy is similar tothat of B. myriophyllus, but it is not possible to compare them due to thedisintegrated character of the specimens. The strobili of Sphenophyllumzwickaviense and B. priveticensis are roughly comparable by the size butthe length of sterile bracts is different. Quadripartite sterile bracts of S.zwickaviense are 7mm long and those of B. priveticensis are 11–14mmlong. Sterile bracts of B. priveticensis are divided in two lanceolate leafylobes and are terminated by two fork teeth. But the main difference isin the shape of sporangiophore and the number and position ofsporangia on them. S. zwickaviense possesses six sterile bracts in awhorl and more than twelve sporangia. One sporangiophore bears 2–3sporangia. Expansion of sporangiophores and position of sporangia isnot mentioned in original diagnosis. B. priveticensis also possesses sixsterile bracts in a whorl, but differs by having peltate sporangiophoreswith four sporangia, i.e., twenty-four sporangia per whorl. Sporangiaof S. zwickaviense are 0.5 × 0.5mm wide and 0.5 × 1mm long whilepear-like sporangia of B. priveticensis are 1–2 mm in diameter.Sphenophyllum simonii Remy and Sphenophyllum laciniatum Storchhave sterile bracts of a different shape. The strobili of Leeites oblongifoliusZodrow andGao are very similar to those of S. oblongifolium and itmay bepossible that both strobili are synonymous because all characteristics aresimilar. Lilpopia crockensis (Remy and Remy) Conert and Schaarschmidtpossesses strobili that are morphologically very different from all otherstrobili of the genus Bowmanites, including the two species describedherein. Characteristics of the abovementioned taxa are given in Table 1.

6. Conclusions

Three new species, Bowmanites myriophyllus, B. priveticensisand Sphenophyllum priveticense are proposed and the diagnosisof S. myriophyllum is emended. New species B. priveticensis is describedusing strobili and epidermal structures. The most characteristic featureis the shape of the sporangiophores having lanceolate expansions bearingsporangia. Another difference is the shedding of sporangiophores withsporangia toward the apex of the cone that results in fertile stemsappearing similar to sterile stems because their sterile bracts are similarto apical leaves (Plate III, 1). These characteristics may suggest that it isa fertile zone and not a true strobilus. However, it is not possible toprove this hypothesis due to the limited number of strobili at differentstages of maturity.

S. myriophyllum and S. priveticense have epidermal structures thatcould enable them to climb on other plants because leaf lobes possesshook-like structures at their ends (Plate V, 6; Plate VI, 1, 2, 4). Thesehook-like structures are not only found on sterile bracts. Both specieshave serrate margins of leaves and teeth oriented toward the end ofleaves. Sterile leaves of S. myriophyllum possess a serrate keel belowthe central vein of the leaf.

The large number ofmonolete-producing sphenophylls and dispersedmonolete spores in this strata is a typical indicator for an environmentaltransitioning from plant assemblages growing at a high watertable level (dominated by arborescent lycopsids of the Lepidodendron/Lepidophloios-type) to plant assemblages growing in drier conditionsandwith a poor nutrient supply (prevalence of sub-arborescent lycopsidsof the Omphalophloios-type).

Acknowledgements

We acknowledge the financial support from the Grant Agencyof the Academy of Sciences of the Czech Republic (P210/12/2053), theResearch Program (AVOZ30130516 and RVO67985831) of theInstitute of Geology, Academy of Sciences and the NationalMuseum, Prague (MKČR DE06P040MG009) and the Researchproject of the Czech Geological Survey No. MZP0002579801, project323000. The authors thank A. Bashforth, Geological Museum,

Copenhagen, Denmark and H. Kerp, University of Münster, Germanyfor linguistic revisions and helpful notes.

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