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REVISION OF THE CONE DISCINITES FROM BOHEMIAN CARBONIFEROUS CONTINENTAL BASINS
by JIŘÍ BEK and ZBYNĚK ŠIMŮNEK
ABSTRACT. Thirty-one specimens of seven Discinites species from the Kladno-Rakovník, Radnice and Lísek basins of the central
and western Bohemian continental basins of the Czech Republic were examined. The stratigraphical position of the studied cones
is Bolsovian to Westphalian D. Emendations of the genus Discinites and species D. hlizae, D. raconicensis and D. bohemicus are
suggested. A new species D. nemejcii sp. nov. is erected. Where possible, descriptions of in situ micro- and megaspores are,
included in diagnoses and descriptions. Isolated microspores represent various stages of maturity. Immature forms have an
outermost exine layer with an irregular primary and secondary reticulum enveloping the inner bodies. Where the outer exine layer
was damaged, destroyed or dissolved, mature laevigate microspores are closely comparable with several species of the dispersed
genus Calamospora. The second type of Discinites microspores, with the inner body having a trilete mark and the outer layer with
an operculum above the proximal pole of the inner body, were isolated from Discinites bohemicus. These microspores of the
Vestispora-type have previously been recorded only from some sphenophyllalean plants, but now they are reported for the first
time from plants of non-sphenophyllalean affinity. Laevigate megaspores are comparable with the dispersed species Calamospora
laevigata.
K. FEISTMANTEL (1879) described a curious cone under the name Discinites bohemicus K. Feistmantel and
believed that it showed sphenopsid affinity. Němejc (1937, 1941) extended the genus Discinites K. Feistmantel with
the addition of several species and found that this cone belonged to fronds of the genera Palaeopteridium Kidston,
Saaropteris Hirmer and Rhacopteris Schimper. These genera were included within the artificial group
Archaeopterides. Němejc (1928) believed that this group belonged to the ferns. Specimens of cones organically
associated with fronds indicate that Discinites is a distinct plant reminiscent of the Noeggerathiales (Němejc 1937) and
also Archaeopteris Dawson (Hirmer 1940) in its features. Hirmer (1940) included Discinites in the suborder
Noeggerathineae, order Filicales, while Němejc (1950) erected a new subclass Psygmophyllineae belonging to
Pteridophyta, and included Discinites in the order Noeggerathiales. Essentially the same classification was used by
Němejc (1963, 1968), who included the Noeggerathiidae in the class Psygmophyllopsida. Zimmermann (1959)
classified these plants as a separate class Noeggerathiopsida of a separate division Noeggerathiophyta. Boureau
(1964) essentially followed the same classification. He erected a new order Discinitales for Discinites and
Saarodiscites Hirmer. Remy and Remy (1977) considered the Noeggerathiae as a taxon of a high order. They were the
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first to use the name Discinitaceae for a family belonging to the order Noeggerathiales from the subdivision
Noeggerathiophytina, division Prospermatophyta. Meyen (1987) referred the order Noeggerathiales to the class
Progymnospermopsida.
Micro- and megaspores isolated from different Discinites species by previous authors are similar to the dispersed
spore genus Calamospora Schopf, Wilson and Bentall (Table 1). All previous authors (except for Bek 1998) evaluated
Discinites fructifications only from their macrofloral point of view and did not compare isolated micro- and
megaspores with dispersed spore species. Almost all isolated spores are closely comparable with several dispersed
calamospores, differing mainly in diameter (Table 1).
MATERIAL AND METHODS
The cones described here come from the Kladno-Rakovník, Radnice and Lísek basins. The stratigraphical position
varies from Bolsovian to Westphalian D. Text-fig. 1 shows the locality inset in a general map of the Czech Republic.
All studied specimens are stored in the National Museum, Prague. The terms used for the descriptions of spores are
those given in the latest edition of the Glossary of Pollen and Spore Terminology (Punt et al., 1994). The spores are
classified according to the system for dispersed spores suggested by Potonié and Kremp (1954, 1955), Dettmann
(1963) and Smith and Butterworth (1967). In situ spores and fructifications were, if possible, compared directly with
the original diagnoses, descriptions and illustrations of the type specimens. The species determinations are based on
these original diagnoses and not on the interpretations of subsequent authors. Spores were recovered by dissolving
small portions of fructifications in nitric acid for 24-32 hours and KOH for 1-2 hours. Most spores were mounted in
glycerine jelly for direct microscopical examination, but some others were coated with gold for SEM examination
using JEOL and CAMECA SX100 scanning electron microscopes. All slides, negatives of photographs and digital
photos of spores are stored in the Institute of Geology, Academy of Sciences, Prague. All slides and negatives of
photographs of fructifications are stored in the Czech Geological Survey, Prague.
SYSTEMATIC PALAEONTOLOGY
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Division PROGYMNOSPERMOPHYTA Beck, 1960
Class NOEGGERATHIOPSIDA Zimmermann, 1959
Order NOEGGERATHIALES Darrah, 1939
Family DISCINITACEAE Remy and Remy, 1977
Genus DISCINITES (Feistmantel) emend.
Type species. Discinites bohemicus K. Feistmantel, 1879
Diagnosis: Gao and Thomas, 1995; p. 190.
Emended diagnosis. Strobilus cylindrical, wide straight axis. Sporophylls circular, bowl-shaped, attached to the axis at
right angles or decurrently to form a sheath; proximal part of sporophylls disc-like and flat. Distal parts of sporophylls
upturned with entire or toothed margins. Sporangia arranged radially on adaxial surface of proximal part of sporophyll.
A thin layer occurs between the outer sporangial wall and the mass of microspores. Trilete microspores with a circular
to oval amb. The diameter of laevigate microspores is 27-130 m. The inner exine layer is 1-2 m thick. Rays of the
trilete marks, sometimes with labrum, one-half to four-fifths of the radius. Immature microspores 71-198 m in
diameter enveloped by an outer exine layer with a strongly irregular reticulum. The primary and secondary reticula
with irregular ridges, lumina and muri. Some microspores with trilete inner body, outer exine layer with operculum
above the trilete mark of inner body. Trilete megaspores have a circular to oval amb, with a diameter of 280-1300 m.
The laevigate exine is 4-6 m thick. Rays of the trilete mark, sometimes with labrum, one-third of the radius.
Remarks. The laevigate microspores are closely comparable with several species of the dispersed genus Calamospora.
Microspores enveloped by an outer exine layer with reticulum may roughly resemble the operculate dispersed species
Pteroretis primum Felix and Burbridge and similar microspores isolated from cones of the Bowmanites dawsonii-type
(T. N. Taylor 1969, 1970; W. A. Taylor 1986; Snigirevskaya 1962), although reticulate Discinites microspores are not
operculate. Operculate laevigate microspores resemble the dispersed species Vestispora laevigata Wilson and
Venkatachala. Megaspores are closely similar to the dispersed species Calamospora laevigata (Ibrahim) Schopf,
Wilson and Bentall.
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Discinites bohemicus (Feistmantel) emend.
Plates 1, Figures 1-9; Text-Figures 2, 5g
1879
Discinites bohemicus; K. Feistmantel, pp. 298-304, fig. 1.
1937
Discinites bohemicus Feistmantel; Němejc, pp. 1-5, text-figs 1, 2a-c; pl. 1, figs 1-3 (non text-fig. 2a-c).
1956
Discinites sp. (cf. bohemicus K. Feistmantel); Remy and Remy, pp. 6-8, text-fig. 1; pls 4, figs 1-6.
1963
Discinites bohemicus Feistmantel; Němejc, p. 247, text-fig. 109a, non text-fig. 109b.
Holotype. K. Feistmantel 1879, pp. 298-304, fig. 1, Coll. National Museum, Prague, E 853 (counterpart E 854), acc. cat. 36675/57
(Plate 1, Figure 1 herein).
Type locality. Stradonice (Lísek Basin). Whetstone horizon, Radnice Group of coal seams, Radnice Member, Kladno Formation,
Bolsovian.
Material. Specimens E 853 (holotype), E 854 (counterpart) and E 856 come from the Stradonice locality. Specimen E 857 comes
from the Pejpina locality, near Beroun. Specimens E 859a-b come from the Kladno locality. Specimen E 866 comes from the
Rakovník-Příčina, ”Na Brantech” locality. All these specimens are preserved as fragments of cones.
Emended diagnosis. Axes of oblong strobili thinly longitudinally striated. Base of sporophylls decurrent forming a
very small sheath; proximal part of disc-like sporophyll at right angles to the axis. Distal parts of sporophylls dissected
into approximately 50 very small teeth. Sporangia oval, large, concentrically arranged on the adaxial surface of the
proximal part of the sporophyll. Trilete microspores with a triangular to oval amb. Rays of the trilete mark from onehalf to two-thirds of the radius. The laevigate inner exine layer 1-2 m thick. Some microspores with laevigate trilete
inner body enveloped by laevigate outer exine layer with circular operculum above the trilete mark of the inner body.
Other microspores lack this layer.
Description. The oblong strobilus possesses an acute (see Němejc 1937, Plate 1, Fig. 3) or obtuse (see Němejc 1937;
text-fig. 1 and Plate 1, Figures 4, 7 herein) apex. The obtuse form is not definite because it is unclear whether the
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entire terminal part of the cone is present (Plate 1; Figs 4, 7). The cone is more than 100 mm long and 23-25 mm wide.
The central axis is 3-4 mm wide and the separation of sporophylls is 3-4 mm. The base of the sporophylls is decurrent
forming a small sheath. The proximal part of the sporophylls rises at a right angle from the axis, the distal part of the
sporophyll is slightly upturned toward the apex of the strobilus so that each sporophyll has bowl-like shape. The edge
of the sporophyll is poorly preserved (according to Němejc 1937 with long teeth). Very short teeth are preserved on
only one specimen (Plate 1, Figure 2). The teeth are 1·0-1·5 mm long and number about 50 per sporophyll. Sporangia
(occuring only as imprints) are oval, slightly deformed due to the pressure, 2·5-3·0 mm long and 1·2-1·5 mm wide.
They are very densely concentrically arranged and number 200-300 per adaxial surface of the proximal part of each
sporophyll. Sporangia (Text-Figure 5g) were preserved on a specimen described by Remy and Remy (1956). They are
egg-shaped or oval, 2·5 mm long and 2.0 mm wide. The scars left by fallen sporangia are usually rounded and about 1
mm in diameter. The sporophyll cuticle (Text-Figure 2) is observable on the cone from the Kladno (Text-Figure 2)
locality. The cells are elongate with straight anticlinal cell walls. They are 20-35 m wide and 100-200 m long.
Trilete microspores have a circular, subcircular to oval amb and a diameter of 56·0 (100·6) 136·0 m. The rays of the
trilete mark reach mostly one-half, sometimes to two-thirds of the radius. The laevigate inner body of most
microspores is enveloped by a very thin laevigate layer with an operculum above the trilete mark of the inner body
(Plate 1, Figures 5-6), but some microspores lack this layer (Plate 1, Figures 8-9).
Remarks. Microspores were isolated only from specimen E 859b. We did not isolate any megaspores although Němejc
(1937) wrote that all Discinites cones were bisporangiate.
Botanical affinity. Palaeopteridium reussii Ettingshausen (Němejc 1937, p. 3).
Distribution. Bolsovian, Kladno Formation Radnice Member, Radnice Group of coal seams. Lísek Basin, Stradonice and Pejpina
localities; Kladno-Rakovník Basin, ”Na Brantech” near Rakovník, Příčina and Kladno localities,.
Discinites cf. bohemicus K. Feistmantel, 1879
1825
Carpolites disciformis Sternberg; Sternberg, p. 15, pl. 7, fig. 13 (Němejc, 1937; pl. 1, fig. 6).
1841
Carpolites discus Corda; Corda, p. 104, pl. 2, fig. 20 (Němejc; 1937, pl. 1, fig. 7).
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1841
Carpolites placenta Corda; Corda, p. 104, pl. 1, fig. 1 (Němejc; 1937, pl. 1, fig. 8).
1962
Discinites discus Corda; Stockmans and Willière, pp. 5-6, pl. 2, figs 1-3.
Material. Specimen E 860 comes from the Ronna Mine, Kladno-Motyčín locality. Specimens E 861 and E 866 come from the
Břasy locality, near Radnice. Specimens E 862 (Carpolites disciformis Sternberg), E 863 (C. discus Corda) and E 864 (C. placenta
Corda) come from the Chomle locality, near Radnice (all these specimens were named as Discinites sp. (cf. bohemicus) by Němejc
1937). All are preserved as isolated sporophylls. Specimens E 867-868 come from the Bílá Hora locality, near Plzeň.
Description. All these specimens are disc-like sporophylls. They are usually flat with a hole or the remains of a nodus in the centre.
They are 22-28 mm in diameter. Carpolites placenta bears faint imprints of sporangia. C. disciformis and C. discus have the same
diameter as Discinites bohemicus. Furthermore, there are many other isolated sporophylls of comparable size but it is dificult to
establish the identity of these specimens without knowledge of sporangial and cone shape.
Remarks. Carpolites disciformis and C. discus are similar to Discinites bohemicus. Very similar specimens were
described by Stockmans and Willière (1962; pl. 2, figs 1-3) from the Carboniferous of Belgium. However, the
Bohemian specimens are bigger and do not usually possess the fine radial striation visible on Belgian specimens. On
the other hand Discinites sp. sensu Remy and Remy resembles Belgian specimens in its width. It is difficult to refer
this specimen with certainty to D. bohemicus because it is narrower and stratigraphically older than all the Bohemian
specimens. The German specimen comes from the upper Langsettian, whereas the parent plant of D. bohemicus is
Palaeopteridium reussii Ettinghausen occuring only within the Duckmantian and Bolsovian in the Czech Republic .
Distribution. Bolsovian, Kladno Formation, Radnice Member, Radnice Basin, Chomle, Břasy and Radnice localities. Plzeň Basin,
Whetstone horizon, Bílá Hora locality and, Kladno-Rakovník Basin, Kladno locality (according to Němejc 1937).
Discinites major Němejc, 1937
Plate 1, Figure 10; Plate 2; Plate 3, Figure 1-4; Text-Figs 3-4, 5e-f
1937
Discinites major Němejc; Němejc, p. 1-8, text-figs 3-7; pl. 1, figs 4-5, 9-11.
1937
Discinites bohemicus Němejc; Němejc, text-figs 2a-c, non text-fig. 1; pl. 1, figs 1-3.
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1963
Discinites bohemicus Feistmantel; Němejc, p. 247, text-fig. 109b, non text-fig. 109a.
Holotype. Němejc 1937, pl. 1, fig. 4. Coll. National Museum Prague (E 869). Acc. cat. P 25135.
Type locality. Lubná, near Rakovník, Rako Mine in Krčelák (Kladno-Rakovník Basin).
Material. Specimens E 858a-858b come from the Pejpina locality, near Beroun. Specimens E 869 (holotype), E 870-871 come
from Rako Mine, Lubná, near Rakovník.
Description. Strobilus at least 90 mm long, but the base is missing. Its maximum width is 35 mm (Plate 2, figure 1). The strobilus
tapers slightly towards the acute apex. Axis is about 4 mm wide. Sporophylls are about 3-4 mm apart. Sporophyll connected to the
axis at a right angle upturned distally. Sporophylls possess a bowl-like shape. The distal edge of the bowl-shaped sporophylls is not
preserved, but it is believed to be undivided. The oval shaped sporangia (Text-Figure 5e-f, Plate 1, Figure 10, Plate 2, Figures 5, 6)
taper slightly towards the apex and have a relatively wide base. They are 2 ·8-3·8 mm long and 13-20 mm wide. All of these large
sporangia are microsporangia with microspores (Plate 1, Figure 10,Plate 2, Figure 2). Megasporangia with only one megaspore
(according to Němejc 1937; text-fig. 5a-g, 6a-c) are smaller than microsporangia, are more rounded and 15-20 mm long. They
occur in the basal half of the cone, are dispersed very rarely among the microsporangia, and become slightly more frequent towards
to the base of the cone. Exceptionally, scars of fallen sporangia are seen on the holotype. They are oval, 08-10 mm long and 250300 m wide. However, these scars are better seen on isolated sporophylls (Němejc 1937; figs 2a-c; pl. 1, fig. 5). The scars are
very prominent and they are submerged under the surface of sporophylls, so that they form scars. These scars are oriented radially
towards the edge of the sporophylls and are 20-25 mm long and 05-08 mm wide infrequently preserved. The outer wall of the
microsporangia (Text-Figure 4, Plate 2, Figure 6) is formed by rather large cells (Němejc 1937). These cells are long, spindleshaped, and oriented parallel to the longitudinal axis of the sporangia. They are 30-50 m wide and 300-600 m long. The shortest
cells, 60-120 m long, are arround the apex of the sporangium. An additional thin inner wall is also occasionally preserved. Its
cells are much smaller, 20-30m wide and 80-160 m long. The sporophyll cuticle (Text-Figure 3a, Plate 2, Figure 3) is preserved
in the basal part of one cone. Since no scars are visible, it is considered to be abaxial. Cells are large, and tetragonal in shape. They
are 35-70 m wide and 200-600 m long. Similar structures are visible under the SEM. Trilete microspores have a circular to oval
amb. Their diameter is 770 (1060) 1410 m. The laevigate exine is 1-2 m thick. Rays of the trilete mark (usually with a labrum)
are 2-4 m wide and high and reach one-half to four-fifths of the radius. Secondary folds of the exine are of irregular shape, size,
number and position. Microspores in clusters are mostly enveloped in a very thin layer (Plate 2, Figure 4). A similar layer occuring
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between the outer sporangial wall and the mass of microspores is also reported in petrified sporangia of the type specimen of
Discinites sinensis Wang (Wang 2000). It may represent some innermost layer of sporangial wall.
Remark. Microspores isolated from specimens E 869 (holotype)-871 are closely comparable with several species of
the dispersed genus Calamospora. All the microspores are similar differing mainly (except for the character of
secondary folds of the exine) in the diameter (the difference of averages of in situ microspores isolated from three
specimens is only 17 m). The diameter of the microspores (77-141 m) is greater than that (95-115 m) given by
Němejc (1937). We did not assign these in situ microspores of Calamospora-type to any dispersed species. The
classification of dispersed calamospores is based on differences in diameter shape, length of laesurae, presence of a
labrum and contact area, and the size, position, number and shape of secondary folding of the exine. It is possible to
compare isolated microspores with several dispersed calamospores (Bek 1998) under these constraints but this is of no
relevance because we cannot distinguish calamospores isolated from parent fructifications of equisetalean,
sphenophyllalean or progymnosperm affinities. Megaspores were not extracted, although Němejc (1937) described
them.
Botanical affinity. Palaeopteridium macrophyllum Němejc (Němejc 1937; p. 4).
Distribution. Bolsovian, Kladno Formation, Radnice Member, Lubná Group of coal seams, Kladno-Rakovník Basin: Rako Mine,
Krčelák, Lubná near Rakovník localities; Radnice Member of the Plzeň Basin: Krimich Mine, Nýřany locality, (West-Bohemian
Museum, Plzeň); (Radnice Member of the Lísek Basin: Pejpina locality).
Discinites raconicensis Němejc, 1941
Plate 3, Figures 5-6; Text-Figure 5h
1941
Discinites raconicensis Němejc; Němejc, pp. 8-9, text-fig. 3.
Holotype. Discinites raconicensis Němejc, 1941, text-fig. 3, Coll. National Museum, Prague, E 876, acc. cat. 9268.
Type locality. Kladno-Rakovník Basin, Moric Mine, near Rakovník.
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Stratigraphical position. Roof shales of the coal seam called ”Věnec”, Nýřany Member, Kladno Formation, Westphalian D.
Material. Specimen E 876, Moric Mine, near Rakovník.
Description. Oblong strobilus over 210 mm long and 46 mm wide with an acute apex (Plate 3, Figure 6). The internal
structure of the cone is not known. Each sporophyll is bowl-shaped with several hundred sporangia on its adaxial
surface. Margin morphology is not known. Sporophylls are about 5 mm apart near the apex of the cone and 8-11 mm
apart in the middle part of the cone. Sporangia (Text-Figure 5h) are very densely distributed and slightly deformed, so
that they have a hexagonal outline (Plate 3, Figure 5). They are about 25-30 mm long and 20-25 mm wide. These
sporangia are preserved only as imprints.
Botanical affinity. Rhacopteris asplenites Gutbier (Němejc 1941; p. 9).
Distribution. Only one cone has been found in the Kladno Formation, Nýřany Member, Westphalian D, the Kladno-Rakovník
Basin, near Rakovník, Moric Mine.
Discinites cf. raconicensis Němejc, 1941
Plate 4; Text-Figure 5d
1941
Discinites sp. 2; Němejc 1941, pp. 6-8, text-fig. 2.
1998
Discinites sp. A; Bek, pp. 176-179; pl. 141, figs 1-11; pl. 141, figs 1-9.
Material. Specimens E 3610-3613 and E 3617, 1st May Mine locality, Lubná, near Rakovník, specimen E 874, Vejvanov locality,
near Radnice.
Description: The largest cone fragment of this species is about 105 mm long and 60-70 mm wide (Plate 4, Figure 1, 2). The
separation of the bowl-like sporophylls is about 60-70 mm. 1000-1400 sporangia are densely spaced on the adaxial surface of
each sporophyll. The margin of the sporophylls is not preserved. Sporangia (Text-Figure 5d) are relatively small, 24-30 mm long
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and 15-22 mm wide, usually slightly elongated and tetragonal with rounded corners. They can be hexagonal or rounded in shape
presumably due to preservational differences (Plate 4, Figures 2, 8). Surfaces of sporangia have several long narrow longitudinally
oriented cells. Another fragment of a relatively wide oblong cone, 160 mm long and 45 mm wide, is preserved in a burnt shale of
the Upper Radnice coal seam and where all the organic coalified matter is damaged. Sporangia are preserved only as hexagonal
imprints. The size of sporangia is 3-4 mm long and 12-18 mm wide. The sporophyll margin, cellular structure, and sporangium
structure are not known. It is difficult to compare specimens from the Lužná locality with the specimen from Vejvanov locality due
to a lack of diagnostic features. Trilete microspores have a circular to oval amb. Their diameter is 410 (810) 1530 m. Rays of
the trilete mark, usually with a labrum about 2-4 m wide and high, reach from one-half to two-thirds of the radius. The laevigate
inner exine layer is 1-2 m thick. Laevigate microspores (Plate 4, Figures 4, 7, 9-10), 41-78 m in diameter, are closely
comparable with several species of the dispersed genus Calamospora. Some microspores are enclosed in an outer layer ranging
with a strongly irregular reticulum (Plate 4, Figure 3). The surface of this outer layer possesses a secondary reticulum within the
primary.Various intermediate forms (Plate 4, Figures 5-6, 11-12) can be recognized. Many microspores of the Calamospora-type
possess various fragments of the outer layer on their surface.
Remarks. Four specimens were recovered from a single locality and stratigraphical horizon. It is not clear if they are fragments of
one larger cone. These cone fragments were collected by Drábek in 1976 and assigned to Discinites cf. raconicensis. They
probably represent fragments of larger cones about 300-400 mm long. They closely resemble Němejc´s D. raconicensis in their
external features including the dimension and shape of sporangia. However, the problem is the stratigraphical positions of both
specimens. Němejc (1941) claimed that D. raconicensis belongs to the fronds called Rhacopteris asplenites restricted to the
Westphalian D strata, whereas the specimen described here comes from the lower Bolsovian strata. For this taxon, Discinites cf.
raconicensis most probably belongs to a different species of Rhacopteris. Table 2 shows the sizes of microspores isolated from five
specimens of Discinites cf. raconicensis.
Distribution. Kladno Formation, Radnice Member in Kladno-Rakovník Basin, Lubná, near Rakovník, 1st May Mine, intercalation
of the Upper Radnice coal seam; spoil piles of the David Mine, Vejvanov locality in the Radnice Basin, intercalation of the Upper
Radnice coal seam, Radnice Member, Kladno Formation, Bolsovian.
Botanical affinity. Rhacopteris elegans Ettingshausen and/or R. speciosa Ettingshausen. Because no organic connection was
obsereved, the association remains questionable.
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Discinites hlizae (Němejc) emend.
Plates 5-8; Text-figure 5a-c
1941 Discinites hlizae Němejc; Němejc, pp. 4-6, pl. 1, figs 1-3; pl. 2, fig. 1a-c.
1998 Discinites hlizae Němejc; Bek, pp. 173-176, pl. 137, figs 1-5; pl. 138, figs 1-8; pl. 139, figs 1-13;
pl. 140, figs 1-9; pl. 150, fig. 5.
Holotype. Němejc 1941, pl. 1, fig. 1, coll. National Museum, Prague, E 872, acc. cat. 25438.
Type locality. Rako Mine, ”Krčelák”, Lubná, near Rakovník, Kladno-Rakovník Basin.
Stratigraphical position. Kaolinic coarse tuff under the lower Lubná coal seam. Lubná Group of coal seams, Radnice Member,
Kladno Formation, upper Bolsovian.
Material. Specimen E 872, Rako Mine, ”Krčelák” locality, Lubná, near Rakovník .
Emended diagnosis. Oblong cones, disc-like sporophylls with oval or circular sporangia (micro- and megasporangia).
Elongated, spindle-shaped or irregularly tetragonal cells form the outer wall of sporangia. Trilete laevigate
microspores with a circular to triangular or oval amb. Rays of the trilete mark, often with a labrum from one-half to
two-thirds of the radius. Some microspores enclosed in an outer exine layer with a strongly irregular primary and
secondary reticulum. Trilete laevigate megaspores with a circular to oval amb. Rays of the trilete mark, with a labrum
about one-third of the radius.
Description. Oblong cones 80-115 mm long and 25-40 mm wide (Plate 5, Figure 1, Plate 6, Figure 1). The separation of
sporophylls is 4-5 mm. The shape of sporophylls is disc-like. Sporangia (Text-Figure 5a-c, Plate 5, Figure 3; Plate 6, Figure 5) and
their imprints (Plate 6, Figure 9) are very well preserved. Therefore the description of the cones is emended to include description
of sporangia and the spore contents. Sporangia are of two types: relatively smaller rounded sporangia are probably megasporangia.
They are 15-20 mm in diameter and each contain 16 megaspores (according to Němejc 1941). Oval sporangia are microsporangia.
They are 20-25 mm long and 14-20 mm wide. The cellular structure is clearly visible on the outer wall of the sporangia. The
cells are elongated, spindle-shaped or irregularly tetragonal, and oriented parallel to the longitudinal axis of the sporangia. Their
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width is 10-30 m and length 80-300 m. Anticlinal cell walls are straight or slightly curved. Trilete microspores have a circular to
oval, rarely triangular amb. The diameter is 400 (906) 1980 m. The laevigate inner layer is 1-2 m thick. Rays of the trilete
mark, often with a labrum, are 2-4 m wide and high. The prominent primary reticulum consists of irregularly distributed ridges
and lumina (Plate 7, Figures 1-5; Plate 8, Figures 1-5). The secondary reticulum possesses irregular ridges, muri and lumina (Plate
8, Figures 1-2). All intermediate forms (Plate 7, Figures 6-9; Plate 8, Figures 1-6) between the laevigate microspores (Plate 7,
Figures 10-11) and forms with strongly reticulate outer layer occur. Laevigate microspores and microspores with various fragments
of the outer layer prevail over the forms with a fully developed outer layer, which comprises only about one per cent of all
described specimens. The smallest laevigate microspores (about 40 m large; Plate 5, Figures 2, 4-6) have a labrum, triangular amb
and the rays of the trilete mark reach four-fifths of the radius or more. Some of the spores may resemble species of the dispersed
genus Leiotriletes (Naumova) Potonié and Kremp and they are interpreted as representing very immature microspores. The
diameter of megaspores is 5100 (63615) 8230 m. The laevigate exine is 4-6 m thick. The rays of the trilete mark, with labrum
about 25 m high and 15 m wide, reach about one-third of the spore radius. These megaspores (Plate 6, Figures 7-8) are classified
as Calamospora cf. laevigata due to the labrum, that is not mentioned in the original diagnosis of C. laevigata (Ibrahim, 1933, pp.
17-18).
Remarks. Němejc (1941) provided a detailed description of a relatively long (450 mm) twig bearing cones - seven
complete and one incomplete. Some cones were deformed during fossilisation. The twig is 35 mm in width and its
surface is smooth, or bears irregular longitudinal cracks. The megaspores size range (510-823 m) is wider than that
(280-770 m) given by Němejc (1941).
Distribution. see ”Type locality” and ”Stratigraphical position”.
Botanical affinity. Němejc (1941) found many pinnules of Saaropteris guthoerlii Hirmer on the same slab with
Discinites hlizae. However, he was very careful in assessing the affinity of these two species, because Hirmer (1941)
had previously assigned the cone of Saarodiscites guthoerlii Hirmer to the species Saaropteris guthoerlii. The cone
Saarodiscites is very similar to that of Discinites. Hirmer (1941) contended that the cones of Saaropteris had
sporangia on the abaxial side of the sporophylls. Unfortunately all sporangia had fallen off and only their imprints are
visible on the cones (see Hirmer 1941, pls 6-7). It is evident from his figures (pls 6-7, figs 1ab and 2aa) that the tissue
of the sporophylls was originally thin and that imprints of sporangia could also be preserved on the abaxial side under
favourable fossilisation conditions. Of course, this must be demonstrated by study of the original specimens. The
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Saarodiscites cones are bigger (45-50 mm in width) than those of Discinites hlizae. Although the cones of Saaropteris
guthoerlii and Discinites hlizae are very similar, it is very difficult to establish their identity. We retain Hirmer`s and
Němejc`s specimens as different species until the revision of Saaropteris guthoerlii from the Saar Basin is completed.
Discinites nemejcii sp. nov.
Plate 9, Text-Figure 6
1941
Discinites sp. 3; Němejc, pp. 9-11, text-fig. 4, pl. 2, figs 3a-c.
1941
Discinites sp. 1; Němejc; p. 6, text-fig. 1; pl. 2, figs 2a-c.
Material. Holotype: Specimens E 875a-b come from Malé Přílepy locality, Malé Přílepy Basin (Plate 9, Figures 1-2 herein).
Paratype. Specimen E 873, Kladno locality.
Type locality. Malé Přílepy in the Malé Přílepy Basin. Whetstone horizon, Radnice coal seam, Radnice Member, Kladno
Formation, Bolsovian.
Stratigraphical horizon. Whetstone horizon, Radnice coal seam, Radnice Member, Kladno Formation, Bolsovian.
Derivation of name. In honour of Prof. František Němejc, a famous Czech palaeobotanist, who also studied Discinites cones.
Diagnosis. Relatively flat disc-like sporophylls. The proximal part of the sporophylls is oriented at right angles to the
axis. Sporangia oval, nearly circular to hexagonal (in compressed form), radially arranged on the adaxial surface of
proximal parts of sporophylls. Trilete microspores with a circular to oval amb. The sculpture of the inner exine layer
laevigate or irregularly sculptured with various fragments of the outer layer. Rays of the trilete mark, from one-half to
two-thirds of the spore radius. Some microspores possess an outer reticulate exine layer enveloping the inner body.
The outer body is sculptured with a primary and secondary reticulum with irregular ridges, muri and lumina. Trilete
laevigate megaspores with a circular to subcircular amb. The rays of the trilete mark, about one-third of the radius.
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Description. The holotype of this species is a fragment of an oblong strobilus (Plate 9, Figures 1, 2) lacking base and apex (92 mm
in length of 38 mm in width). The sporophyll is disc-like. Its margin is not known because it is broken. Sporangia (Text-figure 6)
are radially arranged on the adaxial surface of the sporophyll. They are relatively small, oval to nearly circular or hexagonal (due to
compression). Their length is 15-25 mm and width 10 to 18 mm. According to Němejc (1941), each megasporangium contains
16 megaspores. The surface of the sporangium (outer wall) is flat or bears very fine longitudinal striations. It is very difficult to
distinguish mega- and microsporangia. Megasporangia are probably more circular. The diameter of the microspores is 450 (739)
990 m. The inner layer is 1-2 m thick. The sculpture of the exine is mostly laevigate or irregularly sculptured with various
fragments of an outer layer. The diameter of the forms with an outer layer is 710 (908) 990 m. The outer layer is preserved as an
irregular layer or as fragments when the original sculpture was badly damaged during fossilisation (Plate 9, Figures 5, 7). The
diameter of the megaspores is 5560 (7270) 8250 m. The laevigate exine is 4-6 m thick.
Remarks. Němejc (1941) suspected that the sporophyll (specimen E 873) could belong to Discinites hlizae based on its
dimension and the number of megaspores (16) in the megasporangia. But without knowledge of the whole shape of the
cone, he was unable to make a precise determination. Isolated microspores are closely comparable with several
dispersed species of Calamospora and isolated megaspores (Plate 9, Figure 4) are closely comparable to the dispersed
species C. laevigata.
Distribution. Specimen E 873 was found in the Kladno-Rakovník Basin, Kladno locality (mines in its surroundings); intercalation
”Velká opuka” in the Upper Radnice coal seam, Radnice Member, Kladno Formation, Bolsovian, Carboniferous. Specimens E
875a-b, acc. cat. 3490 have been found in the Malé Přílepy Basin, Malé Přílepy locality, Whetstone horizon, Radnice coal seam,
Radnice Member, Kladno Formation, Bolsovian.
Botanical affinity. Unknown.
Remark on Němejc‘s fertile specimen of Rhacopteris bipinnata Němejc
Němejc (1941; pp 11-12, text-fig. 5) described the basal part of a Discinites cone in organic association with a frond
of Rhacopteris bipinnata. Unfortunately, this is only a very small piece of compressed cone. The sporophylls are very
poorly preserved, only clusters of very densely spaced sporangia typical of Discinites cones are visible. The preserved
base of the cone is 25 mm wide. Our knowledge regarding Bohemian Discinites cones does not allow us to decide to
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which Discinites species this very interesting fructification belongs. Table 3 shows probable botanical affinities of
Bohemian Discinites.
DISCUSSION
Several species and unassigned specimens of Discinites have been described from the Carboniferous of the
Bohemian Massif. The characteristics of those Bohemian cones are listed in Table 4.
Fourteen Discinites species have been described to date. Five species [D. dentilongus Gao and Thomas, D. ?
fimbriata Wang and Wang, D. orientalis Gu and Zhi (Lee, Chow and Deng), D. sunjiaquouensis Wang and Wang and
D. sinensis Wang] were described from the Permian of China. Even though their characteristics are similar to the
European Discinites, we consider them as distinct species living at a different time in a different floristic province than
the European species. Besides the Bohemian specimens, four species of Discinites have been described from Europe
and the USA. D. discus (Corda) Stockmans and Willière was established by Corda (1841) as a seed Carpolites discus.
Later Němejc (1937) recognized that this is a sporophyll of Discinites but without knowledge of the sporangia and fine
structures, he was not able to assign this sporophyll to an exact Discinites species. The specimens described by
Stockmans and Willière (1962) are smaller than the Bohemian specimens, they are only 18 to 20 mm in diameter and
come from the Duckmantian and Bolsovian of Belgium. Despite this, they may belong to D. bohemicus because the
parent plant of D. bohemicus is Palaeopteridium reussii that was also found in the Duckmantian, but it is difficult to
classify them without knowledge of sporangial and cone shapes. Discinites egregius (Grand`Eury) Boureau was
originally described under the name Macrostachya egregia Grand`Eury from the Stephanian of the Loire Basin in
France. This cone is 30 mm wide, with sporophyll margins that are entire or very finely lobed. It is different from the
Bohemian specimens because the sporophyll margins of the Bohemian specimens are usually broken. Discinites
jongmansii Hirmer described by Hirmer (1941) from the Bolsovian in the Saar Basin is a very distinct species and
differs from all Bohemian Discinites in having long teeth on the margins of sporophylls (a sporophyll of this cone was
recently found in the Intra-Sudetic Basin by Šimůnek, pers. comm.). Probably the oldest known species of Discinites is
D. delectus (Arnold) Arnold originally described under the name Bowmanites delectus Arnold from the lower
Pennsylvanian of the Pottsville Formation of the Michigan Coal Basin. It is a relatively narrow cone, only 17 mm
wide, which is even less than the narrowest Discinites bohemicus from the Bohemian Massif. Isolated micro- and
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megaspores of the Calamospora-type are similar to laevigate micro- and megaspores isolated from most Bohemian
Discinites.
Gao and Thomas (1995) compared Discinites with other related genera: Lacoea Read, Noeggerathiaestrobus
Feistmantel, Saarodiscites Hirmer, Heninia Stockmans and Willière and Tongshonia Stockmans and Mathieu. Heninia
(H. belgica Stockmans and Willière) is similar to Discinites (especially D. discus). The difference between Discinites
and Saarodiscites is in the presence of sporangium traces on the abaxial surface of sporophylls of the latter genus. This
may be a feature of preservation. This hypothesis must be verified on Hirmer`s original specimens before the transfer
of Saarodiscites into Discinites. According to Gao and Thomas (1995), Tongshonia seems to be identical with
Discinites. On the other hand Leary (1973) considered Tongshonia to be identical with Lacoea. Lacoea and
Noeggerathiaeostrobus differ from Discinites in having semi-circular sporophylls.
Three degrees of maturity are observed in Bohemian in situ Discinites microspores. The first degree of maturity
(relatively immature microspores) is represented by microspores of the Calamospora-type enveloped by an outer exine
layer with irregular primary and secondary reticulum (Plate 7, Figures 1-5; Plate 8, Figures 1-5). These microspores
are unique and do not resemble any known dispersed spore species. Pteroretis primum is the only species of very
roughly similar morphology (but it is operculate). The second degree of maturity (relatively mature microspores) is
represented by microspores of the Calamospora-type with various fragments of the outer layer still adhering (Plate 4,
Figures 5-6, 11-12; Plate 7, Figures 6-9; Plate 8, Figure 6; Plate 9, Figures 5-7). These microspores cannot be
attributed to any known dispersed miospore species, but are closely similar to microspores isolated from various
equisetalean fructifications by Bek (1998) and Bek and Opluštil (1998). The third degree of maturity (mature
microspores) is represented by laevigate microspores (Plate 1, Figures 8-9; Plate 2, Figure 7; Plate 3, Figures 2-4;
Plate 4, Figures 4, 7, 9-10; Plate 6, Figures 2-4, 6; Plate 7, Figure 10-11) closely comparable with several
calamospores and laevigate operculate microspores of the Vestispora laevigata-type (Plate 1, Figures 5-6).
Four types of microspores of Bohemian Discinites occur. The first type is represented by relatively large operculate
microspores (about 100 m on average) of the Vestispora-type (Plate 1, Figures 5-6) isolated from Discinites
bohemicus. The second type (Plate 2, Figure 7; Plate 3, Figures 2-4) is characterized by laevigate (about 106 m on
average) microspores (isolated from D. major) with a prominent labrum (they are clearly distinguishable based on this
feature). Microspores of the third type (about 90 m on average) were isolated from D. hlizae (Plate 6, Figures 2-4).
They do not possess as prominent a labrum and contact area. Relatively immature microspores with a reticulate outer
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layer and its fragments are well preserved. Microspores of the fourth type (Plate 4, Figures 4-7, 9-12; Plate 9, Figures
5-7) isolated from Discinites cf. raconicensis and D. nemejcii are smaller (about 77 m on average), without a
prominent labrum. The outer layer is more or less destroyed. All Discinites cones were bisporangiate with micro- and
megaspores of the Calamospora-type although we only isolated megaspores from some of them (Plate 6, Figures 7-8;
Plate 9, Figure 4).
Most genera of Carboniferous cones produced one morphological type of microspore comparable with one or
several dispersed species of one or even more genera (Balme 1995, Bek 1998). Discinites cones produced three
morphologically different types of microspores belonging to three different dispersed genera, which is very strange.
Microspores isolated from Discinites bohemicus are the first record of microspores of the Vestispora-type from nonsphenophyllalean plants.
The first petrified Discinites cones, named D. sinensis from Lower Permian of China (Wang 2000), produced
unusual microspores. These microspores have a laevigate trilete inner body (probably endospore) and laevigate middle
layer (probably exospore) with proximal trilete mark with prominent labrum, as well as a circular operculum on the
distal surface. Non-macerated microspores have a scabrate outermost exine layer (perispore). This new type of
microspore (never described and representing a new spore morphotype) has been named Discinispora sinensis Wang
et al. The presence of two types of germination apparatus (proximal trilete mark and distal operculum) (indiatives the
still uncertain taxonomic position of Discinites. Detailed study of Chinese and Bohemian Discinites may lead to a
better understanding of the role these plants played in the evolution of spore and pollen-producing plants.
Acknowledgements. This research was supported in part by the Grant Agency of the Academy of Sciences of the Czech Republic
(grant no. A3013902/1999) and by a Research project of the Institute of Geology (CEZ, Z3013912). The authors are grateful to A.
Gabašová from the Czech Geological Survey, Prague and J. Pšenička from the West-Bohemian Museum, Plzeň for some
photographic help and Prof. B.A. Thomas from the University of Aberystwyth, UK and W.A. Taylor from the University of
Wisconsin, USA for correction of the text.
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— 1941. Noeggerathia, neuendeckte Verwandte Formen und ihre Stellung im System der Farne. Biological
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NĚMEJC, F. 1928. A revision of the Carboniferous and Permian flora of the coal-districts in Central Bohemia.
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— 1937. On Discinites K. Feistmantel. International Bulletin Academie de Science de Bohéme, 1-7.
— 1941. Further indications of the type Discintes in addition to some remarks on the Archaeopteriden of the
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JIŘÍ BEK
Institute of Geology
Academy of Sciences
Rozvojová 135, 165 00 Prague 6
Czech Republic
e-mail: mrbean@gli.cas.cz
ZBYNĚK ŠIMŮNEK
Czech Geological Survey
Klárov 3, 118 21 Prague 1
Czech Republic
e-mail: simunek@cgu.cz
TEXT-FIGURE 1. A map with localities of Discinites species. 1 - Carboniferous sediments exposed and covered by
younger sediments (mainly Cretaceous), 2 - Discinites numbered localities: Kladno-Rakovník Basin - Kladno (D.
bohemicus, D. nemejcii), Rakovník (D. bohemicus, D. raconicensis), 1 - Příčina (D. bohemicus), 2 - Lubná near
Rakovník (D. hlizae, D. major, D. cf. raconicensis); Radnice Basin - 3 - Břasy near Radnice (D. bohemicus), 4 Chomle near Radnice (D. bohemicus), 5 - Vejvanov , (D. cf. raconicensis); Plzeň Basin - 6 Bílá Hora near Plzeň (D.
bohemicus), 7 - Třemošná (D. bohemicus), 8 - Nýřany (D. major); Lísek Basin - 9 Pejpina (D. bohemicus), 10 Stradonice (D. bohemicus); Carboniferous near Malé Přílepy - 11 - Malé Přílepy (D. nemejcii).
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TEXT-FIGURE 2. Discinites bohemicus (K. Feistmantel) emend. (E 859b). Cuticle of sporophyll;  50.
TEXT-FIGURE 3. Discinites major Němejc, 1937 (E 869). a) cuticle of sporophyll;  50. b) surface of
megasporangium;  60.
TEXT-FIGURE 4. Discinites major Němejc, 1937 (E 869). Surface of microsporangium;  30.
TEXT-FIGURE 5. Shapes of microsporangia. All  15. a-c) Discinites hlizae (Němejc) emend. d) D. cf. raconicensis
(Němejc). e-f) D. major Němejc, 1937. g) D. bohemicus (K. Feistmantel) emend. h) D. raconicensis (Němejc) emend.
TEXT-FIGURE 6. Discinites nemejcii sp. nov.
Detail of sporangia from specimen figured on Pl. 9, Fig. 2. Notice different dimension of sporangia caused either by
compression or by different shape of micro- and megasporangia;  25.
EXPLANATION OF PLATES
PLATE 1
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Discinites bohemicus (K. Feistmantel) emend. All figured specimens come from the Kladno Formation, Radnice
Member, Bolsovian.
1.
Holotype (E 853), fragment of a cone, Stradonice locality (Lísek Basin), Radnice group of coals. Coll. K.
Feistmantel;  1.
2.
Fragment of a cone (E 866) with very small teeth preserved on the edge of sporophylls. Lubná, near Rakovník,
”Na Brantech” locality, roofs of 1b Coal Seam, Lubná group of coals;  2.
3.
Counterpart (E 854) of holotype;  1.
4, 7. Terminal part of cone (E 859a-b), Kladno locality, ”Velká opuka” in the upper Radnice coal seam;  2.
5-6. Operculate microspores isolated from specimen E 859b and comparable with dispersed species Vestispora
laevigata; All  500.
8, 9. Isolated inner bodies of microspores with trilete mark from specimen E859b and correlated with dispersed genus
Calamospora. All  500.
10. Discinites major Němejc, 1937. Detail of holotype (E869), Lubná near Rakovník, Rako Mine, Krčelák locality,
Kladno-Rakovník Basin, Lubná group of coals, Radnice Member, Kladno Formation, Upper Bolsovian.
Microsporangium with partly damaged sporangium wall containing microspores;  25.
PLATE 2
1. Discinites major Němejc, 1937. Holotype (E 869), Lubná , near Rakovník, Rako Mine, Krčelák, KladnoRakovník Basin, Lubná group of coals, Radnice Member, Kladno Formation, Upper Bolsovian.  1.
2. Detail of Fig. 1 shows microsporangium with partly damaged sporangium wall containing microspores;  50.
3.
Cuticle of sporophyll;  50.
4.
Edge of sporangium (E870) showing common layer that envelopes microspores;  500.
5.
Microsporangium;  25,
6.
Imprint of sporangium with preserved longitudinally oriented cells;  25.
7.
Microspore of the Calamospora-type isolated from specimen E 869. Note prominent labrum;  500.
8.
Cuticle of sporophyll showing longitudinally oriented cells (isolated from specimen E 870), SEM;  100.
PLATE 3
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23
1. Discinites major Němejc, 1937. Topotype (E 870), Lubná, near Rakovník, Rako Mine, Krčelák, KladnoRakovník Basin, Lubná group of coals, Radnice Member, Kladno Formation, Upper Bolsovian. Part of cone
showing sporangia;  10.
2-4. Microspores of the Calamospora-type isolated from specimen E 870; All  500.
5.
Detail of Discinites raconicensis (Němejc) emend. Specimen E 876 (holotype) showing imprints of
sporangia;  3.
6.
Discinites raconicensis (Němejc) emend. Holotype (E 876). Nearly the whole cone figured by Němejc (1941)
in this text-figure 3. Rakovník, Moric Mine locality, Kladno-Rakovník Basin, roof shale of the coal seam
called „Věnec“, Nýřany Member, Kladno Formation, Westphalian D.  1.
PLATE 4
1.
Discinites cf. raconicensis (Němejc). Fragment of cone (E 3610), Lubná near Rakovník, 1st May Mine locality,
Kladno-Rakovník Basin, upper Radnice coal seam, Radnice Member, Kladno Formation;  1
2.
Discinites cf. raconicensis (Němejc). Detail of Fig. 1 showing sporangia and their imprints;  3.
3-7. Microspores of the Calamospora-type isolated from Discinites cf. raconicensis (Němejc). (E 3610).
Notice relatively immature forms with fragments of outer layer (figs 3, 5-6) and mature microspores without
this layer (figs 4, 7); All  400.
8. Discinites cf. raconicensis (Němejc). (E 3613). Detail of cone showing coalifield sporangia. Lubná near Rakovník,
1st May mine locality, Kladno-Rakovník Basin, upper Radnice coal seam, Radnice Member, Kladno Formation; 
25.
9-12. Microspores of the Calamospora-type isolated from specimen E 3613. Notice forms with fragments of outer
layer (figs 9, 11-12) and mature microspore (Fig. 10) resembling miospores of the Leiotriletes-type; All  400.
PLATE 5
Discinites hlizae (Němejc) emend. Holotype (E 872). Lubná, near Rakovník, Rako Mine, Krčelák, Kladno-Rakovník
Basin, Lubná group of coals, Radnice Member, Kladno Formation, upper Bolsovian.
1.
Fragment of a twig with eight naturally associated cones figured by Němejc (1941, pl. 1, fig. 1);  0.3.
2, 4-6.
Relatively immature microspores of the Leiotriletes-type; All  500.
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24
3.
Detail of isolated sporangia;  20.
PLATE 6
Discinites hlizae (Němejc) emend. Holotype (E 872), Lubná, near Rakovník, Rako Mine, Krčelák, Kladno-Rakovník
Basin, Lubná group of coals, Radnice Member, Kladno Formation.
1.
Detail of Pl. 5, Fig. 1;  1.
2.
Microspores of the Calamospora-type. Notice opened trilete mark and fossilizied inner content of the microspore
going out;  500.
3.
Microspore of the Calamospora-type with two parallel folds of exine resembling the dispersed species
Calamospora breviradiata Kosanke, 1950;  500.
4.
Microspore of the Calamospora-type with one main fold of exine covering one-half of the body and comparing
with the dispersed species Calamospora pedata Kosanke, 1950;  500.
5.
Two sporangia from Fig. 1. Notice the scar at the base of the sporangium and longitudinal striation caused by long
and narrow oriented cells of sporangium wall;  20.
6.
Microspore of the Leiotriletes-type;  500.
7.
Megaspores comparing with the dispersed species Calamospora laevigata (Ibrahim) Schopf, Wilson and Bentall,
1944;  100.
8.
Megaspore comparing with the dispersed species Calamospora laevigata (Ibrahim) Schopf, Wilson and Bentall,
1944 with two microspores of the Calamospora-type on the exine surface, SEM;  135.
9.
Imprint of sporangium from Fig. 1 showing narrow cells;  50.
PLATE 7
Microspores isolated from Discinites hlizae (Němejc) emend. Holotype (E 872). Notice intermediate stages of
microspores from immature forms (Figs 1-8) with strongly reticulate outer layer and relatively relatively less mature
microspores (Figs 9-10) with fragments of outer layer and fully mature microspores (Fig. 11) of the Calamosporatype; All  500.
PLATE 8
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Microspores isolated from Discinites hlizae (Němejc) emend. Holotype (E 872). Notice relatively immature forms
(Figs 1-5) with primary and secondary (Figs 1-3) reticulate outer layer and relatively less mature microspore (Fig. 6)
with fragments of outer layer on inner body of the Calamospora-type; All SEM.
1.
 700.
2.
Detail of Figure 1;  1400.
3.
 600.
4.
 800.
5.
 700.
6.
 700.
PLATE 9
Discinites nemejcii sp. nov. Malé Přílepy, Malé Přílepy Basin, whetstone horizon, Radnice Member, Kladno
Formation, Bolsovian.
1-2. Holotype (E 875a-b). Fragment of cone probably without base and apex figured by Němejc (1941 on textfig. 4.);  1.
3.
Detail of Fig. 1;  3.
4.
Megaspore comparing with the dispersed species Calamospora laevigata (Ibrahim) Schopf, Wilson and Bentall,
1944; 100.
5-7. Relatively immature microspores isolated from Discinites nemejcii sp. nov. (Holotype E 875a) with fragments of
outer layer on the exine surface, Fig. 6 NOMARSKI objective; All  500.
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Fructifications
Discinites major
Size of spores (m)
95-115
1000-1300
D. hlizae
33 (60-85)97
280 (500) 650
D. hlizae
40 (86) 118
510 (636) 823
D. sp. 1 sensu Němejc
60 (69-78) 87
(=D. nemejcii)
450 (530-680) 780
D. sp. 3 sensu Němejc
42 (56-70) 77
(=D. nemejcii)
305-560
D. delectus
75-90
660-750
D. sp. sensu Remy and Remy 45 (85) 130
D. sp. (cf. bohemicus)
33 (54) 90
550-640
D. sp. A sensu Bek
54 (81) 98
(=D. cf. raconicensis)
Noeggerathiostrobus vicinalis 27 (62) 110
450-635
Classification
Calamospora
References
Němejc 1937, 1941
Calamospora
Němejc 1941
C. cf. microrugosa, C. cf. pedata, C. cf. liquida,
Bek 1998
C. cf. breviradiata, C. cf. laevigata, Pteroretis primum
Calamospora
Němejc 1941
Calamospora
Němejc 1941
Calamospora
Arnold 1944
Calamospora
Calamospora
Remy and Remy 1956
Remy and Remy 1956
C. cf. microrugosa, C. cf. pedata, C. cf. breviradiata,
Pteroretis primum
Calamospora
Bek 1998
TABLE 1. Palynologically studied fructifications of Discinites.
Spec. No.
E 3610
E 3611
E 3612
E 3613
E 3617
Size of microspores (m)
620 (825) 1530
650 (755) 1150
660 (808) 1060
610 (826) 990
410 (829) 1320
TABLE 2. The size of microspores isolated from Discinites cf. raconicensis.
Fructifications
Discinites bohemicus
D. major
D. raconicensis
Parent plants
Palaeopteridium reusii
P. macrophyllum
Rhacopteris asplenites
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Remy and Remy 1955
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D. cf. raconicensis
D. hlizae
D. nemejcii
???R. elegans
??? R. speciosa
Saaropteris guthoerlii
??? R. linearis
??? R. postculmica
??? R. bipinnata
TABLE 3. Probable botanical affinity of Bohemian Discinites.
Discinites species
D. bohemicus
Number
of
Shape
specimens
5
oblong
Length (mm)
>100
Cone
Width Axis width
(mm)
(mm)
22-25 3-4
Internodium
length (mm)
3-5
Number / disc Shape
cca 100-150
Sporangia
Length (mm)
elongate elliptical
25-3
Width (mm)
12-2
D. cf. bohemicus (isolated
sporophylls)
D. raconicensis
9
not obtainable not obtainable 22-28
3-45
not obtainable not obtainable not obtainable
not obtainable not obtainable
1
oblong
>210
46
not obtainable
5 (near the
apex), 8-11
several
hundreds
25-3
2-25
D. major
4
ovate
>90
30-35
?3-4
cca 4-5
50 - apex
elongate elliptical
170-200 centr.
28-38
13-2
D. hlizae
7
oblong
80-115
30-40
not obtainable
4-5
not obtainable oval to rounded
? cca 700
(15) 2-25
14-2
D. nemejcii
2
oblong
>92
30-38
? 2-3
cca 4
? 250-700
15-25
1-18
D. cf. raconicensis
5
oblong
>160
60-70
(45)
not obtainable
24-4
12-22
not obtainable not obtainable 18-22
not obtainable > 48
18
4-55
not obtainable
D. discus
3
D. sp. sensu Remy et Remy* 1
oval to isodiametrically hexagonal
elliptical to nearly
isodiametrically
hexagonal
5-7
cca 900-1400 elongated,
isodiametrically
hexagonal
not obtainable not obtainable not obtainable
not obtainable not obtainable
25-3
not obtainable not obtainable
not obtainable not obtainable
TABLE 4. Dimensions of Discinites. Measured values of sporangia may vary from their original natural dimensions
due to different compession of the original three-dimensional sporangia. *-comparable to D. cf. bohemicus.
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