1
Six rare Lepidostrobus species from the Pennsylvanian of the Czech
Republic and their bearing on the classification of lycospores
Jiří Beka*, Stanislav Opluštilb
aLaboratory
of Palaeobiology and Palaeoecology, Institute of Geology, Academy of Sciences, Rozvojová 135, 165 00
Prague 6, Czech Republic, e-mail: mrbean@gli.cas.cz
b
Faculty of Science of the Charles University, Albertov 6, 126 43 Prague 2, Czech Republic, e-mail: oplustil@natur.cuni.cz
Abstract
Grouping of the common and diverse Carboniferous dispersed miospore genus Lycospora is
suggested. Authors divide Carboniferous lycospores into six subgroups, based on their
morphology and the knowledge of their in situ records. The stratigraphical range of the cones
is from the Langsettian to the Stephanian B. Six plant specimens from the Bohemian Late
Palaeozoic continental basins belong to six Lepidostrobus species. Differing morphology of
fructifications and their in situ spores lead to a separation of the cones into the following
species: Lepidostrobus kohoutii sp. nov. containing Lycospora uzunmehmedii spores,
Lepidostrobus cf. haslingdenensis containing Lycospora uber spores, Lepidostrobus sp. A
yielded spores that compare with the dispersed species Lycospora microgranulata,
Lepidostrobus sp. B possessed Lycospora cf. microgranulata spores, Lepidostrobus sp. C
yielded spores of the Lycospora torquifer-type, and Lepidostrobus sp. D contains Lycospora
cf. subjuga spores. It means, that each cone species yielded lycospores comparable to one
dispersed spore species. The palaeoecology of the plant fossils is discussed. Their
environment is typified by either clastic or mixed clastic/peat substrates and standing water.
Therefore, they grew in clastic swamps developed along the lake margins or shallows.
Key words: Lycospora, Lepidostrobus, in situ and dispersed spores, Pennsylvanian.
*
Corresponding author
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1. Introduction
Lycospores belong to the most abundant and the most reported dispersed miospores of
Pennsylvanian age. Authors propose six subgroups of dispersed Carboniferous lycospores (not
new genera, subgenera, species or varieties), based on their morphological characteristics, i.e.
the occurrence of the cingulum and the zona and their widths, the type and the number of
sculpture elements on distal and proximal surfaces and perforations of zona. A comparison
with in situ lycospores from the Carboniferous of the Czech Republic and elsewhere is made.
We compared compressions and petrified (coal-balls) specimens of cones based only on in
situ lycospores due to different mode of the preservation.
Compressed specimens of lycopsid strobili are among the most commonly occurring fertile
parts of plant fossils of the Pennsylvanian coal-bearing deposits. They are found either in
organic connection with leafy shoots of their parent plants or, more often, as isolated cones or
their fragments. Organic connection can provide correlation among the strobilus, its spores
and the parent plant, that can help us to distinguish otherwise morphologically similar cone
species. Otherwise, the determination of the isolated strobili is based only on their
morphology and spore content. Lepidostrobus (Brongniart) Brack-Hanes and Thomas is the
most frequent genus encountered among all the lycophyte fructifications. For a long time, it
has been accepted as a genus of heterogeneous morphology with spores including
bisporangiate and monosporangiate specimens. Accordingly, Chaloner (1953) and Felix
(1954) recommended use of the spores as an important criterion to distinguish different
natural cone species. However, the most important step in understanding the real nature of
Lepidostrobus was achieved by Brack-Hanes and Thomas (1983). They studied the holotype
of the type species of the genus (Brongniart´s original specimen of L. ornatus Brongniart),
which yielded cingulizonate lycospores. Therefore, they restricted the use of the genus
Lepidostrobus only to male monosporangiate cones with lycospores. Those cones, containing
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different microspores or megasporangiate or even bisporangiate cones described formerly as
Lepidostrobus were assigned to different lycopsid genera, which are defined not only on spore
content and general cone morphology or anatomy, but also on the degree of integumentation
expressed by lateral alation or by the occurrence of integument.
Similar to the progress in cone taxonomy, a comparable advance has been made in the
investigation of their parent plants. Most of the studies are based on anatomically preserved
(petrified) specimens (DiMichele 1985; DiMichele and Bateman 1992; DiMichele and
Phillips 1985, 1994; Eggert 1961; Phillips and DiMichele 1992). These authors erected
several new genera and distinguished two principal types of lepidodendrid trees that
dominated the clastic swamps and peat mires during the Late Namurian and Westphalian. The
first type involves the monocarpic genera Lepidodendron Sternberg, Lepidofloyos Sternberg
and Synchysidendron DiMichele and Bateman. These are characterised by a cone-bearing
crown developed only in the final phases of their growth followed by tree death. The second
group represents polycarpic genera that produced small deciduous, cone-bearing, lateralbranch systems throughout life allowing them continuous or repeated reproduction. Polycarpic
forms are represented by several species of the genera Diaphorodendron DiMichele and
Paralycopodites (Morey and Morey) DiMichele. The latter genus is probably an anatomically
defined equivalent of the compression genus Ulodendron sensu Thomas. It was a small tree
that produced lateral deciduous branches with bisporangiate cones of the genus Flemingites
(Carruthers) Brack-Hanes and Thomas.
All the above mentioned genera differ not only in their habitus but also in ecological
constraints concerning the nutrient supply, water-table level and its stability. Lepidodendron
probably favoured clastic to mixed peat/clastic nutrient-rich substrates in habitats with
standing water. Only a few species preferred purely peat substrates (Phillips and DiMichele
1992). Therefore, most species occupied poorly drained floodplains, wet clastic swamps or
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lake margins. Lepidofloyos was also restricted in its occurrence to habitats with standing water
but with peat substrates. Nevertheless, its habitats overlapped to some degree with
Lepidodendron. Most of its occurrence was concentrated in rheotrophic, only occasionally
inundated mires. Diaphorodendron and Synchysidendron were better adapted to habitats with
exposed, to partially submerged, peat substrates being associated with dense undergrowth.
Paralycopodites probably preferred environments with intermittent floodings, clastic input
and peat exposure being usually associated with coal-seam seat-earth clastic partings
(DiMichele and Phillips 1994).
From the above-mentioned short history of lycopsid cone investigation, results dealing with
in situ spores provide important data for the determination of natural species as well as for
ecological constraints of their parent plants. The ideal result would be the correlation of
spores–fructification–parent plant. Such a correlation allows reconstruction of the vegetational
history of coal seams and associated clastic sediments.
Lepidostroboid fructifications from the Czech Republic were described by Feistmantel
(1873 a,b) for the first time, but later mainly by Němejc (1954). The taxonomical studies of
the latter author are based only on cone morphology. In situ spores of some Bohemian
Lepidostrobus species and some related lycopsid cones were first studied by Drábek (1967).
His results, however, have remained unpublished. The most recent research has been carry out
by Bek (1998) and Bek and Opluštil (1998, 2004). Current results are presented in this
contribution, but further investigation is in progress.
2. Material and methods
All of the specimens originated from central and western Bohemian Late Palaeozoic basins
(Plzeň and Kladno-Rakovník basins). The stratigraphical ranges of the studied cones vary
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from the Bolsovian to the Stephanian B. The Lepidostrobus cones are classified in accordance
to the approach of Thomas and Brack-Hanes (1984). Where possible, palynological samples
were taken from various parts of cones (basal, middle and apical) to determine the
morphological variations or ontogenetic stages of the spores. Spores were recovered by
dissolving small portions (separated from the cone species with a mounted needle) of cones in
nitric acid for 24-48 hours and KOH for 1-2 hours. Most of spores were mounted in glycerine
jelly for direct microscopic examination. Others were coated with gold for examination with
the CAMECA SX100 and TESLA scanning-electron microscopes. Measurements of spore
diameters and cingulum and zona widths were taken from 40-50 spores per cone. This number
is sufficient and does not differ from results, based on 100 or 150 specimens. Lycospores
obtained from studied cones were classified according to the system of dispersed spores
suggested by Potonié and Kremp (1954, 1955), Dettmann (1963) and Smith and Butterworth
(1967). In situ spores were compared directly with the original diagnoses (holotypes),
descriptions and illustrations of dispersed lycospore species. Species determinations are based
only on these original diagnoses, and not on the interpretations of subsequent authors.
Comparisons are made with other lycospores isolated from various Lepidostrobus cones. Only
palynological comparisons of compressed and petrified (coal-balls) specimens is evaluated
due to different preservation of fossils.
Studied cone specimens and palynological slides are housed in the palaeobotanical
collection of the National Museum in Prague. Negatives and digital photos of spores are
stored in the Institute of Geology, Academy of Sciences of the Czech Republic, Prague.
Digital photos of cones are in the Faculty of Sciences, Charles University, Prague.
Measurements and the type of sculptures of proximal and distal surfaces of Bohemian in situ
lycospores are given in Tab. 1. It was possible to compare our results only with papers where
a good and precise description or measurements of in situ spores have been made. Therefore,
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we could use for comparison only data reported by Thomas (1965, 1970, 1987, 1988), Willard
(1989b), Thomas and Dytko (1980), Brack-Hanes and Thomas (1983), Hagemann (1966) and
Bek and Opluštil (2004) from compressed specimens and from coal-balls by Felix (1954),
Balbach (1966), Taylor and Eggert (1968), Leisman and Rivers (1974) and Willard (1989a).
Data from these in situ lycospores are given in Table 2 (compressed specimens) and Table 3
(coal-balls specimens). Measurements of Bohemian Lepidostrobus cones are given on Tab. 4.
3. Systematic palaeontology
Class Lycopsida Scott, 1909
Order Lepidocarpales Thomas and Brack-Hanes, 1984
Genus Lepidostrobus (Brongniart) Brack-Hanes and Thomas, 1983
Type species Lepidostrobus ornatus Brongniart, 1828
Lepidostrobus kohoutii, sp. Nov. (Plate I, 1-7).
Holotype: Specimen E 6112, the National Museum, Prague.
Type locality: Ronna Mine in Kladno, Kladno-Rakovník Basin.
Type horizon: The holotype is preserved in roof shale of the Upper Radnice Seam, Radnice
Member, Kladno Formation, Bolsovian, Pennsylvanian.
Material: The only specimen is the holotype.
Etymology: Mr. T. Kohout is the finder of the holotype.
Diagnosis: The strobilus 80 mm long, 35 mm wide. Width without distal laminae 22 mm,
cone axis 4 mm wide. Pedicels perpendicular to the cone axis, 10 mm long, 1 mm high.
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Sporangia oval, 2 mm high along the whole length and as long as pedicels. Distal laminae
narrow, triangular, 23 mm long, with maximum width 5.5 mm at or near the base. Spores of
subtriangular to subcircular amb. The diameter 31-45 m. Laesurae simple reaching 4/5 of the
radius, sometimes labrum 1-3 m wide and high. Cingulum 2-4 m wide. Zona 3-6 m wide.
Proximal and distal part of zona laevigate sometimes punctate or perforated. Proximal surface
of central body laevigate to finely scabrate. Distal surface of central body densely
microspinate to microgranulate.
Description: Although the holotype is 80 mm long, estimated length of L. kohoutii is between
150-250 mm. It is longitudinally split showing the axis, pedicels and distal laminae. However,
the preservation state (specimen is preserved in a grey mudstone with sandy admixture) does
not provide many details on inner morphology. The cone body, including distal laminae has a
total width of 36 mm. Pedicels are attached at an angle of about 85 degree to the cone apex.
Distal laminae are attached to the cone body at an angle of about 30-35 degree, and are
narrow, entire, of triangular to slightly lanceolate shape. They are gently arched curved to the
apex. The apex is, however, not preserved but it may be relatively acute as is indicated by
gradual tapering of the cone in this direction. The midvein is distinct within the whole length
of the laminae. Spores are 31.0 (36.2) 45.0 m in diameter. Labrum is 1.0 (2.1) 3.0 m.
Cingulum is 2.0 (3.3) 4.0 m wide and zona 3.0 (4.5) 6.0 m wide. Zona is sometimes
laevigate, punctate or perforated (Pl. I; 2, 3, 5-7).
Comparison: Spores released from this specimen compare with the dispersed species
Lycospora uzunmehmedii Artűz (Artüz 1957, p. 250). Lepidostrobus kohoutii differs from
other Bohemian lepidostroboid strobili mainly by its spores. In terms the cone can be
comparable with L. sp. D (Plate II, 25) and L. stephanicus (Němejc) Bek and Opluštil, which
yielded different type of lycospores. From coalfields outside the Czech Republic, there are
several species that in size and general morphology seem to be comparable with this new
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species, especially L. meunierii Renault and Zeiller, L. jacksonii Arber and L. kidstonii
Zalessky. L. kohoutii microspores are of the similar morphological type to those isolated from
compression specimens of L. haslingdenensis Thomas and Dytko by Thomas and Dytko
(1980) and Willard (1989b), but differ in the sculpture of distal surface. Similar lycospores
were reported from compression specimens of L. barnsleyensis Thomas, L. dawsonii Thomas,
Bek and Opluštil, L. cf. haslingdenensis and L. jacksonii Arber. Spores named as Lycospora
cf. uzunmehmedii described by Bek and Opluštil (2004) from Bohemian species
Lepidostrobus thomasii Bek and Opluštil possess narrower cingulum and zona (Table 2). The
only very roughly similar in situ petrified (coal-balls) lycospores are described from L.
oldhamius Williamson by Willard (1989a), but these spores differed mainly by broader zona
and narrower cingulum (Tab. 3).
Stratigraphical range and geographic distribution: This species has been known until now
only from the Radnice Member (lower Bolsovian) of central Bohemia (the Kladno-Rakovník
Basin).
Parent plant: Unknown.
Paleoecology: The holotype is preserved in weakly laminated sandy mudstone from the roof
of the Upper Radnice Seam, which is interpreted as lacustrine sediment of an extensive lake
(Opluštil et al. 1999). Spores of L. kohoutii, however, only rarely occur in the Upper Radnice
Seam (Opluštil et al. 2001). Therefore, the parent plant probably preferred only clastic
swamps developed along lake margins or shallows. These habitats were characterised by
permanently standing water.
Lepidostrobus cf. haslingdenensis Thomas and Dytko, 1980. (Plate I, 8-13).
1998 Lepidostrobus cf. haslingdenensis; Bek, pl. 18, figs 15-18; pl. 19.
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Material: The only specimen comes from the Tuchlovice Mine, Roof shale of the Upper
Radnice Seam, Radnice
Member (Bolsovian), Kladno Formation. Kladno-Rakovník Basin. Specimen E 5587 is stored
in the National Museum, Prague.
Description: Specimen is represented by a fragment about 75 mm long of the middle part of a
cone preserved as a longitudinally split impression in grey mudstone. It measures 45 mm
across but only 32 mm without the distal part of the sporophylls (axis and pedicels). Axis is 4
mm wide. Pedicels are arranged into a low helix. They are 11 mm long and approximately 1
mm high being perpendicular or even gently turned backward. Distal laminae triangular,
entire, about 10-11 mm long, gradually tapering to the apex. They are inserted to the pedicels
at an angle of about 45 degree. Spores are of circular to sub-circular amb with a smooth or
slightly undulate outline. Size range is 30.0 (39.1) 45.0 m. The laesurae extends to the outer
margin of central body or onto the zona, often with a labrum 1-4.5 m on average. The
cingulum is 2.0 (2.57) 3.0 m wide and developed as dark ring on the outer margin of central
body. The proximal part of cingulum is laevigate or finely scabrate and the distal part
irregularly scabrate. The zona is 2.0 (4.39) 6.0 m wide, sometimes perforated. The proximal
parts of the zona are laevigate or finely scabrate, the distal parts are finely scabrate. The
sculpture of proximal surface of the central body is laevigate. The distal surface of the central
body is microspinate to microgranulate (Pl. I, 9). The exospore (especially the proximal
surface) sometimes shows various degree of degradation and, consequently it is not easy to
distinguish the original sculpture.
Comparison: Cone morphology of our specimen is similar (Thomas, pers. comm., 1999) to
the holotype of Lepidostrobus haslingdenensis, although it is slightly broader (due perhaps to
taphonomic factors or to different maturity levels). The British holotype is from a different
stratigraphic position (Namurian C) than the Bohemian cone. Due to these facts, we identify
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the Bohemian specimen as L. cf. haslingdenensis. Our lycospores are very similar to those
isolated from Lepidostrobus haslingdenensis by Thomas and Dytko (1980). These authors
compared their spores with the dispersed species Lycospora noctuina Butterworth and
Williams. The sculpture of this dispersed species is rugulate on the distal surface (rugulae
from 3-6 m wide and high, see Butterworth and Williams, 1958, p. 376). British in situ
lycospores possess different densely microspinate distal surfaces and Thomas (1987) emended
the original diagnosis of L. noctuina based on the occurrence of a large grana or verrucae or
rugulae, 1-3 m broad, distally in the central area within the cingulum. Thomas and Dytko
(1980) did not mention these rugulae and they are not seen on their original photos of the
distal surface. We are of the opinion that L. noctuina is defined as a species with a rugulate
distal surface and that it is not necessary to emend this species because a different type of
sculpture was found on in situ spores of Lepidostrobus haslingdenensis. Microspores isolated
from the holotype of L. haslingdenensis do not possess large rugulae, and therefore, they
cannot correspond to the original diagnosis of Lycospora noctuina, given by Butterworth and
Williams (1958). It is possible to compare isolated lycospores to some other, similar and welldefined dispersed species of the same morphological type, but with microspinate or
microgranulate distal surface. Lycospora uber (Hoffmeister, Staplin and Malloy) Staplin
compares (especially in the same type of distal sculpture) with both Bohemian and British
microspores. Lycospores isolated from the Bohemian specimen Lepidostrobus cf.
haslingdenensis are most similar to the dispersed species Lycospora uber. Lycospores of
Lepidostrobus haslingdenensis described by Willard (1989b) possess rugulate, i.e. different,
sculpture on the distal surface, and narrower zona. Willard´s specimen differs from the
Bohemian cone, and also from the holotype of this species, by having a broader cone axis (6-9
mm). We suppose that Willard´s specimen is not L. haslingdenensis what is supported by
different in situ lycospores. It seems that Willard´s spores correspond better to the original
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diagnosis of Lycospora noctuina given by Butterworth and Williams (1958), particularly as
regards the same rugulate distal surface. Morphologically roughly similar spores, but of the
Lycospora uzunmehmedii-type, were isolated from the Bohemian species Lepidostrobus
kohoutii and L. thomasii. Compression strobili of L. dawsonii, L. barnsleyensis and L.
jacksonii yielded this type of lycospores. The only very roughly similar in situ petrified (coalball) lycospores are described from Lepidostrobus oldhamius by Willard (1989a), but these
spores are smaller with narrower cingulum (Tab. 3).
Stratigraphic range and geographic distribution: Lepidostrobus cf. haslingdenensis is known
only from the Late Palaeozoic continental basins of the central and western Bohemia where it
occurs in the Radnice Member of the lower Bolsovian age. The type locality of this species is
Rossendale, Lancashire in Great Britain, and the type horizon is the Millstone Grit of
uppermost Namurian age (Thomas and Dytko 1980).
Parent plant: Unknown.
Palaeoecology: The specimen is preserved in the roof shale of the Upper Radnice Seam that is
of lacustrine origin. Its parent plant therefore probably grew along the lake margins in wet
habitats with standing water. Occurrence of its spores in the dispersed spore association of the
transition phase of the Upper Radnice Seam (Opluštil et al. 2001) indicates that its parent
plants also could have favoured peat substrates of eutrophic planar mires with high water
table.
Lepidostrobus sp. A. (Plate I, 14-20; Plate II, 1-5).
1998 Lepidostrobus sp. B (part); Bek, pl. 14., non pl. 15.
Material: The specimen originates from Kounov locality, near Rakovník, Otruby Member,
Slaný Formation
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(Stephanian B), Kladno-Rakovník Basin. The fragment of this cone is anatomically partly
preserved, fossilized by pyrite and oxidized now. The specimen E 3476 is stored in the
National Museum, Prague.
Description: Fragment of a longitudinally split cone, 30 mm in diameter. The cone axis is
about 7 mm wide, and shows diamond-shaped, helically arranged sporophyll scars. Pedicels
and sporangia are 10 mm long, attached to the axis at an angle of 50-60 degree. Distal laminae
are not preserved. Sporangia are oblong, 12 mm long. Spores are of triangular to subtriangular amb with smooth or finely undulate outlines. Size range is 32.0 (35.0) 42.0 m.
Laesurae are simple, extending to the outer margin of central body, sometimes with a labrum
1-2.5 m large. The cingulum is 1.4 (2.21) 3.5 m wide, developed as dark ring on the outer
margin of the central body. Proximal and distal parts of the cingulum are laevigate, or
irregularly scabrate. The punctate, or often perforated zona, is 1.5 (2.58) 3.5 m wide.
Circular, sub-circular and irregular perforations occur usually on the inner part of zona and
average about 1 m across (Pl. I, 15, 17; Pl. II, 2). The sculpture of the proximal surfaces of
central bodies is microgranulate or granulate (Pl. I; 15, 17), whereas the distal surfaces of
central bodies are densely microgranulate or granulate, in comparison (Pl. I; 18, 19; Pl. II, 1,
5).
Remarks: We do not erect a new species because the fragmentary nature of the specimen
prevents us from giving a precise and complete diagnosis. Isolated lycospores are correlated
with the dispersed species Lycospora microgranulata Bharadwaj (Bharadwaj 1957a, p. 104;
pl. 27, fig. 18).
Comparison: The studied strobilar fragment may be comparable with Lepidostrobus sp. B (Pl.
II, 6) or L. stephanicus. However, L. stephanicus contains different lycospores (Lycospora
punctata Kosanke) and it occurs at a different stratigraphical level. The cone morphology and
the size of Lepidostrobus sp. A is comparable with L. sp. D (Plate II, 25). However, a more
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precise comparison is impossible due to a poor preservation and incompleteness of L. sp. A.
Also, in situ lycospores of both species are roughly similar although not identical. Similar
spores with relatively narrow cingulum and zona belong to the most abundant in situ
lycospores. Spores isolated from Lepidostrobus barnsleyensis Thomas by Thomas (1965)
have the same perforations of the zona, but possess much broader cingulum and zona and
different sculpture elements. Lepidostrobus cf. squarrosus Kidston yielded spores comparable
(Willard 1989b) with Lycospora punctata, which have different density of sculpture elements
on the proximal surface, are smaller and the zona is not perforated. Spores described from
Lepidostrobus praelongus Lesquereux and L. variabilis Lindley and Hutton and comparable
(Willard (1989b) with Lycospora torquifer (Loose) Potonié and Kremp seem to be similar but
differ by smaller diameter, broader cingulum and a zona that is not perforated. Spores isolated
from Lepidostrobus sp. C sensu Hagemann and L. sp. D sensu Hagemann by Hagemann
(1966) and L. ornatus by Brack-Hanes and Thomas (1983) are smaller and lack the
perforations of the zona. Spores of the Bohemian species Lepidostrobus nemejcii Bek and
Opluštil and comparable with Lycospora triangulata Bharadwaj are smaller with a narrower
zona (Tab. 2). Similar spores isolated from petrified (coal-ball) specimens of Lepidostrobus
coulterii Jongmans and L. minor Leisman and Rivers were reported by Balbach (1966) and
Leisman and Rivers (1974). Both possess smaller lycospores, although the sculpture is
similar. L. fayettevillense Taylor and Eggert is another producer of similar cingulizonate
lycospores with prominent perforations of the zona and similar sculpture, only the cingulum
and the zona are broader (Tab. 3).
Stratigraphic range and geographic distribution: This specimen occurs in strata ranging from
the lower Bolsovian to the upper Stephanian B. Such a wide stratigraphical range is not
common among the tree lycopsids. The only known plant with such a long stratigraphical
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range is Asolanus camptotaenia Wood, however, the presence of other rarely occurring
species is not excluded.
Parent plant: Unknown.
Paleoecology: The specimen is preserved in lacustrine sapropelitic roof shale (called Švartna)
of the Kounov Seam. Parent plants probably grew in the wet habitat of lake margins, or
shallows with a high water-table necessary for their reproduction.
Lepidostrobus sp. B. (Plate II, 6-12).
1998 Lepidostrobus sp. B, (part); Bek, pl. 15, non pl. 14.
Material: The single specimen is preserved in whitish tonstein „velká opuka“, intercalated in
the Upper Radnice
Seam. Ronna Mine in Kladno, Radnice Member, Kladno Formation (Bolsovian), KladnoRakovník Basin. The specimen E 3542 is stored in the National Museum, Prague.
Description: The specimen represents only a fragment of the middle part of a longitudinally
split strobilus which is 43 mm wide including distal laminae. The axis is from 2.5 to 3 mm
thick. Pedicels in middle part are about 9 mm long. Distal laminae are rather narrow,
triangular in shape, with maximum width up to 3 mm at the base. They are about 12-13 mm
long being bent arch-like to the apex. Spores are of triangular to sub-triangular amb with
smooth or finely undulate outlines. Size range is 35.0 (40.1) 48.0 m. Laesurae are simple,
extending to the outer margin of the central body, sometimes with labrum 1-3.5 m large. The
cingulum is 2.0 (2.62) 3.5 m wide, developed as dark ring on the outer margin of the central
body. Proximal and distal parts of the cingulum are laevigate or irregularly scabrate. The
laevigate, punctate or sometimes perforated zona is 2.0 (2.73) 4.0 m wide. The sculpture of
the proximal surfaces of central bodies is microgranulate or granulate (Pl. II, 7). The distal
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surfaces of central bodies are more densely microgranulate or granulate than the proximal
surfaces (Pl. II; 11, 12), in comparison.
Remarks: Unfortunately spores of this type were obtained only from this specimen, and
therefore only a limited number of specific characteristics are observable. Consequently, we
do not propose a new species until we obtain additional more complete specimens, which will
allow its ordinary comparison and definition as a new species.
Comparison: The described specimen represents a fragment of a middle-sized cone that is
quite similar to Lepidostrobus lycopoditis Feistmantel, occurring at the same stratigraphical
level. However, these species differ in their spore content. Whereas Lepidostrobus sp. B
provided only microspores, L. lycopoditis is a bisporangiate species, containing both microand megaspores (Bek and Opluštil 1998) and belongs to the genus Flemingites.
Isolated spores are similar to those isolated from Lepidostrobus sp. A and compared with
the dispersed species Lycospora microgranulata. Lycospores of Lepidostrobus sp. B are
bigger with broader cingulum and zona and lower density of distribution of sculptural
elements on both surfaces, which allow classification with Lycospora cf. microgranulata.
Comparison with other compressed and petrified cones and spores is the same as in
Lepidostrobus sp. A due to the similarity of morphological character of the spores in both
cones.
Stratigraphic range and geographic distribution: The only specimen is from the lower
Bolsovian strata of the Radnice Member in the Kladno-Rakovník Basin.
Parent plant: Unknown.
Palaeoecology: The specimen is preserved in the volcanoclastic parting of the Upper Radnice
Seam and was probably buried in situ, i.e. probably within the range of the crown of its parent
plant. It is supposed that the parent plant favoured peat substrates of rheotrophic mires with
high water table.
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Lepidostrobus sp. C. (Plate II, 13-20).
1998 Lepidostrobus sp. C; Bek, Tab 16.
Material: Specimen E 3531 originated from the Lužná Mine near Rakovník, Bolsovian,
Kladno-Rakovník Basin and is stored in the National Museum, Prague.
Description: Small, 70 mm long fragment of the middle part of a longitudinally split strobilus.
The preserved width of the cone is 38 mm, but the original is estimated to be about 48 mm,
including distal laminae and 44 mm without them. The axis measures 8.5 mm in diameter.
Pedicels and sporangia are 19 mm long, being attached at an angle of about 60-70 degree.
Sporangia are 1.8 mm long. Spores are of triangular to sub-triangular amb. Size range is 26.0
(32.5) 36.0 m. Laesurae are simple, extending to the outer margin of central body, sometimes
with labrum 1-2 m large. Cingulum is 2.0 (2.71) 3.5 m wide, developed as dark ring on
outer margin of the central body. Proximal and distal parts of the cingulum and zona finely
microgranulate. The punctate or sometimes perforated zona is 1.4 (1.96) 2.6 m wide. The
sculptures of both surfaces of central bodies are microgranulate or granulate. The density of
distribution of sculptural elements is roughly the same on both surfaces.
Remark: Isolated spores are closely comparable with the dispersed species Lycospora
torquifer (Loose) Potonié and Kremp. The holotype of L. torquifer was established by Loose
(in Potonié, Ibrahim and Loose 1932). Potonié and Kremp (1955; pl. 17, 355) illustrated the
holotype that is not of the Lycospora-type (Somers et al. 1972). All other miospores referred
by Potonié and Kremp (1955; pl. 17, 356-359) to this species evidently belong to Lycospora
(Ibrahim) Schopf, Wilson and Bentall. The Bohemian lycospores are correlated with these
specimens, i.e. L. torquifer sensu Potonié and Kremp (pl. 17, 356-359; non pl. 17, 355).
17
Comparison: Lepidostrobus sp. C is quite similar in size and outer morphology to
Lepidostrobus sp. B (Pl. II, 6), although L. sp. C seems to be slightly more robust. However,
the small differences in sizes are unimportant for their separation into two species. Different
spore content is a sufficient criterion. Spores isolated from L. sp. A and L. sp. B are larger in
diameter and possess broader and perforated zona. Spores from L. sp. D are larger and belong
to a different spore species (Lycospora subjuga-type). Spores isolated from compressed
specimens of Lepidostrobus praelongus and L. variabilis and compared by Willard (1989b)
with the same dispersed species Lycospora torquifer have broader cingulum and zona and
different types of sculpture elements. Very similar spores were described from compressed
specimens of Lepidostrobus sp. D by Hagemann (1966) and L. nemejcii by Bek and Opluštil
(2004). Spores isolated from petrified (coal-ball) specimens of L. coulterii by Balbach (1966)
and L. minor by Leisman and Rivers (1974) are also of similar morphology.
Stratigraphic range and distribution: The only known specimen occurred at the level of the
Radnice Member, Kladno Formation (lower Bolsovian)
Parent plant: unknown.
Palaeoecology: The specimen is from the volcanoclastic partings of the Upper Radnice Seam
mined in a small coalfield in Lužná, near Rakovník. Its parent plant preferred inundated
eutrophic mires.
Lepidostrobus sp. D. (Plate II, 21-25).
1954
Lepidostrobus kidstonii Zalessky; Němejc, pl. 1, fig. 4.
1998
Lepidostrobus sp. A, (part); Bek, pl. 13; non pls 11, 12.
Material: Only one specimen was found. It was referred by Němejc (1954; pl. 1, 4) to
Lepidostrobus kidstonii, preserved in a roof shale of the Upper Radnice Seam, Radnice
Member (Bolsovian), Kladno
18
Formation. The specimen is from the Kralupy upon Vltava (Červená Hůrka Hill) locality in
the south-eastern margin of the Kladno-Rakovník Basin. The specimen, E 5596, is stored
in the National Museum, Prague.
Description: The specimen represents an incomplete 125 mm long part of a slender nearly
cylindrical cone with a blunt apex. Sporophylls are helically arranged. The width of the cone
body is about 20 mm, the length/width ratio is >6:1. The specimen is preserved as a
compression; detached distal laminae are not preserved. Internal morphology including the
axis, is not preserved. Spores are of sub-triangular to sub-circular amb, their outlines are
smooth or finely undulate. The diameter is 28.0 (35.0) 41.0 m. The labrum is 1-2.5 m wide
and high. The cingulum is 2.0 (2.7) 3.5 m wide developed as dark ring on outer margin of
the central body. The zona is 1.0 (1.8) 2.8 m wide. The density of micrograna is higher on
the distal surfaces of central bodies than on the proximal surfaces, in comparison (Plate II,
21). The inner part of the zona is sometimes laevigate, sometimes punctate (perforated on only
one surface) and sometimes it seems to be perforated through (perforations of both, proximal
and distal surfaces).
Comparison: Lepidostrobus sp. D might be comparable with L. nemejcii, L. kidstonii, and also
with L. meunierii. L. sp. D is apparently slender in comparison with L. nemejcii but in situ
lycospores of both species are of the same morphological type. L. kidstonii is another
comparable species, as confirmed by Němejc´s misinterpretation of the holotype of L. sp. D.
Czech specimen fits with Zalessky´s L. kidstonii in size and general morphology. However,
due to limited number of available specimens, as well as just one type of preservation,
detailed comparison is impossible. Nevertheless, the question of exact relationships of these
two species remains unsolved until microspores of L. kidstonii become available. Similarly,
the comparison with other species, such as L. meunierii, is only of limited value due to the
absence of spore content of the holotype. Our specimen is also rather poorly preserved.
19
Isolated lycospores (Pl. II, 21-24) are similar to the dispersed species Lycospora subjuga
Bharadwaj (Bharadwaj 1957b; p. 127; pl. 25, 84). The holotype of L. subjuga has broader
cingulum and zona and we classify our microspores as L. cf. subjuga.
Microspores isolated from Lepidostrobus sp. D are smaller than those from L. sp. A and L.
sp. B and possess narrower zona. Lycospores yielded by L. sp. C are similar only with smaller
diameter. Very similar spores are reported from L. nemejcii by Bek and Opluštil (2004) and L.
sp. D by Hagemann (1966) and the morphology of the cone specimens is the main criterion
for their determination. Roughly similar spores have been isolated from compressed
specimens of L. sp. A by Willard (1989a), which are smaller and possess a much broader
cingulum (Tab. 2). Spores yielded by compressed specimens of L. obovatus (Rénier) Bek and
Opluštil, L. stephanicus by Bek and Opluštil (2004) and L. ornatus by Brack-Hanes and
Thomas (1983) have different sculpture (Tab. 2).
Spores isolated from petrified (coal-ball) specimens of L. coulterii by Balbach (1966) and
L. minor by Leisman and Rivers (1974) are also of similar morphology (Tab. 3).
Parent plant: Unknown.
Stratigraphic range and geographic distribution: This species is known only from lower
Bolsovian continental deposits (Radnice Member) of the central Bohemian Late Palaeozoic
basins.
Palaeoecology: Occurrence of the species in the lacustrine-roof shale of the Upper Radnice
Seam indicates that the parent plant probably occupied habitats with wet clastic substrates and
standing water along the lake margins. However, its spores belong among the commonly
occuring lycospores of the Upper Radnice Seam, in which it is restricted to the lower and
uppermost parts interpreted as eutrophic planar mires with increased clastic input and standing
water above the peat substrate (Opluštil et al 2001).
20
4. Carboniferous dispersed lycospores
In situ lycospores were probably the first described Carboniferous miospores. Lycospores
were often reported together with lycopsid in situ megaspores from cones of the Flemingites
and “Lepidostrobus”-types by, e.g. Kidston (1889), Williamson (1893) and others. To date
about 100 dispersed Carboniferous lycospore “species” have been described in the literature.
Some of them are very similar and can be synonymised (see Somers et al. 1972, Bek and
Opluštil 2004) and others possibly do not belong to this genus (Bek and Opluštil 2004). The
genus Lycospora was proposed by Schopf, Wilson and Bentall in 1944 for: “…trilete radial
spores…equator marked with a short thick tapering ridge, 18-45 μm…nearly smooth;
minutely granulose or rugose; equatorial ridge resembles a typical flange development”
(Schopf, Wilson and Bentall, 1944, p. 54). The chosen type species was Lycospora
micropapillata (Wilson and Coe) Schopf, Wilson and Bentall. Piérart (1964) divided
lycospores into three main groups (Lycospora, Microcingulata and Bizonaria) and identified
some synonymous species.
Somers et al. (1972) published a revision of Carboniferous lycospores and studied and
described several their holotypes. They divided all of them into four main artificial groups,
Lycospora orbicula, L. noctuina (including two varieties), L. rotunda and L. pusilla (including
four varieties). Tens of species of dispersed lycospores were synonymised, based on pure
artificial criteria. Somers et al. (1972) chose L. pusilla as a very important species, giving the
name to the biggest group of lycospores. The problem is that the preservation of the holotype
of Lycospora pusilla is far from perfect, therefore, it is impossible to observe important
morphological details and we can rely only on original descriptions. Ibrahim (1932 in Potonié
et al. 1932) proposed Lycospora pusilla (as Sporonites pusillus Ibrahim) as a cingulate
species, i.e. lacks the zona. Somers et al. (1972) redefined Lycospora pusilla as a
cingulizonate species, i.e. possesses cingulum and zona, although they (Somers et al., 1972,
21
pp. 33-34) stated in the description of the holotype, that the zona is not visible (!) and that the
equatorial structure is only 2-3 μm broad. It is not probable (if the equatorial structure 2-3 μm
would consist of cingulum and zona), that the holotype is really cingulizonate, because the
width of the cingulum and the zona of cingulizonate lycospores is very rarely less than 3-4
μm. Somers et al. (1972) did not follow Ibrahim´s concept of L. pusilla but authors accept L.
pusilla as a cingulate species, i.e. L. pusilla sensu Ibrahim non Somers et al. Authors observed
(Bek 1998, Bek and Opluštil 2004 and herein) several thousands of in situ lycospores, but
cingulizonate specimens, i.e. with so narrow equatorial structure consisting of cingulum and
zona, occur very rarely, if soever.
We propose a new, more detailed grouping of Carboniferous lycospores, based on three
main factors and own knowledge of in situ spores. The first criterion is the width of the zona
and cingulum, the second is the type of sculpture elements on the proximal and distal surface
and the third criterion is the comparison of the number of sculpture elements (“density”) on
the proximal and distal surface. It is very significant feature, because usually the number of
sculpture elements on the distal surface is greater, than on the proximal hemisphere.
It is possible to divide Carboniferous lycospores, based on these criteria, into two principal
types, cingulate and cingulizonate specimens (Thomas 1970). Cingulate lycospores can be
divided into two subgroups.
The illustrations and descriptions of several lycospore species are poor and authors have not
an opportunity to observe all of them (i.e. original slides) and decide their affinity to
subgroups. It is the reason, why we do not mention subgroups of all dispersed lycospore
species (more than 100), but we could choose only the most typical, well defined and often
reported species. On the other hand, the list of all Carboniferous dispersed lycospore species
does not exist.
22
The first subgroup consists of cingulate lycospores with prominent microspinate sculpture of
the distal surface and high number of sculpture elements there. For example, holotypes of the
dispersed species Lycospora orbicula and L. granulata Kosanke belong to this subgroup. All
microspores of this type were produced by cones of the genera Flemingites and probably
Bothrodendrostrobus Hirmer. In situ lycospores of this subgroup were described from several
compression specimens of flemingitalean cones (Bek 1998) from the Pennsylvanian of the
Czech Republic.
The second subgroup of cingulate lycospores is typified by microgranulate to
microverrucate sculpture of the exospore. For example, dispersed species Lycospora parva
Kosanke, L. rugosa Schemel and L. granianellatus Staplin can be assigned to this subgroup.
In situ lycospores of this type were reported from tens of compression specimens of cones
Lepidostrobus sternbergii (Němejc) Bek and Opluštil from the Pennsylvanian of the Czech
Republic and from some petrified and compression Lepidostrobus specimens rom elsewhere.
It is possible to divide cingulizonate Carboniferous lycospores into four subgroups
depending on the width of cingulum and zona and the sculpture of the proximal and distal
surfaces.
The third subgroup consists of cingulizonate lycospores with a relatively narrow cingulum
and narrow zona. The width of the zona is less than 3-3.5 μm. For example, dispersed species
Lycospora denticulata Bharadwaj, L. contacta Habib, L. subjuga Bharadwaj, L. brevijuga
Kosanke, L. brevis Bharadwaj, L. microgranulata Bharadwaj, L. punctata Kosanke and L.
triangulata Bharadwaj can be assigned to this group.
Cingulizonate lycospores of the fourth subgroup are typified by a relatively narrow
cingulum and wide zona. The width of the zona is more than 4 μm. For example, dispersed
species Lycospora loganii (Wilson) Potonié and Kremp, L. pellucida (Wicher) Schopf,
Wilson and Bentall, L. micropapillata (Wilson and Coe) Schopf, Wilson and Bentall, L.
23
micrograna Hacquebard and Barss, L. intermedia (Wilson and Hoffmeister) Wilson and
Hoffmeister, L. pseudoannulata Kosanke and L. perforata Bharadwaj and Venkatachala
belong to this subgroup. Some specimens can possess more or less prominent perforations of
the zona (L. perforata, L. pseudoannulata).
Lycospores with a relatively wide cingulum and wide zona can be divided into two
subgroups, based on the type of sculpture elements. The fifth subgroup is characterized by the
microspinate sculpture of the distal surface and laevigate to very finely scabrate proximal
surface. For example, the dispersed species Lycospora noctuina (Butterworth and Williams),
L. nitida Artüz, L. uzunmehmedii Artüz, L. tenuireticulatus Artüz and L. uber can be assigned
to this subgroup. The affinity of L. paulula Artüz is very questionable, because the holotype
more resembles some densospores rather than lycospores. In situ lycospores of this subgroup
are described mainly from compression Lepidostrobus specimens in the Czech Republic as
well as elsewhere and their occurrence in coal-balls is rare.
Lycospores of the sixth subgroup are characterized by a relatively wide cingulum and wide
zona, prominent microgranulate to granulate sculpture of the distal surface and laevigate to
very finely scabrate sculpture of the proximal surface. For example, the dispersed species
Lycospora rotunda Bharadwaj, L. breviapiculata, L. subtriquetra, L. curtata and L. curtata f.
velata belong to this subgroup. In situ lycospores of this type are known only from
compression strobili of Lepidostrobus ronnaensis from the Pennsylvanian of the Czech
Republic (Bek and Opluštil, 2004).
In situ spores isolated from Bohemian specimens of L. kohoutii and L. cf. haslingdenensis
and compared to the dispersed spore species Lycospora uzunmehmedii and L. uber belong to
the fifth subgroup of cingulizonate lycospores with a relatively broad cingulum and zona. All
other lycospores isolated from Lepidostrobus sp. A-D can be assigned to the third subgroup of
cingulizonate lycospores with a relatively narrow cingulum and zona.
24
5. Conclusions
Six specimens of less commonly known lycopsid cones assigned to six species of the genus
Lepidostrobus are described from the Late Carboniferous continental coal-bearing deposits of
central and western Bohemia. All of the findings represent only isolated strobili, or mainly
their fragments, and their determination is, therefore, based only on external morphology and
spore content, whereas their parent plants are unknown. Despite their fragmentary nature,
these strobili complete demonstrated the high-specific diversity of lycopsids in the continental
basins of the Bohemian Massif during the Westphalian and Stephanian (Bek and Opluštil
2004).
Plants preferred habitats with either clastic or mixed clastic/peat substrate and standing
water. Therefore, they grew in clastic swamps developed along the lake margins or shallows.
More precise characterisation of the habitats of particular species and their synecological
relationships are, however, not known, because all the specimens underwent various degree of
transport before their final burial.
A new detailed grouping of Carboniferous lycospores, based on morphological criteria and
knowledge of in situ, i.e. natural species is proposed. It is possible to recognise six subgroups
of Carboniferous lycospores.
Cingulate species are determined, based on the type of the distal surface. Cingulate
lycospores of the first subgroup possess densely microspinate sculpture of the distal surface
while the second subgroup of cingulated lycospores is characterized by microverrucate,
microgranulate exospore of the distal surface.
25
Cingulizonate lycospores can be divided into four subgroups, based on the width of the
cingulum and the zona and the type of sculpture elements. The third subgroup is typified by a
relatively narrow cingulum and narrow zona.
The fourth subgroup is characterized by a relatively narrow cingulum and wide, sometimes
perforated zona.
Lycospores of the fifth subgroup possess a relatively broad cingulum and broad zona and
microspinate sculpture of the distal surface.
The sixth subgroup of Carboniferous lycospores consists of specimens with a relatively
broad cingulum and broad zona and prominent microverrucate and verrucate distal surface.
All these morphological types of dispersed lycospores were found as in situ from several
Lepidostrobus species from the Pennsylvanian of the Czech Republic (Bek 1998, Bek and
Opluštil 2004, herein).
All in situ lycospores from one cone specimen are of the same type and can be assigned to
one dispersed spore species. We do not know parent plants of all dispersed lycospores. It is
possible to determine only a few of them, especially parent cones of some morphologically
prominent lycospore types.
For example, cingulate lycospores of the first subgroup were produced by Flemingites and
maybe Bothrodendrostrobus cones (Bek 1998). It seems, that cingulate lycospores of the
second subgroup were produced by very robust, i.e. very thick and long (about one meter or
even longer) cones of the Lepidostrobus sternbergii-type (Bek 1998, Bek and Opluštil 2004).
The authors visited several collections of Carboniferous plants in the Czech Republic and in
Museum fűr Natűrkunde in Berlin and Chemnitz, Germany, Natűrwissenschaften Museum
and Geologische Bundesanstaldt in Vienna, Austria, the University of Lille, France and the
Museum in Leiden, Netherlands and all parent cones of such cingulate lycospores were always
of the same (robust) type.
26
Only a few parent plants of lycospores of the third and fourth subgroups can be determined.
It may be possible to determine only those produced cingulizonate lycospore with a narrow
cingulum and a broad perforated zona (Lycospora perforata-type).
Parent plants of lycospores of the fifth subgroup are hardly determined, because all these
lycospores are very similar. On the other hand, we are sure of the identity of the parent plant
of cingulizonate lycospores of the sixth subgroup, i.e. cingulizonate lycospores with a broad
cingulum and a broad zona and microverrucate to verrucate distal surface. These lycospores
were isolated only from Bohemian cones of Lepidostrobus ronnaensis (Bek and Opluštil
2004).
We are aware that the determination of some in situ and dispersed lycospores can be
difficult and subjective, especially those of the third and fourth subgroups thus our division
need not be definitive and perfect.
Acknowledgements. We are grateful to Profs. B.A. Thomas from the University of Lampeter,
D.J. Batten from the University of Aberysthwyth, Wales, U. K., E.L. Zodrow from the
University College of Cape Breton, Sydney, Canada and W.A. DiMichele from the
Smithonian Institute, Washington, USA, Dr. Ch. Wellman from the University of Sheffield,
UK and Prof. M. Streel from the University of Liege, Belgium for correction of the text and
helpful remarks and suggestions. We acknowledge financial support from the Grant Agency
of the Academy of Sciences of the Czech Republic (project A 3013902), the Grant Agency of
the Ministry of Education to the Faculty of Science (Charles University) No. CEZ:
J13/98:113100006 and Research Program (AVOZ30130516) of the Institute of Geology,
Academy of Sciences, Prague. J. Trnka from the Faculty of Sciences of the Charles
27
University, Prague is thanked for the help with preparation of photographs of figured
specimens. Special thanks to Mgr. R. Pátová from the National Museum, Prague.
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Smith, A.H.V. and Butterworth, M.A. 1967. Miospores in the coal seams of the Carboniferous
of Great Britain. Spec. Papers in Palaeont., 1, 1-324.
Somers, Y. et al. 1972. Revision du genre Lycospora Schopf, Wilson et Bentall. Les Spores,
5, 9-110.
Taylor, T.N. and Eggert, D. 1968. Petrified plants from the Upper Mississippian of North
America II. Lepidostrobus fayettevillense sp. nov. Am. J. Bot., 55, 306-313.
Thomas, B.A., 1965. Some studies on Carboniferous lycopods. PhD thesis, Reading
University.
30
Thomas, B.A., 1970. A new specimen of Lepidostrobus binneyanus from the Westphalian B
of Yorkshire. Pollen et Spores, 12, 217-234.
Thomas, B.A., 1987. The use of in-situ spores for defining species of dispersed spores. Rev.
Palaeobot. Palynol., 51, 227-233.
Thomas, B.A., 1988. The fine structure of the Carboniferous lycophyte microspore Lycospora
perforata Bharadwaj and Venkatachala. Pollen et Spores, 30, 81-88.
Thomas, B.A. and Brack-Hanes, S.D., 1984. A new approach to family groupings in the
lycophytes. Taxon, 33, 247-255.
Thomas, B.A. and Dytko, A., 1980. Lepidostrobus haslingdenensis: a new species from the
Lancashire Millstone Grit. Geol. J., 15, 137-142.
Willard, D.A., 1989a. Source plants for Carboniferous microspores: Lycospora from
permineralized Lepidostrobus. Am. J. Bot., 76, 820-826.
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Bot., 76, 1429-1440.
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Roy. Soc. London, Phil. Trans. 184B, 1-38.
Explanation of plates
PLATE I
1. Lepidostrobus kohoutii, sp. nov., holotype E 6112, Mine Ronna in Kladno, the KladnoRakovník Basin, roof shale of the Upper Radnice Seam, Radnice Member (lower Bolsovian).
 0.75.
2-7. Spores isolated from Lepidostrobus kohoutii, sp. nov., holotype E 6112, and comparable
with the dispersed species Lycospora uzunmehmedii Artűz, 1957. Fig. 4. Tetrad of
microspores. All  500. Notice wide and often perforated cingulum.
8. Lepidostrobus cf. haslingdenensis Thomas and Dytko, 1980, E 5587, Tuchlovice Mine near
Kladno, the Kladno-Rakovník Basin, roof shale of the Upper Radnice Seam, Radnice Member
(lower Bolsovian).  1.
9. Densely microspinate distal surface of spore isolated from Lepidostrobus cf.
haslingdenensis Thomas and Dytko, 1980, E 5587, and comparable with the dispersed species
Lycospora uber Artűz 1957, SEM.  1600.
10-13. Spores isolated from Lepidostrobus cf. haslingdenensis Thomas and Dytko, 1980, E
5587, and comparable with the dispersed species Lycospora uber Artűz 1957. All  500.
31
14. Lepidostrobus sp. A, E 3476, Kounov near Rakovník, Otruby Member, Slaný Formation
(Stephanian B), the Kladno-Rakovník Basin, roof shale of the Kounov Seam. General view on
the longitudinally split fragment of the strobilus fossilised by pyrite.  1.
15. Lateral view of spore isolated from Lepidostrobus sp. A, E 3476, and comparable with the
dispersed species Lycospora microgranulata Bharadwaj, 1957, SEM.  1600.
16., 18. Tetrad of spores isolated from Lepidostrobus sp. A, E 3476, and comparable with the
dispersed species Lycospora microgranulata Bharadwaj, 1957. All  500.
17. Proximal surface of spore isolated from Lepidostrobus sp. A, E 3476, and comparable
with the dispersed species Lycospora microgranulata Bharadwaj, 1957, SEM.  800. Notice
the perforation of the zona.
19. Two spores isolated from Lepidostrobus sp. A, E 3476, and comparable with the dispersed
species Lycospora microgranulata Bharadwaj, 1957.  500.
20. Three spores isolated from Lepidostrobus sp. A, E 3476, and comparable with the
dispersed species Lycospora microgranulata Bharadwaj, 1957.  500.
PLATE II
1-5. Spores isolated from Lepidostrobus sp. A, E 3476, and comparable with the dispersed
species Lycospora microgranulata Bharadwaj, 1957. Figs. 1 and 5 showing lateral views and
the differences of the number of sculpture elements on the proximal and distal surface. All 
500.
6. Lepidostrobus sp. B, E 3542, Ronna Mine in Kladno, Radnice Member, Kladno Formation
(lower Bolsovian), tonstein “velká opuka” intercalated in the Upper Radnice Seam.  1.1.
7. Proximal surface of spore isolated from Lepidostrobus sp. B, E 3542, and named as
Lycospora cf. microgranulata Bharadwaj, 1957. SEM.  1700.
8-12. Spores isolated from Lepidostrobus sp. B, E 3542, and named as Lycospora cf. subjuga
Bharadwaj, 1957, SEM. All  500. Figs. 11. and 12. show lateral views and the sculpture of
the proximal and the distal surface.
13-14. Spores isolated from Lepidostrobus sp. C, E 3531, and comparable with the dispersed
species Lycospora torquifer (Loose) Potonié and Kremp, 1955. All  500.
32
15. Proximal surface of spore isolated from Lepidostrobus sp. C, E 3531, and comparable
with the dispersed species Lycospora torquifer (Loose) Potonié and Kremp, 1955, SEM. 
1300.
16. Lepidostrobus sp. C, E 3531, Lužná Mine near Rakovník, the Kladno-Rakovník Basin,
Radnice Member (lower Bolsovian).  1.
17-18, 20. Spores isolated from Lepidostrobus sp. C, E 3531, and comparable with the
dispersed species Lycospora torquifer (Loose) Potonié and Kremp, 1955. All  500.
19. Proximal surface of spore isolated from Lepidostrobus sp. C, E 3531, and comparable
with the dispersed species Lycospora torquifer (Loose) Potonié and Kremp, 1955, SEM. 
2000.
21-24. Spores isolated from Lepidostrobus sp. D, E 5596, and named as Lycospora cf.
subjuga Bharadwaj, 1957. All  500.
25. Lepidostrobus sp. D, E 5596, Červená hůrka hill in Kralupy nad Vltavou, the KladnoRakovník Basin, roof shale of the Upper Radnice Seam, Radnice Member (lower Bolsovian).
 0.75.
Cones
r (m)
Lepidostrobus kohoutii
15.5(18.1)22.5
Lepidostrobus cf. haslingdenensis 150(196)225
Lepidostrobus sp.A
190(223)250
Lepidostrobus sp. B
170(225)260
Lepidostrobus sp. C
140(163)180
Lepidostrobus sp. D - basal part 165(185)205
- apical part 140(172)205
c (m)
2.0(3.3)4.0
20(26)30
14(22)30
20(26)35
20(27)35
20(26)35
20(26)30
z (m)
3.0(4.5)6.0
20(44)60
15(26)35
20(27)40
14(20)26
15(20)28
10(19)25
In situ lycospores
Sculpture of proximal surface
Laevigate, finely scabrate
Laevigate, finely scabrate
Microgranulate, granulate
Microgranulate, granulate
Microgranulate, granulate
Microgranulate
Sculpture of distal surface
Microspinate, microgranulate
Densely microspinate
Densely microgranulate, granulate
Densely microgranulate, granulate
Microgranulate, granulate
Densely microgranulate
Table 1. The radius (r), cingulum (c) and zona widths (z), sculptures of proximal and distal
surfaces of Bohemian in situ lycospores and their parent cones.
33
Lycospore /
Parent cone
Lycospora perforata /Lepidostrobus barnsleyensis
Lycospora sp./ Lepidostrobus comosus
Lycospora sp./ Lepidostrobus sp. C
Lycospora sp./ Lepidostrobus jacksonii
r (m) c (m) z (m) Sculpture of
proximal surface
14
2.5
3
Microgranulate
17
5
2
Microgranulate
16
2
2
Microgranulate
18
2.5
2.5
Microganulate
Lycospora/ Lepidostrobus sp. D
17
2
1.5
Microgranulate
Lycospora cf. uber/ Lepidostrobus dawsonii
20.5
4.5
4
Laevigate
Lycospora noctuina/ Lepidostrobus haslingdenensis
1645
225
29
Lycospora punctata/ Lepidostrobus cf. squarrosus
1545
34
34
Granulate,
microgranulate
Microgranulate
Lycospora rotunda/ Lepidostrobus sp. A
Lycospora torquifer/ Lepidostrobus praelongus,
L. variabilis
Lycospora noctuina/ Lepidostrobus haslingdenensis
1515
157
56
38
24
37
1713
27
3
Lycospora perforata/ Lepidostrobus binneyanus
Lycospora granulata/ Lepidostrobus ornatus
135
14
2
2.5
4
3
Lycospora punctata/ Lepidostrobus stephanicus
18
2.6
2.1
Lycospora triangulata/ Lepidostrobus nemejcii
18
2.5
1.9
Lycospora cf. uzunmehmedii/ Lepidostrobus thomasii
18
2.9
3.9
Lycospora rotunda/ Lepidostrobus ronnaensis
19
3.9
3.8
Lycospora loganii/ Lepidostrobus obovatus
18
2.7
2
Sculpture of distal
surface
Laevigate
Microgranulate
Microgranulate
Microgranulate,
granulate
Densely
microgranulate
Microgranulate,
microverrucate
Verrucate, rugulate
Densely
microgranulate
Finely granulate
Granulate, rugulate
Verrucate, rugulate, Densely rugulate,
baculate
baculate, verrucate
Scabrate, laevigate Microspinate
Microgranulate
Granulate,
microgranulate
Microspinate,
microgranulate
Microverrucate,
microgranulate
Laevigate, finely
scabrate
Laevigate, finely
scabrate
Microverrucate,
verrucate
Reference
Thomas 1965
Thomas 1965
Hagemann 1966
Thomas 1965
Hagemann 1966
Thomas, Bek and
Opluštil in press
Willard 1989b
Willard 1989b
Willard 1989b
Willard 1989b
Thomas and
Dytko 1980
Laevigate
Thomas 1988
Densely granulate,
Brack-Hanes and
microgranulate
Thomas 1983
Densely microspinate, Bek and Opluštil
microgranulate
(2004)
Densely
Bek and Opluštil
microverrucate,
2004
microgranulate
Microgranulate
Bek and Opluštil
2004
Microverrucate,
Bek and Opluštil
verrucate
2004
Microverrucate
Bek and Opluštil
2004
Table 2. The radius (r), cingulum (c) and zona (z) widths and sculptures of proximal and distal
surfaces of selected compressed in situ lycospores and their parent cones.
Lycospore /
r (m) C (m) z (m)
Parent cone
4
6
Lycospora sp./Lepidostrobus oldhamius (associated with 17
Lepidofloyos harcourtii)
Lycospora cf. perforata/ Lepidostrobus fayettevillense
23
3
4
Sculpture of
proximal surface
laevigate
Sculpture of distal
surface
Microspinate,
microgranulate
Reference
Densely
microgranulate
Densely
microgranulate
microgranulate
Taylor and Eggert
1968
Balbach 1966
Lycospora sp./ Lepidostrobus coulterii
13
1.5
2.5
Laevigate, finely
microgranulate
microgranulate
Lycospora sp./ Lepidostrobus minor
13
2
3
microgranulate
Willard 1989a
Leisman and
Rivers 1974
Table 3. The radius (r), cingulum (c) and zona (z) widths and sculptures of proximal and distal
surfaces of selected coal-balls in situ lycospores and their parent cones.
34
Cones
Lepidostrobus kohoutii
Lepidostrobus cf. haslingdenensis
Lepidostrobus sp. A
Lepidostrobus sp. B
Lepidostrobus sp. C
Lepidostrobus sp. D
Width of the
cone (mm)
35
45
26-30
43
38-48
20
Width of the
axis (mm)
4
4
7
2.5-3
8.5
-
Length of
pedicels (mm)
Length of
distal laminae
(mm)
Estimated length of
the cone (mm)
17
11
10
9
19
-
23
10-11
12-13
-
150-250
300-500
?
?
?
200-400
Table 4. Measurements of Czech Lepidostrobus species described herein.