Foss. Rec., 19, 1–9, 2016
www.foss-rec.net/19/1/2016/
doi:10.5194/fr-19-1-2016
© Author(s) 2016. CC Attribution 3.0 License.
Two new species of megasporangiate Sigillariostrobus Schimper
(Sigillariostrobaceae) fructifications from the British Coal Measures
B. A. Thomas1 and L. J. Seyfullah2
1Institute of Biological, Environmental and Rural Sciences, Aberystwyth University, Penglais, Aberystwyth, Ceredigion, UK
2Department of Geobiology, Georg-August-Universität Göttingen, 37077 Göttingen, Germany
Correspondence to: L. J. Seyfullah (leyla.seyfullah@geo.uni-goettingen.de)
Received: 12 August 2015 – Revised: 2 November 2015 – Accepted: 3 November 2015 – Published: 16 November 2015
Abstract. The arborescent lycophyte group known as the sig-
illarians bore pedunculate fructifications, including Sigillar-
iostrobus Schimper, which has a known propensity to disag-
gregate, making comparisons of cones and in situ spores of-
ten difficult if not impossible. Sigillariostrobus is monospo-
rangiate, containing either megaspores or microspores. Two
new species of Sigillariostrobus with megaspores are de-
scribed from the British Coal Measures. Two cones of
Sigillariostrobus saltwellensis sp. nov. are described from
Langsettian–Duckmantian strata of Great Britain, contain-
ing in situ Laevigatisporites glabratus (Zerndt) Potonié and
Kremp spores, making this the first British Sigillariostrobus
species described containing such spores. Sigillariostrobus
barkeri sp. nov. is given here to the previously described
cone with in situ Tuberculatisporites brevispiculus (Schopf)
Potonié and Kremp spores. This is the first Sigillariostrobus
cone with this megaspore species in situ, and thus shows
that T. brevispiculus and T. mamillarius (Bartlett) Potonié
and Kremp are not conspecific as others have previously sug-
gested.
1 Introduction
Sigillarian trees are one of the distinctive groups of Car-
boniferous arborescent lycophytes that are not as tall, or as
branched, as Lepidodendron Sternberg. Arnold (1947) men-
tions Sigillaria reniformis to be 18 ft (ca. 6 m) tall and 6 ft (ca.
2 m) in diameter at the base, tapering to 1 foot (30.5 cm) at
the top. This description suggests that the stem was incom-
plete. Balfour (1872) states that they can be 40–50 ft (12–
15 m) tall and 5 ft (ca. 1.5 m) in diameter. Balfour also gives
measurements of 20 stems as 10–20 ft (3–6 m) high and 1–
3 ft (30.5 cm–1 m) in diameter, standing erect in alternating
strata of shales and sandstones, near Morpeth in Northum-
berland (UK). They bear cones (fructifications) on peduncles
off the main stem or branches. The first figures of a fruc-
tification belonging to Sigillaria Brongniart were given by
Goldenberg (1855, pl. 4, Fig. 3; 1857, pl. 10, Figs. 1, 2), but
he gave no name to them. Schimper (1870, pl. 47, Figs. 12–
14) refigured some of Goldenberg’s illustrations and gave
them the name Sigillariostrobus but did not propose a holo-
type. Andrews (1970) included Sigillariostrobus (Schimper)
Geinitz 1873 in the index that he published because the first
specific name S. bifidus was given by Geinitz. However, Se-
ward (1910) and Chaloner (1967) believed S. bifidus to be a
conifer cone belonging to the genus Gomphostrobus Marion,
1890. For this reason Chaloner (1967) took S. goldenbergi of
Feistmantel (1876) as the type species and gave the generic
authorship as Sigillariostrobus (Schimper 1870) Feistman-
tel, 1876, non Geinitz. Feistmantel (1876) had merely given
the specific name to specimens figured by Goldenberg (1855)
and Schimper (1870), leaving it to Zeiller (1884) to give the
first clear description and figure of S. goldenbergi. For this
reason Chaloner (1967) proposed the type species authorship
as Sigillariostrobus goldenbergi (Feistmantel) Zeiller, 1884.
Feistmantel’s other species, S. feistmanteli, was first regarded
to be not a cone but a fertile zone and, therefore, is the type of
Sporangiostrobus Bode – a genus which has since been syn-
onymised with Omphalophloios White (Opluštil et al., 2010;
Bek et al., 2015).
In situ spores have become an integral part of describing
fructifications, e.g. Chaloner (1953), Thomas (1970), Brack-
Hanes and Thomas (1983), Bek and Opluštil (1998, 2004),
Thomas and Bek (2014). This leads onto a comparison of the
in situ spores with named genera and species of dispersed
Published by Copernicus Publications on behalf of the Museum für Naturkunde Berlin.
2 B. A. Thomas and L. J. Seyfullah: Two new species of megasporangiate Sigillariostrobus Schimper
spores and then to using this information for palaeoecologi-
cal interpretations.
Although the spores of the type specimens of Sigillar-
iostrobus (S. goldenbergi) are unknown, Sen (1958) de-
scribed the spores of a specimen he identified to be S. gold-
enbergi as Laevigatisporites mamillarius. Examination of
the spore contents of other species has shown the genus to
be made up of monosporangiate cones that contain either
megaspores or microspores. Of the megasporangiate cones,
the megaspores spore walls are generally about 20–30 µm
with triradiate marks that are approximately half the radius of
the spore. If the entire surface of the spore is smooth and usu-
ally with prominent curvaturae, it is of the type referred to the
dispersed spore genus Laevigatisporites (Ibrahim) Potonié
and Kremp. If the spore wall is ornamented with tubercles on
the dorsal surface it would be referred to the dispersed spore
genus Tuberculatisporites (Ibrahim) Potonié and Kremp. The
number of dispersed megaspores species belonging to these
two genera far exceeds the number of known cones, which
must be partially due to the fact that most, if not all, Sigillar-
iostrobus cones fall to pieces by losing their sporophylls and
can even be reduced to a bare axis (Fig. 1).
The rarity of finding two more or less intact cones al-
lows us here to describe a new Sigillariostrobus species
from the British Coal Measures, with spores referable to the
smooth-surfaced dispersed spore genus Laevigatisporites.
We also give a name to another Sigillariostrobus cone
from the British Coal Measures (Bolsovian) that has previ-
ously been described (Thomas, 1980) with spores referable
to the tuberculate-surfaced dispersed spore genus Tubercu-
latisporites.
Megaspores that are trilete and circular in outline, with
an originally concavo-convex shape, and having a complex
exine ornamentation of tubercles, would clearly, if found
isolated, be included within the genus Tuberculatisporites
(Ibrahim) Potonié and Kremp (1954).
There are two species of Tuberculatisporites that con-
cern us in this study: T. mamillarius (Bartlet) Potonié and
Kremp (1955) and T. brevispiculus (Schopf) Potonié and
Kremp (1955). Some authors, including Zerndt (1932), Di-
jkstra (1946, 1955a, 1955b), Arnold (1950) and Winslow
(1959), believed the two species to be conspecific, while Po-
tonié and Kremp (1955), Bhardwaj (1957a, b), Arnold (1961)
and Selosse (1953) kept them separate. Spinner (1964) re-
examined and redefined the genus Tuberculatisporites and
included a list of species in which he distinguished T . mamil-
larius from T. brevispiculus. Following Spinner (1968) the
two differ in their ornamentation. The tubercles are larger in
T. mamillarius, being up to 130 µm tall and 94 µm in basal
diameter on the distal surface, with them up to 28 µm tall
and 25 µm in basal diameter on the proximal contact faces.
In T. brevispiculus they are only 10 µm tall and vary from 25
to 30 and 40 to 45 µm in basal diameter on the distal surface,
while the proximal contact faces are covered in apiculi that
are 15 µm in diameter and less than 5 µm tall.
Figure 1. Sigillariostrobus fructification with only a few basal
sporophylls (lower arrow), having shed most of its sporophylls ex-
posing the upper part of the axis (upper arrow) as illustrated by
Kidston (1897) (scale bar 50 mm). From the Kidston Collection at
the British Geological Survey. Catalogue number: P686530. Repro-
duced with permission of the British Geological Survey ©NERC.
All rights reserved.
2 Materials and methods
The cones were investigated using a Carl Zeiss Stemi 2000
microscope with incident light equipped with a Canon 450D
digital camera. Spore preparations were made by macera-
tion with Schulze solution and cleared using dilute (4 %) am-
monia solution. The resulting spores were slide-mounted in
glycerine jelly as with dispersed spores. These spore prepa-
rations were observed with a transmitted light microscope
(Carl Zeiss Axioscope A1) also equipped with a Canon 450D
digital camera, and will be deposited with the specimens.
There are three cones that form the basis of this paper. One
cone (Pb. 53-C.203, Hunterian Museum, Glasgow) comes
from the southern part of the South Staffordshire coalfield
from Saltwells Pit (now part of the Saltwells National Nature
Reserve). The exact horizon of the specimen is unknown, al-
though Besley (1988) refers to the general horizon as alluvial
coal-bearing facies of the upper Langsettian–Duckmantian,
which represent continuously emergent conditions allowing
the extensive development of swamp soils. A second cone
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B. A. Thomas and L. J. Seyfullah: Two new species of megasporangiate Sigillariostrobus Schimper 3
(Pb. 707, Hunterian Museum, Glasgow) has no record of
where it was collected. The third specimen, no. 1971.I28
from the Barker collection, Sheffield City Museum, was col-
lected from above the Shafton Coal, Brierley Colliery, York-
shire – top of the Similis-Pulchra Zone (Bolsovian).
3 Systematic descriptions
(B. A. Thomas, L. J. Seyfullah)
Family Sigillariostrobaceae Thomas and Brack-Hanes,
1984
Genus Sigillariostrobus (Schimper 1870) Feistmantel, 1876, non
Geinitz E.
Type species: Sigillariostrobus goldenbergi (Feistmantel) Zeiller,
1884
Sigillariostrobus saltwellensis sp. nov.
(Figures 2–3)
Holotype: Specimen Pb. 53-C.203, stored in the Hunterian
Museum, Glasgow.
Type locality: Saltwells Pit, near Cradley Heath, Stafford-
shire, Great Britain.
Type stratum: Alluvial coal-bearing facies – upper
Langsettian–Duckmantian.
Etymology: Named after the locality of the specimen that
we designate as the type.
Diagnosis: Cone at least 86 mm long, ca. 17–20 mm in
diameter, axis 1.2 mm, pedicel ca. 8 mm long, sporangia
7 mm long, sporophyll laminae 12 broad at base, 6.3 mm
long spreading outwards from the cone. Sporangia 7 mm
long, 2.5 mm high. Megaspores trilete, probably originally
plano-convex, average diameter 2000 µm in their equatorial
planes, smooth walls with a thickened area between the arms
of the trilete mark. Lesurae approximately half the spore
radius, ca. 15 µm broad at the centre of the spore.
Description: Both cones are preserved as dark compression
(Figs. 2a–b, 3a–b), although they are broken somewhere
above their bases (Figs. 2a, c, 3a, c), so their complete
length is unknown. The megasporophylls have free tips
approximately 6.3 mm long (Fig. 3d), only slightly diverging
from the sides of the cone. The megasporophylls are not well
preserved (Figs. 2b–c, 3d) and the compression material is
very brittle, so the cuticle could not be prepared. Numerous
obvious large red-brown megaspores are visible to the naked
eye (Figs. 2d, 3e). They all measure approximately 2000 µm
in diameter in their equatorial plane, with lesurae 50 µm
long. The entire surfaces of the spores are smooth (Figs. 2e,
3f).
Comments: A second specimen, Pb. 707, also stored in the
Hunterian Museum, Glasgow, is referred to this species, but
the locality and horizon information for this specimen is not
known.
Discussion: The megaspores clearly fit within the param-
eters of the dispersed spore genus Laevigatisporites and
are closest to the species L. glabratus (Zerndt) Potonié
and Kremp, 1956. We agree with Karczewska (1967) that
the spores described by Zerndt (1930) as Types 9 and 10
should be referred to Laevigatisporites glabratus as a result
of Bochenski’s (1936) description of the spores from his
Sigillariostrobus czarnockii. Bochenski (1936) described the
spores as 440 to 2700 µm in diameter, and underdeveloped
spores which were triangular, thicker than the normal ones
and with strongly marked lesurae. Zerndt (1932) agreed
that Bochenski’s finding of his megaspores of Types 9
and 10 (Zerndt, 1930) in the same cone was considerably
important to the understanding of Triletes glabratus (now
Laevigatisporites glabratus).
We also agree that there are no reliable grounds to dis-
tinguish such species as Laevigatisporites primus Wicher
and Laevigatisporites reinschi Ibrahim, as has been done
by Potonié and Kremp (1955). Erecting the species Sporites
primus, Wicher (1934) assigned only the specific name to
the spores, described by Bennie and Kidston (1886) and
Zerndt (1930) as Triletes Type I Kidston. Wicher (1934) in-
terpreted the species Sporonites reinschi Ibrahim to be iden-
tical to Sporites primus; he erected himself and – contrary to
the law of priority – included Sporonites reinschi Ibrahim in
the synonymy of S. primus Wicher. After the publication of
Bochenski’s work, it became clear that the spores of Triletes
Type I Kidston (= Sporonites reinschi = Sporites primus)
and Triletes glabratus Zerndt belong to the same species.
Triletes glabratus Zerndt, 1930, is the oldest specific name
and, therefore, has specific priority.
In situ spores identified as being identical to Laevi-
gatisporites glabratus (Zerndt) Potonié and Kremp have
been also described from four species of Sigillariostrobus.
They are S. tieghemi Zeiller, 1884 (Schenk 1885, Zeiller
1888), S. quadrangularis (Lesquereux) White, 1903 (Wood,
1957), S. czarnockii Bochenski, 1936, and S. leiosporous Ab-
bott, 1963. The features of these cones together with those of
the British cones are summarised in Table 1.
Both S. czarnockii and S. leiosporous have sporophyll lam-
inae with ciliate edges, which is a feature not observed in
the cones from localities in Britain. S. quadrangularis has its
sporophylls in whorls, which is a most unusual character for
Sigillariostrobus. This again is different from the helical ar-
rangement in the cones from localities in Britain. S. tieghemi
is described as having downward-projecting heels at the dis-
tal end of the pedicel, a feature which is again not shown by
the cones from localities in Britain. Although all these cones
contain megaspores that would be included in the same dis-
persed spore species, Laevigatisporites glabratus, there are
characters of the cones which lead us to the conclusion that
they are all different in some respects. For that reason we ac-
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4 B. A. Thomas and L. J. Seyfullah: Two new species of megasporangiate Sigillariostrobus Schimper
Figure 2. Type specimen of Sigillariostrobus saltwellensis sp. nov. Specimen Pb. 53-C.203, Hunterian Museum, Glasgow. Alluvial coal-
bearing facies – upper Langsettian–Duckmantian. (a) Cone (scale bar 10 mm). (b) Detail of apical portion of the cone (scale bar 5 mm). (c)
Detail of basal portion of the cone (scale bar 5 mm). (d) In situ glabrous megaspores (scale bar 2 mm). (e) Spore referable to Laevigatisporites
glabratus macerated from the cone (scale bar 1000 µm).
cept them all as distinct species and therefore refer the British
specimens to a new species.
The anatomically preserved monosporangiate Mazocar-
pon Benson, 1918 includes two species that contain
spores referable to Laevigatisporites glabratus. They are
M. oedipterum Schopf, from the McLeansboro Group, in
Illinois (Schopf, 1941, late Pennsylvanian (Stephanian A)),
and M. bensonii Pigg, from the Duquesne Coal, Cone-
maugh Group, Ohio (late Pennsylvanian (Stephanian B)).
Pigg (1983) showed the two species to have sufficiently dif-
ferent anatomical features to be separated. Although it is al-
most certain that Sigillariostrobus and Mazocarpon represent
the same type of sigillarian cone but are merely preserved
differently, we hesitate to include our S. saltwellensis within
either of these two species because of the lack of anatomical
evidence and their rather different ages.
Sigillariostrobus barkeri sp. nov
(Figure 4)
Holotype: Specimen number 1971.I28 held in the Barker col-
lection, Sheffield City Museum.
Type locality: Brierly Colliery, Yorkshire, Great Britain.
Type horizon: Shale above the Shafton Coal, top of the
Similis-Pulchra Zone, Bolsovian.
Etymology: Named after the collection where the speci-
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B. A. Thomas and L. J. Seyfullah: Two new species of megasporangiate Sigillariostrobus Schimper 5
Table 1. Morphological comparison of Sigillariostrobus cones with in situ spores referable to Laevigatisporites glabratus.
S. tieghemi
Zeiller, 1884
S. quadrangularis (Les-
quereux) White, 1903
S. czarnockii Bochen-
ski, 1936
S. leiosporous Abbott,
1963
Sigillariostrobus
saltwellensis sp.
nov.
Cone size 160 mm long
25–50 mm
broad
Up to 160 mm long, av-
erage width 11.6 mm
ca. 180 mm long, 17–
25 mm broad
Up to 180 mm long,
20 mm broad
At least 86 mm
long, 17–20 mm
broad
Cone axis 1 mm 7–8 mm 4–7 mm 1.2 mm
Sporophyll
arrangement
Alternating
whorls with
8–10 per whorl
In whorls Helical with sporo-
phylls lying above one
another
Alternating verticils
of 4–5 sporophylls;
3–4 mm between
verticils
Helical
Sporophyll
shape
With prominent
heel
Sporophyll laminae
9 mm long
Sporophylls 20–25 mm
long; upper part
sharply pointed with a
hollowed-out deltoid
shape, margin ciliate.
Pedicels: 5–7 mm long,
1 mm broad at attach-
ment point, widening to
4 mm
Laminae: broadly
triangular, acuminate,
5–8 mm long, 4–6 mm
broad, margin ciliate
Laminae ca.
6.5 mm long,
spreading out-
wards from
cone
Sporangia None observed 4 mm long, 5 mm high,
4 mm broad
4–7 mm long, breadth
2–4.5 mm;
contain three tetrads of
megaspores
4–5 mm long, 2 mm
high, 4–4.5 mm wide;
contains one tetrad of
megaspores
7 mm long,
2.5 mm high
Peduncle Acicular leaves
or bracts
attached to the
upper portion
None observed At least 40 mm long
and 7–8 mm broad,
uppermost 30 mm with
long, triangular and
pointed sterile leaves
25 mm long and 25 mm
wide at the base
At least 65 mm long
and 2–5 mm broad,
covered with minute
stiff spines up to 1 mm
long
None observed
Age and
locality
Age unknown,
Valenciennes,
France
Brazil Formation,
Upper Pottsville,
Duckmantian/
Bolsovian
Indiana, USA
Duckmantian, Upper
Silesian Basin, Poland
Upper Freeport (no.
7) Coal Allegheny,
mid-Pennsylvanian,
Bolsovian/Asturian,
southeastern Ohio,
USA
Upper Langsettian–
Duckmantian
men had been deposited, the Barker collection held at the
Sheffield Museum, Great Britain.
Diagnosis: Cone at least 13 mm in diameter without sporo-
phyll laminae; axis 3 mm in diameter. Sporangia ca. 5.5 mm
long and 2 mm high, megasporangiate. Megaspores trilete,
probably originally plano-convex with greatest diameters of
1400–1800 µm in their equatorial planes. Lesurae one-third
to one-half the spore radius, ca. 15 µm broad. Spore walls are
about 20 µm thick, covered with numerous conical apiculi.
Apiculi on distal walls ca. 10 µm high, varying in basal di-
ameter and in numbers from 300 apiculi, 40–45 µm broad at
the base, to 550 apiculi, 25–30 µm broad at the base. Apiculi
on the contact faces are about 15 µm broad at the base and
less than 5 µm high.
Description: The specimen is a fragment of a cone with lit-
tle remaining except the megaspores arranged in the posi-
tions they must have occupied in the sporangia (Fig. 4a–
b). This is a single Sigillariostrobus cone 13 mm in diam-
eter that lacks sporophyll laminae (Fig. 4a), with an axis
3 mm in diameter. The sporangial remains are ca. 5.5 mm
long and 2 mm high, containing in situ spores, referable to the
dispersed spore genus Tuberculatisporites (Ibrahim) Potonié
and Kremp. Megaspores recovered from the specimen have
a maximum diameter of 1400–1800 µm in their equatorial
planes, with lesurae one-third to one-half of the spore radius,
and are clearly trilete (Fig. 4c) and tuberculate (Fig. 4d). The
spore walls are about 20 µm thick and are covered with nu-
merous conical apiculi (Fig. 4d–e). The size and distribution
of the apiculi vary over the surface of the megaspores, with
the apiculi on the contact faces being smaller than those else-
where on the surface. On the distal walls of the megaspores,
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6 B. A. Thomas and L. J. Seyfullah: Two new species of megasporangiate Sigillariostrobus Schimper
Figure 3. Second cone of Sigillariostrobus saltwellensis sp. nov. Specimen Pb. 707, Hunterian Museum, Glasgow. (a) The cone (scale bar
20 mm). (b) Detail of the upper part of the cone (scale bar 5 mm). (c) Detail of the mid-region of the cone, showing the megasporophyll
impressions at the edge of the cone, often without the cuticle preserved (scale bar 5 mm). (d) Detail of (c), showing the megasporophylls
with fissile compression and scattered megaspores (scale bar 5 mm). (e) Detail of (d), showing details of megaspores within the cone (scale
bar 2 mm). (f) Megaspore referable to Laevigatisporites glabratus macerated from the spore mass shown in (e) (scale bar 2000 µm).
the apiculi are ca. 10 µm high, varying in basal diameter and
in numbers from 300 apiculi, 40–45 µm broad at the base to
550 apiculi, 25–30 µm broad at the base. On the contact faces
of the megaspores, the apiculi are about 15 µm broad at the
base and less than 5 µm high.
Discussion: This Sigillariostrobus cone was originally de-
scribed by Thomas (1980: Figs. 3–6). Although the megas-
pores were closely compared with Tuberculatisporites bre-
vispiculus (Schopf) Potonié and Kremp, which was the first
record of this spore, the cone was not given a specific name.
Spores belonging to the genus Tuberculatisporites, like those
referred to the smooth-walled Laevigatisporites (Zerndt) Po-
tonié and Kremp (1954), have been recovered several times
from sigillarian cones but from no others. Spores referable
to Tuberculatisporites mamillarius have been previously de-
scribed from Sigillariostrobus rhombibracteatus by Bochen-
ski (1936), Chaloner (1953) and Wood (1957), but this is the
only record of in situ spores referable to T. brevispiculus.
As explained above, Tuberculatisporites brevispiculus has
been discussed many times in relation to its possible affin-
ity with T. mamillarius (Bartlett) Potonié and Kremp (1955).
We are following Spinner (1968), who kept them separate.
As Thomas (1980) regarded the present cone as sigillarian
and referred it to Sigillariostrobus, we now consider that it
would be preferable to make it the basis of a new species.
The age of the new cone is also of some importance
since Spinner (1968) gives the stratigraphical range of
T. brevispiculus as generally Asturian and early Stepha-
nian. In contrast, T. mamillarius and Sigillariostrobus rhom-
bibracteatus have been only described from the Langsettian
and Duckmantian. The present cone, in coming from the
early Bolsovian, seems to fall between the two zones men-
tioned by Spinner (1968). Nevertheless, no real problem is
posed by this new record as this extended stratigraphical
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B. A. Thomas and L. J. Seyfullah: Two new species of megasporangiate Sigillariostrobus Schimper 7
Figure 4. Sigillariostrobus barkeri sp. nov. No. 1971.I28, Barker collection, Sheffield City Museum, from the shales above the Shafton Coal,
Brierly Colliery, Yorkshire (top of the Similis-Pulchra Zone, Bolsovian, Moscovian). (a) The remains of the cone showing the arrangement of
the megaspores (scale bar 10 mm). (b) Camera lucida drawing of the cone in (a) (scale bar 10 mm). (c) Macerated in situ megaspore assigned
to Tuberculatisporites mamillarius (Bartlett) Potonié and Kremp (scale bar 500 µm). (d) Contact face of the in situ megaspore (scale bar
150 µm). (e) Apiculi on the distal surface of the in situ megaspore (scale bar 10 µm). Figure previously published by Thomas (1980).
range of T. brevispiculus still does not overlap with the range
of T. mamillarius.
4 Conclusions
Despite the tendency of the sigillarian fructification Sig-
illariostrobus to fall apart, two cones of Sigillariostrobus
saltwellensis sp. nov. are described from the British upper
Langsettian–Duckmantian strata, containing spores referable
to the dispersed spore species Laevigatisporites glabratus,
and these have been compared to the four other Sigillar-
iostrobus cones that contain L. glabratus spores. This is also
the first Sigillariostrobus cone species described containing
L. glabratus spores from localities in Great Britain.
Sigillariostrobus barkeri sp. nov. of Bolsovian age is given
to the previously described but unnamed cone with in situ
spores referable to T. brevispiculus. This is the first cone with
this megaspore species in situ, and thus shows that T. bre-
vispiculus and T. mamillarius are not conspecific.
Author contributions. B. A. Thomas and L. J. Seyfullah prepared
the specimens, made the descriptions and comparisons, and pre-
pared the manuscript together.
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8 B. A. Thomas and L. J. Seyfullah: Two new species of megasporangiate Sigillariostrobus Schimper
Acknowledgements. We thank the Hunterian Museum for the
opportunity to examine their specimens, and also both Jirˇi Bek and
Stanislav Opluštil for their helpful comments on the manuscript.
We gratefully acknowledge support by the German Research Foun-
dation and the Open Access Publication Funds of the Göttingen
University. L. J. Seyfullah thanks the University of Göttingen for
their continuing support.
Edited by: D. Korn
Reviewed by: J. Bek and S. Opluštil
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