Pteridospermophyta in the Mesozoic floras of Abyek and Jirande, Western Alborz, Iran

Abstract

This paper deals with the Mesozoic seed ferns from the Shemshak Group from two localities in the Western Alborz Mountains, Northern Iran (i.e., the Abyek area in Qazvin Province and the Jirande area in Gilan Province). The materials were obtained from two coal-bearing horizons in the Shemshak Group, Late Triassic-Middle Jurassic in age. Five species from the Abyek and two species from the Jirande area, belonging to the phylum Pteridospermophyta, were identified. Vegetative and reproductive organs of Caytoniales (i.e., Sagenopteris sp. A and Sagenopteris sp. B leaves, Caytonia sp. ovuliferous organ, and Caytonanthus arberi Thomas (Harris) emend. Harris, 1964 microsporangiate organ, as well as two form-species of incertae sedis seed ferns, Pachypteris lanceolata Brongniart, 1828 and Ptilozamites tenuis Ôishi 1932 emend. Ôishi and Huzioka, 1938) are described in the present paper. This paper discusses the morphological characters of these species, together with discussions on their previous palaeobotanical descriptions, as well as the stratigraphic and palaeogeographical distribution of these taxa. This is the first record of Caytonanthus arberi in Asia and Ptilozamites tenuis from Iran. Pachypteris lanceolata is described for the first time in the Western Alborz.

Full text

Vol.:(0123456789)
1 3
Arabian Journal of Geosciences (2022) 15:1573
https://doi.org/10.1007/s12517-022-10691-5
ORIGINAL PAPER
Pteridospermophyta intheMesozoic floras ofAbyek andJirande,
Western Alborz, Iran
SeyedKamyarKamran1· MasoomehSohrabiMollayouse2 · MehranArian1· MajidMirzaie‑Ataabadi3· AliSolgi1
Received: 13 December 2021 / Accepted: 12 August 2022
© Saudi Society for Geosciences 2022
Abstract
This paper deals with the Mesozoic seed ferns from the Shemshak Group from two localities in the Western Alborz Moun-
tains, Northern Iran (i.e., the Abyek area in Qazvin Province and the Jirande area in Gilan Province). The materials were
obtained from two coal-bearing horizons in the Shemshak Group, Late Triassic-Middle Jurassic in age. Five species from
the Abyek and two species from the Jirande area, belonging to the phylum Pteridospermophyta, were identified. Vegetative
and reproductive organs of Caytoniales (i.e., Sagenopteris sp. A and Sagenopteris sp. B leaves, Caytonia sp. ovuliferous
organ, and Caytonanthus arberi Thomas (Harris) emend. Harris, 1964 microsporangiate organ, as well as two form-species
of incertae sedis seed ferns, Pachypteris lanceolata Brongniart, 1828 and Ptilozamites tenuis Ôishi 1932 emend. Ôishi
and Huzioka, 1938) are described in the present paper. This paper discusses the morphological characters of these species,
together with discussions on their previous palaeobotanical descriptions, as well as the stratigraphic and palaeogeographical
distribution of these taxa. This is the first record of Caytonanthus arberi in Asia and Ptilozamites tenuis from Iran. Pachyp-
teris lanceolata is described for the first time in the Western Alborz.
Keywords Seed ferns· Shemshak Group· Plant macrofossils· Mesozoic· Caytoniales
Introduction
The Shemshak Group is an Upper Triassic-Middle Jurassic
unit exposed across the Alborz Mountains and Central Iran
(Fürsich etal. 2009). This Group contains rich and diver-
sified Triassic and Jurassic plant macrofossils throughout
the Alborz mountain range, studied by Göppert (1861),
Stur (1886), Schenk (1887), Krasser (1891), Zeiller (1905),
Potonié (1933), Boureau etal. (1950), Lorenz (1964), Kilp-
per (1964, 1968, 1971, 1975), Barnard (1965, 1967), Alavi
and Barale (1970); Barnard and Miller (1976), Sadovnikov
(1976, 1980), Corsin and Stampfli (1977), Fakhr (1977),
Schweitzer (1977, 1978), Schweitzer and Kirchner (1995,
1996, 1998, 2003), Schweitzer etal. (1997, 2000, 2009),
Vaez-Javadi and Ghavidel-Syooki (2002), Vaez-Javadi
(2006, 2011), Vaez-Javadi and Allameh (2015), and Farahi-
manesh etal. 2021. However, few studies were performed on
the Mesozoic macrofloras of the Western Alborz; especially,
little attention was given to the seed ferns in this region.
Stur (1886), Schenk (1887), and Krasser (1891) have men-
tioned the presence of seed ferns in the Western Alborz (see
Table2). The first taxonomical work on the Western Alborz
plant macro-remains was carried out by Fakhr (1977),
who described two seed fern species, namely Ptilozamites
abiekensis and Ptilozamites (Ctenozamites) iranica. Subse-
quently, Sadovnikov (1983) described one seed fern species
(Amdrupia stenodonta) in the Western Alborz. Schweitzer
and Kirchner (1998) have described six species of seed ferns
from five areas in the Western Alborz, namely Scytophyl-
lum waehneri (= Scytophyllum persicum, see Kustatscher
etal. 2011) from Aghusbin, Kaman, Kalashtar, and Ekrasar;
Sagenopteris cf. nilssoniana from Zangerud; Anthrophyop-
sis crassinervis from Aghusbin, Kalashtar, and Ekrasar;
Ptilozamites (Ctenozamites) cycadea from Zangerud;
Responsible Editor: Attila Ciner
* Masoomeh Sohrabi Mollayousefi
m.mollayousefi@iauctb.ac.ir
1 Department ofEarth Sciences, Science andResearch
Branch, Islamic Azad University (IAU), Tehran, Iran
2 Department ofPetroleum, Mining andMaterials
Engineering, Islamic Azad University, Central Tehran
Branch, Tehran, Iran
3 Department ofGeology, Faculty ofScience, University
ofZanjan, Zanjan, Iran

Arab J Geosci (2022) 15:1573
1 3
1573 Page 2 of 14
Ptilozamites (Ctenozamites) iranica from Aghusbin; and
Ptilozamites (Ctenozamites) caspiensis from Ekrasar. The
present work is based on two newly discovered plant-bearing
horizons of the Shemshak Group in the Abyek and Jirande
areas, which improves our understanding of the Mesozoic
floral composition in the Western Alborz Mountains.
Material andmethods
Samples were collected from two coal-bearing horizons of
the Shemshak Group in the Abyek and the Jirande area. The
Abyek section is located at 60km SE of Qazvin, and the
Jirande section is situated between Sangroud and Pakdeh vil-
lages, 119km NW of Qazvin, Western Alborz, Iran (Fig.1).
Samples are preserved as impressions, yielding no cuticle.
The samples are stored in the Earth Sciences Department
of Islamic Azad University, Science and Research branch,
labeled under successive numbers (1–109) in Abyek and
(1–93) in Jirande, preceded by the KA prefix for Abyek and
KJ for Jirande samples. The samples were studied and pho-
tographed using 20 × magnification binocular microscopy
and Panasonic FZ200 Digital Camera. Oblique lighting was
used to enhance contrast and fine details. The systematic
classification followed Taylor etal. 2009.
Systematic palaeobotany
Phylum: Pteridospermophyta, Oliver and Scott, 1904
Order: Caytoniales Gothan, 1932.
Family: Caytoniaceae Thomas, 1925
Genus: Sagenopteris Presl, emend. Harris, 1964
Type species: Sagenopteris nilssoniana (Brongniart)
Ward, 1900
Sagenopteris sp. A
Figure2a–c
Material: KA 41–44
Description:
Leaves are palmately compound, with a petiole bearing
four leaflets. The petiole is more than 10.5mm long and
1.5mm wide, possessing a central furrow. Leaflets are long-
lanceolate with entire margin, 36mm long, and 4 to 8mm
wide at the broadest part, tapering gradually to an acute-
acuminate apex. Leaflets are five to six times longer than
wide. The midvein extends to the apex, giving rise to lateral
veins at an angle of 25°. The vein meshes are 0.5mm wide.
Remarks:
Four specimens assigned to Sagenopteris sp. A were sam-
pled in the Abyek area, two of them associated with Cay-
tonia sp. ovuliferous organ. The morphological characters
of specimens described above are similar to Sagenopteris
phillipsi (Brongniart) Presl, 1838 in having long-lanceolate
leaflets (five to six times longer than wide), tapering gradu-
ally to an acute apex, and a midvein extending to the end of
lamina; however, due to lack of cuticle evidence, they are
described here as Sagenopteris sp. A.
Comparisons:
Following are comparisons with most similar species:
Sagenopteris nilssoniana (Brongniart) Ward, 1900 differs
from the Abyek material in having ovate/obovate to lanceolate
leaflets (with a length/width ratio of less than 5:1), an entire to
undulating margin, and an acute to obtuse apex versus more
Fig. 1 Location map of the studied areas in the Western Alborz, Iran

Arab J Geosci (2022) 15:1573
1 3
Page 3 of 14 1573
elongated leaflets with entire margins and acute-acuminate
apices in the Abyek material. Moreover, the midvein usually
reaches 2/3 of the leaflet’s length in S. nilssoniana, while in
the Abyek Sagenopteris, the midvein extends to the apex.
Sagenopteris colpodes Harris, 1940 is an aggregate of
two similar species, including smaller and larger forms (Har-
ris 1964); so far, only the smaller form has been reported
in Iran (Barnard and Miller 1976; Schweitzer and Kirchner
1998), which differs from the present material in having
broadly ovate to broadly obovate leaflets two times longer
than wide with an obtuse apex (vs. Sagenopteris leaflets
in the Abyek material with long-lanceolate leaflets, up to
six times longer than wide, and an acute-acuminate apex).
Moreover, in the Abyek material, the midvein extends to the
apex, while in S. colpodes, the midvein disappears consider-
ably below the apex.
Sagenopteris iranica Fakhr, 1977, from the Eastern
Alborz, is similar to the present material in the leaflet out-
line (entire margin, long-lanceolate with an acute-acumi-
nate apex) and the venation pattern (a distinct midvein that
extends to the apex and gives off lateral vein at an angle of
20°) but differs in having much larger leaflets (145 × 25mm).
S. iranica was established based on the gross -morphology
of a single detached leaflet; this species can be considered
Fig. 2 a–c Sagenopteris sp. A,
Abyek area

Arab J Geosci (2022) 15:1573
1 3
1573 Page 4 of 14
as a synonym of S. phillipsi, as proposed by Schweitzer and
Kirchner (1998).
Sagenopteris nariwaensis Huzioka, 1970, from the Upper
Triassic of Japan, is similar to the Abyek Sagenopteris in
leaflets' outline (long-lanceolate leaflets gradually tapering
at both ends) and venation (prominent midvein giving off
lateral veins at an acute angle), but its leaf is composed by
six leaflets, which is not observed in our material.
Distribution:
So far, four species of the genus Sagenopteris have
already been described from Iran: S. phillipsi (Sadovnikov
1976, 1980 from Karmozd in the Central Alborz and Taza-
reh in the Eastern Alborz; Schweitzer and Kirchner 1998
from Zirab in the Central Alborz; Vassiliev 1984 from Bab-
nizu in Kerman basin, Central Iran), S. nilssoniana (Fakhr
1977 from Shemshak in the Central Alborz; Schweitzer and
Kirchner 1998 from Zangerud in the Western Alborz), S.
colpodes (Barnard and Miller 1976 from Khatunbargah in
the Central Alborz; Schweitzer and Kirchner 1998 from
Zirab in the Central Alborz), and S. iranica (Fakhr 1977
from Ferizi in the Eastern Alborz). The determinations of
all these materials were based on gross morphology; among
them, only the S. colpodes material described by Barnard
and Miller (1976) have yielded cuticle fragments (see Bar-
nard and Miller 1976, pp. 45–47).
Sagenopteris sp. B
Figure3a, b
Material: KJ 34, 72–75, 80, 81
Description:
Several detached leaflets were found. They are variable in
size and shape. Smaller leaflets are ovate, up to18 mm long,
and 4–8mm wide in the middle of leaflets. Larger leaflets
are lanceolate in outline and 12–15mm wide in the middle
of leaflets. No complete leaflets were preserved. The longest
fragment is up to 35mm long. Leaflets are 2.3 to 4 times as
long as broad, entire-margined, and attenuated to an acute
apex. The midvein extends to the leaflet apex, giving off
lateral anastomosing veins at 20°. The width of vein meshes
varies between 0.5 and 1mm.
Remarks:
Seven specimens of Sagenopteris sp. B were sampled in
the Jirande area. The leaflets of Sagenopteris sp. B are most
similar to those of S. nilssoniana (i.e., ovate to lanceolate,
two to four times longer than wide, with an acute apex).
The Jirande specimens correspond well with records of
S. nilssoniana described before from Iran (Fakhr 1977; Sch-
weitzer and Kirchner 1998) in the leaf size and the width of
vein meshes; however, due to the absence of cuticle, these
materials are described here as Sagenopteris sp. B.
Comparisons:
The Jirande material (Sagenopteris sp. B) differs from the
Abyek material (Sagenopteris sp. A) in having less elongated
leaflets (ovate to lanceolate in shape) versus long-lanceolate
leaflets of the Abyek Sagenopteris. Moreover, leaflets in the
Jirande material taper more abruptly to the apex.
The Jirande Sagenopteris resembles some leaflets of S.
trapialensis Elgoriaga etal. 2019 from the Lower Jurassic of
Patagonia in the Southern Hemisphere in having larger lan-
ceolate and smaller ovate leaflets as well as a similar length/
width leaflet ratio (2.5:1 in average). However, it differs from
S. trapialensis in having leaflets with entire margins (vs.
entire to slightly undulate margins of S. trapialensis), acute
apex (vs. obtuse to subacute apices of S. trapialensis), mid-
veins extending to the apex (vs. midveins extended up to
3–4/5 of leaflet length in S. trapialensis), and larger vein
meshes (0.5 to 1mm vs. 0.3 to 0.5mm in S. trapialensis).
Morphological characters of selected Sagenopteris spe-
cies are compared with the studied materials in Table1.
Genus: Caytonia Thomas emend Harris 1940, nom. Cons.
(Herendeen 2011).
Type species: Caytonia sewardi Thomas, 1925
Caytonia sp.
Figures3c–e and 4a, b
Material: KA 41, 42, 45–48
Description:
The main axis is 1mm wide, bearing two longitudinal
grooves, and dichotomized twice. Each terminal axis (stalk)
expands at the distal part, ending in an orthotropous hemispheri-
cal cupule. Fourteen longitudinal furrows traverse the cupule
surface on each side. Compressed cupules are 3mm in diameter.
Remarks:
Six specimens of this species were collected in the Abyek
area. The Abyek Caytonia is characterized by grooved
axes, the hemispherical shape and orthotropous position
of cupules, and longitudinal furrows on the cupule surface
(Fig.3c–e, arrowhead).
Comparisons:
The encountered specimens are most similar to Caytonia
nathorsti in having an axis with longitudinal grooves and rounded
cupules with longitudinal furrows (Harris 1964, p. 23, Fig.10 A, C,
I), but it has slightly smaller cupules (i.e., 3mm wide vs. 4mm in
C. nathorsti, see Harris 1964). Our material differs from C. thomasi
in having round cupules versus elliptical ones and from C. sewardi
by its grooved axis in contrast to the smooth axis of the latter.
Genus Caytonanthus Harris emend. Harris, 1964
Type species: Caytonanthus arberi Thomas (Harris)
emend. Harris, 1964
Caytonanthus arberi (Thomas) Harris emend. Harris, 1964
Selected synonyms:
1964 Caytonanthus arberi (Thomas) Harris; Harris,
p. 15; pl. 1. Figures11, 12, 16–19; pl. 2, figs.2, 3; text-
figs.7A, B, D, E; 8C-F, I.
1994 Caytonanthus arberi (Thomas) Harris; Osborn, p.
1520, figs.1, 2.
Figure4c
Material. KA 44; KJ 47, 75, 80, 81

Arab J Geosci (2022) 15:1573
1 3
Page 5 of 14 1573
Description:
The microsporophyll rachis (main axis) is 0.6mm wide
and more than 1.2mm long, dichotomizing twice. The first-
order branches are 1.5mm long and 0.4mm wide, arranged at
an angle of 45° to each other. The second-order branches are
0.3mm wide, bearing a distinct tubercle near the distal end.
Each secondary branch bears an elongated elliptical synangium
(or anther) with an acute apex. The attached synangia are 3mm
long and 1mm wide, while detached ones are 3 × 1mm to
5.5 × 2mm. Two pollen sacs are preserved on each synangium.
Pollen sacs are longitudinally striated and attached to each other
at both ends.
Remarks:
Five specimens of this species were sampled, including four
samples from the Jirande and one from the Abyek area. These
specimens are attributed to Caytonanthus arberi (Thomas)
Harris 1964 based on elongated elliptical synangia with acute
apices, the longitudinally striated surface of pollen sacs, and
the fusion of adjacent pollen sacs at the base and apex. In the
present material, the attached synangia are 3 × 1mm in size, the
same size as the emended diagnosis of C. arberi in Harris (1964)
based on Yorkshire specimens; likewise, the detached synangia
reach up to 5.5 × 2mm in the present material, similar to addi-
tional material from Yorkshire (5.1 × 1.8mm in Osborn 1994).
Fig. 3 a, b Sagenopteris sp. B,
Jirande area; c-e Caytonia sp.
with cupules ornamented by
longitudinal furrows (arrow-
heads), Abyek area

Arab J Geosci (2022) 15:1573
1 3
1573 Page 6 of 14
The presence of a tubercle near the distal end of ultimate
branches is distinguishable in other Caytonanthus species such as
C. oncodes Harris (1941), C. Kochi Harris (1937), C. rewaensis
Prakash and Das (2017), and Caytonanthus sp. described by Elgor-
riaga etal. (2019), although less distinct than the present material.
Comparisons:
Following are comparisons with most similar species:
Caytonanthus kochi Harris, 1937, from the Lower Jurassic
of Greenland, is similar to the Jirande Caytonanthus in the axis
branching pattern (dichotomizing twice), the width of the first-
order branch (0.4mm), and elongated synangia with acute apices
but differs in having slightly narrower synangia (0.8–1.2mm vs.
1–2mm in the present material) with abruptly contracted bases.
Caytonanthus oncodes Harris, 1964 is also somewhat
similar to the present material, but C. oncodes has smaller
synangia with less length/width ratio and rounded synangia
apex versus acute apex in the herein described material.
Elgorriaga etal. (2019) have reported numerous detached
synangia alongside two samples of branched axes with
attached synangia, from the Lower Jurassic of Patagonia in
southern Argentina, under the name Caytonanthus sp. The
Patagonia Caytonanthus is similar to the Jirande material in
(i) the number of axes dichotomizing (branching twice), (ii)
elliptical shape of synangia, (iii) the ensiform outline and the
longitudinally striated surface of pollen sacs, (iv) pollen sacs
length/width ratio (six times as long as wide), and (v) the size
of pollen sacs (i.e., the average size in the Patagonia material
is 5.1 × 0.9mm versus the range of 3–5.5 × 0.5–1mm in the
Jirande material). However, the Patagonia material differs in
having pollen sacs fused only at the base (vs. pollen sacs fused
at both ends in the present material), rounded to sub-acute
synangia apices (vs. acute apex in the current material), and
the presence of a dorsal keel on each pollen sac.
Distribution:
So far, Caytonanthus arberi has been recorded from
the Jurassic of Yorkshire, England (Thomas 1925; Harris
1941, 1964; Osborn 1994). This is the first record of the
genus Caytonanthus from the Middle East and the species
C. arberi in Asia.
Order: incertae sedis
Family: incertae sedis
Genus: Pachypteris Brongniart 1828 emend. Harris 1964
Pachypteris lanceolata Brongniart, 1828
Figure4d
Selected synonyms:
1965 Pachypteris shemshakensis Barnard, p. 1134–1139; pl.
96, figs.3, 4; pl. 97, Fig.5;pl. 98, Fig.3; text-figs.2A-B, 3 A.
1967 Pachypteris lanceolata Brongniart; Delle, p. 84–85;
pl. 11, figs.4–14.
1976 Pachypteris lanceolata (Brongniart) Harris; Bar-
nard and Miller, p. 41–45; pl. 1, figs.10–13; pl. 2, Fig.1;
text-Fig.3 A-H; text-Fig.4A–E.
1998 Pachypteris lanceolata Brongniart; Schweitzer and
Kirchner, p. 26–28; pl. 5, figs.1-5; text-Fig.5, 6.
1998 Pachypteris shemshakensis Barnard; Schweitzer and
Kirchner, p. 28–29; pl. 6, figs.1–7.
2008 Pachypteris cf. indica (Oldham and Morris) Bose
and Roy; Birkenmajer and Ociepa, p. 47–49, Fig.19 B.
Material. KA 44
Description:
Two pinnae are preserved. Pinna fragments are up to 28mm
long and 14.5mm wide. The pinna rachis is 0.4 to 0.9mm
Table 1 Comparison of Sagenopteris sp. A and Sagenopteris sp.
B with relevant Sagenopteris species, according to the literature by
Brongniart (1825), Ward (1900), Harris (1940), Huzioka (1970),
Barnard and Miller (1976), Fakhr (1977), Schweitzer and Kirchner
(1998), Elgorriaga etal. (2019)
Species Leaflet outline Length/width ratio Margin Apex Midvein
extension
Vein mesh width
Sagenopteris sp. A
(This study)
Long lanceolate 5–6:1 Entire Acute-acuminate Up to the apex 0.5mm
Sagenopteris phil-
lipsi Long lanceolate 5–10:1 Entire Acute-acuminate Up to the apex 0.5mm
Sagenopteris
iranica Long lanceolate 6:1 Entire Acute-acuminate Up to the apex Up to 1mm
Sagenopteris nari-
waensis Long lanceolate 6:1 Entire Acute-acuminate Below the apex Up to 1mm
Sagenopteris col-
podes Ovate-Obovate 2:1 Entire Obtuse Below the apex 0.75–1mm
Sagenopteris sp. B
(This study)
Ovate-Lanceolate 2–4:1 Entire Acute Up to the apex 0.5–1mm
Sagenopteris nilsso-
niana Ovate-Obovate-
Lanceolate
< 5:1 Entire-undulate Acute-obtuse Up to 90% of leaflet
length
0.2–0.7mm
Sagenopteris trapi-
alensis Ovate-Variable 1.3–3.8:1 Entire-slightly
undulate
Obtuse-subacute Up to 60–70% of
leaflet length
0.3–0.6mm

Arab J Geosci (2022) 15:1573
1 3
Page 7 of 14 1573
wide and ornamented by fine longitudinal striae. Pinnules are
inserted oppositely to sub-oppositely, at 1-mm intervals and at
an angle of 30°–45°. Pinnules are 5 to 10mm long and 1.8 to
2mm wide, lanceolate to ovate, decurrent on the basiscopic
margin, and slightly constricted on the acroscopic side. Pinnule
margins are entire, gradually converging to a rounded apex.
The venation pattern is not visible, but a faint impression of
veins is recognizable, including a midvein extended to the apex
and lateral veins arising from it at low angles.
Remarks:
One specimen of this species was sampled from the
Abyek area. The specimen is closely allied to P. lanceolata
Brongniart, described by Harris (1964) based on Yorkshire
specimens, including the holotype. The diagnostic characters
of the species (see Harris 1964, p.137) agree perfectly with
the Abyek material in (i) the pinnule attachment angle (i.e.,
30° vs. 30°–45° in the Abyek material), (ii) the pinnule out-
line (lanceolate to ovate shape, decurrent basiscopic margin,
and rounded to obtuse apex), and (iii) the pinnule dimension
(2–15 × 1.5–5mm vs. 5–10 × 1.8–2 in the Abyek material).
Comparisons:
The following species show close resemblance with
Pachypteris lanceolata:
Pachypteris shemshakensis Barnard, 1965, from the
Lower Jurassic of Dorud in the Central Alborz, presents the
same characters as P. lanceolata (i.e., pinnules with entire
Fig. 4 a, b Caytonia sp., Abyek
area c Caytonanthus arberi,
Jirande area; d Pachypteris
lanceolata, Abyek area

Arab J Geosci (2022) 15:1573
1 3
1573 Page 8 of 14
margin, ovate to lanceolate shape, 6–15 × 3–5mm in size, as
well as decurrent basiscopic margin and rounded to obtuse
apex). The lowest pinnule on the basiscopic side is partly
attached to the main rachis in both species. P. shemshakensis
was established by Barnard (1965) based on gross morphol-
ogy due to the presence of two pinnules on rachis between
adjacent pinnae instead of one in P. lanceolata. However,
since this character was observed hitherto only in one sin-
gle specimen (Barnard 1965, pl. 98, Fig.3; also see retaken
photos of this specimen in Schweitzer and Kirchner 1998, pl.
6, figs.1, 2) and regarding the remarkable agreement with P.
lanceolata characters, it would be more reliable to consider
P. shemshakensis as a synonym of P. lanceolata.
Pachypteris rhomboidalis (Andrae) Doludenko, 1974 is
similar to P. lanceolata in the pinnule outline (ovate to lan-
ceolate or rhomboidal shape with decurrent catadromous
and contracted anadromous base), but P. rhomboidalis is
distinguished by its larger pinnule size (15–20 × 5–7mm
versus 5–10 × 1.8–2mm in the Abyek material). Further-
more, in P. rhomboidalis, the pinnule midvein is quite clear
and prominent, and the secondary veins are recognizable in
detail, whereas in P. lanceolata, the midvein and secondary
veins are hard to distinguish.
Pachypteris papillosa (Thomas and Bose) Harris, 1964
is similar to P. lanceolata in pinnules shape (ovate to lan-
ceolate) but differs in having pinnate frond versus bipin-
nate frond of P. lanceolata. P. papillosa has a larger pinnule
attachment angle (70° vs. 30°–45° in the Abyek material)
and includes larger pinnules (up to 45 × 10mm) as well as
pinnules lobed once in the acroscopic margin versus the
entire margin in P. lanceolata.
Distribution:
Up to now, this species has been recorded from the Middle
Jurassic (Bajocian-Bathonian) of Yorkshire (Brongniart 1828;
Phillips 1829; Harris 1964), the Bathonian of Hissar Range in
Tajikistan, the Callovian of Ak-Djar in Kyrgyzstan, the Callovian
of Shirkent and Karatag in Tajikistan (Gomolitzky etal. 1962),
the Bajocian of Taurkyr in southern Turkmenistan (Baranova
etal. 1963), the Middle Jurassic of Tkvarchelian coal-basin in
Transcaucasia (Georgia) (Delle 1967), the Callovian of North-
ern Caucasus (Vakhrameev and Samylina 1958), the Callovian
of Georgia (Doludenko 1969; Doludenko and Svanidze 1969),
and the Middle Jurassic of Dahana-i-Tor in northern Afghani-
stan (Schweitzer and Kirchner 1998). In Iran, P. lanceolata was
described from the Middle Jurassic of the Central Alborz (Bar-
nard 1965; Kimyai 1971; Barnard and Miller 1976; Fakhr 1977),
the Eastern Alborz (Sadovnikov 1976, 1980), and the Kerman
basin in Central Iran (Schweitzer and Kirchner 1998). This is the
first record of this species in the Western Alborz.
Genus Ptilozamites Nathorst, 1878 a, b emend.
Type-species: Ptilozamites heeri Nathorst, 1878.
Ptilozamites tenuisÔishi 1932, emend. Ôishi and Huz-
ioka 1938
Figure5a–c
Selected synonyms:
1931 Pterophyllum contiguum (non Schenk), Ôishi, p. 359.
1932 Ptilozamites tenuis Ôishi, p. 321; pl. 43 (25),
figs.1–3.
1938 Ptilozamites tenuis Ôishi, Ôishi and Huzioka, p. 93,
94; pl. 12 (6), Fig.2, 2a.
1940 Ptilozamites tenuis Ôishi, Ôishi, p. 365.
? 1976 Ctenozamites usnadzei (non Doludenko and Sva-
nidze), Barnard and Miller; pl. 1, Fig.1.
1977 Ctenozamites iranica Fakhr, p. 87; pl. 26, Fig.1.
1977 Ptilozamites abiekensis Fakhr, 87; pl. 56, figs.2, 3.
1998 Ctenozamites iranica Fakhr; Schweitzer and Kirch-
ner, p. 51–53; pl. 11, figs.7–9; text- Fig.22.
Material. KA 6, 49–53
Description:
Pinna fragments are up to 50mm long and vary between 14
and 20mm wide. The pinna rachis is 0.8 to 1.3mm wide, bearing
longitudinal ribs. Pinnules arise alternately at an angle of 40°–60°,
attached by their whole base to the rachis, slightly decurrent on
the basiscopic margin. The pinnules are 10.5 to 14mm long, 3
to 3.5mm wide, with parallel and entire margins and truncated
apex. Veins are prominent and coarse, simple or forked at the base
(Fig.5c, arrowhead). The number of veins in the middle part of
pinnules is 5 to 6. The vein concentration is 14 to 16 per centimeter.
Remarks:
Six specimens of this species were sampled in the Abyek
area. Ptilozamites tenuis Ôishi, 1932 can be distinguished
from other Ptilozamites species by its linear, parallel-sided
pinnules with truncated apex and the few number of veins
(4 to 7) with a parallel arrangement.
Comparisons:
The characters of the species Ptilozamites abiekensis Fakhr,
1977 and Ptilozamites (Ctenozamites) iranica Fakhr, 1977
coincide with the diagnosis of Ptilozamites tenuis Ôishi, 1932
emended by Ôishi and Huzioka 1938. These characters are
briefly as follows: the angle of pinna attachment (55° to 75°);
the pinnule outline (linear, parallel‐sided with decurrent base
and truncated apex ending to an acute point on the acroscopic
margin); the venation pattern (parallel veins, simple or forked at
the base); and vein numbers (4 to 6 in the middle of pinnules).
Accordingly, Ptilozamites abiekensis and Ptilozamites (Cteno-
zamites) iranica should be regarded as synonyms of P. tenuis.
Following are comparisons with most similar species:
Ptilozamites (Ctenozamites) caspiensis Schweitzer and Kirch-
ner, 1998, from the Upper Rhaetian of Ekrasar in Western Alborz
and Zirab in the Central Alborz, is similar to Ptilozamites tenuis
in having equal pinnule length (11–16mm in average), and the
same venation pattern (parallel veins, simple or rarely forked near
the base), but it differs in having larger pinnule attachment angle
(70° vs. 40°–60° in the Abyek material), wider pinnules (5–8mm
vs. 3–3.5mm in the Abyek material), and higher vein density (20
veins per centimeter vs. 14–16 in the Abyek material).

Arab J Geosci (2022) 15:1573
1 3
Page 9 of 14 1573
Ptilozamites (Ctenozamites) usnadzei, from the Upper
Jurassic of Georgia (Doludenko and Svanidze 1969) and the
Middle Jurassic of the Central Alborz (Barnard and Miller
1976), is similar to P. tenuis in pinnules length (10–18mm
long) and the pinnule outline (rather parallel-sided with
rounded to truncated apex); however, P. usnadzei has nar-
rower pinnules (6–11mm wide), and more divergent veins
that are unbranched or forked once in the middle of pin-
nules, versus parallel veins, simple or forking near the base
in P. tenuis.
Fig. 5 a–c Ptilozamites tenuis,
Abyek area; c close-up of vena-
tion with lateral vein dichoto-
mized in the base (arrowhead)

Arab J Geosci (2022) 15:1573
1 3
1573 Page 10 of 14
Table 2 Pteridospermophyta species so far described from Iran
Species Author Location Age
Scytophyllum waehneri Stur 1886 Western Alborz Rhaetian
Schenk 1887 Western Alborz
Krasser 1891 Western Alborz
Zeiller 1905 Central Alborz
Kilpper 1975 Central Alborz
Sadovnikov 1976 Central Alborz
Schweitzer 1978 Central Alborz
Sadovnikov 1980, 1983 Eastern Alborz
Schweitzer and Kirchner 1998 Central Alborz
Eastern Alborz
Western Alborz
Central Iran
Vaez-Javadi 2006 Eastern Alborz
Farahimanesh etal. 2021 Eastern Alborz
Ctenozamites cycadea Schenk 1887 Western Alborz Early Jurassic
Schweitzer and Kirchner 1998 Eastern and Western Alborz
Ctenozamites usnadzei Barnard and Miller 1976 Central Alborz Middle Jurassic
Ctenozamites iranica Fakhr 1977 Western Alborz Rhaetian
Ctenozamites whitei Schweitzer and Kirchner 1998 Central Alborz Middle Jurassic
Ctenozamites caspiensis Schweitzer and Kirchner 1998 Western Alborz Rhaetian
Vaez-Javadi 2011 Central Alborz Middle Jurassic
Ptilozamites sp. Potonié 1933 Western and Central Alborz Rhaetian-Early Jurassic
Ptilozamites abiekensis Fakhr 1977 Western Alborz Rhaetian
Ptilozamites ctenoides Sadovnikov 1976, 1980 Eastern Alborz Rhaetian
Ptilozamites nilssoni Sadovnikov 1976, 1980, 1983, 1991 Eastern Alborz Rhaetian
Ptilozamites tenuis This study Western Alborz Rhaetian-Early Jurassic
Lepidopteris ottonis Lorenz 1964 Central Alborz Rhaetian
Schweitzer and Kirchner 1998 Eastern Alborz and Central Iran Norian
Lepidopteris sp. Kilpper 1975
Sadovnikov 1976, 1983
Central Alborz
Eastern Alborz
Rhaetian
Lepidopteris shemshakensis Sadovnikov 1980 Eastern Alborz Rhaetian
Pachypteris shemshakensis Barnard 1965
Schweitzer and Kirchner 1998
Central Alborz Rhaetian
Pachypteris lanceolata Kimyai 1971 Central Alborz Middle Jurassic
Barnard and Miller 1976 Central Alborz
Sadovnikov 1976, 1980 Eastern Alborz
Schweitzer and Kirchner 1998 Central Iran
This study Western Alborz Rhaetian-Early Jurassic
Keraiaphyllum brevifolium Kilpper 1975 Central Alborz Rhaetian
Schweitzer and Kirchner 1998 Central Alborz Norian-Rhaetian
Keraiaphyllum rarinervium Sadovnikov 1976, 1983 Central Alborz Rhaetian
Keraiaphyllum tazarense Sadovnikov 1983 Eastern Alborz Rhaetian
Keraiaphyllum intersulcatum Sadovnikov 1983 Eastern Alborz Rhaetian
Drepanozamites tietzei Schenk 1887 Western Alborz Rhaetian
Kilpper 1975 Central Alborz Rhaetian
Schweitzer and Kirchner 1998 Central Alborz Norian-Rhaetian
Sagenopteris colpodes Barnard and Miller 1976 Central Alborz Middle Jurassic
Schweitzer and Kirchner 1998 Central Alborz
Sagenopteris iranica Fakhr 1977 Eastern Alborz Early-Middle Jurassic

Arab J Geosci (2022) 15:1573
1 3
Page 11 of 14 1573
Ptilozamites chinensis Hsü, 1950, from the Late Triassic of
Hunan in China, is similar to P. tenuis in having elongated pin-
nules, but it differs due to the linear-falcate shape of pinnules and
the acute apex of pinnules with two to four apical teeth (see Hsü
1950, and Zhou 1981).
Ptilozamites (Ctenozamites) cycadea (Berger 1832) Schenk,
1887 shows similarities with P. tenuis in having entire margin pin-
nules with straight upper margin, parallel veins, and similar vein
concentration, but it differs by the convex lower margin of pinnules,
the larger pinnule size (typically 20 × 10cm), less length/width ratio
(two times as long as broad), and the number of vein dichotomies
(forking once or twice vs. simple or forked once in P. tenuis).
Ptilozamites nilssoni Nathorst, 1878 shows some elongated pin-
nules like those of P. tenuis and a similar venation pattern to the
latter (simple or occasionally bifurcated in the base) but differs by its
falcate form of pinnules with a convex lower margin, in contrast to
linear, parallel-sided pinnules of P. tenuis. Furthermore, P. nilssoni
shows lower vein density (i.e., 10/cm vs. 14–16 in P. tenuis).
Ptilozamites tenuis is somewhat similar to some species of Nilsso-
nia (i.e., N. acuminata, N. compta, and N. muensteri). But, contrary to
Nilssonia, adjacent pinnules are equal in width, and pinnules’ lamina
never covers the rachis. It is also distinguished from Nilssonia by its
coarse and forked veins.
Nilssonia acuminata (Presl) Göppert, 1844 resembles the pre-
sent material in having similar pinnae width and elongated pin-
nules with truncated apex but differs in pinnules’ falcate shape,
lower length/width ratio, and variable widths. Moreover, in N.
acuminata, veins are fine and simple with a higher concentration
(18–23 veins per centimeter vs. 14–16 in the present material).
Nilssonia compta (Phillips) Göppert, 1844 is similar to Ptilo-
zamites tenuis in having parallel-margined pinnules with truncated
apex but differs by its much larger pinnae width (40–60mm vs.
14–20mm in the present material), the variable shape of pinnules,
and the higher vein concentration (20 veins per centimeter).
Nilssonia muensteri (Presl) Schimper 1872 is also similar to
the present material in the pinnae size and elongated pinnules,
but it differs in having falcate pinnules with a lower length/width
ratio and fine simple veins with a higher concentration (20–22
veins per centimeter).
Distribution:
So far, Ptilozamites tenuis has been reported from the Rhae-
tian of Southern Japan (Ôishi 1931, 1932, 1940; Ôishi and Huz-
ioka 1938). This is the first record of this species from Iran.
Discussion
Comparison withseed ferns fromother regions
inIran
Thirty-five seed fern species have so far been described from four
regions in Iran (i.e., Western Alborz, Central Alborz, Eastern
Alborz, and Central Iran), including the present materials (Table2).
Inspection of Table2 reveals the following results:
Central Alborz has recorded the highest number of seed
fern taxa in Iran (20 species).
Fifteen seed fern taxa have been documented in the
Western Alborz; among them, nine species were recorded
in Iran only from this region (Ctenozamites cycadea, C
iranica, Ptilozamites abiekensis, P. tenuis, Sagenopteris
sp. A, Sagenopteris sp. B, Caytonanthus arberi, Caytonia
sp., and Amdrupia stenodonta), and six taxa (Sagenopteris
Table 2 (continued)
Species Author Location Age
Sagenopteris nilssoniana Fakhr 1977 Central Alborz Early-Middle Jurassic
Schweitzer and Kirchner 1998 Central Alborz
Vaez-Javadi and Mirzaei-Ataabadi 2006 Central Iran
Sagenopteris undulata Fakhr 1977 Eastern Alborz Early-Middle Jurassic
Sagenopteris phillipsi Sadovnikov 1976, 1980 Eastern and Central Alborz Middle Jurassic
Sagenopteris sp. A This study Western Alborz Rhaetian-Early Jurassic
Sagenopteris sp. B This study Western Alborz Rhaetian-Early Jurassic
Peltaspermum decipiense Schweitzer and Kirchner 1998 Central Alborz Norian-Rhaetian
Farahimanesh etal. 2021 Eastern Alborz Rhaetian
Caytonia sp. Schweitzer and Kirchner 1998 Central Alborz Middle Jurassic
Caytonanthus arberi This study Western Alborz Rhaetian-Early Jurassic
Scoresbya dentate Schweitzer and Kirchner 1998 Central Alborz
Central Iran
Rhaetian-Early Jurassic
Scoresbya nariwaensis Schweitzer and Kirchner 1998 Central Alborz Rhaetian?
Anthrophyopsis crassinervis Schweitzer and Kirchner 1998 Central Alborz
Eastern Alborz
Central Iran
Rhaetian
Amdrupia stenodonta Sadovnikov 1983 Western Alborz Rhaetian

Arab J Geosci (2022) 15:1573
1 3
1573 Page 12 of 14
sp. A, Sagenopteris sp. B, Caytonia sp., Caytonanthus
arberi, Pachypteris lanceolata, and Ptilozamites tenuis)
are recorded in this paper for the first time from the West-
ern Alborz. Also, thirteen seed fern taxa from the Eastern
Alborz and six from Central Iran have been recorded.
Two seed fern species (Scytophyllum waehneri and
Pachypteris lanceolata) occur in all mentioned regions.
The seed fern from the Western Alborz shares more taxa (5
species, namely Scytophyllum waehneri, Ctenozamites caspien-
sis, Pachypteris lanceolata, Drepanozamites tietzei, and Ptilo-
zamites sp.) with the flora from the Central Alborz than with the
Eastern Alborz (3 species, namely Scytophyllum waehneri, Pti-
lozamites nilssoni, and Pachypteris lanceolata) and Central Iran
floras (2 species, namely Scytophyllum waehneri and Pachypt-
eris lanceolata) (Table2). We can see that the Western Alborz
seed ferns are most similar to those of the Central Alborz, while
they represent the least similarities to the seed ferns encountered
in Central Iran; this is consistent with their paleogeographic
distances.
Palaeogeographical andpalaeoecological
implications
The floral composition is generally influenced by two main
factors: geography and climate (see Abbink etal. 2004). Dur-
ing the Late Triassic-Middle Jurassic time, the Alborz basin
was located in the peripheral foreland basin of the Cimme-
ride orogenic belt, along the southern margin of the Eurasia
super-continent (Golonka 2004; Fürsich etal. 2009; Wilmsen
etal. 2009). The formation of the peripheral foreland basin
in the Western Alborz region, and the appearance of a nar-
row marine embayment within this basin (Fürsich etal. 2009;
Wilmsen etal. 2009), provided favorable conditions for habi-
tats hosting plants such as encountered seed ferns.
Caytoniales are considered to have inhabited shadow places
of inland and undisturbed areas near the seashore (Meyen 1988)
under a warm climate with high air humidity and low soil mois-
ture (Abbink etal. 2004; Barbacka 2011; Götz etal. 2011).
Pachypteris has been considered a subtropical genus, colonizing
coastal areas (Vakhrameev 1991; Abbink etal. 2004; Barbacka
2011). Ptilozamites is also considered to have thrived in coastal
regions (Kustatscher etal. 2010; Barbacka 2011).
Conclusion
In the studied areas, the Shemshak Group has yielded the
remains of Caytonialean seed ferns and Pteridospermophyta
with unknown botanical affinity. They are represented here by
four foliage species (Sagenopteris sp. A, Sagenopteris sp. B,
Pachypteris lanceolata, and Ptilozamites tenuis) as well as two
reproductive species (Caytonia sp. and Caytonanthus arberi).
Caytonanthus arberi has been recorded for the first time from
Asia; likewise, it is the first record of Ptilozamites tenuis from
Iran and Pachypteris lanceolata from the Western Alborz. The
seed fern taxa encountered in Iran vary considerably across dif-
ferent regions; however, the seed ferns recorded in the Western
Alborz are most similar to those from Central Alborz.
Pachypteris lanceolata and Ptilozamites tenuis may
represent coastal areas, while encountered Caytoniales
can reflect more inland areas under warm, humid climates.
Declarations
Conflict of interests The authors declare that they have no competing
interests.
References
Abbink OA, Van Konijnenburg-van Cittert JH, Visscher H (2004) A spo-
romorph ecogroup model for the Northwest European Jurassic-Lower
Cretaceous: concepts and framework. Neth J Geosci 83(1):17–31
Alavi M, Barale G (1970) Etude preliminaire de la flore de la Forma-
tion de Shemshak dans la region de Djam (Iran). Bull. Mus. Soc.
Linneene de Lyon 39(8, 2–3): 241–252
Baranova ZE, Burakova AI, Bekasova NB (1963) Stratigraphy,
lithology and flora of the Jurassic sediments in Tuarkyra. Trudy
VSEGEI 88:1–232
Barbacka M (2011) Biodiversity and the reconstruction of Early
Jurassic flora from the Mecsek Mountains (southern Hungary).
Acta Palaeobot 51(2):127–179
Barnard PDW (1965) The geology of the upper Djadjerud and Lar
valleys (North Iran) II. Palaeontology. Flora of the Shemshak
Formation Part 1. Liassic plants from Dorud. Rivista Italiana
di Paleontologia e Stratigrafia 71 (4): 1123–1168
Barnard PDW (1967) The geology of the Upper Djadjerud and Lar
Valleys (North Iran). II. Palaeontology. Flora of the Shemshak
Formation. Part 2: Liassic plants from Ashtar and Shemshak.
Rivista Italiana De Paleontologia e Stratigrafia 73:539–568
Barnard PDW, Miller JC (1976) Flora of the Shemshak Formation
(Elburz, Iran). Part 3: Middle Jurassic (Dogger) plants from
Katumbargah, Vasek Gah and Imam Manak. Palaeontographica
B 155:6–117
Berger AC (1832) Die Versteinerungen der Fischer und Pflanzen im
Sandsteine der Coburger Gegend. Coburg
Boureau É, Furon R, Rosset LF (1950) Contribution à l’étude des
floresjurassiques d’Asie. Mus Natl Hist Nat 30:207–242
Brongniart A (1825) Observations sur les végétaux fossiles renfer-
més dans les grès de Hoer en Scanie. Ann Sci Nat 6:200–224
Brongniart A (1828) Histoire des végétaux fossiles ou recherches
botanique et géologique surles végétaux renfermes dans
les diverses couches du globe. T. 1. Paris: G. Dufour et
Ed.d’Ocagne: 1–136
Corsin P, Stampfli G (1977) La Formation de Shemshak dans
L’elburz Oriental (Iran) Flore-Stratigraphie-Paléogéographie.
Geobios 10(4):509–571
Delle GV (1967) The Middle Jurassic flora of the Tkvarchelian coal-
basin (Transcaucasia). Paleobotanika 6:51–132
Doludenko MP (1969) On the relation between the genera Pachypt-
eris and Thinnfeldia. Tr Geol Inst Akad Nauk SSSR 190:14–34

Arab J Geosci (2022) 15:1573
1 3
Page 13 of 14 1573
Doludenko MP (1974) On the relation of the genera Pachyperis,
Thinfeldia and Cycadopteris, Symposium on Morphological and
Stratigraphical Paleobotany. Lucknow 2:8–16
Doludenko MP, Svanidze TI (1969) The Late Jurassic flora of Geor-
gia. Transactions of the Geological Institute. Acad Sci USSR
178:1–116
Elgorriaga A, Escapa IH, Cúneo NR (2019) Southern Hemisphere
Caytoniales: vegetative and reproductive remains from the
Lonco Trapial Formation (Lower Jurassic). Patagonia J Sys-
tematic Palaeontol 17(17):1477–1495
Fakhr MS (1977) Flore Jurassic de l’Iran, Contribution à l’étude
de la flore Rhéto-Liasique de la Formation de Shemshak de
l’Elbourz (Iran): Comite des travaux historiques et scientifiques.
Mém Sect Sci 5:9–284
Farahimanesh M, Ghavidel-Syooki M, Sadeghi A (2021) The Rhae-
tian ferns and seed ferns from the Shemshak Group, Ghoznavi
area, NE Alborz Mountain. Iran Arabian J Geosci 14(1):1–13
Fürsich FT, Wilmsen M, Seyed-Emami K, Majidifard MR (2009)
Lithostratigraphy of the Upper Triassic-Middle Jurassic Shem-
shak Group of Northern Iran. Geol Soc London Spec Publ
312(1):129–160
Golonka J (2004) Plate tectonic evolution of the southern margin
of Eurasia in the Mesozoic and Cenozoic. Tectonophysics
381(1–4):235–273
Gomolitzky NP, Kurbator VV, Sikstel TA (1962) New material char-
acterizing the genus Pachypteris. Paleontology 2:166–167
Göppert HR (1844) Ueber die fossilen Cykadeen überhaupt. mit
Rücksicht auf die in Schlesien vorkommenden Arten. Uebersi-
cht der Arbeiten und Veränderungen der schlesischen Gesells-
chaft für vaterländische Kul-tur1844:114–144
Göppert HR (1861) Über das Vorkommen von Lias-Pflanzen im Kau-
kasus und in der Alborus-Kette. Académie des sciences
Götz AE, Ruckwied K, Barbacka M (2011) Palaeoenvironment of
the Late Triassic (Rhaetian) and Early Jurassic (Hettangian)
Mecsek Coal Formation (south Hungary): implications from
macro-and microfloral assemblages. Palaeobiodiversity Pal-
aeoenvironments 91(2):75–88
Harris TM (1937) The fossil flora of Scoresby Sound East Greenland.
Part 5: stratigraphic relations of the plant beds. Meddelelser om
Gronland, Alexander Doweld
Harris TM (1940) Caytonia. Ann Bot 4(16):713–734
Harris TM (1941) Caytonanthus, the microsporophyll of Gaytonia.
Ann Bot 5(17):47–58
Harris TM (1964) The Yorkshire Jurassic Flora. Vol. 2. Caytoniales,
Cycadales and Pteridosperms. Trustees of the British Museum
(Natural History) London: 1–191
Herendeen PS (2011) Report of the Nomenclature Committee for
Fossil Plants: 7. Taxon 60:902–905
Hsü J (1950) Some Rhaetic plants from Liling, east Hunan. J Indian
Bot Soc 29:1–46
Huzioka K (1970) A new species of Sagenopteris from Nariwa,
southwest Honshu, Japan. Trans Proc Paleontol Soc Japan
77:229–234
Kilpper K (1964) Über die Rät/Lias-Flora aus dem nördlichen Abfall
des Alburs-Gebirges in Nordiran: Teil 1: Bryophyta und Pteri-
dophyta. Palaeontographica B 114:1–78
Kilpper K (1968) Einige Bennettiteen-Blatter aus dem Lias von
Karmuzd-Zirab (N-Iran). J Linnean Soc (botany) 61:129–135
Kilpper K (1971) Uber line Rat/Lias Flora aus dem Nordlichen
Abfall des Alburs-Gerbiges in NordIran; Teil. 2: Ginkgophyten-
Belaubungen. Palaeontogr Abt B 135:89–102
Kilpper K (1975) Paleobotanische Untersuchungen in Nord-Iran. I.
Nachweis nichtmariner Obertrias am Nordabfall des Alburs-
Gebirges. 1. Grossform der Pflanzenfunde von seltenen Gattun-
gen. Rev Palaeobot Palynol 19:139–153
Kimyai A (1971) A study of plant fossil of Shemshak Formation in
Garmabdar. J Sci Tehran University 3(2):1–15
Krasser F (1891) Über die fossile Flora der rhätischen Schichten Per-
siens. Sitzungsberichte der österreichischen Akademie der Wis-
senschaften, mathematisch-naturwissenschaftliche Klasse 100:
413–432
Kustatscher E, Pott C, Van Konijnenburg-van Cittert JH (2011) Scy-
tophyllum waehneri (Stur) nov. comb., the correct name for Scy-
tophyllum persicum (Schenk) Kilpper, 1975. Zitteliana 51:9–18
Kustatscher E, Van Konijnenburg-van Cittert JH (2010) Seed ferns
andcycadophytes from the Triassic flora of Thale (Germany).
NeuesJahrbuch für Geologie und Paläontologie-Abhandlungen
258(2):195–217
Lorenz C (1964) Die Geologie des oberen Karadj-Tales (Zentral-
Elburz), Iran. Offsetdruck, Schmidberger & Müller
Meyen SV (1988) Reconstructions of selected Seed Ferns. Ann Mis-
souri Bot Gard 75:1010–1057
Ôishi S (1931) Yabeiella sp. from the Japanese Triassic. Jap J Geol
Geogr 8(4):357–359
Ôishi S (1932) The Rhaetic plants from the Nariwa District, Prov.
Bitchu (Okayama Prefecture), Japan. J Faculty Sci Hokkaido Imp
Univ Ser 4 Geol Mineral 1(3):257–380
Ôishi S (1940) The Mesozoic floras of Japan. Journ. Fac. Sci. Hokkaido
Imp Univ 5:123–480
Ôishi S, Huzioka K (1938) Fossil plants from Nariwa A supple-
ment. J Faculty Sci Hokkaido Imp Univ Ser 4 Geol Mineral
4(1–2):69–101
Oliver FW, Scott DH (1904) On the structure of the Palaeozoic seed,
Lagenostoma lomaxi, with a statement of the evidence upon
which it is referred to Lyginodendron. Philos Trans R Soc Lond
B197:193–247
Osborn JM (1994) The morphology and ultrastructure of Caytonan-
thus. Can J Bot 72(10):1519–1527
Phillips J (1829) Illustrations of the Geology of Yorkshire, or, a
description of the strata and organic remains of the Yorkshire
Coast. York, Thomas Wilson
Potonié R (1933) Über einige Pflanzenreste aus dem Jura Persiens.
Arbeit Inst Paläobot Petrogr Brennstein 3:247–250
Prakash N, Das N (2017) First record of microsporophyll genus Cay-
tonanthus Thomas from Early Cretaceous beds of South Rewa
Gondwana Basin, India: Its evolutionary and palaeogeographical
significance. Island Arc 26(1):1–11
Sadovnikov GN (1976) The Mesozoic flora of Alborz and central Iran
and its stratigraphicimportance. National Iran Steel Company of
Iran, Tehran, pp 1–118
Sadovnikov GN (1980) Flora of the Mesozoic coal formation in the
Elburz Mountains. Proceedings of the USSR, Academy of Science
Sadovnikov GN (1983) Die Flora des Kohlenformation des Elburs.
III. Atlas of the Gheshlagh-Flora. Akad. Nauk SSSR Inst Geol
Moscow 83 (in Russian)
Sadovnikov GN (1991) On some gymnosperms from the late Triassic
of North Iran. Paleontol J 4:95–106
Schenk A (1887) Fossile Pflanzen aus der Albourskette. Bibliotheca
Botanica Wiesbaden 6:1–12
Schimper WP (1872) Traité de paléontologie végétale, ou La flore du
monde primitif dans ses rapports avec les formations géologiques
et la flore du monde actuel. Baillière et fils, Paris
Schweitzer HJ (1977) Die räto-jurassischen Floren des Iran und
Afghanistans. 4. Die rätische Zwitterblüte Irania hermaphroditica
nov. spec. und ihre Bedeutung für die Phylogenie der Angiosper-
men. Palaeontographica B 161:98–145
Schweitzer HJ (1978) Die räto-jurassischen Floren des Iran und
Afghanistans. 5. Todites princeps, Thaumatopteris brauniana
und Phlebopteris polypodioides. Palaeontographica B 168:17-60

Arab J Geosci (2022) 15:1573
1 3
1573 Page 14 of 14
Schweitzer HJ, Kirchner M (1995) Die Rhäto-Jurassischen Floren
des Iran und Afghanistans: 8. Ginkgophyta. Palaeontographica
B 237:1–58
Schweitzer HJ, Kirchner M (1996) Die rhäto-jurassischen Floren des
Iran und Afghanistans. 9. Coniferophyta. Palaeontographica B
238:77–139
Schweitzer HJ, Kirchner M (1998) Die rhäto-jurassischen Floren des
Iran und Afghanistans. 11. Pteridospermophyta und Cycadophyta
I. Cycadales.Palaeontographica B 248:1–85
Schweitzer HJ, Kirchner M (2003) Die rhato-Jurassischen Floren des
Iran und Afghanistans. 13. Cycadophyta III. Bennettitales. (Mit
40 Tafeln, 67 Abbildungen und 3 Tabellen im Text und auf 4
Beilagen). Palaeontographica B 264:1–166
Schweitzer HJ, Van Konijnenburg-van Cittert JHA, Van der Burgh J
(1997) The Rhaeto-Jurassic flora of Iran and Afghanistan. 10.
Bryophyta, Lycophyta, Sphenophyta, Pterophyta-Eusporangiatae
and-Protoleptosporangiatae. Palaeontographica B 243:103–192
Schweitzer HJ, Kirchner M, Van Konijnenburg-Van Cittert JH (2000)
The Rhaeto-Jurassic flora of Iran and Afghanistan. 12. Cycado-
phyta II. Nilssoniales. Palaeontographica B 254:1–63
Schweitzer HJ, Schweitzer U, Van Konijnenburg-van Cittert JHA, Van
der Burgh J, Ashraf RA (2009) The Rhaeto-Jurassic flora of Iran
and Afghanistan. 14. Pterophyta-Leptosporangiatae. Palaeonto-
graphica B 279:1–108
Stur D (1886) Vorlage der von Dr. Wähner aus Persien mitgebrachten
fossilen Pflanzen. Verhandlungen Der Kaiserlich-Königlichen
Geologischen Reichsanstalt 16:431–436
Taylor EL, Taylor TN, Krings M (2009) Paleobotany: the biology and
evolution of fossil plants. Academic Press
Thomas HH (1925) The Caytoniales, a new group of angiospermous
plants from the Jurassic rocks of Yorkshire. Phil Trans R Soc
Lond 213B:299–363
Vakhrameev VA (1991) Jurassic and Cretaceous floras and climates of
the Earth. Cambridge University Press
Vakhrameev VA, Samylina VA (1958) The first discovery of a
representative of genus Pachypteris in the U.S.S.R. Bot Zh
43:1611–1612
Vaez-Javadi F (2006) Plant fossil remains from the Rhaetian of Shem-
shak Formation, Narges-Chal area, Alborz, NE Iran. Riv Ital Pale-
ontol Stratigr 112(3):397–416
Vaez-Javadi F (2011) Middle Jurassic flora from the Dansirit For-
mation of the Shemshak Group, Alborz, north Iran. Alcheringa
35(1):77–102
Vaez-Javadi F, Allameh M (2015) Biostratigraphy of the Bazehowz
Formation at its Type section, South West Mashhad based on plant
macrofossils. Geopersia 5(1):27–44
Vaez-Javadi F, Ghavidel-Syooki M (2002) Plant megafossil remains
from Shemshak Formation of Jajarm area, NE Alborz Iran. The
Palaeobotanist 51(1–3):57–72
Vassiliev Y (1984) Mesozoic plant fossils from coal areas in Iran. Atlas
Min Met 2(2):1–97
Ward LF (1900) Status of the Mesozoic floras of the United States,
First paper: the older Mesozoic. Reports of the United States Geo-
logical Survey (Vol. 20). US Government Printing Office
Wilmsen M, Fürsich FT, Seyed- Emami K, Majidifard MR, Taheri J
(2009) The Cimmerian Orogeny in northern Iran: Tectono- strati-
graphic evidence from the foreland. Terra Nova 21(3):211–218
Zeiller R (1905) Sur les plantes rhétiennes de la Perse recueillies par
MJ de Morgan. Société géologique de France 5:190–197
Zhou Z (1981) Some Late Triassic and Early Jurassic pteridosperms
from Hunan. Acta Palaeontol Sin 20(1):15–26
Springer Nature or its licensor holds exclusive rights to this article under
a publishing agreement with the author(s) or other rightsholder(s);
author self-archiving of the accepted manuscript version of this article
is solely governed by the terms of such publishing agreement and
applicable law.