Content uploaded by Philippe Moisan
Author content
All content in this area was uploaded by Philippe Moisan on Nov 02, 2023
Content may be subject to copyright.
Research paper
Lycopsids from the Madygen Lagerstätte (Middle to Late Triassic, Kyrgyzstan,
Central Asia)
Philippe Moisan
a,
⁎, Sebastian Voigt
b
a
Forschungsstelle für Paläobotanik, Institut für Geologie und Paläontologie, Westfälische Wilhelms-Universität Münster, Schlossplatz 9, 48143 Münster, Germany
b
Geologisches Institut, Technische Universität Bergakademie Freiberg, Bernhard-von-Cotta-Straße 2, 09599 Freiberg, Germany
abstractarticle info
Article history:
Received 18 February 2012
Received in revised form 5 December 2012
Accepted 14 December 2012
Available online 4 January 2013
Keywords:
Isoetites
Lepacyclotes
Ferganodendron
Mesenteriophyllum
Triassic
Madygen
Abundant lycopsid remains from the Middle–Late Triassic Madygen Formation in southwestern Kyrgyzstan,
Central Asia are described in detail, based on macromorphological and epidermal features. The lycopsid
assemblage consists of the subarborescent morphotaxa Mesenteriophyllum kotschnevii,Ferganodendron
sauktangensis and Pleuromeiopsis kryshtofovichii, and the herbaceous morphotaxon Lepacyclotes zeilleri that
is described by the first time from Central Asia. In addition, two new species are introduced, Isoetites
madygensis Moisan et Voigt sp. nov. and Isoetites sixteliae Moisan et Voigt sp. nov., which show a remarkable
similarity to extant Isoëtes species, i.e., a short corm and leaves with air channels, suggesting that this lineage
goes back to the Early Mesozoic at least. The Madygen lycopsid flora consists of a mixture of subarborescent
and herbaceous forms that has not been previously documented from the Ladinian–Carnian onward. The
Madygen biota thrived at mid-northern latitudes in an intramontane basin, while most coeval occurrences
represent tropical and subtropical biotas. The geographic position and climate are considered to be the
main reasons for the unique flora and fauna found in the Madygen Formation, which shows a remarkably
high abundance of lycopsids. The Madygen lycopsids show several characteristics indicating (semi-)aquatic
habitats.
© 2013 Elsevier B.V. All rights reserved.
1. Introduction
Lycopsids reached their maximum diversity in the Pennsylvanian
(Niklas et al., 1985; Meyen, 1987; DiMichele and Skog, 1992; Thomas,
1992) with as major representatives the arborescent lepidodendrids
and sigillarians that dominated tropical wetlands. In palaeotropical
Euramerica most arborescent lycopsids became extinct in the Late
Pennsylvanian (Phillips et al., 1985), but in Cathaysia arborescent forms
persisted into the Permian (Van Waveren et al., 2007). Nevertheless,
lycopsids are extremely rare in the uppermost Permian, but they were
dominant again in the Lower Triassic. After the worldwide end-Permian
biotic crisis lycopsids played a crucial role in the repopulation of the
habitats as opportunistic pioneer plants (Retallack, 1975; Looy et al.,
1999; Grauvogel-Stamm and Ash, 2005). The heterogeneity of the
terrestrial plant communities increased markedly between the Anisian–
Ladinian as a result of the resurgence and proliferation of several
plant groups such as sphenopsids, ferns, pteridosperms, cycadophytes,
ginkgophytes, and conifers (e.g., Retallack, 1975; Krassilov, 1981;
Kustatscher et al., 2007; Hermann et al., 2011). The Madygen flora
is not only one of the most diverse Middle–Late Triassic Northern
Hemisphere floras known to date (Dobruskina, 1995; Moisan et al.,
2011, 2012), it also comprises a high abundance of lycopsids unlike
most other contemporaneous floras.
In Eurasia, especially in central and western Europe, Russia, China,
the northern Caucasus and western Kazakhstan, the terrestrial
vegetation was dominated by the lycopsid Pleuromeia Corda during
the Early Triassic (Vakhrameev et al., 1978; Dobruskina, 1987, 1994;
Looy et al., 1999, 2001). Pleuromeia occurs mostly in monotypic asso-
ciations in the Lower–Middle Triassic, occasionally associated with
conifers, ferns and sphenopsids (e.g., Kryshtofovich, 1923; Kon'no,
1973; Dobruskina, 1974; Grauvogel-Stamm, 1978; Wang and Wang,
1982; Fuchs et al., 1991; Dobruskina, 1994).
Although occurrences of lycopsid macrofossils are relatively scarce
in late Middle–Late Triassic floras worldwide, the Ladinian–Carnian
Madygen Formation from Kyrgyzstan has yielded several lycopsid
forms that occur abundantly. Lycopsid remains from the Madygen
Formation were first identified by Brick (1936), in a small collection
of fossil plants made by E. A. Kochnev during geological mapping of
the Madygen area. She identified two small fragments as Pleuromeia
oculina (Blankenhorn) Potonié, and dated the Madygen Formation
as Early Triassic. Based on a larger collection of fossil plants from
Madygen, Sixtel (1962) reported Sigillaria sauktangensis Sixtel,
Review of Palaeobotany and Palynology 192 (2013) 42–64
⁎Corresponding author.
E-mail address: f_mois01@uni-muenster.de (P. Moisan).
0034-6667/$ –see front matter © 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.revpalbo.2012.12.003
Contents lists available at SciVerse ScienceDirect
Review of Palaeobotany and Palynology
journal homepage: www.elsevier.com/locate/revpalbo
Sigillaria sp., Pleuromeiopsis kryschtofovichii Sixtel and Pleuromeiopsis
sp. Dobruskina (1974) revised Sixtel's and Brick's material and
established the monotypic genus Ferganodendron to accommodate
the above-mentioned lycopsids described by Sixtel and Brick. In addi-
tion, Sixtel (1961) described the genus Mesenteriophyllum that she
interpreted as a gymnosperm, but which is now recognised as a
lycopsid (Bomfleur et al., 2011).
Newly collected material from the Madygen Formation shows that
the composition of the flora was quite different than suggested previ-
ously, in terms of numbers and taxa, particularly with regard to
lycopsids. Here, we describe the rich and abundant lycopsid flora from
Madygen, which comprises the endemic morphotaxa Ferganodendron
sauktangensis (Sixtel) Dobruskina and Pleuromeiopsis kryshtofovichii
Sixtel as well as the widespread morphotaxa Mesenteriophyllum
kotschnevii Sixtel, Lepacyclotes zeilleri (Fliche) Retallack. In addition,
two new quillworts, Isoetites sixteliae Moisan et Voigt sp. nov. and
Isoetites madygensis Moisan et Voigt sp. nov. are described. The
lycopsids from the Madygen Formation are preserved as impressions
without cuticle. However, the excellent preservation of the fossils,
which were embedded in fine-grained sediments made it possible to
prepare silicone replicas that show minute details of the plant surfaces
under the SEM.
2. Geological setting, material and methods
The Madygen Formation is exposed on the northern rim of the
Turkestan Mountains in southwestern Kyrgyzstan. The Madygen
Formation comprises a c. 580 m thick succession of alluvio-fluvial
and lacustrine deposits exposed in three outcrop areas in the immedi-
ate vicinity of the village of Madygen in the Fergana Valley (Voigt
et al., 2006; Voigt and Hope, 2010). The Madygen Formation uncon-
formably overlies the Palaeozoic basement and is overlain by trans-
gressive Jurassic and Cretaceous deposits. In the southwestern
outcrop area (Urochishche Madygen, Fig. 1), the most completely
preserved succession of the stratotype area, lacustrine deposits
occur in three different levels (lakes 1–3; Fig. 1). Successions are
less complete in the other outcrop areas and comprise sediments of
the two lower lake phases at most. The lycopsid fossils described
Fig. 1. A: Exposures of the Madygen Formation in southwestern Kyrgyzstan and its geographical location in Central Asia. B: Geological map of the Madygen Formation showing
geographic and stratigraphic position of the lycopsid localities.
43P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
herein were collected in ten different localities (Fig. 1;Appendix A)in
the southwestern outcrop area (Urochishche Madygen), the north-
western outcrop area (Urochishche Dzhaylyaucho), and the eastern
outcrop area (Sauk Tanga). All fossils were found in lacustrine deposits,
five sites in lake level 1 in Urochishche Madygen and Sauk Tanga, two
sites in lake level 2 in Urochishches Madygen and Dzhaylyaucho, and
Plate I. Mesenteriophyllum kotschnevii from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan.
1. Epitype of M.kotschnevii showing the wide midvein and undulated leaf margin. FG 596/X/828. Scale bar =1 cm.
2–6, 8–10. Linear leaves of M.kotschnevii showing the wide midvein and undulated margins. Note the serrated margin in Fig. 3 (arrows). 2: FG 596/X/777; 3: FG 596/X/837; 4:
FG 596/X/495; 5: FG 596/X/395a; 6: FG 596/X/394; 8: FG 596/X/499; 9: FG 596/X/752a; 10: FG 596/X/769b. All scale bars =1 cm.
7. A sporophyll with undulated margin and basal expansion (arrow) associated to M.kotschnevii FG 596/X/938. Scale bar =1 cm.
11–12. Leaves of M.kotschnevii showing a surface transversely folded, representing collapsed air channels. 11: FG 596/X/631b; 12: FG 596/X/281. Scale bars=1 cm.
13. A portion of leaf showing serrated margin (arrow). FG 596/X/383. Scale bar = 1 cm.
Plate II. Ferganodendron sauktangensis from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan. (see on page 46)
1. Portion of a F.sauktangensis stem showing the scar of a lateral branch (arrow). FG 596/X/279. Scale bar=2 cm.
2. Small and rhomboidal leaf bases on the stem of F.sauktangensis. FG 596/X/1032. Scale bar=1 cm.
3–4, 7. Irregular longitudinal striae on the stem surfaces of F.sauktangensis. 3: FG 596/X/256. Scale bar= 2 cm; 4: FG 596/X/391. Scale bar=1 cm; 7: FG 596/X/442. Scale
bar= 1 cm.
5. Rhomboidal leaf bases of F.sauktangensis showing three scars (arrows). FG 596/X/767. Scale bar=1 cm.
6. Detail of Fig. 5. Scale bar=5 mm.
8–10. Portion of stems of F.sauktangensis with elliptical leaf bases more sparsely distributed. 8: FG 596/X/587; 9: FG 596/X/321; 10: FG 596/X/538. All scale bars= 1 cm.
Plate III. Ferganodendron sauktangensis and Pleuromeiopsis kryshtofovichii from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan. (see on page 47)
1. Conical rhizomorph of F.sauktangensis with rounded rootlet scars. FG 596/X/538. Scale bar =2 cm.
2. Conical rhizomorph of F.sauktangensis. FG 596/X/1062. Scale bar = 1 cm.
3–4. Stem and rhizomorph of F.sauktangensis. Note the irregular longitudinal striae on the stem. FG 596/X/1005a–b. Scale bars= 1 cm.
5. A portion of a cast of the rhizomorph of F.sauktangensis. FG 596/X/553. Scale bar =1 cm.
6–7. Stems of Pleuromeiopsis kryshtofovichii showing striated surface and elliptical leaf bases (arrows). 6: FG 596/X/554. Scale bar= 1 cm; 7. FG 596/X/584. Scale bar =
2 cm.
8–9. Fragments of indeterminate stems with scars arranged in parastichies. FG 596/X/150a–b. Scale bars= 1 cm.
Plate IV. Lycopsid leaves and rhizomorphs from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan. (see on page 48)
1–3. Lycopsid leaves showing serrated (?spines) margins. The arrows in Fig. 2 indicate the midvein. 1: FG 596/X/506; 2: FG 596/X/916. Scalebars = 1 cm. 3: FG 596/X/912.
Scale bar= 5 mm.
4. Detail of Fig. 1. Scale bar=2 mm.
5. Detail of Fig. 3. Scale bar=2 mm.
6. Apical of portion of a lycopsid leaf tapering towards the apex. FG 596/X/682. Scale bar =5 mm.
7. A smaller lycopsid leaf with serrated (?spines) margin. FG 596/X/909. Scale bar =5 mm.
8. Lycopsid leaves with entire and serrated margins. FG 596/X/464. Scale bar= 1 cm.
9–11. Fragments of lycopsid rhizomorphs with attached rootlets. FG 596/X/1008a–c. Scale bars= 1 cm.
Plate V. Isoetites sixteliae from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan. (see on page 49)
1–2. Serrated leaves of I.sixteliae ending in a sharp point. 1: FG 596/X/772; 2: FG 596/X/1023b. Scale bars=5 mm.
3–6. Leaves of I.sixteliae showing a fine midvein and serrated margin. 3: FG 596/X/591; 4: FG 596/X/508; 5: FG 596/X/919; 6: FG 596/X/603. All scale bars =5 mm.
7. Holotype of I.sixteliae showing a fine midvein and serrated margin. FG 596/X/918. Scale bar= 5 mm.
8. A large leaf of I.sixteliae showing serrated margin (arrow). FG 596/X/367. Scale bar = 1 cm.
9. Detail of Fig. 8 showing isodiametric cells at the leaf margin. Scale bar= 500 μm.
10–11. Detail of the leaf of I.sixteliae showing the midvein, serrated margin and epidermal cells. FG 596/X/234. Scale bar for fig. 10: 2 mm. Scale bar for fig. 11: 500 μm.
12. Leaf of I.sixteliae showing the epidermal cells arranged in longitudinal rows and serrated margin. FG 596/X/42. Scale bar= 1 mm.
13. Leaf of the holotype of I.sixteliae with an expanded base representing the sporangial portion. FG 596/X/918. Scale bar=1 cm.
14–15. Leaves expanded at the base of I.sixteliae. 14: FG 596/X/519; 15: FG 596/X/316. Scale bars =1 cm.
Plate VI. Isoetites madygensis from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan. (see on page 50)
1. Several sporophylls in growth position of I.madygensis radiating from the central part, where the corm was located. FG 596/X/985a. Scale bar = 5 mm.
2. Detail of Fig. 1 showing the base of the sporophylls with the sporangium and ligule (arrows). Scale bar =5 mm.
3. Detail of Fig. 1, the arrows indicates the position of the sporangium and ligule, and a fragment of a corm. Scale bar =5 mm.
4. Non-elongated, compact corm of I.madygensis bearing leaves/?sporophylls. FG 596/X/740. Scale bar =5 mm.
5. Detail of Fig. 4 showing the rootlet scars and the corm. Scale bar =2 mm.
6. Lanceolate leaves/?sporophylls of I.madygensis. FG 596/X/776. Scale bar =5 mm.
7. Holotype of I.madygensis showing the compact, non-elongated corm bearing several leaves. Isolated leaves show a thin midvein. FG 596/X/54. Scale bar =1 cm.
8. Detail of Fig. 7 showing the non-elongated, compact corm with attached leaves and rootlets scars. Scale bar= 5 mm.
9–13. Sporophylls of I.madygensis showing the basal sporangium. 9: FG 596/X/1004a; 10: FG 596/X/384; 11: FG 596/X/392; 12: FG 596/X/687; 13: FG 596/X/666a. Scale bar
for figs. 9–11: 2 mm. Scale bar for figs. 12–13: 5 mm.
Plate VII. Isoetites sixteliae,I.madygensis and Lepacyclotes zeilleri from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan. (see on page 51)
1, 3. Air channels in the serrated leaves of I.sixteliae. 1: FG 596/X/614; 2: FG 596/X/371a. Scale bars =2 mm.
2. Leaf of a recent Isoëtes showing the internal air channels. Scale bar= 2 mm.
4–7. Collapsed air channels in the entire-margined leaves of I.madygensis. 4: FG 596/X/669b; 5: FG 596/X/978a; 6: FG 596/X/978b; 7: FG 596/X/619. Scale bar for fig. 4:
2 mm. Scale bars for figs. 5–7: 5 mm.
8. Isolated sporophylls of L.zeilleri. FG 596/X/50a. Scale bar=1 cm.
9. Base of a sporophyll of L.zeilleri. FG 596/X/742. Scale bar=5 mm.
10. Isolated sporophylls of L.zeilleri showing the presence of a ligule (arrow). FG 596/X/49. Scale bar=1 cm.
11. Several well-preserved sporophylls of L.zeilleri showing the large sporangium divided by a vertical midline, and the presence of a ligule (arrows). FG 596/X/947.
Scale bar= 1 cm.
12–13. Isolated sporophylls of L.zeilleri. 12: FG 596/X/518; 13: FG 596/X/628. Scale bars = 1 cm.
44 P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
three sites in the uppermost lake level (Fig. 1). Plant remains have been
fluvially transported and deposited in nearshore to offshore parts of the
respective lakes. This study is based on about 200 lycopsid specimens.
A Ladinian–Carnian age has been inferred for the Madygen
Formation, based on the macroflora (Dobruskina, 1995). However,
Shcherbakov (2008a,b) suggested that a Ladinian age is more likely
since in Madygen, some insect groups such as Blattodea, Orthoptera
and Homoptera appear to be more primitive than their relatives
from the Carnian of South Africa and Australia.
The plant fossils are preserved as impressions, lacking organic
material.However, most of the lycopsids occur in very fine-grained sed-
iments in which minute details of the plant surfaces have been pre-
served as casts (e.g., Plate V, 11, 12; Plate VIII, 4, 7). Silicone replicas of
the plant surfaces showing cell structure were prepared and examined
using SEM in order to obtain information of the epidermal cell patterns
(Plate IX, X). For details of this technique see Moisan et al. (2011)
The silicone replicas were analyzed and photographed with a JEOL
840 scanning electron microscope at the Interdisciplinary Centre for
Electron Microscopy and Analysis (ICEM), Westfälische Wilhelms-
Universität Münster, using the image-analysis software analySIS 3.2
(Soft Imaging System, Münster, Germany). Plant surfaces were examined
with a Leica MZ 16 stereomicroscope and photographed with a Nikon
DS-5 M digital camera. Hand specimens were photographed with a
Canon EOS 7D digital camera equipped with a macro photo lens MP-E
65 mm and by using the software Canon EOS Utility version 2.8. In
order to enhance contrast and avoid cast shadows, indirect polarised
light was used. All specimens are deposited in the palaeontological collec-
tion of the Geologisches Institut, Technische Universität Bergakademie
Freiberg, Germany.
3. Systematic palaeobotany
Division Lycophyta
Order Pleuromeiales
Fossil genus Mesenteriophyllum Sixtel, 1961 emend. Moisan et
Voigt
Type species Mesenteriophyllum kotschnevii Sixtel, 1961 emend.
Moisan et Voigt
Remarks: Mesenteriophyllum was erected by Sixtel (1961) who
included two species, M. kotschnevii and M. serratum; she designated
M. kotschnevii as the type species. The genus was classified as
“Gymnospermae incertae sedis”.Dobruskina (1995) also classified
Mesenteriophyllum among the gymnosperms, but arguments to support
a gymnospermous affinity were not given. Mesenteriophyllum was
recognised as a lycopsid by Bomfleur et al. (2011) who demonstrated
Plate VIII. Lycopsids from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan.
1. A sporophyll of L.zeilleri showing the sporangium with several impressions of in situ megaspores and the presence of the ligule (arrow). FG 596/X/1011b. Scale
bar= 5 mm.
2. A sporophyll of L.zeilleri showing the sporangium divided by a vertical midline and the ligule (arrow). FG 596/X/551. Scale bar =5 mm.
3. Detail of Fig. 2 showing the ligule. Scale bar= 2 mm.
4, 6. Pentagonal, hexagonal epidermal cells on the sporophyll of L.zeilleri. 4: FG 596/X/49b; 6: FG 596/X/39. Scale bar for fig. 4 =250 μm. Scale bar for fig. 6 =250 μm
5. Epidermal cells arranged in longitudinal rows (arrows) at the distal portion of a sporophyll of L.zeilleri. FG 596/X/130b. Scale bar =1 mm.
7. Isodiametric to rectangular epidermal cells on the leaf of M.kotschnevii. FG 596/X/425b. Scale bar =1 mm.
8–9. Smaller sporophylls of L.zeilleri with a long pointed apex and the characteristic sporangium divided vertically. 8: FG 596/X/992b; 9: FG 596/X/985c. Scale bars=
2 mm.
52 P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
the presence of air channels, a typical feature for certain lycopsids,
and found this type of leaf in close association with a pleuromeialean
fructification. In Madygen Mesenteriophyllum leaves occur almost
exclusively in lycopsid-dominated assemblages, which include an
abundance of sterile leaves, sporophylls, putative strobili, stems and
rhizomes.
Emended diagnosis: Leaves simple, linear, and large with a distinct, wide
central vein. Leaves with undulated margin, serrated. Epidermal cells
isodiametric, polygonal to rectangular; oriented in distinct longitudinal
rows.
Mesenteriophyllum kotschnevii Sixtel, 1961 emend. Moisan et Voigt
(Plate I,1–13, Plate VIII,7;Plate X,7)
Synonymy and selected references
1936 Taeniopteris spathulata, Brick, p. 169–170; pl. 1, 6.
1960 Nilssonia mesentheriformis,Sixtel,p.72–73; pl. 10, 3.
1961 Mesenteriophyllum kotschnevii, Sixtel, p. 156–157; text-fig. 5.
1961 Mesenteriophyllum serratum, Sixtel, p. 156–158; text-fig. 6.
1962 Thallites insolitus, Sixtel, p. 293–294; pl. 1, 4, 5a–b.
1962 Taeniopteris (?)plicata, Sixtel, p. 362–363; pl. 25, 7; text-fig. 34.
1962 Mesenteriophyllum kotschnevii, Sixtel, p. 398–399; pl. 29, 1–8; text-fig. 47.
1962 Mesenteriophyllum serratum, Sixtel, p. 400; pl. 29, 9–10, text-fig. 48.
1995 Thallites sp., Dobruskina, p. 20; pl. 43, 5–11.
1995 Taeniopteris spathulata, Dobruskina, p. 31; pl. 32, 11, 13–16.
1995 Mesenteriophyllum kotschnevii, Dobruskina, p. 32; pl. 42, 1–8, 12.
1995 Mesenteriophyllum serratum, Dobruskina, p. 32; pl. 42, 11, 18.
2010 Mesenteriophyllum serratum, Bomfleur et al., p. 198; pl. 4, A–G; text-fig. 5.
Emended diagnosis: Leaves simple, linear, and large with a distinct, wide
central vein. Leaves with undulated and serrated margins. Leaves
with transversely folded surfaces. Epidermal cells tetra- to pentagonal,
arranged in regular, longitudinal rows. Cells at the leaf margins trans-
versely rectangular elongate.
Holotype: No 14398/398; p. 155–156; text-fig. 5 (No 14398/938).
Sixtel, 1961.
Epitype: FG 596/X/828 (Plate I, 1).
Material studied: FG 596/X/281, X/350a–b, X/383, X/394–395a–b,
X/425b–c, X/495, X/499, X/610, X/631b, X/752a, X/769a–c, X/777a–b,
X/828, X/837a–b, X/938.
Repository: Palaeontological collection of the Geologisches Institut,
Technische Universität Bergakademie Freiberg, Germany.
Locality: Vicinity of the village of Madygen, approximately 250 km
west of Osh, southwest Kyrgyzstan, Central Asia (Fig. 1).
Stratigraphic horizon: Madygen Formation.
Age: Middle–Late Triassic (Ladinian–Carnian).
Description: All the specimens of Mesenteriophyllum kotschnevii are
incomplete, apex and base of the leaves are not preserved. The largest
leaf is 14 cm long and 2.5 cm wide (Plate I, 1). Smaller leaves between
3–10 cm long and 1–2.5 cm wide. Leaves with a single midvein of
3–5 mm wide. Leaf margins variable in outline, commonly with undu-
lated margins (Plate I,1–2, 4–6, 8–10, 13) and in some portions with
slightly serrated margins (Plate I, 3, 13; arrows). Two specimens show
regular, evenly spaced folds, perpendicular to the midvein (Plate I,
11–12). The folds are at regular distances of c. 1 mm, and nearly reach
the midvein. A putative sporophyll was found in close association
with the sterile leaves. The sporophyll (Plate I, 7) is 8 cm long and
7–13 mm wide, slightly tapering towards the apex and with undulated
margin; a basal expansion of 1.5 cm long and 1.3 cm wide (Plate I,7;
arrow), represents the sporangial region.
Epidermal cells are isodiametric, mostly tetra- to pentagonal in
outline, 30–72 μmlong and 33–74 μm wide, arranged in regular lon-
gitudinal rows (Plate VIII,7;X, 7). The cells towards the leaf margins
are transversely rectangular, elongate in outline, 33–67 μm long μm
and 46–105 μm wide. Stomata were not observed.
Remarks and comparisons: Sixtel (1961) gave a schematic line-
drawing of the holotype of Mesenteriophyllum kotschnevii (No
14398/938, p. 157, text-fig. 5), but the protologue mentions specimen
No 14398/398 as the holotype (Sixtel, 1961, p. 155–156). In 1962 she
gave another description of Mesenteriophyllum and published the
same line-drawing as in 1961, but with different specimen numbers
for the holotype: No 289, p. 399, text-fig. 47 in the caption of the
line drawing, and No 1398 in the text (p. 398–399). A photograph
of the latter specimen (No 1398) (Sixtel, 1962, pl. 29, 1) shows a dif-
ferent aspect than the line drawing, obviously because both only
show a part of the specimen. Unfortunately, neither the photograph
nor the line drawing shows the typical features indicated in the orig-
inal diagnosis (Sixtel, 1961). According to the original diagnosis the
leaf margin is serrated to spiny. However, the line drawing of the ho-
lotype shows an irregular margin with two large spines (Fig. 2). The
photograph of the holotype obviously shows another portion with
an irregular leaf margin, which apparently seems to be affected by in-
sect damage. Because of the confusion regarding the identity of the
holotype and because all the specimens indicated by Sixtel (1961,
1962) as types of M. kotschnevii are ambiguous, we introduce an
epitype in addition to the holotype in accordance with ICBN Art. 9.7
(McNeill et al., 2006).
Sixtel (1961, 1962) distinguished two species within
Mesenteriophyllum,i.e.,M. serratum and M. kotschnevii.Leavesof
M. serratum would be smaller and with less spiny, more regular, serrated
margins. The new material collected from Madygen shows that these
two species are conspecific, because both margin types are found on
the same leaves (Plate I, 3, 13; arrows). The smaller entirely serrated
leaves appear to belong to a different taxon that is here described as
anewspeciesIsoetites sixteliae.Isoetites sixteliae occurs in a differ-
ent lycopsid assemblage that consists predominantly of herbaceous
lycopsids like Lepacyclotes and Isoetites. All species respectively speci-
mens here placed in the synonymy of M.kotschnevii show the diagnos-
tic features, viz., linear and simple leaves with a single, wide midvein,
no lateral veins and with undulating to serrated margins. Recently,
Mesenteriophyllum has been found in two other Triassic localities.
Sadovnikov (2008) reported Mesenteriophyllum in association with
Pleuromeia and Tundrodendron from Lower Triassic in the eastern
Taimyr. Bomfleur et al. (2011) described a single slab from the Upper
Triassic of the central Transantarctic Mountains containing leaves of
M.serratum associated by an isolated, fragmentary Pleuromeia-like
cone and megaspores. According to the latter authors, the sterile leaves
from Antarctica have undulated margins macroscopically showing a
serrated appearance of the leaf margins. This feature was interpreted
as the result of the collapse of internal air channels. However, in the
Madygen material the leaf margins of Mesenteriophyllum are serrated
at least in some portions (Plate I, 3, 13; arrows). The interpretation
given for the Antarctic specimens seems correct, because specimens
from Madygen of M. kotschnevii show that the marginal portions of
the leaves are undulated (Plate I, 8, 10) or partially even completely
folded (Plate I,11–12). Hence, the nature of the lateral margins is appar-
ently a secondary feature and not a primary character.
So far, leaves of M.kotschnevii have not been found attached to any
stems in the Madygen Formation. However, due to the repeated
co-occurrence with Ferganodendron sauktangensis and Pleuromeiopsis
kryshtofovichii is seems likely that M.kotschnevii might have been the
foliage of one of these taxa.
53P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
Fossil genus Ferganodendron (Sixtel) Dobruskina, 1974 emend. Moisan
et Voigt
Type species Ferganodendron sauktangensis (Sixtel) Dobruskina, 1974
emend. Moisan et Voigt
Emended diagnosis: Stem having longitudinal discontinuous striae.
Leaf bases spirally arranged transversely rhomboidal to elliptical, with
a vascular bundle and two parichnos scars. Rhizomorph conical with
rounded rootlet scars spirally arranged.
Ferganodendron sauktangensis (Sixtel) Dobruskina, 1974 emend.
Moisan et Voigt (Plate II,1–10; Plate III,1–5)
Synonymy and selected references
1936 Pleuromeia oculina, Brick, p. 165–166; pl. 1, 1, 3.
1962 Sigillaria sauktangensis, Sixtel, p. 302–304; pl. 4, 1–6; text-figs. 3–4.
1962 Sigillaria (?) sp., Sixtel, p. 304–305; pl. 4, 7–9.
1962 Pleuromeiopsis kryshtofovichii, Sixtel; p. 305–310; pl. 5, 1, 3–5; pl. 6, 1–2.
1962 Pleuromeiopsis (?) sp., Sixtel, p. 310; pl. 7, 1–3.
1974 Ferganodendron sauktangensis, Dobruskina, p. 389–390; pl. 10, 1–7.
1995 Ferganodendron sauktangensis, Dobruskina, p. 20–22; pl. 1, 1–7; pl. 2, 1–13;
pl. 3, 1–7; pl. 4, 2–5; pl. 5, 5, 8–10; text-fig. 15.
Emended diagnosis: Stem surface with longitudinal discontinuous
striae. Leaf bases variable in shape and size along the stem, transversely
rhomboidal to elliptical, with a vascular bundle and two parichnos
scars. Narrow leaves attached to the terminal portion of the stem.
Rhizomorph conical. Rootlet scars spirally arranged.
Syntypes: No 1371 (?1341), p. 302; pl. 4, 1 (?fig. 4). Sixtel, 1962.
Material studied: FG 596/X/259, X/279, X/305, X/321, X/332, X/379, X/391,
X/442, X/538, X/553, X/587, X/767, X/942, X/1005a–b, X/1032, X/1062.
Repository: Palaeontological collection of the Geologisches Institut,
Technische Universität Bergakademie Freiberg, Germany.
Locality: Vicinity of the village of Madygen, approximately 250 km
west of Osh, southwest Kyrgyzstan, Central Asia (Fig. 1).
Stratigraphic horizon: Madygen Formation.
Age: Middle–Late Triassic (Ladinian–Carnian).
Description: The largest specimen of Ferganodendron sauktangensis
is 14 cm long and 5.5 cm wide (Plate II, 1). The stem bears spirally
arranged, transversely rhomboidal (Plate II,2,4)toelliptical(Plate II,
8–10), 1–1.5 mm long and 2–2.5 mm wide leaf bases. Irregular and dis-
continuous, longitudinal striae are present along the stem surfaces in
stems with smaller leaf bases (Plate II,1,3–4, 7), and at the stem base
where they areinterrupted by the rhizomorph (Plate III,3–4). Specimen
FG 596/X/279 is a stem with a circular scar of a lateral branch that is
1.8 cm in diameter (Plate II, 1; arrow). The shape and size of the leaf
bases are variable, depending on their position on the stem and the
decortication level. The specimen FG 596/X/767 shows well preserved
rhomboidal leaf bases with a central vascular bundle flanked by two
parichnos scars (Plate II,5–6; arrows). Complete rhizomorphs are con-
ical in shape (Plate III,1–2). Rhizomorphs are preserved as moulds
(Plate III, 1, 4) and casts (Plate III, 2, 3, 5); they are densely covered by
oval rootlet scars that are 2–3 mm in diameter, arranged in irregular
parastichies due to the presence of partial orthostichies (Plate III,1,4).
Remarks and comparisons: Sixtel (1962) erected Sigillaria sauktangensis
based on fragmentary material from the Madygen Formation. Sixtel
(1962) indicated in the text the specimen No 1371 (illustrated in pl. 4,
1) as the holotype, but in the figure caption the specimen No 1341 (pl.
4, 4) is mentioned as holotype. Subsequently, Dobruskina (1974) revised
Sixtel's material and transferred Sigillaria sauktangensis to the new mono-
typic genus Ferganodendron. Furthermore, she designated a lectotype for
F.sauktangensis, disregarding Sixtel's choice of the holotype. According to
the ICBN (Art. 9.8) this is an error and should be corrected into an epitype.
The newly collected specimens provide additional data on the morphol-
ogy of F.sauktangensis. New features include the presence of three scars
on the leaf bases, striated stem surfaces and organically connected
rhizomorphs.
The attribution to Sigillaria by Sixtel (1962) was based on the
presence of leaf bases that were interpreted as having three scars.
Dobruskina (1974) described F.sauktangensis as having only one scar
located on the upper part of the leaf bases. The new material collected
in Madygen from the same localities show that some leaf bases indeed
have three leaf scars, representing the vascular bundle and parichnos.
In some aspects Sigillaria vaguely resembles Ferganodendron, i.e., in
the structure and shape of the leaf bases. However, Ferganodendron
apparently lacks a ligule and had lateral branches, as is suggested by an
oval scar basically similar to the so-called ‘ulodendroid scars’in some
lepidodendralean genera (cf. Jonker, 1976), whereas Sigillaria never
have lateral branches. The branch scars in Palaeozoic Lepidodendrales
are much larger (4.5–16 cm long and 4–11 cm wide) than in
F.sauktangensis, which is logical as the former were tall trees. In the
Pennsylvanian lepidodendralean that produce these branch scars
DiMichele (1985) hypothesized that the branches were deciduous, and
it is probable that Ferganodendron also had deciduous lateral branches.
Based on size differences of the leaf bases, Sixtel (1962) established
Pleuromeiopsis kryshtofovichii. According to Dobruskina (1974, 1995)
Plate IX. SEM images of silicone replicas of the surfaces of Isoetites madygensis and Lepacyclotes zeilleri from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan.
1. Proximal portion of the sporophyll of I.madygensis showing the elliptical, basal sporangium. FG 596/X/1004. Scale bar = 1 mm.
2. Epidermal cells of the sporangial portion of I.madygensis showing in the middle portion irregularly arranged rectangular-elongated cells, and isodiametric cells at
the margins. FG 596/X/771. Scale bar=500 μm.
3. Epidermal cells of I.madygensis showing variation in shape and arrangement. FG 596/X/1004. Scale bar= 500 μm.
4, 6. Longitudinally elongated epidermal cells in the middle portion of the sporophyll of I.madygensis. FG 596/X/1004. Scale for fig. 4 = 250 μm. Scale bar for fig. 6 =
100 μm.
5. Mostly tetragonal to pentagonal epidermal cells at the margin of the sporophyll of I.madygensis. FG 596/X/1004. Scale bar = 200 μm.
7. Impressions of in situ megaspores in the sporangium of L.zeilleri. FG 596/X/1011b. Scale bar= 500 μm.
Plate X. SEM images of silicone replicas of lycopsids from the Madygen Formation, Middle–Late Triassic, SW Kyrgyzstan. (see on page 56)
1–2. Epidermal cell pattern of leaves of I.sixteliae showing mostly isodiametric cells and some rectangular cells. FG 596/X/340a. Scale bar for fig. 1 =500 μm. Scale bar for
fig. 2= 200 μm.
3. Detail of Fig. 1 showing isodiametric and rectangular epidermal cells. Scale bar= 50 μm.
4. Rectangular epidermal cells in the central portion of a leaf of I.sixteliae. FG 596/X/745a. Scale bar =100 μm.
5. Irregularly distributed pentagonal-hexagonal to elongate epidermal cells of L.zeilleri. FG 596/X/50. Scale bar =200 μm.
6. Portion of a leaf of I.sixteliae showing the epidermal cell pattern and serrated margin. FG 596/X/918b. Scale bar = 500 μm.
7. Rectangular and isodiametric epidermal cells arranged in longitudinal rows of M.kotschnevii.FG 596/X/769c. Scale bar =200 μm.
8. Epidermal cells of I.sixteliae showing smooth and flat periclinal cell walls. FG 596/X/614. Scale bar= 200 μm.
54 P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
the size and shape of the leaf bases is highly variable in Ferganodendron
sauktangensis, depending on their position along the stem or due to
the level of decortication. All specimens previously assigned to
P.kryshtofovichii appear to belong to F.sauktangensis, except for the
holotype of P.kryshtofovichii which is indeed different from the other
lycopsids from Madygen (see below).
No new specimens of Ferganodendron sauktangensis with attached
leaves have been found. Some specimens attributed to P.kryshtofovichii
by Sixtel (1962) possess a cluster of narrow leaves; these are terminal
portions of F.sauktangensis stems (cf. Dobruskina, 1974, 1995). The
newly collected material of F.sauktangensis also comprises rhizomorphs
found in organic connection with stems. These rhizomorphs show
some superficial similarities to those of Pleuromeia.Thefigured rootlet
scars (Plate III, 1) resemble those of Pleuromeia sternbergii from the
Buntsandstein of Germany (Mader, 1990; Grauvogel-Stamm, 1993,
1999). The three-dimensional preservation of P.sternbergii shows that
its rhizomorph consists of a more or less rounded base with four
upward-directed lobes. However, it is not possible to determine
whether the rhizomorphs F.sauktangensis were divided into lobes or
not. Although, Ferganodendron shows some similarities to Pleuromeia,
we agree with Dobruskina (1974, 1995) that both taxa can be clearly
separated based on structure of the leaf bases and the size of the
plant. Moreover, typical Pleuromeia fructifications have never been
found in Madygen, neither strobili nor sporophylls that are commonly
found in other localities from which Pleuromeia has been reported.
Dobruskina (1974) indicated that Ferganodendron stems could reach a
diameter of 20–30 cm, but it is evident that she overestimated the
size of the plant. The subarborescent lycopsid Lycopia dezanchi from
the Anisian of the Dolomites (Kustatscher et al., 2010) differs mainly
with Ferganodendron by having a dichotomous rhizome, stems bifurcat-
ing apically, and leaves more closely spaced along thestem. Moreover, L.
dezanchi lacks longitudinal striae on the stem surface, and the leaf scars
are generally similar in size and shape along the stem and parichnos and
vascular bundle scars are unknown.
Fossil genus Pleuromeiopsis Sixtel, 1962
Type species Pleuromeiopsis kryshtofovichii Sixtel, 1962
Pleuromeiopsis kryshtofovichii Sixtel, 1962 (Plate III,6–7)
Selected references
1936 Pleuromeia oculina, Brick, p. 165–166; pl. 1, 2.
1962 Pleuromeiopsis kryshtofovichii, Sixtel, p. 305–310; pl. 5, 2;
text-figs. 5–6.
1967 Pleuromeiopsis kryshtofovichii,Chaloner et Boureau,p.654–655;
text-figs. 450A–B.
1995 Pleuromeiopsis kryshtofovichii, Dobruskina, pl. 5, 1.
Material studied: FG 596/X/554, X/584.
Description: Fragments of stems 10 cm long and 6 cm wide. Leaf bases
5–6mm long and 1–1.3 cm wide, rhomboidal-elliptical, separated by
distinct transverse striae, which more or less follow the parastichy
arrangement of the leaf scars (Plate III,6–7). The centre of the leaf bases
showsanellipticalscar(Plate III, 6; arrows), c oncave or convex depending
onthetypeofpreservation.Vascular bundle and parichnos are not
observable. Reproductive structures and epidermal features unknown.
Remarks and comparisons: Pleuromeiopsis was introduced by Sixtel
(1962) with the single species P.kryshtofovichii, based on stem frag-
ments with attached leaves and roots with large scars. However, all
the specimens assigned to P.kryshtofovichii, except for the holotype,
were misinterpreted by Sixtel (1962), viz., the stems belong to the
lycopsid Ferganodendron sauktangensis, the roots with large scars are
galls on gymnosperm leaves, and the strobili are ovulate organ of the
seedfern Peltaspermum (Dobruskina, 1995). Sixtel's holotype is the only
specimen that is not comparable to any other lycopsid from Madygen.
The leaf bases on the stem of P.kryshtofovichii are considerably larger
than those of F.sauktangensis, and their structure is different having a
single elliptical scar. The stem shows striae like F.sauktangensis,butin
P.kryshtofovichii striae are transversely oriented and continuous. Our
new collection from Madygen includes two specimens that show the fea-
tures to be seen in Sixtel's holotype of P.kryshtofovichii. The specimens
were found associated by M.kotschnevii,F.sauktangensis and other inde-
terminate lycopsids. However, because of the very limited number of
specimens it is difficult to prove the true identity of P.kryshtofovichii;
we agree with Dobruskina (1974, 1995) that the genus Pleuromeiopsis
should be retained until more information becomes available.
Order Isoetales
Fossil genus Isoetites Münster, 1842 emend.Kustatscher et al., 2010
Type species Isoetites crociformis Münster, 1842
Isoetites madygensis sp. nov. Moisan et Voigt (Plate VI,1–13; Plate
VII,4–7; Plate IX,1–6)
Synonymy
?1995 Annalepis (Tomiostrobus)leae, Dobruskina, p. 22; pl. 4, 6.
Diagnosis: Corm very short, wide, unlobed, bearing spirally arranged
sterile leaves and sporophylls. Sporophylls and sterile leaves ovate,
Fig. 2. Original illustration of Mesenteriophyllum kotschnevii by Sixtel (1961, 1962).
57P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
lanceolate. Ligulate sporophylls with a basal and ellipsoid-ovate sporangi-
um. Sterile leaves with air channels and a thin midvein. Normal epidermal
cells of sporophylls and sterile leaves more or less rectangular, slightly
elongate, aligned in longitudinal rows; cells surrounding the sporangium
isodiametric (pentagonal) and rectangular. Stomata and papillae absent.
Holotype: FG 596/X/54 (Plate VI,7–8).
Material studied: FG 596/X/52–54, X/384, X/392, X/601, X/619,
X/666a–b, X/669a–b, X/687, X/732, X/740(a–b)–741, X/749, X/755,
X/763, X/771, X/776, X/780, X/845, X/913a–b, X/978a–b, X/985a–b,
X/1004a–b, X/1009.
Repository: Palaeontological collection of the Geologisches Institut,
Technische Universität Bergakademie Freiberg, Germany.
Locality: Vicinity of the village of Madygen, approximately 250 km
west of Osh, southwest Kyrgyzstan, Central Asia (Fig. 1).
Stratigraphic horizon: Madygen Formation.
Age: Middle–Late Triassic (Ladinian–Carnian).
Etymology: The specific epithet refers to the Madygen Lagerstätte
from which the fossils were collected.
Description: Isoetites madygensis consists of nearly complete speci-
mens including three corms with attached sporophylls and sterile
leaves (Plate VI,1–5, 7–8), isolated sporophylls with a basal sporangium,
sterile leaves with air channels and well-preserved epidermal cells. The
holotype FG 596/X/54 (Plate VI,7–8) consists of a corm 8 mm long and
1.5 cm wide, with attached sterile leaves being 4–5mm wide atthe
base; rootlets scars are visible as small depressions of about 1 mm in
diameter (Plate VI,8).Thesameslabshowsseveralisolated,upto3cm
long ovate-lanceolate leaves with a thin midvein (Plate VI, 7). Specimen
FG 596/X/740 consists of a 6 mm long and 1.5 cm wide corm, bearing
three sterile 5–6 mm wide, basally expanded leaves (Plate VI,4–5); the
corm shows circular rootlet scars. Specimen FG 596/X/985a–b consists
of at least fifteen sporophylls arranged in the form of a ring (incomplete)
that is c. 2.5 cm in diameter (Plate VI,1).Thesporophyllsareovate-
spatulate of 6–9mm wide, bearing a basal c. 2 mm wide ovate-ellipsoid
sporangium. Sporophylls show a ligule distal to the sporangium (Plate
VI,2–3 arrows). The same slab also shows a small corm fragment with
sporophylls showing the helically overlapping bases (Plate VI, 3; arrow).
Leaves or sporophylls are ovate-lanceolate, 4–5mm wideat thebase
and at least 3.5 cm long; they lack a midvein (Plate VI, 6). Several isolated
sporophylls basally show an elliptical, 2–4mmlongand1–2mmwide
sporangium (Plate VI,9–13; Plate IX, 1). Sterile leaves show a regular sur-
face pattern of marginal undulations (Plate VII,4–7).
Epidermal cells in the basal portion of the sporophylls, around
the sporangia, irregularly distributed, isodiametric (pentagonal) to
elongate, 90–111 μm long and 53–64 μm wide (Plate IX,1–3, 5).
Other cells are tetragonal to rectangular-elongate, 59–117 μm long
and 34–60 μm wide, arranged in longitudinal rows (Plate IX, 4, 6).
Stomata, papillae and trichomes absent.
Isoetites sixteliae sp. nov. Moisan et Voigt (Plate V,1–15; Plate VII,1,
3; Plate X,1–4, 6, 8)
Synonymy
1995 Mesenteriophyllum serratum, Dobruskina, p. 32; pl. 42, 9–10, 13–17.
Diagnosis: Leaves linear, simple and narrowly lanceolate, tapering
to a sharp point (grass-like), with a thin midvein. Leaves with an
expanded base (?sporophyll). Leaf margins serrated. Leaves with longi-
tudinal air channels separated by transversesepta. Epidermal cells rect-
angular to isodiametric arranged in rows. Stomata and papillae absent.
Holotype: FG 596/X/918 (Plate V, 7, 13; Plate X, 6).
Material studied: FG 596/X/42–43, X/231(a–b)–234, X/308a–b, X/310,
X/316, X/318, X/322b, X/334, X/340a–b, X343, X/367a–b, X/371a–b,
X/378, X/382, X/384a, X/396–397, X/427(a–b)–429, X/508–513, X/
519, X/541, X/549, X/560, X/576, X/590–591, X/603, X/606–607, X/
614–615a–b, X/616, X/629, X/660, X/675b, X/680, X/686a–b, X/689,
X/712, X/730, X/735, X/744–745a–b, X/748a–b, X/759, X/765(a–c)–
766, X/772, X/785, X/896, X/915, X/918(a–b)–919, X/980, X/983a–
b, X/1015, X/1018, X/1023a–b.
Repository: Palaeontological collection of the Geologisches Institut,
Technische Universität Bergakademie Freiberg, Germany.
Locality: Vicinity of the village of Madygen, approximately 250 km
west of Osh, southwest Kyrgyzstan, Central Asia (Fig. 1).
Stratigraphic horizon: Madygen Formation.
Age: Middle–Late Triassic (Ladinian–Carnian).
Etymology: The specific epithet is proposed in honour of Tatiana A.
Sixtel, for her contributions to the study of fossil plants in Madygen.
Description: Eighty-three specimens are assigned to Isoetites sixteliae
including sterile leaves with well-preserved epidermal cells, air chan-
nels and putative sporophylls. The largest leaf is about 12 cm and be-
tween 3–5 mm wide (Plate V, 8). Leaf margins are serrated with
pointed teeth about 0.5 mm which are longitudinally separated
from each other by 2–3mm. Leaves ending in a pointed apex (Plate
V,1–2). All leaves have a thin midvein (Plate V,1–8, 10, 12). Air chan-
nels are represented by rectangular, 1–1.5 mm long and 0.5 mm wide
imprints that are separated by septa (or diaphragms) at regular inter-
vals (Plate VII, 1, 3). Air channels and septa are connected to the
midvein. Leaves with expanded bases (?sporangial portion) of a max-
imum width of 1 cm (Plate V,13–15) were found in the same slabs
with the sterile leaves.
Epidermal cells are 38–70 μm long and 25–48 μm wide. The epi-
dermal cell pattern and arrangement are simple, and the cells do
not show significant variations in shape and size (Plate V,11–12;
Plate X, 1, 6). Isodiametric cells are mostly found near the leaf margin
(Plate V, 9, 11; Plate X,2–3) and rectangular-elongated cells near the
central portion (Plate V, 12; Plate X, 4). Periclinal cell walls are
smooth and flat (Plate X,1–4, 8), without any evidence of papillae
or trichomes.
Remarks and comparisons: The genus Isoetites Münster has been used
to designate fossils resembling the extant genus Isoëtes L. (often
misspelled as Isoetes; see ICBN Art. 60.6, McNeill et al., 2006). Some
authors use the name Isoëtes/Isoetes for fossil specimens, e.g., Bock
(1962, 1969),Hill (1987),Banerji (1988),Wang (1991),Retallack
(1997),Srivastava et al. (2004),Naugolnykh and Mogutcheva (2006).
Other authors use the name Isoetites for fossils that are reminiscent of
the extant genus Isoëtes but that are incompletely known or differ in
some respects, e.g., Brown (1939, 1958),Teixeira (1948),Bose and
Roy (1964),Bose and Banerji (1984),Ash and Pigg (1991),Barale
(1999),Kustatscher et al. (2010). Because our material is still incom-
pletely known, i.e., the megaspores and microspores are unknown
and not all specimens show a ligule, we classify our material in Isoetites.
However, I.madygensis so far seems tobe the fossil species that is most
similar to extant Isoëtes, particularly with regard to the presence of air
channels and the compact, non-elongated corm especially such as in
58 P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
the young stages of the extant Isoëtes tuckermanii (Karrfalt and Eggert,
1977). Only a single previously described specimen from Madygen, a
sporophyll, can be attributed to Isoetites. To accommodate this speci-
men Dobruskina (1995) established a new species of Annalepis,A.
leae, but it does not show the diagnostic features of Annalepis.This
specimen might belong to Isoetites madygensis.Isoetites sixteliae dif-
fers clearly from Isoetites madygensis because of the serrated margin
in their leaves that is absent in the latter species. Several fossil spe-
cies of Isoëtes and Isoetites have been described; for similarities and
differences with Isoetites madygensis and Isoetites sixteliae we refer
to Table 1.
The overall appearance of Isoetites is similar to that of the
Palaeozoic Cormophyton (Skog and Hill, 1992). Sporophylls described
as Sadovnikovia belemnoides from the Lower Permian of the Urals
(Naugolnykh, 1994)andViatcheslavia vorcutensis from the Upper
Permian of the Urals and Russian Platform (Naugolnykh, 2001;
Naugolnykh and Zavialova, 2004) are similar to I.madygensis in having
a reduced basal sporangium and lanceolate sporophylls. However, both
Russian species are associated with stem fragments and were regarded
as arborescent pleuromeialeans. Similarities between these Russian
lycopsids and Isoëtes were already pointed out by Naugolnykh (1994),
who postulated an evolutionary series based on sporophyll morphol-
ogy, beginning with S.belemnoides as the oldest known ancestor of
Isoëtes.
The epidermal pattern of the sporangium wall is considered one of
the typical characters of Isoëtes (Prada and Rolleri, 2005; Rolleri and
Prada, 2007). In I.madygensis the epidermal cells surrounding the
sporangium are irregular in shape, isodiametric to pentagonal. Variations
in shape and size have been documented for several Isoëtes species,
e.g., Pant and Srivastava (1962),Budke et al. (2005),Prada and Rolleri
(2005),Rolleri and Prada (2007). Epidermal cells in the sporophylls of
I.madygensis are rectangular arranged in longitudinal rows. This cell
patternisalsocommoninrecentIsoëtes species.
Isoetites madygensis and Isoetites sixteliae lack stomata on its leaf
surface. When stomata are present in extant Isoëtes species, they
are distributed in longitudinal rows (Masarati and Thomas, 1982;
Musselman and Roux, 2002; Prada and Rolleri, 2003), and commonly
restricted to the apical portion of the leaves (Rolleri and Prada, 2007).
However, the presence or absence of stomata in Isoëtes appears to be
dependent of the growing conditions (Kott and Britton, 1985). Prada
and Rolleri (2003) demonstrated that the absence of stomata is not
exclusively related to adaptation to aquatic habitats, since they
found stomata in all the amphibious species they studied, but only
two of the six aquatic species have stomata. The vegetative characters
of Isoëtes are considered of limited taxonomic value due to infraspecific
variability or interspecific uniformity (Hickey, 1986a; Taylor and Hickey,
1992; Prada and Rolleri, 2003).
The plant surfaces of I.sixteliae show excellently preserved air
channels identical to those seen in modern Isoëtes (Plate VII,1–3; cf.
Liu et al., 2006; Roux et al., 2009). We interpret the transverse undu-
lations of I.madygensis as the result of the collapse of internal air
channels, as has already been suggested for other lycopsids with sim-
ilar structures (Skog et al., 1992; Pigg, 2001; Bomfleur et al., 2011).
Air channels are present in both extant Isoëtes and fossil Isoëtes/
Isoetites species indicating that primitive forms were aquatic (Taylor
and Hickey, 1992). Air channels facilitate the diffusion of CO
2
taken
up from the surrounding soil to the leaves for photosynthesis
(Madsen et al., 2002; Pedersen et al., 2011). The absence of stomata
in fossils Isoëtes/Isoetites is also considered as an indication that fossil
taxa grew in aquatic habitats (Srivastava et al., 2004). However, it
remains uncertain whether Isoetites sixteliae and Isoetites madygensis
were aquatic plants, because the above mentioned extant aquatic
Isoëtes species also lack stomata (Prada and Rolleri, 2003). The sporo-
phylls of Isoetites sixteliae lack a ligule, however, even some recent
species such as Isoëtes eludens have eligulate sporophylls (Roux et
al., 2009).
Serrated margins in modern Isoëtes are unknown, even in the fully
alate I.gigantea and I.bradei (R.J. Hickey, personal communication,
2010). These species belong to the subgenus Euphyllum (Hickey,
1990). Isoetites sixteliae represents the first and oldest evidence of
serrated margins in Isoetites.Cidarophyton rewanense described from
the Early Triassic of Australia by Chaloner and Turner (1987) consists
of a spheroidal unbranched axis interpreted as a lycopsid cone axis.
More recently, Cantrill and Webb (1998) pointed out that C. rewanense
is best interpreted as a corm, due to its similarity to Isoëtes. The corm of
Isoetites madygensis is similar in shape and size to those of C. rewanense,
however, sporophylls or leaves are unknown in the Australian species
which precludes further comparisons.
Nathorstiana arborea from the Lower Cretaceous of Germany
(Richter, 1909; Mägdefrau, 1932) is known from different ontogenetic
stages of plant growth (Karrfalt, 1984). Young developmental stages
of the unlobed corm of Nathorstianaare similar to those of I.madygensis,
but in Nathorstiana the corm is notoriously elongated and its reproduc-
tive structures are unknown.
Fossil genus Lepacyclotes Emmons, 1856
Type species Lepacyclotes circularis Emmons, 1856
Lepacyclotes zeilleri (Fliche) Retallack, 1997 (Plate VII,8–13; Plate
VIII,1–6, 8–9; Plate IX,7;Plate X,5)
Synonymy and selected references
1910 Annalepis zeilleri, Fliche, p. 267–273; pl. 27, 3–5.
1983 Annalepis zeilleri, Grauvogel-Stamm et Duringer, p. 28–39; pl. 1-6; text-figs.
2a–b; text-fig. 3e.
1991 Isoetes ermayinensis, Wang, p. 13–14: text-figs. 3a–k, 4a–b, 5a–c, 6a–j, pl. 1;
1–15; pl. 2, 11–19; pl. 3, 1–2, 4–12.
1995 Annalepis (Tomiostrobus)leae, Dobruskina, p. 22; pl. 4, 7.
1995 Annalepis zeilleri, Kelber et Hansch, p. 54; figs. 112–115; p. 95; figs. 199–200.
1997 Lepacyclotes zeilleri, Retallack, p. 508; fig. 1; table 1.
1998 Annalepis zeilleri, Meng, p. 772–773; pl. 2, 1–22; text-figs. 1b–c, 3–4.
2000 Annalepis zeilleri, Meng, p. 155–161; pl. 2, 17–29; pl. 3, 1–17; pl. 4, 19.
2001 Annalepis zeilleri, Grauvogel-Stamm et Lugardon, p. 127–132; figs. 5a–f, 6a, 7a, 8a.
2004 Annalepis zeilleri, Kustatscher et al., p. 58–59; pl. 1, 1.
2005 Annalepis zeilleri, Kustatscher et Van Konijnenburg-van Cittert, p. 34.
2008 Annalepis zeilleri, Kustatscher et VanKonijnenburg-van Cittert, p. 68; figs.1A–B.
2009 Annalepis zeilleri, Bernard et al., p. 255–257; pl. 1, 1–3; pl. 4, 1.
2010 Annalepis zeilleri, Kustatscher et al., table 3.
Material studied: FG 596/X/39–41, X/48–50, X/130a–b, X/518, X/
551, X/585, X/628, X/741–742, X/771, X/947, X/985c, X/992a–b,
X/1011a–b.
Description: Complete sporophylls, 3–3.9 cm long and 5–14 mm
wide, with an acuminate to mucronate apex (Plate VII,8,10–11)
and a trapezoidal proximal portion (Plate VII, 9, 11). Smaller sporo-
phylls are 1–1.2 cm long and 4–6 mm wide with a much longer
apex (Plate VIII,8–9). The widest portion of the sporophyll is near
the apex (distal). The sporangium is 3–5 mm wide, central, large
and oblong to slightly obovate reaching a length of c. 3/4 of the
total sporophyll length, where the sporophyll has its maximum
width (Plate VII, 11; Plate VIII,1–2, 8–9). In some specimens the spo-
rangium is symmetrically divided in two by a vertical midline (Plate
VII, 11 arrows; Plate VIII,2,8–9). The ligule is elliptical, 2–2.5 mm
long and 1–1.5 mm wide (Plate VII,10–11; Plate VIII,1–3 arrows).
Impressions of megaspores on the sporophyll surface are 485–550 μm
in diameter (Plate VIII,1;Plate IX,7).
Epidermal cells, 48–71 μm in diameter, mostly pentagonal-
hexagonal (Plate VIII,4–6; Plate X, 5), occasionally rectangular
(Plate X, 5), not aligned in distinct rows, only near the apex that has
a distinct vertical band of c. 700 μmwide of rectangular-elongated
cells (Plate VIII, 5; arrows) surrounded by pentagonal-hexagonal
cells.
59P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
Table 1
Comparison of Mesozoic species of Isoëtes/Isoetites (data taken from: Saporta, 1894; Brown, 1939, 1958; Teixeira, 1948; Bose and Roy, 1964; Bose and Banerji, 1984; Banerji, 1988; Ash and Pigg, 1991; Wang, 1991; Skog and Hill, 1992; Skog et
al., 1992; Retallack, 1997; Barale, 1999; Naugolnykh and Mogutcheva, 2006; Kustatscher et al., 2010).
Taxon Leaves Leaf
margin
Air
channels
Sporangium Ligula Rhizomorph Epidermal cells Occurrence Age
Isoetites madygensis sp. nov. Ovate-lanceolate Entire Present Ovate Present Non-elongated corm Isodiametric-pentagonal,
rectangular
Madygen, Kyrgyzstan Middle–Late Triassic
Isoetites sixteliae sp. nov. Lanceolate, pointed apex Serrate Present ––– Rectangular-isodiametric Madygen, Kyrgyzstan Middle–Late Triassic
Isoëtes beestonii Retallack, 1997 Cordate, transverse
undulations
Entire –Elliptical Sunken
Glossopodium
Rounded to crudely
tetragonal corm
–Blackwater, Queensland,
Australia
Early Triassic
Isoetites brandneri
Kustatscher et al., 2010
Elongated, folded Entire –– – Unlobed, bulb-like Isodiametric, elongated Kühwiesenkopf, Dolomites,
Italy
Middle Triassic
Isoetites choffati Saporta, 1894 2–3 cm long with a pointed
lamina
Entire –Oblong –Bulbous base –Cercal, Portugal Early Cretaceous
Isoetites daharensis Barale, 1999 Ribbon shaped, dilated at
base, spatulate
Entire –Elongate Present Unlobed spherical or
conical corm
–Tataouine, Tunisia Early Cretaceous
Isoëtes ermayinensis Wang,
1991
Lanceolate, spatulate Entire Present Obovate or
wedge-shaped
Present Dichotomously branched –Shanxi, China Middle Triassic
Isoetites horridus Brown, 1939 Spatulate-tipped Entire Present Elliptical –Unlobed corm –Wyoming, USA Late Cretaceous
Isoëtes innae Naugolnykh
and Mogutcheva, 2006
Linear, ovoid, spine-like Entire –Ovoid –Elongated corm –Tunguska Basin, Siberia,
Russia
Early Triassic
Isoëtes janaianus Banerji, 1988 Lanceolate Entire –Oval-oblong –Five-lobed Rectangular Bhuj, India Middle–Late Jurassic
Isoetites phyllophila Skog
et al., 1992
Dilated at base, tips
acuminate
Entire Present ––Erect, elongate, conical
corm
Elongate, rectangular Central Kansas, southcentral
Nebraska, USA
Cretaceous
Isoetites rolandii Ash and
Pigg, 1991
Elongate, linear Entire –Oval-oblong –Cormlike plant base –Hells Canyon, Oregon, Idaho,
USA
Middle Jurassic
Isoetites serratus Brown, 1939 Strap-shaped with
spatulate ends
Serrate Present ––?Rounded corm –Wyoming, USA Late Cretaceous
Isoetites serratifolius Bose and
Roy, 1964
Elongate Serrate –Obovate –– – Northwestern India Early Jurassic
60 P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
Remarks and comparisons: Fliche (1910) originally described the
genus Annalepis from the Lettenkohle (Lower Keuper, late Middle
Triassic) of eastern France, and suggested a coniferalean affinity.
Grauvogel-Stamm and Duringer (1983) demonstrated that Annalepis
zeilleri was a lycopsid sporophyll, based on the presence of micro-
and macrosporophylls containing Aratrisporites microspores and
Tenellisporites megaspores. Retallack (1997) transferred Annalepis
zeilleri to the genus Lepacyclotes based on similarities of the sporo-
phylls, i.e., pointed/mucronate sporophyll apex. Kustatscher et al.
(2010) accepted that Lepacyclotes and Annalepis are congeneric,
the former having priority. Lepacyclotes circularis Emmons, the
type of the genus, consists of a rosette of apically serrated leaves
attached to a short spherical to lobed corm (Bock, 1969; Retallack,
1997).
Lepacyclotes (al. Annalepis)zeilleri is a typical sporophyll found in
the Triassic of France, (e.g., Grauvogel-Stamm and Duringer, 1983),
Germany (Kelber and Hansch, 1995; Kelber, 1998; Kustatscher and
Van Konijnenburg-van Cittert, 2008) China (Meng, 1994, 1996,
1998, 2000; Yu et al., 2010) and Italy (Kustatscher et al., 2004;
Kustatscher and Van Konijnenburg-van Cittert, 2005). Our material
of L. zeilleri is the first unequivocal record of Lepacyclotes (al. Annalepis)
in Central Asia. Although organic remains are not preserved,
numerous impressions of megaspores can be seen on the surface
of the sporangium. They are similar in size to those described
by Grauvogel-Stamm and Duringer (1983) and Bernard et al.
(2009).
The lycopsid genus Tomiostrobus originally described from the
Permian of the Kuznetsk Basin (Neuburg, 1936) was defined for
sporophylls with a distal limb. Sadovnikov (1982) gave an emended
diagnosis and described five additional species. Sadovnikov (1982)
and Retallack (1997) included Skilliostrobus (Ash, 1979)inTomiostrobus,
because they regarded the sporophylls as almost identical. Kustatscher
et al. (2010) doubted whether Tomiostrobus (Skilliostrobus) represents
only isolated sporophylls or sporophylls helically attached to a peduncu-
late cone-like structure. The smaller sporophylls from Madygen (Plate
VIII,8–9) with a longer acuminate apex are assigned to Lepacyclotes
zeilleri, because they have a vertical midline that symmetrically divides
the sporangium, a typical feature for L.zeilleri (Grauvogel-Stamm and
Duringer, 1983; Grauvogel-Stamm and Lugardon, 2001). Cylostrobus
sydneyensis represents sporophylls spirally arranged in a cone (Helby
and Martin, 1965) which resemble those of L.zeilleri but the sporangia
lack of the distinctive midline.
Indeterminate lycopsid remains
Lycopsid stems (Plate III,8–9)
Material studied: FG 596/X/150a–b, X/474.
Description: Two specimens and a counterpart, 11 cm long and
8–11 mm wide, consist of stem fragments (Plate III,8–9). Leaf scars,
2–3 mm long and 1 mm wide, are vertically rhomboidal in outline,
arranged in parastichies (Plate III,8–9).
Isolated rhizomorphs (Plate IV,9–11)
Material studied: FG 596/X/1008a–c.
Description: The single specimen and its counterpart consists of a
rhizomorph structure of c. 4 cm in diameter, with long and thin root-
lets of more than 3 cm long and 2–4 mm wide (Plate IV,9–11).
Isolated lycopsid leaves (Plate IV,1–8)
Material studied: FG 596/X/386, X/424, X/464, X/506, X/632, X/682,
X/909, X/912, X/916.
Description: Leaves simple, 6.5 cm long and 8 mm wide (Plate IV,
1–2), gradually tapering towards the apex (Plate IV, 6). Both margins
of the leaves are strongly and regularly serrated (?spines). Teeth
between 1 to 3 mm long, depending on the size of the leaves (Plate IV,
1–8). Some specimens appear to have a midvein (Plate IV,2,8;arrows).
Leaves, simple and entire margined, were found in the same slab with
these leaves (Plate IV,8).
Comments: These specimens are described as indeterminate lycopsids,
due to the absence of distinctive features. All these lycopsid remains
occur in the same beds with Ferganodendron sauktangensis,Pleuromeiopsis
kryshtofovichii and Mesenteriophyllum kotschnevii.
4. Discussion
4.1. Diversity and evolutionary significance of the Madygen lycopsids
The new lycopsid fossils f rom the Madygen Formation represent a
diverse suite of forms, including the subarborescent Ferganodendron
sauktangensis,Pleuromeiopsis kryshtofovichii and Mesenteriophyllum
kotschnevii that resemble the typical Pleuromeia-type from the
Early Triassic in their general appearance, and the herbaceous
lycopsids Isoetites madygensis,Isoetites sixteliae and Lepacyclotes
zeilleri that resemble the extant Isoëtes. The Anisian flora from
the Dolomites also shows a mixture of subarborescent and herba-
ceous forms (Kustatscher et al., 2010). However, the Madygen flora
is younger, being of Ladinian–Carnian age. This is the singularity of
the lycopsid flora of Madygen, considering that from the Ladinian
onward, the diversity and abundance of lycopsids decreased
considerably. According to Dobruskina (1995) lycopsids are the
poorest represented group in the Madygen flora, and she assigned
all lycopsid remains to Ferganodendron. New information on
F.sauktangensis reveals that this taxon shares some features with
Pleuromeia. However, it should be noted that F.sauktangensis show
some significant differences with Pleuromeia such lateral branching,
structure of the leaf bases and root system. So far, Ferganodendron
sauktangensis appears to be endemic to Madygen. Several organisms
from Madygen also show a restricted geographical distribution such
as various vertebrates (Sharov, 1970, 1971; Tatarinov, 2005; Voigt et
al., 2006; Buchwitz and Voigt, 2010; Schoch et al., 2010), insects
(e.g., Shcherbakov, 2008a,b; Béthoux et al., 2010)andpterido-
sperms (Sixtel, 1962; Dobruskina, 1995). The proportionally high
number of endemic taxa in Madygen may be explained by the fact
that our current knowledge of terrestrial Triassic ecosystems is
based primarily on data from tropical and subtropical latitudes,
whereas the Madygen Formation was deposited an intramontane
basin at a palaeolatitude of 35°–40°N (Voigt et al., 2006; Moisan
et al., 2011).
The unequivocal presence of Lepacyclotes (al. Annalepis)zeilleri in
the Madygen biota fills a gap on its populations that showed a dis-
junct distribution between China and Western Europe. The occur-
rence of L.zeilleri in Madygen suggests a uniform geographical
distribution of this taxon from China through Central Asia to Western
Europe. Lepacyclotes (al. Annalepis)zeilleri has been considered by
some authors within the Pleuromeiaceae/Pleuromeiales (Thomas
and Brack-Hanes, 1984; Taylor et al., 2009) or Lepidodendrales
(Grauvogel-Stamm and Duringer, 1983; Skog and Hill, 1992). We
agree with Retallack (1997) that L.zeilleri must be included in the
Isoetales, considering that it is ligulate, heterosporous, with an herba-
ceous habit, and moreover shares some specific similarities with Isoëtes
such as ultrastructure of the spores (Grauvogel-Stamm and Lugardon,
2001). It has been suggested that Lepacyclotes (al. Annalepis)maybe
ancestral to Isoëtes,viaIsoetites (e.g., Meng, 1998; Grauvogel-Stamm
and Lugardon, 2001; Liu et al., 2004). However, it must also be consid-
ered that the contemporary Triassic species Isoetites madygensis
and I.sixteliae already had lacunate leaves and a reduced, short
corm as in Isoëtes. The evolutionary history of Isoëtes is complex in-
volving a combination of convergence, divergence, parallel and retic-
ulate evolution and gene silencing (Taylor and Hickey, 1992; Caplen
and Werth, 2000).
61P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
According to Pigg (1992) no Triassic Isoëtes-like forms show
evidence of a reduced corm, which is typical for modern Isoëtes.
Nevertheless, the reduced corm appears to be an ancestral condi-
tion in the evolutionary lineage of Isoëtes that first evolved in the
Middle–Late Triassic in Isoetites madygensis. Throughout the entire
Mesozoic, taxa assigned to Isoëtes/Isoetites show an elongated
corm, except for the Middle Jurassic Isoetites rolandii (Pigg, 2001).
All 150–200 extant species of Isoëtes have also a short, reduced
corm, except for the terrestrial I.andicola that was formerly in-
cludes in Stylites, which has an elongated, branched corm (Keeley
et al., 1984). Hickey (1990) pointed out that certain fossil Isoetites
appear to belong to the subgenus Euphyllum. This implies that I.
sixteliae and I.madygensis can be considered as the oldest known
ancestral taxa of this subgenus that had a cosmopolitan distribution
during the Mesozoic and it is represented today only by three
relictual species (i.e., Isoëtes gigantea,I.bradei and I.baculata)in
South America. It has been postulated that the subgenus Isoëtes
had its origin in Gondwana and subsequently radiated northward
into Laurasia (Hickey, 1986b; Taylor and Hickey, 1992). However,
both new species described here show several characters (i.e., air chan-
nels, non-elongated corm) that are typical of extant Isoëtes.Thismay
suggest that the origin of Isoëtes may have been in Asia during the
Triassic.
4.2. Palaeoenvironmental implications
Lycopsids were dominant elements of wetland plant communities
in Madygen. Wetland areas include a variety of habitats characterized
by a high diversity and they can act as refugia for many plants and
animals species (Greb et al., 2006; Gopal, 2009). Hydrophytes
adapted to moist and wet conditions are the typical plants of wetland
environments (Greb et al., 2006).
Hydrophytic species of Isoëtes have internal air channels and lack
stomata or only have non-functional stomata. These features are
regarded as an adaptation to semi-aquatic and aquatic environments
(Taylor and Hickey, 1992). Green (2010) proposed the Lycopsid
Photosynthetic Pathway (LPP) for hydrophytes that involves
the concentration of sedimentary CO
2
in internal air channels
(aerenchyma) in environments with high O
2
and low CO
2
concen-
trations in order to reduce photorespirative loss as well as root
aeration via the parichnos system. This unusual metabolism is
common in Isoëtes inhabiting environments with low CO
2
,and
was probably also utilized by Carboniferous arborescent lycopsids
(Green, 2010). The morphological adaptations related to this me-
tabolism are present in the Madygen lycopsids, and therefore it is
probable that this metabolism was applied by Madygen lycopsids
in wetland ecosystems. This interpretation is supported by the presence
of air channels and absence of stomata in Isoetites madygensis,Isoetites
sixteliae and Mesenteriophyllum kotschnevii,andbythepenetrativeroot
system in Ferganodendron sauktangensis that can be interpreted
as being embedded in the peat/soil to increase efficiency of CO
2
fixation.
Acknowledgments
This research was partially supported by the Deutscher Akademischer
Austausch Dienst (DAAD grant A/06/27956 to P.M.) and the Deutsche
Forschungsgemeinschaft (DFG grant VO 1466/1–1toS.V.).Thefirst
author thanks Hans Kerp (Münster) for the valuable discussion
and for critically reading the manuscript. He also wishes to thank
Serge V. Naugolnykh (Moscow), R. James Hickey (Oxford, OH) and
Benjamin Bomfleur (Lawrence, KS) for providing valuable informa-
tion and literature, and to Ilja Kogan (Freiberg) for the translations
of the original Russian diagnoses and descriptions of F.sauktangensis,
P.kryshtofovichii and M.kotschnevii. This manuscript was im-
proved by detailed and constructive comments of two anonymous
reviewers.
Taxa Locality Horizon Interpretation
Lepacyclotes zeilleri;Isoetites madygensis;
Ferganodendron sauktangensis;
Mesenteriophyllum kotschnevii;
lycopsid stem
LI/3: Urochishche Madygen,
R 0606721/H 4431875
Lower Grey Member (T2) Lacustrine, near-shore
Isoetites madygensis;Isoetites sixteliae LI/7: Urochishche Madygen,
R 0604653/H 4432504
Brown-grey Member (T5) Lacustrine, offshore
Isoetites madygensis;Isoetites sixteliae;
Lepacyclotes zeilleri
LI/8: Urochishche Madygen,
R 0603654/H 4431998
Brown-grey Member (T5) Lacustrine, offshore
Isoetites madygensis LI/9: Urochishche Madygen,
R 0604647/H 4432160
Upper Grey Member (T4) Lacustrine, proximal to distal delta-front
Isoetites madygensis; isolated rhizomorphs LI/13: Urochishche Madygen,
R 0604998/H 4431663
Upper Grey Member (T4) Lacustrine, littoral to sublittoral (delta-front)
Lepacyclotes zeilleri LI/21: Urochishche Madygen,
R 0604526/H 4432601
Brown-grey Member (T5) Lacustrine, sublittoral
Lepacyclotes zeilleri;Pleuromeiopsis
kryshtofovichii;Mesenteriophyllum
kotschnevii;Ferganodendron
sauktangensis; isolated lycopsid leaves
LI/38: Urochishche Madygen,
R 0605629/H 4432382
Lower Grey Member (T2) Lacustrine, littoral to sublittoral
Lepacyclotes zeilleri; lycopsid stem L II/1: Urochishche Dzhaylyaucho,
R 0602032/H 4435521
Upper Grey Member (T4) Lacustrine, littoral to sublittoral (delta-front)
Mesenteriophyllum kotschnevii L III/4: Sauk Tanga,
R 0608750/H 4432594
Lower Grey Member (T2) Lacustrine, littoral to sublittoral
Mesenteriophyllum kotschnevii L III/5: Sauk Tanga,
R 0608737/H 4432629
Lower Grey Member (T2) Lacustrine, littoral to sublittoral
Appendix A. Locality specification data of the described lycopsids from the Madygen Formation
62 P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
References
Ash, S.R., 1979. Skilliostrobus gen. nov., a new lycopsid cone from the Early Triassic of
Australia. Alcheringa 3, 73–89.
Ash, S.R., Pigg, K.B., 1991. A new Jurassic Isoetites (Isoetales) from the Wallowa terrane
in Hells Canyon, Oregon and Idaho. American Journal of Botany 78, 1636–1642.
Banerji, J., 1988. Some Mesozoic plant remains from Bhuj Formation with remarks on
the depositional environment of beds. The Palaeobotanist 37, 159–168.
Barale, G., 1999. Sur la présence d'une nouvelle espèce d'Isoetites dans la flore du
Crétacé inférieur de la région de Tataouine (Sud tunisien): implications
paléoclimatiques et phylogénétiques. Canadian Journal of Botany 77, 189–196.
Bernard, S., Benzerara, K., Beyssac, O., Brown Jr., G.E., Grauvogel-Stamm, L., Duringer, P.,
2009. Ultrastructural and chemical study of modern and fossil sporoderms by
Scanning Transmission X-ray Microscopy (STXM). Review of Palaeobotany and
Palynology 156, 248–261.
Béthoux, O., Voigt, S., Schneider, J.W., 2010. A Triassic palaeodictyopteran from Kyrgyzstan.
Palaeodiversity 3, 9–13.
Bock, W., 1962. A study on fossil Isoetes. Journal of Paleontology 36, 53–59.
Bock, W., 1969. The AmericanTriassic Flora and Global Distribution,vols. 3–4. Geological
Center, North Wales, Pennsylvania (406 pp.).
Bomfleur, B., Krings, M., Taylor, E.L., Taylor, T.N., 2011. Macrofossil evidence for
pleuromeialean lycophytes from the Triassic of Antarctica. Acta Palaeontologica
Polonica 56, 195–203.
Bose, M.N., Banerji, J., 1984. The fossil floras of Kachchh. I –Mesozoic megafossils. The
Palaeobotanist 33, 1–189.
Bose, M.N., Roy, S.K., 1964. Studies on the upper Gondwana of Kutch –2. Isoetaceae.
The Palaeobotanist 12, 226–228.
Brick, M.I.,1936. The First Finding of the Lower Triassic Flora inMiddle Asia. Transactions
of Geological Institute of the USSR Academy of Sciences 5, 161–174 (in Russian, with
English Abstract).
Brown, R.W., 1939. Some American fossil plants belonging to the Isoetales. Journal of
the Washington Academy of Sciences 29, 261–269.
Brown, R.W., 1958. New occurrences of the fossil quillworts called Isoetites. Journal of
the Washington Academy of Sciences 48, 358–361.
Buchwitz, M., Voigt, S., 2010. Peculiar carapace structure of a Triassic chroniosuchian
implies evolutionary shift in trunk flexibility. Journal of Vertebrate Paleontology
30, 1697–1708.
Budke, J.M., Hickey, R.J., Heafner, K.D., 2005. Analysis of morphological and anatomical
characteristics of Isoetes using Isoetes tennesseensis. Brittonia 57, 167–182.
Cantrill, D.J., Webb, J.A., 1998. Permineralized pleuromeid lycopsid remains from the
Early Triassic Arcadia Formation, Queensland, Australia. Review of Palaeobotany
and Palynology 102, 189–211.
Caplen, C.A., Werth, C.R., 2000. Isozymes of the Isoetes riparia complex, II. Ancestry and
relationships of polyploids. Systematic Botany 25, 260–280.
Chaloner, W.G., Boureau, E., 1967. Lycophyta. In: Boureau, E. (Ed.), Traité de
Paléobotanique, Tome II, Bryophyta, Psilophyta, Lycophyta. Masson et Cie, Paris,
pp. 435–802.
Chaloner, W.G., Turner, S., 1987. An enigmatic Triassic lycopod axis from Australia.
Review of Palaeobotany and Palynology 51, 51–58.
DiMichele, W.A., 1985. Diaphorodendron, gen. nov., a segregate from Lepidodendron
(Pennsylvanian age). Systematic Botany 10, 453–458.
DiMichele, W.A., Skog, J.E., 1992. The Lycopsida: a symposium. Annals of the Missouri
Botanical Garden 79, 447–449.
Dobruskina, I.A., 1974. Triassic lepidophytes. Paleontological Journal 3, 384–397.
Dobruskina, I.A., 1987. Phytogeography of Eurasia during the Early Triassic.
Palaeogeography, Palaeoclimatology, Palaeoecology 58, 75–86.
Dobruskina, I.A., 1994. Triassic Floras of Eurasia. Österreichische Akademie der
Wissenschaften, Wien (422 pp.).
Dobruskina, I.A., 1995. Keuper (Triassic) Flora from Middle Asia (Madygen, South
Fergana). New Mexico Museum of Natural History and Science Bulletin 5, 1–49.
Emmons, E., 1856. Geological report of the Midland counties of North Carolina. George
P. Putnam & Co., New York (352 pp.).
Fliche, P., 1910. Flore fossile du Trias en Lorraine et Franche-Comté avec des considérations
finales par M.R. Zeiller. Bulletin de la Société des Sciences de Nancy III, XI , 1, pp. 222–286.
Fuchs, G., Grauvogel-Stamm, L., Mader, D., 1991. Une remarquable flore à Pleuromeia
et Anomopteris in situ du Buntsandstein Moyen (Trias inférieur) de l'Eifel (R. F.
Allemagne) –morphologie, paléoécologie et paléogéographie. Palaeontographica
Abteilung B 222, 89–120.
Gopal, B., 2009. Biodiversity in wetlands. In: Maltby, E., Barker, T. (Eds.), The wetlands
handbook. Wiley-Blackwell, Oxford, pp. 65–95.
Grauvogel-Stamm, L., 1978. La flore du Grès a Voltzia (Buntsandstein supérieur) des
Vosges du Nord (France). Morphologie, anatomie, interprétations phylogénique
et paléogéographique. Sciences Géologiques Mémoires 50, 1–225.
Grauvogel-Stamm, L., 1993. Pleuromeia sternbergii (Münster) Corda from the Lower
Triassic of Germany –further observations and comparative morphology of its
rooting organ. Review of Palaeobotany and Palynology 77, 185–212.
Grauvogel-Stamm,L., 1999. Pleuromeiasternbergii (Münster) Corda, einecharakteristische
Pflanze des deutschen Buntsandsteins. In: Hauschke, N., Wilde, V. (Eds.), Trias. Eine
ganz andere Welt. Dr. Friedrich Pfeil, München, pp. 271–281.
Grauvogel-Stamm, L., Ash, S.R., 2005. Recovery of the Triassic land flora from the end-
Permian life crisis. Comptes Rendus Palevol 4, 593–608.
Grauvogel-Stamm, L., Duringer, P., 1983. Annalepis zeilleri Fliche, 1910 emend., un
organe reproducteur de Lycophyte de la Lettenkohle de l'Est de la France.
Morphologie, spores in situ et paléoécologie. Geologische Rundschau 72, 23–51.
Grauvogel-Stamm, L., Lugardon, B., 2001. The Triassic lycopsids Pleuromeia andAnnalepis:
relationships, evolution, and origin. American Fern Journal 91, 115–149.
Greb, S.F., DiMichele, W.A., Gastaldo, R.A., 2006. Evolution and Importance of Wetlands
in Earth History. In: Greb, S.F., DiMichele, W.A. (Eds.), Wetlands through timeThe
Geological Society of America, Special Paper 399, 1–40.
Green, W.A., 2010. The function of the aerenchyma in arborescent lycopsids: evidence
of an unfamiliar metabolic strategy. Proceedings of the Royal Society B 277,
2257–2267.
Helby, R., Martin, A.R.H., 1965. Cylostrobus gen. nov., cones of lycopsidean plants from
the Narrabeen Group (Triassic) of New South Wales. Australian Journal of Botany
13, 389–404.
Hermann, E., Hochuli, P.A., Bucher, H., Brühwiler, T., Hautmann, M., Ware, D., Roohi, G.,
2011. Terrestrial ecosystems on North Gondwana following the end-Permian mass
extinction. Gondwana Research 20, 630–637.
Hickey, R.J., 1986a. On the identity of Isoëtes triquetra A. Braun. Taxon 35, 243–246.
Hickey, R.J., 1986b. The early evolutionary and morphological diversity of Isoetes, with
descriptions of two new neotropical species. Systematic Botany 11, 309–321.
Hickey, R.J., 1990. Studies of neotropical Isoëtes L. I.Euphyllum, a new subgenus. Annals
of the Missouri Botanical Garden 77, 239–245.
Hill, R.S., 1987. Tertiary Isoetes from Tasmania. Alcheringa 12, 157–162.
Jonker, F.P., 1976. The Carboniferous “genera”Ulodendron and Halonia –an assessment.
Palaeontographica Abteilung B 157, 97–111.
Karrfalt, E., 1984. Further observations on Nathorstiana (Isoetaceae). American Journal
of Botany 71, 1023–1030.
Karrfalt, E.E., Eggert, D.A., 1977. The comparative morphology and development of
Isoetes L. I. Lobe and furrow development in I.tuckermanii A. Br. Botanical Gazette
138, 236–247.
Keeley, J.E., Osmond, C.B., Raven, J.A., 1984. Stylites, a vascular land plant without
stomata absorbs CO
2
via its roots. Nature 310, 694–695.
Kelber, K.P., 1998. Phytostratigraphische Aspekte der Makrofloren des süddeutschen
Keupers. Documenta Naturae 117, 89–115.
Kelber, K.P., Hansch, W., 1995. Keuperpflanzen. Die Enträtselung einer über 200
Millionen Jahre alten Flora. Museo 11, 1–157.
Kon'no, E., 1973. New species of Pleuromeia and Neocalamites from the upper Scythian
bed in the Kitakami Massif, Japan, with a brief note on some equisetacean plants
from the Upper Permian bed in the Kitakami Massif. Tohoku University Scientific
Reports, 2nd Series. Geology 43, 99–115.
Kott, L.S., Britton, D.M., 1985. Role of morphological characteristics of leaves and the
sporangial region in the taxonomy of Isoetes in northeastern North America.
American Fern Journal 75, 44–55.
Krassilov, V.A., 1981. Changes of Mesozoic vegetation and the extinction of dinosaurs.
Palaeogeography, Palaeoclimatology, Palaeoecology 34, 207–224.
Kryshtofovich, A., 1923.Pleuromeia and Hausmannia in EasternSiberia, with a summaryof
recent contributions to the paleobotany of the region. American Journal of Science 5,
200–208.
Kustatscher, E., Van Konijnenburg-van Cittert, J.H.A., 2005. The Ladinian flora (Middle
Triassic) of the Dolomites: Palaeoenviromental reconstructions and palaeoclimatic
considerations. Géologie Alpine 2, 31–51.
Kustatscher, E., Van Konijnenburg-van Cittert, J.H.A., 2008. Lycophytes and horsetails
from the Triassic flora of Thale (Germany). Neues Jahrbuch für Geologie und
Paläontologie Abhandlungen 250, 65–77.
Kustatscher, E., Wachtler, M., Van Konijnenburg-van Cittert, J.H.A., 2004. A number of
additional and revised taxa from the Ladinian flora of the Dolomites, Northern
Italy. Géologie Alpine 1, 57–69.
Kustatscher, E., Wachtler, M., Van Konijnenburg-van Cittert, J.H.A., 2007. Horsetails and
seed ferns from the Middle Triassic (Anisian) locality Kühwiesenkopf (Monte Prà
della Vacca), Dolomites, Northern Italy. Palaeontology 50, 1277–1298.
Kustatscher, E., Wachtler, M., Van Konijnenburg-van Cittert, J.H.A., 2010. Lycophytes
from the Middle Triassic (Anisian) locality Kühwiesenkopf (Monte Prà della
Vacca) in the Dolomites (northern Italy). Palaeontology 53, 595–626.
Liu, X., Gituru, W.R., Wang, Q.F., 2004. Distribution of basic diploid and polyploid
species of Isoetes in East Asia. Journal of Biogeography 31, 1239–1250.
Liu, H., Wang, Q.F., Taylor, W.C., 2006. Morphological and anatomical variation in
sporophylls of Isoetes sinensis Palmer (Isoetaceae), and endangered quillwort in
China. American Fern Journal 96, 67–74.
Looy, C.V., Brugman, W.A., Dilcher, D.L., Visscher, H., 1999. The delayed resurgence of
equatorial forests after the Permian–Triassic ecologic crisis. Proceedings of the
National Academy of Sciences of the United States of America 96, 13857–13862.
Looy, C.V., Twitchett, R.J., Dilcher, D.L., Van Konijnenburg-van Cittert, J.H.A., Visscher,
H., 2001. Life in the end-Permian dead zone. Proceedings of the National Academy
of Sciences of the United States of America 98, 7879–7883.
Mader, D., 1990. Palaeoecology of the flora in Buntsandstein and Keuper in the Triassic
of Middle Europe, vols. 1–2. Gustav Fischer Verlag, Stuttgart (1582 pp.).
Madsen, T.V., Olesen, B., Bagger, J., 2002. Carbon acquisition and carbon dynamics by
aquatic isoetids. Aquatic Botany 73, 351–371.
Mägdefrau, K., 1932. Über Nathorstiana, eine Isoëtacee aus dem Neokom von
Quedlinburg a. Harz. Beihefte zum Botanischen Centralblatt 49, 706–718.
Masarati, D.L., Thomas, B.A., 1982. The stomata of Isoetes.In:Nautiyal,D.D.(Ed.),
Phyta, Studies on Living and Fossil Plants. Society of Plant Taxonomists, Allahabad,
pp. 155–162.
McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Hawksworth, D.L., Marhold, K.,
Nicolson, D.H., Prado, J., Silva, P.C., Skog, J.E., Wiersema, J.H., Turland, N.J., 2006.
International Code of Botanical Nomenclature (Vienna Code). A.R.G. Gantner
Verlag, Ruggell (568 pp.).
Meng, F., 1994. Discovery of Pleuromeia-Annalepis flora in South China and its signifi-
cance. Chinese Science Bulletin 39, 130–134.
Meng, F., 1996. Middle Triassic lycopsid flora of South China and its palaeoecological
significance. Palaeobotanist 45, 334–343.
63P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
Meng, F., 1998. Studies on Annalepis from Middle Triassic along the Yangtze River and
its bearing on the origin of Isöetes. Acta Botanica Sinica 40, 768–774 (in Chinese,
with English Abstract).
Meng, F., 2000. Advances in the study of Middle Triassic plants of the Yangtze Valley of
China. Acta Palaeontologica Sinica 39, 154–166.
Meyen, S.V., 1987. Fundamentals of Palaeobotany. Chapman and Hall, New York (432
pp.).
Moisan, P., Voigt, S., Pott, C., Buchwitz, M., Schneider, J.W., Kerp, H., 2011. Cycadalean
and bennettitalean foliage from the Triassic Madygen Lagerstätte (SW Kyrgyzstan,
Central Asia). Review of Palaeobotany and Palynology 164, 93–108.
Moisan, P., Voigt, S., Schneider, J.W., Kerp, H., 2012. New fossil bryophytes from the
Triassic Madygen Lagerstätte (SW Kyrgyzstan). Review of Palaeobotany and Palynology
187, 29–37.
Münster, G.G.zu., 1842. Beiträge zur Petrefacten-Kunde. Der Buchner'schen Buchhandlung,
Bayreuth (131 pp.).
Musselman, L.J., Roux, J.P., 2002. Isoetes toximontana (Isoetaceae), a new quillwort with
green megaspores from the Northern Cape of South Africa. Novon 12, 504–507.
Naugolnykh, S.V., 1994. A new lepidophyte from the Kungurian of the Central Urals.
Paleontological Journal 28, 168–175.
Naugolnykh, S.V., 2001. The morphology, systematics and paleoecology of the lycopsid
Viatcheslavia vorcutensis. Paleontological Journal 35, 204–210.
Naugolnykh, S.V., Mogutcheva, N.K., 2006. A new representative of Isoetes (Lycopodiopsida)
from the Lower Triassic of the Tunguska Basin. News of Paleontology and Stratigraphy.
Geologiya i Geofizika 47, 81–93 (supplement to journal, (in Russian, with English
Abstract)).
Naugolnykh, S.V., Zavialova, N.E., 2004. Densoisporites polaznaensis sp. nov.: with com-
ments on its relation to Viatcheslavia vorcutensis Zalessky. The Palaeobotanist 53,
21–33.
Neuburg, M.F., 1936. On the stratigraphy of the coal-bearing deposits of the Kuznetsk
Basin. Bulletin of the Academy of Sciences of the USSR, Geologic Series 4, 469–510.
Niklas, K.J., Tiffney, B.H., Knoll, A.H., 1985. Patterns in vascular land plant diversification: an
analysis at the species level. In: Valentine, J.W.(Ed.),Phanerozoicdiversity patterns:
profiles in macroevolution. Princeton University Press, Princeton, pp. 97–128.
Pant, D.D., Srivastava, G.K., 1962. The genus Isoetes in India. Proceedings of the National
Institute of Sciences of India 28, 242–280.
Pedersen, O., Pulido, C., Rich, S.M., Colmer, T.D., 2011. In situ O
2
dynamics in submerged
Isoetes australis: varied leaf gas permeability influences underwater photosynthesis
and internal O
2
. Journal of Experimental Botany 62, 4691–4700.
Phillips, T.L., Peppers, R.A., DiMichele, W.A., 1985. Stratigraphic and interregional
changes in Pennsylvanian coal-swamp vegetation: environmental inferences.
International Journal of Coal Geology 5, 43–109.
Pigg, K.B., 1992. Evolution of isoetalean lycopsids. Annals of the Missouri Botanical
Garden 79, 589–612.
Pigg, K.B., 2001. Isoetalean lycopsid evolution: from the Devonian to the present.
American Fern Journal 91, 99–114.
Prada, C., Rolleri, C.H., 2003.Caracteres diagnósticos foliares en táxones ibéricos de Isoetes
L. (Isoetaceae, Pteridophyta). Anales Jardín Botánico de Madrid 60, 371–386.
Prada, C., Rolleri, C.H., 2005. A new species of Isoetes (Isoetaceae) from Turkey, with a
study of microphyll intercellular pectic protuberances and their potential taxo-
nomic value. Botanical Journal of the Linnean Society 147, 213–228.
Retallack, G.J., 1975. The life and times of a Triassic lycopod. Alcheringa 1, 3–29.
Retallack, G.J., 1997. Earliest Triassic origin of Isoetes and quillwort evolutionary radia-
tion. Journal of Paleontology 71, 500–521.
Richter, P.B., 1909. Beiträge zur Flora der unteren Kreide Quedlinburgs. II. Die Gattung
Nathorstiana P. Richter and Cylindrites spongioides Goeppert.Wilhelm Engelmann,
Leipzig (12 pp.).
Rolleri, C.H., Prada, C., 2007. Caracteres diagnósticos foliares en Isoetes (Pteridophyta,
Isoetaceae). Annals of the Missouri Botanical Garden 94, 202–235.
Roux, J.P., Hopper, S.D., Smith, R.J., 2009. Isoetes eludens (Isoetaceae), a new endemic
species from the Kamiesberg, Northern Cape, South Africa. Kew Bulletin 64,
123–128.
Sadovnikov, G.N., 1982. The morphology, systematics, and distribution of the genus
Tomiostrobus. Paleontological Journal 1, 100–109.
Sadovnikov, G.N., 2008. On the global stratotype section and point of the Triassic base.
Stratigraphy and Geological Correlation 16, 31–46.
Saporta, G.De, 1894. Flore fossile du Portugal. Nouvelles Contributions á la Flore
Mesozoique. Academie Royale des Sciences, Lisbonne, pp. 1–288.
Schoch, R.R., Voigt, S.,Buchwitz, M., 2010. Achroniosuchid from theTriassic of Kyrgyzstan
and analysis of chroniosuchian relationships.Zoological Journal of the Linnean Society
160, 515–530.
Sharov, A.G., 1970. A specific reptile from the Lower Triassic of Fergana. Paleontologicheskii
Zhurnal 1, 127–131 (in Russian).
Sharov, A.G., 1971. New flying reptiles from the Mesozoic of Kazakhstan and Kyrgyzstan.
Trudy Paleontologiceskogo Instituta AkademijaNauk SSSR 130, 104–113 (inRussian).
Shcherbakov, D.E., 2008a. Madygen, Triassic Lagerstätte number one, before and after
Sharov. Alavesia 2, 113–124.
Shcherbakov, D.E., 2008b. On Permian and Triassic insect faunas in relation to biogeog-
raphy and the Permian–Triassic crisis. Paleontological Journal 42, 15–31.
Sixtel, T.A., 1960. The Stratigraphy of the Continental Sediments of the Upper Permian
and Triassic of Middle Asia. Trudy Tashkent. Universiteta, nov. ser. Geol. nauki.,
176, pp. 1–146 (in Russian).
Sixtel, T.A., 1961. The representatives of the gigantopteris and some accompanying
plants from the Madygen Formation of Fergana. Paleontological Journal 1, 151–158
(in Russian).
Sixtel, T.A., 1962. Flora of the Late Permian and Early Triassic in Southern Fergana.
Stratigrafia i paleontologia Uzbekistana i sopredelnychrayonov. Tashkent 271–414
(in Russian).
Skog, J.E., Hill, C.R., 1992. The Mesozoic herbaceous lycopsids. Annals of the Missouri
Botanical Garden 79, 648–675.
Skog, J.E., Dilcher, D.L., Potter, F.W., 1992. A new species of Isoëtites from the mid-
Cretaceous Dakota Group of Kansas and Nebraska. American Fern Journal 82,
151–161.
Srivastava, G.K., Srivastava, P.C., Shukla, P.K., 2004. Antiquity and diversification of
Isoëtes through the ages. In: Srivastava, P.C. (Ed.), Vistas in Palaeobotany and
Plant Morphology: Evolutionary and E nvironmental Perspectives. U.P. Offset, Lucknow,
pp. 263–279.
Tatarinov, L.P., 2005. A new cynodont (Reptilia, Theriodontia) from the Madygen
Formation (Triassic) of Fergana, Kyrgyzstan. Paleontological Journal 39, 192–198.
Taylor, W.C., Hickey, R.J., 1992. Habitat, evolution, and speciation in Isoetes. Annals of
the Missouri Botanical Garden 79, 613–622.
Taylor, T.N., Taylor, E.L., Krings, M., 2009. Paleobotany. The biology and evolution of
fossil plants.Elsevier/Academic Press, New York (1230 pp.).
Teixeira, C., 1948. Flora Mesozóica Portuguesa. I. Parte. Serviços Geológicos de Portugal,
Lisboa (118 pp.).
Thomas, B.A., 1992. Paleozoic herbaceous lycopsids and the beginnings of extant
Lycopodium sens. lat.and Selaginella sens.lat. Annals of the MissouriBotanical Garden
79, 623–631.
Thomas, B.A., Brack-Hanes, S.D., 1984. A new approach to family groupings in the
lycophytes. Taxon 33, 247–255.
Vakhrameev, V.A., Dobruskina, I.A., Meyen, S.V., Zaklinskaja, E.D., 1978. Paläozoische
und mesozoische Floren Eurasiens und die Phytogeographie dieser Zeit. VEB
Gustav Fischer Verlag, Jena (300 pp.).
Van Waveren, I.M., Iskandar, E.A.P., Booi, M., Van Konijnenburg-van Cittert, J.H.A., 2007.
Composition and palaeogeographic position of the Early Permian Jambi flora from
Sumatra. Scripta Geologica 135, 1–28.
Voigt, S., Hope, D., 2010. Mass occurrence of penetrative trace fossils in Triassic lake
deposits (Kyrgyzstan, Central Asia). Ichnos 17, 1–11.
Voigt, S., Haubold, H., Meng, S., Krause, D., Buchantschenko, J., Ruckwied, K., Götz, A.E.,
2006. Die Fossil-Lagerstätte Madygen: Ein Beitrag zur Geologie und Paläontologie
der Madygen-Formation (Mittel- bis Ober- Trias, SW- Kirgisistan, Zentralasien).
Hallesches Jahrbuch für Geowissenschaften 22, 85–119.
Wang, Z., 1991. Advances on the Permo-Triassic lycopods in North China. I. An Isoetes
from the mid-Triassic in Shaanxi Province. Palaeontographica Abteilung B 222,
1–30.
Wang, Z., Wang, L., 1982. A new species of the lycopsid Pleuromeia from the Early
Triassic of Shanxi, China, and its ecology. Palaeontology 25, 215–225.
Yu, J.X., Broutin, J., Huang, Q., Grauvogel-Stamm, L., 2010. Annalepis, a pioneering
lycopsid genus in the recovery of the Triassic land flora in South China. Comptes
Rendus Palevol 9, 479–486.
64 P. Moisan, S. Voigt / Review of Palaeobotany and Palynology 192 (2013) 42–64
