American Journal of Botany 96(1): 237–251. 2009.

SEED FERNS FROM THE LATE PALEOZOIC AND MESOZOIC:

ANY ANGIOSPERM ANCESTORS LURKING THERE?1
Edith L. Taylor2 and Thomas N. Taylor
Department of Ecology and Evolutionary Biology, and Natural History Museum and Biodiversity Research Center, University of
Kansas, Lawrence, Kansas 66045-7534 USA

Five orders of late Paleozoic–Mesozoic seed ferns have, at one time or another, figured in discussions on the origin of angio-
sperms, even before the application of phylogenetic systematics. These are the Glossopteridales, Peltaspermales, Corystosper-
males, Caytoniales, and Petriellales. Although vegetative features have been used to suggest homologies, most discussion has
focused on ovulate structures, which are generally interpreted as megasporophylls bearing seeds, with the seeds partially to almost
completely enclosed by the megasporophyll (or cupule). Here we discuss current information about the reproductive parts of these
plants. Since most specimens are impression-compression remains, homologizing the ovulate organs, deriving angiospermous
homologues, and defining synapomorphies remain somewhat speculative. Although new specimens have increased the known
diversity in these groups, a reconstruction of an entire plant is available only for the corystosperms, and thus hypotheses about
phylogenetic position are of limited value. We conclude that, in the case of these seed plants, phylogenetic analysis techniques
have surpassed the hard data needed to formulate meaningful phylogenetic hypotheses. Speculation on angiosperm origins and
transitional stages in these fossils provides for interesting discussion, but currently it is still speculation, as the role of these groups
in the origin of angiospermy continues to be cloaked in Darwin’s mystery.

Key words: angiosperm ancestors; caytonialeans; corystosperms; fossil plants; glossopterids; Mesozoic; peltasperms; Perm-
ian; seed plant phylogeny.

The Permian glossopterids, along with the three major groups
of late Paleozoic–Mesozoic seed ferns (peltasperms, corys-
tosperms, caytonialeans), all represent gymnosperms that have
historically figured prominently in discussions on the origin of
angiosperms (e.g., Krassilov, 1977b). The Glossopteridales are
considered to represent a monophyletic group, and all members
have similar foliage, i.e., Glossopteris Brongniart and Ganga-
mopteris McCoy, although more recent work suggests some
diversity in reproductive organs (e.g., Taylor et al., 2007;
Prevec et al., 2008). They represent the dominant elements in
Permian floras from the supercontinent of Gondwana (includ-
ing Antarctica, Australia-New Zealand, parts of Africa and
South America, and peninsular India) and appear to have gone
extinct around the Permo–Triassic boundary (McManus et al.,
2002). The other, post-Carboniferous seed ferns, however, were
widespread from the Permian to the Cretaceous, and unlike the
Carboniferous forms, appear to have few morphological char-
acters in common. Rather, they are included as a collection of
either orders or families in which the relationships among the
various taxa are uncertain (Taylor et al., 2006). In most seed
plant phylogenies, each of these groups is included as a single
terminal branch, often a composite terminal. Although a few of
these seed plants are known from anatomically preserved speci-
mens, most are represented by impression and compression fos-
sils and therefore are not understood with the same degree of
1 Manuscript received 19 June 2008; revision accepted 27 October 2008.
The authors thank the following colleagues for kindly supplying figures:
J. M. Anderson, C. P. Daghlian, H. Kerp, S. McLoughlin, H. Nishida, P.
Ryberg, and M. Zavada, as well as students and postdoctoral scholars in
our laboratory who contributed to this work. This research was partially
supported by the National Science Foundation (OPP-0229877, ANT-
0635477).
2 E-mail: etaylor@ku.edu; tntaylor@ku.edu
doi:10.3732/ajb.0800202
237
resolution as the late Paleozoic pteridosperms, many of which
are understood, at least in part, in anatomical detail. Neverthe-
less, some of these “younger” seed ferns possess morphological
features that have been used to suggest affinities with some of
the Paleozoic seed ferns (Petriella, 1981). As research has con-
tinued, our knowledge of the diversity within some of these
groups has increased, which often makes it difficult to define
the group according to its original concept. Nevertheless, this
additional diversity needs to be accommodated in seed plant
phylogenies.
Historically, and for obvious reasons, many of these gymno-
sperms have been the subject of considerable interest as poten-
tial angiosperm progenitors. These assumptions have been
based on a number of features, most notably the enclosure of
the ovules in some type of leaflike structure that has generally
been termed a cupule. In the sections that follow, we provide a
discussion of several seed fern groups and offer our perspective
on the relationships of these fossil groups and possible homolo-
gies with the angiosperms.
The general concept of the seed ferns was proposed by Oli-
ver and Scott (1904) for Paleozoic (Carboniferous) plants that
possessed fernlike foliage, but with seeds and pollen sacs, rather
than sporangia, attached to the leaves. The initial concept of the
Pteridospermae was based on structurally preserved organs of
the Carboniferous lyginopterids, but was subsequently ex-
panded to other Carboniferous–Permian orders, and today also
includes the late Paleozoic Glossopteridales. In Mesozoic rocks,
four groups have been delimited. One of these, the Corystosper-
males (Upper Permian–Cretaceous), is known in some detail,
based on permineralized and compression fossils from several
continents. The Peltaspermales (Pennsylvanian–Triassic) and
the Caytoniales (Triassic–Cretaceous) are known only from im-
pression and compression specimens; both structurally pre-
served and compressed fossils have been assigned to the
Petriellales (Triassic). Although foliage and other vegetative parts
of these seed plants are known, we have limited the discussion

238 American Journal of Botany
that follows to the reproductive parts of these plants because
they have served as the focal point in discussions linking them
to the flowering plants.
GLOSSOPTERIDALES
There is a considerable body of literature on the Glossop-
teridales, beginning with the description of the vegetative leaf
Glossopteris by Adolphe Brongniart in 1828. Today, the group
is known from stem, root, and leaf genera (Fig. 1), that are pre-
served in a variety of modes. The reproductive organs of the
glossopterids, however, have remained a continuing source of
controversy since the first report of “linear sori” on some
Glossopteris leaves (Feistmantel, 1886).
Ovulate organs— Today there are more than 50 different
types of ovulate reproductive structures that have been de-
scribed (McLoughlin, 1990), with additional ones continually
being reported or reexamined (e.g., Adendorff, 2005; Prevec et
al., 2008). A few reproductive organs have been found attached
to the parent plant in association with vegetative leaves (Fig. 2)
(e.g., Pant and Singh, 1974). Other reproductive organs are at-
tributed to the group based on venation patterns of the me-
gasporophyll that are similar to those of the vegetative leaves
and on co-occurrence at the same locality. One of the major
obstacles in the interpretation of glossopterid reproductive or-
gans has been the fact that most are preserved as impression-
compression (e.g., Fig. 3) or mold-cast specimens and thus,
interpreting the homology of these organs has been controver-
sial. In attempting to document diversity, some authors have
used an artificial system of morphofamilies based on macro-
morphological features (e.g., Arberiaceae, Dictyopteridiaceae,
Lidgettoniaceae, Rigbyaceae) to classify and categorize these
structures (see Adendorff, 2005, for a history of glossopterid
classification). Others have used different names for higher-
level taxonomy in the mistaken belief that families of fossil
plants should be named after an ovulate organ rather than the
oldest validly published genus in the group.
Plumstead (1956) initially described the ovulate fructifica-
tions of the glossopterids and thus demonstrated that they were
gymnosperms and not ferns. She erected the genus Hirsutum
Plumstead for shield-shaped, “bivalvate” structures that she in-
terpreted as bisexual (Plumstead, 1956, 1958). Plumstead sug-
gested that the fructifications produced hairlike, pollen-producing
organs on one of the two valves. She also interpreted flat, bract-
like structures in Scutum Plumstead (Fig. 4) as pollen bearing
(for details, see Prevec et al., 2008). Thus far, no glossopterid
reproductive organs have been conclusively demonstrated as
containing both pollen and seeds (Pant, 1987). Despite the nu-
merous ovulate structures attributed to the glossopterids, many
represent variations on a basic theme. These consist of a me-
gasporophyll bearing seeds; the megasporophyll has been vari-
ously termed a capitulum, cupule, fertiliger, or cladode. In
instances where the reproductive structure is attached to the
parent plant, it appears in an axillary position (e.g., Holmes,
1987) or fused or adpressed to the petiole or lamina of a
Glossopteris leaf.
Perhaps the most important discovery contributing to under-
standing the homology of the glossopterid ovulate reproductive
organs is the report of Gould and Delevoryas (1977) of ana-
tomically preserved Glossopteris megasporophylls with at-
tached ovules (Fig. 5). Although the megasporophylls were not
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[Vol. 96
found organically attached to axes or leaves, anatomical simi-
larities between of the megasporophylls and vegetative leaves
of Glossopteris suggest that they were borne on the same plant
(see Nishida et al., 2007). Attached to one surface of the fleshy
megasporophyll are numerous ovoid seeds with the edges of the
megasporophyll slightly enrolled and partially covering the
seeds. Now named Homevaleia gouldii Nishida et al. (2007),
the megasporophyll lacks the sclerenchymatous hypodermal fi-
bers found in the vegetative leaf of G. homevalensis. Each or-
thotropous ovule is 1.2–1.3 mm long by 0.8–0.9 mm wide and
attached by a small stalk to the adaxial surface of the megasporo-
phyll, based on the anatomy. The integument is thickened in the
micropylar region, and the outer layer (sarcotesta) extends out
from the ovules to form a meshwork of filaments between the
seeds. Gould and Delevoryas (1977) suggested that this mesh-
work may have been involved in some way in directing pollen
flow, possibly aided by a pollination droplet. Another sugges-
tion is that perhaps the spongy meshwork served to insulate the
ovules in the cool, temperate environment in which this plant
lived (Nishida et al., 2007). Many of the ovules contain well-
preserved megagametophytes, each with a single archegonium
(Fig. 6). Bisaccate pollen of the Protohaploxypinus Samoilo-
vitch type (Fig. 7) has been reported in the pollen chambers of
these ovules (Gould and Delevoryas, 1977; Nishida et al., 2004).
Especially significant is the report of pollen tubes in various
stages of releasing flagellated sperm in the region of the arche-
gonium (Nishida et al., 2003). Sperm are top shaped (Fig. 8),
small (~12 µm in diameter), and characterized by spiral bands
of dark dots at one end that are interpreted as corresponding to
the positions of the basal bodies of numerous flagella aligned
along a multilayered structure (MLS). Interestingly, the glossop-
terid sperm are morphologically like those of cycads and
Ginkgo, but much smaller (Nishida et al., 2007). In general or-
ganization, the Homevaleia megasporophyll shows some simi-
larities to the impression-compression ovulate structures
Dictyopteridium Feistmantel ex Zeiller or Plumsteadia Rigby
(Fig. 9), in part based on the arrangement of ovules in the per-
mineralized and compressed forms (Nishida et al., 2007).
Anatomically preserved glossopterid megasporophylls with
attached ovules are also known from permineralized peat in the
Late Permian Buckley Formation of Antarctica (Taylor and
Taylor, 1992; Taylor et al., 2007). One of these is ~6.0 mm
wide and 1.0 mm thick and is thought to have been produced by
the same plant that bore Glossopteris schopfii Pigg (1990a)
leaves, based on similar anatomy (Taylor and Taylor, 1992).
The seeds are attached to the adaxial surface of the megasporo-
phyll (Fig. 10) (Taylor and Taylor, 1992). This character is
based on the position of the tissues that make up the vascular
bundles and contrasts with the suggested abaxial attachment of
ovules described from impression fossils (Adendorff, 2005).
Another permineralized specimen from the Late Permian of
Antarctica shows some similarity to several of the so-called cu-
pulate glossopterid reproductive structures (Surange and Chan-
dra, 1975), such as Arberia White or Rigbya Lacey et al. (Fig.
4). A preliminary analysis suggests that it is a branching struc-
ture bearing at least four uniovulate cupules. Each cupule is
~3.0 mm long and contains a sessile ovule with two flattened
wings extending from the integument. The stalk at the base of
the organ contains a C-shaped vascular strand like that pro-
duced in some leaves (Taylor et al., 2007).
Dispersed ovules that show details of embryo development
have been described from the Antarctic permineralized peat.
Although they occur in close association with Glossopteris

January 2009] Taylor and Taylor—Seed ferns from the late Paleozoic–Mesozoic
leaves, none have been found attached. They are considered to
represent glossopterid seeds based on the overwhelming domi-
nance of this group in the peat and in nearby floras of compa-
rable age (e.g., Cúneo et al., 1993). Seeds of Plectilospermum
Taylor and Taylor are ~5.0 mm long and platyspermic with the
nucellus and integument attached only at the base (Taylor and
Taylor, 1987). Many seeds have a cellular megagametophyte
with two archegonia, each showing evidence of a multicellular
embryo, confirming the existence of polyembryony (Fig. 11).
Development had progressed in some embryos so that the pres-
ence of a suspensor could be distinguished (Smoot and Taylor,
1986).
As noted, one of the problems in discussing the ovulate struc-
tures in the glossopterids is the use of nonstandard terms to de-
scribe their morphology. Schopf (1976) suggested the term
fertiliger for the megasporophyll, a designation that Pant (1987)
also used. Schopf (1976) proposed that the ovulate complex in
the glossopterids represented an axillary shoot and suggested a
distant relationship with the cordaites, an idea that was termed
the cladode concept by Retallack and Dilcher (1981). Perminer-
alized reproductive structures that have been described to date,
however, clearly indicate that the ovules are borne on the adax-
ial surface of a megasporophyll, although reconstructions based
on impression fossils suggest attachment was abaxial. What re-
mains to be determined is whether the megasporophyll is at-
tached to a reduced and modified axillary branch system or to
the base of the subtending vegetative or scale leaf (Pigg and
Trivett, 1994). It is also possible that the structures that make up
the glossopterid ovulate complex do not involve the same ho-
mologies in every case. This hypothesis is further strengthened
by the variety of morphologic forms, including those that have
multiple ovules (Fig. 12) and those with apparently solitary
ovules. Seed morphology is also variable, with both radially
and bilaterally symmetrical forms known. The presence or ab-
sence of wings on some ovules may be related to dispersal syn-
dromes. Moreover, some glossopterid ovules have only a single
archegonium, while in others, two are present. The situation
may in part parallel the extraordinary variation demonstrated
among the seeds of Paleozoic seed ferns, which were produced
on leaves, in cupules, and on naked branching systems. Thus,
the glossopterids, despite the evident uniformity of their vege-
tative organs, apparently represent a rather diverse group of late
Paleozoic seed ferns that may ultimately rival the Carbonifer-
ous taxa in the variability of their ovulate reproductive struc-
tures. Because the group dominated Gondwana for more than
30 Myr, perhaps it is not surprising that the glossopterids are
more diverse than their vegetative morphology would suggest.
While at least some of the glossopterid ovulate structures are
homologous with a megasporophyll, with ovules attached to the
adaxial surface, we echo the comments of others who have ex-
amined and discussed the glossopterid reproductive structures
in noting that additional permineralized specimens will be criti-
cal in understanding the homologies of these unusual reproduc-
tive organs.
One interesting feature is the apparent absence of any wing-
like structure on the megasporophyll in the permineralized
specimens, a structure that is commonly found in impression-
compression forms (Fig. 4). It is doubtful that the enrolled
edges of the megasporophyll of Homevaleia gouldii could cre-
ate such a structure if compressed. The absence of the wing
may reflect a developmental stage of the megasporophyll or a
difference in taxa between South Africa and Australia. The
presence of this structure on many of the impression specimens
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239
suggests that there is some function for the wing. Protection of
the developing ovules may be one obvious adaptation. Another
might involve dispersal, with the entire megasporophyll being
shed as a unit (Adendorff et al., 2002), although this is unlikely
because most impression-compression specimens appear to
have already shed their seeds. The presence of some dispersed
seeds with integumentary wings is also not consistent with the
idea of the entire unit being shed.
Pollen organs— Unlike the ovulate organs, there is little
variability in the pollen organs of the Glossopteridales. The two
most common genera are Glossotheca Surange and Mahesh-
wari (1970) and Eretmonia Du Toit (1932), although it has been
suggested that they actually represent a single taxon (Pant,
1987). In both, clusters of pollen sacs are borne on two stalks,
which are attached to a subtending leaf. In Eretmonia, which is
more completely known, the pollen sacs are elongate (~1.0 mm
long), and the subtending leaf is small and rhombohedral. Arbe-
riella Pant et Nautiyal is used for isolated pollen sacs similar to
those of Glossotheca Surange and Maheshwari. Pollen of the
glossopterids is bisaccate and striate (taeniate). It has generally
been compared to the sporae dispersae genus Protohaploxypi-
nus, although Lindström et al. (1997) found four different mor-
phogenera within a single pollen sac of Arberiella.
CAYTONIALES
The Caytoniales are a small group of Triassic–Cretaceous
seed ferns (Krassilov, 1977a) that was erected in 1925 by the
British paleobotanist H. H. Thomas, based on compression
specimens from the Middle Jurassic plant-bearing beds along
the coast of Cayton Bay in Yorkshire, Great Britain. These
plants possessed such interesting features that Thomas regarded
them as a new group of angiosperms in his initial description
(Thomas, 1925). Since that time, the seed-bearing structure
Caytonia Thomas has captured the imagination of paleobota-
nists and has been variously used as a link to the angiosperm
carpel (e.g., Gaussen, 1946; Doyle, 1978, 1996, 2006). As a
result, the Caytoniales have figured prominently in a number of
phylogenetic analyses, despite the fact that they continue to be
one of the most poorly known groups of Mesozoic seed ferns.
Ovulate organs— Caytonia consists of an axis ~5.0 cm long
bearing stalked, multiovulate cupules in subopposite pairs (Fig.
13) (Thomas, 1925; Harris, 1964). This structure, which has
been interpreted as a megasporophyll, bears scars along the sur-
face, suggesting that the cupules were shed at maturity. Each
cupule is compressed to be nearly circular in outline (Fig. 14)
and recurved with a liplike projection directed toward the point
of attachment (Harris, 1940). A single cupule contains from 8–
30 seeds, depending on the species, and each seed is attached by
a delicate stalk in an orthotropous position (Fig. 15). Seeds are
~2.0 mm long and radially symmetrical with an integument
composed of an outer, uniseriate epidermis that covers a row of
radially aligned, thick-walled cells (e.g., Harris, 1958). Harris
(1951) suggested that perhaps the outer portion of the integu-
ment was fleshy.
In the initial description of Caytonia, pollen grains were
found on the liplike portion of the cupule (Fig. 15), a structure
that Thomas (1925) termed the stigmatic flap or region. This was
no doubt the principal reason that the cupule was described as
an angiospermous fruit containing numerous seeds. According
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240 American Journal of Botany [Vol. 96

Figs. 1–12. Glossopterid vegetative and reproductive organs. 1. Glossopteris leaf, Permian of Antarctica, showing anastomosing venation (courtesy
H. Kerp). Bar = 2.5 cm. 2. Suggested reconstruction of Ottokaria zeilleri Pant and Nautiyal showing relationship between vegetative leaf and megasporo-
phyll (redrawn from Pant and Nautiyal, 1984). 3. Rigbya arberioides Lacey et al. showing lobes and cupules (courtesy P. Ryberg). Bar = 5.0 mm. 4. Scutum
rubidgeum Plumstead (courtesy S. McLoughlin). Bar = 5.0 mm. 5. Megasporophyll of H. gouldii Nishida et al. showing several ovules (courtesy P. Ry-
berg). Bar = 1.0 mm. 6. Seed of Homevaleia gouldii with archegonium (arrow) in megagametophyte tissue (courtesy P. Ryberg). Bar = 0.25 mm. 7. Bisac-
cate pollen grain. Bar = 25 µm. 8. Sperm from H. gouldii ovule (courtesy H. Nishida). Bar = 100 µm. 9. Plumsteadia semnes Rigby showing numerous

January 2009] Taylor and Taylor—Seed ferns from the late Paleozoic–Mesozoic
to this interpretation, fertilization would have taken place via a
pollen tube that grew from the pollen grain on the stigmatic
surface to the pollen chamber of an ovule. Fine strands of cuti-
cle that extended from the cupule to the seeds were interpreted
as the remnants of pollen tubes or as extensions of the seed
micropyles. Later research, however, revealed that pollen grains
were actually present inside the cupule, perhaps drawn in by
pollination droplets that originated at the micropylar end of
each seed. The next step in elucidating the reproductive biology
of Caytonia was provided by Maria Reymanówna (1973). Her
detailed work indicated that each seed was associated with an
elongate canal extending from the seed micropyle to the outer
lip of the cupule and that the number of these canals was equal
to the number of seeds produced in each cupule. Today, Cayto-
nia is interpreted as a multiovulate cupule and not an angiosper-
mous fruit as originally thought.
Caytonia demonstrates at least one method in which seeds
became enclosed and in this feature parallels the morphology,
although not the function, of an angiosperm carpel (Krassilov,
1977a). Doyle (1978, 2006) has supported the idea, proposed
earlier by Gaussen (1946), that the Caytoniales represent some
grade of angiosperm ancestors. According to this theory, there
is a reduction in the number of ovules in Caytonia to one per
cupule. The cupule wall would be homologous with the second
integument in an angiosperm ovule and the cupule-bearing axis
would develop to grow over and enclose the cupule and is thus
homologous with the carpel. This idea requires that the main
Caytonia axis develop into a flattened, dorsiventral structure,
and cuticular differences on the upper and lower sides of the
axis have been used to support this scenario. Because all cayto-
nialean fossils represent flattened compressions and no internal
anatomy is known, however, it is difficult to extrapolate the
three-dimensional morphology of the reproductive structures.
In addition, in some specimens of Caytonia, the ovules are
borne on long stalks (e.g., Fig. 13), a configuration which would
presumably make it difficult for overgrowth of the stalk.
Pollen organs— Caytonanthus Harris (originally Antholi-
thus) is a pollen-bearing structure that was also found in the
Middle Jurassic Yorkshire flora in association with Caytonia
cupules and foliage of Sagenopteris Presl in Sternberg. Thomas
included it in the Caytoniales on the basis of the pollen, which
was identical to that found in Caytonia. Caytonanthus consists
of a slender axis bearing flattened, pinnate lateral branches (Fig.
16); each branch bears from 1–3 elongate synangia (Osborn,
1994). Synangia are about 1.0 cm long, pointed at the distal end
(Fig. 17), and contain 1–4 pollen sacs (originally termed loc-
ules) that are arranged around a central zone of tissue (Harris,
1941). These structures (which have been termed anthers in
some treatments) are circular in cross section, with dehiscence
taking place toward the center of each synangium. The epider-
mis is composed of delicate fusiform cells, perhaps with thick-
er-walled fibrous cells beneath.
Pollen grains of Caytonanthus are small and bisaccate, and if
found dispersed would be included in Vitreisporites Leschik.
Grains are 25–40 µm in diameter and contain endoreticulations
lining the interior of the sacci, thus making these grains eusac-
←
seeds (courtesy P. Ryberg). Bar = 1.7 cm. 10. Suggested reconstruction of a Glossopteris megasporophyll with seeds attached to adaxial surface (from
Taylor and Taylor, 1992). 11. Cross section of Plectilospermum seed showing two archegonial chambers (arrows) containing embryos. Bar = 850 µm. 12.
Diagrammatic reconstruction of Denkania indica (redrawn from Surange and Chandra, 1975).
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241
cate (Zavada and Crepet, 1986; Osborn, 1994). Another inter-
pretation is that the grains are protosaccate and the
endoreticulations are continuous between the saccus wall and
surface of the corpus (Pedersen and Friis, 1986). The ultrastruc-
ture of the exine suggests that the sexine is alveolate, and there
is a conspicuous sulcus on the distal surface (Zavada and Crepet,
1986).
CORYSTOSPERMALES
The Corystospermales have a worldwide distribution based
on foliage and, in many cases, reproductive structures. In Gond-
wana, foliage of Dicroidium Gothan is often the dominant ele-
ment in Middle–Late Triassic floras, and the genus is known
from localities in Antarctica, South Africa, Australia, Argen-
tina, Tasmania, and India. In Laurasia, foliage of Pachypteris
Brongniart (Harris, 1964) and Thinnfeldia von Ettingshausen is
assigned to the corystosperms. Umkomasia Thomas and Pteru-
chus Thomas are known from the Jurassic of Europe associated
with Thinnfeldia foliage (Kirchner and Müller, 1992). Recently,
three species of Dicroidium leaves have been described from
the Upper Permian of Jordan (Abu Hamad et al., 2008), sug-
gesting that the group may have originated in the late Paleozoic
and spread into Gondwana through Africa. The family was es-
tablished by Thomas in 1933 to include Dicroidium foliage,
Umkomasia (and other) ovulate organs, and Pteruchus pollen
organs, with the familial name based on the characteristic hel-
met shape of the uniovulate cupules. Based on the morphologi-
cal diversity within the corystosperms, Archangelsky (1996)
suggests that they were a diverse and rapidly evolving group
during the Triassic. The plants were probably small to large
woody shrubs and trees and bore pinnate leaves with open di-
chotomous venation. Although the various plant parts have not
been found attached, their consistent occurrence in the same
beds, similarity in epidermal anatomy, and identical pollen
found in both pollen organs and seeds have been used to indi-
rectly associate the various organs (Petriella, 1983). When per-
mineralized, the plants have been reconstructed based on the
occurrence of unique secretory cells in the various organs (Tay-
lor, 1996).
Ovulate organs— One of the interesting aspects of the corys-
tosperms is that the reproductive organs are morphologically
consistent within the group. The most common seed-containing
structure is Umkomasia (Fig. 18) (Thomas 1933), which is
known widely from Gondwanan deposits (Taylor, 1996), but is
also known from Europe (Kirchner and Müller, 1992) and
China (Figs. 19, 20) (Zan et al., 2008). It consists of a fertile
branch, more than 15 cm long in some taxa, e.g., U. quadripar-
tita Anderson and Anderson (2003), which produces laterals.
Each lateral bears one to several pairs of recurved, helmet-like,
generally uniovulate cupules. Some specimens have a pair of
subtending bracts at the base of the branch (Fig. 21); in others,
they are not present and may have been shed (Holmes, 1987).
The branching in Umkomasia was originally interpreted to be
flattened in a single plane (Thomas, 1933), but permineralized
15372197, 2009, 1, Downloaded from https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.0800202 by CAPES, Wiley Online Library on [11/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
[Vol. 96
American Journal of Botany
242
 15372197, 2009, 1, Downloaded from https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.0800202 by CAPES, Wiley Online Library on [11/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
January 2009] Taylor and Taylor—Seed ferns from the late Paleozoic–Mesozoic 243

specimens from the Middle Triassic of Antarctica indicate
that the cupulate branch has stem-like anatomy and produces
paired traces to the laterals. The cupule-bearing branches of U.
resinosa Klavins et al. (2002) are helically arranged and cu-
pules contain either one or two ovules attached to the abaxial
surface (Fig. 22) of the cupule wall. Ovules of this species are
small, possess a single integument, and are characterized by a
bifid micropylar extension at the distal end.
Cupules preserved as compressions and impressions often
appear urn-like in morphology with two prominent lobes (Fig.
19). The surface of the cupule is wrinkled, suggesting that it
may have been fleshy (Thomas, 1933). On well-preserved spec-
imens, stomata are present on both surfaces of the cupule. The
seeds of U. macleanii are small (5.0 mm long) and borne either
singly or in pairs in each cupule (Thomas, 1933). Compressed
cupules have also been found attached to short shoots, which in
turn arise from longer branches bearing D. odontopteroides
leaves (Fig. 23) (Axsmith et al., 2000). Cupules of U. uniramia
Axsmith et al. (2000) are borne in whorls of five to eight. The
cupules attached to short shoots (Fig. 24) and leaves attached to
long shoots from the Upper Triassic of Antarctica has perplexed
some who challenge whether long shoots would retain leaves
(e.g., Artabe and Brea, 2003; Holmes and Anderson, 2005). In
modern Ginkgo L., however, leaves are borne on relatively
large stems as well as on short shoots (Axsmith et al., 2007). It
is also important to note that this Dicroidium plant grew at high
polar latitudes, and thus the parameters affecting growth were
no doubt much different than they are in temperate regions to-
day (Axsmith et al., 2007). The report of Umkomasia cupules
(Figs. 19, 20) in association with Thinnfeldia-type foliage from
the Upper Triassic of China (Zan et al., 2008) further expands
the diversity within the corystosperms. Umkomasia has also re-
cently been described from the Upper Permian of India (Chan-
dra et al., 2008), providing some support for the idea that the
corystosperms arose in the late Paleozoic (Kerp et al., 2006;
Abu Hamad et al., 2008).
Several additional genera of cupulate structures are often in-
cluded in the corystosperms. These include Spermatocodon
Thomas (1933), Pilophorosperma Thomas (1933) (Fig. 25),
and Karibacarpon Lacey (1976). Judging from the number of
species described, Pilophorosperma is perhaps the most com-
mon seed-producing member of this group (Thomas, 1933).
Some cupules are less hemispherical in outline than are those in
Umkomasia. Specimens of P. granulatum have three cupules at
the end of each branch, while two are common in P. gracile. On
the outer surface of the cupule are short papillae, and the inner
surface is covered with long, pointed hairs. In P. crassum the
cupules are borne in an overlapping fashion along the axis so
that the fertile branch appears compact. Holmes (1987) places
Pilophorosperma and Karibacarpon in the genus Umkomasia.
On the basis of consistent co-occurrence in the Middle Triassic
of eastern Australia, he suggests that U. feistmantelii was the
cupulate organ of the plant that produced Dicroidium zuberi
foliage.
Some seeds of the corystosperms possessed a slightly curved,
bifid micropylar canal, which often extended beyond the cupule
margin (Fig. 25) (Thomas, 1933). Macerations suggest that the
integument contained no fibrous cells and that the seed epider-
mis was covered with a thick cuticle. Embedded in the pollen
chambers are numerous saccate pollen grains similar to those
found in the pollen-producing organ Pteruchus (Townrow,
1962).
Pollen organs— The most common pollen organ attributed
to the Corystospermales is Pteruchus (Thomas, 1933) (Fig. 26).
The genus is known throughout Gondwana (Townrow, 1962;
Taylor, 1996), but there are also scattered reports from Laur-
asia, e.g., P. septentrionalis from the Rhaeto–Liassic (Late
Triassic–Early Jurassic) of Germany (Kirchner and Müller,
1992). Most specimens have alternately arranged microsporo-
phylls attached to an axis about 4.0 cm long (Thomas, 1933;
Townrow, 1962), although in P. indicus the microsporophylls
appear to be helically arranged (Pant and Basu, 1973). Each
microsporophyll terminates in a flattened head (Fig. 27) that
bears numerous, elongate pollen sacs on the abaxial surface;
these are partially protected by the tissue of the head. The num-
ber of sacs per head is variable (20–200), and dehiscence is
longitudinal. The cuticle on all parts shows epidermal cells with
slightly undulate margins and few stomata (Townrow, 1962).
Permineralized specimens of Pteruchus are known from peat
deposits from the lower Middle Triassic of Antarctica (Yao et
al., 1995). Pteruchus fremouwensis Yao et al. (1995) consists of
a branch bearing helically arranged microsporophylls (Fig. 28).
Each microsporophyll has a flattened head bearing ~20 elon-
gate, sessile, unilocular pollen sacs. Pollen sacs are ≤2.0 mm
long and possess walls that contain secretory cells (Fig. 29) like
those in the leaves of D. fremouwensis (Pigg, 1990b) from the
same peat deposit (Pigg and Taylor, 1990). If found dispersed,
the pollen of Pteruchus would be assigned to Alisporites Daugh-
erty, Pteruchipollenites Couper, or Falcisporites Leschik. Pol-
len is bisaccate, with the sacci slightly inclined and partially
covering the distal sulcus. Pollen of P. dubius ranges from 80–
115 µm in the primary plane (Taylor et al., 1984). The sacci
have endoreticulations on their inner surfaces, but the outer sur-
faces are smooth (Zavada and Crepet, 1985).
Another pollen organ believed to belong to the corystosperms
is Pteroma Harris. Pteroma is known to date only from the
Middle Jurassic of Yorkshire, where it is associated with Pac-
hypteris foliage (Harris, 1964). In Pteroma thomasii, the mi-
crosporophyll is flattened and bears two rows of pollen sacs
from what is interpreted as the lower surface. Each head is
shield-shaped, with the pollen sacs partially embedded in the
tissue. Pollen is bisaccate and up to 107 µm long. It is not known
whether these grains are protosaccate or eusaccate like those of
Pteruchus (Osborn and Taylor, 1993). Other putative corys-
tosperm pollen organs include Nidiostrobus from India (Bose
and Srivastava, 1973) and Kachchhia from India and Antarctica
(Bose and Banerji, 1984; Gee, 1989).

←
Figs. 13–22. Reproductive organs of Caytoniales and Corystospermales. 13. Caytonia cupules attached to axis. Bar = 5.0 mm. 14. Cupule of Caytonia
sewardii Thomas emend. Harris. Bar = 2.5 mm. 15. Suggested reconstruction of Caytonia cupule showing attachment of seeds and “stigmatic lip” (arrow).
16. Reconstruction of Caytonanthus arberi (Thomas) Harris showing synangia attached to pinnate branches (from Crane, 1985). 17. Branch bearing several
Caytonanthus synangia. Bar = 3.0 mm. 18. Several Umkomasia cupules (C) attached to branch. Bar = 1.0 mm. 19. Suggested reconstruction of U. asiatica
Zan et al. (courtesy B. Axsmith). 20. Axis of U. asiatica bearing cupules at multiple nodes. Bar 2.0 mm. (courtesy B. Axsmith) 21. Diagrammatic recon-
struction of U. uniramia (from Axsmith et al., 2000). 22. Cross section of U. resinosa cupule (C) showing ovule (O) and cupule stalk (B). Note distinctive
secretory cavities, e.g., below “C” (from Klavins et al., 2002). Bar = 1.0 mm.
 15372197, 2009, 1, Downloaded from https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.0800202 by CAPES, Wiley Online Library on [11/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
244 American Journal of Botany [Vol. 96

Figs. 23–33. Reproductive organs of corystosperms and Petriellales. 23. Short shoot (SS) of a corystosperm bearing several pedicels terminating in
cupules (arrows) of Umkomasia uniramia. Bar = 1.0 cm. 24. Dicroidium axis showing leaf attached to stem and short shoot (SS), which bears cupules of
U. uniramia. Bar = 2.0 mm. 25. Suggested reconstruction of Pilophorosperma geminatum Thomas (redrawn from Thomas, 1933). 26. Branch bearing
several Pteruchus sp. microsporophylls (courtesy C. P. Daghlian). Bar = 5.0 mm. 27. Pteruchus microsporophyll showing attached pollen sacs (arrows)

January 2009] Taylor and Taylor—Seed ferns from the late Paleozoic–Mesozoic
PETRIELLALES
This order of seed ferns was defined based on a single genus
of permineralized cupule containing seeds from the lower Mid-
dle Triassic of Antarctica (Taylor et al., 1994). A new family,
Kannaskoppiaceae, based on impression-compression speci-
mens, was later included in the order (Anderson and Anderson,
2003).
Ovulate organs— Cupules of Petriellaea Taylor et al. (1994)
are bilateral, elongate, and borne in clusters on a dichotomizing
axis (Fig. 30). A vascular bundle extends along the midrib of
the cupule with a trace given off to each ovule. The cupule wall
is thin, but there is no evidence of a preformed suture. Each
cupule contains from 2–6 small (1.0 mm in diameter), triangu-
lar ovules (Fig. 31), each attached to the adaxial surface of the
cupule surface at differing levels (Fig. 32). Ovules are orthotro-
pous with the integument thickened only in the corners; extend-
ing from the distal end of the seed is a short micropylar tube.
Morphologically, there is some superficial similarity between
the cupules of Petriellaea and those of Caytonia. While the
structure of the Caytonia cupules remains unknown, in Petriel-
laea the organization suggests that the cupule has evolved by a
proximal-distal folding of the megasporophyll (Fig. 32) (Taylor
et al., 1994), rather than by enrolling or folding laterally along
the midrib, the hypothesized ancestral conduplicate carpel or-
ganization sometimes attributed to the earliest angiosperms.
Nothing is known about pollen organs that were associated with
the Petriellaea cupules.
The Kannaskoppiaceae includes impression fossils of pollen
organs (Kannaskoppianthus), ovulate structures (Kannaskop-
pia) (Fig. 33), and foliage (Kannaskoppifolia) from the Upper
Triassic Molteno Formation of South Africa (Anderson and
Anderson, 2003). Leaves are cuneate to flabellate and lack a
petiole; the lamina is entire to deeply divided into segments
with anastomosing venation. Similar leaves from the Upper
Triassic of Argentina and Chile have been described as Rochip-
teris by Herbst et al. (2001). The seed-bearing organs Kan-
naskoppia are termed strobili with each ~2.0 cm long and are
attached in groups of two or three in the axils of the leaves.
Each consists of a short, proximal axis segment, which then
forks to form two secondary axes. Each of these bears two rows
of megasporophylls, with each sporophyll containing a single
cupulate ovule (Fig. 33); at maturity, the cupule splits into three
lobes. The pollen organ Kannaskoppianthus is up to 45 mm
long and constructed of two rows of microsporophylls arising
from a forked axis, each with five recurved, longitudinal pollen
sacs in a distal concavity protected by an operculum.
PELTASPERMALES
The peltasperms were initially described from the Upper Tri-
assic of Greenland and South Africa based on associated foli-
age (Lepidopteris Schimper), pollen organs (Antevsia Harris),
and seed-bearing organs (Peltaspermum Harris) (Thomas,
←
(courtesy C. P. Daghlian). Bar = 2.0 mm. 28. Suggested reconstruction of Pteruchus fremouwensis fertile axis (from Yao et al., 1995). 29. Section of per-
mineralized Pteruchus fremouwensis pollen sac containing bisaccate grains. Bar = 250 µm. 30. Suggested reconstruction of Petriellaea triangulata T.
Taylor et al. showing position of cupules on fertile axis. 31. Cross section of P. triangulata cupule containing three triangular seeds. Bar = 1.0 mm. 32. Dia-
grammatic cutaway of P. triangulata cupule showing seed attachment and proximal-distal folding of megasporophyll to form cupule (from Taylor et al.,
1994). 33. Branches bearing numerous Kannaskoppia vincularis Anderson and Anderson cupules (courtesy J. M. Anderson). Bar = 2.0 cm.
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245
1933; Harris, 1937). As defined by Townrow (1960), the group
included bipinnate foliage of Lepidopteris, peltate, ovulate re-
productive structures of Peltaspermum, and pinnately branched
pollen organs of Antevsia, which bore small, oval pollen grains.
Townrow (1960) was able to put the three parts together be-
cause L. ottonis Schimper, A. zeilleri (Nathorst) Harris, and P.
rotula Harris all bore characteristic blisters on the epidermis
and had the same stomatal structure. A unique feature of the
foliage of the peltasperms is the presence of intercalary pin-
nules on the rachis, termed Zwischerfiedern. Because of the
peltate megasporophylls, the group was at one time thought to
be morphologically isolated from other seed ferns. Today, how-
ever, the order encompasses a much wider range of morpholo-
gies and reproductive biologies than in the original concept.
The geographic and geologic range of the group has also ex-
panded, and peltasperms are now known from North America,
Europe (Kerp, 1988; Kerp et al., 2001), and the Russian plat-
form (Gomankov and Meyen, 1986; Naugolnykh and Kerp,
1996), and from the Pennsylvanian (Kerp et al., 2001) into the
Triassic. As currently circumscribed, the group was probably
more widespread during the Permian than the Triassic.
Ovulate organs—The ovulate organs of the peltasperms dem-
onstrate a wide range of morphological variability. Those from
the Rotliegend (Upper Pennsylvanian–Lower Permian) of Eu-
rope are unlike the structures for which the group was erected.
Based on repeated co-occurrences of material from the Rotlieg-
end of central Europe, the well-known foliage type Callipteris
conferta (Sternberg) Brongniart and the ovule-bearing reproduc-
tive organ Autunia Krasser (Fig. 34) were combined into Autunia
conferta (Sternberg) Kerp (Kerp, 1982, 1988). Autunia was orig-
inally introduced for strobili with helically arranged, fan-shaped
megasporophylls, each bearing one or two ovules on the lower
surface (Figs. 35, 36). Pollen-producing organs of Arnhardtia
(Zeiller) Haubold et Kerp are believed to belong to A. conferta
and consist of 5–9 pollen sacs borne on a peltate microsporophyll
(Barthel, 2006). Unlike previously described peltasperms, how-
ever, these pollen sacs contain monosaccate grains assignable to
Vesicaspora Schemel; grains are up to 50 µm in diameter.
The genus Peltaspermum, for which the group was estab-
lished, is known from both Gondwana and Laurasia, and in-
cludes ovulate organs (Harris, 1937) that consist of elliptical to
radial, peltate, umbrellalike discs (Figs. 37, 38), which are
sometimes termed peltoids (e.g., Meyen 1984; Naugolnykh,
2001). The discs (megasporophylls) are up to 1.0 cm in diame-
ter and helically arranged into simple linear-cylindrical strobili
(Fig. 39); ovules are borne in a ring on the lower surface of the
megasporophylls (Townrow, 1960). The number of ovules per
disc is variable.
Based on association, a concept which was expanded by
Meyen (1987) and termed assemblage genera, Poort and Kerp
(1990) emended the genus Peltaspermum, so that foliage and
ovulate structures are both circumscribed by the “natural” ge-
nus Peltaspermum. “Natural” taxon concepts similar to the one
suggested for A. conferta have been proposed for several other
Early Permian taxa, including Autunia naumannii (Gutbier)
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246 American Journal of Botany [Vol. 96

Figs. 34–42. Reproductive organs of peltasperms. 34. Suggested reconstruction of Autunia conferta ovuliferous organ (from Kerp, 1988). 35. Autunia
conferta ovulate organ showing megasporophylls (courtesy H. Kerp). Bar = 1.8 cm. 36. Suggested reconstruction of two A. conferta megasporophylls (from
Kerp, 1988). 37. Suggested reconstruction of Peltaspermum rotula megasporophyll showing several ovules (from Crane, 1985). 38. Peltaspermum martin-
sii (Kurtze) Poort et Kerp peltate megasporophyll showing scalloped margin (courtesy H. Kerp). Bar = 4.5 mm. 39. Suggested reconstruction of Peltasper-
mum thomasii axis bearing numerous megasporophylls with attached ovules (redrawn from Dilcher, 1979). 40. Suggested reconstruction of Peltaspermopsis
polyspermis (from Naugolnykh, 2001). 41. Suggested reconstruction of Lepidopteris frond with pollen organs of the Antevsia-type at the tip (courtesy M.
Zavada). 42. Suggested reconstruction of Antevsia zeilleri pollen organ showing pinnate axis bearing clusters of pollen sacs (from Crane, 1985).

Kerp (ovulate organs), Rhachiphyllum schenkii (Heyer) Kerp
(foliage), and Arnhardtia scheibei (Gothan) Haubold et Kerp
(pollen organs) (e.g., Kerp, 1988; Kerp and Haubold, 1988;
Barthel, 2001, 2006). Peltaspermum reproductive organs, de-
scribed from a site in Morocco, have certain features, i.e., resin-
ous bodies, that also occur on Rhachiphyllum Kerp foliage from
the same site (Kerp et al., 2001).
Peltaspermopsis polyspermis Gomankov is a natural genus
concept used for Late Permian peltasperms from Russia. The
plant includes vegetative stems, some with conspicuous
nodes, which are interpreted as evidence of seasonal growth
interruptions. Lanceolate leaves assigned to Pursongia
Zalessky are also considered to be part of the same plant,
along with reproductive organs in the form of seed-bearing

January 2009] Taylor and Taylor—Seed ferns from the late Paleozoic–Mesozoic
discs and their “racemose” aggregations (Fig. 40) (Naugol-
nykh, 2001).
The genus Meyenopteris Poort et Kerp has also been sug-
gested to be a “natural” genus of peltasperms (but see Karasev
and Krassilov, 2007), using the concept of Poort and Kerp
(1990), and includes the form taxa Lepidopteris natalensis and
Peltaspermum thomasii. Lepidopteris natalensis foliage is typi-
cally bipinnate, with small blister-like swellings on the axes
and intercalary pinnules (Zwischerfiedern), characters that are
diagnostic for the genus. The blister-like swellings also occur
on the ovule-bearing organs of P. thomasii, which made it pos-
sible to connect the two plant parts. In this species, two ovules
(3.0 mm long) occur on the lower surface of each disc-shaped
megasporophyll. Lopadangium Zhao et al. emend. Gomankov
et Meyen is used for peltaspermaceous ovulate organs that can-
not be correlated with foliage or other organs.
Pollen organs— In the Mesozoic forms, such as Antevsia,
pollen sacs are borne on branches. Antevsia includes branched
axes that bear lateral groups of 4–12 elongate pollen sacs at
their distal tips (Fig. 41). Townrow (1960) described Antevsia
as pinnate, with the main axis bearing alternate laterals in one
plane and secondary branches produced irregularly. Although
these branches are described as microsporophylls, the branches
are not expanded at their tips (Fig. 42). Branches show the char-
acteristic blister-like swellings that also occur on Lepidopteris
fronds. Pollen sacs are up to 5 mm long and dehisce longitudi-
nally. Like the majority of the Mesozoic Gondwanan pel-
tasperms, pollen grains are small (23–40 µm) and oval, with a
distal sulcus; if dispersed, these grains would be assigned to
Cycadopites Wodehouse ex Wilson and Webster.
In the Late Pennsylvanian–Early Permian peltasperms, the
pollen organs are not known in great detail. Perhaps the best-
known form is Callipterianthus arnhardtii Roselt (1962) from
the Rotliegend (Pennsylvanian–Early Permian) of Germany.
This fossil has a frond-like (pinnate) organization with pollen
sacs attached to reduced pinnules in the distal part of the frond.
Sterile pinnae occur in the proximal portion of the frond, and
intercalary pinnules are present on the rachis. Although the
specimen is partially sterile, a correlation with sterile foliage is
difficult. Callipterianthus arnhardtii has been suggested to rep-
resent the pollen-producing organ of either Autunia naumannii
or Arnhardtia scheibei (see Kerp, 1996). Barthel (2006), how-
ever, assigned pollen organs of the Pterispermostrobus type,
which is composed of elongate pollen sacs attached to a peltate
microsporophyll, to Arnhardtia scheibei.
DISCUSSION
We have provided information about the reproductive parts
of the glossopterids and the major groups of late Paleozoic–
Mesozoic seed ferns so as to provide base-level data that can be
used in discussing either real or inferred relationships with the
angiosperms. Although there are, of course, still gaps in our
knowledge of these groups, a large amount of data has been
gathered in the last ten years, especially for the peltasperms and
the corystosperms. Except for the Petriellales, these groups
have been considered as possible angiosperm ancestors because
all have ovules attached to a more or less leaflike megasporo-
phyll. Some groups have cupules that partially enclose their
ovules, i.e., the corystosperms, Petriellales, and caytonialeans.
Others, such as the peltasperms, do not have enclosed ovules.
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247
Nevertheless, at one time or another, taxa in all of these groups
have been suggested as demonstrating morphological stages of
angiospermy based on the implied homologies of the reproduc-
tive organs or as showing some intermediate stage in the trans-
formation of an angiosperm organ, usually a carpel. Space does
not permit a discussion of what does or does not constitute a
unique angiosperm feature, and all are familiar with the limita-
tions imposed on defining such a feature in the fossil record.
The problem of real or implied homology using fossils is exac-
erbated by the absence of structurally preserved fossils in some
of the groups and by the use of composite terminals in some
phylogenetic analyses, rather than whole plants reconstructed
on the basis of attachment, shared anatomical characters, or
consistent and exclusive co-occurrence.
Like many other Paleozoic and Mesozoic pteridosperms, the
glossopterids have been suggested as possible ancestors to the
angiosperms. Perhaps the earliest advocate of this relationship
was Plumstead (1956), who interpreted the glossopterid ovulate
organs as bisexual. The glossopterid-angiosperm connection
has periodically gained support as additional information about
the ovulate organs has been published and hypotheses advanced
that attempt to homologize these organs with those of other
seed plants (e.g., Melville, 1983). For example, Retallack and
Dilcher (1981) championed the idea initially suggested by Steb-
bins (1974) of the glossopterids as flowering plant progenitors.
They proposed a transformational series in which the number of
ovules on the megasporophyll is reduced to one. The me-
gasporophyll would then enclose the ovule to form a second
integument, and the subtending leaf would be transformed into
the carpel. Doyle (2006) also discussed this idea as a way to
evolve the angiosperm carpel, with perhaps Caytonia as an in-
termediate. Geologic time remains an obstacle to the acceptance
of glossopterids as angiosperm ancestors, however, because the
glossopterids disappeared around the Permian–Triassic bound-
ary (~251 million years ago [Ma]), while the first recognizable
angiosperm megafossils do not occur until the Lower Creta-
ceous (Barremian–Aptian, ~145 Ma; Friis et al., 2006).
Phylogenetic analyses have treated the glossopterids as a
single, composite terminal, reconstructed of various organs, of-
ten from different sites, thus providing little resolution of the
group’s diversity in the analysis (e.g., Nixon et al., 1994; Doyle,
2006; Hilton and Bateman, 2006). There has been some attempt
to relate isolated organs based on associations and anatomy,
and in some cases, the vegetative parts of plants have been re-
constructed, e.g., Glossopteris schopfii Pigg and G. skaarensis
Pigg from Antarctica (Pigg and Taylor, 1993) and G. homeva-
lensis Pigg et McLoughlin from Australia (Pigg and Nishida,
2006). A permineralized ovulate structure from the Late Perm-
ian of Antarctica could be attributed to G. schopfii leaves based
on similar anatomy (Taylor and Taylor, 1992), and Nishida et
al. (2007) suggested that Homevaleia gouldii ovulate organs
belong to leaves of G. homevalensis. These studies represent a
first step in taking what may be a heterogeneous group and de-
veloping a framework that can be used to test subsequent hy-
potheses of relationships, not only within the glossopterids, but
with other seed plants. Until homologies can be established,
there is currently insufficient evidence to link the glossopterids
with any other major group of plants, either as progenitors or
descendants.
The late Paleozoic–Mesozoic seed ferns have never repre-
sented a natural group, and whether each of the four orders dis-
cussed represents a monophyletic group is also far from settled,
because no complete phylogeny has been presented for any of

248 American Journal of Botany
these plants. It is clear from our current state of knowledge of
these groups that the discovery of additional specimens, espe-
cially those in different modes of preservation, and research
that focuses directly on these groups is what is needed in the
years ahead if more robust relationships are to be fully realized.
In almost all phylogenetic analyses of seed plants published to
date, each of these seed fern orders has been treated as a clade
and is often represented as a single, composite terminal, even
when reconstructions have been based only on association.
Some reconstructions of “whole” plants have used organs from
different continents! In the case of the corystosperms, which
currently represent the best-known group overall, the remark-
able uniformity of both pollen and ovulate organs throughout
Gondwana suggests that at least the Gondwanan representa-
tives may constitute a single clade. Artabe and Brea (2003) con-
clude that the group is not a natural one, but their analysis was
based primarily on stem anatomy, a character that is variable
and may simply reflect habitat. It is interesting to note that the
Dicroidium plants reconstructed from South America and Ant-
arctica suggest the presence of two quite different growth archi-
tectures (based on stem anatomy), although their reproductive
organs and foliage are very similar (Petriella, 1978; Taylor,
1996).
The Caytoniales are perhaps the most poorly known of the
Mesozoic seed ferns, yet it is the cupule-bearing megasporo-
phyll of Caytonia that has received the greatest amount of at-
tention in scenarios leading to the origin of the angiosperm
carpel and seeds with two integuments (e.g., Doyle, 2006).
While important details about the reproductive biology of the
Caytoniales are known, there is nothing to suggest that they
possessed anything other than a distinctly gymnospermous
seed-plant reproductive system. As is the case with some of the
peltasperms, the Caytoniales produced saccate pollen, a mor-
phological character state that both groups share with the Paleo-
zoic callistophytalean seed ferns. It is important to point out
that prior to the recognition of the Pennsylvanian Callisto-
phytales, pollen of this morphotype would have implied affini-
ties with some conifer or cordaite seed-plant group. If in fact any
of these groups are distant progenitors of flowering plants, then
the fact that many possess saccate pollen may simply reflect that
the pollination syndrome did not keep pace with developmental
changes in the function of the mature microgametophyte. The
primitive nature of the male gametophyte is also underscored
by the discovery of flagellated sperm in the glossopterid Hom-
evaleia that, aside from their reduced size, morphologically re-
semble the male gametes of cycads and Ginkgo.
At the present time, the Peltaspermales are perhaps the most
perplexing of all of these late Paleozoic–Mesozoic seed plants.
The morphology of the Autunia ovulate organ is distinct from
that of the Peltaspermum known from numerous other sites,
including Siberia and Gondwana; however, the absence of any
anatomical features significantly impedes progress in resolving
this seed-plant group. This is also the case for the Petriellales,
although structurally preserved cupules and seeds are known,
no other plant parts have been found to date. These isolated
organs are included within the assemblage known as seed ferns
both for convenience and necessity, because all produce ovules
on a modified, leaflike structure.
We have noted earlier that many of the ideas regarding the
origin of the flowering plants have been based on scenarios in
which megasporophylls become transformed so that the lamina
covers the ovule(s), and this stage has been cited as a precursor
to the evolution of a carpel. Although this was the original con-
15372197, 2009, 1, Downloaded from https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.0800202 by CAPES, Wiley Online Library on [11/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
[Vol. 96
cept of Thomas when he described the Caytoniales, subsequent
theories have used the cupule to form a second integument of
the seeds, and other tissue to derive the carpel. All these sce-
narios suggest that megasporophyll closure is longitudinal, a
theoretical basis that is supported by conduplicate folding in
certain extant flowering plants and even in some Early Creta-
ceous angiosperms (e.g., Archaeanthus Dilcher et Crane,
Lesqueria Crane et Dilcher, Protomonimia Nishida et Nishida).
What is interesting about Petriellaea is that megasporophyll
closure may have proceeded in a different pattern in which the
tip of the megasporophyll is bent back toward the base or peti-
ole (Taylor et al., 1994). This may also represent the pattern in
Caytonia.
Beginning in the late Paleozoic and continuing into the Me-
sozoic, almost all seed-plant groups demonstrate increasing
levels of cupule and megasporophyll closure. Historically, this
has been interpreted as a response to predation or other selec-
tive advantages associated with ovule protection. While this
structure–function relationship may have some validity, another
hypothesis suggests that intercalating sterile tissue between
ovules and pollen was a key physiological innovation in the
transition from gymnospermy to angiospermy (Zavada and
Taylor, 1986a). In this context, closure of the megasporophyll
was directly linked to the evolution of self incompatibility and
a change from postzygotic to prezygotic selection. There would
have been strong selective pressure on pollen to develop alter-
native mechanisms to insure compatibility with the stigma in
angiosperms. More efficient pollen transfer, the evolution of
accessory floral parts to influence potential pollinator behavior,
and the loss of sacci may have been some of the evolutionary
stimuli in the transition to angiospermy. Interestingly, to the
extent where pollen for a group is known, all the late Paleozo-
ic–Mesozoic seed ferns, including the glossopterids, possess
saccate pollen, with the exception of Cycadopites grains recov-
ered from the peltasperm Antevsia (Townrow, 1960). This is in
contrast to the situation in angiosperms where sacci are found
only in the extant genus Lactoris Philippi (Zavada and Taylor,
1986b)
While the morphological features showing a transition to the
angiosperm carpel may be difficult to resolve based on fossils,
the fossil record can sometimes offer another dimension—the
gametophyte phase—in deciphering the transition from gymno-
spermy to angiospermy. The discovery of male gametes similar
to those in cycads and Ginkgo in the glossopterid Homevaleia
(Nishida et al., 2003) contributes important details about one
group of plants thought to represent potential flowering plant
progenitors. The inclusion of this character state in a data ma-
trix may suggest glossopterid affinities with the cycads and
Ginkgo—a conclusion that is predictable, but may not be cor-
rect if the glossopterids are more heterogeneous than previously
thought.
We believe that our understanding of all these fossil groups
will significantly advance when anatomically preserved speci-
mens are found and when there are better reconstructions that
either combine organs unique to a site or that link organs based
on distinctive synapomorphies (Taylor et al., 2006). With the
exception of the Petriellales and Corystospermales, which are
known from permineralized and compressed fossils, we know a
great deal about the morphology and even the epidermal anat-
omy of some groups like the Caytoniales and Peltaspermales,
but nothing about the internal anatomy. These data will be criti-
cal in characterizing homologies with other seed plant groups.
In addition, details about where the reproductive organs are

January 2009] Taylor and Taylor—Seed ferns from the late Paleozoic–Mesozoic
produced, how they are attached, and how much diversity they
demonstrate within each group are important areas that require
additional attention. This is especially true of the Glossopterid-
ales. As to the question posed in the title of this paper—the
same question that has been addressed countless times since
Darwin’s declaration regarding the “abominably perplexing”
problem of angiosperm origins (Crepet, 2000)—from our per-
spective, the mystery remains intact. We would offer one ca-
veat, however. In our opinion, it will be more productive to
attempt to solve Darwin’s mystery if there were greater atten-
tion directed at mining the rock record in the hope of discover-
ing more informative and new specimens, than to continue to
construct new phylogenies using the same, often ambiguous
characters.
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