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itteilungen des
ärntner otanikzentrums
lagenfurt
ulfenia 17 –
An advanced peltasperm Permoxylocarpus trojanus Naug. from the
Lower Permian of the Urals (Russia): an ancient case of
entomophily in gymnosperms?
erge augolnykh lexei skolski
Summary: he evolutionarily advanced gymnosperm of peltaspermalean a nity Permoxylocarpus
trojanus aug from the ower ermian deposits of the rals ussia is described emale reproductive
organs of P. trojanus are spherical semiclosed peltate capsules with – enclosed seeds oth
macromorphology and microstructure including epidermalcuticular characters and the anatomical
structure of conducting tissues are characterized terile leaves of Praephylladoderma leptoderma aug
provisionally belonging to the same parent plant are also described ome aspects of peltasperm
evolution during the ate alaeozoic – arly esozoic are brie y summarized he occurrence of
enclosed ovuliferous organs capsules in some peltasperms the structure of their pollen grains as well
as some palaeoentomological evidences suggest the gradual shift from anemophily to entomophily in
evolution of these gymnosperms
Keywords: peltasperms evolution ermian seedbearing organs preangiosperms fossil records
entomophily
teridosperms sensu lato were widely distributed seed plants during the ate alaeozoic his
very diverse group is commonly regarded as a grade rather than a wellsupported clade espite
that a general evolutionary pattern of this group can be recognized entailing the morphology
of the fertile organs
he most ancient pteridosperms known from the pper evonian agenostomales Elkinsia
polymorpha, Moresnetzia zalesskyi and some other related forms
see these papers for further references had
relatively simple fertile fronds or monopodially branched shoots without extended leaf lamina
heir ovules andor ovulate cupules were attached to the branch tips urther pteridosperm
evolution entailed the development of fertile fronds which were very similar or almost identical
to the ordinary vegetative fronds but with the ovules attached to the leaf lamina or frond rachis
rigonocarpales allystophytales see these papers
for further references teridosperms of this grade were common in arboniferous vegetation
especially in the equatorial belt
uring the ermian the pteridosperms became widely distributed more diverse and advanced than
their arboniferous members ithin these plants the modi ed fertile fronds with reduced leaf
laminae were gradually transformed into specialized ovuliferous organs n some taxa belonging
mainly to the order eltaspermales these organs provided better protection for the ovules
partially covered by the derivates of leaf lamina ie cupules and capsules of di erent kinds
eltaspermalean pteridosperms or peltasperms probably gave rise to the esozoic pteridosperm
orders orystospermales and aytoniales aytoniales are often regarded as preangiosperms
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etc vuliferous organs of
corystosperms can be interpreted as modied megasporophylls that originated
from the fertile fronds of primitive pteridosperms he ovules of many esozoic pteridosperms
or at least some of them eg Petriellaea aylor et al Caytonia arris see for review
were almost completely enclosed by modied capsules linking these plants
morphologically close to the angiosperms
ermian peltaspermalean pteridosperms may have linked to the relatively primitive arboniferous
and progressive esozoic forms and because of this intermediate position they are of special
palaeobotanical interest ermian peltaspermalean pteridosperms possessed a unique syndrome
of mixed characters represented by peltate discoid seedbearing organs aggregated into
racemose structures and pinnate or simple leaves ig herefore this group is important for
understanding the evolutionary trends in gymnosperms and other seed plants
n the present paper new peltaspermalean pteridosperm material collected from the ower
ermian deposits of the rals ussia is described he plant characterized here was attributed
to the order eltaspermales family ngaropeltaceae, and named Permoxylocarpus trojanus aug
riginal description of the material was published in ussian only with the exception of an
nglish diagnosis as it is required by the nternational ode of otanical
omenclature his plant shows some characters that are more typical of esozoic preangiosperms
rather than of alaeozoic pteridosperms n addition to the general morphological observations
some anatomical and epidermalcuticular characters of the plant were studied omparison
were drawn between P. trojanus and the most closely related taxa among morphologically similar
ermian and riassic plants ie Peltaspermum arris Angaropeltum oweld Caytonia arris
Umkomasia homas Spermatocodon homas Pilophorosperma homas Petriellaea aylor et al
and Ktalenia circularis rchangelsky
aterials and methods
he main part of the material studied was collected from hekarda locality ungurian of the
israls erm district ussia by the rst author during several eld seasons in –
he holotype of Permoxylocarpus trojanus aug was provided by the late rof ofronitsky
erm tate niversity ome additional specimens were collected at the hekarda locality
by harov alaeontological nstitute oscow in – and provided by
omankov omarov otanical nstitute t etersburg ne specimen from the simbal
collection oscow ig was also studied or exact position of the plantbearing
strata cited in the text see pecimens characterizing the pper
ermian peltasperms of echora coal basin ig – were provided by ukhonto
tate ernadsky eological useum oscow the racemose aggregation of seedbearing discs
ig from the pper ermian of ussian arast was given to the authors by urago
ladivostok
Figure 1. – Peltaspermum sp a racemose aggregation of seedbearing discs found in close association with leaves of
callipterid morphology albeiskian ormation rzumskian stage pper ermian echora coal basin dzva iver
outcrop – – Comia sp leaf with unicoherent venation eidinskian ormation azanian stage
pper iddle ermian echora coal basin yraga iver erkhesyryaginskoe locality – Peltaspermum buragoae
aug in manuscr a racemose aggregation of seedbearing discs found in close association with leaves of callipterid
morphology itza oristic assemblage godinzinskian ormation azanian stage pper ermian central part of
>>>
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Permoxylocarpus trojanus: an ancient case of entomophily in gymnosperms
rimorie ussian arast inegorka iver left bank of amenisty spring – Rhachiphyllum = Callipteris adzvense
alessky aug last order pinna eidinskian ormation azanian stage pper iddle ermian echora coal
basin yraga iver erkhesyryaginskoe locality – Rhachiphyllum = Callipteris adzvense alessky aug two
last order pinnae eidinskian ormation azanian stage pper iddle ermian echora coal basin izhne
yryaginskoe locality borehole depth m cale bar = cm
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he specimens were macerated in the chulzes reagent according to standard procedure
and the cuticles obtained were studied by means of the light microscope and the
tereoscan scanning electron microscope ambridge
erminology
upule is a commonly used term for the description of closed or semiclosed derivates of planate
seedbearing structures his term is ambiguous however in relation to dierent representatives
of rigonocarpales eltaspermales aytoniales and some other groups of gymnosperms whose
cupulelike organs of independent origin may not be considered as homologues ecause of that
we prefer to use term capsule instead of cupule for any closed or semiclosed ovuliferous organs
of gymnosperms his term is welldened morphologically but it is not loaded by phylogenetic
connotations he term capsule was used by in a very similar way
esults
eltaspermales aylor
ngaropeltaceae oweld
= ardiolepidaceae eyen emend nom illeg
Permoxylocarpus Naugolnykh, 2007. emale generative organs are spherical capsules with central
stalk and with seeds disposed inside the capsule uter surface of the capsule bears radial ribs
he genus diers from the most closely related genera Sylvocarpus aug and Angaropeltum oweld
= Cardiolepis euburg nom illeg in showing well developed radial ribs disposed on the surface
of the capsule from Ktalenia rchangelsky in a considerably larger number of enclosed ovules
– instead of – and also in another type of associated leaves lanceolate leaves with the
parallel venation of Praephylladoderma leptoderma aug instead of pinnate leaves of Ruorinia
sierra rchangelsky from Petriellaea aylor et al in more numerous ovules per capsule –
instead of – and in the central position of the capsule stalk stalk or pedicel of Petriellaea
attached to the side part of the cupule from the corystosperm ovulate organs belonging to
Umkomasia homas and closely related genera in having more numerous ovules – instead
of – per ovuliferous organ and also in the position of the stalk central for Permoxylocarpus
and marginal for Corystospermum, Umkomasia and related forms like Pilophorosperma ssociated
leaves of Permoxylocarpus and corystosperms are also very dierent such as simple lanceolate
leaves of Praephylladoderma provisionally linked to Permoxylocarpus and compoundly pinnate
leaves of Dicroidium characteristic of corystosperms here is a certain similarity between
Permoxylocarpus and Caytonia cupules he most important dierence between these genera is the
position of the stalk which is central for Permoxylocarpus and marginal for Caytonia. oreover
Caytonia had compound leaves of Sagenopteris type with the net venation whereas Permoxylocarpus
had simple lanceolate leaves
Permoxylocarpus trojanus Naugolnykh. emiclosed capsules spheroid with umbrellashaped
peltate round lamina on central stalk arginal parts of the lamina strongly turned downward
uter surface of the capsule bears smooth radial ribs disposed around the place of the stalk
attachment void areas between the ribs probably corresponding to the places of seed
attachment seed scars urface of the capsule which is more distant from the stalk is smoother
or bears unclear concentric folds ach capsule bears – seeds located inside the capsule and
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Permoxylocarpus trojanus: an ancient case of entomophily in gymnosperms
surrounding the central stalk ome seeds could be immature because of the asymmetrical shape
of the capsule
he collection studied includes six capsules ig ig attributed to this species he mode
of their preservation is compressionimpression ome parts of compressed material are missing
therefore one can see the imprint of the outer surface of the generative organ
he holotype ig ig exposed from its adaxial surface has a slightly deformed
asymmetrical spheroid shape ther specimens have same macromorphological features but
some of the capsules ig ig expose their adaxial surface like the holotype and
others show their abaxial surface ig ig
Figure 2. – macromorphology of Permoxylocarpus trojanus aug seedbearing capsules – holotype
rutaya atushka locality – syntypes hekarda locality layer oshelevskian ormation
ungurian stage ower ermian ylva iver asin erm region – general morphology of angaropeltidian seed
bearing capsule – abaxial surface – radial rib – stalk with conducting tissues – way for pollen grains to seed
micropyle cale bar = cm
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Figure 3. – – macromorphology of Permoxylocarpus trojanus aug seedbearing capsules – holotype
rutaya atushka locality – syntypes – specimen from simbal collection
hekarda locality layer oshelevskian ormation ungurian stage ower ermian ylva iver asin
erm region – Praephylladoderma leptoderma aug leafy shoot hekarda locality layer oshelevskian
ormation ungurian stage ower ermian ylva iver asin erm region cale bar = cm
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Permoxylocarpus trojanus: an ancient case of entomophily in gymnosperms
here is a small unclear round depression at the capsules upper part his depression corresponds
to the place of the stalk attachment ig he adaxial surface of the capsule has nearly
alternating smooth furrows and wide radial ribs he ribbed surface forms the rst concentric
band closest to the stalk he next concentric band being farther away from the stalk and located
nearer the outer margin of the capsule is separated from the previous band bearing the lobes by
the welldeveloped denitive fold he fold is represented as a concentric rib on the abaxial outer
surface of the fructication his second band bears poorly developed concentric and radial folds
ext the most outer band third band forming margins of the capsule has a smoother surface
than the second band here is one more fold between the second and the third concentric band
of the capsule his fold corresponds to the place of curving of the capsules umbrella he lower
adaxial surface of the capsules is covered by radial ribs he margin of the capsule is entirely
smooth
he capsule is spherical with the margins of the capsules umbrella directed downwards
ostdiagenetic compression of the sediment leads to a atter and smoother shape of the
fructication he original form of the capsule was more threedimensional almost regularly
spherical vules disposed inside the capsule and places of their attachment correspond to the
lobes of the rst capsule band he ovules surround the stalk of the capsule in a radial arrangement
ig
Anatomical structure of conductive tissue. piece of the capsule pedicel of the holotype was
macerated umerous tracheids of the conductive bundle were extracted and studied using
ig – wo types of tracheids were distinguished ome tracheids are narrow their
diameter – m in transsection with helical thickenings on the cell walls ig – he
tracheids of another type are wider – m in diameter with circular or oval bordered pits
diameter of their borders is ca – m arranged in opposite to alternate pattern into rarely
vertical rows on lateral cell walls ig sometimes cooccurring with helical ig
or reticular ig thickenings robably the narrow tracheids of the rst type belonged to
protoxylem whereas the wide ones were taken from metaxylem
Epidermal-cuticular structure of the capsule. uticles from the three dierent areas of the
capsule shown on ig and ig were studied ach area has its own specic cuticular
characters
uticles of the rst type were taken from the lower part of the capsule near the margin of the
capsule ig he cuticle is relatively thick with numerous cells of isometric outlines
mostly hexagonal t the central part of almost every cell there are small papillalike structures
with shallow folds or depressions on the uplifting top verage diameter of the cells is – m
sometimes slightly more
uticles of the second type were taken from the outer side part of the capsule his cuticle is
also quite thick robust with well developed polygonal epidermal cells of prolonged outlines
ig ell size is × m in average apillae or stomata were not found imilar cuticles
are known for Phylladoderma tscheremuschca saulova from the ower atarian rzhumskian of
the olga iver basin close vicinity of azan right bank of the olga iver echischi village
heremushka outcrop l and P. sentjakensis saulova from the ower
azanian entjak village ama iver l
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Figure 4. natomy of Permoxylocarpus trojanus aug seedbearing capsules preparation was made from the specimen
shown on ig and ig – wide tracheid with circular bordered pits – narrow tracheid with helical to
reticulate thickenings no bordered pits – narrow tracheid with helical thickenings left and wide tracheid with
circular bordered pits and helical thickenings right – wide tracheid with circular bordered pits and reticular
thickenings – – epidermalcuticular structure hekarda locality layer oshelevskian ormation
ungurian stage ower ermian ylva iver asin erm region cale bar = m – m –
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Permoxylocarpus trojanus: an ancient case of entomophily in gymnosperms
uticles of the third type were taken from inner surface of the capsule hese cuticles are very
thin only some weakly preserved cell walls are seen ig urface of the cuticle is relatively
smooth with some small but distinct papillae
Associated organs. he ungurian oristic assemblage with Permoxylocarpus trojanus is taxonrich
everal wholeplantconcept reconstructions
have been proposed for some of the ungurian higher plants he best documented taxa from
these deposits are Sadovnikovia belemnoides augolnykh sporophylls Viatscheslaviophyllum
sp phylloids lepidophyte — Bowmanites biarmensis augolnykh strobiles Sphenophyllum
biarmicum alessky sterile leaves and stems sphenophyll — Equisetinostachys sp fertile
zones and sporophylls Phyllotheca stenophylloides alessky sterile leafy shoots Sachyogyrus
multifarius alessky fertile stems and sporophylls Phyllotheca biarmica alessky sterile leafy
shoots sphenophytes — Ptychocarpus distichus augolnykh fertile fronds Pecopteris uralica
alessky sterile fronds marattialean fern — Alternopsis stricta augolnykh ovuliferous organs
and specialized fertile leaves Psygmophyllum cuneifolium utorga chimper sterile leaves
pteridosperms — Rhachiphyllum al Callipteris retensorium alessky augolnykh sterile
fronds Peltaspermum sp ovuliferous organs – seedbearing discs Permotheca disparis alessky
augolnykh polliniferous organs peltaspermalean pteridosperms — or Peltaspermum
retensorium alessky species promoted to the natural status
Gaussia imbricata augolnyh ovuliferous organs Ruoria spp normal well developed
leaves Nephropsis Sulcinephropsis crinitus luchova bracts — Scirostrobus pterocerum
augolnykh oweld et augolnykh ovuliferous organs Sylvella sp seeds Ruoria spp
normal well developed leaves Nephropsis Sulcinephropsis sp bracts Lepeophyllum sp short
scalelike leaves vojnovskyaleans he two last mentioned reconstructed wholeplantconcept
taxa are probably ecological varieties or morphs of one and the same natural species hus a
general type of the associated sterile leaves is known for almost all generative organs of the most
common plants of the ora
f generative organs are found the sterile leaves belonging to the same plant are already present
in collections containing many plant megafossils collected from one and the same locality his
corresponds to the socalled arrisrule which is often cited in palaeobotanical literature and
is based on the simple empirical observation that commonly each plant produces many more
sterile leaves than generative organs
n the case of Permoxylocarpus trojanus we can also use a typological model as a base for
reconstruction of relationship between dierent organs or such a model we can use some
closely related forms belonging to the same taxonomic group eg Angaropeltum Cardiolepis,
nom illeg from the slightly younger deposits azanian of the ussian platform and is
rals ll of the earlier described species of Angaropeltum A. piniformis euburg oweld
A. sentjakensis saulova oweld had the sterile leaves of Phylladoderma type owever leaves
of this type have never been described from the ungurian of the israls before ence after
careful reinvestigation of the previously collected material and additional eld work the sterile
leaves of the ungurian phylladodermas were found hey are described below and attributed
to Praephylladoderma leptoderma aug ost probably these leaves belonged to the same parent
plant as those of Permoxylocarpus trojanus.
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Praephylladoderma Naugolnykh. traplike narrow leaves with wedgelike base and round apex
ig enation parallel veins robust at the leaf base but narrower in the leaf apex wo
main veins emerging from the leaf base eins never come out to the leaf margins eins always
bifurcating three to four times simple veins are absent eaf lamina very thin unresisting to
maceration pidermal structure is unknown his genus diers from the most closely related
genus Phylladoderma alessky in very narrow leaf lamina and thin cuticles unresistent to
maceration in chulzes reagent
pecies composition type species P. leptoderma aug and probably some undescribed forms from
the fumian and lowermost azanian of the israls and amaolga ivers basin
Praephylladoderma leptoderma Naugolnykh. ong narrow lanceolate leaves supercially similar
to Cordaites but having another pattern of venation eaves simple linear with wedgelike
narrow base aximal width of the leaf disposed near its apex eaf apex round apexes of the
young undeveloped leaves can be slightly acute enation regularly dichotomous proximal parts
of veins robust middle and apical parts of the veins rather thin wo distinctive basal veins
entering the leaf base ach vein dichotomously branching up to four times verage length of the
leaves is – cm average width of the leaves is cm here are four leafy shoots in the collection
studied ne of them is shown on ig hey possess leaves preserved in organic connection
to the shoots eaves arranged in loose spiral order lace of leaf attachment marked on the shoot
by small uplifting having a small subtriangular scar of the leaf attachment in its upper part
iscussion
he plants most similar to Permoxylocarpus trojanus are Sylvocarpus armatus aug
a and Angaropeltum oweld such as A. piniformis euburg oweld initially described
by and later reinterpreted by as well as the closely
related but less known species A. sentjakensis saulova oweld he most important dierence
between Sylvocarpus and Permoxylocarpus is the absence of radial ribs for the rst genus and its
almost completely closed seedbearing capsule Angaropeltum piniformis is a characteristic species
for azanian of the echora israls
emale generative organs of A. piniformis are spherical capsules enclosing numerous seeds of
Nucicarpus euburg type he seeds are also very distinctive and characterized by long and
narrow micropylar parts he stalk of the capsule is attached to its adaxial surface at the center
hus the general structure of both A. piniformis and Permoxylocarpus trojanus is very similar he
only important dierence is the presence of the welldeveloped sculpture of the Permoxylocarpus
trojanus capsules surface consisting of the above mentioned radial ribs and furrows as well as a
stronger vascularization and a smaller size of the P. trojanus capsules terile leaves of A. piniformis
Phylladoderma alessky and P. trojanus Praephylladoderma aug are basically very similar too
A. piniformis and some poorly documented forms from the mian oadian of the israls
may be regarded as evolutionary descendants of P. trojanus.
very similar pattern of macromorphological characters is known for another gymnosperm
Ktalenia circularis rchangelsky his species originates from the arly retaceous of anta ruz
rovince in rgentina K. circularis has ovuliferous capsules which were also previously described
as cupules attached to the fertile axis in opposite order probably at the basal area of several linear
bracts ne or two ovules were present
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Permoxylocarpus trojanus: an ancient case of entomophily in gymnosperms
in each capsule he short undeveloped pedicel of the capsule appears to be lateral ie attaching
to the capsules lateral side
he most notable dierence between K. circularis and Permoxylocarpus trojanus is the considerably
greater number of seedsovules which are present in the latter species he type of associated
leaves is also very dierent Ktalenia circularis had sterile tripinnate fronds of Ruorinia sierra
rchangelsky, in contrast to the simple lanceolate leaves of Praephylladoderma leptoderma, which
probably belong to the plant with the capsules of Permoxylocarpus trojanus.
nother plant which is morphologically similar to Permoxylocarpus trojanus has been described
from the riassic of ntarctica et al as Petriellaea triangulata aylor elueyo et
aylor emale generative organs of P. triangulata were interpreted as small cupules with vascular
strand of reticulatescalariform tracheids in the stalk pedicel dividing into discrete ve or six
bundles run to the ovules he ovules are arranged into a single oblique row or two rows inside the
cupule hese ovules are small × mm and ovoid et al similar to the ngaran
Nucicarpus. he most important dierence between P. triangulata and Permoxylocarpus trojanus is
the asymmetrical position of the cupule pedicel of P. triangulata attached to the side part of the
cupule noter dierence is the smaller number of seeds incapsulated in the cupule of Petriellaea
triangulata and the smaller size of the cupule homologous organ to the Permoxylocarpus trojanus
capsule
here are some characters in common between Permoxylocarpus and Caytonia
he latter genus was studied in detail by excellent works of
he basic similarity between Permoxylocarpus and Caytonia, ie general shape of the capsule as
well as similarity of Permoxylocarpus with some other esozoic pteridosperms especially of the
orystospermaceae family shows that these groups can be regarded as one grade in pteridosperm
evolution racheids with bordered porepairs as known for Caytonia
are similar to the tracheids of Permoxylocarpus.
he most important dierences between ermian Permoxylocarpus and Angaropeltum on the
one hand and esozoic Corystospermum, Umkomasia and other related corystosperm taxa
based on seedbearing organs, Caytonia, Petriellaea and Ktalenia on the other hand are the
mode of the stalk attachment as well as the general shape of exostomium ie the aperture for
penetrating of pollen grains inside the capsule Permoxylocarpus, Sylvocarpus, and Angaropeltum
had a central stalk with the ovules attached to the adaxial surface of the capsule in radial order
ike Caytonia, Corystospermum, Umkomasia, Petriellaea and Ktalenia they had a marginally
disposed stalk and the ovules of the latter two genera were attached to the internal surface of the
incurvated laminalike seedbearing capsule enclosed to a dierent extent semiclosed or even
almost open for Corystospermum and Umkomasia and almost completely closed for Petriellaea,
Caytonia, and Ktalenia. he exostomium of Permoxylocarpus and Angaropeltum was circular and
distinctly larger the one of Sylvocarpus was splitlike the one of Caytonia was semilunar, and it
was rounded in Ktalenia. Corystospermum and Umkomasia had no exostomium at all because of
a reduction of the capsule and rather large ovules n general all the forms listed above show a
pattern characterized by a decrease in seed number and reduction of the capsule size
volutionary predecessors of the angaropeltidians were representatives of eltaspermaceae recorded
from uppermost arboniferous and ower ermian deposits
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hese plants had open peltate seedbearing discs which most probably were windpollinated he
evolution of female reproductive organs in the peltasperms appears as a gradual transition from
the open discs to almost enclosed capsules which provide additional protection
for ovules his trend shows a certain resemblance to the transformations
of pistilate inorescences from simple spikes and globose heads to discshaped inorescences
in the tribes orstenieae and astilleae and then to the enclosed syconia in the tribe iceae of
the angiosperm family oraceae
iglike owers with enclosed perigone formed by completely connate tepals are also found
in Aspidistra locii rnautov ogner belonging to the family onvallariaceae
lthough the seedbearing organs of peltasperms are not homologues of the
angiosperm owers or inorescences we may compare them as organs of similar functions
ithin extant plants the capsulelike female reproductive organs occur only in taxa which
are highly specialized in pollination by arthropods like g wasps of the family goninae in
Ficus or soil amphipods or collembolans in Aspidistra
t is no wonder because these enclosed organs obstruct access for winddispersed pollen
to the ovules s molecular phylogenetic reconstructions show the shift from anemophily to
entomophily in oraceae is nearly associated with the transition from globose to discshaped
inorescences and to syconia ertain
functional similarity of seedbearing capsules in the peltasperms with such structures as syconia
of Ficus and the enclosed perigone of Aspidistra locii suggest that evolutionary transformations
of female reproductive organs in these extinct gymnosperms were attended with their shift to
insect pollination lthough the microstrobiles of peltasperms does not show evident traits
of entomophily this condition the male generative organs which indicate wind pollination
occur in some extant plant taxa pollinated by insects eg in Brosimum a member of oraceae
his hypothesis is in good agreement with the evidences of close interactions between peltasperms
and insects eltasperm pollen were found in guts of some ermian insects
and the microsporangia with the wall perforations and bites made by insects
occur on the microsporangium walls in some peltasperms he seed
bearing capsules damaged by insects are also known in some ngaropeltidaceae
a b
oreover the gradual shift from anemophily to entomophily in peltasperms can be indirectly
conrmed by correlation between the structure of their female reproductive organs and the types of
pollen grains he early ermian peltasperms Autunia conferta ternberg erp and Peltaspermum
retensorium alessky augolnykh erp had open seedbearing organs in combination with
distinct air sacs on pollen the typical feature of anemophilous plants
n many younger taxa belonging to the family ngaropeltidaceae however the
female reproductive organs became partially or almost completely enclosed and the air sacs on
their pollen grains were less distinct quasisaccate or protomonosaccate pollen of the Vesicaspora
type sensu inally the air sacs are completely lost pollen of the Vittatina type
sensu in some late ermian members of the family eltaspermaceae
evertheless our suggestion on the insect pollination of peltasperms remains hypothetic and it
should be tested by new evidences ndeed the peltasperms show one of the most ancient events
© Landesmuseum für Kärnten; download www.landesmuseum.ktn.gv.at/wulfenia; www.biologiezentrum.at
Permoxylocarpus trojanus: an ancient case of entomophily in gymnosperms
of coevolution between higher plants and insects b and their interactions
were apparently even more complex than it is commonly considered e can conclude that
the trend of seedbearing organ specialization from open discs to enclosed capsules displays the
last phases of pteridosperm sl evolution during alaeozoicesozoic transition he female
reproductive organs of the pteridosperms became more and more specialized which lead to
the formation of closed ovuliferous carpellike structures similar to those of the most primitive
preangiospermous plants
cknowledgements
e express our sincere gratitude to rof ofronitsky harov omankov
simbal ukhonto and urago for their great help in obtaining materials for
our study
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© Landesmuseum für Kärnten; download www.landesmuseum.ktn.gv.at/wulfenia; www.biologiezentrum.at
Permoxylocarpus trojanus: an ancient case of entomophily in gymnosperms
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ddresses of the authors
erge augolnykh
eological nstitute ussian cademy of ciences
yzhevsky per
oscow
ussia
mail naugolnykh@ramblerru
naugolnykh@listru
lexei skolski
omarov otanical nstitute
otanical useum
rof opov str
t etersburg
ussia
mail aoskolski@gmailcom
© Landesmuseum für Kärnten; download www.landesmuseum.ktn.gv.at/wulfenia; www.biologiezentrum.at
© Landesmuseum für Kärnten; download www.landesmuseum.ktn.gv.at/wulfenia; www.biologiezentrum.at
