Morphology and growth habit of Dicksonites pluckenetii from the Upper Carboniferous of Graissessac (France)

Morphological variability of the vegetative fronds of Dicksonites pluckenetii (SCHLOTHEIM ex STERNBERG) STERZEL is described from new material collected in the Stephanian Basin of Graissessac (Southern Massif Central France). Complete bipartite fronds sometimes still attached to their parent stem allow us to reconstruct the plant as a scrambler/climber. This interpretation is supported by the relatively small slender stems with long internodes and densely covered by short spines, which are also present along the entire length of petioles. A fertile pinna with slightly reduced pinnules bearing up to six radially symmetrical synangia is interpreted as a portion of male frond of the same species; its organization confirms the relationships of Dicksonites with the Callistophytales as was already suggested by features of the female fronds.     

The current status of medullosan seed ferns

The compression and permineralized fossil record of presumed medullosan plants is reviewed and evaluated. Recent studies of stelar architecture suggest that stems are monostelic rather than polystelic as commonly believed. Imparipinnate fronds are mostly of the Neuralthopteris/Alethopteris type with Myeloxylon anatomy; the affinity between medullosan stems and neuropterid fronds is uncertain. It is suggested that the reproductive organs of the family should be restricted to those pollen organs believed to have produced Monoletes pollen and to ovules, for the most part, of the Trigonocarpus/Pachytesta type. It is doubtful that reproductive organs replaced lateral pinnules; it is more likely that they terminated pinnae or were borne abaxially along the pinna rachis. In some cases they may have been borne directly on penultimate axes or were, as in Codonotheca and Goldenbergia, borne on fertile fronds or fertile portions of vegetative fronds. Recent research involving pollen organs is emphasized and a phylogeny of Monoletespollen organs is proposed. It is suggested that the fossil record of medullosans is best interpreted from a punctuational perspective.     

Pollen Organ Telangiopsis sp. of Late Devonian Seed Plant and Associated Vegetative Frond

Pollen organ Telangiopsis sp., associated with but not attached to vegetative fronds, has been collected from the Upper Devonian (Famennian) Wutong Formation, Dongzhi County, Anhui Province, China. Fertile axes with terminal pollen organs are dichotomous for 2–4 times and may be proximally attached by fragmentary pinnules. Pollen organs are synangiate and borne on the top of a short stalk. Synangia are radial in symmetry and each consists of 4–8 elongate microsporangia fused at base. Microsporangia have a longitudinal dehiscence line and show a tapered apex. The associated stem is spiny and bears a vegetative frond which bifurcates once at the basalmost part. Frond rachises possess one order of pinna arranged alternately. Pinnules are borne alternately, planate, highly dissected, and equally dichotomous for 2–3 times. Comparisons among Late Devonian seed plants recognize several branching patterns in the fertile fronds/axes bearing terminal pollen organs. Telangiopsis sp. reinforces that the Late Devonian pollen organs are synangiate usually with basally fused microsporangia. It is suggested that the evolutionary divergence of radial and bilateral symmetries of pollen organs may have occurred in the Famennian, when the earliest seed plants evolved planate and sometimes laminate pinnules.     

The callistophytales (Pteridospermopsida): Reproductively sophisticated paleozoic gymnosperms

The Callistophytales nov. order is proposed for a small family of Pennsylvanian and possibly Permian pteridosperms. Plants conform to two species of Callistophyton and their reproductive organs. The small, eustelic sporophytes have a scrambling, shrub-like habit, profuse branching, adventitious roots and pinnately compound fern-like leaves with sphenopterid pinnules. Reproductive organs are borne on the abaxial surface of the pinnules, and consist of cardiocarpalean ovules and synangiate pollen organs with saccate pollen. Gametophyte development and reproductive biology conform closely to those of extant gymnosperms in the Coniferales. Anatomically preserved reproductive organs of the major groups of Paleozoic gymnosperms are compared, and a closer than currently recognized relationship of Paleozoic seed plants is proposed.     

Korallipteris,a new genus for Mesozoic Gleichenia-like fern fronds

A new morphogenus, Korallipteris, is proposed to accommodate fertile or sterile fern fronds possessing essentially Gleichenia-like morphology (e.g., pinnate fronds with small pecopteroid pinnules) but lacking diagnostic features that allow their classification at fern family-level. The aim is thus, to solve an old nomenclatural problem that involves the use of illegitimate genera, such as Gleichenites Goeppert and Microphyllopteris Arber. Several species of Gleichenites described from the Mesozoic of Argentina and Chile, and Microphyllopteris unisora Cantrill and Nagalingum from Antarctica are here reassigned to Korallipteris gen. nov. The proposal of Nagalingum and Cantrill to preserve Gleicheniaceaephyllum Crabtree emend. Nagalingum and Cantrill for true Gleicheniaceous ferns, diagnosed by having an arrested laminar bud, is here accepted.     

Early Cretaceous Gleicheniaceae and Matoniaceae (Gleicheniales) from Alexander Island,Antarctica

The first Cretaceous representatives of Matoniaceae and Gleicheniaceae in the Antarctic are described from the Albian flora of Alexander Island. Two new species are recognized, one is assigned to Matonia, and another to the emended gleicheniaceous form-genus Gleicheniaceaephyllum. The taxonomy of fossil genera assigned to Gleicheniaceae is problematic, and as an alternative to the invalid genus Gleichenites, we propose that Microphyllopteris be utilised as a form-genus for species that have either an unknown branching pattern or pseudo-dichotomous branching without resting buds; Microphyllopteris is reserved for ferns that cannot be ascribed conclusively to Gleicheniaceae. Alternatively, Gleicheniaceaephyllum should be used for gleicheniaceous material that has a resting bud amongst two to four primary branches. Gleicheniaceaephyllum acutum sp. nov. is preserved as sterile and fertile fronds, pinnae and pinnules. The other new species, Matonia jeffersonii sp. nov., occurs as sterile and fertile pinnae and pinnules, and is aligned to Matoniaceae based on the presence of circular-oval sori with peltate indusia. The two taxa are inferred to have either a pedate or scrambling habit. These morphologies are relatively common at Alexander Island, compared to other high latitude fossil floras, and are interpreted as an adaptation to the high disturbance, polar conditions and unusual photoperiod experienced at Alexander Island during the Early Cretaceous. The theory that angiosperms caused the demise of ferns is discussed in relation to the Matoniaceae and Gleicheniaceae, and there does not appear to be association between angiosperm expansion and the decline of these two fern families.     

A marattialean fern,Scolecopteris libera n. sp., from the Asselian (Permian) of Inner Mongolia,China

Scolecopteris libera n. sp. is established on partial three-dimensionally preserved materials containing both sterile and fertile fronds collected from the volcanic tuff at the top of the Taiyuan Formation at the Wuda Coalfield, Inner Mongolia, China. It is characterized by tripinnate fronds with rachises over 200 mm in diameter; and long lanceolate or falciform pinnules with thick veins. Pinnules are of the pecopterid-type, with those at the base of the ultimate pinnae usually divided into small lobes. Eight to twelve circular synangia are arranged in two rows along the midvein. Synangia are borne on a short pedicel attached to the middle of lateral veins. A synangium is composed of 7–10 fusiform exannulate sporangia with pointed apices. The sporangia are free among each other beyond the base of synangia. In situ microspores of the Cyclogranisporites leopoldii type differ from all other in situ Paleozoic marattialean spores.     

Fertile pinnules of Danaeites rigida Gu and Zhi (Marattiales) from the Upper Permian of south China

Recently collected specimens of Danaeites rigida Gu and Zhi from the Upper Permian of south China have been subjected to detailed morphological investigations in order to reveal features of their fertile pinnules. The specimens are preserved as compression/impressions and possess pecopteroid-type pinnules with a single row of synangia on either side of the pinnule midvien. Individual synangia are bilaterally symmetrical, sessile, and their bases are embedded in the tissues of the pinnules. Synangia possess 18–24 sporangia that are laterally fused to one another throughout their entire length. Sporangial dehiscence is through a longitudinal slit and sporangia contain trilete spores with a granular ornamentation, referable to the dispersed spore Cyclogranisporites. This combination of characters is unique in specimens preserved by compression/impression although they are similar to those known in permineralized marattialean fertile fronds. However, anatomical details salient to the identification of these permineralized taxa are not identifiable in impression/compression fossils such as Danaeites. The taxonomic and phylogenetic implications of these findings are considered in detail and we conclude that Danaeites has closer links with Marattia than with Danaea.     

Reconstructing the Pennsylvanian-Age Filicalean Fern Botryopteris tridentata (Felix) Scott

Numerous anatomically preserved fragments of the Middle Pennsylvanian age filicalean fern, Botryopteris tridentata, occur in coal balls collected at the Pittsburgh and Midway Coal Company mine near Baxter Springs, Kansas. Included are the first fertile specimens of the species, evidence of complete vegetative frond architecture, and fronds that are specialized for vegetative propagation. Rhizomes are erect and unbranched, have helical phyllotaxis and short internodes, and typically display an ectophloic solenostele. Fronds are tripinnately compound with lobed pinnules that have open, dichotomous venation. Fertile pinnae or individual pinnules are interspersed among vegetative frond segments and produce sori of annulate sporangia beneath veins on the abaxial pinnule surface. Fertile pinnule lobes are rolled toward the abaxial surface to enclose the sori. Sporangia have a horizontally elongated biseriate annulus located near the short broad stalk and produce tetrahedral-shaped trilete spores with coarse spines. Epiphyllous branches diverge from the stipe or rachis, and some fronds produce only branches. This fern is reconstructed as having short stems. Helically arranged fronds are either pinnately dissected with lobed vegetative pinnules and abaxially rolled fertile pinnules or are specialized for vegetative propagation. The latter functioned as the foliar equivalent of stolons. While some characters of the B. tridentata plant are similar to those of Botryopteris forensis, generitype of the Botryopteridaceae, others are more comparable to those of Psalixochlaena cylindrica, generitype of Psalixochlaenaceae, suggesting the need for reevaluation of systematic relationships among species of the Botryopteridaceae and Psalixochlaenaceae.     

The photosynthetic performance of sterile and fertile sporophytes in a natural population of the fern Dryopteris affinis

We studied the photosynthetic performance of sterile and fertile sporophytes in a natural population of the fern Dryopteris affinis growing within a riparian forest (Central Italy) using chlorophyll (Chl) a fluorescence transients, the OJIP phase, where O is for the minimum fluorescence, P is for the peak (the maximum), and J and I are inflections. The “vitality” of the samples was assessed by the maximum quantum yield of primary photochemistry obtained indirectly from the fluorescence data (F_v/F_m); in the same way, the so-called performance index (PI_ABS) was obtained from fluorescence data. The photosynthetic performance (inferred from PI_ABS) of D. affinis changed significantly with the seasonal development of the fronds. The highest photosynthetic performance was recorded in the summer, corresponding to the period of spore release. The photosynthetic performance decreased in the winter, down to the minimal values of senescent fronds reached at the end of the seasonal cycle (May-June). On the whole, during the seasonal development, sterile and fertile fronds had a similar photosynthetic behaviour, as inferred from fluorescence data. At the end of spore maturation and dispersal (September-October), the fertile fronds showed somewhat lower photosynthetic performance than the sterile fronds, as revealed by PI_ABS. Being a long-lived fern, confined to humid and undisturbed sites in the Mediterranean, D. affinis deserves to be further investigated as a potential indicator of ecological continuity in Mediterranean riparian forests.     

ARCHAEOPTERIS MACILENTA,ANATOMY AND MORPHOLOGY OF ITS FROND

The stem, rachides, and pinnae of Archaeopteris macilenta, formerly considered to be a fern of Devonian age, comprise a branch system in which the ultimate divisions heretofore referred to as pinnules are the leaves. The primary vascular system of the “frond” is a lobed siphonostele from which leaf traces arise in a spiral sequence. The anatomy of the “rachis” and of the “pinnae” is shown to be similar to that of the stem, Callixylon, which bore these “fronds.” Branching, epidermal pattern and stomates are described for the spirally arranged leaves (fertile pinnules). Attachment and dehiscence of sporangia as well as their stomates are reported. Archaeopteris is retained in the Class Progymnospermopsida which includes plants with gymnospermous anatomy and pteridophytic reproduction. It is suggested that Actinopodium, Svalbardia and Siderella are related closely to Archaeopteris and that this group of genera shows evolutionary stages in webbing of leaves and planation of branch systems. The opportunities for ontogenetic studies of the arborescent genus Archaeopteris are pointed out.     

Ashicaulis woolfei n. sp.: additional evidence for the antiquity of osmundaceous ferns from the Triassic of Antarctica

Numerous small fern trunks and dispersed osmundaceous frond fragments occur within a Middle Triassic silicified peat near Fremouw Peak in the Transantarctic Mountains of Antarctica. These specimens form the basis of a new species of osmundaceous ferns that further helps to characterize the early Mesozoic vegetation of high latitude Gondwana. Ashicaulis woolfei n. sp. consists of small, upright trunks with a persistent armor of frond bases, adventitious roots, and vegetative frond parts. In cross section the trunks are ～2.5 cm in diameter and include up to 45 frond bases. Stems range from 5 to 8 mm in diameter with a xylem cylinder of 8-9 xylem segments separated by leaf gaps. Phyllotaxy is variable, approaching 2/5 or 3/8, with 10-12 frond traces in the cortex. Stipes have parenchymatous, stipular wings that are usually devoid of sclerenchyma; fronds are pinnate with alternate-subopposite pinnatifid pinnules. Although the absence of fertile pinnules and sporangia precludes assigning the fossils to a living genus, this species demonstrates that ferns with stelar architecture and histology similar to Osmunda subgenus Osmundastrum (Osmundaceae) were present in the Southern Hemisphere by the mid-Triassic.     

Latisemenia longshania,gen. et sp. nov., a new Late Devonian seed plant from China

The earliest known ovules in the Late Devonian (Famennian) are borne terminally on fertile branches and are typically enclosed in a cupule. Among these ovules are some that have terete integumentary lobes with little or no fusion. Here, we report a new taxon, Latisemenia longshania, from the Famennian of South China, which bears cupulate ovules that are terminal as well as opposite on the fertile axis. Each ovule has four broad integumentary lobes, which are extensively fused to each other and also to the nucellus. The cupule is uniovulate, and the five flattened cupule segments of each terminal ovule are elongate cuneate and shorter than the ovule. Associated but not attached pinnules are laminate and Sphenopteris-like, with an entire or lobate margin. Latisemenia is the earliest known plant with ovules borne on the side of the fertile axis and may foreshadow the diverse ovule arrangements found among younger seed plant lineages that emerge in the Carboniferous. Following the telome theory, Latisemenia demonstrates derived features in both ovules and cupules, and the shape and fusion of integumentary lobes suggest effective pollination and protection to the nucellus. Along with other recent discoveries from China, Latisemenia extends the palaeogeographic range of the earliest seed plants.     

Phenology and photosynthetic activity in sterile and fertile sporophytes of Dryopteris filix-mas (L.) Schott

Summary This study describes the phenology of sporophytes of the fern Dryopteris filix-mas in relation to whole plant development. Sterile and fertile potted sporophytes were set out at an exposed site and the seasonal development of the fronds was measured from the commencement of unfolding, through the phase of increasing length, up to discoloration. The physiological activity of the fronds was determined by measuring photosynthetic gas exchange. The fronds of sterile sporophytes unfolded in April, about a week earlier than those of fertile plants, but the colour had already begun to turn in September and their life span was 1–2 months shorter. However, between mid-June and the end of August the sterile sporophytes put out several sets of new fronds: these overwintered without changing color and were still photosynthetically active in the following spring. All types of fronds were fully expanded 1–2 months from the beginning of unfolding and, with a natural supply of CO₂, had similar maximum net photosynthetic rates of 8–9 mol/m² · s. The decline in photosynthetic performance began before symptoms of senescence were visible and was due to decreased efficiency of the mesophyll. It is concluded that the phenology of D. filix-mas changes with transition from the sterile to the fertile phase. Whereas fertile sporophytes are genuinely summergreen, the sterile sporophytes with their summer fronds remain green throughout the winter and should therefore be termed semi-evergreen. The formation of overwintering summer shoots clearly extends the period of photosynthetic productivity of sterile sporophytes.     

Leaf evolution in early-diverging ferns: insights from a new fern-like plant from the Late Devonian of China

Background and Aims With the exception of angiosperms, the main euphyllophyte lineages (i.e. ferns sensu lato, progymnosperms and gymnosperms) had evolved laminate leaves by the Late Devonian. The evolution of laminate leaves, however, remains unclear for early-diverging ferns, largely represented by fern-like plants. This study presents a novel fern-like taxon with pinnules, which provides new insights into the early evolution of laminate leaves in early-diverging ferns.Methods Macrofossil specimens were collected from the Upper Devonian (Famennian) Wutong Formation of Anhui and Jiangsu Provinces, South China. A standard degagement technique was employed to uncover compressed plant portions within the rock matrix.Key Results A new fern-like taxon, Shougangia bella gen. et sp. nov., is described and represents an early-diverging fern with highly derived features. It has a partially creeping stem with adventitious roots only on one side, upright primary and secondary branches arranged in helices, tertiary branches borne alternately or (sub)oppositely, laminate and usually lobed leaves with divergent veins, and complex fertile organs terminating tertiary branches and possessing multiple divisions and numerous terminal sporangia.Conclusions Shougangia bella provides unequivocal fossil evidence for laminate leaves in early-diverging ferns. It suggests that fern-like plants, along with other euphyllophyte lineages, had independently evolved megaphylls by the Late Devonian, possibly in response to a significant decline in atmospheric CO₂ concentration. Among fern-like plants, planate ultimate appendages are homologous with laminate pinnules, and in the evolution of megaphylls, fertile organs tend to become complex.     

A synopsis of the Neotropical species of <Emphasis Type="Italic">Cyathea</Emphasis> (Cyatheaceae; Polypodiopsida) with bipinnate fronds

A synopsis of all Neotropical species of Cyathea with bipinnate fronds is presented. This artificial group includes 27 species with entire or very shallowly lobed pinnules. The provided key includes together, for the first time, species that had been previously treated in three different genera: Sphaeropteris, Trichipteris, and Cyathea. Full synonymy and geographic distribution is provided for each species. A second-step lectotype is designated for Cyathea marginalis.     

Ontogeny and ecology of the filicalean fern Oligocarpia gothanii (Gleicheniaceae) from the Middle Permian of China

Reinvestigation of Oligocarpia gothanii Halle (Gleicheniaceae) from the Permian Period of China has uncovered a rare demonstration of ontogenetic succession in numerous intact plants and has led to emendation of the diagnosis and typification of the species. Reconstruction of the fern's vegetative life cycle shows a small rosette-shaped juvenile plant with immature fronds of pinnae undifferentiated into pinnules. The second stage is a series of leafy shoots increasing in size and complexity and bearing fronds essentially comparable with those of the adult plants. Mature plants have sphenopteroid-type fronds differentiated into vegetative and fertile fronds. Close association of plants on the same bedding plane and the presence of a connecting root network between shoots, show that juvenile plants spread by vegetative propagation using underground stolons. Sedimentological information and co-occurrence with the bryophyte Thallites hallei Lundblad suggests that O. gothanii occupied a wetland habitat and was preserved in an obrution event consistent with flooding of marginal areas in a fluvio-deltaic setting. Although recent studies on Oligocarpia have focused exclusively on reproductive features, new ontogenetic information suggests that plants growing in rapidly changing environments may have been reliant on vegetative propagation and would not have needed to reach sexual maturity for successful reproduction.     

Variation in gametophyte dominance in populations of Chondrus verrucosus (Gigartinaceae,Rhodophyta)

We describe the abundance, including spatial and temporal variability, of phases of the isomorphic Chondrus verrucosus Mikami from Japan. Chondrus verrucosus occurred in a dense (～90% cover) and temporally stable bed on a small, isolated rocky outcrop (Oyakoiwa) in Shizuoka Prefecture. Small vegetative fronds were always much more abundant than large vegetative and fertile fronds over the spring to late summer periods in 1999 and 2000. Over the same period, fertile carposporophytic fronds were generally more abundant than fertile tetrasporophytic fronds, and fertile male fronds appeared infrequently at low densities. Using the resorcinol‐acetal test, we determined the proportion of gametophytes and tetrasporophytes in three populations of C. verrucosus: Oyakoiwa and Noroshi (Shizuoka) in the summers of 1999 and 2000 and Kamehana Point (Miyagi) in autumn 2000. All populations had a significantly higher proportion of gametophytes than tetrasporophytes in both years, although gametophytic proportions were lower at Noroshi (～70%) than at Oyakoiwa (～80%) and Kamehana Point (～97%). However, examination of all isolated individuals sampled on Noroshi showed equal proportions of each phase in 1999, but gametophyte dominance (74%) in 2000. Differences in dispersal and spore production between phases are discussed as mechanisms potentially contributing to variation in gametophyte dominance.     

The demography of fronds of Chondrus crispus Stackhouse

A demographic analysis was done of an intertidal stand of Chondrus crispus Stackhouse at Pubnico Point, Nova Scotia. The distribution of gametophytic and tetrasporophytic fronds was assessed. Fronds were placed in one of four size-classes. Size-specific measurements for fronds are presented. Gametophytic fronds dominated the stand. Fronds in Size-Class 2 exhibited high survivorship. Survivorship fell sharply when fronds entered Size-Class 3 and Size-Class 4. Fronds became fertile and highly branched when in the third and fourth size-classes. It is suggested that these two factors contribute to the higher mortality rates of such individuals.Monthly spore release rates·m^?2 for gametophytes and tetrasporophytes were measured. The annual release rate of carpospores exceeded the annual release rate of tetraspores by a ratio of 4.7:1. Rates did not fall below 10⁶ spores released·m^?2·month^?1 for either phase.     

中国东北古新世乌云植物群: Woodwardia bureiensis, Dryopteris sp.和 Osmunda sachalinensis

在黑龙江省嘉荫县乌云镇古新世乌云植物群中新发现的Woodwardia bureiensis, Dryopteris sp. 和 Osmunda sachalinensis在中国也是新记录。Woodwardia bureiensis Fedotov与其他种的主要区别是小羽片排列紧密且边缘彼此靠合。Osmunda sachalinensis小羽片基部宽楔, 而以前乌云植物群所报道的Osmunda greenlandica小羽片基部平展。