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.     

Cyathea cranhamii sp. nov. (Cyatheaceae), anatomically preserved tree fern sori from the Lower Cretaceous of Vancouver Island, British Columbia

Permineralized cyatheaceous sori occur among remains of conifers, fungi, and other plants in newly discovered calcareous concretions from Early Cretaceous (Barremian) marine sediments of Vancouver Island, British Columbia, Canada. Sori are superficially attached in two rows to narrow pinnules and display a globose sphaeropteroid indusium. Annulate sporangia with multicellular stalks diverge from a basal, vascularized receptacle. The nearly vertical uniseriate annulus is not interrupted by the stalk. The sporangia bear 64 trilete spores with perispore sculpturing that ranges from irregular granulate/echinate to prominent rodlets. These specimens, described as Cyathea cranhamii sp. nov., are the first anatomically preserved tree fern sori from the fossil record. They represent the most ancient evidence for fertile structures of the Cyatheaceae and demonstrate that essentially modern species of cyatheaceous tree ferns had evolved by the Early Cretaceous.     

A review of selected triassic to Early Cretaceous ferns

After becoming nearly extinct during the Permian, the ferns began a slow recovery during the Triassic as the climate of the earth moderated. As a result, a considerable number and variety were present and widely distributed during the Jurassic and Early Cretaceous. However, with the rapid expansion of the angiosperms during the Late Cretaceous, the ferns once again became reduced in variety and greatly restricted in distribution. Some of the Mesozoic ferns are rather primitive and obviously are closely related descendants of Paleozoic taxa. Such ferns are assigned mostly to the Marattiaceae, Guaireaceae, Osmundaceae, and Gleicheniaceae. The majority of the Mesozoic ferns, however, are distinctive and appear to have originated during that era. These fossil ferns generally fit into modern orders and families such as the Matoniaceae or the Dipteridaceae. In some cases, it is difficult to clearly distinguish some of the Mesozoic ferns from living genera. A portion was presented as an invited paper to the symposium: Evolution of pteridophytes and gymnosperms at the XV International Botanical Congress, Yokohama, Japan (1993).     

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.     

Fossil evidence of eupolypod ferns in the mid-Cretaceous of Myanmar

Divergence time estimates based on DNA sequence variation of extant species suggest that eupolypod ferns were diverse already in the Cretaceous; however, fossil evidence to support this assumption remains exceedingly rare. Holttumopteris burmensis gen. et sp. nov., a fertile fern foliage fragment preserved in a piece of Albian–Cenomanian Burmese amber from Myanmar, is characterized by divided fertile leaves with catadromous, free lateral veins. Sporangia possess a vertical annulus interrupted by the sporangium stalk and contain monolete spores with a lophate perine. The sporangia occur clustered in discrete sori overarched by a laterally attached, reniform indusium. Reconstruction of ancestral character states suggests affinities of Holttumopteris to the Thelypteridaceae; however, several taxonomically relevant characters are not preserved. This discovery is nevertheless important because H. burmensis represents the first unequivocal fossil of a representative of the eupolypods from the middle Cretaceous.     

Diversity and significance of late cretaceous permineralized plant remains from Hokkaido,Japan

Middle to Late Cretaceous permineralized plants hitherto described from Hokkaido, Japan are summarized. The fossil flora comprises fungi, ferns, gymnosperms and angiosperms. Many modern fern families have been recognized including Anemiaceae, Cyatheaceae, Dennstaedtiaceae, Gleicheniaceae Loxsomaceae, Lygodiaceae and Matoniaceae. Gymnosperms are most abundant in the flora. Some recently-found materials are tentatively introduced with brief comments emphasizing their morphological and taxonomical significance. A bisporangiate flower ofCycadeoidella japonica Ogura shows fine internal anatomy and provides evidence that the cycadeoidalean ovule was a cupulate, unitegmic structure. Vascular tracheids in the synangial wall support the evolution of cycadeoidalean synangia from Paleozoic seed-fern synangia. A new gymnosperm female fructification has a thick envelope comparable to an angiosperm carpel around a large seed. The angiosperms contain various morphologies that require further extensive study.     

中国中生代的马通蕨科植物:化石记录、多样性与时空分布特征*

现生真蕨目马通蕨科(Matoniaceae)植物仅存Matonia和Phanerosorus两属, 集中分布于马来西亚、印度尼西亚等热带地区。马通蕨科植物在中生代时期全球广布, 且主要分布于热带、亚热带地区, 有近9个属, 被作为热带、亚热带气候的标志性植物化石之一。本文梳理总结了中国中生代的马通蕨科化石记录并分析其多样性特征, 共计有2属16种, 包括异脉蕨属(Phlebopteris) 15种和准马通蕨属(Matonidium) 1种。对其化石记录和地质地理分布分析表明, 晚三叠世时期, 异脉蕨属植物广泛分布于热带—亚热带湿热气候区, 包括华南一带的四川、湖北、 云南、西藏、福建等地; 早侏罗世时期, 其分布逐渐向北方扩展, 在南、北方植物区系界线附近均有发现; 中侏罗世局限于湖北、青海等地; 早白垩世时仅在黑龙江和西藏少量发现。准马通蕨属仅在黑龙江地区的早白垩世地层中发现。整体上, 马通蕨科在中国中生代的分布范围变迁与气候带范围变化相吻合。     

The coal-forming plants of rhabdopissites in the Lipovtsy coal field (Lower Cretaceous of Southern Primorye)

The plants that produced unique resinous coals (rhabdopissites) of the Lipovtsy coal field are revealed. They belong mainly to the group Miroviaceae (Oswaldheeria). Pseudotorellia (Ginkgoales) played an important role in the formation of humic coals of this Early Cretaceous coal field. The coal-forming plants comprise also cyatheaceous and gleicheniaceous ferns. A leafy shoot of Pseudotorellia has been found in this locality for the first time. It is assigned to the new species Pseudotorellia krassilovii Bugdaeva sp. nov.     

Singular taphonomic record of a wildfire event from middle Albian deposits of Escucha Formation in northeastern of Spain

A fossil plant assemblage composed of a great amount of macro, meso and microscopical charcoalifed remains occur in a single layer in Albian deposits of the Escucha Formation in northeastern Spain. This assemblage consists fundamentally of fragments of pinnae and pinnules corresponding to the Matoniaceous ferns Weichselia reticulata and Phlebopteris dunkeri also with some gymnosperm wood remains. The features of both the fossil plants and the deposit itself indicate accurate paleoenvironmental conditions related to the action of wildfires over the vegetation growing in freshwater swamp plains during Albian in Southwestern Eurasia.     

Ellesmeris sphenopteroides,gen. ET SP. NOV., a new zygopterid fern from the Upper Devonian (Frasnian) of Ellesmere,N.W.T., Arctic Canada

A new fern-like fossil plant is described from the lower Upper Devonian of southern Ellesmere Island, Canadian Arctic Archipelago. The plant occurs in an Archaeopteris-dominated flora preserved in the Nordstrand Point Formation (Mid-Late Frasnian) near Bird Fiord. The plant has a pinnate vegetative system with three branch orders and laminate sphenopteroid pinnules. Primary pinnae usually diverge from the main axis in distichous pairs (quadriseriate), but can depart singly (biseriate). Each primary pinna bears a basal catadromic aphlebia. Anatomically, the plant exhibits a mesarch, bipolar protostele that is ribbon- to clepsydropsoid-shaped in the main axis. Primary pinna traces are also initially bipolar and crescent-shaped, but may become four-ribbed before dividing into a pair of bipolar traces. The morphology and anatomy of this plant are nongymnospermous and are most similar to Zygopteridales (particularly Rhacophytaceae and Zygopteridaceae). The Frasnian age of Ellesmeris shows that laminated foliage had evolved in some zygopterid ferns much earlier than previously recognized. The Sphenopteris-like pinnules of Ellesmeris indicate the need for caution when attributing such a convergent foliar design to other plant groups, such as the Devonian gymnosperms.     

Paralygodium meckertii sp. nov. (Schizaeaceae) from the Upper Cretaceous (Coniacian) of Vancouver Island, British Columbia, Canada

More than 50 specimens of permineralized fertile pinnules with abaxially borne sporangia have been discovered in calcareous marine nodules from the Upper Cretaceous (Coniacian) Comox Formation from the Eden Main localities on Vancouver Island, British Columbia, Canada. Isolated pinnules 1.6–3.0 mm wide × 1.6–2.8 mm long are lobed and abaxially enrolled to form irregular globose structures. Pyriform sporangia 216–300 μm wide × 360–468 μm long occur in two rows on the abaxial surface of pinnule lobes. Sporangia have an apical annulus of 15–18 cells. Spores are tetrahedral and trilete, 33–42 μm in diameter, with straight to concave interradial sides, laesurae extending nearly to the equator, and a psilate exine. Spores are assignable to the sporae dispersae genus Deltoidospora. Fertile pinnules are compared to fossils of Anemia poolensis and two previously described species of Paralygodium, and show closest similarities to P. vancouverensis from the Eocene of British Columbia. The Cretaceous Eden Main specimens differ in number of pinnule lobes and their morphology and are described as a new taxon: P. meckertii sp. nov. This discovery extends the Cretaceous geographic range of Paralygodium from Japan to North America and adds to our knowledge of the diversity of extinct schizaeaceous ferns.     

New Middle Jurassic record of fern genus Eboracia (Dicksoniaceae,Filicales) and its spatio-temporal distribution in China

As an important component of the Mesozoic flora, the extinct fern genus Eboracia Thomas (Dicksoniaceae, Filicales) is widely reported in China with diverse fossil records. New material of Eboracia lobifolia, represented by a nearly intactly preserved fossil frond, is described herein from the Middle Jurassic Haifanggou Formation in Beipiao of western Liaoning, Northeast China. The frond is lanceolate in gross outline, at least 38.0 cm long, and can be divided into the basal sterile part, the upper fertile part and a transitional part in between. The new discovery confirms for the first time that the frond of E. lobifolia is hemidimorphic rather than holodimorphic. Many in-situ spores were detached from the sori of the fertile pinnae, which are characterized by rounded-tetrahedral shape, smooth surface, distinct border, and a wide, long triradiate crack almost reaching the equator. Comparing with dispersed spores in the same horizon, these in-situ spores seem to be most similar to Cyathidites minor Couper in morphology. A spatio-temporal analysis of Eboracia in China shows that Eboracia with totally four species occurred in a time interval ranging from the Late Triassic to the Early Cretaceous in China, and mostly flourished in the Middle Jurassic; the genus was widely distributed in both the Northern and Southern Phytofloristic Provinces of China, particularly in southern China during the Late Triassic to Early Jurassic, while more abundant and diverse in northern China during the Middle Jurassic to Early Cretaceous.     

Anatomically preserved Woodwardia virginica (Blechnaceae) and a new filicalean fern from the middle Miocene Yakima Canyon flora of central Washington, USA

Anatomically preserved Woodwardia virginica (Blechnaceae) and a newly recognized onocleoid fern are described from the middle Miocene Yakima Canyon flora of central Washington State, USA. Identification of the W. virginica fossils is based on a combination of vegetative pinnules, rhizome and stipe anatomy, and fertile pinnules with indusiate sori and sporangia like those of extant W. virginica. Fronds are isomorphic. Vegetative pinnae are elongated and pinnatifid, with a secondary vein paralleling the midvein. Secondary veins of the pinnule lobe anastomose to form primary areoles and are either simple or dichotomize toward the margin. Rhizomes have a simple dictyostele with 3-5 cauline vascular bundles and often a sclerotic hypodermis. Leaf traces contain two large adaxial vascular bundles that occur laterally and adaxially, flanking an arc of 4-6 smaller bundles. Fertile pinnules have linear sori that are somewhat embedded in the laminae and are enclosed by a thin indusium. Leptosporangia display a vertical annulus and an elongated stalk. A second fern, Wessiea yakimaensis gen. et sp. nov., is represented by anatomically preserved branching rhizomes and attached frond bases that conform to the Onoclea-type pattern of rhizome and frond-base vasculature. Rhizomes have a simple dictyostele of 4-5 cauline meristeles. Leaf divergence is helical, with paired hippocampiform rachial traces. These two ferns occur in the same matrix with specimens of Osmunda wehrii. They demonstrate that filicalean fern assemblages similar to those of extant temperate floras were well established in western North America by the middle Miocene and further emphasize the exceptional species longevity of some homosporous pteridophytes.     

First fossil record of the genus Nageia (Podocarpaceae) in south China and its phytogeographic implications

The first fossil record of the genus Nageia Gaertner from the Tertiary of southern China is reported. Nageia hainanensis sp. nov. is described from the Eocene Changchang Formation of Hainan Island. The extant species of Nageia are widely distributed in southeastern Asia, from northeastern India to southeastern China and southern Japan, and southward through Indochina to the Malay Archipelago and New Guinea. The fossil evidence of Nageia has been discovered from the Lower Cretaceous of the Far East of Russia and Japan. The distribution of both modern and fossil species of Nageia indicates that this genus could have originated in the northeastern part of Asia in the Early Cretaceous and spread to south China at least in the Eocene. Discovery of Nageia fossil record in south China provides important evidence for the geological and phytogeographic history of the genus.     

How diverse were ferns in the Baltic amber forest?

Diverse temperate forest types and a high atmospheric humidity have recently been suggested for the Eocene source area of Baltic amber. However, ferns are astonishingly rare as inclusions in this amber, which is in contrast to other seed‐free land plants, fungi, and lichens. Moreover, the identities of some of the few described putative fern taxa are dubious, and some fossils were even assigned to the Paleozoic seed fern genera Alethopteris, Pecopteris and to the form genus Sphenopteris containing Paleozoic and Mesozoic fern‐like leaf fossils. Here, we review previously described fern inclusions from Baltic amber and identify further fern‐like leaf inclusions as belonging to the extant angiosperm genus Comptonia (sweet ferns, Myricaceae). We conclude that only one taxon, Matonia striata (Matoniaceae), can with confidence be identified as a Polypodiopsida representative. Although “Pecopteris” humboldtiana is so far only known as sterile foliage, its leaf morphology strongly suggests that also this taxon belongs to the Polypodiopsida rather than to any other tracheophyte lineage. We propose accommodating “Pecopteris” humboldtiana in the new genus Berendtiopteris. “Alethopteris” serrata and “Sphenopteris” phyllocladoides are not to be regarded as evidence of ferns from Baltic amber. Reinvestigation of the holotypes of these two taxa did not reveal to which tracheophyte lineages these fossils belong. We suggest that the scarcity of fern remains from Baltic amber may reflect both a relatively low fern diversity in the source area of the fossil resin, and an absence or rarity of epiphytic and climbing ferns as observed in modern temperate forest ecosystems.     

Deux fougères fertiles du Stéphanien du Massif Central (France)

Two fertile ferns from the Stephanian of the Massif Central (France) are described in detail. One has been found in the Blanzy basin and is assigned to Oligocarpia (Sphenopteris) leptophylla (Bunbury) nov. comb., according to the morphology of the sterile pinnae and of the fructifications. Its spores agree with the «sporae dispersae Granulatisporites parvus (Ibr.) Potonié and Kremp. The other fern has been collected in the St Etienne basin and is assigned to Senftenbergia plumosa (Artis) Radforth var. ligerensis nov. var. Its spores agree with the morphographic genus Raistrickia (S., W. and B.) Potonié and Kremp but do not look like any described species; therefore the new species name R. polymorpha nov. sp. has been given to that spore.     

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.     

THE CRETACEOUS PTERIDOSPERMS RUFLORINIA AND KTALENIA AND IMPLICATIONS ON CUPULE AND CARPEL EVOLUTION

The vegetative (Ruflorinia sierra) and fertile (Ktalenia circularis) organs of an Early Cretaceous pteridosperm collected from Santa Cruz Province in Argentina are described. The sterile leaf is at least tripinnate and bears decurrent secondary pinnae with obliquely attached, sharply pointed pinnules. The fertile member arises from the base of the vegetative rachis and bears two types of appendages, cupules and bracts. Bracts are attached to the main axis near cupules and are present in clusters of up to six. Cupules are sessile, spherical, and arranged in opposite or subopposite pairs along the axis. A small lip is present on one surface of the cupule. The number of seeds per cupule may be one or two, with each characterized by a distal nucellar beak and circular, chalazal scar. Cuticular anatomy, including the fine structure of the stomatal complex, is described for both vegetative and reproductive organs. The cupules of Ktalenia and other Mesozoic seed plants are compared, and a discussion presented regarding the possible function of the cupule.     

Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences

The phylogenetic structure of ferns (= monilophytes) is explored here, with a special focus on the early divergences among leptosporangiate lineages. Despite considerable progress in our understanding of fern relationships, a rigorous and comprehensive analysis of the early leptosporangiate divergences was lacking. Therefore, a data set was designed here to include critical taxa that were not included in earlier studies. More than 5000 bp from the plastid (rbcL, atpB, rps4) and the nuclear (18S rDNA) genomes were sequenced for 62 taxa. Phylogenetic analyses of these data (1) confirm that Osmundaceae are sister to the rest of the leptosporangiates, (2) resolve a diverse set of ferns formerly thought to be a subsequent grade as possibly monophyletic (((Dipteridaceae, Matoniaceae), Gleicheniaceae), Hymenophyllaceae), and (3) place schizaeoid ferns as sister to a large clade of "core leptosporangiates" that includes heterosporous ferns, tree ferns, and polypods. Divergence time estimates for ferns are reported from penalized likelihood analyses of our molecular data, with constraints from a reassessment of the fossil record.