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Premise: An anatomically preserved fossil fern sporeling has been discovered in a Lower Cretaceous marine concretion from Vancouver Island, British Columbia, Canada, providing an opportunity to characterize rhizome growth from an extinct species., Methods: The specimen was studied from serial transverse sections prepared by the cellulose acetate peel technique., Results: The rhizome ranges from ~0.7 to 1.1 mm in diameter, has a sclerenchymatous pith, a stele that attains a dictyoxylic architecture, and sclerenchymatous outer cortex, features that are characteristic of osmundaceous rhizomes. Cauline xylem forms a medullated protostele or solenostele at some levels, but is dissected into discrete xylem bundles at others. Fronds diverge in a helical phyllotaxis, range up to 1.1 mm in greatest dimension, and have a C-shaped trace and outer cortex of sclerotic cells. Inside the sclerenchyma of the petioles are two lateral sclerotic strands and a sclerotic bundle adaxial to the trace. Together, these characters reveal the occurrence of a new species, Todea minutacaulis and provide evidence for developmental changes that occur in the rhizome as the sporeling increased in size., Conclusions: Small size of the specimen, medullated protostelic-dictyoxylic solenostelar vascular architecture, and incompletely sclerified cells apically reveal that osmundaceous sporeling development has remained constant since at least the Early Cretaceous. Together with Todea tidwellii and Osmunda vancouverensis that also are present in the Apple Bay flora, this sporeling documents the occurrence of a multispecies assemblage of osmundaceous ferns and demonstrates that the genus Todea was diversifying rapidly by the Early Cretaceous., (© 2023 Botanical Society of America.)

Premise of the Study: Recent clarification of the distribution of Marattiales through time provides the impetus for "total evidence" phylogenetic analyses of a major fern clade with a rich fossil record. These analyses serve as empirical tests for results from systematic analyses of living species and also of the belief that relationships among living species accurately reflect the overall pattern of phylogeny for clades with an extensive fossil record and a large percentage of extinction., Methods: Species of living and fossil Marattiaceae are analyzed employing a "total evidence approach" via maximum parsimony. Analyses were conducted using TNT implemented through WinClada., Key Results: Systematic analyses of living species and of living + extinct species provide roughly concordant topologies for living taxa. However, living species of Marattiales are only one component of a much larger clade with two major subclades. One consists of Psaroniaceae and extends through time to at least the Early Cretaceous. The other consists of Marattiaceae and includes all living species. Various analyses support the generic-level clades of living species from earlier analyses, but the arrangement of such clades varies from analysis to analysis., Conclusions: Marattiales is a monophyletic group that is extremely common in late Paleozoic and early Mesozoic deposits, with a stem group Psaroniaceae and a crown group Marattiaceae. Because Marattiaceae represents only a small component of overall marattialean diversity, living species alone neither account for evolutionary changes within the clade over time, nor accurately reflect the overall pattern of marattialean fern phylogeny., (© 2018 Botanical Society of America.)

An anatomically preserved fossil fern rhizome with diverging stipe bases and root traces is described from the Lower Cretaceous (Valanginian to Hauterivian) Apple Bay locality, Vancouver Island, British Columbia, Canada. The specimen is assignable to Solenostelopteris Kershaw, a morphogenus with six previously described species. The Apple Bay fossil is 1.3-1.6 mm in diameter, with parenchymatous pith and cortex, and is described as S. skogiae sp. nov. The xylem of the solenostele is exarch and one to six cells thick. Successive stipes diverge from only one side of the rhizome, implying a dorsi-ventral symmetry and prostrate habit. No trichomes or scales are produced. Diarch root traces emerge from all sides of the rhizome, some associated with leaf trace divergence. The pith and cortex are made up of uniform, thin-walled cells. The Apple Bay rhizome is most similar to S. nipanica Vishnu-Mittre from the Lower Cretaceous Nipania Flora, India, but differs in size and in distinctive tissue zonation in the cortex. This new species is the youngest record of the genus Solenostelopteris in North America, and it emphasizes that both new specimens of fossils and more complete descriptions of living ferns are needed to help clarify our concepts of Mesozoic ferns.

Anatomically preserved fossil gleicheniaceous fern remains in carbonate marine concretions from Vancouver Island, British Columbia, Canada support the development of a whole plant concept for an Eocene species of Gleichenia , and provide data to develop the first organismal concept for an extinct species of Gleichenia from the Cenozoic fossil record. New information for completing the plant reconstruction includes anatomically preserved pinnule fragments with attached radial exindusinate sori of six to seven annulate sporangia containing trilete spores. While the overall pattern of evolution within Gleicheniaceae remains incompletely understood because well-documented fossils from pre-Cretaceous deposits are inconsistently preserved, this organismal plant concept strengthens evidence that all diagnostic features of the family Gleicheniaceae were present, and that essentially modern species were well-established and diversifying during the Cretaceous and Paleogene. • First whole plant reconstruction of a Cenozoic species of Gleichenia. • Characterized by both external morphology and internal anatomy. • Evidence of stems, frond fragments, and sori of sporangia containing spores. • Essentially modern species of Gleicheniaceae evolved no later than the Paleogene. • Family Gleicheniaceae is inconclusively present in pre-Cretaceous deposits. [ABSTRACT FROM AUTHOR]

Abstract: 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. [Copyright &y& Elsevier]

An anatomically preserved leptosporangiate filicalean that combines characters of the Pteridaceae and tree ferns has been discovered in the Lower Cretaceous (Valanginian-Hauterivian boundary) Apple Bay flora on Vancouver Island, Canada. Exindusiate sori are radial, with numerous annulate sporangia diverging from a conical receptacle that is positioned below a pinnule vein. No paraphyses or other trichomes are present. Sporangia have a nearly vertical annulus that is not interrupted by the long, quadriseriate stalk. Spores are trilete and angular, with a distinctive equatorial flange and a distal deltoid structure that contacts the rim at the corners of the spore. The suture extends three-fourths of the distance to the margin and is flanked by prominent ridges. Spore walls consist of two dense layers, and the surface is psilate. Pterisorus radiata gen. et sp. nov. has characters that are reminiscent of both tree ferns and the most highly derived of modern species, providing direct evidence for the third major radiation of Filicales. [ABSTRACT FROM AUTHOR]

The discovery of two anatomically preserved rhizomes with attached stipe bases and adventitious roots contributes to our knowledge of filicalean fern diversity and evolution. Specimens are preserved in marine carbonate nodules at the Lower Cretaceous (Valanginian to Hauterivian) Apple Bay locality on Vancouver Island, British Columbia, Canada. They represent a previously unknown filicalean fern with transitional solenostelic-dictyostelic stelar architecture and are described as Stramineopteris aureopilosus gen. et sp. nov. Rhizomes range from 2.5 to 5.5 mm in diameter and are amphiphloic. Arc-shaped leaf traces display a slightly asymmetrical divergence pattern and radiate from all sides of the stele. Primary xylem maturation is exarch. Multicellular trichomes are present throughout but are produced in greater numbers at rhizome dichotomies. Both pith and cortex are of a distinctive heterogeneous construction. A comparison of this distinctive combination of characters to previously described fern rhizomes reveals that vegetative anatomy may be more taxonomically diagnostic for a broader range of ferns than is commonly thought. This realization provides the impetus to explore more closely the distribution of several anatomic characters among leptosporangiate ferns and to reexamine the value of vegetative anatomy for developing a clearer understanding of fern systematics. [ABSTRACT FROM AUTHOR]

Several fertile pinnules with abaxially borne sporangia have been discovered in calcareous marine nodules from the Middle or Late Eocene Appian Way locality on the east coast of Vancouver Island, British Columbia, Canada. Fertile pinnules are attached in groups of two or three. They are deeply lobed, with each lobe rolled toward the abaxial surface and the entire pinnule rolled toward the abaxial surface to form an irregularly globose structure. Sporangia occur abaxially and form two rows over the entire surface of each pinnule lobe. The midrib of each pinnule lobe is thickened, forming a receptacle from which broadly attached sporangia and numerous paraphyses diverge. As is characteristic of many schizaeaceous ferns, sporangia of the fossil are flask shaped with a short stalk and have a uniseriate, transverse annulus at the apex. Spores are tetrahedral and trilete, like those of the type species for the extinct genus Paralygodium. The new specimens are described as Paralygodium vancouverensis sp. nov. and are compared to both living genera of schizaeaceous ferns and the extinct species Paralygodium yezoense and Anemia poolensis. The unique combination of characters displayed by each of the fossil species reveals the presence of several extinct schizaeaceous lineages and supports interpretations that the diversity of Schizaeaceae s.l. was greater during the Cretaceous and Paleogene than it is today. [ABSTRACT FROM AUTHOR]

A distinctive new permineralized stem from marine deposits of western North America provides additional evidence for the diversity of Upper Cretaceous tree ferns. The fossil occurs in a calcareous concretion from the Late Campanian Spray Formation at Shelter Point, Vancouver Island, British Columbia, Canada. It measures 20 cm long and 7 cm in maximum diameter, with adventitious roots diverging between persistent, helically arranged fronds. Frond traces are derived from an amphiphloic dictyostele, and the stem produces no medullary or cortical bundles. The pith has a sclerenchymatous center and a parenchymatous outer zone. Sclerenchyma sheaths accompany both cauline and foliar vasculature. Frond traces diverge as six to ten bundles, most often eight. At the stem periphery, the cortex produces a homogeneous sclerenchymatous hypodermis, and is clothed by a dense ramentum of both uniseriate and large multiseriate trichomes. This novel combination of characters reveals the presence of a new genus and species of tree ferns, Rickwoodopteris hirsuta gen. et. sp. nov. Cladistic analysis of stem characters infers that R. hirsuta conforms to the dicksoniaceous grade of Cyatheaceae s.l., and further clarifies our understanding of evolutionary diversification among Mesozoic filicalean tree ferns. [Copyright &y& Elsevier]