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Premise: Anatomically preserved evidence for a novel clade of gymnosperms emphasizes diversity of seed plants immediately prior to the appearance of angiosperm fossils in the paleontological record. Methods: Cupulate seeds from the Early Cretaceous Apple Bay locality (Vancouver Island) are described from serial cellulose acetate peels and three‐dimensional reconstruction. Phylogenetic context is assessed through the comparative analysis of gymnosperm seed producing fructifications and maximum parsimony analysis of a revised morphological data set for seed plant phylogeny. Results: Xadzigacalix quatsinoensis gen. et sp. nov. is characterized by an orthotropous ovule with an elongated micropyle and complex integument, enclosed within a radial cupule. The micropylar canal is elongated; and the nucellus extends into the micropyle to seal the post pollination ovule. Except at the apex of the micropyle, the seed is completely enclosed by a parenchymatous cupule with ca. 20 axially elongated secretory ducts. The cupulate seed is produced upon a triangular woody stele, consisting of a parenchymatous pith surrounded by radially aligned tracheids. The stele produces three short terete traces that terminate within the base of the cupule as transfusion tissue at the seed chalaza. Conclusions: Organography, vascularization, nature of the integument and nucellus, and configuration of the micropylar canal distinguish Xadzigacalix quatsinoensis from all other gymnosperm clades. Cladistic analyses suggest the new plant may have affinities with gnetophytes or angiosperms. These results are complemented with a critical re‐evaluation of ovulate structures for Mesozoic gymnosperms, providing new insight into plant diversity immediately antecedent to the explosive diversification of flowering plants. [ABSTRACT FROM AUTHOR]

A new genus and species, Geminispermum virginiense, is described based on a well-preserved coalified cupulate reproductive unit recovered from the Early Cretaceous (early–middle Albian) Puddledock locality, Virginia, U.S.A. The reproductive unit is bisymmetrical and consists of an axis that bifurcates into two cupule-bearing stalks, each in the axil of a bract. Each cupule stalk bears a single non-valvate cupule recurved towards the center of the reproductive unit. The cupule opens distally by a short transverse slit with a distinct upper margin. Each cupule almost completely encloses a single orthotropous seed that is free from the cupule except at the base. The nucellus is also free from the integument except at the basal point of attachment. Geminispermum combines features of the ovulate structures of Caytoniales, Umkomasiales (= Corystospermales, including Doyleales) and Petriellales, but cannot be included in any of these existing orders as they are currently understood. The recurved, closed, non-valvate cupules are particularly similar to those of Caytonia, Petriellaea and Reymanownaea in external morphology, but differ in being one-seeded. The cupules of Geminispermum differ from the one-seeded cupules of Umkomasiales in being non-valvate and in having only a single cupule per bract. Geminispermum is perhaps most similar to the one- or two-seeded non-valvate cupules of Ktalenia from the Early Cretaceous of Argentina, but Ktalenia is poorly preserved, details of cupule architecture are uncertain, and the cupules appear to be associated with a single strongly dissected bract. Geminispermum is currently the only unequivocal seed plant cupule recovered from the Early Cretaceous Potomac Group and is distinct from all previously described cupulate reproductive structures.

A new genus and species, Geminispermum virginiense, is described based on a well-preserved coalified cupulate reproductive unit recovered from the Early Cretaceous (early–middle Albian) Puddledock locality, Virginia, U.S.A. The reproductive unit is bisymmetrical and consists of an axis that bifurcates into two cupule-bearing stalks, each in the axil of a bract. Each cupule stalk bears a single non-valvate cupule recurved towards the center of the reproductive unit. The cupule opens distally by a short transverse slit with a distinct upper margin. Each cupule almost completely encloses a single orthotropous seed that is free from the cupule except at the base. The nucellus is also free from the integument except at the basal point of attachment. Geminispermum combines features of the ovulate structures of Caytoniales, Umkomasiales (= Corystospermales, including Doyleales) and Petriellales, but cannot be included in any of these existing orders as they are currently understood. The recurved, closed, non-valvate cupules are particularly similar to those of Caytonia, Petriellaea and Reymanownaea in external morphology, but differ in being one-seeded. The cupules of Geminispermum differ from the one-seeded cupules of Umkomasiales in being non-valvate and in having only a single cupule per bract. Geminispermum is perhaps most similar to the one- or two-seeded non-valvate cupules of Ktalenia from the Early Cretaceous of Argentina, but Ktalenia is poorly preserved, details of cupule architecture are uncertain, and the cupules appear to be associated with a single strongly dissected bract. Geminispermum is currently the only unequivocal seed plant cupule recovered from the Early Cretaceous Potomac Group and is distinct from all previously described cupulate reproductive structures. [ABSTRACT FROM AUTHOR]

PREMISE OF THE STUDY: Discovery of cupulate ovules of Doylea tetrahedrasperma within a compact, compound seed cone highlights the rich diversity of fructification morphologies, pollination biologies, postpollination enclosure of seeds, and systematic diversity of Early Cretaceous gymnosperms. METHODS: Specimens were studied using the cellulose acetate peel technique, three-dimensional reconstructions (in AVIZO), and morphological phylogenetic analyses (in TNT). KEY RESULTS: Doylea tetrahedrasperma has bract/fertile short shoot complexes helically arranged within a compact, compound seed cone. Complexes diverge from the axis as a single unit and separate distally into a free bract tip and two sporophylls. Each sporophyll bears a single, abaxial seed, recurved toward the cone axis, that is enveloped after pollinaton by sporophyll tissue, forming a closed cupule. Ovules are pollinated by bisaccate grains captured by micropylar pollination horns. CONCLUSIONS: The unique combination of characters shown by D. tetrahedrasperma includes the presence of cupulate seeds borne in conifer-like compound seed cones, an ovuliferous scale analogue structurally equivalent to the ovulate stalk of Ginkgo biloba, gymnospermous pollination, and nearly complete enclosure of mature seeds. These features characterize the Doyleales ord. nov., clearly distinguish it from the seed fern order Corystospermales, and allow for recognition of another recently described Early Cretaceous seed plant as a second species in genus Doylea. A morphological phylogenetic analysis highlights systematic relationships of the Doyleales ord. nov. and emphasizes the explosive phylogenetic diversification of gymnosperms that was underway at the time when flowering plants may have originated and/or first began to radiate. [ABSTRACT FROM AUTHOR]