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Premise of research. Early Cretaceous (Valanginian) rocks exposed at Apple Bay (Vancouver Island, Canada) have yielded numerous anatomically preserved bryophytes that have expanded our knowledge of the pre-Cenozoic fossil record of this group. A new moss type with tricostate leaves from Apple Bay combines characters not encountered in moss species recorded previously, warranting in-depth characterization of a new taxon. Methodology. Carbonate concretions containing permineralized fossil material were sectioned using the cellulose acetate peel technique. The resulting acetate peels providing serial sections through the bryophyte specimens were used to reconstruct the habit of the plant and its leaves in two and three dimensions. Pivotal results. We describe a new species of moss in the family Tricostaceae, Tricosta priapiana sp. nov. The anatomy and morphology of the new species justify expanding and refining the diagnoses of Tricostaceae and Tricosta , especially with respect to perigonial branches and the morphology of branch buds. The presence of pseudoparaphyllia surrounding the branch buds supports the pleurocarpous affinities proposed previously for the Tricostaceae. Conclusions. Tricosta priapiana adds to the diversity of tricostate mosses, an extinct group that was intriguingly diverse in the Mesozoic. Nevertheless, the tricostate state is probably a convergent trait, or a trait that may have evolved independently in acrocarpous and pleurocarpous moss lineages in response to similar selective pressures. Still, it remains unclear what such selective pressures may have been, as the paleoclimates and habitats that hosted the tricostate mosses are not well circumscribed, precluding comparisons among the different species and their closest analogs among living multicostate mosses. [ABSTRACT FROM AUTHOR]

Premise of research. Two permineralized inflorescences have been found in calcium carbonate concretions from the Eocene Appian Way locality on Vancouver Island, British Columbia, increasing sampling density and helping clarify the pattern of phylogeny for Alismatalean monocots. Methodology. Serial sections of specimens were prepared by the cellulose acetate peel technique and studied with light microscopy. Phylogenetic analyses using a morphological matrix of living and fossil Araceae and Acoraceae were performed using TNT version 1.5 to help establish relationships of the fossil inflorescences. Pivotal results. Inflorescences are cylindrical spikes, >37 mm long and 8.1 mm wide, with a peduncle at least 14.5 mm long, bearing numerous helically arranged, bisexual sessile flowers with hooded tepals. Fruits are trilocular, with basal axile placentation and one seed per locule. Remnants of probable stamens are present in most flowers. Conclusions. Flower structure was compared to those of other petalloid, spicate, bisexual flowers in the monocot families Acoraceae and Araceae (Alismatales). The number of carpels is similar to that in Acorus (Acoraceae) and Spathiphyllum (Araceae, Monsteroideae) and some Pothoideae. Single-seeded ovaries are common in Araceae, while the locules in Acorus contain several apically attached seeds. Phylogenetic analyses place Appianospadix in Proto-Araceae near the base of the aroid tree. This inflorescence with a unique combination of characters is described as Appianospadix bogneri , a new genus and species of Alismatales (family: Araceae). [ABSTRACT FROM AUTHOR]

Premise of research. The Late Cretaceous Yezo Group in Hokkaido, Japan, yields abundant plant debris in calcareous concretions. Nearly 150 species of plants have been described since 1910. Here we add a new genus of a probable stem polypodioid leptosporangiate fern to the plants of the Yezo Group based on a permineralized fertile pinnule from the Mikasa locality. Methodology. The fertile pinnule was detected in a calcareous concretion as an X-ray computed tomography image. Specimens were studied from serial cellulose acetate peels. Three-dimensional reconstructions of the entire pinnule and the soral, sporangial, and indusial morphologies were rendered using Molcer Plus and Amira. Pivotal results. The pinnule is 11.7 mm long, less than 7 mm wide, and deeply pinnatisect, with an abaxially grooved midrib or "costule" bearing one terminal and six lateral segments with catadromic pinnate venation. Lateral veins are simple or bifurcate once near the segment margin. The sorus is indusiate and abaxial and occurs slightly proximal to the end of the single vein or, when the lateral vein bifurcates, near the end of a much shorter anadromic vein. The indusium is reniform, is attached close to a massive receptacle only at its base, and is not laterally fused with the lamina. Approximately 17 sporangia with a slightly oblique, incomplete annulus, each estimated to contain ca. 64 spores, are formed per sorus. Spores are tetrahedral and trilete, have a triangular amb and slightly convex sides, are 42–45 μm in equatorial diameter, have a smooth exine, and are comparable to Biretisporites -type dispersed spores. Conclusions. The unique characters of the fossil are shared separately by extant basal polypods, most abundantly in Dennstaedtiaceae. However, the oblique annulus of the fossil does not occur in any extant members of Polypodiales unless it appears homoplastically. We therefore describe the fossil as a new taxon of a probable stem polypod, Mikasapteris rothwellii gen. et sp. nov., without any order and family assignments. The presence of Mikasapteris during the Late Cretaceous shows that pre-Polypodiales ferns were more diverse during this period. [ABSTRACT FROM AUTHOR]

Premise of research. Sixteen compression/impression specimens of a new floating aquatic angiosperm that enrich our knowledge of Late Cretaceous vegetation have been identified from the early Maastrichtian St. Mary River Formation near Cardston, Alberta, Canada. Methodology. Fossils were studied using dégagement and photographed under reflected light. Leaves were morphotyped using methodology in the Manual of Leaf Architecture and compared to other extant and extinct rosette-forming floating aquatic angiosperms. Pivotal results. Stems of this aquatic dicot show determinate growth of floating shoots with opposite/decussate attachment of heterophyllous leaves. Leaves are simple and sessile at the apicalmost node, simple and petiolate at the second node, and compound with three leaflets at nodes 3–6. Compound leaves are petiolate and ternate (trifoliate). Leaflet shape is unlobed, obovate, and symmetrical; base is acute-obtuse; apex angle is obtuse; base shape is convex; petiolar attachment is marginal; apex shape is rounded; and margin is crenate. Primary veins are flabellate to pinnate with three to five basal veins that dichotomize distally. Secondary veins are craspedodromous with uniform spacing and angles. Teeth are of a single order, three per centimeter near apex, and convex-convex; the sinus is angular, and the apex is foraminate. Conclusions. While the basic growth habit of this plant is similar to that of several extant taxa in the Onagraceae, Lythraceae, Plantaginaceae, and Hydrocharitaceae, leaf venation and the presence of compound leaves differ greatly from the living species. In growth habit and the presence of both simple and compound leaves, the new plant is similar to Quereuxia angulata , also found at this fossil site. These fossils are described as a new genus and species, Tolmania aquatica Edmonds, Stockey & Rothwell, which differs from Quereuxia in pattern of heterophylly, venation, larger plant size, stem morphology, and the presence of hydathodes in the leaves. [ABSTRACT FROM AUTHOR]

PREMISE: Fossils can reveal long‐vanished characters that inform inferences about the timing and patterns of diversification of living fungi. Through analyzing well‐preserved fossil scutella, shield‐like covers of fungal sporocarps, we describe a new taxon of early Dothideomycetes with a combination of characters unknown among extant taxa. METHODS: Macerated clays from the Potomac Group, lower Zone 1, from the Lower Cretaceous (Aptian, 125–113 Ma) of Virginia USA yielded one gymnospermous leaf cuticle colonized by 21 sporocarps of a single fungal morphotype. We inferred a tree from nuclear ribosomal DNA of extant species, and coded morphological characters to evaluate alternative, equally parsimonious placements of the fossil in a molecular constraint tree of extant species. RESULTS: Bleximothyrium ostiolatum gen. et sp. nov. has an ostiolate scutellum of radiate, dichotomizing hyphae. Unlike otherwise similar extant and fossil taxa, B. ostiolatum has tangled hyphae at its scutellum margin. Scutella of B. ostiolatum are connected to superficial mycelium, to intercalary and lateral appressoria, and to extensive subcuticular "mycélium en palmettes". The gymnospermous host has characters consistent with identity as a non‐papillate ginkgophyte or cycad. CONCLUSIONS: Bleximothyrium ostiolatum is the oldest known fossil fly‐speck fungus that occurs on plant cuticles and has the radiate, ostiolate scutellum known only from Dothideomycetes. Its combination of characters, its scutellum margin, and mycélium en palmettes are unknown in other extant and fossil species, and Bleximothyrium ostiolatum likely represents a new group of fly‐speck fungi that may now be extinct. [ABSTRACT FROM AUTHOR]

Premise of research. A single cylindrical conifer seed cone has been discovered in a calcium carbonate marine concretion from the Upper Cretaceous Eden Main Quarry, dated as early Coniacian (∼89 Ma), 35 km northwest of Courtenay on Vancouver Island, British Columbia, Canada. Methodology. The specimen was sectioned with a rock saw, and serial sections were prepared using the classical cellulose acetate peel technique. Peel sections were mounted on microscope slides. Reconstructions of the cone and seeds were rendered with Avizo. Pivotal results. The cone is somewhat abraded, at least 2.3 cm long and 1 cm wide, with the apex missing and most of the base coalified. The well-preserved midsection shows bract/scale complexes inserted at a steep angle. The bract/scale complex has a central rhomboidal shaft and two lateral winglike extensions. Xylem of the cone axis is robust, with secondary xylem tracheids that have circular-bordered pits. A single circular vascular trace enters the bract/scale complex. Three resin canals enter the base of the scale and divide to form up to eight or nine canals distally. Six overlapping, inverted winged seeds, ∼8 mm long and 1 mm thick, with two large, fleshy wings are borne on the adaxial surface of each scale. The nucellus is free from the endotesta to its base. Megagametophytes and embryos occur in some seeds. Cells of the megagametophyte have granular contents. This cone shows similarities in overall structure and vasculature to those of living Taiwania (Cupressaceae); however, the fossil cone size is generally larger, with six seeds per scale versus two, and seeds are extremely large and fleshy with a unique submicropylar structure. Conclusions. This cone is compared with living and fossil taxa of Cupressaceae and described as Comoxostrobus rossii gen. et sp. nov. (Cupressaceae subfamily Taiwanioideae), revealing that taiwanioid Cupressaceae were much more diverse and widespread during the Cretaceous. [ABSTRACT FROM AUTHOR]

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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]

PREMISE OF THE STUDY: Mosses, very diverse in modern ecosystems, are currently underrepresented in the fossil record. For the pre-Cenozoic, fossil mosses are known almost exclusively from compression fossils, while anatomical preservation, which is much more taxonomically informative, is rare. The Lower Cretaceous of Vancouver Island (British Columbia, Canada) hosts a diverse anatomically preserved flora at Apple Bay. While the vascular plant component of the Apple Bay flora has received much attention, the numerous bryophytes identified at the locality have yet to be characterized. METHODS: Fossil moss gametophytes in more than 20 carbonate concretions collected from the Apple Bay locality on Vancouver Island were studied in serial sections prepared using the cellulose acetate peel technique. KEY RESULTS: We describe Tricosta plicata gen. et sp. nov., a pleurocarpous moss with much-branched gametophytes, tricostate plicate leaves, rhizoid-bearing bases, and delicate gametangia (antheridia and archegonia) borne on specialized branches. A new family of hypnanaean mosses, Tricostaceae fam. nov., is recognized based on the novel combination of characters of T. plicata. CONCLUSIONS: Tricosta plicata reveals pleurocarpous moss diversity unaccounted for in extant floras. This new moss adds the first bryophyte component to an already diverse assemblage of vascular plants described from the Early Cretaceous at Apple Bay and, as the oldest representative of the Hypnanae, provides a hard minimum age for the group (136 Ma). [ABSTRACT FROM AUTHOR]

Premise of the study: Two ovulate conifer cones, one of which is attached terminally to a short leafy shoot, reveal the presence of a new species of Hughmillerites in the Early Cretaceous Apple Bay flora of Vancouver Island, British Columbia, Canada. This ancient conifer expands the diversity of Cupressaceae in the Mesozoic and reveals details about the evolution of Subfamily: Cunninghamioideae. • Methods: Specimens were studied from anatomical sections prepared using the cellulose acetate peel technique. • Key results: Vegetative shoots have helically arranged leaves that are CunninghamiaAike. Seed cones have many helically arranged bract/scale complexes in which the bract is larger than the ovuliferous scale. Each ovuliferous scale has three free tips that separate from the bract immediately distal to an inverted seed. Several ovuliferous scales show interseminal ridges between seeds. • Conclusions: This study documents a new extinct species of cunninghamioid conifers, Hughmillerites vancouverensis, expanding the record of the genus from the Late Jurassic to the Early Cretaceous. This new extinct species emphasizes the important role that conifers from subfamily Cunninghamioideae played in the initial evolutionary radiation of Cupressaceae. In light of recent findings in conifer regulatory genetics, we use H. vancouverensis to hypothesize that variations of expression in certain gene homologues played an important role in the evolution of the cupressaceous ovuliferous scale. [ABSTRACT FROM AUTHOR]

More than 800 permineralized fagaceous fruits have been studied from the Eocene Appian Way locality of Vancouver Island, British Columbia. These cupulate nuts are the most common fruits preserved at the locality. They occur in calcareous concretions and were studied using the cellulose acetate peel technique. Cupules are borne on a spiny stalk and are broadly ovate in both longitudinal and transverse section. Cupules are evalvate and have both branching and simple spines. A single, ovoid, sclerotic nut is enclosed entirely by the cupule, except at the apex, where a stylar protrusion is free from any surrounding tissues. The nut is bilocular with a glabrous endocarp lining. At maturity, a single seed occupies the locular cavity. The embryo is straight and no endosperm is evident. The single-fruited, spiny cupule is most similar to fruits of Fagaceae subfamily Castaneoideae. Bicarpellate fruits and a glabrous endocarp place them within the fossil genus Cascadiacarpa; however, they differ from Cascadiacarpa spinosa in nut wall anatomy, cupule ornamentation, shape, and size. Fruits of Cascadiacarpa exilis sp. nov. are compared with spiny, small compression and impression fossil fruits from the Eocene Taneum Creek locality of Washington State. The characters of the Washington State fossils overlap with those observed in the anatomically preserved Appian Way cupulate fruits and, as such, may represent compression and impression fossils of the same genus. The permineralized fruits document that evalvate, spiny, cupulate nuts of Fagaceae were present and common in the Eocene of western North America. [ABSTRACT FROM AUTHOR]

Premise of the study: Colonists of even the most inhospitable environments, lichens are present in all terrestrial ecosystems. Because of their ecological versatility and ubiquity, they have been considered excellent candidates for early colonizers of terrestrial environments. Despite such predictions, good preservation potential, and the extant diversity of lichenized fungi, the fossil record of lichen associations is sparse. Unequivocal lichen fossils are rare due, in part, to difficulties in ascertaining the presence of both symbionts and in characterizing their interactions. This study describes an exceptionally well-preserved heteromerous lichen from the Lower Cretaceous of Vancouver Island. Methods: The fossil occurs in a marine carbonate concretion collected from the Apple Bay locality on Vancouver Island, British Columbia, and was prepared for light microscopy and SEM using the cellulose acetate peel technique. Key results: The lichen, Honeggeriella complexa gen. et sp. nov., is formed by an ascomycete mycobiont and a chlorophyte photobiont, and exhibits heteromerous thallus organization. This is paired with a mycobiont-photobiont interface characterized by intracellular haustoria, previously not documented in the fossil record. Conclusions: Honeggeriella adds a lichen component to one of the richest and best characterized Early Cretaceous floras and provides a significant addition to the sparse fossil record of lichens. As a heteromerous chlorolichen, it bridges the >350 million-year gap between previously documented Early Devonian and Eocene occurrences. [ABSTRACT FROM AUTHOR]

* Premise of the study: Pinaceae and nonpinoid species are sister groups within the conifer clade as inferred from molecular systematic comparisons of living species and therefore should have comparable geological ages. However, the fossil record for the nonpinoid lineage of extant conifer families is Triassic, nearly 100 million years older than the oldest widely accepted Lower Cretaceous record for Pinaceae. An anatomically preserved fossil conifer seed cone described here extends the stratigraphic range of Pinaceae nearly 30 million years, thus reducing the apparent discrepancy between evidence from the fossil record and inferences from systematic studies of living species. * Methods: Material was prepared as serial thin sections by the cellulose acetate peel technique, mounted on microscope slides, and viewed and photographed using transmitted light. * Key results: A large cylindrical cone consisting of bract-scale complexes that diverge from the cone axis in a helical phyllotaxis has bracts and scales that separate from each other in the midregion and are of equal length and of nearly equal width. The cone has two inverted and winged seeds that are attached to the adaxial surface of each cone scale and, thus, represents an early member of the Pinaceae. * Conclusions: Eathiestrobus mackenziei gen. et sp. nov. extends the fossil record for well-documented members of the family Pinaceae from the Lower Cretaceous to the Kimmeridgian Stage of the Upper Jurassic. This species also clarifies the set of characters that are diagnostic for seed cones of Pinaceae and reveals possible plesiomorphic characters for seed cones of the family. [ABSTRACT FROM AUTHOR]

• Premise of the study: Morphology is a reflection of evolution, and as the majority of biodiversity that has lived on Earth is now extinct, the study of the fossil record provides a more complete picture of evolution. This study investigates anatomically preserved bryophyte fossils from the Eocene Oyster Bay Formation of Vancouver Island. While the bryophyte fossil record is limited in general, anatomically preserved bryophytes are even more infrequent; thus, the Oyster Bay bryophytes are a particularly significant addition to the bryophyte fossil record. • Methods: Fossils occur in two marine carbonate nodules collected from the Appian Way locality on the eastern shore of Vancouver Island, British Columbia, and were prepared using the cellulose acetate peel technique. • Key results: The fossils exhibit a novel combination of characters unknown among extinct and extant liverworts: (1) three-ranked helical phyllotaxis with underleaves larger than the lateral leaves; (2) fascicled rhizoids associated with the leaves of all three ranks; (3) Anomoclada-type endogenous branching. • Conclusions: A new liverwort family, Appianacae fam. nov., is established based upon the novel combination of characters. Appiana gen. nov. broadens the known diversity of bryophytes and adds a hepatic component to one of the richest and best characterized Eocene floras. [ABSTRACT FROM AUTHOR]

Abstract: This paper provides a whole plant concept for a new species, Emporia cryptica sp. nov. Hernandez-Castillo, Stockey, Rothwell & Mapes (Emporiaceae: Voltziales), the fourth conifer to be reconstructed from the rich fossil biota at the Late Pennsylvanian, Hamilton Quarry, Kansas. E. cryptica has an orthotropic stem, lateral plagiotropic branches with simple leaves, simple pollen cones, and compound ovulate cones. Branches have an endarch eustele with dense wood surrounding a parenchymatous pith with sclerotic nests/plates, and secondary xylem tracheids with multiseriate hexagonal bordered pits. Leaves on both penultimate and ultimate branches are simple and amphistomatic with two adaxial stomatal bands, monocyclic and dicyclic stomata, and two narrow abaxial rows of stomata with numerous trichome bases. Pollen cones are simple with helically arranged microsporophylls and adaxial pollen sacs. Prepollen is monolete, eusaccate, and monosaccate (Potonieisporites Bharadwaj). Ovulate cones are compound with bilaterally symmetrical dwarf shoots in the axils of helically arranged forked bracts. Axillary dwarf shoots bear numerous sterile scales interspersed with two megasporophylls, each bearing a single inverted terminal ovule. This plant displays morphological and cuticular characters similar to several morphogenera of Paleozoic walchian conifers but is most comparable to the fossil plant species Emporia lockardii and Hanskerpia hamiltonensis. E. cryptica is the only walchian conifer where ovules and seeds with megagametophytes, immature embryos and mature embryos have been documented, demonstrating that the most ancient conifers possessed seed dormancy and polycotyledonary embryos. This reproductive biology is similar to that of many Mesozoic and extant conifers with saccate pollen except for the presence of prepollen grains that are common among Paleozoic walchian conifers. [Copyright &y& Elsevier]

A diverse assemblage of fossil conifer leaves was identified in calcareous marine concretions from the Lower Cretaceous (Valanginian–Hauterivian) Apple Bay locality, Vancouver Island. Of the hundreds of isolated leaf fragments, most show affinities to Pinaceae. Leaves with closest similarities to Picea (spruce) were studied using cellulose acetate peels. Picea-like leaves vary in cross-section from rhomboidal, pentagonal, triangular, to ovoid. One fused vascular bundle with a centrally located ray and abaxial sclerenchyma is surrounded by a circular endodermis and transfusion tissue. Mesophyll is plicate containing two lateral external resin canals surrounded by a sclerenchyma sheath. Hypodermal fibres are one to three layers thick, except in areas of stomata. Leaves are amphistomatic, with sunken guard cells. Vascular bundles are identical anatomically to Picea; however, plicate mesophyll is similar to that in leaves of Pinus. Extensive sclerenchyma in the hypodermis and surrounding resin canals differs from that in most extant Picea. The major difference between these leaves and those of Picea is leaf shape. These fossil leaves probably belong to an extinct pinaceous conifer, and are described as Midoriphyllum piceoides gen. et sp. nov. Similar evidence from Cretaceous seed cones suggests that like the angiosperms, the Pinaceae were undergoing rapid mosaic evolution during the Lower Cretaceous. [ABSTRACT FROM AUTHOR]

Large numbers of permineralized juglandaceous fruits were identified in calcareous nodules from the Eocene Appian Way locality on Vancouver Island, British Columbia, Canada. The fruits, small dorsiventrally flattened nutlets, 4.5–7.0 mm long and 5.5–9.0 × 3–5 mm in diameter, were studied using cellulose acetate peels. They are wingless, ribbed, and have a lobed epicarp that surrounds the hurler. Cells of the inner epicarp are thin-walled and traversed by a system of branching vascular strands. The stony nutlet wall is composed of fibers, with an outer layer of distinctive idioblasts. The fruits have a symmetry like that in Juglandaceae, subfamily Juglandoideae, tribe Platycaryeae, while the fibrous taut walls are like those of subfamily Engelhardioideae. This unique combination of characters indicates that these fruits represent a new genus and species of Juglandaceae: Beardia vancouverensis gen. et sp. nov. The excellent preservation of the Appian Way specimens has allowed a unique view of the internal fruit anatomy and external morphology. As the only wingless, flattened nuts known in the family, they further extend the range of morphological variation in fruits in the family. These fossils further support the hypothesis that North America was an important center of generic diversity for Juglandaceae during the early Tertiary. [ABSTRACT FROM AUTHOR]

Relationships of living and fossil Pinaceae were inferred using parsimony and Bayesian inference of morphological characters and plastid and nuclear DNA sequences. When considering extant taxa only, adding molecular to morphological characters resulted in markedly increased resolution and branch support compared with analysis of morphology alone. Including 45 fossil taxa resulted in drastically decreased resolution in morphology-based consensus trees. We evaluated the effect on branch support and resolution of including DNA sequences, deleting fossils lacking information for cone scale apices and seeds, using reduced consensus methods, and using implied weighting, and found that the greatest improvements were found by including DNA sequences and using implied weighting. The tree topologies from parsimony and Bayesian inference confirm previous findings that the fossil genus Pseudoaraucaria and a few species of Pityostrobus from the Lower Cretaceous are related to abietoid genera, and that other species of Pityostrobus are pinoid and closely related to Pinus. Focusing phylogenetic analyses on the most complete fossil cones, specifically those that are anatomically preserved and include both cone scale apices and seeds, and taking into account homoplasy, resulted in the clearest hypotheses for the timing and sequence of diversification in the family. [ABSTRACT FROM AUTHOR]