Fossil evidence for sporeling development of a Mesozoic osmundaceous fern.

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.

A novel cupulate seed plant, Xadzigacalix quatsinoensis gen. et sp. nov., provides new insight into the Mesozoic radiation of gymnosperms.

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.

Fossil evidence for Paleocene diversification of Araceae: Bognerospadix gen. nov. and Orontiophyllum grandifolium comb. nov.

PREMISE: Nearly 200 araceous leaves and two spadices have been identified among Paleocene fossils from the Blindman River locality near Blackfalds, Alberta, Canada. Although not found in attachment, these probably represent parts of the same extinct plant species. METHODS: Specimens were studied using light microscopy. Phylogenetic analyses using a morphological matrix of living and fossil Araceae were performed using TNT version 1.5 to help establish relationships of the fossil leaves and spadices within Araceae and to each other. RESULTS: Leaves are simple with a broad petiole, entire margin, and elliptic to ovate or oblong blade with an acute to slightly rounded apex. A multi-veined midrib extends into the basal region of the blade. Parallelodromous primary veins of two orders diverge at acute angles, converging with a submarginal vein or at the apex. Transverse veins are opposite percurrent, producing rectangular to polygonal areoles. Higher-order veins are mixed opposite/alternate. Spadices are cylindrical, with helically arranged, bisexual, perigoniate flowers, each with six free tepals and a protruding style. Fruits are trilocular, with axile placentation and one seed per locule. CONCLUSIONS: Leaves are assignable to the fossil genus Orontiophyllum J. Kvacek & S.Y. Sm. as O. grandifolium comb. nov. Spadices are described as Bognerospadix speirsiae gen. et sp. nov. Leaves and spadices each conform to an early-diverging lineage of Araceae, increasing the known diversity of Proto-Araceae (viz., subfamilies Gymnostachydoideae and Orontioideae). Together, they provide strong evidence for extinct Proto-Araceae with novel combinations of characters shortly after the Cretaceous-Paleogene floral transition.

Cynodontium luthii sp. nov.: a permineralized moss gametophyte from the Late Cretaceous of the North Slope of Alaska.

PREMISE: Mosses are a major component of Arctic vegetation today, with >500 species known to date. However, the origins of the Arctic moss flora are poorly documented in the fossil record, especially prior to the Pliocene. Here, we present the first anatomically preserved pre-Cenozoic Arctic moss and discuss how the unique biology of bryophytes has facilitated their success in polar environments over geologic time. METHODS: A permineralized fossil moss gametophyte within a block of Late Cretaceous terrestrial limestone, collected along the Colville River on the North Slope of Alaska, was studied in serial sections prepared using the cellulose acetate peel technique. RESULTS: The moss gametophyte is branched and has leaves with a broad base, narrow blade, and excurrent costa. We describe this fossil as Cynodontium luthii sp. nov., an extinct species of a genus that is known from the High Arctic today. Cynodontium luthii is the oldest evidence of the family Rhabdoweisiaceae (by ≥18 Ma) and reveals that genera of haplolepideous mosses known in the extant Arctic flora also lived in high-latitude temperate deciduous forests during the Late Cretaceous. CONCLUSIONS: The occurrence of C. luthii in Cretaceous sediments, together with a rich Pliocene-to-Holocene fossil record of extant moss genera in the High Arctic, suggests that some moss lineages have exploited their poikilohydric, cold- and desiccation-tolerant physiology to live in the region when it experienced both temperate and freezing climates.

Extending the fossil record for foliicolous Dothideomycetes: Bleximothyrium ostiolatum gen. et sp. nov., a unique fly-speck fungus from the Lower Cretaceous of Virginia, USA.

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.

Evolutionary diversification of taiwanioid conifers: evidence from a new Upper Cretaceous seed cone from Hokkaido, Japan.

A single cylindrical seed cone 2 cm long, 1.1 cm wide has been found preserved in a calcium carbonate marine concretion from the Hakobuchi Formation (late Campanian-early Maastrichtian) of Hobetsu, Hokkaido, Japan. The cone, attached to a bent peduncle lacking leaves, has helically arranged bract/scale complexes that arise at right angles from the cone axis in the middle of the cone. The cone axis, ca. 1 mm wide, has a broad cylinder of secondary vascular tissue, and lacks a continuous resin canal system. Bract-scale complexes are laminar, cordate-orbiculate, and upturned distally, consisting primarily of bract tissue with no visible scale tip. The vascular trace to the bract/scale complex originates as a rod that divides laterally into several traces at the level of seed attachment. A single resin canal originates at the base of the bract-scale complex abaxial to the vascular strand, but more distally there are up to ca. 15 large resin canals that form a single row. Two to three inverted winged seeds are attached adaxially near the cone periphery. Cone structure and vascularization are most similar to those in the Cupressaceae, Subfamily Taiwanioideae, differing from living Taiwania cryptomerioides by having up to three seeds/scale rather than two, an abruptly upturned bract tip, in details of bract/scale vasculature, and a cone peduncle lacking leaves. This cone is described as Mukawastrobus satoi Stockey, Nishida and Rothwell. Together with previously described Early to Late Cretaceous taiwanioid seed cones from Mongolia and Hokkaido the new species demonstrates that the taxonomically diagnostic characters of such conifers are as subtle as those of Cretaceous and Cenozoic sequoioid Cupressaceae. This realization emphasizes that evolutionary diversification and turnover among taiwanioid conifers during the Cretaceous and Paleogene are probably far greater than currently recognized.

Diversification of crown group Araucaria: the role of Araucaria famii sp. nov. in the mid-Cretaceous (Campanian) radiation of Araucariaceae in the Northern Hemisphere.

PREMISE: Exceptional anatomical preservation of a fossil araucarian seed cone from a marine carbonate concretion from Vancouver Island, British Columbia, Canada provides unusually complete evidence for cone structure including seeds, megagametophytes, microgametophytes, and embryos of an Upper Cretaceous (Campanian) species of Araucaria, providing important new insights into the structure and relationships of Cretaceous Northern Hemisphere Araucariaceae. METHODS: The cone was studied from serial thin sections prepared by the coal ball peel technique. Phylogenetic analysis using a modified morphological matrix with both discrete and continuous characters was performed using TNT version 1.5. RESULTS: The nearly spherical cone, 6 × 6 cm in diameter, has helically arranged cone-scale complexes, consisting of a large bract with an upturned tip and a small, fleshy ovuliferous scale. Vascularization of the cone-scale complex is single at its origin. Widely winged bracts, with a bulging base, contain numerous vascular bundles, interspersed with transfusion tissue, and a large number of resin canals. Seeds are ovoid, 1.2 cm long, 1.2 cm in diameter. Nucellus is free from the integument, except at its base, with a convoluted apex, containing possible pollen tubes. Megagametophytes and mature cellular embryos occur in several seeds. CONCLUSIONS: This small cone with attached, imbricate leaves, wide bracts, and unusually large seeds, most closely resembles those of Araucaria Section Eutacta. Width and continuity of secondary xylem in the cone axis, and intact cone-scale complexes indicate that this cone probably did not disarticulate readily at maturity. When added to a modified, previously published phylogenetic analysis, Araucaria famii sp. nov. enhances our understanding of the Cretaceous radiation of Northern Hemisphere Araucaria Section Eutacta.

A new epiphyllous fly-speck fungus from the Early Cretaceous Potomac Group of Virginia (125-112 Ma): Protographum luttrellii, gen. et sp. nov.

Fly-speck fungi reproduce via thyriothecia that consist of sporogenous tissue appressed to cuticle surfaces of plant leaves and covered by a shield-like scutellum. Thyriothecial scutella likely evolved repeatedly in Dothideomycetes (Ascomycota), and their morphology varies by lineage. Fly-speck fungi have an exceptionally good fossil record that begins in the Mesozoic. The interpretation of scutellum characters in fossils may provide insights into origins of Dothideomycetes and help calibrate the timing of ascomycete evolution. From sediments of the Lower Cretaceous (125-112 Ma) Potomac Group of Virginia, from Dutch Gap Canal, lower Zone 1, we found scutella similar to those of extant Aulographaceae (Dothideomycetes), attached to a single piece of dispersed coniferous cuticle. We analyze hyphae and scutellum development among four extant Aulographaceae species for comparison with the fossil. The excellent preservation of fungi on the leaf cuticle surface allows us to infer a developmental sequence for the fossil. Scutellum development begins with coordinated growth of multiple neighboring generator hyphae and continues with hyphae producing two-dimensional pseudomonopodial, dichotomous, radial growth. Asci and ascospores were not found. We coded states for seven morphological characters using direct observations of the fossil and eight extant taxa, and using the literature for 28 others. We inferred a phylogeny using nuclear 18S and 28S rDNA of 36 extant taxa, 34 Dothideomycetes and two Arthoniomycetes. The phylogeny includes newly determined sequences from five species, two from Aulographaceae. With a branch-and-bound search, we inferred the most parsimonious placements of the fossil given the molecular tree topology. The parsimony analysis constrained by the rDNA phylogeny places the fossil taxon among stem lineages near Aulographaceae or among the known living members of Aulographaceae. We describe the fossil morphotype as Protographum luttrellii, gen. et sp. nov. The fossil provides the oldest evidence of morphological characters restricted among extant fungi to Aulographaceae.

Resolving the overall pattern of marattialean fern phylogeny.

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.

Grimmiaceae in the Early Cretaceous: Tricarinella crassiphylla gen. et sp. nov. and the value of anatomically preserved bryophytes.

Background and Aims: Widespread and diverse in modern ecosystems, mosses are rare in the fossil record, especially in pre-Cenozoic rocks. Furthermore, most pre-Cenozoic mosses are known from compression fossils, which lack detailed anatomical information. When preserved, anatomy significantly improves resolution in the systematic placement of fossils. Lower Cretaceous (Valanginian) deposits on Vancouver Island (British Columbia, Canada) contain a diverse anatomically preserved flora including numerous bryophytes, many of which have yet to be characterized. Among them is the grimmiaceous moss described here. Methods: One fossil moss gametophyte preserved in a carbonate concretion was studied in serial sections prepared using the cellulose acetate peel technique. Key Results: Tricarinella crassiphylla gen. et sp. nov. is a moss with tristichous phyllotaxis and strongly keeled leaves. The combination of an acrocarpous condition (inferred based on a series of morphological features), a central conducting strand, a homogeneous leaf costa and a lamina with bistratose portions and sinuous cells, and multicellular gemmae, supports placement of Tricarinella in family Grimmiaceae. Tricarinella is similar to Grimmia, a genus that exhibits broad morphological variability. However, tristichous phyllotaxis and especially the lamina, bistratose at the base but not in distal portions of the leaf, set Tricarinella apart as a distinct genus. Conclusions: Tricarinella crassiphylla marks the oldest record for both family Grimmiaceae and sub-class Dicranidae, providing a hard minimum age (136 million years) for these groups. The fact that this fossil could be placed in an extant family, despite a diminutive size, emphasizes the considerable resolving power of anatomically preserved bryophyte fossils, even when recovered from allochthonous assemblages of marine sediments, such as the Apple Bay flora. Discovery of Tricarinella re-emphasizes the importance of paleobotanical studies as the only approach allowing access to a significant segment of biodiversity, the extinct biodiversity, which is unattainable by other means of investigation.

Extending the fossil record of Polytrichaceae: Early Cretaceous Meantoinea alophosioides gen. et sp. nov., permineralized gametophytes with gemma cups from Vancouver Island.

PREMISE OF THE STUDY: Diverse in modern ecosystems, mosses are dramatically underrepresented in the fossil record. Furthermore, most pre-Cenozoic mosses are known only from compression fossils, lacking detailed anatomical information. When preserved, anatomy vastly improves resolution in the systematic placement of fossils. Lower Cretaceous deposits at Apple Bay (Vancouver Island, British Columbia, Canada) contain a diverse anatomically preserved flora that includes numerous bryophytes, many of which have yet to be characterized. Among them is a polytrichaceous moss that is described here. METHODS: Fossil moss gametophytes preserved in four carbonate concretions were studied in serial sections prepared using the cellulose acetate peel technique. KEY RESULTS: We describe Meantoinea alophosioides gen. et sp. nov., a polytrichaceous moss with terminal gemma cups containing stalked, lenticular gemmae. Leaves with characteristic costal anatomy, differentiated into sheathing base and free lamina and bearing photosynthetic lamellae, along with a conducting strand in the stem, place Meantoinea in family Polytrichaceae. The bistratose leaf lamina with an adaxial layer of mamillose cells, short photosynthetic lamellae restricted to the costa, and presence of gemma cups indicate affinities with basal members of the Polytrichaceae, such as Lyellia, Bartramiopsis, and Alophosia. CONCLUSIONS: Meantoinea alophosioides enriches the documented moss diversity of an already-diverse Early Cretaceous plant fossil assemblage. This is the third moss described from the Apple Bay plant fossil assemblage and represents the first occurrence of gemma cups in a fossil moss. It is also the oldest unequivocal record of Polytrichaceae, providing a hard minimum age for the group of 136 million years.

Cretaceous origin of dogwoods: an anatomically preserved Cornus (Cornaceae) fruit from the Campanian of Vancouver Island.

BACKGROUND: Cornaceae consists of 58 species, all within the genus Cornus. The Cenozoic record of Cornus is extensive and well documented. Molecular divergence-time studies suggest that crown-group Cornus may have originated by the Late Cretaceous. However, there has been no formal report of Cornus from Cretaceous deposits. Here, we characterize a permineralized fossil fruit assignable to Cornus subg. Cornus from the Upper Cretaceous (Campanian) Shelter Point locality of Vancouver Island, British Columbia, Canada. METHODS: Serial sections of the specimen were made using the cellulose acetate peel technique. Peels were mounted onto microscope slides and studied by light microscopy. RESULTS: The fossil fruit consists of a tri-locular woody endocarp with dorsal germination valves. The locules are sub-triangular to ellipsoidal in transverse section and are separated by thin septa. Endocarp tissue consists of elongated and isodiametric sclereids and secretory cavities. Internal vascular tissue was not observed, but is interpreted to have been located along the outer periphery of the septa for some length, common in many cornalean taxa. There is one seed in each locule, one of which was found to have endosperm and a dicotyledonous embryo. DISCUSSION: Woody endocarps with germination valves, without central vascular bundles, and with one seed per locule are characteristic of several families within the order Cornales. The interpreted vascular pattern and presence of secretory cavities indicates that the fossil fruit is assignable to Cornus subg. Cornus. Comparative analysis suggests that the fossil is most similar to Cornus piggae, a species described from the Paleocene of North Dakota. This fossil is the first evidence of crown-group Cornaceae from the Cretaceous and sheds light on both the plesiomorphic fruit characters and the timing of the initial diversification of the family and basal asterid lineage, Cornales.

Anatomically preserved fossil cornalean fruits from the Upper Cretaceous of Hokkaido: Eydeia hokkaidoensis gen. et sp. nov.

PREMISE OF THE STUDY: The basal asterid clade Cornales radiated during the Late Cretaceous. However, our understanding of early evolutionary patterns and relationships remain obscure. New data from five permineralized fruits in calcareous concretions from the Upper Cretaceous (Coniacian-Santonian) Haborogawa Formation, Hokkaido, Japan provide anatomical details that aid our knowledge of the group. METHODS: Specimens were studied from cellulose acetate peels, and three-dimensional reconstructions were rendered using AVIZO. KEY RESULTS: Fruits are drupaceous, roughly pyriform, 2.9-4.3 mm in diameter, with a fleshy mesocarp, transition sclereids, and a stony endocarp of four to five locules, with the septa forming a cross or star-like pattern in transverse section, distinct germination valves, and one apically attached anatropous seed per locule. Vascular tissue occurs in zones between the mesocarp and exocarp, in two rows within the septa, and prominent seed bundles can be traced throughout the fruit sections. Seeds have a single integumentary layer of radially flattened square to rectangular cells and copious cellular endosperm. A fully formed, straight, cellular dicotyledonous embryo, with closely appressed, spathulate cotyledons, is present within each seed. CONCLUSIONS: The unique combination of characters shown by these fruits is found in Cornaceae, Curtisiaceae, and Davidiaceae and allows us to describe a new taxon of Cornales, Eydeia hokkaidoensis gen. et sp. nov., with many similarities to extant Davidia involucrata. These fossils underscore the phylogenetic diversification of Cornales that was underway during the Late Cretaceous and support the hypothesis that a Davidia-like fruit morphology is plesiomorphic within Cornales.

Phylogenetic diversification of Early Cretaceous seed plants: The compound seed cone of Doylea tetrahedrasperma.

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.

Exploring the fossil history of pleurocarpous mosses: Tricostaceae fam. nov. from the Cretaceous of Vancouver Island, Canada.

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).

Hughmillerites vancouverensis sp. nov. and the Cretaceous diversification of Cupressaceae.

UNLABELLED: • 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 Cunninghamia-like. 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.

Honeggeriella complexa gen. et sp. nov., a heteromerous lichen from the Lower Cretaceous of Vancouver Island (British Columbia, Canada).

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.

Seed cone anatomy of Cheirolepidiaceae (Coniferales): reinterpreting Pararaucaria patagonica Wieland.

PREMISE OF THE STUDY: Seed cone morphology and anatomy reflect some of the most important changes in the phylogeny and evolutionary biology of conifers. Reexamination of the enigmatic Jurassic seed cone Pararaucaria patagonica reveals previously unknown systematically informative characters that demonstrate affinities with the Cheirolepidiaceae. This paper documents, for the first time, internal anatomy for seed cones of this important extinct Mesozoic conifer family, which may represent the ghost lineage leading to modern Pinaceae. METHODS: Morphology and anatomy of cones from the Jurassic La Matilde Formation in Patagonia are described from a combination of polished wafers and thin section preparations. New photographic techniques are employed to reveal histological details of thin sections in which organic cell wall remains are not preserved. Specific terminology for conifer seed cones is proposed to help clarify hypotheses of homology for the various structures of the cones. KEY RESULTS: Specimens are demonstrated to have trilobed ovuliferous scale tips along with a seed enclosing pocket of ovuliferous scale tissue. Originally thought to represent a seed wing in P. patagonica, this pocket-forming tissue is comparable to the flap of tissue covering seeds of compressed cheirolepidiaceous cones and is probably the most diagnostic character for seed cones of the family. CONCLUSIONS: Pararaucaria patagonica is assigned to Cheirolepidiaceae, documenting anatomical features for seed cones of the family and providing evidence for the antiquity of pinoid conifers leading to the origin of Pinaceae. A list of key morphological and anatomical characters for seed cones of Cheirolepidiaceae is developed to facilitate assignment of a much broader range of fossil remains to the family. This confirms the presence of Cheirolepidiaceae in the Jurassic of the Southern Hemisphere, which was previously suspected from palynological records.

A lower Cretaceous (Valanginian) seed cone provides the earliest fossil record for Picea (Pinaceae).

PREMISE OF STUDY: Sequence analyses for Pinaceae have suggested that extant genera diverged in the late Mesozoic. While the fossil record indicates that Pinaceae was highly diverse during the Cretaceous, there are few records of living genera. This description of an anatomically preserved seed cone extends the fossil record for Picea A. Dietrich (Pinaceae) by ∼75 Ma. METHODS: The specimen was collected from the Apple Bay locality of Vancouver Island (Lower Cretaceous, Valanginian) and is described from anatomical sections prepared using cellulose acetate peels. Cladistic analyses of fossil and extant pinaceous seed cones employed parsimony ratchet searches of an anatomical and morphological matrix. KEY RESULTS: This new seed cone has a combination of characters shared only with the genus Picea A. Dietr. and is thus described as Picea burtonii Klymiuk et Stockey sp. nov. Bisaccate pollen attributable to Picea is found in the micropyles of several ovules, corroborating the designation of this cone as an early spruce. Cladistic analyses place P. burtonii with extant Picea and an Oligocene representative of the genus. Furthermore, our analyses indicate that Picea is sister to Cathaya Chun et Kuang, and P. burtonii helps to establish a minimum date for this node in hypotheses of conifer phylogeny. CONCLUSIONS: As an early member of the extant genus Picea, this seed cone extends the fossil record of Picea to the Valanginian Stage of the Early Cretaceous, ca. 136 Ma, thereby resolving a ghost lineage predicted by molecular divergence analyses, and offers new insight into the evolution of Pinaceae.

The seed cone Eathiestrobus gen. nov.: fossil evidence for a Jurassic origin of Pinaceae.

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.