Ruth A. Stockey

A new fossil species in the moss family Tricostaceae, Tricosta angeiophoros sp. nov. (Early Cretaceous, Valanginian, ca. 136 Ma), possesses characters (stem and costa conducting strands) that prompt revision of the familial and generic diagnoses. In turn, discovery of this new species motivates a broader re‐examination of the relationships of tricostate mosses and the significance of the tricostate condition, which we undertake here. Tricostate mosses are an entirely extinct group known exclusively from gametophytes possessing leaves with unique morphology, characterized by a central costa and two symmetric lateral costae that originate separately at the leaf base. The stratigraphic range of tricostate mosses spans the Mesozoic and they are known from eastern Asia and western North America. The group now includes seven species: four species included in the two genera of the family Tricostaceae and three species included in the genus Tricostium. Whereas the Tricostaceae are unequivocally a hypnanaean family, likely belonging to the order Hypnales, the affinities of Tricostium remain obscure, due to a lack of family‐level diagnostic characters. Morphological characters available in the fossils are insufficient for distinguishing Tricostium from the Tricostaceae unequivocally. Uncertainty about the relationships among tricostate mosses and their position in the broader moss systematics leaves unanswered the question of whether tricostate leaves, which are known only in these extinct mosses, have a single origin or have evolved convergently in different groups. The answer to this question also bears on the role of adaptive change in the evolution of tricostate leaves. Taken together, the habitats inferred for tricostate mosses and those that favor their closest living analogs suggest that tricostate leaves may have evolved in response to the selective pressures of variable rheophytic conditions, to minimize leaf damage due to friction and to desiccation‐induced shrinkage. Aside from the evolutionary origins of the tricostate condition and independent tests for the hypothesized drivers of tricostate leaf evolution, another major unanswered question concerns the causes and timing of the post‐Cretaceous extinction of tricostate mosses. The fossil record is likely to provide answers to many of the questions concerning the tricostate condition as additional tricostate mosses are currently under study and fossiliferous strata with good potential for preserving bryophyte fossils have been identified and will be targeted by future studies.

Summary: Fossil woods are preserved as calcareous permineralizations in the Upper Cretaceous Northumberland Formation on Collishaw Point, Hornby Island, British Columbia, Canada. They are well preserved, often including peripheral tissues, and reveal the diversity and structure of vegetation in the region ca. 72 million years ago. We describe a new genus and species based on two specimens. Two specimens were cut using a rock saw to expose the fossil wood in transverse and longitudinal sections. Thin sections were prepared using the cellulose acetate peel technique. Peels were mounted on slides with Eukitt mounting medium. Anatomical description follows the IAWA guidelines. We compared the anatomy of the fossils with that of other plants using data available in the published literature and the InsideWood database. The combination of wide vessels and high vessel frequency given the narrow diameter of the stem, combined with the abrupt increase in vessel diameter near the pith indicates this plant had hydraulic properties typical of a climbing plant (liana). The anatomy of the rays, the arrangement of the vessels and their pitting, and the thickness of the bark are also consistent with the climbing habit, but not diagnostic of any extant family. This is the second species of liana to be described from the Hornby Island assemblage, providing new information about the composition of structurally complex mixed conifer–angiosperm forests in the Late Cretaceous of western North America.

Anatomically preserved lycophytes of the Lycopodiales and Selaginellales have been discovered among a diverse assemblage of plants and fungi in carbonate marine concretions at the Apple Bay locality along the shore of Holbert Inlet near the northern end of Vancouver Island, British Columbia, Canada. Lycopodialean stems are plectostelic and actinostelic, branch dichotomously, and are similar to both Lycopodicaulis oellgaardii and Lycoxylon spp. The Selaginella specimens represent the first anatomically preserved Selaginellales with excellent internal cellular preservation in the fossil record, and are described as Selaginella quatsinoense Rothwell et Stockey sp. nov. Stems have three and five exarch, monarch stelar segments, each of which is surrounded by an aerenchymatous endodermis with trabeculae. The leaf base is indented on the adaxial surface, suggesting the position of a ligule. These fossils document that species with diagnostic internal anatomy of modern Lycopodiales and Selaginellales evolved no later than the Valanginian of the early Cretaceous.

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.

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.

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.

Premise: Fossils show that fly‐speck fungi have been reproducing with small, black thyriothecia on leaf surfaces for ~250 million years. We analyzed morphological characters of extant thyriothecial fungi to develop a phylogenetic framework for interpreting fossil taxa.Methods: We placed 59 extant fly‐speck fungi in a phylogeny of 320 Ascomycota using nuclear ribosomal large and small subunit sequences, including newly determined sequences from nine taxa. We reconstructed ancestral character states using BayesTraits and maximum likelihood after coding 11 morphological characters based on original observations and literature. We analyzed the relationships of three previously published Mesozoic fossils using parsimony and our morphological character matrix, constrained by the molecular phylogeny.Results: Thyriothecia evolved convergently in multiple lineages of superficial, leaf‐inhabiting ascomycetes. The radiate and ostiolate scutellum organization is restricted to Dothideomycetes. Scutellum initiation by intercalary septation of a single hypha characterizes Asterinales and Asterotexiales, and initiation by coordinated growth of two or more adjacent hyphae characterizes Aulographaceae. Scutella in Microthyriales are initiated apically on a lateral hyphal branch. Patterns of hyphal branching in scutella contribute to distinguishing among orders. Parsimony resolves three fossil taxa as Dothideomycetes; one is further resolved as a member of a Microthyriales‐Zeloasperisporiales clade within Dothideomycetes.Conclusions: This is the most comprehensive systematic study of thyriothecial fungi and their relatives to date. Parsimony analysis of the matrix of character states of modern taxa provides an objective basis for interpreting fossils, leading to insights into morphological evolution and geological ages of Dothideomycetes clades.

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.

Premise of research. Five permineralized specimens of infructescences and dispersed fruits assignable to the Altingiaceae have been identified from the Late Cretaceous (Coniacian) Eden Main Quarry on Vancouver Island, British Columbia, Canada. Two infructescences and three isolated two-loculed fruits are preserved in calcium carbonate marine concretions. Methodology. Concretions were sectioned using the cellulose acetate peel technique and were included in a phylogenetic analysis employing morphological characters using TNT. Pivotal results. Infructescences are globose ca. 1 cm in diameter with at least 24 two-loculed fruits and a broad central axis. We have been unable to identify any perianth parts, but there are knob-like extrafloral processes similar to the phyllomes of other altingiaceous infructescences. The inner carpel lining consists of a single palisade layer of elongate rectangular cells. The endocarp also contains several layers of thick-walled, elongate sclerenchyma. The mesocarp is well differentiated, with fiber bundles interspersed with parenchyma and resin canals. There are at least 16 seeds (ovules) per carpel with axile placentation, but most of these are immature or aborted. Fruits have septicidal and ventricidal dehiscence. Specimens are assigned to Protoaltingia comoxense Scharfstein, Stockey & Rothwell gen. et. sp. nov. Conclusions. These fossils represent the second-most ancient occurrence of Altingiaceae, and phylogenetic analysis of morphological characters confirms that P. comoxense is most similar to species of Altingia. Results of morphological analyses resolve Altingia and Liquidambar as sister clades, placing P. comoxense as the basal member of the Altingia clade and sister to Altingia spp., thus documenting that evolutionary diversification of the core eudicot family Altingiaceae (Saxifragales) was well underway by the mid–Upper Cretaceous.

PREMISE OF THE STUDY: Fossils show that fly-speck fungi have been reproducing with small, black thyriothecia on leaf surfaces for ∼250 million years. We analyze morphological characters of extant thyriothecial fungi to develop a phylogenetic framework for interpreting fossil taxa. METHODS: We placed 59 extant fly-speck fungi in a phylogeny of 320 Ascomycota using nuclear ribosomal large and small subunit sequences, including newly determined sequences from nine taxa. We reconstructed ancestral character states using BayesTraits and maximum likelihood after coding 11 morphological characters based on original observations and literature. We analyzed the relationships of three previously published Mesozoic fossils using parsimony and our morphological character matrix, constrained by the molecular phylogeny. KEY RESULTS: Thyriothecia evolved convergently in multiple lineages of superficial, leaf- inhabiting ascomycetes. The radiate and ostiolate scutellum organization is restricted to Dothideomycetes. Scutellum initiation by intercalary septation of a single hypha characterizes Asterinales and Asterotexiales, and initiation by coordinated growth of two or more adjacent hyphae characterizes Aulographaceae (order incertae sedis ). Scutella in Microthyriales are initiated apically on a lateral hyphal branch. Patterns of hyphal branching in scutella contribute to distinguishing among orders. Parsimony resolves three fossil taxa as Dothideomycetes; one is further resolved as a member of a Microthyriales-Zeloasperisporiales clade within Dothideomycetes. CONCLUSIONS: This is the most comprehensive systematic study of thyriothecial fungi and their relatives to date. Parsimony analysis of the matrix of character states of modern taxa provides an objective basis for interpreting fossils, leading to insights into morphological evolution and geological ages of Dothideomycetes clades.