Zirabia cylindrica comb. nov. provides evidence of Doyleales in the Jurassic.

PREMISE: In recent years, Doyleales have played an important role in the discussion and exploration of seed plant relationships and the origin of angiosperms. This order comprises a diversity of recently described genera with cupule-bearing compound ovulate cones recovered from Early Cretaceous deposits in North America and Asia. Their relatively late appearance in the fossil record, stratigraphically near the appearance of angiosperms in the Early Cretaceous, has been noteworthy. Here, we report a new genus of Doyleales, Zirabia gen. nov. from the Early Jurassic of Iran, that was originally described as the ginkgophyte Karkenia. METHODS: We reinvestigated material previously assigned to Karkenia cylindrica from the Lower Jurassic of the Zirab locality, Alborz Mountains, northern Iran. RESULTS: The studied specimen features a main axis bearing helically to irregularly arranged bract-cupule complexes, each composed by a long laminar bract subtending and sheathing a cupule stalk that bears a single-seeded cupule with a dorsal protrusion. The morphological features of this taxon do not conform with those of Karkenia, and suggest affinities with Doyleales rather than Ginkgoales. Within Doyleales, this fossil has a unique combination of characters indicating that it is a new genus within the order; thus, a new combination is erected, Zirabia cylindrica. CONCLUSIONS: Our results indicate that Doyleales is significantly older than previously thought, with their stratigraphic range now extending from the Lower Jurassic to the Cretaceous. The Early Jurassic occurrence of Doyleales provides important data on the emergence of seed-enclosing structures seen in seed plants throughout the Mesozoic.

Cretaceous pollen cone with three-dimensional preservation sheds light on the morphological evolution of cycads in deep time.

The Cycadales are an ancient and charismatic group of seed plants. However, their morphological evolution in deep time is poorly understood. While molecular divergence time analyses estimate a Cretaceous origin for most major living cycad clades, much of the extant diversity is inferred to be a result of Neogene diversifications. This leads to long branches throughout the cycadalean phylogeny that, with few exceptions, have yet to be rectified by unequivocal fossil cycads. We report a permineralized pollen cone from the Campanian Holz Shale located in Silverado Canyon, CA, USA (c. 80 million yr ago). This fossil was studied via serial sectioning, SEM, 3D reconstruction and phylogenetic analyses. Microsporophyll and pollen morphology indicate this cone is assignable to Skyttegaardia, a recently described genus based on disarticulated lignitized microsporophylls from the Early Cretaceous of Denmark. Data from this new species, including a simple cone architecture, anatomical details and vasculature organization, indicate cycadalean affinities for Skyttegaardia. Phylogenetic analyses support this assignment and recover Skyttegaardia as crown-group Cycadales, nested within Zamiaceae. Our findings support a Cretaceous diversification for crown-group Zamiaceae, which included the evolution of morphological divergent extinct taxa with unique traits that have yet to be widely identified in the fossil record.

Icacinaceae fossil provides evidence for a Cretaceous origin of the lamiids.

Today the asterids comprise over 80,000 species of flowering plants; however, relatively little is known about the timing of their early diversification. This is particularly true for the diverse lamiid clade, which comprises half of asterid diversity. Here, a lamiid fossil fruit assigned to Icacinaceae from the Campanian of western North America provides important macrofossil evidence indicating that lamiids diverged at least 80 million years ago and sheds light on potential Cretaceous rainforest-like ecosystems.

Paleoclimate-induced stress on polar forested ecosystems prior to the Permian-Triassic mass extinction.

The end-Permian extinction (EPE) has been considered to be contemporaneous on land and in the oceans. However, re-examined floristic records and new radiometric ages from Gondwana indicate a nuanced terrestrial ecosystem response to EPE global change. Paleosol geochemistry and climate simulations indicate paleoclimate change likely caused the demise of the widespread glossopterid ecosystems in Gondwana. Here, we evaluate the climate response of plants to the EPE via dendrochronology snapshots to produce annual-resolution records of tree-ring growth for a succession of late Permian and early Middle Triassic fossil forests from Antarctica. Paleosol geochemistry indicates a shift in paleoclimate towards more humid conditions in the Early and early Middle Triassic relative to the late Permian. Paleosol morphology, however, supports inferences of a lack of forested ecosystems in the Early Triassic. The plant responses to this paleoclimate change were accompanied by enhanced stress during the latest Permian as determined by high-resolution paleoclimate analysis of wood growth intervals. These results suggest that paleoclimate change during the late Permian exerted significant stress on high-latitude forests, consistent with the hypothesis that climate change was likely the primary driver of the extinction of the glossopterid ecosystems.

Extending beyond Gondwana: Cretaceous Cunoniaceae from western North America.

Cunoniaceae are important elements of rainforests across the Southern Hemisphere. Many of these flowering plants are considered Paleo-Antarctic Rainforest Lineages that had a Gondwanan distribution since the Paleocene. Fossils of several modern genera within the family, such as Ceratopetalum, have indicated biogeographical connections between South America and Australia in the Cenozoic. Here, we report a dramatic geographical range extension for Ceratopetalum, and Cunoniaceae as a whole, based on two exceptionally preserved fossil winged fruits from Campanian (c. 82-80 Ma old) deposits on Sucia Island, Washington, USA. The fossils were studied using physical sectioning, light microscopy, micro-computed tomography scanning and multiple phylogenetic analyses. The fossil fruits share diagnostic characters with Ceratopetalum such as the presence of four to five persistent calyx lobes, a prominent nectary disk, persistent stamens, a semi-inferior ovary and two persistent styles. Based on morphological comparisons with fruits of extant species and support from phylogenetic analyses, the fossils are assigned to a new species Ceratopetalum suciensis. These fossils are the first unequivocal evidence of crown Cunoniaceae from the Cretaceous of North America, indicating a more complicated biogeographical history for this important Gondwanan family.

Taxon sampling and alternative hypotheses of relationships in the euphyllophyte plexus that gave rise to seed plants: insights from an Early Devonian radiatopsid.

An abrupt transition in the fossil record separates Early Devonian euphyllophytes with a simple structure from a broad diversity of structurally complex Middle-Late Devonian plants. Morphological evolution and phylogeny across this transition are poorly understood due to incomplete sampling of the fossil record. We document a new Early Devonian radiatopsid and integrate it in analyses addressing euphyllophyte relationships. Anatomically preserved Emsian fossils (402-394 Ma) from the Battery Point Formation (Gaspé, Quebec, Canada) are studied in serial sections. The phylogenetic analysis is based on a matrix of 31 taxa and 50 characters emphasising vegetative morphology (41 discrete, nine continuous). The new plant, Kenrickia bivena gen. et sp. nov., is one of very few structurally complex euphyllophytes documented in the Early Devonian. Inclusion of Kenrickia overturns previously established phylogenetic relationships among Radiatopses, reiterating the need for increased density of Early Devonian taxon sampling. Kenrickia is recovered as the sister lineage to all other radiatopsids, a clade in which paraphyletic Stenokoleales led to a lignophyte clade where archaeopterids and seed plants fall into sister clades. Our results shed light on early euphyllophyte relationships and evolution, indicating early exploration of structural complexity by multiple lineages and reiterating the potential of a single origin of secondary growth in euphyllophytes.

Fossil evidence for a Cretaceous rise of the mahogany family.

PREMISE: The mahogany family (Meliaceae) is an angiosperm lineage comprising many species that are important elements in tropical ecosystems, and is often used as a study system to understand the evolution of tropical rainforests. While divergence time studies have estimated a Cretaceous origin for the family, no unequivocal fossils of that age have been described. Here, the first Cretaceous evidence for Meliaceae is reported, based on an exceptionally well-preserved fruit from the Upper Cretaceous (79-72 Ma, Campanian) of North America. METHODS: The fossil fruit was prepared using traditional paleobotanical techniques. Bayesian phylogenetic analyses using morphological and molecular data were conducted to assess the phylogenetic position of the Cretaceous fruit in Meliaceae and to assess the effect of morphology for inferring the overall pattern of phylogeny for the family. RESULTS: The fruit consists of a fleshy mesocarp and a woody endocarp with a hollow center, nine locules, loculicidal sutures, and one subapically attached seed per locule that has an enlarged sarcotesta near the hilum. The combination of characters in this fruit is strikingly similar to the genus Melia L. Phylogenetic analyses recover the Cretaceous fruit as being closely related to Melia and highlights the effect of fruit morphological data for inferring the overall pattern of phylogeny in Meliaceae. There are a few structural differences between the fossil fruit of this study and Melia; thus, the newly characterized Cretaceous taxon is named Manchestercarpa vancouverensis gen. et sp. nov. DISCUSSION: These results clearly confirm a Cretaceous origin for Meliaceae and that important tropical families were present prior to the development of modern tropical ecosystems in the Cenozoic.

Cretaceous asterid evolution: fruits of Eydeia jerseyensis sp. nov. (Cornales) from the upper Turonian of eastern North America.

BACKGROUND AND AIMS: The asterids (>80 000 extant species) appear in the fossil record with considerable diversity near the Turonian-Coniacian boundary (~90 Ma; Late Cretaceous) and are strongly represented in the earliest diverging lineage, Cornales. These early asterid representatives have so far been reported from western North America and eastern Asia. In this study, we characterize a new cornalean taxon based on charcoalified fruits from the upper Turonian of eastern North America, a separate landmass from western North America at the time, and identify early palaeobiogeographical patterns of Cornales during the Cretaceous. METHODS: Fossils were studied and imaged using scanning electron microscopy and micro-computed tomography (micro-CT) scanning. To assess the systematic affinities of the fossils, phylogenetic analyses were conducted using maximum parsimony. KEY RESULTS: The charcoalified fruits are represented by tri-locular woody endocarps with dorsal apically opening germination valves. Three septa intersect to form a robust central axis. Endocarp ground tissue consists of two zones: an outer endocarp composed of isodiametric sclereids and an inner endocarp containing circum-locular fibres. Central vasculature is absent; however, there are several small vascular bundles scattered within the septa. Phylogenetic analysis places the new taxon within the extinct genus Eydeia. DISCUSSION: Thick-walled endocarps with apically opening germination valves, no central vascular bundle and one seed per locule are indicative of the order Cornales. Comparative analysis suggests that the fossils represent a new species, Eydeia jerseyensis sp. nov. This new taxon is the first evidence of Cornales in eastern North America during the Cretaceous and provides insights into the palaeobiogeography and initial diversification of the order.

The critical role of fossils in inferring deep-node phylogenetic relationships and macroevolutionary patterns in Cornales.

PREMISE OF THE STUDY: The basal asterid order, Cornales, experienced a rapid radiation during the Cretaceous, which has made it difficult to elucidate the early evolution of the order using extant taxa only. Recent paleobotanical studies, however, have begun to shed light on the early diversification of Cornales. Herein, fossils are directly incorporated in phylogenetic and quantitative morphological analyses to reconstruct early cornalean evolution. METHODS: A morphological matrix of 77 fruit characters and 58 taxa (24 extinct) was assembled. Parsimony analyses including and excluding fossils were conducted. A fossil inclusive tree was time-scaled to visualize the timing of the initial cornalean radiation. Disparity analyses were utilized to infer the morphological evolution of cornaleans with drupaceous fruits. KEY RESULTS: Fossil inclusive and exclusive parsimony analyses resulted in well-resolved deep-node relationships within Cornales. Resolution in the fossil inclusive analysis is substantially higher, revealing a basal grade including Loasaceae, Hydrangeaceae, Hydrostachyaceae, Grubbiaceae, a Hironoia+Amersinia clade, and Curtisiaceae, respectively, that leads to a "core" group containing a clade comprising a Cretaceous grade leading to clade of Nyssaceae, Mastixiaceae, and Davidiaceae that is sister to a Cornaceae+Alangiaceae clade. The time-scaled tree indicates that the initial cornalean diversification occurred before 89.8 Ma. Disparity analyses suggest the morphological diversity of Cornales peaked during the Paleogene. CONCLUSIONS: Phylogenetic analyses clearly demonstrate that novel character mosaics of Cretaceous cornaleans play a critical role in resolving deep-node relationships within Cornales. The post-Cretaceous increase of cornalean disparity is associated with a shift in morphospace occupation, which can be explained from ecological and developmental perspectives.

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.

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.