Jason Hilton

Jason Hilton
  • BSc (Sheffield) PhD (Cardiff)
  • Professor at University of Birmingham

Research on palaeobotany and palaeoenvironments through geological time

About

188
Publications
100,670
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Introduction
Jason is a research focused palaeobotanist and evolutionary plant biologist with broad interests in the consequences of plant evolution in deep time on geological phenomena including climate and environmental change. Research addresses periods of plant innovation and radiation, diversity changes, mass extinctions, geo-engineering and the co-evolution of plant life and the environment.
Current institution
University of Birmingham
Current position
  • Professor
Additional affiliations
September 2003 - April 2021
University of Birmingham
Position
  • Various
Description
  • Jason does stuff. Lots of different stuff, really. He is not actively looking for more stuff.
January 2000 - October 2003
National Museums Scotland
Position
  • Curator of Palaeobotany
May 1997 - December 1998
Chinese Academy of Sciences
Position
  • Researcher

Publications

Publications (188)
Article
The Early Devonian witnessed a major phase in the terrestrialization of land by plants. Understanding the implications for organic matter formation and accumulation during this interval is key to understanding global carbon burial. Existing research on the Early Devonian primarily focuses on marine carbonate records that do not permit evaluation of...
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The Permian-Triassic mass extinction (PTME) is the most severe biological crisis in Earth history and is closely linked to massive contemporaneous volcanism. However, there is currently limited evidence of Mercury (Hg) enrichment directly from volcanic sources in terrestrial strata, necessitating evidence from different regions and latitudes to con...
Article
Premise of research. Reconstructed whole plants provide high-quality information on extinct plant species comparable to the morphological and anatomical information available from living plants but are rarely recognized from the fossil record. On the basis of fossils preserved under a blanket of ash in the earliest Permian-aged Wuda Tuff Flora, Inn...
Article
A marattialean fern with organically connected fertile and vegetative parts is described from the early Permian Wuda Tuff Flora in Inner Mongolia, China. The tripinnate fronds have typical pecopterid vegetative pinnules but do not match any known Pecopteris species. The fertile pinnules are distinctly different from the vegetative ones, with an ext...
Article
The recognition of plant biogeographic ranges over time and migration routes is crucial for understanding ancient global plate tectonic configurations. This paper deals with a new fossil plant from the uppermost part of the Taiyuan Formation (earliest Asselian) in the Wuda Coalfield, Inner Mongolia, China. It has a taeniopterid morphology comprisin...
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Frequent wildfires associated with emplacement of the Central Atlantic Magmatic Province (CAMP) are thought to have been important drivers of two significant changes in terrestrial plant communities and diversity during the Triassic-Jurassic Mass Extinction (TJME, ca. 201.51 Ma). However, it remains to be investigated whether these two changes are...
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The arid climate of the Late Triassic was interrupted by a particularly humid episode known as the Carnian Pluvial Episode (CPE; ca. 234–232 million years ago). The CPE is often linked to eruptions in the Wrangellia Large Igneous Province (LIP), and is assumed to have led to global warming, enhanced weathering, water deoxygenation, and biotic chang...
Article
The formation and accumulation of lacustrine organic-rich shales are strongly related to palaeoclimate and palaeoenvironmental conditions. Middle Jurassic organic-rich lacustrine shales in the Qaidam Basin, NW China, are important source rocks for unconventional oil and gas exploration, but factors controlling their organic matter (OM) abundance an...
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The Late Triassic Carnian Pluvial Episode (CPE; ca. 234–232 Ma) was characterized by dramatic global temperature and humidity increases, which in many terrestrial settings was accompanied by changes from arid to humid vegetation types. This study reviews current evidence of terrestrial floral composition and distribution during the CPE and analyzes...
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The deep-time geological record can provide insights into the processes and mechanisms of glacier retreat. Ice sheets of the Late Paleozoic Ice Age (LPIA) collapsed extensively during the early Artinskian (early Permian) approximately 290 million years ago through massive glacial melting that was associated with dramatic increases in global tempera...
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The Permian-Triassic Mass Extinction (PTME) is the greatest biotic crisis of the Phanerozoic. In terrestrial settings, the PTME appears to have been diachronous and it has been suggested that losses initiated before the marine crisis. We examine organic carbon-isotope (δ¹³Corg) and geochemical proxies for environmental change in a palaeotropical we...
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The breakup of Pangaea and the rapid opening of the Ligurian and Central Atlantic oceans during the Middle Jurassic resulted in widespread volcanism accompanied by significant shifts in global environments, climates, and floras. Although major volcanism is a plausible driver of such global changes, linking these phenomena in the Middle Jurassic is...
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The Permian-Triassic Mass Extinction (ca. 252 Ma; PTME) is the most severe biocrisis of the Phanerozoic in both the oceans and on land. The crisis saw the collapse of terrestrial ecosystems in low, mid and high latitudes. Although terrestrial plant losses have been implicated as a driver of concurrent changes in terrestrial sedimentary environments...
Article
The global pattern of plant evolution through the Permian−Triassic mass extinction is uncertain, and the extent to which land plants were affected is debated. Detailed studies undertaken at a regional scale can help evaluate this floral transition, and thus we provide a detailed account of floral evolution from the Permian to Middle Triassic of Nor...
Article
Green-grey coloured reduction spots are common in continental red beds through geological history and occur in a range of different lithologies and depositional environments, but their timing and mode of formation remain controversial. We investigate the Late Cretaceous to earliest Paleogene Jiaozhou Formation using borehole data from the Jiaolai B...
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The Late Paleozoic Ice Age (LPIA; ca. 335–260 million years ago) was one of the most significant glacial events in Earth’s history. It records cycles of ice advance and retreat in southern high-latitude Gondwana and provides a deep-time perspective for climate–glaciation coevolution. However, climate records using clay mineral proxies from the LIPA...
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Initial breakup of Pangaea and opening of the Central Atlantic Ocean during the Early Jurassic resulted in widespread volcanism and was accompanied by significant changes in atmospheric composition, climate and environment of the Earth's surface system. Although profound changes in global geochemical cycles, palaeoclimate, and palaeoenvironments du...
Article
Wildfires are an important source of disturbances in the Earth's system and are of great significance for understanding the interactions between environmental, atmospheric and vegetation changes over deep time. The early Cretaceous was a “high-fire” interval with frequent and widespread wildfires globally, but the timing and global significance of...
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The End-Triassic Mass Extinction (ETME) saw the catastrophic loss of ca. 50% of marine genera temporally associated with emplacement of the Central Atlantic Magmatic Province (CAMP). However, the effects of the ETME on land is a controversial topic. Evaluation of the disparate cause(s) and effects of the extinction requires additional, detailed ter...
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The Permian-Triassic Mass Extinction (PTME) is the greatest biodiversity crisis in Earth history and while the marine crisis is increasingly well constrained, the timing and cause(s) of terrestrial losses remain poorly understood. There have been suggestions that the End- Permian Terrestrial Collapse (EPTC) pre-dated, was synchronous with or post-d...
Article
The link between the Permian–Triassic mass extinction (252 million years ago) and the emplacement of the Siberian Traps Large Igneous Province (STLIP) was first proposed in the 1990s. However, the complex cascade of volcanically driven environmental and biological events that led to the largest known extinction remains challenging to reconstruct. I...
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Gigantopterid plants share common traits of megaphyllous leaves with multi-ordered venation and have a stratigraphic distribution restricted to the Permian Period. They display a large variety of leaf morphologies which may indicate affinities from more than one plant groups including ferns and pteridosperms such as the peltasperms. Here we describ...
Article
The extreme rarity of soft-tissue preservation in ammonoids has meant there are open questions regarding fundamental aspects of their biology. We report an exceptionally preserved Middle Jurassic ammonite with unrivaled information on soft-body organization interpreted through correlative neutron and X-ray tomography. Three-dimensional imaging of m...
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The Wuda Tuff Flora from the Taiyuan Formation of Inner Mongolia is a palaeobotanical lagerstätte that is key to our current understanding of the composition and ecology of the earliest Permian Cathaysian floras. We investigate an anatomically preserved axis that represents an additional plant family recorded in this flora and considers its palaeob...
Article
Alethopteris ascendens Halle is an endemic element in the Permian Cathaysian floras and is assumed to be a member of the Medullosales based on its alethopterid type foliage. Previous accounts have documented only its pinnae and bipinnate fronds, but other parts of the plant are unknown. Here we describe new specimens from the earliest Permian (Asse...
Preprint
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The Late Paleozoic Ice Age (LPIA; ca. 360–260 million years ago) was one of the most significant glacial events in Earth history that records cycles of ice advance and retreat in southern high-latitude Gondwana and provides a deep-time perspective for climate-glaciation coevolution. However, climate records from the LIPA are poorly understood in lo...
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Significance The Carnian Stage of the Triassic Period marks one of the most significant intervals of the past 250 My. Within the space of ∼2 My, the world’s biota underwent major changes with dinosaurs becoming the notable incumbents. These events coincide with a remarkable interval of intense rainfall known as the Carnian Pluvial Episode (CPE). He...
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The Late Paleozoic Icehouse (LPI) provides a deep-time perspective for climate-glaciation-environment coevolution and offers potential insights into future climatic and environmental predictions. Most previous studies attributed climatic and environmental changes during the LPI to perturbations of atmospheric pCO2, yet the driving mechanism for pCO...
Preprint
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Ensuring taught fieldwork is a positive, generative, collective, and valuable experience for all participants requires considerations beyond course content. To guarantee safety and belonging, participants’ identities (backgrounds and protected characteristics) must be considered as a part of fieldwork planning and implementation. Furthermore, getti...
Article
Determining the diversity of past floras helps with interpreting both the history and predicting the future of vegetation change. For global-scale and regional-scale diversity studies especially, secondary data are often used but local-scale studies tend to be based on survey data that require rigorous sampling. The correct sampling strategies depe...
Article
A new species of the Marattialean fern frond Rothwellopteris is proposed for specimens preserving both morphology and anatomy from the Wuchiapingian–Changshingian (late Permian) aged Xuanwei Formation of South China. Fronds are at least bipinnate and bear falcate, pecopteroid pinnules with a thin lamina. The penultimate pinna rachis has a polycycli...
Article
The purpose of this research was to examine paleoenvironments, hydrothermal activity, and seawater restriction of the lower Cambrian Niutitang Formation shale gas reservoir in the eastern Xuefeng uplift and to determine factors affecting organic matter (OM) enrichment. In the studied borehole Xiangan 1 well in western Hunan Province, the Niutitang...
Article
Palaeobotany and palynology are the main direct sources of evidence for studying vegetation diversity dynamics through geological time. However, plant fossil diversity is affected by various factors other than vegetation diversity, which need to be taken into account in such studies. The use of fossil-taxa will potentially inflate perceived plant d...
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Significance There were two heterosporous lignophyte lineages of which only one, the seed plants, survived the Permian–Triassic mass extinction. Based on exceptionally complete fossil trees from a 300-My-old volcanic ash, the enigmatic Noeggerathiales are now recognized as belonging to the other lineage. They diversified alongside the primary seed...
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The late Paleozoic ice age (LPIA) during the Carboniferous and Permian (ca. 360–260 Ma) was vegetated Earth's only recorded icehouse-to-greenhouse transition and provides a deep-time perspective for climate-glaciation-environment coevolution and future climate change. During the LPIA, changes in glacial-interglacial cycles have been closely linked...
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The origin of angiosperms is a classic macroevolutionary problem, because of their rapid rise in the Early Cretaceous fossil record, beginning about 139 Ma ago, and the conflict this creates with older crown-group ages based on molecular clock dating. Silvestro et al. use a novel methodology to model past angiosperm diversity based on a Bayesian Br...
Article
This paper investigates the taxonomic diversity of marattialean stems from the family Psaroniaceae preserved in-situ within the Wuda Tuff Flora. Three species of the permineralized stem Psaronius and eight species of the compression/impression stem Caulopteris are documented, but it has not been possible to associate the anatomically preserved spec...
Article
The Miocene aged Shanwang Formation from the Shanwang National Geopark in China represents a succession of lacustrine diatomaceous shales containing an abundant and diverse biota with lagerstätte fossilization of soft tissues. To date, the Shanwang Formation has not been investigated for cyclostratigraphy nor has it been dated with high precision m...
Article
Hydrasperman pteridosperms (seed ferns) were widespread and diverse in Carboniferous equatorial wetlands of Euramerica, and although present in Permian equatorial wetlands in Cathaysia, little is known about their ovules (unfertilised seeds) from this phytogeographical realm. Hydrasperman ovules are recognisable from their distinctive pollen chambe...
Article
The late Paleozoic genera Chansitheca Regè, Oligocarpia Göppert and Szea Z. Yao et T. N. Taylor are small ferns that represent putative early members of the Gleicheniaceae based on their morphology and the anatomy of their fertile organs. However, the rachis and cauline anatomy are unknown, rendering their systematic affinities controversial. Here...
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This study focuses on Early Cretaceous mudstones from the Shahai and Fuxin formations in the Fuxin continental basin. We analyse chemical weathering, land surface temperatures and palaeoclimates based on chemical weathering indices, and emphasize the implications of continental chemical weathering on nutrient fluxes into lakes and oceans. According...
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The Early Jurassic break-up of Pangea and the formation of the Central Atlantic Ocean are associated with changes to the atmosphere, climate, and environment, as recorded in marine carbonates from the western Tethys Ocean. However, the expression of these global changes in terrestrial successions is less well known. To better understand terrestrial...
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How plant seeds originated remains unresolved, in part due to disconnects between fossil intermediates and developmental genetics in extant species. The Carboniferous fossil Genomosperma is considered among the most primitive known seeds, with highly lobed integument and exposed nucellus. We have used this key fossil taxon to investigate the evolut...
Article
Climbing plants are important components of modern-day tropical forests and primarily comprise angiosperms, but in Paleozoic tropical forests, pteridosperms were the dominant climbers. Climbing pteridosperms are well known from the Paleozoic floras of Euramerica, while much less is known about climbing pteridosperms from other phytogeographic regio...
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The late Permian was the acme of Pangea assembly, with collision and subduction of global plates accompanied by major changes in atmospheric composition, paleoclimates and paleoenvironments of the Earth's surface system. These events are extensively recorded in marine successions from the Tethys, but much less are known from continental successions...
Article
The early Cambrian (541–514 Ma) was a crucial interval for the evolution of life on Earth, popularly known as the “Cambrian Explosion”. Here, we report the timing of changes in hydrothermal and depositional inputs, as well as paleo-redox state, which may have influenced biogeochemical changes. According to high-resolution petrology, fossil distribu...
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The Permo-Triassic marine mass extinction has been blamed on a range of culprits including anoxia, acidification, high temperature and increased sedimentation and nutrient influx, the last two being a direct consequence of terrestrial biomass die-off and climatic changes. In marine settings, the role of these kill mechanisms is likely to be depth-d...
Article
Much of our existing knowledge of Middle Jurassic paleoclimate is based on well-dated marine isotopic records that show fluctuations between warm (greenhouse effected) and cool climates. In contrast, much less is known from contemporaneous terrestrial deposits that are often difficult to correlate stratigraphically with marine successions, and are...
Article
We describe a new species of the Paleozoic fern genus Botryopteris in volcanic tuffs from the Lopingian (upper Permian) aged Junlian Formation in SW China's Sichuan Province. The species has a large stem and stele compared to those of some other species of the genus. Xylem strands of the leaf trace and rachis are “ω”-shaped in cross section and com...
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Premise of research. Mississippian (Lower Carboniferous) anatomically preserved ovules are pivotal to our present understanding of the Paleozoic primary seed plant radiation, but few are known from the late Viséan stratigraphic interval approximately 330 million years ago. Here, we document an exceptionally well-preserved mesoscopic charcoalified o...
Article
Coal seams preserve high‐resolution records of ancient terrestrial water table (base level) fluctuations in ancient peat accumulations, but little is known about base level change in anomalously thick coal seams. Using the Early Cretaceous 91 m anomalously thick No. 6 coal (lignite) seam in the Erlian Basin (north‐east China) as a case study, the o...
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The timing of the origin of angiosperms is a hotly debated topic in plant evolution. Molecular dating analyses that consistently retrieve pre‐Cretaceous ages for crown‐group angiosperms have eroded confidence in the fossil record, which indicates a radiation and possibly also origin in the Early Cretaceous. Here, we evaluate paleobotanical evidence...
Article
Peat formed in mire settings sensitively records environmental fluctuations during deposition including changes in water table or base level and accommodation. On this basis coal seams, as geologically preserved peats, can provide evidence of high-resolution paleoclimatic fluctuations as well as paleobotanical evolution through periods of peat-form...
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We have investigated the geologic features of the lower Cambrian-aged Niutitang Shale in the northwestern Hunan province of South China. Our results indicate that the Niutitang Shale has abundant and highly mature algal kerogen with total organic carbon (TOC) content ranging from 0.6% to 18.2%. The equivalent vitrinite reflectance (equal-Ro) value...
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As one of the most important source rocks and reservoirs of unconventional natural gas, the sedimentary environment and mode of peat swamp (the predecessor of coal seam) is important to the coal seam’s spatial distribution, material composition, hydrocarbon generation potential, reservoir physical properties, etc. To reveal the depositional charact...
Article
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Most knowledge concerning Mesozoic Era floras has come from compression fossils. This has been augmented in the last 20 years by rarer permineralized material showing cellular preservation. Here, we describe a new genus of anatomically preserved gymnosperm seed from the Callovian–Oxfordian (Jurassic) Oxford Clay Formation (UK), using a combination...
Article
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Premise of the study: Noeggerathiales are an extinct group of heterosporous shrubs and trees that were widespread and diverse during the Pennsylvanian-Permian Epochs (323-252 Ma) but are of controversial taxonomic affinity. Groups proposed as close relatives include leptosporangiate ferns, sphenopsids, progymnosperms, or the extant eusporangiate f...
Article
In their study of charred conifer twigs from the Lower Cretaceous Chaswood Formation of Canada, Falcon-Lang et al. (2016) established the species Pinus mundayi that they interpreted as a “two needled” pine and the oldest stratigraphic evidence for the extant genus Pinus. This is based on their interpretation of what they thought was distinctive woo...
Article
Plant macrofossil remains of Becklesia Seward, 1895 emend. Watson and Cusack, 2005, a rare genus of extinct cycad, have been known for over a century from Great Britain and Libya. Historically the genus was poorly characterized leading to many fossil cycads that resembled Becklesia being placed in other genera including Cycadites and Paracycas. We...
Article
Late Permian (Lopingian) volcanoclastic lithologies from the Huopu Mine near Fuyuan, Guzihou Province, SW China have yielded konservat lagerstätte-grade plant macrofossils. These fossils derive from a stratigraphic interval bounded by the mid-Capitanian extinction below and the end Permian extinction above and globally, few anatomically preserved f...
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Background and Aims. The largely Mississippian strata of the Kilpatrick Hills, located at the western end of the Scottish Midland Valley, enclose several macrofossil floras that together contain ca 21 organ-species of permineralised plants and ca 44 organ-species of compressed plants, here estimated to represent 25 whole-plant species (Glenarbuck =...
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• Triassic and Jurassic fossils record structural changes in conifer seed cones through time, provide the earliest evidence for crown-group conifer clades, and further clarify sister-group relationships of modern conifer families. A new and distinct seed-cone from the Isle of Skye in western Scotland provides the oldest detailed evidence for the an...
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Diminutive, silica-permineralized lycopsid axes, from a Guadalupian (Middle Permian) silicified peat in the Bainmedart Coal Measures of East Antarctica are described and assigned to Paurodendron stellatum sp. nov. Axes consist only of primary-growth tissues with a vascular system characterized by an exarch actinostele with 6–20 protoxylem points. S...
Presentation
The Cheirolepidiaceae are an extinct group of conifers with an extensive Mesozoic fossil record well-documented by foliage, wood, pollen and reproductive organs. Three-dimensionally preserved Cheirolepidiaceous ovuliferous cones are rare, with few known species. Here we use X-Ray computed micro-tomography at the Diamond Light Source Synchrotron (Ox...
Article
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Male cones of Ixostrobus hailarensis from Lower Cretaceous strata of the Hailar region of north-east China were re-examined for features of their cuticle and in situ pollen. The cones are loosely aggregated and possess helically arranged peltate microsporophylls containing resin bodies. Microsporophyll cuticle is thick, papillate and has elliptical...
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The genus name Schimperstrobus was first used by Deng et al. (2014)1. Deng, S., Jason Hilton, J., Glasspool, I. J. & Dejax, J. 2014. Pollen cones and associated leaves from the Lower Cretaceous of China and a re-evaluation of Mesozoic male cycad cones. Journal of Systematic Palaeontology, http://dx.doi.org/10.1080/14772019.2013.819817[Web of Scienc...
Article
The Permian aged osmundalean fern Palaeosmunda yunnanense Tian et Chang has been re-investigated based on the original specimens and demonstrated to represent a new genus within the extinct family Guaireaceae that we name Tiania yunnanense (Tian et Chang) gen. et. comb. nov. The stem of T. yunnanense is small, c. 30 mm in diameter, and comprises an...
Article
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We document a new species of ovulate cone (Pararaucaria collinsonae) on the basis of silicified fossils from the Late Jurassic Purbeck Limestone Group of southern England (Tithonian Stage: ca. 145 million years). Our description principally relies on the anatomy of the ovuliferous scales, revealed through X-ray synchrotron microtomography (SRXMT) p...
Presentation
Full-text available
The Cheirolepidiaceae are an extinct group of conifers with an extensive Mesozoic fossil record well-documented by foliage, wood, pollen and reproductive organs. Three-dimensionally preserved Cheirolepidiaceous ovuliferous cones are rare, with few known species. Here we use X-Ray computed micro-tomography (XMT) at the Diamond Light Source Synchrotr...
Article
Full-text available
Based upon anatomical evidence, Permian aged gigantopterid fossils are in general reconstructed as climbing or scrambling plants. Gigantonoclea, a genus of adpressed gigantopterid foliage from the Permian of northern China, has been reported to co-occur with hook-like organs that were interpreted as indicating a scrambling/climbing habit. We reinve...
Article
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Reinvestigation of the anatomically preserved stem Palaeosmunda plenasioides from the Lopingian (Late Permian) of China has led to the establishment of Zhongmingella gen. nov. within the extinct family Guaireaceae (Osmundales). Zhongmingella has a rhizomatous stem with heterogeneous pith and cortex comprising parenchyma and uniformly distributed se...
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The Toploje Member chert is a Roadian to Wordian autochthonous–parautochthonous silicified peat preserved within the Lambert Graben, East Antarctica. It preserves a remarkable sample of terrestrial life from high latitude central Gondwana prior to the Capitanian mass extinction event from both mega- and microfossil evidence that includes cryptic co...
Article
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The strong positive relationship evident between cell and genome size in both animals and plants forms the basis of using the size of stomatal guard cells as a proxy to track changes in plant genome size through geological time. We report for the first time a taxonomic fine‐scale investigation into changes in stomatal guard‐cell length and use thes...
Article
Fossil lignophyte stems show remarkable diversity including non-sympodial and sympodial forms and a variety of pith and tracheid structures. Most lack scalariform bordered pits on radial walls of secondary xylem tracheids, while these occur in the putative progymnosperm Protopitys and some (but not all) cycads, bennettitaleans, and angiosperms. We...
Article
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A new species of permineralised marattialean fern trunk is described from Lopingian (late Permian) aged volcaniclastic tuffs from Panxian County, western Guizhou Province, China and named Psaronius xuii sp. nov. The stem has vascular bundles that are encircled by a bi-layered sheath and has a ground tissue consisting of thicker-walled and thinner-w...
Article
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A small but diverse fossil plant assemblage from Gardeshwar in Gujarat Province of western India is reinvestigated, based on analysis of recently collected specimens that represent previously unrecognised taxa in combination with a critical review of previously reported taxa from the site. The assemblage is dominated by conifers including Brachyphy...
Article
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The fossil record of Peronosporomycetes (water moulds) is rather sparse, though their distinctive ornamentation means they are probably better reported than some true fungal groups. Here we describe a rare Palaeozoic occurrence of this group from a Guadalupian (Middle Permian) silicified peat deposit in the Bainmedart Coal Measures, Prince Charles...
Article
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I. II. III. IV. V. VI. VII. VIII. References SUMMARY: We evaluate stomatal development in terms of its primary morphogenetic factors and place it in a phylogenetic context, including clarification of the contrasting specialist terms that are used by different sets of researchers. The genetic and structural bases for stomatal development are well co...
Article
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Premise of research. A third genus of anatomically preserved conifer seed cones has been recognized from a Late Jurassic deposit in northeastern Scotland. This cone is described as Bancroftiastrobus digitata Rothwell, Mapes, Stockey et Hilton. Methodology. The cone was sectioned with the classic coal ball peel technique and studied and photographed...
Article
Full-text available
Premise of research. Although the most comprehensively known Devonian seeds were borne in a “telomic” cupule, those of some species have been postulated as being borne terminally on naked axes lacking a cupule. Uncertainty remains as to whether such seeds were shed from a cupule before preservation. We reinvestigate the Upper Devonian fossils Pseud...

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