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The abstract is available here: https://uscholar.univie.ac.at/o:1637808

Background Understanding the natural variability of plant resins is crucial for evaluating the chemical information stored in ambers that may support inferring palaeoenvironmental conditions. However, even among extant resin-producing plants, the variation of resinous exudates within and between tree genera and species is still poorly understood. Methods We analysed plant exudates from across the Araucariaceae in New Caledonia using Fourier-transform infrared spectroscopy in attenuated total reflection mode (FTIR-ATR). Both, wild (in situ) and arboretum grown (ex situ) populations were used to uncover the effects of intergeneric, inter- and intraspecific variation of the araucariacean plants on the resin chemistry. Results We show that even resins from the same species in the same habitat do have natural (intraspecific) chemical variation. Some of the variation can result from the degree of resin polymerisation, but this is not the only source of variation. Wild sourced resins have greater natural intraspecific chemical variation than the arboretum sourced ones. Interspecific and intergeneric differences were not easy to distinguish in the resins sampled. This has strong implications for the evaluation of the chemical information from worldwide ambers: multiple samples should be analysed to give a more accurate picture of the natural chemical variation present and how this may or may not overlap with the chemistries of other resin types. Additionally we discovered that the Araucariacean species can produce up to three distinct types of exudate (gum, resin, and gum resin). This is the first time that gum resins have been identified and characterized with FTIR-ATR. We also provide a guide on how to distinguish the exudate types when using FTIR-ATR.

This article presents re-investigated Crato Formation sample material from David J. Batten’s palynological legacy using a combination of light microscopy (LM) and scanning electron microscopy (SEM)...

The abstract is available here: https://uscholar.univie.ac.at/o:1090994

Here we present LM and SEM data of pollen of extant Sciadopitys verticillata, fossil Cerebropollenites from Aptian/Albian strata (Austria, Mongolia) and Sciadopityspollenites from Campanian/Maastrichtian and Oligocene/Miocene strata (Siberia, Germany). Measurements and image comparisons show that the investigated fossil pollen taxa range from somewhat comparable to very similar to extant Sciadopitys verticillata, and that a previous affiliation of Cerebropollenites taxa with Tsuga cannot be corroborated. Additionally, it can be speculated that either the Rhaetian to Lower Cretaceous Cerebropollenites taxa are the pollen equivalent of the Eurasian Miroviaceae macrofossils that have unresolved relationships with Sciadopityaceae, or that they might belong to a basal group in the Sciadopityaceae, which is quite recognizable due to the similarity of the pollen morphology of Cerebropollenites thiergartii and Sciadopitys verticillata. However, until in situ pollen within cones attributed to the Miroviaceae are found, we will not know for sure.

Species belonging to the Carboniferous genera Lepidodendron and Lepidophloios are well known from the coalfields of Pennsylvania (Lower Asturian (Westphalian D) substage of the Pennsylvanian) through the publications of Lesquereux and White. They believed that the lycophyte flora was rather distinct from other contemporaneous coal floras elsewhere due to its isolation. A full revision of the lepidodendroid lycophyte species is given here and this reassessment permitted comparisons with those floras known from the European and Canadian Maritimes coalfields. Differences in species distribution suggest there was a barrier (Arcadian Highlands) to species distribution to and from the coalfields of Pennsylvania. Several species are recognized only from the coalfields of Pennsylvania, although the number has been significantly reduced from the original publications.

Diverse temperate forest types and a high atmospheric humidity have recently been suggested for the Eocene source area of Baltic amber. However, ferns are astonishingly rare as inclusions in this amber, which is in contrast to other seed‐free land plants, fungi, and lichens. Moreover, the identities of some of the few described putative fern taxa are dubious, and some fossils were even assigned to the Paleozoic seed fern genera Alethopteris, Pecopteris and to the form genus Sphenopteris containing Paleozoic and Mesozoic fern‐like leaf fossils. Here, we review previously described fern inclusions from Baltic amber and identify further fern‐like leaf inclusions as belonging to the extant angiosperm genus Comptonia (sweet ferns, Myricaceae). We conclude that only one taxon, Matonia striata (Matoniaceae), can with confidence be identified as a Polypodiopsida representative. Although “Pecopteris” humboldtiana is so far only known as sterile foliage, its leaf morphology strongly suggests that also this taxon belongs to the Polypodiopsida rather than to any other tracheophyte lineage. We propose accommodating “Pecopteris” humboldtiana in the new genus Berendtiopteris. “Alethopteris” serrata and “Sphenopteris” phyllocladoides are not to be regarded as evidence of ferns from Baltic amber. Reinvestigation of the holotypes of these two taxa did not reveal to which tracheophyte lineages these fossils belong. We suggest that the scarcity of fern remains from Baltic amber may reflect both a relatively low fern diversity in the source area of the fossil resin, and an absence or rarity of epiphytic and climbing ferns as observed in modern temperate forest ecosystems.

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 plant radiation and constitute seed plants’ closest relatives. Noeggerathiales are reconstructed as members of a plexus of free-sporing woody plants called progymnosperms, extending their age range by 60 My. Following the origin of seed plants, progymnosperms were previously thought to have become gradually less abundant before dying out in Carboniferous. We show they diversified and evolved complex morphologies including cone-like structures from modified leaves before going extinct at the Permian–Triassic extinction., Noeggerathiales are enigmatic plants that existed during Carboniferous and Permian times, ∼323 to 252 Mya. Although their morphology, diversity, and distribution are well known, their systematic affinity remained enigmatic because their anatomy was unknown. Here, we report from a 298-My-old volcanic ash deposit, an in situ, complete, anatomically preserved noeggerathialean. The plant resolves the group’s affinity and places it in a key evolutionary position within the seed plant sister group. Paratingia wuhaia sp. nov. is a small tree producing gymnospermous wood with a crown of pinnate, compound megaphyllous leaves and fertile shoots each with Ω-shaped vascular bundles. The heterosporous (containing both microspores and megaspores), bisporangiate fertile shoots appear cylindrical and cone-like, but their bilateral vasculature demonstrates that they are complex, three-dimensional sporophylls, representing leaf homologs that are unique to Noeggerathiales. The combination of heterospory and gymnospermous wood confirms that Paratingia, and thus the Noeggerathiales, are progymnosperms. Progymnosperms constitute the seed plant stem group, and Paratingia extends their range 60 My, to the end of the Permian. Cladistic analysis resolves the position of the Noeggerathiales as the most derived members of a heterosporous progymnosperm clade that are the seed plant sister group, altering our understanding of the relationships within the seed plant stem lineage and the transition from pteridophytic spore-based reproduction to the seed. Permian Noeggerathiales show that the heterosporous progymnosperm sister group to seed plants diversified alongside the primary radiation of seed plants for ∼110 My, independently evolving sophisticated cone-like fertile organs from modified leaves.

International audience; 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 diversities, requiring taxonomic lists to be normalised. Autochthonous floras provide the most direct evidence of vegetation diversity but these are rare; most plant beds are allochthonous with plant remains that have been subjected to varying levels of fragmentation, transportation and time averaging. Local-scale vegetation diversity is especially difficult to determine from the fossil record, even with rigorous sampling protocols and detailed sedimentological analysis. Landscape-scale and regional-scale vegetation diversities are more reliably determined but usually at the rank of family. Macrofossil and palynological data tend to reveal evidence of different aspects of plant diversity, and the best results are obtained if the two diversity signals are integrated. Despite the inherent difficulties, the plant fossil record provides clear evidence of the dynamic history of vegetation through geological times, including the effects of major processes such as climate changes and mass extinctions

International audience; Amber, a natural polymer, is fossil tree resin derived from diverse botanical sources with varying chemical compositions. As such, all amber is susceptible to the effects of light, temperature, relative humidity, and oxygen, as well as exposure to certain chemicals, and will deteriorate over time in collections if left unprotected. Here we review approaches for the conservation, preparation, and imaging of amber specimens and their inclusions, and address indications and causes of amber degradation, as well as recommendations for a suitable storage environment. We also provide updated preparation and embedding protocols, discuss several techniques for imaging inclusions, and address digitization efforts. A stable storage environment is essential to mitigate or avoid deterioration of amber, which often manifests as crazing, spalling, breaking and colour changes. Based on previous conservation studies of fossil resins, we generally recommend storage in a climate-monitored environment with a relative humidity of ca. 50%, 18 °C, and stress that light exposure must be kept to a minimum. For stabilization and anoxic sealing, amber specimens should ideally be embedded in an artificial epoxide resin (EpoTek 301-2 or similar is currently recommended). Amber should not be treated with or stored in vegetable or mineral oils (even for a short time for examination or photography), or come into contact with alcohol, disinfecting agents, hydrogen peroxide, or other destructive solvents or mixtures, since any of these materials can irreversibly damage the amber. Most photography of inclusions for research and digitization purposes can be successfully accomplished using light microscopy. Scanning electron microscopy (SEM) is sometimes used to uncover fine details, but is an invasive method. However, X-ray based methods (utilizing micro computed tomography, or micro-CT) are becoming more frequently used and increasingly indispensable in the examination of minute internal structures of inclusions, and to fully visualize important structures in opaque amber. Micro-CT makes it possible to digitize an inclusion three-dimensionally, and thus enables digital specimen ‘loans.’ Light microscopal images are still widely used in the digitization of amber specimens and are an essential alternative to micro-CT imaging when resources or time are limited. Overall, due to the vulnerability of all fossil resins, we recommend that conservation of amber samples and their inclusions be prioritized.

Amber is chiefly known as a preservational medium of biological inclusions, but it is itself a chemofossil, comprised of fossilised plant resin. The chemistry of today's resins has been long investigated as a means of understanding the botanical sources of ambers. However, little is known about the chemical variability of resins and consequently about that of the ambers that are derived from particular resins. We undertook experimental resin production in Araucariacean plants to clarify how much natural resin variability is present in two species, Agathis australis and Wollemia nobilis, and whether different resin exudation stimuli types can be chemically identified and differentiated. The latter were tested on the plants, and the resin exudates were collected and investigated with Fourier-transform infrared attenuated total reflection (FTIR-ATR) spectroscopy to give an overview of their chemistry for comparisons, including multivariate analyses. The Araucariacean resins tested did not show distinct chemical signatures linked to a particular resin-inducing treatment. Nonetheless, we did detect two separate groupings of the treatments for Agathis, in which the branch removal treatment and mimicked insect-boring treatment-derived resin spectra were more different from the resin spectra derived from other treatments. This appears linked to the lower resin viscosities observed in the branch- and insect-treatment-derived resins. However the resins, no matter the treatment, could be distinguished from both species. The effect of genetic variation was also considered using the same stimuli on both the seed-grown A. australis derived from wild-collected populations and on clonally derived W. nobilis plants with natural minimal genetic diversity. The variability in the resin chemistries collected did reflect the genetic variability of the source plant. We suggest that this natural variability needs to be taken into account when testing resin and amber chemistries in the future.

BackgroundAmber has been reported from the Early Cretaceous Crato Formation, as isolated clasts or within plant tissues. Undescribed cones of uncertain gymnosperm affinity have also been recovered with amber preserved in situ. Here, we provide multiple lines of evidence to determine the botanical affinity of this enigmatic, conspicuous cone type, and to better understand the diversity of amber-source plants present in the Crato Formation and beyond.ResultsA new taxon of amber-bearing pollen coneAraripestrobus resinosusgen. nov. et sp. nov. is described here from complete cones and characteristic disarticulated portions. The best-preserved cone portion has both in situ amber infilling the resin canals inside the preserved microsporophyll tissues and pollen of theEucommiidites-type. This places this genus within the Erdtmanithecales, an incompletely known gymnosperm group from the Mesozoic.FTIR analysis of the in situ amber indicates a potential araucariacean conifer affinity, although affinity with cupressacean conifers cannot be definitely ruled out. Pyr-GC-MS analysis of theAraripestrobus resinosusgen. nov. et sp. nov. in situ fossil resin shows that it is a mature class Ib amber, thought to indicate affinities with araucariacean and cupressacean, but not pinaceous, conifers. This is the first confirmed occurrence of this class of amber in the Crato Formation flora and in South America, except for an archaeological sample from Laguna Guatavita, Colombia.ConclusionsThe combined results of the cones’ novel gross morphology and the analyses of the in situ amber and pollen clearly indicate that the new taxon of resinous gymnosperm pollen cones from the Crato Formation is affiliated with Erdtmanithecales. The cone morphology is very distinct from all known pollen cone types of this extinct plant group. We therefore assume that the plant group that producedEucommiidites-type pollen is much more diverse in habits than previously thought. Moreover, the diversity of potential amber source plants from the Crato Formation is now expanded beyond the Araucariaceae and the Cheirolepidiaceae to include this member of the Erdtmanithecales. Despite dispersedEucommiiditespollen being noted from the Crato Formation, this is the first time macrofossils of Erdtmanithecales have been recognized from the Early Cretaceous of South America.

Motivation for this comment Recently, the Society of Vertebrate Paleontology (SVP) has sent around a letter, dated 21st April, 2020 to more than 300 palaeontological journals, signed by the President, Vice President and a former President of the society (Rayfield et al. 2020). The signatories of this letter request significant changes to the common practices in palaeontology. With our present, multi-authored comment, we aim to argue why these suggestions will not lead to improvement of both practice and ethics of palaeontological research but, conversely, hamper its further development. Although we disagree with most contents of the SVP letter, we appreciate this initiative to discuss scientific practices and the underlying ethics. Here, we consider different aspects of the suggestions by Rayfield et al. (2020) in which we see weaknesses and dangers. It is our intent to compile views from many different fields of palaeontology, as our discipline is (and should remain) pluralistic. This contribution deals with the aspects concerning Myanmar amber. Reference is made to Haug et al. (2020a) for another comment on aspects concerning amateur palaeontologists/ citizen scientists/private collectors.

The roots of modern-style ecosystems go back to the Carnian (ca. 233 Ma), a time of global climate change and extinction., The Carnian Pluvial Episode (Late Triassic) was a time of global environmental changes and possibly substantial coeval volcanism. The extent of the biological turnover in marine and terrestrial ecosystems is not well understood. Here, we present a meta-analysis of fossil data that suggests a substantial reduction in generic and species richness and the disappearance of 33% of marine genera. This crisis triggered major radiations. In the sea, the rise of the first scleractinian reefs and rock-forming calcareous nannofossils points to substantial changes in ocean chemistry. On land, there were major diversifications and originations of conifers, insects, dinosaurs, crocodiles, lizards, turtles, and mammals. Although there is uncertainty on the precise age of some of the recorded biological changes, these observations indicate that the Carnian Pluvial Episode was linked to a major extinction event and might have been the trigger of the spectacular radiation of many key groups that dominate modern ecosystems.

Amber is fossilised plant resin. It can be used to provide insights into the terrestrial conditions at the time the original resin was exuded. Amber research thus can inform many aspects of palaeontology, from the recovery and description of enclosed fossil organisms (biological inclusions) to attempts at reconstruction of past climates and environments. Here we focus on the resin itself, the conditions under which it may have been exuded, and its potential path to fossilisation, rather than on enclosed fossils. It is noteworthy that not all plants produce resin, and that not all resins can (nor do) become amber. Given the recent upsurge in the number of amber deposits described, it is time to re-examine ambers from a botanical perspective. Here we summarise the state of knowledge about resin production in modern ecosystems, and review the biological and ecological aspects of resin production in plants. We also present new observations on conifer-derived resin exudation, with a particular focus on araucarian conifer trees. We suggest that besides disease, insect attacks and traumatic wounding from fires and storms, other factors such as tree architecture and local soil conditions are significant in creating and preserving resin outpourings. We also examine the transformation of resin into amber (maturation), focusing on geological aspects of amber deposit formation and preservation. We present new evidence that expands previous understanding of amber deposit formation. Specific geological conditions such as anoxic burial are essential in the creation of amber from resin deposits. We show that in the past, the production of large amounts of resin could have been linked to global climate changes and environmental disruption. We then highlight where the gaps in our knowledge still remain and potential future research directions.

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 depend on the types of fossils being investigated, including their physical size, and whether the aim is to determine taxonomic richness or relative abundance. Describing and comparing diversities can use a range of different metrics, depending on whether binary presence/absence or abundance data are available. Each metric provides a different insight into the diversities and the choice of which to use depends on the research question being investigated. Various numerical approaches are available for identifying spatial and stratigraphical diversity patterns, mainly classificatory techniques (e.g., cluster and parsimony analyses) and ordination (e.g., Detrended Correspondence Analysis, Nonmetric Dimensional Scaling). The choice of technique again depends on the research question, but often it has proved useful to run both types of analysis in tandem. This article is illustrated by past biodiversity case studies from throughout the fossil record, dealing with floras ranging in age from the Devonian to the last few centuries.

PREMISE OF THE STUDY Extant dwarf mistletoes (Arceuthobium M. Bieb., Viscaceae) are hemiparasites with complex roles in nature. They are one of the most severe pests in northern hemisphere conifer forests, but they also enhance the structural complexity and species diversity of the forests. Here, we describe the first pre-Miocene macrofossils of dwarf mistletoes. The fossils from Eocene Baltic amber provide new insights into the morphological evolution of the Arceuthobium lineage and its paleobiogeography. METHODS The amber inclusions were investigated with light microscopy and compared with extant Viscaceae and to historic descriptions of lost Baltic amber fossils with affinities to Viscaceae. KEY RESULTS Six fossil species of the Arceuthobium lineage, A. johnianum comb. nov., A. mengeanum comb. nov., A. conwentzii sp. nov., A. groehnii sp. nov., A. viscoides comb. nov. and A. obovatum sp. nov., occurred in source forests of Baltic amber, representing the oldest macrofossil evidence of dwarf mistletoes. They share morphological features of their bracts, internodes, fruits, and stomata with extant Arceuthobium. Differences from extant dwarf mistletoes, such as the perianth merosity, the nonfusion of squamate bracts and presence of oblanceolate expanded leaves, indicate their affiliation to an ancient lineage of the genus. CONCLUSIONS The occurrence of six species of dwarf mistletoes in a single amber deposit suggests Arceuthobium was a keystone taxon of the Baltic amber source area. As in extant conifer forests, they probably influenced the structural complexity of the forest, not only leading to more open woodlands but also increasing species diversity, at least at a microhabitat scale.

Amber is a valuable source of Mesozoic and Cenozoic fossil fungi. The earliest amber-preserved fungi were described in the nineteenth and early twentieth centuries from Eocene Baltic amber. In 1907, Richard Klebs summarized the legacy of Robert Caspary based on his specimens, research notes and publications. This treatise contains descriptions and illustrations of 13 morphologies of fossil fungi which have not been investigated since. We managed to locate six of Caspary’s amber specimens and imaged and re-investigated the fossil fungi within. We provide amended descriptions of these six specimens, select new specimens from historic and recent collections which are probably conspecific with five fossils that appear to have been lost, and finally also describe and evaluate five newly found fossil fungi from Baltic amber. The fungi belong to the phylum Ascomycota (Subkingdom Dikarya). Only two of Caspary’s fungi can be confidently assigned to modern genera, Metacapnodium (Metacapnodiaceae) and Calicium (Caliciaceae). The new combination Calicium succini (Caspary) Rikkinen & A. R. Schmidt is made. The fossils originally placed in Acremonium, Cetraria, Gonatobotrys, Ramularia, Stilbum and Torula cannot be assigned to these genera, and should not be used as minimum age constrains for the respective lineages.

Some liquid plant exudates (e.g. resin) can be found preserved in the fossil record. However, due to their high solubility, gums have been assumed to dissolve before fossilisation. The visual appearance of gums (water-soluble polysaccharides) is so similar to other plant exudates, particularly resin, that chemical testing is essential to differentiate them. Remarkably, Welwitschiophyllum leaves from Early Cretaceous, Brazil provide the first chemical confirmation of a preserved gum. This is despite the leaves being exposed to water twice during formation and subsequent weathering of the Crato Formation. The Welwitschiophyllum plant shares the presence of gum ducts inside leaves with its presumed extant relative the gnetalean Welwitschia. This fossil gum presents a chemical signature remarkably similar to the gum in extant Welwitschia and is distinct from those of fossil resins. We show for the first time that a water-soluble plant exudate has been preserved in the fossil record, potentially allowing us to recognise further biomolecules thought to be lost during the fossilisation process.

Two new species of ants (Hymenoptera: Formicidae) collected from New Caledonia are described and figured based on worker specimens: Leptogenys loarelae Ramage sp. nov. (Ponerinae, Ponerini) and Lioponera neocaledonica Jouault, Ramage & Perrichot sp. nov. (Dorylinae, Cerapachyini). All specimens were collected from the South Province of Grande Terre. These two new species are primarily distinguished from the other New Caledonian relatives by the size and shape of petiole for L. loarelae Ramage sp. nov. and by the presence of dorsolateral margins on the mesosoma for L. neocaledonica Jouault, Ramage & Perrichot sp. nov. Keys to New Caledonian Leptogenys and Lioponera are provided.

We report the first bona fide graminid spikelet inclusions found in Eocene Baltic amber. The most informative anatomically preserved specimen is assigned to the genus Rhynchospora Vahl (Cyperaceae), whereas two others show affinities with sedges (Cyperaceae) or grasses (Poaceae). Examination of historic descriptions of putative graminid inclusions from Baltic amber suggests that one is of coniferous origin, while the affinities of the other fragmentary specimens remain uncertain as they have been lost. The graminid inclusions described here challenge previous notions of the Baltic amber source area being a dark, close canopy forest and rather indicate at least some open and light habitats with swampy and wet areas within the ‘Baltic amber forest’, and thus enlighten its hitherto obscure palaeoenvironment and floristic composition.

During the past two decades, a plethora of fossil micro-organisms have been described from various Triassic to Miocene ambers. However, in addition to entrapped microbes, ambers commonly contain microscopic inclusions that sometimes resemble amoebae, ciliates, microfungi, and unicellular algae in size and shape, but do not provide further diagnostic features thereof. For a better assessment of the actual fossil record of unicellular eukaryotes in amber, we studied equivalent inclusions in modern resin of the Araucariaceae; this conifer family comprises important amber-producers in Earth history. Using time-of-flight secondary ion mass spectrometry (ToF-SIMS), we investigated the chemical nature of the inclusion matter and the resin matrix. Whereas the matrix, as expected, showed a more hydrocarbon/aromatic-dominated composition, the inclusions contain abundant salt ions and polar organics. However, the absence of signals characteristic for cellular biomass, namely distinctive proteinaceous amino acids and lipid moieties, indicates that the inclusions do not contain microbial cellular matter but salts and hydrophilic organic substances that probably derived from the plant itself. Rather than representing protists or their remains, these microbe-like inclusions, for which we propose the term 'pseudoinclusions', consist of compounds that are immiscible with the terpenoid resin matrix and were probably secreted in small amounts together with the actual resin by the plant tissue. Consequently, reports of protists from amber that are only based on the similarity of the overall shape and size to extant taxa, but do not provide relevant features at light-microscopical and ultrastructural level, cannot be accepted as unambiguous fossil evidence for these particular groups.

The Lower Cretaceous Crato Formation of north-east Brazil yields a diverse plant assemblage. It has yielded many macrofossils thought to be related to the enigmatic gymnosperm group Gnetales, including the long leaf Welwitschiophyllum brasiliense Dilcher et al. 2005. This fossil plant is considered to be related to the extant gnetalean Welwitschia mirabilis Hooker 1863, despite lacking many gnetalean characteristics. Presently, this macrophyte fossil is known only from detached leaves and much anatomical information is currently unavailable. The reproductive structures assigned to the family Welwitschiaceae in the Crato Formation, as well as several key morphological features of the leaves are currently thought to relate fossil Welwitschiophyllum to extant Welwitschia. These leaf characters include: isobilateral leaf form, triangular elongated leaf shape with a wide base, longitudinal splitting from a frayed leaf apex, numerous parallel veins and possible thickening of the epidermis. However, several of these leaf characteristics also occur in many other macrophytes, perhaps as a result of convergence. Anatomical and morphological data described here from fossil Welwitschiophyllum leaves is compared with extant Welwitschia. Our results show that Welwitschiophyllum can only be placed tentatively in Gnetales, as many of the features we report are not diagnostic, and may have resulted from convergent evolution (e.g., gum). Fossils with better anatomical preservation and the reconstruction of the whole plant are really needed to better understand the affinities of Welwitschiophyllum.

International audience; Amber is fossilised plant resin. It can be used to provide insights into the terrestrial conditions at the time the original resin was exuded. Amber research thus can inform many aspects of palaeontology, from the recovery and description of enclosed fossil organisms (biological inclusions) to attempts at reconstruction of past climates and environments. Here we focus on the resin itself, the conditions under which it may have been exuded, and its potential path to fossilisation, rather than on enclosed fossils. It is noteworthy that not all plants produce resin, and that not all resins can (nor do) become amber. Given the recent upsurge in the number of amber deposits described, it is time to reexamine ambers from a botanical perspective. Here we summarise the state of knowledge about resin production in modern ecosystems, and review the biological and ecological aspects of resin production in plants. We also present new observations on conifer-derived resin exudation, with a particular focus on araucarian conifer trees. We suggest that besides disease, insect attacks and traumatic wounding from fires and storms, other factors such as tree architecture and local soil conditions are significant in creating and preserving resin outpourings. We also examine the transformation of resin into amber (maturation), focusing on geological aspects of amber deposit formation and preservation. We present new evidence that expands previous understanding of amber deposit formation. Specific geological conditions such as anoxic burial are essential in the creation of amber from resin deposits. We show that in the past, the production of large amounts of resin could have been linked to global climate changes and environmental disruption. We then highlight where the gaps in our knowledge still remain and potential future research directions.

Amber is nearly ubiquitous in lignites from Otago and Southland and cornmon throughout New Zealand; however, no amber inclusions have been reported previously. We studied amber from 22 Cretaceous to Miocene sites in southem New Zealand, recovering inclusions at three localities: Cosy Dell (late Oligocene), Roxburgh (early Miocene), and Hyde (?early Miocene). Preparation of New Zealand amber to expose inclusions for study under incident and transmitted light is challenging and time-consuming, with most samples brittle and opaque. Thus, we stabilize and clear the amber lumps using epoxy preparation under vacuum before grinding and imaging under light microscopy. To date we have recovered 63 arthropods, as weIl as plant remains, fungi, and nematodes. Arachnids include diverse mites (Mesostigmata, Oribatida, Astigmata, and Prostigmata), a variety of spiders and web remains with prey, and a pseudoscorpion. Sorne Collembola were identified as belonging to the family Entomobryidae (Entomobryomorpha). Insects include members of the families Dermestidae (Coleoptera), Mymaridae and Scelionidae (Hymenoptera), Veliidae (Heteroptera), Ceratopogonidae (Forcipomyia) and Mycetophilidae (Diptera), as well as Psocoptera, and Lepidoptera. The most abundant fungi in New Zealand amber are hyphomycetes similar to the genus Casparyotorula from European Palaeogene ambers and we discovered similar fungi growing on resin of the extant Agathis australis, the iconic New Zealand Kauri. Furthermore, specimens of the genus Metacapnodium (Capnodiales) represent the first Southem Hemisphere fossil sooty moulds; saprophytic ascomycetes with brown hyphae, often forming extensive subicula on living plant surfaces. These fungi are ubiquitous and diverse in New Zealand today. Many of these new amber fossils represent groups with an otherwise poor fossil record for the entire Southem Hemisphere. The systematic and ecological diversity of the inclusions highlights the potential of New Zealand amber for reconstructing past terres trial ecosystems of Zealandia, one of the biogeographically crucial former Gondwanan landmasses.

The Baltic amber deposit represents the largest accumulation of any fossil resin worldwide and hundreds of thousands of entrapped arthropods have been recovered so far. The source plants of Baltic amber, however, are still controversial, and the botanical composition of the ‘Baltic amber forest’ remains poorly studied. Here, we provide the first unequivocal Baltic amber inclusions of the umbrella pine Sciadopitys (Sciadopityaceae), a genus that has been suggested as the source of succinite (the main variety of Baltic amber) based on chemical analyses. As previously suggested sciadopitoid inclusions must be reconsidered as being notional, representing angiosperm leaves instead, the new fossils are the first unambiguous macrofossil evidence of Sciadopitys from the ‘Baltic amber forest’, and the first pre-Oligocene macrofossil record of Sciadopitys from Europe. The fossil Sciadopitys cladodes provide new insights into the conifer diversity of the ‘Baltic amber forest’ and broaden the picture of its palaeoecology, indicating the presence of humid swamp to raised bog habitats. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180, 258–268.

Stigmaria is one of the iconic plant fossils of the Carboniferous and fragments of the narrower parts of the rhizomorph are found in most museum collections. However, very few almost entire specimens have been found and preserved. A new specimen of Stigmaria from Brymbo, North Wales is described and compared with other preserved examples from Europe and North America. The Brymbo specimen shows a large portion of trunk still attached to the large stigmarian base, which is a rare find, and this specimen supports our ideas of how these impressively large casts were formed. Stigmarias were preserved by the deposition of minerals around them following a sediment inundation, which gave sufficient support while the tissues rotted and filled with sediments. Remnants of the outer tissues were compressed to form a thin surrounding coal layer.

International audience; The world’s sole ‘parasitic’ gymnosperm Parasitaxus usta (Podocarpaceae) is endemic to the island of Grande Terre, New Caledonia. It is a threatened species because of its limited geographic range and progressing habitat fragmentation. Here, we report a novel scale insect outbreak on a Parasitaxus sub-population from Monts Dzumac in the southern part of Grande Terre. The identity of the scale insect was determined through combining morphological and molecular methods. The field collection of scale insects and their secretions from infested Parasitaxus specimens allowed morphological identification of the superfamily Coccoidea. Subsequent genetic sequencing using CO1 markers allowed phylogenetic placement of the wax scale insects to the genus Ceroplastes (Coccoidea, Coccidae), a widespread pest genus. The identified species, C. pseudoceriferus, has not been previously recorded from New Caledonia. As Parasitaxus is already vulnerable to extinction, this new threat to its long-term survival needs to be monitored. Other New Caledonian endemic plant species are potentially at risk of this new species, although it was not observed on Falcatifolium taxoides, the host of Parasitaxus.

International audience; Conifers of the endemic species Araucaria humboldtensis on Mont Humboldt in New Caledonia exhibit extensive resin exudation. The resin flows of these threatened trees are here shown to be induced by two beetle species, which bore into branches and branchlets, leading to abundant outpouring of resin, which gradually solidifies into often drop-shaped resin bodies. The exudate is colonized by a resinicolous and likely insect-vectored ascomycete, Resinogalea humboldtensis, which is only known from Mont Humboldt. The fungus grows into fresh resin and eventually develops ascomata on the surface of solidifying resin. The solidified resin is also colonized by another fungus, a dematiaceous hyphomycete. Based on protein coding (CO1, CAD, ArgK) and ribosomal (LSU) genes, the larger branch-boring beetle is a weevil of the tribe Araucariini, which represents the sister group of all other cossonine weevils. The smaller beetle species belongs to the longhorn beetles (Cerambycidae). The strong host specificity of the Araucariini, along with the occurrence of two unique fungi, suggests that the resin-associated community is native and has evolved on the endemic conifer host. The formation of large amber deposits indicates massive resin production in the past, but the environmental triggers of exudation in Mesozoic and Cenozoic ecosystems remain unclear. Our observations from Mont Humboldt support the notion that the occurrences of small drop-shaped amber pieces in Triassic to Miocene amber deposits were linked to ancient insect infestations.

Stable carbon-isotope geochemistry of fossilized tree resin (amber) potentially could be a very useful tool to infer the composition of past atmospheres. To test the reliability of amber as a proxy for the atmosphere, we studied the variability of modern resin δ 13 C at both local and global scales. An amber δ 13 C curve was then built for the Cretaceous, a period of abundant resin production, and interpreted in light of data from modern resins. Our data show that hardening changes the pristine δ 13 C value by causing a 13 C-depletion in solid resin when compared to fresh liquid–viscous resin, probably due to the loss of 13 C-enriched volatiles. Modern resin δ 13 C values vary as a function of physiological and environmental parameters in ways that are similar to those described for leaves and wood. Resin δ 13 C varies between plant species and localities, within the same tree and between different plant tissues by up to 6‰, and in general increases with increasing altitudes of the plant-growing site. We show that, as is the case with modern resin, Cretaceous amber δ 13 C has a high variability, generally higher than that of other fossil material. Despite the high natural variability, amber shows a negative 2.5–3‰ δ 13 C trend from the middle Early Cretaceous to the Maastrichtian that parallels published terrestrial δ 13 C records. This trend mirrors changes in the atmospheric δ 13 C calculated from the δ 13 C and δ 18 O of benthic foraminiferal tests, although the magnitude of the shift is larger in plant material than in the atmosphere. Increasing mean annual precipitation and p O 2 could have enhanced plant carbon-isotope fractionation during the Late Cretaceous, whereas changing p CO 2 levels seem to have had no effect on plant carbon-isotope fractionation. The results of this study suggest that amber is a powerful fossil plant material for palaeoenvironmental and palaeoclimatic reconstructions. Improvement of the resolution of the existing data coupled with more detailed information about botanical source and environmental growing conditions of the fossil plant material will probably allow a more faithful interpretation of amber δ 13 C records and a wider understanding of the composition of the past atmosphere.

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 reinvestigated these hook-like structures and re-evaluated the nature of the co-occurrences in context with the flora preserved in each plant-bearing fossil ‘bed’ in the North China sedimentary succession. New findings show that the species Gigantonoclea hallei probably climbed using specially adapted clusters of compound grappling hook-like shoots borne on the stems. This structural arrangement comprising shoots of hooks is new to the scrambling/climbing concept in gigantopterids. However, a key figured specimen previously reported as showing intermediate hook-tipped leaf morphology on a sole pinnule tip is discounted as such and is reinterpreted as a ‘normal’ pinnule partially hidden under sediment that results in an unusual appearance to this pinnule tip. Adaptations for climbing or scrambling based upon ‘hooked leaves’ observed in Gigantonoclea lagrelii are no longer supported and are reinterpreted as incompletely expanded leaves where the vernation process was interrupted. These data weaken prior interpretations of G. lagrelii as a climber/scrambler and raise doubts about the ubiquity of hooks amongst the gigantopterids as structures enabling them to climb or scramble their way through the Permian world.

Triassic plant remains are uncommon globally, with few Early-Middle Triassic floras well documented. Thus, the Middle Triassic (Anisian) of Bromsgrove, Worcestershire, UK provides new insights into the terrestrial biota at this time and is extremely valuable since it provides the majority of fossil plants from the UK terrestrial Triassic sequence. This small but diverse flora comprises typical gymnospermous (Willsiostrobus, Pelourdea) and sphenopterid (Schizoneura, Neocalamites) elements of an Anisian-age flora. Reinvestigation of megafossil remains led to the discovery of a previously unknown and undescribed plant fossil with in situ spores, Bromsgrovia willsii gen. et sp. nov. The in situ spores were extracted and examined by light, scanning electron and transmission electron microscopy. The Bromsgrove Anisian flora is summarized and illustrated along with the first occurrence of in situ Verrucosisporites applanatus spores, a marker for the Middle Triassic. The parent plant of V. applanatus is suggested to be a horsetail with an unusual morphology.

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 secretory cells, and is dictyostelic. In order to evaluate the systematic and phylogenetic position of Zhongmingella within Guaireaceae, and Guaireaceae within Osmundales, we conducted a preliminary cladistic analysis of a broad range of Osmundales and related taxa based on 18 extinct and six extant genera and subgenera. Results do not support the traditionally defined family Thamnopteroideae (Bathypteris, Chasmatopteris, Iegosigopteris, Thamnopteris, Zalesskya) and demonstrate that Thamnopteroideae is not a subfamily of Osmundaceae as previously thought. Guaireaceae is monophyletic but in addition to its traditionally defined members (Guairea, Lunea, Donwellicaul...

Extant dwarf mistletoes (The amber inclusions were investigated with light microscopy and compared with extant Viscaceae and to historic descriptions of lost Baltic amber fossils with affinities to Viscaceae.Six fossil species of theThe occurrence of six species of dwarf mistletoes in a single amber deposit suggests

International audience; A novel species of ascomycetes is described from resin of Araucaria humboldtensis on Mont Humboldt in New Caledonia. The fungus is placed in the new genus Resinogalea Rikkinen & A.R. Schmidt, with the species name R. humboldtensis Rikkinen & A.R. Schmidt. It has only been found growing on semi-hardened resin flows on branches of its endemic and endangered conifer host. The morphology and anatomy of the new fungus are compared with those of ecologically similar taxa, including Bruceomyces castoris. The new family Bruceomycetaceae Rikkinen & A.R. Schmidt is described toaccommodate Resinogalea and Bruceomyces.

Agaricomycetes are major components of extant terrestrial ecosystems; however, their fruiting bodies are exceedingly rare as fossils. Reinvestigation of a peculiar fossil from Late Triassic sediments of southern Germany interpreted as a bracket fungus revealed that this fossil in fact represents a wood abnormality, resulting from injury to the cambium and subsequent callus growth in a Baieroxylon -like ginkgoalean wood. As a result, the fossil record of the Agaricomycetes does not yet pre-date the Early Cretaceous, suggesting a late diversification of basidiomycetes possessing large fruiting bodies. doi:10.1002/mmng.201200006

The only known specimen of Lepidophloios ichthyoderma Lesquereux, from the Pennsylvanian of Illinois, is a stem fragment that has numerous unusual large raised tubercles surrounded by smaller scales. The relationship of these structures to each other was unclear. Reinvestigation by cuticular analysis shows the epidermal structure of both the large protuberant tubercles and the small irregularly interposed small leaves share the same epidermal cell morphology and stomatal pitting. Both the tubercles and the leaves appear vegetative as no reproductive structures were recovered and the presence of stomata on these tubercles may also indicate a vegetative role. This combination of vegetative tubercles and leaves leads to a renaming of the specimen as Halonia ichthyoderma comb. nov. and a redefinition of the genus Halonia Lindley and Hutton is given.

Recent investigations into Permian aged floras from China have highlighted the widespread occurrence of callistophytalean pteridosperms that challenge previous understanding of their spatial and temporal distribution and diversity. In China, the group spans the Permian period and constitutes a distinctive but rare component in many peat- forming environments. The stratigraphically earliest calli- stophytalean occurs in the Asselian-Sakmarian stages with fossils from the Taiyuan Formation of northern China including ovules of Callospermarion undulatum in coal ball assemblages, and ovulate fronds of Norinosperma shanxiensis and synangiate fronds of Norinotheca shanxiensis in adpres- sion assemblages. More abundant in the fossil records are ad- pression remains from the Roadian-Wordian stages with the Lower Shihhotse Formation preserving abundant vegetative and ovulate remains of Emplectopteris triangularis that is now considered to represent a callistophytalean. The youngest callistophytalean recognised is from the Wuchaipingian- Changhsingian stages with the Xuanwei Formation of southern China containing a single stem of Callistophyton boyssetii that provides indisputable evidence of the group in the lead up to the end-Permian mass extinction. These accounts are augmented by analysis of pollen records that demonstrate the callistophytalean pollen genus Vesicaspora to be widespread through palynological assemblages from the Permian period in both North and South China, including the Upper Shihhotse Formation, Shihchienfeng Group, Xuanwei Formation, and possibly also in the mid-Pennsylva- nian Benxi Formation. Although macrofossil specimens are uncommon elements in the assemblages that contain them, they demonstrate the continuity of callistophytalean pterido- sperms from the Pennsylvanian sub-period into the early Guadalupian epoch of the Permian in North China and into the Lopingian epoch of the Permian in South China. Of the species present, both Callistophyton boyssetii and Callosper- marion undulatum are known from the Pennsylvanian-earli- est Permian age floras of Euramerica, whereas Norinosperma, Norinotheca and Emplectopteris appear to represent endemic Cathaysian elements. Results imply that callistophytalean pteridosperms can no longer be excluded from theories of post-Carboniferous plant evolution and floristics, appearing to have played an important role in both Permian and Car- boniferous aged plant communities. The presence of Vesicas- pora in several formations from which macro-remains have not been identified is a hopeful indicator that further calli- stophytalean pteridosperms are yet to be found.

Isolated ovules occur in many fossil plant assemblages, where they provide important insights into seed-plant diversity and evolution. However, in many cases, the ovules cannot be attributed to individual groups of seed plants, restricting systematic and evolutionary assessments that can be made from otherwise well-characterized fossil taxa. In the present paper, we describe a new kind of ovule discovered in tuffaceous sediments from the Permian-aged Xuanwei Formation in Guizhou Province, China. This ovule has 180° rotational symmetry and an integument comprising a variably thick sarcotesta, a uniformly thick sclerotesta and a uniformly thin endotesta. The nucellus is attached to the integument at least basally and contains a collapsed seed megaspore; a nucellar apex is absent. Both the integument and nucellus are vascularized by paired bundles in the major plane of the ovule; the integumentary bundles are considerably larger than the nucellar bundles and the nucellar bundles emerge from a conical vascular pad. Generation of a three-dimensional reconstruction based on serial peels revealed the gross morphology and organization of the ovule and highlighted the presence of features consistent with cardiocarpalean-type ovules (ovule shape, histological features of the integument) and also features more typical of lagenostomalean- and trigonocarpalean-type ovules (large integumentary bundles, presence of nucellar bundles). To assess the affinity and evolutionary significance of the ovule, it has been included in a cladistic matrix of cardiocarpalean-, lagenostomalean- and trigonocarpalean-type ovules. Results place the ovule within the cardiocarpalean group of ovules known to have been produced by several plant groups, including cordaitean coniferophytes, pteridosperms and Palaeozoic conifers. The cladistic topology supports generic level distinction of the present species, requiring the establishment of Muricosperma guizhouensis Seyfullah & J.Hilton gen. & sp. nov. Lagenostomalean ovules produced by hydrasperman pteridosperms form a basal paraphyletic grade, whereas trigonocarpalean ovules produced by medullosan pteridosperms form a monophyletic group in which Stephanospermum is paraphyletic with respect to Rhynchosperma and Pachytesta. The results also place the Mississippian ovule Mitrospermum bulbosum apart from all of the Pennsylvanian species of Mitrospermum that form a strongly supported clade. Consequently, M. bulbosum is transferred to the new genus Whitaddera Seyfullah & J.Hilton as W. bulbosa (Long) Seyfullah & J.Hilton. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164, 84–108.

In 1927, T. G. Halle published an extensive synthesis of the Permian fossil plant assemblages from China’s Shanxi Province that included five genera and species of seed plant that either had ovules in organic attachment or closely and repeatedly associated with foliage. These fertile pteridosperms were the subject of a later publication (1929) that provided additional information on each species, but since that time only Nystroemia pectiniformis Halle has been subjected to detailed investigation based on the original materials. Reinvestigation of the remaining four species has yielded additional examples and new information for each species, and has confirmed the pteridospermalean affinity for three of the four species. Alethopteris norinii Halle shows organic attachment of a pollen organ similar to Wittleseya (Newberry) Halle and is reinterpreted as a microsporangiate medullosan pteridosperm. Emplectopteris triangularis Halle and Sphenopteris tenuis (Schenk) Halle both bear numerous ovules attached directly to the leaf laminae, with Emplectopteris conforming with the gross-organisation of callistophytalean pteridosperms and S. tenuis being interpreted as a derived lagenostomalean pteridosperm. However, discovery of the counterpart to the key specimen of Pecopteris wongii Halle that Halle interpreted as bearing ovules demonstrates this species to be a sterile marattialean frond that agrees with its assignment to Pecopteris. In addition, two further specimens are demonstrated to be fertile organs of previously unknown pteridosperms; Norinosperma shanxiensis gen. et sp. nov. bears cardiocarpalean ovules abaxially on the proximal portions of the frond with ovules positioned directly on the veins of a leaf lamina; Norinotheca shanxiensis gen. et sp. nov. bears large pollen organs abaxially and singly on the vein of the leaf lamina. Collectively Halle’s pteridosperms remain important today as they are some of the few specimens that actually demonstrate organic connection of fertile organs to foliage, helping to characterise the morphology and organisation of this seed plant group. These species also demonstrate the persistence of seed plant families and genera in the Permian of China that became regionally extinct at the end of the Carboniferous in Euramerica, and show that pteridosperms remained an important component of Permian as well as Carboniferous floras globally.

Some higher plants, both angiosperms and gymnosperms, can produce resins and some of these resins can polymerize and fossilize to form ambers. Various physical and chemical techniques have been used to identify and profile different plant resins and have then been applied to fossilized resins (ambers), to try to detect their parent plant affinities and understand the process of polymerization, with varying levels of success. Here we focus on resins produced from today's most resinous conifer family, the Araucariaceae, which are thought to be the parent plants of some of the Southern Hemisphere's fossil resin deposits. Fourier transform infrared (FTIR) spectra of the resins of closely related Araucariaceae species were examined to test whether they could be distinguished at genus and species level and whether the results could then be used to infer the parent plant of a New Zealand amber. The resin FTIR spectra are distinguishable from each other, and the three Araucaria species sampled produced similar FTIR spectra, to which Wollemia resin is most similar. Interspecific variability of the FTIR spectra is greatest in the three Agathis species tested. The New Zealand amber sample is similar in key shared features with the resin samples, but it does differ from the extant resin samples in key distinguishing features, nonetheless it is most similar to the resin of Agathis australis in this dataset. However on comparison with previously published FTIR spectra of similar aged amber and older (Eocene) resinites both found in coals from New Zealand and fresh Agathis australis resin, our amber has some features that imply a relatively immature resin, which was not expected from an amber of the Miocene age. peerReviewed