Notes on a fertile rhyniophytoid from the Lower Devonian of Guangxi,southwestern China

A fertile rhyniophytoid is discovered from the Lower Devonian of Guangxi, southwestern China, which is characterised by small size, dichotomous branchings and terminal sporangia. This plant is similar to the genus Aberlemnia Gonez et Gerrienne and also, is comparable to some mesofossil morphotypes of early land plants from the Early Devonian (Lochkovian) Old Red Sandstone floras. This study adds new data to the generally zosterophyll-dominated Early Devonian floras of South China and sheds some lights on the palaeophytogeography of rhyniophytoids.     

Leaf evolution in early-diverging ferns: insights from a new fern-like plant from the Late Devonian of China

Background and Aims With the exception of angiosperms, the main euphyllophyte lineages (i.e. ferns sensu lato, progymnosperms and gymnosperms) had evolved laminate leaves by the Late Devonian. The evolution of laminate leaves, however, remains unclear for early-diverging ferns, largely represented by fern-like plants. This study presents a novel fern-like taxon with pinnules, which provides new insights into the early evolution of laminate leaves in early-diverging ferns.Methods Macrofossil specimens were collected from the Upper Devonian (Famennian) Wutong Formation of Anhui and Jiangsu Provinces, South China. A standard degagement technique was employed to uncover compressed plant portions within the rock matrix.Key Results A new fern-like taxon, Shougangia bella gen. et sp. nov., is described and represents an early-diverging fern with highly derived features. It has a partially creeping stem with adventitious roots only on one side, upright primary and secondary branches arranged in helices, tertiary branches borne alternately or (sub)oppositely, laminate and usually lobed leaves with divergent veins, and complex fertile organs terminating tertiary branches and possessing multiple divisions and numerous terminal sporangia.Conclusions Shougangia bella provides unequivocal fossil evidence for laminate leaves in early-diverging ferns. It suggests that fern-like plants, along with other euphyllophyte lineages, had independently evolved megaphylls by the Late Devonian, possibly in response to a significant decline in atmospheric CO₂ concentration. Among fern-like plants, planate ultimate appendages are homologous with laminate pinnules, and in the evolution of megaphylls, fertile organs tend to become complex.     

A new group of Early Devonian plants with valvate sporangia containing sculptured permanent dyads

As part of a study to explore diversity and disparity in Early Devonian terrestrial vegetation, several hundreds of sporangia with in situ spores have been isolated from a Lochkovian locality in Shropshire. These include a small number (seven) of sporangia showing dehiscence into four valves and containing permanent sculptured dyads, belonging to the Cymbohilates horridus complex and C. cymosus, which are recorded in coeval dispersed spore assemblages. A further, previously described, mesofossil comprises an incomplete sporangium containing C. horridus that terminates a naked isotomously branching stem with stomata. The valvate sporangia are placed in a new genus, Partitatheca, containing four species, P. splendida (type), P. horrida, P. densa and P. cymosa, their names reflecting the names of the dispersed spore species and varieties. Complex ultrastructure in the walls of the dyads is similar to that in earlier dyads in the Dyadospora complex where it provides evidence for a hepatic affinity of the earliest embryophytes, but the new taxa present a combination of bryophyte and tracheophyte characters and are considered to represent a new embryophyte lineage. General discussion includes the development of dyads, more particularly their relevance to understanding the diversity in meiotic processes, and the disappearance of dyads from the dispersed spore record prior to the Middle Devonian. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 229–257.     

Composition and dynamics of the great Phanerozoic Evolutionary Floras

Factor analysis of a data set representing the global distribution of vascular plant families through time shows the broad pattern of vegetation history can be explained in terms of five Evolutionary Floras. The Rhyniophytic (=Eotrachyophytic) Flora represents the very earliest (Silurian and earliest Devonian) vascular plants, notably the Rhyniophytopsida. The Eophytic Flora represents the early (Early–Middle Devonian) mainly homosporous land plants, notably the Zosterophyllopsida, Trimerophytopsida and early Lycopsida. The Palaeophytic Flora represents the Late Devonian and Carboniferous vegetation, which saw the introduction of heterospory among the spore producing plants and of early gymnosperms. The Mesophytic Flora first appeared in the Late Carboniferous and Permian macrofossil record, although there is palynological evidence of these plants having grown earlier in extra‐basinal habitats and was dominated by gymnosperms with more modern affinities. The Cenophytic Flora that first appeared during Cretaceous times was overwhelmingly dominated by angiosperms. The end‐Devonian, end‐Triassic and end‐Cretaceous mass‐extinction events recognized in the marine fossil record had little impact on the diversity dynamics of these Evolutionary Floras. Rather, the changes between floras mainly reflect key evolutionary innovations such as heterospory, ovules and angiospermy.     

Ultrastructure of laevigate hilate spores in sporangia and spore masses from the Upper Silurian and Lower Devonian of the Welsh Borderland

Spore masses and isolated sporangia, containing laevigate hilate cryptospores attributable to the dispersed taxon Laevolancis divellomedia sensu lato, have been recovered on bulk maceration of Upper Silurian (Pridoli) and Lower Devonian (Lochkovian) deposits from the Welsh Borderland. Detailed morphological, anatomical and ultrastructural analysis, using light microscope, scanning electron microscope and transmission electron microscope techniques, reveals subtle differences between the specimens and they can be grouped into five distinct types. The different groups are distinguished principally by using sporangia-spore mass characteristics, presence or absence of extra-exosporal material and nature of spore-wall ultrastructure. Of the groups, one has a uniformly homogeneous exospore and the other four groups have a bilayered exospore. In the former the spores lack extra-exosporal material and occur in a discoidal sporangium. Of the bilayered groups, two have exospores of homogeneous composition but with the two layers differing in electron density. They occur in discoidal sporangia and spore masses and are distinguished on the presence or absence of extra-exosporal material and differences in the widths of the two layers. Finally, two bilayered groups possess a lamellate inner layer, but vary in presumed sporangial shape. Elongate sporangia have spores with concentric continuous lamellae, lacking further ultrastructure. In contrast, spores from a discoidal spore mass have white-line-centred, presumably tripartite, lamellae which are laterally discontinuous, overlapping and irregularly spaced. These findings, which suggest that morphologically similar spores were produced by a number of plant taxa, have important implications regarding the assessment of early land-plant diversity. The affinities of hilate cryptospore-producing plants are unknown and problematic, particularly as no extant non-angiosperm plants produce dyads, other than through meiotic irregularity, and spore-sporangial characters have no exact counterpart in coeval plants. Studies of specimens with in situ hilate cryptospores suggest that they derive from rhyniophytoids, i.e. plants that resemble the simplest of vascular plants but lack evidence of vascular tissue, although hilate cryptospore-containing examples show no axial branching. It might be argued, based on evidence from spore wall ultrastructure, that some of the plants have more in common with lycopsids and filicopsids than bryophytes, a surprising finding bearing in mind the stratigraphic distribution of hilate cryptospores-dyads and inferences that the producers were bryophyte-like. Detailed studies of wall structure in the hilate cryptospores permit consideration of spore wall development. It is suggested that extra-exosporal material derives from a tapetum and is thus produced by the diploid sporophyte. The white-line-centred lamellae in a single specimen provide the earliest evidence for the presence of such structures in early land plant spores and provide further evidence that sporopollenin deposition on such structures is the most primitive mode of sporopollenin deposition among land plants.     

New palaeobotanical data from the Lower Devonian of Mezquita de Loscos (Teruel Province,Spain)

Recent fieldwork has uncovered three new localities from the Lower Devonian of Mezquita de Loscos (Teruel Province, Spain) with further plant mega-fossils and the first record of micro-fossils. Such plant remains have been interpreted as belonging to a basal euphyllophyte, Taeniocrada-like stems, Hostinella genus and paired sporangia. Fourteen spore taxa were recovered, including Ambitisporites, Aneurospora, Brochotriletes, Chelinospora, Emphanisporites, Gneudnaspora and Retusotriletres, among others. New evidence confirms a Lochkovian age for this outcrop and suggests that the plant diversity was more complex than originally documented.     

Exceptional preservation in Lower Devonian coalified fossils from the Welsh Borderland: a new genus based on reniform sporangia lacking thickened borders

Reniform sporangia, comprising two equal valves and containing retusoid spores, recovered from Lower Old Red Sandstone strata of Devonian age (micrornatus-newportensis Spore Biozone: lower Gedinnian lower Lochkovian) on North Brown Clee Hill in the Welsh Borderland are placed in Resilitheca salopensis gen. et sp. nov. Conventional compression fossils from Targrove, Ludlow of fertile axes showing isotomous branching with limited overtopping are considered conspecific because the terminal reniform sporangia contain the same spores. Spore ultrastructure is described using scanning and transmission electron microscopy. Sections show faint traces of lamellae. Particles associated with spores and sporangium wall are compared with the globules of pteridophytes and Ubisch bodies of angiosperms, and related to the development of the sporangium. The new plants are compared with Cooksonia caledonica Edwards known only from impressions, and with Renalia Gensel showing far more pronounced pseudomonopodial branching.     

Tetrads in sporangia and spore masses from the Upper Silurian and Lower Devonian of the Welsh Borderland

Prior to the mid-Silurian, evidence for the earliest embryophytes comes from dispersed spores, particularly permanent tetrads, there being no fossils showing gross morphology or anatomy of the producers. The fragmentary coalified mesofossils described here from the uppermost Silurian (Pridoli) and basal Devonian (Lochkovian) of the Welsh Borderland contain tetrads assigned to Tetrahedraletes, Velatitetras and Cheilotetras. These spores together with examples from spore masses have been examined by scanning and transmission electron microscopy and display diversity in ultrastructure of the exospore and envelope. Tetrads have been found, together with a putative elater, in the forking apex of an axial Lochkovian fossil, named Grisellatheca salopensis gen. et sp. nov., that anatomically, apart from spore characters, reveals no unequivocal evidence for hepatic affinity. The remaining fossils are equally as uninformative as regards affinity. Tetrads with ornamented envelopes are recorded in an isolated discoidal sporangium and in the bases of incomplete sporangia borne terminally on a bifurcating axis. Both ornament and ultrastructure suggest that the spores belong to quite distinct species within Velatitetras. Tetrahedraletes is recorded in an incomplete sporangium subtended by a forking axis, in which no cellular detail is preserved. Naked unfused tetrads also assigned to Tetrahedraletes are recorded in spore masses from both localities and again exospore ultrastructure demonstrates diversity. A final Lochkovian sporangium contains naked tetrads with sporadic Papiculate ornament and shows a unique trilayered exospore. Comparisons of exospore ultrastructure in these tetrads, which it is argued are mature and dispersed as such, provide no unequivocal evidence for affinities, be they tracheophyte or bryophyte. The bifurcating sporophytes are evidence against similarities with extant bryophytes. It is concluded that these very fragmentary fossils derive from small plants comprising relict populations of the vegetation that flourished on land in turfs through the greater part of the Ordovician and early Silurian, but that was gradually replaced by the tracheophytes.     

New species and biostratigraphic significance of middle Palaeozoic brachiopod genus Linguopugnoides Havliček

Linguopugnoides Havli?ek 1960, originally based on 3 Early Devonian (Lochkovian‐Pragian) species from the Czech Republic, is now construed as embracing at least 25 species distributed over all northern hemisphere continents as well as Australia, and extending from late Llandovery to early Emsian. Species‐diversity peaked during the Lochkovian when the genus was widespread, with 15 species having been reported. Four species are described as new. L. praecarens sp.nov. from the latest Silurian (Pridoli) of the Turkestan Range (South Tien Shan) is considered to be ancestral to the numerous, mainly late Lochkovian‐Pragian forms that appeared during the great diversification of the genus during the early Devonian. Three new species are described from the Lochkovian of the Zeravshan Range, Uzbekistan: L. rectiplicatus, L. diversiplicatus as well as the previously figured but not described L. kimi Rzhonsnitskaya. Species of Linguopugnoides have biostratigraphic and biogeographic significance and have obvious value for inter‐regional correlation.     

亚轮生奇异蕨(Adoketophyton subverticillatum)在贵州都匀泥盆系蟒山群的发现及其意义

Adoketophyton subverticillatum (Li et Cai) Li et Edwards (亚轮生奇异蕨)是云南早泥盆世坡松冲植物群中颇具代表性的、研究较为详细的一种植物, 目前对其整体形态特征和解剖结构都有较深入的认识。本文通过系列针修法, 将采自贵州都匀包阳剖面蟒山群下段的一块植物化石新材料归入A. subverticillatum。基于目前对 Adoketophyton时空分布的认识, 认为该属可作为早泥盆世植物群的代表分子, 并据此推断蟒山群下段的时代应为早泥盆世。Adoketophyton subverticillatum在都匀的发现, 表明贵州都匀蟒山群中的植物与云南坡松冲植物群在属、种一级上存在扩散与交流现象。Adoketophyton以及其他相关植物的研究表明, 在高大气CO₂浓度的早泥盆世, 叶性结构如孢子叶、苞片、枝叶复合体等已广泛分布于华南板块。     

Early Devonian (Lochkovian) trilobites and brachiopods from the west Balkhash Region,Kazakhstan and their significance for regional terrane dynamics

A moderately diverse Early Devonian (Lochkovian) trilobite and brachiopod association has been recovered from the lower part of the volcano-sedimentary Shakshagaily Formation, exposed on the western side of Lake Balkhash in central Kazakhstan. Its discovery demonstrates the presence of Lower Devonian marine deposits in the region. The trilobite assemblage includes representatives of six genera: the dalmanitid Kasachstania alperovichi nov. sp. is the dominant taxon, with other components being Ananaspis?, Leonaspis?, Maurotarion, Trimerus (Edgillia), and Warburgella. Associated brachiopods include species of Glossoleptaena, Leptaena, Protolepstostrophia, Pseudostrophochonetes, and Resserella. Whilst a significant proportion of these taxa show distinct links with the contemporaneous trilobite and brachiopod faunas of the north Balkhash Region, the occurrence of Trimerus (Edgillia) aff. kinglakensis Gill, 1949 suggests a possible link with the Lochkovian trilobite fauna of the Australasian sector of Gondwana. The west Balkhash Region preserves an extensive record of late Silurian to Early Devonian island arc volcanism, slope-rise sedimentation and olistostrome formation, which has invariably been ignored in models favouring early amalgamation of the Kazakhstanian terranes into a single continent by the end of the Ordovician.     

Permanent dyads in sporangia and spore masses from the Lower Devonian of the Welsh Borderland

Dyads, interpreted as cryptospore permanent dyads, are reported from sporangia and spore masses recovered from fluviatile deposits of Lochkovian (Early Devonian) age from the Welsh Borderland. The morphology, anatomy and ultrastructure of the specimens have been analysed using LM, SEM and TEM. Two specimens consist of a single sporangium terminating an axis: one is cup-shaped and attached to an unbranched axis and named Culullitheca richardsonii gen. et sp. nov., the other comprises an isotomously branching axis with one sporangium preserved and is named Fusitheca fanningiae gen. et sp. nov. Two specimens show dyads adhering to cuticular fragments presumably derived from sporangia. In one the cuticle is of irregular shape, in the other it is elliptical. The final specimen comprises an elongate spore mass. In all cases ultrastructural analysis reveals that the spore walls are essentially homogeneous. Our findings suggest that cryptospore permanent dyads were produced by rhyniophytoid plants of small stature. The affinities of these plants remain conjectural, as is the relationship of the cryptospore permanent dyads with other sporomorph morphotypes (cryptospores and trilete spores). However, the production of dyads by a plant with a bifurcating sporophytic axis is evidence against affinity with extant bryophytes. The mode of formation, adaptive significance, affinities, phylogenetic relationships and stratigraphical history (including demise) of cryptospore permanent dyads is discussed.     

New insights into early land ecosystems: a glimpse of a lilliputian world

Small, relatively uncompressed, very fragmentary plant remains (mesofossils) are described from a Silurian (P ídolí) and a Lower Devonian (Lochkovian) locality in the Welsh Borderland. Excellent cellular preservation provides characters leading to the demonstration of diversity in plants of simple gross morphology and allows deliberations on functional anatomy (e.g. of stomata), and reproductive biology (including development and dehiscence of sporangia). A survey of in-situ spores is presented, and preliminary comparisons made with dispersed spore assemblages especially in relation to reconstruction of vegetation on local and regional scales. The earliest body fossils of unequivocal terrestrial arthropods isolated from the same locality as the P ídolí plants suggest that the decomposer/microherbivore/predator soil and litter communities found in the Lower and Middle Devonian extend back at least into the Silurian. Evidence for plant—animal interaction in the Lower Devonian comes from spore-dominated coprolites believed to have been produced by litter-feeding myriapods.     

The phylogeny of aglaspidid arthropods and the internal relationships within Artiopoda

The phylogenetic position of aglaspidids, a problematic group of Lower Palaeozoic arthropods of undetermined affinities, is re‐examined in the context of the major Cambrian and Ordovician lamellipedian arthropod groups. A cladistic analysis of ten genera of aglaspidids sensu stricto, six aglaspidid‐like arthropods and 42 Palaeozoic arthropod taxa indicates that Xenopoda, Cheloniellida, Aglaspidida sensu lato and Trilobitomorpha form a clade (Artiopoda Hou and Bergström, 1997 ) nested within the mandibulate stem‐lineage, thus discarding previous interpretations of these taxa as part 'of the chelicerate stem‐group (Arachnomorpha Heider, 1913 ). The results confirm an aglaspidid identity for several recently described arthropods, including Quasimodaspis brentsae, Tremaglaspis unite, Chlupacaris dubia, Australaglaspis stonyensis and an unnamed Ordovician Chinese arthropod. The problematic Bohemian arthropod Kodymirus vagans was recovered as sister taxon to Beckwithia typa, and both form a small clade that falls outside Aglaspidida sensu stricto, thus discarding eurypterid affinities for the former. The analysis does not support the phylogenetic position of Kwanyinaspis maotianshanensis at the base of Conciliterga as proposed in recent studies, but rather occupies a basal position within Aglaspidida sensu lato. The results indicate a close association of aglaspidid arthropods with xenopods (i.e. Emeraldella and Sidneyia) and cheloniellids (e.g. Cheloniellon, Duslia); the new clade “Vicissicaudata” is proposed to encompass these arthropods, which are characterized by a differentiated posterior region. The phylogenetic position of aglaspidid arthropods makes them good outgroup candidates for analysing the internal relationships within the groups that form Trilobitomorpha. This work provides a much clearer picture of the phylogenetic relationships among Lower Palaeozoic lamellipedians.     

A diverse assemblage of early land plants from the Lower Devonian of the Welsh Borderland

FANNING, U., EDWARDS, D. & RICHARDSON, J. B., 1992. A diverse assemblage of early land plants from the Lower Devonian of the Welsh Borderland. Nine rhyniophytoid taxa are described from an early Gedinnian locality ( middle micromatus-newportensis spore Biozone) near Ludlow, England. They include Cooksonia pertoni, C. hemisphaerica, C. cambrensis, Tortilicaulis transwalliensis and three new taxa, Salopella marcensis sp. nov., Uskiella reticulata sp. nov. and Tarrantia salopensis gen. et sp. nov. Isolated sporangia of reniform shape, and those subtended by short lengths of axis, contain spores of Apiculiretusispora type and may belong to C. caledonica or Renalia . Morphologically distinctive forking, terminal sporangia lacking identifiable spores are not placed in a new taxon, because evidence based on in situ spores from elsewhere suggests they may belong to Salopella . Structures previously interpreted as clusters of sporangia of Yarravia- type are shown to be ± globular sporangia longitudinally split into valves. Sterile axes are dominated by smooth forms; although rare examples possess small enations. Tracheids have not been seen in axes of fertile specimens nor in sterile coalified compressions. A single pyrite permineralization contains tracheids of zosterophyll type. The assemblage demonstrates diversity among rhyniophytoids in the early Devonian and the existence of low vegetation 'alongside' the much larger zosterophyll dominated type.     

Teruelia diezii gen. et sp. nov.: an early polysporangiophyte from the Lower Devonian of the Iberian Peninsula

A new basal land plant, Teruelia diezii gen. et sp. nov., is described from the shallow‐water marine deposits of the Lower Devonian (Lochkovian–Pragian) Nogueras Formation of the Iberian Peninsula (north Gondwana palaeocontinent). Teruelia is preserved as a compression fossil and consists of isotomously branched, robust stems terminated in large, fusiform, twisted sporangia. This morphology suggests that Teruelia is very probably equivalent to Aglaophyton, a permineralized early polysporangiophyte known up to now only from the Lower Devonian (early Pragian to ?earliest Emsian) Rhynie Chert in Scotland (Laurussia palaeocontinent), which represents an early terrestrial hot‐spring ecosystem. Accepted phylogenies identify Aglaophyton as sister to vascular plants. Our phylogeny‐based results identify the Aglaophyton/Teruelia biological entity (i.e. Aglaophyton anatomical characters plus Teruelia external morphology) as the most direct vascular plant precursor. It shows that at least one Rhynie Chert type plant had a much wider distribution than previously known and suggests that Aglaophyton was not restricted to hydrothermal environments, unlike other Rhynie Chert plants.     

Multilocus sequence analysis of homologous recombination and diversity in Arthrobacter sensu lato named species and glacier-inhabiting strains

Members of the bacterial genus Arthrobacter sensu lato are Gram-positive actinomycetes distributed worldwide and found in numerous environments including soil, water, glacier ice, and sewage. Homologous recombination is an important driving force in bacterial evolution, but its impact on Arthrobacter sensu lato evolution is poorly understood. We evaluated homologous recombination among 41 Arthrobacter sensu lato named species, using multilocus sequence analysis (MLSA). A high level of recombination was found, associated with strong diversification and a reticulate evolutionary pattern of Arthrobacter sensu lato. We also collected a total of 31 cold-adapted Arthrobacter sensu lato strains from two cold glaciers located in northwest China and two temperate glaciers in southwest China, and evaluated their diversity and population structure by MLSA. The glacier strains displayed high diversity, but rates of recombination among the four glacier groups were quite low, indicating that barriers to homologous recombination formed in the past among the populations on different glaciers. Our findings indicate that historical glaciation events shaped the contemporary distributions, taxonomic relationships, and phylogeographic patterns of Arthrobacter sensu lato species on glaciers.     

Phylogeny and evolution of Mesozoic and extant lineages of Histeridae (Coleoptera), with discovery of a new subfamily Antigracilinae from the Lower Cretaceous

In order to place a newly discovered species Antigracilus costatus gen. sp. n. from the Lower Cretaceous Yixian Formation (China) and to assess previously unplaced fossil taxa, we investigated the relationships of extant and extinct lineages of Histeridae based on three data sets: (i) 69 morphological characters belonging to 48 taxa (representing all 11 subfamilies and 15 of 17 tribes of modern Histeridae); (ii) partitioned alignment of 6030 bp from downloaded nucleotide sequences (28S, CAD, COI, 18S) of 50 taxa (representing 10 subfamilies and 15 of 17 tribes of modern Histeridae); and (iii) a combined morphological and molecular dataset for 75 taxa. Phylogenetic analyses of the morphology and combined matrices recovered the new Lower Cretaceous taxon as a sister group to remaining Histeridae and it is placed in †Antigracilinae subfam. n. †Antigracilinae constitutes the earliest record of Histeridae from the Lower Cretaceous Yixian Formation (∼125 Myr), backdating the minimum age of the family by 25 Myr from the earliest Cenomanian (~99 Myr) to the Barremian of the Cretaceous Period. Our molecular phylogeny supports Histeridae to be divided into seven different clades, with currently recognised subfamilies Abraeinae (sensu lato), Saprininae, Chlamydopsinae, and Histerinae (sensu lato) recovered as monophyletic, while Dendrophilinae, Onthophilinae, and Tribalinae are polyphyletic taxa. The Burmese amber species †Pantostictus burmanicus Poinar & Brown is placed as a sister group to the tribe Plegaderini (Abraeinae) and was assigned as a new tribe Pantostictini trib. n. Both molecular and combined phylogenies recovered the subfamilies Trypanaeinae and Trypeticinae deeply within the subfamily Abraeinae (sensu lato), and they are downgraded into Trypanaeini stat. n. and Trypeticini stat. n.     

Geography determines genetic relationships between species of mountain pine (Pinus mugo complex) in western Europe

Aim Our aims were to test whether morphological species of mountain pines were genetically supported in the western part of the distribution range of the Pinus mugo species complex (Pinus mugo Turra sensu lato), to resolve genetically homogeneous clusters of populations, to determine historical demographic processes, and to assess the potential hybridization of mountain pines with Scots pine, Pinus sylvestris L. Location Populations were sampled in the Iberian System, the Pyrenees, the French Mont Ventoux, Vosges and Jura mountains, the German Black Forest and throughout the Alps. This corresponded to a range‐wide sampling for mountain pine sensu stricto (Pinus uncinata Ram.) and to a sampling of the western parts of the ranges of dwarf mountain pine (Pinus mugo Turra sensu stricto) and bog pine/peatbog pine [Pinus rotundata Link/Pinus × pseudopumilio (Willk.) Beck]. Methods In total, 786 individuals of P. mugo sensu lato from 29 natural populations, and 85 individuals of P. sylvestris from four natural populations were genotyped at three chloroplast microsatellites (cpSSRs). Populations were characterized for standard genetic diversity statistics and signs of demographic expansion. Genetic structure was explored using analysis of molecular variance, differentiation statistics and Bayesian analysis of population structure (BAPS). Results One hundred haplotypes were identified in P. mugo sensu lato. There was a stronger differentiation between geographical regions than between morphologically identified taxa (P. mugo sensu stricto, P. uncinata and P. rotundata/P. ×pseudopumilio). Overall genetic differentiation was weak (G_ST = 0.070) and displayed a clear phylogeographic structure [N_ST = 0.263, N_ST > N_ST (permuted), P < 0.001]. BAPS identified a Pyrenean and an Alpine gene pool, along with several smaller genetic clusters corresponding to peripheral populations. Main conclusions The core regions of the Pyrenees and Alps were probably recolonized, respectively by P. uncinata and P. uncinata/P. mugo sensu stricto, from multiple glacial refugia that were well connected by pollen flow within the mountain chains. Pinus rotundata/P. × pseudopumilio populations from the Black Forest, Vosges and Jura mountains were probably recolonized from various glacial populations that kept their genetic distinctiveness despite late glacial and early Holocene expansion. Marginal P. uncinata populations from the Iberian System are compatible with elevational shifts and long‐term isolation. The causes of haplotype sharing between P. mugo sensu lato and P. sylvestris require further research.     

Exceptional preservation in Lower Devonian coalified fossils from the Welsh Borderland: a new genus based on reniform sporangia lacking thickened borders

Reniform sporangia, comprising two equal valves and containing retusoid spores, recovered from Lower Old Red Sandstone strata of Devonian age ( micrornatus-newportensis Spore Biozone: lower Gedinnian lower Lochkovian) on North Brown Clee Hill in the Welsh Borderland are placed in Resilitheca salopensis gen. et sp. nov. Conventional compression fossils from Targrove, Ludlow of fertile axes showing isotomous branching with limited overtopping are considered conspecific because the terminal reniform sporangia contain the same spores. Spore ultrastructure is described using scanning and transmission electron microscopy. Sections show faint traces of lamellae. Particles associated with spores and sporangium wall are compared with the globules of pteridophytes and Ubisch bodies of angiosperms, and related to the development of the sporangium. The new plants are compared with Cooksonia caledonica Edwards known only from impressions, and with Renalia Gensel showing far more pronounced pseudomonopodial branching.