Reconstruction of an enigmatic Pennsylvanian cone reveals a relationship to Sphenophyllales.

PREMISE: We studied the 3D morphology of a small, well-preserved cone from the Pennsylvanian Mazon Creek Lagerstätte to characterize its structure and determine its systematic affinity. Previously tentatively assigned to the enigmatic Tetraphyllostrobus, we show that it differs in key respects from that genus as described. METHODS: We systematically compared the new fossil with relevant Paleozoic cone genera and employed advanced imaging techniques, including scanning electron microscopy, Airyscan confocal super-resolution microscopy, optical microscopy, and X-ray microcomputed tomography to visualize and reconstruct the fossil cone in 3D. RESULTS: The analyses demonstrate unequivocally that the sporophylls of the new Mazon Creek cone are arranged in whorls of six and have characters typical of Sphenophyllales, including epidermal cells with undulatory margins and in situ spores assignable to Columinisporites. The combination of characters, including sporophyll arrangement, anatomy, and spore type, supports the establishment of Hexaphyllostrobus kostorhysii gen. et sp. nov. within Sphenophyllales. Furthermore, we show that Tetraphyllostrobus, although originally described as possessing smooth monolete spores, actually possesses Columinisporites-type spores, indicating that it, too, was most likely a sphenophyll. CONCLUSIONS: The recognition of Hexaphyllostrobus contributes to our knowledge of Pennsylvanian sphenophyll diversity, and in particular increases the number of species with in situ Columinisporites-type spores. Attribution of Hexaphyllostrobus to Sphenophyllales calls into question current interpretations of Tetraphyllostrobus suggesting that future research on better-preserved macrofossil material may demonstrate a sphenophyllalean relationship.

NCBI Taxonomy: a comprehensive update on curation, resources and tools.

The National Center for Biotechnology Information (NCBI) Taxonomy includes organism names and classifications for every sequence in the nucleotide and protein sequence databases of the International Nucleotide Sequence Database Collaboration. Since the last review of this resource in 2012, it has undergone several improvements. Most notable is the shift from a single SQL database to a series of linked databases tied to a framework of data called NameBank. This means that relations among data elements can be adjusted in more detail, resulting in expanded annotation of synonyms, the ability to flag names with specific nomenclatural properties, enhanced tracking of publications tied to names and improved annotation of scientific authorities and types. Additionally, practices utilized by NCBI Taxonomy curators specific to major taxonomic groups are described, terms peculiar to NCBI Taxonomy are explained, external resources are acknowledged and updates to tools and other resources are documented. Database URL: https://www.ncbi.nlm.nih.gov/taxonomy.

Life habits and evolutionary biology of new two-winged long-proboscid scorpionflies from mid-Cretaceous Myanmar amber.

Long-proboscid scorpionflies are enigmatic, mid-Mesozoic insects associated with gymnosperm pollination. One major lineage, Aneuretopsychina, consists of four families plus two haustellate clades, Diptera and Siphonaptera. One clade, Pseudopolycentropodidae, from mid-Cretaceous Myanmar amber, contains Parapolycentropus. Here, we newly establish Dualula, assigned to Dualulidae, constituting the fifth lineage. Parapolycentropus and Dualula lineages are small, two-winged, with unique siphonate mouthparts for imbibing pollination drops. A cibarial pump provides siphonal food inflow; in Dualula, the siphon base surrounds a hypopharynx housing a small, valved pump constricted to a narrow salivary duct supplying outgoing enzymes for food fluidization. Indirect evidence links long-proboscid mouthpart structure with contemporaneous tubulate ovulate organs. Direct evidence of gymnospermous Cycadopites pollen is associated with one Parapolycentropus specimen. Parapolycentropus and Dualula exhibit hind-wing reduction that would precede haltere formation, likely caused by Ultrabithorax. Distinctive, male Aneuretopsychina genitalia are evident from specimens in copulo, supplemented by mixed-sex individuals of likely male mating swarms.

Paleoecological and paleoenvironmental interpretation of three successive macrofloras and palynofloras from the Kola Switch locality, lower Permian (Archer City Formation, Bowie Group) of Clay County, Texas, USA.

Fossil floras have been recovered from a unique deposit of early Permian age in North-Central Texas. The site, Kola Switch, preserves three distinct floras in different lithofacies, in a succession from a single outcrop. The sedimentary environment appears to be a floodplain channel fill of primarily siltstones and claystones. The lowermost flora, preserved in a kaolinitic siltstone, indicates active water flow. It is dominated by plants typical of well-drained substrates, dominated by Sphenopteris germanica, and contains no wetland elements. The middle flora is from a finely laminated carbonaceous claystone and is dominated by marattialean tree ferns, with no elements from habitats typical of seasonal moisture availability. It contains no roots and appears to have formed as a floating peat mat. The upper flora is a mixed assemblage of wetland taxa and those typical of well-drained soil environments or a seasonal rainfall regime. Unlike the two lower floras, it has a relatively even distribution of dominance and is the most diverse of the three assemblages. Palynofloras also were recovered from each of these beds. The palynofloras, although varying between and even within the beds, indicate a common background species pool during the time interval sampled, suggesting that these distinct floras reflect local changes in microhabitat conditions under a constant climatic background. The palynoflora from each bed has characteristics in common with the macroflora of that bed, but also distinct differences. Together, the macroflora and microflora provide an unusually broad picture of this site through time. Kola Switch compares favorably with the recently described flora from the nearby Sanzenbacher Ranch site of approximately the same age and also with floras of Rotliegend age from Central Europe.

The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies.

Mid-Mesozoic kalligrammatid lacewings (Neuroptera) entered the fossil record 165 million years ago (Ma) and disappeared 45 Ma later. Extant papilionoid butterflies (Lepidoptera) probably originated 80-70 Ma, long after kalligrammatids became extinct. Although poor preservation of kalligrammatid fossils previously prevented their detailed morphological and ecological characterization, we examine new, well-preserved, kalligrammatid fossils from Middle Jurassic and Early Cretaceous sites in northeastern China to unravel a surprising array of similar morphological and ecological features in these two, unrelated clades. We used polarized light and epifluorescence photography, SEM imaging, energy dispersive spectrometry and time-of-flight secondary ion mass spectrometry to examine kalligrammatid fossils and their environment. We mapped the evolution of specific traits onto a kalligrammatid phylogeny and discovered that these extinct lacewings convergently evolved wing eyespots that possibly contained melanin, and wing scales, elongate tubular proboscides, similar feeding styles, and seed-plant associations, similar to butterflies. Long-proboscid kalligrammatid lacewings lived in ecosystems with gymnosperm-insect relationships and likely accessed bennettitalean pollination drops and pollen. This system later was replaced by mid-Cretaceous angiosperms and their insect pollinators.

Spatiotemporal relationships among Late Pennsylvanian plant assemblages: Palynological evidence from the Markley Formation, West Texas, U.S.A.

The Pennsylvanian lowlands of western Pangea are best known for their diverse wetland floras of arborescent and herbaceous ferns, and arborescent horsetails and clubmosses. In apparent juxtaposition, a very different kind of flora, dominated by a xerophilous assemblage of conifers, taeniopterids and peltasperms, is occasionally glimpsed. Once believed to represent upland or extrabasinal floras from well-drained portions of the landscape, these dryland floras more recently have been interpreted as lowland assemblages growing during drier phases of glacial/interglacial cycles. Whether Pennsylvanian dryland and wetland floras were separated spatially or temporally remains an unsettled question, due in large part to taphonomic bias toward preservation of wetland plants. Previous paleobotanical and sedimentological analysis of the Markley Formation of latest Pennsylvanian (Gzhelian) age, from north central Texas, U.S.A, indicates close correlation between lithofacies and distinct dryland and wetland megaflora assemblages. Here we present a detailed analysis one of those localities, a section unusual in containing abundant palynomorphs, from the lower Markley Formation. Paleobotanical, palynological and lithological data from a section thought to represent a single interglacial/glacial phase are integrated and analyzed to create a complex picture of an evolving landscape. Megafloral data from throughout the Markley Formation show that conifer-dominated dryland floras occur exclusively in highly leached kaolinite beds, likely eroded from underlying soils, whereas a mosaic of wetland floras occupy histosols, ultisols, and fluvial overbank deposits. Palynological data largely conform to this pattern but reveal a more complex picture. An assemblage of mixed wetland and dryland palynofloral taxa is interpolated between a dryland assemblage and an overlying histosol containing wetland taxa. In this section, as well as elsewhere in the Markley Formation, kaolinite and overlying organic beds appear to have formed as a single genetic unit, with the kaolinite forming an impermeable aquiclude upon which a poorly drained wetland subsequently formed. Within a single inferred glacial/interglacial cycle, lithological data indicate significant fluctuations in water availability tracked by changes in palynofloral and megafloral taxa. Palynology reveals that elements of the dryland floras appear at low abundance even within wetland deposits. The combined data indicate a complex pattern of succession and suggest a mosaic of dryland and wetland plant communities in the Late Pennsylvanian. Our data alone cannot show whether dryland and wetland assemblages succeed one another temporally, or coexisted on the landscape. However, the combined evidence suggests relatively close spatial proximity within a fragmenting and increasingly arid environment.

Conservatism of Late Pennsylvanian vegetational patterns during short-term cyclic and long-term directional environmental change, western equatorial Pangea.

Patterns of plant distribution by palaeoenvironment were examined across the Pennsylvanian-Permian transition in North-Central Texas. Stratigraphically recurrent packages of distinct lithofacies, representing different habitats, contain qualitatively and quantitatively different macrofloras and microfloras. The species pools demonstrate niche conservatism, remaining closely tied to specific habitats, during both short-term cyclic environmental change and a long-term trend of increasing aridity. The deposits examined principally comprise the terrestrial Markley and its approximate marine equivalent, the Harpersville Formation and parts of lower Archer City Formation. Fossiliferous deposits are lens-like, likely representing fill sequences of channels formed during abandonment phases. Palaeosols, represented by blocky mudstones, comprise a large fraction of the deposits. They suggest progressive climate change from minimally seasonal humid to seasonal subhumid to seasonal dry subhumid. Five lithofacies yielded plants: kaolinite-dominated siltstone, organic shale, mudstone beds within organic shale, coarsening upward mudstone-sandstone interbeds and channel sandstone. Both macro- and microflora were examined. Lithofacies proved compositionally distinct, with different patterns of dominance diversity. Organic shales (swamp deposits), mudstone partings (swamp drainages) and coarsening upward mudstone-sandstone interbeds (floodplains) typically contain Pennsylvanian wetland vegetation. Kaolinite-dominated siltstones and (to the extent known) sandstones contain taxa indicative of seasonally dry substrates. Some kaolinite-dominated siltstones and organic shales/coals yielded palynomorphs. Microfloras are more diverse, with greater wetland-dryland overlap than macrofloras. It appears that these two floras were coexistent at times on the regional landscape.

A probable pollination mode before angiosperms: Eurasian, long-proboscid scorpionflies.

The head and mouthpart structures of 11 species of Eurasian scorpionflies represent three extinct and closely related families during a 62-million-year interval from the late Middle Jurassic to the late Early Cretaceous. These taxa had elongate, siphonate (tubular) proboscides and fed on ovular secretions of extinct gymnosperms. Five potential ovulate host-plant taxa co-occur with these insects: a seed fern, conifer, ginkgoopsid, pentoxylalean, and gnetalean. The presence of scorpionfly taxa suggests that siphonate proboscides fed on gymnosperm pollination drops and likely engaged in pollination mutualisms with gymnosperms during the mid-Mesozoic, long before the similar and independent coevolution of nectar-feeding flies, moths, and beetles on angiosperms. All three scorpionfly families became extinct during the later Early Cretaceous, coincident with global gymnosperm-to-angiosperm turnover.