Climate change and C4 and C3 grasses in a midlatitude dryland steppe.

Climate change is projected to alter the structure of plant communities due to increasing temperatures and changes to precipitation patterns, particularly in midlatitude dryland ecosystems. Modifications to climatic suitability may lead to major community changes such as altered dominant plant functional types. Previous studies have indicated that climatic suitability is likely to increase for C4 grasses and decrease for C3 grasses in the Western United States. However, if no C4 grass species currently exist to serve as a propagule source, expansion into areas of increased suitability will be limited. We conducted a field and modeling study in the Upper Green River Basin (UGRB) of Western Wyoming to determine if (1) C4 grasses are present to provide a propagule source and (2) C4 grasses are likely to increase in importance relative to C3 grasses due to climatic changes. We searched 44 sites for C4 grasses to establish presence, and modeled suitability at 35 sites using 17 Global Climate Models, two greenhouse gas Representative Concentration Pathways (RCPs; 4.5 and 8.5), and two time periods (mid- and late century; 2030-2060 and 2070-2099, respectively). We found C4 grasses at 10 of the 44 sites, indicating that there is a present propagule source. Our model projected increases in suitability for both C3 and C4 grasses across sites for all RCPs and time periods. In the mid-century RCP 4.5 scenario, the C3 functional type increased in projected biomass in 29 of 35 sites, and the C4 type increased in 31 sites. In this scenario, C3 grasses increased in projected biomass by a median 4 g m-2 (5% change), and C4 grass biomass increased by a median 8 g m-2 (21% change). Our study suggests that climate change will increase climatic suitability for grasses across the UGRB, and that all requirements are in place for C4 grasses to increase in abundance.

Phylogenetic assessment of endangered and look-alike Pigtoe species in a freshwater mussel diversity hotspot.

The Green River in Kentucky in the eastern United States is a freshwater mussel biodiversity hotspot, with 71 known species. Among them, the endangered Pleurobema plenum coexists with other morphologically similar species in the genera Fusconaia and Pleurobema, known colloquially as "pigtoes." Identification of species in these genera is challenging even for mussel experts familiar with them. In our study, the correct identification of these species by experts ranged from 57% to 83%. We delineated taxonomic boundaries among seven species and tested for cryptic biodiversity among these look-alike mussels utilizing mitochondrial and nuclear DNA sequence variation. Phylogenetic analysis of combined (1215 bp) mitochondrial DNA cytochrome oxidase I (COI) and NADH dehydrogenase 1 (ND1) genes showed five well-diverged groups that included F. flava, F. subrotunda, P. cordatum, and P. plenum as distinct clades, with P. sintoxia and P. rubrum grouped into a single clade. While our mitochondrial DNA analyses did not distinguish P. sintoxia and P. rubrum as phylogenetically distinct species, the typical shell forms of these two nominal taxa are very distinct. Further phylogenetic analysis using nuclear ribosomal transcribed spacer region subunit I (ITS1) DNA sequences also showed that P. sintoxia and P. rubrum were not distinct lineages. No cryptic species were detected in the Fusconaia and Pleurobema samples analyzed from the Green River. The highest haplotype diversity (h), average number of nucleotide differences (k), and nucleotide diversity (π) were observed for F. subrotunda at both the COI (h = 0.896, k = 3.805, π = 0.00808) and ND1 (h = 0.984, k = 6.595, π = 0.00886) markers, with similarly high genetic diversity in the other taxa. Our results give managers confidence that cryptic taxa do not occur within or among these morphologically similar species in the Green River, and populations appear genetically diverse, indicative of large and healthy populations.

Bizarre morphology in extinct Eocene bugs (Heteroptera: Pentatomidae).

Newly discovered fossil bugs (Insecta: Hemiptera: Heteroptera: Pentatomidae) from the Eocene of Messel (Germany) and Green River (North America) exhibit an exaggerated morphology including prominent spiny humeral and anterolateral angles of the pronotum and a spiny lateral abdominal margin. Especially the humeral angles are unique; they consist of expansive, rounded projections with strong spines, which is a rare trait among pentatomids. A hypothesis for the function of this extreme morphology is defence against small vertebrate predators, such as birds or reptiles. The same protuberances also produce a disruptive effect camouflaging the specimen in its environment and provide additional protection. Therefore, the extreme morphology provides primary as well as secondary anti-predator defence. The morphology of Eospinosus peterkulkai gen. et sp. nov. and E. greenriverensis sp. nov. resembles that of Triplatygini, which today occur exclusively in Madagascar, as well as that of Discocephalinae or Cyrtocorinae, which today occur in the Neotropics. Due to a lack of conclusive characters, it cannot be excluded that the fossil species may represent a case of remarkable convergence and are not related to either taxon. Phylogenetic analyses using parsimony as well as Bayesian algorithms confirmed that the new genus is a member of Pentatomidae, but could not solve its phylogenetic relationships within Pentatomidae.

Estimating Flight Style of Early Eocene Stem Palaeognath Bird Calciavis grandei (Lithornithidae).

Lithornithids are volant stem palaeognaths from the Paleocene-Eocene. Except for these taxa and the extant neotropical tinamous, all other known extinct and extant palaeognaths are flightless. Investigation of properties of the lithornithid wing and its implications for inference of flight style informs understood locomotor diversity within Palaeognathae and may have implications for estimation of ancestral traits in the clade. Qualitative comparisons with their closest extant volant relatives, the burst-flying tinamous, previously revealed skeletal differences suggesting lithornithids were capable of sustained flight, but quantitative work on wing morphology have been lacking. Until comparatively recently, specimens of lithornithids preserving wing feather remains have been limited. Here, we reconstruct the wing of an exceptionally preserved specimen of the Early Eocene lithornithid Calciavis grandei and estimate body mass, wing surface area, and wing span. We then estimate flight parameters and compare our estimates with representatives from across Aves in a statistical framework. We predict that flight in C. grandei was likely marked by continuous flapping, and that lithornithids were capable of sustained flight and migratory behavior. Our results are consistent with previous hypotheses that the ancestor of extant Palaeognathae may also have been capable of sustained flight. Anat Rec, 303:1035-1042, 2020. © 2019 Wiley Periodicals, Inc.

Phosphatized tungsten-metabolizing coccoid microbes interpreted from oil shale of an Eocene lake, Green River Formation, Utah, USA.

Microscopic globular structures have been observed in some beds of oil shale from eastern Utah. These beds comprise carbonate-dominated mud that is interlaminated with variably thick and continuous organic-rich layers. Collectively they are enriched in phosphorus, REEs, and actinides. The beds are considered of lacustrine origin and assigned to the Eocene Green River Formation. The globules themselves are of microcrystalline carbonate fluorapatite (µCFA), often contain concentric internal structures, and usually group together in clusters of up to 80, possibly more. Detailed SEM and microprobe analyses have revealed tungsten (W) to be almost exclusively associated with the globular clusters found within the more organic-rich laminae, often at concentrations of over 200 ppm, two orders of magnitude above shale standards. The globular structures are present in freshly cut sections where they occasionally grade into a µCFA matrix cement. This, together with the draping of the clusters by stringers of organic matter that would have accumulated in the Eocene lake, confirms that the structures are not a contaminant. The limited range of sizes and globular shapes is consistent with the morphology of coccoidal bacteria: Concentric internal structures may represent remnants of the nucleoid and cell wall. Paired concentric structures may indicate cell division (reproduction) processes were occurring until mineralization. The phosphate mineralization itself may have been promoted by release of phosphate from the stressed cells, bringing porewaters to supersaturation, or by the cells acting as nucleation sites. The recording of trace amounts of W almost exclusively in globular clusters preserved in the most organic-rich stringers (anoxia prone) further suggests facultative use of W-enzymes in a microbial metabolism. Combined, their context, morphology, and indication of biogenic process are strong evidence that the structures are fossilized (phosphatized) microbes, possibly sulfate-reducing bacteria, or methanogenic archaea.

Systematics and phylogeny of the Zygodactylidae (Aves, Neognathae) with description of a new species from the early Eocene of Wyoming, USA.

Zygodactylidae are an extinct lineage of perching birds characterized by distinct morphologies of the foot and wing elements. Although the clade has a complex taxonomic history, current hypotheses place Zygodactylidae as the sister taxon to Passeriformes (i.e., songbirds). Given the rather sparse fossil record of early passeriforms, the description of zygodactylid taxa is important for inferring potentially ancestral states in the largest radiation of living birds (i.e., the ∼6,000 species of extant passeriforms). Despite the exceptional preservation of many specimens and considerable species diversity in Zygodactylidae, the relationships among species have not been previously evaluated in a phylogenetic context. Herein, we review the fossil record of Zygodactylidae from North America and describe five new well-preserved fossils from the early Eocene Green River Formation of Wyoming. Two specimens are identified as representing a new species and the first records of the taxon Zygodactylus outside Europe. Anatomical comparisons with previously named taxa and the results of phylogenetic analysis including newly described specimens and previously named zygodactylid taxa provide the first hypothesis of the species-level relationships among zygodactylids. The monophyly of Zygodactylidae is supported in these new analyses. However, the monophyly of Primozygodactylus and the taxonomic distinction between Zygodactylus and Eozygodactylus remain unresolved and would likely benefit from the description of additional specimens.

Oldest new genus of Myrmeleontidae (Neuroptera) from the Eocene Green River Formation.

Epignopholeon sophiae gen. et sp. nov. (Neuroptera: Myrmeleontidae) is described from the early Eocene of the Green River Formation (Colorado, U.S.A.). It represents the oldest confident record of the family. The new genus is remarkable in that tergite 7 of the female is much shorter than its long sternite 7. The preserved wing venation shows that the genus belongs to the subfamily Myrmeleontinae, and most probably to the tribe Gnopholeontini. The discovery of this species is consistent with estimations of relatively dry and warm conditions during deposition of the upper Parachute Creek Member of the Green River Formation.

An interesting new genus of Berothinae (Neuroptera: Berothidae) from the early Eocene Green River Formation, Colorado.

Xenoberotha angustialata gen. et sp. nov. (Neuroptera: Berothidae) is described from the early Eocene of the Parachute Creek Member of the Green River Formation (U.S.A., Colorado). It is assigned to Berothinae as an oldest known member of the subfamily based on the presence of scale-like setae on the foreleg coxae. Distal crossveins of the fourth (outer) gradate series which are located very close to the wing margin in Xenoberotha gen. nov. is a character state previously unknown in Berothinae.