Past ecosystems drive the evolution of the early diverged Symphyta (Hymenoptera: Xyelidae) since the earliest Eocene

Paleoxyela nearctica gen. et sp. nov., is described from the upper Eocene of Florissant Formation in Colorado. We placed Paleoxyela gen. nov. in the subfamily Macroxyelinae and the tribe Macroxyelini based on the numerous wing venation characters visible on the specimen. Proxyelia pankowskii gen. et sp. nov. is described from the lower Eocene Fossil Lake deposits of the Green River Formation in Wyoming. We placed Proxyelia gen. nov. in the subfamily Macroxyelinae and the tribe Xyeleciini based on the numerous wing venation characters visible on the specimen. These new records of the family Xyelidae are of particular importance to better understand the past diversity of the clade and propose hypotheses about their diversification. Extant Xyelidae inhabit temperate Northern Hemisphere forests, and most of their larvae feed on conifers, which may explain why they are relatively poorly diversified compared to the other symphytan families. We suggest that the global decline in conifers and the reduced diversity of extant host trees partly explain the diversity of extant Xyelidae. We correlate the biome repartition during the Eocene to that of the extant xyelid.

The Fossil Record of Elateridae (Coleoptera: Elateroidea): Described Species, Current Problems and Future Prospects

The Elateridae (click-beetles) are the largest family in Elateroidea; however, their relationships, systematics and classification remain unclear. Our understanding of the origin, evolution, palaeodiversity and palaeobiogeography of Elateridae, as well as reconstruction of a reliable time-calibrated phylogeny for the group, are hampered by the lack of detailed knowledge of their fossil record. In this study, we summarize the current knowledge on all described fossil species in Elateridae, including their type material, geographic origin, age, bibliography and remarks on their systematic placement. Altogether, 261 fossil species classified in 99 genera and nine subfamilies are currently listed in this family. The Mesozoic click-beetle diversity includes 143 species, with most of them described from the Jurassic Karatau, and 118 described species are known from the Cenozoic deposits, mainly from the Eocene North American Florissant Formation and European Baltic amber. Available data on the described past diversity of Elateridae suggest that almost all fossil lineages in this group are in urgent need of revision and numerous Mesozoic species might belong to different families. Our study is intended to serve as a comprehensive basis for all subsequent research focused on the click-beetle fossil record.

The first described turtle beetles from Eocene Baltic amber, with notes on fossil Chelonariidae (Coleoptera: Byrrhoidea)

Chelonariidae, or turtle beetles, are rarely represented in the fossil record. Two new extinct species of this thermophilous coleopteran family, Chelonarium andabata Alekseev and Bukejs sp. nov. and Ch. dingansich Alekseev and Bukejs sp. nov., are described and illustrated from Eocene Baltic amber using X-ray micro-computed tomography (micro-CT). They are the first formally described species of turtle beetles from Eocene Baltic amber and the first known European representatives of this family. Based on modern habitats of the group, the presence of the plants with which their larvae are associated (epiphytic orchids) is proposed in the Eocene amber forest. The Eocene Florissant Formation fossil Chelonarium montanum Wickham, 1914, which was originally placed within Chelonariidae, is discussed based on its original description, and placement as incertae sedis within Byrrhoidea is proposed for this compression fossil (http://zoobank.org/References/C2EE164D-59DD-42FE-937D-B01C78DCD228, last access: 8 February 2021).

The Fossil Record of Darkling Beetles (Insecta: Coleoptera: Tenebrionidae)

The fossil record of Tenebrionidae (excluding the Quartenary) is presented. In total, 122 fossil species, clearly belonging to the family, are known; some beetles were determined only to genus; 78 genera are listed in the fossil record, including 29 extinct genera. The great diversity of tenebrionids occurs in the Lower Cretaceous Lagerstätte of China (Yixian Formation), Middle Paleocene of France (Menat), Lower Eocene deposits of Germany (Geiseltal), Upper Eocene Baltic amber (Eastern Europe), Upper Eocene deposits of Florissant Formation (USA) and Miocene (Dominican amber). Tenebrionids of the following major lineages, including seven subfamilies, are currently known in the fossil record. These include the lagrioid branch (Lagriinae, Nilioninae), pimelioid branch (Pimeliinae), and tenebrioid branch (Alleculinae, Tenebrioninae, Diaperinae, Stenochiinae). The importance of the fossil record for evolutionary reconstructions and phylogenetic patterns is discussed. The oldest Jurassic and Early Cretaceous darkling beetles of the tenebrionoid branch consist of humid-adapted groups from the extant tribes Alleculini, Ctenopodiini (Alleculinae), and Alphitobiini (Tenebrioninae). Thus, paleontological evidence suggests that differentiation of the family started at least by the Middle Jurassic but does not indicate that xerophilic darkling beetles differentiated much earlier than mesophilic groups.

Paleoclimate and paleoecology of the latest Eocene Florissant flora of Central Colorado, U.S.A.

The uppermost Eocene Florissant Formation of central Colorado, U.S.A. contains a diverse flora and fauna preserved in lacustrine facies and represents a key episode in Earth history immediately preceding the Eocene-Oligocene boundary. Laminated shales contain impressions of non-monocot angiosperm leaves that were used to estimate paleoecological and paleoclimatic parameters using leaf physiognomic methods including: leaf mass per area (MA), digital leaf physiognomy (DiLP), leaf margin analysis (LMA), and leaf area analysis (LAA). The majority (58%) of the morphotypes analyzed for MA suggested a semi-evergreen leaf lifespan, whereas another 27% indicated a deciduous habit and 15% an evergreen habit. There was no significant relationship between MA and insect damage based on a small subset of Florissant’s leaves. Higher MA values (~73% of leaves ≥ one-year lifespan), coupled with a tendency toward long and narrow leaf shapes and small leaf areas, indicate the existence of sclerophyllous vegetation. Using the global regression for mean annual temperature (MAT), the DiLP estimate of MAT was anomalously cold: 5.5 ± 4 °C. However, using a Northern Hemisphere regression the DiLP MAT estimate of 11.6 ± 3.3 °C was more plausible. Using DiLP, mean annual precipitation (MAP) was estimated at 740 +608/-334 mm∙yr-1, which supports dry conditions. Estimates for MAT and MAP using the univariate LMA and LAA methods overlapped within uncertainty of the DiLP results. In addition, those taxa classified as growing in wet areas (riparian) had significantly more teeth than non-riparian taxa. These paleoclimatic and paleoecological results suggest that outside the riparian forest, the Florissant flora sampled a seasonally dry temperate sclerophyllous shrubland to woodland, perhaps similar to modern chaparral forests, in the western interior of the U.S.A. just before the transition into the cooler Oligocene.

Freshwater molluscan fauna from the Florissant Formation, Colorado: paleohydrologic reconstruction of a latest Eocene lake

The freshwater molluscan assemblage from the uppermost Eocene Florissant Formation (34.07 ± 0.10 Ma), Colorado, USA, provides a reliable proxy in reconstructing past ecology and environmental characteristics of ancient Lake Florissant. In particular, stable-isotope analyses of aragonitic shell material contribute to our understanding of the paleohydrologic history of this ancient lake. Re-examination of molluscan taxonomy in the middle shale and caprock conglomerate (informal) units produces three sphaeriid bivalves (family Sphaeriidae, genus Sphaerium) and two pulmonate gastropod genera (family Planorbidae, genus Gyraulus; and family Lymnaeidae, genus Lymnaea). The middle-shale assemblage, representing quiet-lake deposition, is dominated by pulmonate gastropods; the shell material in all specimens in this unit has been replaced by silica. The caprock conglomerate assemblage, representing redeposition by a debris flow, is dominated by bivalves; specimens within the caprock conglomerate unit are aragonite, interpreted to be biogenic (original). Carbon and oxygen isotope analyses of the aragonite show a strong covariance (all molluscan data: r = 0.83; sphaeriids: r = 0.76) with consistent grouping (δ13C: 0‰ to −5.5‰, δ18O: +2‰ to −5‰) from all families. This result indicates that ancient Lake Florissant was “closed” and that evaporation had a stronger effect on isotopic values than precipitation, for the duration of the sampled interval. This finding is in agreement with our current understanding of how the sediments of the Florissant Formation were deposited within a lake that formed because of a paleoriver being dammed by debris flows.

A New (Hovenia) and an old (Chadronoxylon) Fossil Wood from the Late Eocene Florissant Formation, Colorado, U.S.A.

A fossil wood with features similar to those of the Oligocene Hovenia palaeodulcis Suzuki (Rhamnaceae) from Japan is described from the late Eocene Florissant Fossil Beds National Monument, Colorado, U.S.A. This is the first report of fossil wood of this Asian genus in North America and is further documentation of Tertiary exchange between East Asia and North America. The affinities of Chadronoxylon florissantensis, the most common angiosperm wood at Florissant, are reevaluated; its combination of features suggests relationships with two families in the Malpighiales, the Salicaceae and Phyllanthaceae. Chadronoxylon is compared with Paraphyllanthoxylon Bailey. The Eocene P. hainanensis from China has notable differences from the original diagnosis of Paraphyllanthoxylon, but shares features with Chadronoxylon warranting transfer of P. hainanensis to Chadronoxylon and the creation of Chadronoxylon hainanensis (Feng, Yi, Jen) Wheeler & Meyer, comb. nov.