Temperate Genera Research Articles

The oldest Rhantus (Coleoptera, Dytiscidae) from the earliest Eocene Fur Formation, Denmark

†Rhantus villumi sp. nov. is described and illustrated on the basis of a single specimen from the earliest Eocene Fur Formation, Denmark. With an estimated age of ca. 55.4 Ma, it represents the oldest member of an extant genus of the family Dytiscidae. However, the presence of Rhantus in the early Eocene is only slightly older than previous estimates, and generally agrees well with phylogenetic analyses of the subfamily Colymbetinae. The presence of a predominantly temperate genus in the presumably warm Lagerstätte is briefly discussed, supporting the hypothesis of temporary cooling, as suggested by several other invertebrate records from the Fur Formation. It is suggested that the new species could feed on mosquito larvae, which are known from the Lagerstätte. Finally, a record of another Dytiscidae species from the same locality is mentioned, but, due to poor preservation and lack of diagnostic characters, this fossil remains unidentified to genus or species level.

A reappraisal of the phylogeny and historical biogeography of Sparganium (Typhaceae) using complete chloroplast genomes

BackgroundSparganium (Typhaceae) is a widespread temperate genus of ecologically important aquatic plants. Previous reconstructions of the phylogenetic relationships among Sparganium species are incompletely resolved partly because they were based on molecular markers comprising < 7,000 bp. Here, we sequenced and assembled the complete chloroplast genomes from 19 Sparganium samples representing 15 putative species and three putative subspecies in order to explore chloroplast genome evolution in this genus, clarify taxonomic lineages, estimate the divergence times of Sparganium species, and reconstruct aspects of the biogeographic history of the genus.ResultsThe 19 chloroplast genomes shared a conserved genome structure, gene content, and gene order. Our phylogenomic analysis presented a well-resolved phylogeny with robust support for most clades. Non-monophyly was revealed in three species: S. erectum, S. eurycarpum, and S. stoloniferum. Divergence time estimates suggest that the two subgenera of Sparganium split from each other ca. 30.67 Ma in the middle Oligocene. The subgenus Xanthosparganium diversified in the late Oligocene and Miocene, while the subgenus Sparganium diversified in the late Pliocene and Pleistocene. Ancestral area reconstruction suggested that the two subgenera may have originated in East Eurasia and North America.ConclusionThe non-monophyletic nature of three putative species underscores the necessity of taxonomic revision for Sparganium: S. stoloniferum subsp. choui may be more appropriately identified as S. choui, and subspecies of S. erectum may be in fact distinct species. The estimated diversification times of the two subgenera correspond to their species and nucleotide diversities. The likely ancestral area for most of subgenus Xanthosparganium was East Eurasia and North America from where it dispersed into West Eurasia and Australia. Most of subgenus Sparganium likely originated in North America and then dispersed into Eurasia. Our study demonstrates some of the ways in which complete chloroplast genome sequences can provide new insights into the evolution, phylogeny, and biogeography of the genus Sparganium.

Shoot Phenology in Bambusoideae: A Review

The study of plant phenology is important nowadays since global climate-changing phenomena are impacting the growing patterns and growing periods of plants. Bamboo is of great importance to the agriculture and forestry of temperate, subtropical to tropical regions, especially of Asia. Although some temperate genera can thrive under different climatic conditions, from the Korean Peninsula to South China, it is not known how bamboo will be affected by climate change, so the collection of data related to bamboo phenology could be of interest to research related to climate change. In this review, we describe available data on the phenology of 8 temperate genera, including 79 species, varieties and forms and 4 subtropical–tropical bamboo genera, including 19 species. Primarily, culm shoot physiology is discussed with some reference to leaf phenology data, where available, as well as their interaction. Since the data available in Western literature is often limited to the definition of season rather than exact dates and periods of given months, there is still a great need to explore more about the exact phenology of individual bamboo species to be able to determine the impact of periodic changes in weather patterns or climate change on bamboo phenology in the future.

Raising awareness through the Global Tree Assessment: Extinction risk of six fully assessed tree groups

Societal Impact StatementTrees are essential to life on earth—playing important economic, ecological and cultural roles. However, the world is facing a biodiversity crisis, and to be able to effectively conserve tree species with limited funds, we need to know which tree species are threatened. The Global Tree Assessment has addressed this gap in knowledge through accelerating the production of International Union for Conservation of Nature (IUCN) Red List assessments. Analysing the results of completed taxonomic groups allows for targeted conservation. Here, we explore the extinction risk of six understudied genera with ethnobotanical uses, allowing for conservation priorities to be set.Summary To highlight the ongoing progress of the Global Tree Assessment, a synthesis of the International Union for Conservation of Nature (IUCN) Red List data has been produced for six genera (Gilbertiodendron, Humiriastrum, Liquidambar, Nectandra, Teijsmanniodendron and Tilia) which have all their tree species assessed. This was used to analyse the extinction risk within these groups with the aim of setting conservation priorities and raising awareness of genera not previously publicised in the Global Tree Assessment. The genera were chosen based on completeness, in terms of number of tree species with published IUCN Red List assessments. IUCN Red List categories, threats and uses from each species' IUCN Red List assessment were analysed. An ex situ survey was also carried out to identify species found in botanic gardens, arboreta and seed banks. Higher threat levels were found in the predominately tropical genera (Gilbertiodendron, Humiriastrum, Nectandra and Teijsmanniodendron) compared with the broadly more temperate genera (Tilia and Liquidambar). Habitat loss due to agriculture is a major threat in all genera with exploitation also a key threat to Gilbertiodendron and Tilia. Gilbertiodendron, Humiriastrum, Nectandra and Teijsmanniodendron have no threatened species conserved in ex situ collections. We have identified a need for a range of conservation measures, including collecting threatened species of these groups for ex situ conservation, to ensure no species becomes extinct.

Phylogeny and biogeography of the northern temperate genus Dracocephalum s.l. (Lamiaceae).

The northern temperate genus Dracocephalum consists of approximately 70 species mainly distributed in the steppe-desert biomes of Central and West Asia and the alpine region of the Qinghai-Tibetan Plateau (QTP). Previous work has shown that Dracocephalum is not monophyletic and might include Hyssopus and Lallemantia. This study attempts to clarify the phylogenetic relationships, diversification patterns, and the biogeographical history of the three genera (defined as Dracocephalum s.l.). Based on a sampling of 66 taxa comprising more than 80% from extant species of Dracocephalum s.l., morphological, phylogenetic (maximum parsimony, likelihood, and Bayesian inference based on nuclear ITS and ETS, plastid rpl32-trnL, trnL-trnF, ycf1, and ycf1-rps15, and two low-copy nuclear markers AT3G09060 and AT1G09680), molecular dating, diversification, and ancestral range estimation analyses were carried out. Our results demonstrate that both Hyssopus and Lallemantia are embedded within Dracocephalum and nine well-supported clades can be recognized within Dracocephalum s.l. Analyses of divergence times suggest that the genus experienced an early rapid radiation during the middle to late Miocene with major lineages diversifying within a relatively narrow timescale. Ancestral area reconstruction analyses indicate that Dracocephalum s.l. originated in Central and West Asia and southern Siberia, and dispersed from Central and West Asia into the QTP and adjacent areas twice independently during the Pliocene. The aridification of the Asian interior possibly promoted the rapid radiation of Dracocephalum within this region, and the uplift of the QTP appears to have triggered the dispersal and recent rapid diversification of the genus in the QTP and adjacent regions. Combining molecular phylogenetic and morphological evidence, a revised infrageneric classification of Dracocephalum s.l. is proposed, which recognizes nine sections within the genus.

Phylogenetic relationships and sectional delineation within Gentiana (Gentianaceae)

AbstractGentiana is a sub‐cosmopolitan temperate genus among the most species‐rich in Gentianaceae. Although molecular data (produced via Sanger sequencing) allowed the resolution of phylogenetic relationships between Gentiana and other genera in subtribe Gentianinae, the validity of sections within the genus remains largely untested. In this study, we evaluated the monophyly of all 14 sections attributed to Gentiana, using 294 unlinked anchored loci, the nuclear ribosomal DNA (rDNA) cistron as well as plastid genomes, all produced by anchored hybrid enrichment. We reconstructed phylogenetic relationships by conducting maximum likelihood and Bayesian analyses. These analyses represent a significant improvement over previous taxonomic studies using molecular tools. Our results partly correspond to traditional taxonomic treatments, with several sections being well supported as monophyletic, including Gentiana sect. Calathianae, sect. Ciminalis, sect. Cruciata, sect. Frigida, sect. Gentiana and sect. Pneumonanthe. In contrast, G. sect. Isomeria, sect. Microsperma and sect. Monopodiae were found to be polyphyletic, whereas sect. Dolichocarpa and sect. Fimbricorona were nested within sect. Chondrophyllae. We here provide new taxonomic treatments for these sections, mostly based upon the traditional delineation of their series, which were recovered as monophyletic. In our new treatment, Gentiana encompasses 13 sections. A new determination key to the sections of Gentiana is provided.

Jumping plant lice of the genus Aphalara (Hemiptera, Psylloidea, Aphalaridae) in the Neotropics.

The Neotropical species of the predominantly north temperate genus Aphalara are reviewed. Four species are recorded here from this region, two of which are described as new. Aphalara ritterisp. nov. occurs in southern Brazil (Paraná, Rio Grande do Sul, Santa Catarina) and represents the first and only species reported from South America. A second new species, Aphalara ortegaesp. nov., is described from Mexico and Puerto Rico. Another two species, Aphalara persicaria Caldwell, 1937 and A. similaCaldwell, 1937, have been previously reported from Mexico and the USA, and one of them also from Cuba. The two new species are both associated with Persicaria hydropiperoides and P. punctata (Polygonaceae) on which the immatures induce leaf roll galls. The two new species are morphologically similar to A. persicaria, to which they are probably closely related. A key is provided for the adults and immatures of the Neotropical species of Aphalara.

Determinants of spatial patterns of plant diversity depend on the biogeographical affinities of the taxa − a case study in Northwest Yunnan, Southwest China

The influence of the biogeographical affinities of taxa on the spatial determinants of plant diversity remains largely unknown. I explored the spatial patterns of tropical and temperate plant diversity in Northwest Yunnan and their primary determinants. Generalized linear models (GLMs) and variance partitioning were used to investigate the influences of predictors. Temperate genus diversity significantly increased with latitude, whereas tropical genus diversity did not show any obvious latitudinal trends. Proportions of tropical genera decreased with latitude, whereas those of temperate genera were on the contrary. Environment heterogeneity explained higher percentage of the spatial variation of temperate genus diversity than for that of tropical genus diversity. It may imply that the influence of environment heterogeneity on plant diversity may depend on the taxa's biogeographic affinities, which might modify the exploitation of environment heterogeneity, though the flora diversification induced by the uplifting of the Tibet Plateau may also play important roles. Climate predictors explained higher percentages of variation in the proportions of tropical genera than for the variations in tropical genus diversity. It may suggest that compared with the associations between genus diversity with tropical affinities and climatic conditions, the associations between the proportions of tropical plants and climatic conditions might more accurately reflect niche conservatism of tropical plants in the study area.

The current situation and countermeasures for conservation of ancient and famous trees in Laoshan, Qingdao, China

ABSTRACT: This study aimed to examined the characteristics of ancient and famous trees in Laoshan District, Qingdao City, Shandong Province, China, including species composition, number of individuals, origin, distribution, and age structure, to highlight the values of inheriting history and culture, improving urban environment, protecting regional biodiversity, promoting tourism economy and so on. The analyses was made through field surveys and quantitative analyses of statistical data and relevant literature, The main issues in their conservation are investigated and priority conservation measures are proposed. Results showed that there are 290 ancient and famous trees, comprising 42 species, 34 genera, and 26 families. These included four types of ancient and famous trees with different origins, namely religious trees planted by Buddhists, naturally preserved wild trees, trees with agricultural backgrounds for providing food or used as offerings, and exotic trees introduced from other places. There are relatively more local species and comparatively more elder trees. Ancient and famous trees are distributed in large numbers in Mountain Lao Scenic Area and many of them fall into temperate genera. Currently, ancient and famous tree conservation in Laoshan District is challenging because of issues such as habitat deterioration, severe effects of natural hazards, pests and diseases, weakening physiological function, and inadequate management. Based on all these analyses, countermeasures are proposed, which include regularly inspecting ancient and famous trees, restoring their habitats, reinforcing pest and disease controls, promoting studies on conservation technologies, improving management practices, increasing grants, and reforming the ownership system of ancient and famous trees.

Climate change and the latitudinal selectivity of ancient marine extinctions

AbstractGeologically rapid climate change is anticipated to increase extinction risk nonuniformly across the Earth's surface. Tropical species may be more vulnerable than temperate species to current climate warming because of high tropical climate velocities and reduced seawater oxygen levels. To test whether rapid warming indeed preferentially increased the extinction risk of tropical fossil taxa, we combine a robust statistical assessment of latitudinal extinction selectivity (LES) with the dominant views on climate change occurring at ancient extinction crises. Using a global data set of marine fossil occurrences, we assess extinction rates for tropical and temperate genera, applying log ratios to assess effect size and Akaike weights for model support. Among the classical “big five” mass extinction episodes, the end-Permian mass extinction exhibits temperate preference of extinctions, whereas the Late Devonian and end-Triassic selectively hit tropical genera. Simple links between the inferred direction of climate change and LES are idiosyncratic, both during crisis and background intervals. More complex models, including sampling patterns and changes in the latitudinal distribution of continental shelf area, show tropical LES to be generally associated with raised tropical heat and temperate LES with global cold temperatures. With implications for the future, our paper demonstrates the consistency of high tropical temperatures, habitat loss, and the capacity of both to interact in generating geographic patterns in extinctions.

Using phylogenomics to reconstruct phylogenetic relationships within tribe Polygonateae (Asparagaceae), with a special focus on Polygonatum

Polygonatum is a widespread temperate genus with approximately 75 species centered in the Eastern Himalaya and Indo-Burma biodiversity hotspots. A complete assessment of the remarkable diversity of Polygonatum in these areas requires an accurate circumscription of the genus, as well as a clear understanding of generic and infrageneric relationships, both of which have been problematic in the past. In this study, we reconstruct phylogenetic relationships within Polygonatum and test its monophyly using a phylogenomic approach. For that, we built a comprehensive dataset that includes complete or nearly-complete plastid genomes of 19 species of Polygonatum, one of Disporopsis, and four of Heteropolygonatum. Their plastid genomes do not present any major structural differences and range from 153,821 to 155,580 bp in length. Molecular phylogenetic analyses of the chloroplast coding regions indicate that Polygonatum and Heteropolygonatum are monophyletic, providing support for their recognition as distinct genera and corroborating recent adjustments of their circumscriptions. An expanded analysis with higher species sampling using the petA-psbJ plastid gene region combined with the nuclear ribosomal ITS provided support for the recognition of three distinct sections within Polygonatum. These same sections are further supported by chromosome data: Polygonatum sect. Sibirica (x = 12); Polygonatum sect. Polygonatum (x = 9–11); and, Polygonatum sect. Verticillata (x = 13–15). Populations of P. multiflorum from northwestern Himalaya are here shown to be best treated as a separate taxon, P. govanianum. Furthermore, P. verticillatum is shown to be polyphyletic, indicating that it represents a species-complex that includes multiple Asiatic species. Despite that, additional studies are still needed until the proper nomenclatural adjustments can be made.

Structural characteristics and niches of dominant tree populations in Tetracentron sinense communities: implications for conservation

Background: Tetracentron sinense Oliv., is an endangered, broad-leaved tree distributed in south-central China. Structural characteristics of the community and dominant tree population niches in T. sinense communities are currently unclear.Questions: (1) What is the status of T. sinense populations in the community? (2) What are the interspecies interactions between T. sinense populations and its associated species. Study sites and dates: T. sinense communities in Meigu Dafengding Nature Reserve in southwest China in 2014.Methods: Six sites of 400 m2 were studied for importance value, special diversity, niche breadth, and overlap of dominant tree populations. Results: There were 80 species, 67 genera, and 41 families of vascular plants. Sixty-one genera of seed plants in T. sinense communities were classified into 9 types. The North Temperate genus was the largest, accounting for 44.26 %. Species richness, species diversity, and evenness indexes were all low. There were 6 dominant tree populations in T. sinense communities, and the importance values and niche breadths of T. sinense populations were significantly higher than other species. Horizontal niche overlaps between T. sinense and Pterocarya stenoptera, Acer pictum subsp. mono , and Cercidiphyllum japonicum were high; the vertical niche overlaps between T. sinense and these species were all greater than 0.7.Conclusions: T. sinense communities have North Temperate character, and are immature and unstable communities, still in the early stage of succession. T. sinense is the edificator tree in the communities. A high similarity of habitat demand and strong interspecific competition for light exist between T. sinense and its associated trees.

Clarified relationship between Dactylorhiza viridis and Dactylorhiza iberica renders obsolete the former genus Coeloglossum (Orchidaceae: Orchidinae)

SummaryTwo decades have passed since DNA evidence first demonstrated an intimate relationship between the circumboreal species 'Coeloglossum' viride and the temperate Eurasian genus Dactylorhiza s.s. Most subsequent molecular phylogenies showed 'C.' viride to diverge after the D. incarnata group. The law of monophyly therefore dictated inclusion in Dactylorhiza of 'C.' viride, irrespective of its undeniably distinctive morphology. Those orchid enthusiasts still determinedly seeking reasons for retaining the genus Coeloglossum have often used as a justification the one published molecular study that suggested that D. viridis diverged earlier than D. incarnata. Interestingly, these respective phylogenetic positions are supported by recent data-rich studies based on next-generation sequencing. However, recent DNA phylogenies also show that D. iberica — long regarded as morphologically distinct but nonetheless universally accepted as belonging within the genus — diverged penecontemporaneously with D. viridis. Thus, in order to justify maintaining 'Coeloglossum' as a separate monotypic genus it would also be necessary to transfer D. iberica to a new monotypic genus, thus recognising two genera that are not only monotypic but also show only modest molecular divergence from the remaining dactylorchids. Examining in greater detail the morphology and micromorphology of D. viridis and D. iberica, we show that both species possess multiple morphological character states that are unique within the genus Dactylorhiza, but argue that greater phenotypic disparity is commonly the case in early-divergent lineages per se. Review of previous publications discussing D. iberica revealed little knowledge of its autecology, and contradictory DNA-based inferences that can be traced back to just two original specimens. We also suggest that morphological and molecular variation within both species has been under-estimated and under-explored.

Why are there so many sedges? Sumatroscirpeae, a missing piece in the evolutionary puzzle of the giant genus Carex (Cyperaceae)

For over a century, the origins and mechanisms underlying the diversification of the enormous temperate genus Carex (>2100 species; Cariceae, Cyperaceae) have remained largely speculative. Characteristics such as its diverse ecology, varied biogeography, and intriguing cytology have made Carex a powerful model for studying plant evolution, but its uncertain sister-group relationships hinder its use in studies that depend on accurate ancestral state estimates and biogeographic inferences. To identify the sister to Carex, we estimated the phylogeny of all genera in the Cariceae-Dulichieae-Scirpeae clade (CDS) using three plastid and two nuclear ribosomal markers. Ancestral state reconstructions of key characters were made, and a time-calibrated tree estimated. Carex is strongly supported as sister to the rare East Asian Sumatroscirpus, sole genus of a new tribe, Sumatroscirpeae trib. nov. Believed to be unique to Carex, the perigynium (prophyllar bract enclosing a flower) is in fact a synapomorphy shared with this small tribe (∼4 species) that appeared 36 Mya. We thus suggest the initial key innovation in the remarkable diversification of Carex is not the perigynium, but could be the release of mechanical constraints on perigynia through spikelet truncation, resulting in novel adaptive morphologies. Monoecy, chromosomal change, and rapid inflorescence development enabling phenological isolation may also be involved. The tiny tribe Sumatroscirpeae will provide unprecedented insights into the inflorescence homology, evolution, diversification, and biogeographic history of its sister-group Carex, one of the world’s most diverse plant lineages.

A new species of Paris sect. Axiparis (Melanthiaceae) from Yunnan, China

Paris Linnaeus (1573: 367) is a temperate genus of about 27 species of perennial herbs distributed in Eurasia (Li 1998, Ji et al. 2007). Most species are restricted to East Asia, chiefly to China (19 species), with the Yunnnan-Guizhou Plateau in southwest China as the centre of species diversity (Li 1998). Paris is well known in China for its medicinal value. Those species with a thick rhizome have been used as medicinal herbs for more than 2,000 years in China owing to its analgesic, haemostatic, anti-tumor, and anti-inflammatory activities (Long et al. 2003). More than 40 commercial drugs and health products have been developed in China with Paris used as raw material (Li et al. 2015).

Phytogeographical patterns of genera of endemic flora in relation to latitudinal and climatic gradients in China

This paper aims at detecting the relationships between phytogeographical patterns of genera of Chinese endemic seed plants and latitude or climatic factors. The landmass of China was divided into four latitudinal zones, each of c. 8°. Based on a total of 1664 indigenous genera of Chinese endemic seed plants which were grouped into fifteen geographical elements, belonging to three major categories (cosmopolitan, tropical and temperate) and which were absent or present in 28 provinces in China, we analyzed the phytogeographical patterns of genera of Chinese endemic seed plants and detected the relationships between them and main climatic factors. Our results showed that the proportion of tropical genera decreases with the increase in latitude; the proportion of temperate genera increases with the increase in latitude; and the proportion of cosmopolitan genera increased gradually increased gradually with latitude. There are a slow decrease in the proportion of tropical genera and slow increase in the proportion of temperate genera at latitudes 35°–40°. Alternatively, the tropical genera and the temperate genera have the same proportion at latitude c. 25°. These changes and issues about the different genera also appeared in main climate factors. In general, the genera present in a more northerly flora are a subset of the genera present in a more southerly flora. In summary, the large-scale patterns of phytogeography of endemic flora in China are strongly related to latitude, which covary with several climatic variables such as temperature.

Phlegmariurus lehnertii, a new name for Phlegmariurus lancifolius (Lycopodiaceae)

The pantropical lycophyte genus Phlegmariurus Holub (1964: 21) (Lycopodiaceae) comprises approximately 250 species (PPG I, 2016) and despite extensive study, its taxonomy remains poorly understood (Øllgaard 1992, Field &amp; Bostock 2013). In recent years, many species have been transferred to Phlegmariurus from the closely related temperate genus Huperzia Bernhardi (1801: 126) (Øllgaard 2012a,b, Field &amp; Bostock 2013, Arana 2016, Duy et al. 2016, Field et al. 2016) and new species continue to be described from both the New World (Øllgaard 2015, 2016a,b) and Old World tropics (Hsieh et al. 2012, Duy et al. 2016).

A summary of hybrids detected in the genus Hedera (Araliaceae) with the provision of three new names

Hedera L. (Araliaceae) is a northern temperate genus of around twelve, largely allopatric, species. No hybrids have been confirmed from the few areas where species distributions naturally intersect...

Phyllosiphon ari sp. nov. (Watanabea clade, Trebouxiophyceae), a new parasitic species isolated from leaves of Arum italicum (Araceae)&lt;br /&gt;

The trebouxiophycean genus Phyllosiphon contains unique green algae that thrive as endophytic parasites in the leaves of various members of the Araceae. The DNA sequences of the parasitic populations were originally acquired from infected leaves of subtropical Arisarum vulgare. However, several previous studies showed that the members of the Phyllosiphon clade also occur as free-living algae on various subaerial substrates across Europe. Phyllosiphon infection was also observed on the leaves of the temperate genus Arum, but no molecular data was available for these parasites. We recently found these parasites at a single sub-Mediterranean locality. These algae were genetically different from those previously obtained from Arisarum leaves on the basis of their 18S rDNA and rbcL gene sequences. In the present study, we describe this organism as a new species, Phyllosiphon ari. Phylogenetic differentiation of Phyllosiphon taxa, correlated with host specificity to different genera of the Araceae, suggests concerted host-pathogen co-evolution driving species diversification within this peculiar green algal lineage.

Paleoenvironment of the Quilchena flora, British Columbia, during the Early Eocene Climatic Optimum

The Quilchena fossil locality is dated (51.5 ± 0.4 Ma) to the Early Eocene Climatic Optimum, and this locality is reconstructed as the warmest and wettest of the Early Eocene upland sites from the Okanagan Highlands of British Columbia and northern Washington State. Mean annual temperature (MAT) is estimated from leaf margin analysis, using 55 dicot morphotypes, as 16.2 ± 2.1 °C/14.6 ± 4.8 °C. Using bioclimatic analysis of 45 nearest living relatives, a moist mesothermal climate is indicated (MAT 12.7–16.6 °C; cold month mean temperature (CMMT) 3.5–7.9 °C; mean annual precipitation (MAP) 103–157 cm/year. Leaf size analysis estimates MAP at 121 ± 39 cm/year. Estimates from the climate leaf analysis multivariate program corroborate these results, although with a slightly cooler MAT (13.3 ± 2.1 °C). Plants that support an interpretation of warm winters with minimal or no frost include Azolla, Glyptostrobus, Taxodium, Keteleeria, Pseudolarix, Eucommia, Dipteronia, Hovenia, Ternstroemia, and others. These thermophilous elements occur together with temperate genera such as Alnus, Betula, Ulmus, Calocedrus, and Fraxinus. Palynological assemblages at Quilchena are dominated by bisaccate conifers and Cupressaceae. Common angiosperms include Ulmus type, triporates, Pterocarya, and Alnus. Insect fossils at Quilchena that today inhabit tropical and subtropical regions also support warm and equable climate without significant frost, and include obligate palm-feeding beetles (Pachymerina), which indicate CMMT perhaps as high as 8 °C. These are found together with temperate aphids, wasps, giant lacewings, brown lacewings, and a panorpoid scorpionfly, supporting an interpretation of equable climatic conditions during the Early Eocene Climatic Optimum.