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  1. Abstract

    Recently discovered amber-preserved fossil Cicadellidae exhibit combinations of morphological traits not observed in the modern fauna and have the potential to shed new light on the evolution of this highly diverse family. To place the fossils explicitly within a phylogenetic context, representatives of five extinct genera from Cretaceous Myanmar amber, and one from Eocene Baltic amber were incorporated into a matrix comprising 229 discrete morphological characters and representatives of all modern subfamilies. Phylogenetic analyses yielded well resolved and largely congruent estimates that support the monophyly of most previously recognized cicadellid subfamilies and indicate that the treehoppers are derived from a lineage of Cicadellidae. Instability in the morphology-based phylogenies is mainly confined to deep internal splits that received low branch support in one or more analyses and also were not consistently resolved by recent phylogenomic analyses. Placement of fossil taxa is mostly stable across analyses. Three new Cretaceous leafhopper genera, Burmotettix gen. nov., Kachinella gen nov., and Viraktamathus gen. nov., consistently form a monophyletic group distinct from extant leafhopper subfamilies and are placed in Burmotettiginae subfam. nov. Extinct Cretaceous fossils previously placed in Ledrinae and Signoretiinae are recovered as sister to modern representatives of these groups. Eomegophthalmus Dietrich and Gonçalves from Baltic amber consistently groups with a lineage comprising treehoppers, Megophthalminae, Ulopinae, and Eurymelinae but its position is unstable. Overall, the morphology-based phylogenetic estimates agree with recent phylogenies based on molecular data alone suggesting that morphological traits recently used to diagnose subfamilies are generally informative of phylogenetic relationships within this group.

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  2. Abstract Aims

    Mexico harbours a diverse fauna comprising more than 1,400 leafhopper species, 60% of which appear to be strictly endemic, with many apparently restricted to particular habitats and host plants. The aims of this study were to identify areas of high species richness and endemism, and determine the biogeographic affinities of species in the diverse arboreal tribe Athysanini to provide data useful for conservation prioritization.




    A data set of 643 historical occurrence records based on authoritatively identified specimens from museums, recent fieldwork, literature and digital repositories was analysed. Analyses of species richness and areas of endemism were conducted using geographic information systems.


    A total of 164 species belonging of 50 genera were included, of which 145 species of 31 genera are considered to be endemic. The Mexican state of Guerrero yielded the most species records (48%). The highest numbers of taxa and endemic species were concentrated along the Mexican Transition Zone (MTZ) within which the Trans‐Mexican Volcanic Belt (TVB) province had the most species records. Endemicity analyses showed two different geographical patterns but similar species richness weights with overlapping values over the MTZ. Distribution across vegetation types was not uniform, with most athysanine species concentrated in the dry tropical forest (65%). Species were documented at elevations between sea level and 3,200 m a.s.l. with three altitudinal preference classes. Conservation assessments applying IUCN criteria categorized a majority of species (145) as endangered or critically endangered.

    Main conclusions

    Our findings suggest that most identified areas of high species richness throughout the territory have predominantly endemic taxa. Distributional patterns found are non‐random, influenced by richness and endemism values over the TVB province and in the MTZ with a variable dispersion among species. Data highlight a greatly threatened status by habitat loss, remarking an urgent need for an improved conservation framework.

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  3. Abstract

    The suborder Auchenorrhyncha (“true hoppers”) comprises nearly half of known Hemiptera, with >43,000 known species of sap‐sucking herbivores distributed worldwide, including many important agricultural pests and vectors of plant disease. More than half of the known Auchenorrhyncha belong to superfamily Membracoidea (leaf‐ and treehoppers), which has been a source of phylogenetic contention for many years. To construct an improved backbone phylogeny of this superfamily, we obtained transcriptome data for multiple representatives of all 5 previously established extant families and nearly all subfamilies to test their monophyly and relationships. 138 taxa (132 Membracoidea and 6 outgroups) were sampled with an emphasis on families Cicadellidae and Membracidae, which were paraphyletic as previously defined by most authors, several problematic subfamilies (Aphrodinae, Eurymelinae, Ledrinae, Nicomiinae, Stegaspidinae and Tartessinae). We analysed different combinations of data sets (amino acid, complete nucleotide and degeneracy‐coded nucleotide) using different modelling schemes. The resultant trees based on different analyses are congruent in most nodes. Discordant nodes mainly pertain to relationships among cicadellid subfamilies and tribal relationships within Aphrodinae and Eurymelinae. Analyses of gene‐ and site concordance factors and quartet scores indicate that this instability is largely attributable to an overall lack of informative characters across genes and sites rather than strongly supported conflict among genes. According to the congruent nodes, we make the following revisions: combine Stegaspidinae and Centrotinae into a single subfamily, Centrotinae sensu lato; restore Stenocotini from Tartessinae to its original position in the Ledrinae; and transformHoldgatiellaEvans from Nicomiinae to Melizoderinae. In addition, to solve the paraphyly of both Cicadellidae and Membracidae, a preferred option would be to combine all five previously recognized families into a single family, Membracidae sensu lato; the other option could be to render Cicadellidae monophyletic by excluding Megophthalminae and Ulopinae from Cicadellidae and elevating them to status as separate families.

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  4. Abstract

    Phytoplasmas (Mollicutes,Acholeplasmataceae), vector‐borne obligate bacterial plant parasites, infect nearly 1,000 plant species and unknown numbers of insects, mainly leafhoppers (Hemiptera, Deltocephalinae), which play a key role in transmission and epidemiology. Although the plant–phytoplasma–insect association has been evolving for >300 million years, nearly all known phytoplasmas have been discovered as a result of the damage inflicted by phytoplasma diseases on crops. Few efforts have been made to study phytoplasmas occurring in noneconomically important plants in natural habitats. In this study, a subsample of leafhopper specimens preserved in a large museum biorepository was analyzed to unveil potential new associations. PCR screening for phytoplasmas performed on 227 phloem‐feeding leafhoppers collected worldwide from natural habitats revealed the presence of 6 different previously unknown phytoplasma strains. This indicates that museum collections of herbivorous insects represent a rich and largely untapped resource for discovery of new plant pathogens, that natural areas worldwide harbor a diverse but largely undiscovered diversity of phytoplasmas and potential insect vectors, and that independent epidemiological cycles occur in such habitats, posing a potential threat of disease spillover into agricultural systems. Larger‐scale future investigations will contribute to a better understanding of phytoplasma genetic diversity, insect host range, and insect‐borne phytoplasma transmission and provide an early warning for the emergence of new phytoplasma diseases across global agroecosystems.

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  5. Abstract

    The hemipteran suborder Auchenorrhyncha is a highly diverse, ecologically and agriculturally important group of primarily phytophagous insects which has been a source of phylogenetic contention for many years. Here, we have used transcriptome sequencing to assemble 2139 orthologues from 84 auchenorrhynchan species representing 27 families; this is the largest and most taxonomically comprehensive phylogenetic dataset for this group to date. We used both maximum likelihood and multispecies coalescent analyses to reconstruct the evolutionary history in this group using amino acid, nucleotide, and degeneracy‐coded nucleotide orthologue data. Although many relationships at the superfamily level were consistent between analyses, several differing, highly supported topologies were recovered using different datasets and reconstruction methods, most notably the differential placement of Cercopoidea as sister to either Cicadoidea or Membracoidea. To further interrogate the recovered topologies, we explored the contribution of genes as partitioned by third‐codon‐position guanine‐cytosine (GC) content and heterogeneity. We found consistent support for several relationships, including Cercopoidea + Cicadoidea, most often in genes that would be expected to be enriched for the true species tree if recombination‐based dynamics in GC content have contributed to the observed GC heterogeneity. Our results provide a generally well‐supported framework for future studies of auchenorrhynchan phylogeny and suggest that transcriptome sequencing is likely to be a fruitful source of phylogenetic data for resolving its clades. However, we caution that future work should account for the potential effects of GC content heterogeneity on relationships recovered in this group.

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  6. The grassland leafhopper genus Aconurella is widespread in the Old World. Species of this genus are difficult to identify by traditional morphological characters but the morphology-based species classification in this genus has not previously been tested using molecular data. This study analysed DNA sequence data from two mitochondrial genes (COI, 16S) and one nuclear gene (ITS2) to infer the phylogenetic relationships and status of five previously recognized Aconurella species and compare the performance of different molecular species-delimitation methods using single and multiple loci. The analysis divided the included haplotypes into five well-supported subclades, most corresponding to existing morphology-based species concepts. However, different molecular species delimitation methods (jMOTU, ABGD, bPTP, GMYC and BPP) yielded somewhat different results, suggesting the presence of between 4 and 8 species, sometimes lumping the haplotypes of Aconurella diplachnis and Aconurella sibirica into a single species or recognizing multiple putative species within Aconurella prolixa. Considering the different results yielded by various methods employing single loci, the BPP method, which combines data from multiple loci, may be more reliable for delimiting species of Aconurella. Our results suggest that the morphological characters previously used to identify these species are reliable and adequately reflect boundaries between genetically distinct taxa. 
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  7. Based on study of the holotype of Scaphytopius (Tumeus) rubidus DeLong, 1980, it was determined that this species does not belong in any genus of the subfamily Deltocephalinae described so far. To accommodate this species, we describe and illustrate a new genus in the tribe Bahitini, Anavilhanas gen. nov., and propose the new combination, Anavilhanas rubida (DeLong) comb. nov. 
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  8. Biodiversity informatics workbenches and aggregators that make their data externally accessible via application programming interfaces (APIs) facilitate the development of customized applications that fit the needs of a diverse range of communities. In the past, the technical skills required to host web-facing applications placed constraints on many researchers: they either needed to find technical help, or expand their own skills. These limits are now significantly reduced when free or low-cost web-site hosting is combined with small, well-documented applications that require minimal configuration to setup. We illustrate two applications that take advantage of this approach: an interactive key engine (presently named "distinguish") and TaxonPages, a taxon page service application. Both applications make use of TaxonWorks' API. We discuss the limits, e.g., the user must be online to access the data behind the application, and advantages of this approach, e.g., the application server can be served locally, on the users' own computer, and the underlying data are all accessible in more technical formats. 
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  9. Large systematic revisionary projects incorporating data for hundreds or thousands of taxa require an integrative approach, with a strong biodiversity-informatics core for efficient data management to facilitate research on the group. Our original biodiversity informatics platform, 3i (Internet-accessible Interactive Identification) combined a customized MS Access database backend with ASP-based web interfaces to support revisionary syntheses of several large genera of leafhopers (Hemiptera: Auchenorrhyncha: Cicadellidae). More recently, for our National Science Foundation sponsored project, “GoLife: Collaborative Research: Integrative genealogy, ecology and phenomics of deltocephaline leafhoppers (Hemiptera: Cicadellidae), and their microbial associates”, we selected the new open-source platform TaxonWorks as the cyberinfrastructure. In the scope of the project, the original “3i World Auchenorrhyncha Database” was imported into TaxonWorks. At the present time, TaxonWorks has many tools to automatically import nomenclature, citations, and specimen based collection data. At the time of the initial migration of the 3i database, many of those tools were still under development, and complexity of the data in the database required a custom migration script, which is still probably the most efficient solution for importing datasets with long development history. At the moment, the World Auchenorrhyncha Database comprehensively covers nomenclature of the group and includes data on 70 valid families, 6,816 valid genera, 47,064 valid species as well as synonymy and subsequent combinations (Fig. 1). In addition, many taxon records include the original citation, bibliography, type information, etymology, etc. The bibliography of the group includes 37,579 sources, about 1/3 of which are associated with PDF files. Species have distribution records, either derived from individual specimens or as country and state level asserted distribution, as well as biological associations indicating host plants, predators, and parasitoids. Observation matrices in TaxonWorks are designed to handle morphological data associated with taxa or specimens. The matrices may be used to automatically generate interactive identification keys and taxon descriptions. They can also be downloaded to be imported, for example, into Lucid builder, or to perform phylogenetic analysis using an external application. At the moment there are 36 matrices associated with the project. The observation matrix from GoLife project covers 798 taxa by 210 descriptors (most of which are qualitative multi-state morphological descriptors) (Fig. 2). Illustrations are provided for 9,886 taxa and organized in the specialized image matrix and could be used as a pictorial key for determination of species and taxa of a higher rank. For the phylogenetic analysis, a dataset was constructed for 730 terminal taxa and >160,000 nucleotide positions obtained using anchored hybrid enrichment of genomic DNA for a sample of leafhoppers from the subfamily Deltocephalinae and outgroups. The probe kit targets leafhopper genes, as well as some bacterial genes (endosymbionts and plant pathogens transmitted by leafhoppers). The maximum likelihood analyses of concatenated nucleotide and amino acid sequences as well as coalescent gene tree analysis yielded well-resolved phylogenetic trees (Cao et al. 2022). Raw sequence data have been uploaded to the Sequence Read Archive on GenBank. Occurrence and morphological data, as well as diagnostic images, for voucher specimens have been incorporated into TaxonWorks. Data in TaxonWorks could be exported in raw format, get accessed via Application Programming Interface (API), or be shared with external data aggregators like Catalogue of Life, GBIF, iDigBio. 
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  10. Despite several decades’ effort to detect and identify phytoplasmas (Mollicutes) using PCR and Sanger sequencing focusing on diseased plants, knowledge of phytoplasma biodiversity and vector associations remains highly incomplete. To improve protocols for documenting phytoplasma diversity and ecology, we used DNA extracted from phloem-feeding insects and compared traditional Sanger sequencing with a next-generation sequencing method, Anchored Hybrid Enrichment (AHE) for detecting and characterizing phytoplasmas. Among 22 of 180 leafhopper samples that initially tested positive for phytoplasmas using qPCR, AHE yielded phytoplasma 16Sr sequences for 20 (19 complete and 1 partial sequence) while Sanger sequencing yielded sequences for 16 (11 complete and 5 partial). AHE yielded phytoplasma sequences for an additional 7 samples (3 complete and 4 partial) that did not meet the qPCR threshold for phytoplasma positivity or yielded non-phytoplasma sequences using Sanger sequencing. This suggests that AHE is more efficient for obtaining phytoplasma sequences. Twenty-three samples with sufficient data were classified into eight 16Sr subgroups (16SrI-B, I-F, I-AO, III-U, V-C, IX-J, XI-C, XXXVII-A), three new subgroups (designated as 16SrVI-L, XV-D, XI-G) and three possible new groups. Our results suggest that screening phloem-feeding insects using qPCR and AHE sequencing may be the most efficient method for discovering new phytoplasmas. 
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