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      Towards a comprehensive taxonomic revision of the Neotropical dung beetle subgenus Deltochilum ( Deltohyboma) Lane, 1946 (Coleoptera: Scarabaeidae: Scarabaeinae): Division into species-groups

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          Abstract

          Deltochilum Eschscholtz, 1822 is perhaps the most speciose genus of the tribe Deltochilini sensu Tarasov & Dimitrov (2016) (Scarabaeidae: Scarabaeinae) and has been traditionally divided into eight subgenera. Among them, the subgenus Deltohyboma Lane, 1946, is the most speciose with 47 species, five of which are described here ( D. genieri sp. nov., D. gilli sp. nov., D. susanae sp. nov., D. bolivariensis sp. nov. and D. inesae sp. nov.), and at least 165 species still undescribed. Due to the large number of species, and for practical purposes, the subgenus is here divided into 19 species-groups, with D. inesae sp. nov. left as incertae sedis. This division into species-groups will help in the reliable identification of species and will aid in the completion of the revision of subgenus. This report is the first part of the taxonomic revision of the subgenus Deltohyboma; it is based on the examination of all type specimens and almost 9,800 specimens of which approximately 1,200, mostly males, had their genitalia studied. The 19 species-groups recognized here are based mainly on characters described for the first time for Deltohyboma, namely, the state of a) the anterior margin of the clypeus (between the clypeal teeth), b) the internal margin of hypomera, c) the ventral face of the protibia, d) the posterior margin of the metafemur, and e) several new characters resulting from the first detailed study of the aedeagus and the endophallus of the group. Diagnosis, description, geographic distribution, composition and identifications keys (for males as well as males and females) for the species-groups are provided.

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          Biogeographical regionalisation of the Neotropical region.

          A biogeographic regionalisation of the Neotropical region is proposed as a hierarchical classification of sub-regions, dominions, provinces and districts. This regionalisation is based on biogeographic analyses of terrestrial plant and animal taxa, and seeks to provide universality, objectivity and stability, such that it can be applied when describing distributional areas of particular taxa or comparing different biogeographic analyses. The Neotropical region is currently comprised of three sub-regions (Antillean, Brazilian and Chacoan), two transition zones (Mexican and South American), seven dominions (Mesoamerican, Pacific, Boreal Brazilian, Southwestern Amazonian, Southeastern Amazonian, Chacoan and Parana) and 53 provinces. For some of the latter, sub-provinces and districts are recognized. Complete synonymies and brief descriptions of the areas are provided, as well as the endemic taxa that diagnose the different provinces.
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            Scarabaeine Dung Beetles (coleoptera: Scarabaeidae: Scarabaeinae): An Invertebrate Focal Taxon for Biodiversity Research and Conservation

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              New data from the Middle Jurassic of China shed light on the phylogeny and origin of the proboscis in the Mesopsychidae (Insecta: Mecoptera)

              Background The Mesopsychidae is an extinct family of Mecoptera, comprising eleven described genera from Upper Permian to Lower Cretaceous deposits. In 2009, several well-preserved mesopsychids with long proboscides were reported from the mid Mesozoic of Northeastern China, suggesting the presence of pollination mutualisms with gymnosperm plants and highlighting their elevated genus-level diversity. Since that time, additional mesopsychid taxa have been described. However, the phylogeny of genera within Mesopsychidae has not been studied formally, attributable to the limited number of well-preserved fossils. Results Here, we describe two new species, Lichnomesopsyche prochorista sp. nov. and Vitimopsyche pristina sp. nov. and revise the diagnosis of Lichnomesopsyche daohugouensis Ren, Labandeira and Shih, 2010, based on ten specimens from the latest Middle Jurassic Jiulongshan Formation of Inner Mongolia, China. After compiling data from these new fossil species and previously reported representative taxa, we conducted phylogenetic analyses and geometric morphometric studies that now shed light on the taxonomy and phylogeny of Mesopsychidae. We also evaluate the recurring origin of the siphonate proboscis in the Mecoptera and propose an evolutionary developmental model for its multiple origins. Conclusions Phylogenetic and geometric morphometric results confirm the establishment of two new species, each to Lichnomesopsyche and Vitimopsyche. Vitimopsyche pristina sp. nov. extends the existence of the genus Vitimopsyche Novokshonov and Sukacheva, 2001, from the mid Lower Cretaceous to the latest Middle Jurassic. Two methods of analyses indicate an affiliation of Mesopsyche dobrokhotovae Novokshonov, 1997 with Permopsyche Bashkuev, 2011. A phylogenetic analysis of the Mesopsychidae supports: 1), Mesopsychidae as a monophyletic group; 2), Mesopsyche as a paraphyletic group, to be revised pending future examination of additional material; and 3), the independent origin of the proboscis in the Pseudopolycentropodidae, its subsequent loss in earliest Mesopsychidae such as Epicharmesopsyche, its re-origination in the common ancestor (or perhaps independently) in the Vitimopsyche and Lichnomesopsyche clades of the Mesopsychidae. The third conclusion indicates that the proboscis originated four or five times within early Mecoptera, whose origin is explained by an evolutionary developmental model. Electronic supplementary material The online version of this article (doi:10.1186/s12862-015-0575-y) contains supplementary material, which is available to authorized users.
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                Author and article information

                Contributors
                Role: ConceptualizationRole: InvestigationRole: MethodologyRole: VisualizationRole: Writing – original draft
                Role: ConceptualizationRole: Funding acquisitionRole: ResourcesRole: SupervisionRole: Validation
                Role: Editor
                Journal
                PLoS One
                PLoS One
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                6 January 2021
                2021
                : 16
                : 1
                : e0244657
                Affiliations
                [1 ] Programa de Pós-graduação em Ecologia e Conservação da Biodiversidade, Universidade Federal de Mato Grosso, Cuiabá, Mato Grosso, Brazil
                [2 ] Departamento de Biologia e Zoologia, Instituto de Biociências, Universidade Federal de Mato Grosso, Cuiabá, Mato Grosso, Brazil
                Nanjing Agricultural University, CHINA
                Author notes

                Competing Interests: The authors have declared that no competing interests exist

                Author information
                https://orcid.org/0000-0002-0343-3704
                Article
                PONE-D-20-30471
                10.1371/journal.pone.0244657
                7787713
                33406525
                7e1553ef-af9f-45d9-b637-d591429b8610
                © 2021 González-Alvarado, Vaz-de-Mello

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 27 September 2020
                : 12 December 2020
                Page count
                Figures: 33, Tables: 16, Pages: 79
                Funding
                Funded by: funder-id http://dx.doi.org/10.13039/100004457, The World Academy of Sciences;
                Award ID: Process No.190752/2015-7
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/501100003593, Conselho Nacional de Desenvolvimento Científico e Tecnológico;
                Award ID: Process No.190752/2015-7
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/501100003593, Conselho Nacional de Desenvolvimento Científico e Tecnológico;
                Award ID: CNPq 306745/2016-0, 302997/2013-0, 484035/2013-4, 440604/2015-0, 248299/2012-3
                Award Recipient :
                Funded by: Synthesys
                Award ID: BE-TAF-3985
                Award Recipient :
                Funded by: ISYEB
                Award Recipient :
                Funded by: UMR
                Award ID: 7205
                Award Recipient :
                Funded by: CNRS/MNHN
                Award Recipient :
                Funded by: Brazilian Biodiversity Research Programme (PPBio)
                Award Recipient :
                Funded by: Pró-reitora de Pesquisa (PROPeq) and Pró-Reitoria de Ensino de Pós-Graduação (PROPG)
                Award Recipient :
                Funded by: Pró-reitora de Pesquisa (PROPeq) and Pró-Reitoria de Ensino de Pós-Graduação (PROPG)
                Award Recipient :
                AG was supported by a PhD fellowship from CNPq-TWAS (Conselho Nacional de Desenvolvimento Científico e Tecnológico-The World Academy of Sciences) [Process No.190752/ 2015-7] and the Canadian Museum of Nature Visiting Scientist Award. F.Z.V. was supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq 306745/2016-0, 302997/2013-0, 484035/2013-4, 440604/2015-0, 248299/2012-3, 431760/2018-7), Synthesys BE-TAF-3985, ISYEB, UMR 7205 CNRS/MNHN, FAPEMAT PRONEM 2014 and the Brazilian Biodiversity Research Programme (PPBio). AG and F.Z.V. received financial support from Pró-reitora de Pesquisa (PROPeq) and Pró-Reitoria de Ensino de Pós-Graduação (PROPG), Universidade Federal de Mato Grosso (UFMT). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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