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      From micropterism to hyperpterism: recognition strategy and standardized homology-driven terminology of the forewing venation patterns in planthoppers (Hemiptera: Fulgoromorpha)

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          Abstract

          Following recent advances in the morphological interpretations of the tegmen basal cell margins in the Paraneoptera, a standardized and homology-driven groundplan terminology for tegmina types, structures and vein patterns in Hemiptera Fulgoromorpha, including fossils, is proposed. Each term is listed with a morphological definition, compared and linked to the main systems of planthopper forewing description that have been reviewed. The importance of a standardized and homology-driven terminology is stressed to enhance the quality of data in taxonomic descriptions and to strengthen phylogenetic morphological analysis results. When the interpretation of the origin of vein branches is render difficult, a three-step strategy for pattern recognition of the vein is proposed based on two principles: (1) vein forks are more informative than topology of the vein branches: a search for homologous areas, the nodal cells in particular, must first guide the recognition rather the number of branches of a vein, and (2) minimum of ad hoc evolutionary events should be invoked in the understanding of a modified vein pattern. Examples of some conflicting interpretations of venation patterns in planthoppers are discussed within different families for both extant and extinct taxa. For the first time, the concept of brachypterism is defined in a non-relative way independently from other structures, and the new one of hyperpterism is proposed; a reporting system is proposed for each of them.

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          Most cited references 67

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          Functional Morphology of Insect Wings

           Robin Wootton (1992)
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            Origin of the insect wing and wing articulation from the arthropodan leg

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              The earliest known holometabolous insects.

              The Eumetabola (Endopterygota (also known as Holometabola) plus Paraneoptera) have the highest number of species of any clade, and greatly contribute to animal species biodiversity. The palaeoecological circumstances that favoured their emergence and success remain an intriguing question. Recent molecular phylogenetic analyses have suggested a wide range of dates for the initial appearance of the Holometabola, from the Middle Devonian epoch (391 million years (Myr) ago) to the Late Pennsylvanian epoch (311 Myr ago), and Hemiptera (310 Myr ago). Palaeoenvironments greatly changed over these periods, with global cooling and increasing complexity of green forests. The Pennsylvanian-period crown-eumetabolan fossil record remains notably incomplete, particularly as several fossils have been erroneously considered to be stem Holometabola (Supplementary Information); the earliest definitive beetles are from the start of the Permian period. The emergence of the hymenopterids, sister group to other Holometabola, is dated between 350 and 309 Myr ago, incongruent with their current earliest record (Middle Triassic epoch). Here we describe five fossils--a Gzhelian-age stem coleopterid, a holometabolous larva of uncertain ordinal affinity, a stem hymenopterid, and early Hemiptera and Psocodea, all from the Moscovian age--and reveal a notable penecontemporaneous breadth of early eumetabolan insects. These discoveries are more congruent with current hypotheses of clade divergence. Eumetabola experienced episodes of diversification during the Bashkirian-Moscovian and the Kasimovian-Gzhelian ages. This cladogenetic activity is perhaps related to notable episodes of drying resulting from glaciations, leading to the eventual demise in Euramerica of coal-swamp ecosystems, evidenced by floral turnover during this interval. These ancient species were of very small size, living in the shadow of Palaeozoic-era 'giant' insects. Although these discoveries reveal unexpected Pennsylvanian eumetabolan diversity, the lineage radiated more successfully only after the mass extinctions at the end of the Permian period, giving rise to the familiar crown groups of their respective clades.
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                Author and article information

                Contributors
                bourgoin@mnhn.fr
                wangrr_2008@163.com
                manfred.asche@mfn-berlin.de
                hannelore.hoch@mfn-berlin.de
                soulier@mnhn.fr
                adam@miiz.waw.pl
                sayap@uplb.edu.ph
                jacek.szwedo@biol.ug.edu.pl
                Journal
                Zoomorphology
                Zoomorphology
                Zoomorphology
                Springer Berlin Heidelberg (Berlin/Heidelberg )
                0720-213X
                1432-234X
                21 September 2014
                21 September 2014
                2015
                : 134
                : 1
                : 63-77
                Affiliations
                [ ]Département Systématique et Evolution, UMR 7205-ISyEB, MNHN-CNRS-UPMC-EPHE, Muséum National d’Histoire Naturelle, CP 50, 45 rue Buffon, 75005 Paris, France
                [ ]Institute of Zoology, Chinese Academy of Sciences, 1 Beichen West Road, Chaoyang District, Beijing, 100101 China
                [ ]Museum für Naturkunde Leibniz-Institut für Evolutions und Biodiversitätsforschung an der Humboldt Universität zu Berlin, AG Biosystematik Invalidenstr. 43, 10115 Berlin, Germany
                [ ]Department of Palaeozoology, Museum and Institute of Zoology, Polish Academy of Sciences, 64, Wilcza Street, 00-679 Warsaw, Poland
                [ ]Crop Protection Cluster and Museum of Natural History, University of the Philippines Los Baños, 4030 Los Baños, Laguna Philippines
                [ ]Department of Invertebrate Zoology and Parasitology, University of Gdansk, 59, Wita Stwosza Street, 80-308 Gdańsk, Poland
                Author notes

                Communicated by Andreas Schmidt-Rhaesa.

                Article
                243
                10.1007/s00435-014-0243-6
                4326643
                25705075
                © The Author(s) 2014

                Open AccessThis article is distributed under the terms of the Creative Commons Attribution License which permits any use, distribution, and reproduction in any medium, provided the original author(s) and the source are credited.

                Categories
                Original Paper
                Custom metadata
                © Springer-Verlag Berlin Heidelberg 2015

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