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      Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicoidea)

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          Abstract

          Fossils provide primary material evidence for the pattern and timing of evolution. The newly discovered “beast ants” from mid-Cretaceous Burmite, † Camelosphecia gen. nov., display an exceptional combination of plesiomorphies, including absence of the metapleural gland, and a series of unique apomorphies. Females and males, represented by † C. fossor sp. nov. and † C. venator sp. nov., differ in a number of features which suggest distinct sexual biologies. Combined-evidence phylogenetic analysis recovers † Camelosphecia and † Camelomecia as a clade which forms the extinct sister group of the Formicidae . Notably, these genera are only known from alate males and females; workers, if present, have yet to be recovered. Based on ongoing study of the total Aculeata informed by the beast ant genera, we provide a brief diagnosis of the Formicoidea . We also provide the first comprehensive key to the major groupings of Mesozoic Formicoidea , alongside a synoptic classification in which † Zigrasimeciinae stat. nov. and † Myanmyrma maraudera comb. nov. are recognized. Finally, a brief diagnosis of the Formicoidea is outlined.

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          The Ants

          From the Arctic to South Africa - one finds them everywhere: Ants. Making up nearly 15% of the entire terrestrial animal biomass, ants are impressive not only in quantitative terms, they also fascinate by their highly organized and complex social system. Their caste system, the division of labor, the origin of altruistic behavior and the complex forms of chemical communication makes them the most interesting group of social organisms and the main subject for sociobiologists. Not least is their ecological importance: Ants are the premier soil turners, channelers of energy and dominatrices of the insect fauna. TOC:The importance of ants.- Classification and origins.- The colony life cycle.- Altruism and the origin of the worker caste.- Colony odor and kin recognition.- Queen numbers and domination.- Communication.- Caste and division of labor.- Social homeostasis and flexibility.- Foraging and territorial strategies.- The organization of species communities.- Symbioses among ant species.- Symbioses with other animals.- Interaction with plants.- The specialized predators.- The army ants.- The fungus growers.- The harvesters.- The weaver ants.- Collecting and culturing ants.- Glossary.- Bibliography.- Index.
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            Phylogeny of the ants: diversification in the age of angiosperms.

            C. Moreau (2006)
            We present a large-scale molecular phylogeny of the ants (Hymenoptera: Formicidae), based on 4.5 kilobases of sequence data from six gene regions extracted from 139 of the 288 described extant genera, representing 19 of the 20 subfamilies. All but two subfamilies are recovered as monophyletic. Divergence time estimates calibrated by minimum age constraints from 43 fossils indicate that most of the subfamilies representing extant ants arose much earlier than previously proposed but only began to diversify during the Late Cretaceous to Early Eocene. This period also witnessed the rise of angiosperms and most herbivorous insects.
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              Evolutionary History of the Hymenoptera.

              Hymenoptera (sawflies, wasps, ants, and bees) are one of four mega-diverse insect orders, comprising more than 153,000 described and possibly up to one million undescribed extant species [1, 2]. As parasitoids, predators, and pollinators, Hymenoptera play a fundamental role in virtually all terrestrial ecosystems and are of substantial economic importance [1, 3]. To understand the diversification and key evolutionary transitions of Hymenoptera, most notably from phytophagy to parasitoidism and predation (and vice versa) and from solitary to eusocial life, we inferred the phylogeny and divergence times of all major lineages of Hymenoptera by analyzing 3,256 protein-coding genes in 173 insect species. Our analyses suggest that extant Hymenoptera started to diversify around 281 million years ago (mya). The primarily ectophytophagous sawflies are found to be monophyletic. The species-rich lineages of parasitoid wasps constitute a monophyletic group as well. The little-known, species-poor Trigonaloidea are identified as the sister group of the stinging wasps (Aculeata). Finally, we located the evolutionary root of bees within the apoid wasp family "Crabronidae." Our results reveal that the extant sawfly diversity is largely the result of a previously unrecognized major radiation of phytophagous Hymenoptera that did not lead to wood-dwelling and parasitoidism. They also confirm that all primarily parasitoid wasps are descendants of a single endophytic parasitoid ancestor that lived around 247 mya. Our findings provide the basis for a natural classification of Hymenoptera and allow for future comparative analyses of Hymenoptera, including their genomes, morphology, venoms, and parasitoid and eusocial life styles.
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                Author and article information

                Contributors
                Journal
                Zookeys
                Zookeys
                2
                urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E
                urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2020
                18 December 2020
                : 1005
                : 21-55
                Affiliations
                [1 ] Department of Entomology & Nematology, University of California, Davis, One Shields Ave, Davis, CA 95616, USA University of California Davis United States of America
                [2 ] Friedrich-Schiller-Universität Jena, Institut für Zoologie und Evolutionsforschung, 1 Erberstraße, 07743 Jena, Thüringen, Germany Friedrich-Schiller-Universität Jena Germany
                [3 ] Univ. Rennes, CNRS, Géosciences – UMR 6118, F-35000, Rennes, France Univ. Rennes, CNRS Rennes France
                [4 ] Pós-Graduação em Ecologia, Departamento de Biologia Geral, Universidade Federal do Viçosa, 36570-900, Viçosa, MG, Brazil Universidade Federal do Viçosa Viçosa Brazil
                Author notes
                Corresponding author: Brendon E. Boudinot ( boudinotb@ 123456gmail.com )

                Academic editor: M. Borowiec

                Author information
                https://orcid.org/0000-0002-4588-0430
                https://orcid.org/0000-0002-7973-0430
                Article
                57629
                10.3897/zookeys.1005.57629
                7762752
                33390754
                27d15926-8c86-416c-966d-bcf3a5d1bfcc
                Brendon E. Boudinot, Vincent Perrichot, Júlio C. M. Chaul

                This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 14 August 2020
                : 14 November 2020
                Funding
                Funded by: Museum of Comparative Zoology, Harvard University 100010952 http://doi.org/10.13039/100010952
                Categories
                Research Article
                Formicidae
                Identification Key
                Nomenclature
                Palaeontology
                Taxonomy
                Mesozoic
                World

                Animal science & Zoology
                classification,identification key,mesozoic revision,morphology,paleontology,phylogeny,taxonomic synopsis

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