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      Inordinate Spinescence: Taxonomic Revision and Microtomography of the Pheidole cervicornis Species Group (Hymenoptera, Formicidae)

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          Abstract

          The ant genus Pheidole—for all of its hyperdiversity and global ubiquity—is remarkably conservative with regard to morphological disparity. A striking exception to this constrained morphology is the spinescent morphotype, which has evolved multiple times across distantly related lineages of Indoaustralian Pheidole. The Pheidole cervicornis group contains perhaps the most extraordinary spinescent forms of all Pheidole. Here we present a taxonomic revision of the P. cervicornis group, and use microtomographic scanning technology to investigate the internal anatomy of the thoracic spines. Our findings suggest the pronotal spines of Pheidole majors, are possibly skeletomuscular adaptations for supporting their disproportionately large heads. The ‘head support hypothesis’ is an alternative to the mechanical defense hypothesis most often used to explain spinescence in ants. The P. cervicornis group is known only from New Guinea and is represented by the following four species, including two described here as new: P. barumtaun Donisthorpe, P. drogon sp. nov., P. cervicornis Emery, and P. viserion sp. nov. The group is most readily identified by the minor worker caste, which has extremely long pronotal spines and strongly bifurcating propodeal spines. The major and minor workers of all species are illustrated with specimen photographs, with the exception of the major worker of P. cervicornis, which is not known.

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          Most cited references 28

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          Exceptional convergence on the macroevolutionary landscape in island lizard radiations.

          G. G. Simpson, one of the chief architects of evolutionary biology's modern synthesis, proposed that diversification occurs on a macroevolutionary adaptive landscape, but landscape models are seldom used to study adaptive divergence in large radiations. We show that for Caribbean Anolis lizards, diversification on similar Simpsonian landscapes leads to striking convergence of entire faunas on four islands. Parallel radiations unfolding at large temporal scales shed light on the process of adaptive diversification, indicating that the adaptive landscape may give rise to predictable evolutionary patterns in nature, that adaptive peaks may be stable over macroevolutionary time, and that available geographic area influences the ability of lineages to discover new adaptive peaks.
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            Convergent evolution within an adaptive radiation of cichlid fishes.

            The recurrent evolution of convergent forms is a widespread phenomenon in adaptive radiations (e.g., [1-9]). For example, similar ecotypes of anoles lizards have evolved on different islands of the Caribbean, benthic-limnetic species pairs of stickleback fish emerged repeatedly in postglacial lakes, equivalent sets of spider ecomorphs have arisen on Hawaiian islands, and a whole set of convergent species pairs of cichlid fishes evolved in East African Lakes Malawi and Tanganyika. In all these cases, convergent phenotypes originated in geographic isolation from each other. Recent theoretical models, however, predict that convergence should be common within species-rich communities, such as species assemblages resulting from adaptive radiations. Here, we present the most extensive quantitative analysis to date of an adaptive radiation of cichlid fishes, discovering multiple instances of convergence in body and trophic morphology. Moreover, we show that convergent morphologies are associated with adaptations to specific habitats and resources and that Lake Tanganyika's cichlid communities are characterized by the sympatric occurrence of convergent forms. This prevalent coexistence of distantly related yet ecomorphologically similar species offers an explanation for the greatly elevated species numbers in cichlid species flocks. Copyright © 2012 Elsevier Ltd. All rights reserved.
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              Micro-computed tomography: Introducing new dimensions to taxonomy

              Abstract Continuous improvements in the resolution of three-dimensional imaging have led to an increased application of these techniques in conventional taxonomic research in recent years. Coupled with an ever increasing research effort in cybertaxonomy, three-dimensional imaging could give a boost to the development of virtual specimen collections, allowing rapid and simultaneous access to accurate virtual representations of type material. This paper explores the potential of micro-computed tomography (X-ray micro-tomography), a non-destructive three-dimensional imaging technique based on mapping X-ray attenuation in the scanned object, for supporting research in systematics and taxonomy. The subsequent use of these data as virtual type material, so-called “cybertypes”, and the creation of virtual collections lie at the core of this potential. Sample preparation, image acquisition, data processing and presentation of results are demonstrated using polychaetes (bristle worms), a representative taxon of macro-invertebrates, as a study object. Effects of the technique on the morphological, anatomical and molecular identity of the specimens are investigated. The paper evaluates the results and discusses the potential and the limitations of the technique for creating cybertypes. It also discusses the challenges that the community might face to establish virtual collections. Potential future applications of three-dimensional information in taxonomic research are outlined, including an outlook to new ways of producing, disseminating and publishing taxonomic information.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                27 July 2016
                2016
                : 11
                : 7
                Affiliations
                [1 ]Okinawa Institute of Science & Technology Graduate University, 1919–1 Tancha, Onna-son, Okinawa, 904–0495, Japan
                [2 ]Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, Michigan, United States of America
                University of Innsbruck, AUSTRIA
                Author notes

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

                Conceived and designed the experiments: EMS GF EPE. Performed the experiments: EMS GF. Analyzed the data: EMS GF EPE. Wrote the paper: EMS GF EPE.

                Article
                PONE-D-16-11940
                10.1371/journal.pone.0156709
                4963106
                27463644
                © 2016 Sarnat et al

                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.

                Page count
                Figures: 14, Tables: 2, Pages: 24
                Product
                Funding
                Funded by: funder-id http://dx.doi.org/10.13039/100000001, National Science Foundation;
                Award ID: DEB-1145989
                Award Recipient :
                This work was supported by subsidy funding to OIST and NSF (DEB-1145989).
                Categories
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                People and places
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                Oceania
                Papua New Guinea
                Biology and Life Sciences
                Anatomy
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                Custom metadata
                All relevant data are within the paper and its Supporting Information files. The full volumetric datasets have been archived at the Dryad Data Repository ( http://datadryad.org, doi: 10.5061/dryad.1j41m).

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