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      Revision and Microtomography of the Pheidole knowlesi Group, an Endemic Ant Radiation in Fiji (Hymenoptera, Formicidae, Myrmicinae)

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          Abstract

          The Fijian islands, a remote archipelago in the southwestern Pacific, are home to a number of spectacular endemic radiations of plants and animals. Unlike most Pacific archipelagos, these evolutionary radiations extend to social insects, including ants. One of the most dramatic examples of ant radiation in Fiji has occurred in the hyperdiverse genus Pheidole. Most of the 17 native Fijian Pheidole belong to one of two species groups that descended from a single colonization, yet have evolved dramatically contrasting morphologies: the spinescent P. roosevelti species group, and the more morphologically conservative P. knowlesi species group. Here we revise the knowlesi group, in light of recent phylogenetic results, and enhanced with modern methods of X-ray microtomography. We recognize six species belonging to this group, including two of which we describe as new: Pheidole caldwelli Mann, Pheidole kava sp. n., Pheidole knowlesi Mann, P. ululevu sp. n., P. vatu Mann, and P. wilsoni Mann. Detailed measurements and descriptions, identification keys, and high-resolution images for queens, major and minor workers are provided. In addition, we include highly detailed 3D surface reconstructions for all available castes.

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          Most cited references 16

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          Arthropods on islands: colonization, speciation, and conservation.

          Islands have traditionally been considered to be any relatively small body of land completely surrounded by water. However, their primary biological characteristic, an extended period of isolation from a source of colonists, is common also to many situations on continents. Accordingly, theories and predictions developed for true islands have been applied to a huge array of systems, from rock pools, to single tree species in forests, to oceanic islands. Here, we examine the literature on islands in the broadest sense (i.e., whether surrounded by water or any other uninhabitable matrix) as it pertains to terrestrial arthropods. We categorize islands according to the features they share. The primary distinction between different island systems is "darwinian" islands (formed de novo) and "fragment" islands. In the former, the islands have never been in contact with the source of colonists and have abundant "empty" ecological niche space. On these islands, species numbers will initially increase through immigration, the rate depending on the degree of isolation. If isolation persists, over time species formation will result in "neo-endemics." When isolation is extreme, the ecological space will gradually be filled through speciation (rather than immigration) and adaptive radiation of neo-endemics. Fragment islands are fundamentally different. In these islands, the ecological space will initially be filled as a consequence of connection to the source of colonists prior to insularization. Species numbers will decrease following fragmentation through the process of relaxation. If these islands become more isolated, species will eventually arise through relictualization with the formation of "paleo-endemics." Given sufficient time, this process can result in generic level endemism on ancient fragment islands, a phenomenon well illustrated in Madagascar and New Zealand. Recognizing the distinction between the different kinds of islands is fundamental for understanding emerging patterns on each, in particular speciation, biodiversity (e.g., neo-endemics versus paleo-endemics), and conservation (e.g., naiveté in interactions with alien species).
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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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              Imaging applications of synchrotron X-ray phase-contrast microtomography in biological morphology and biomaterials science. I. General aspects of the technique and its advantages in the analysis of millimetre-sized arthropod structure.

              Synchrotron-generated X-rays provide scientists with a multitude of investigative techniques well suited for the analysis of the composition and structure of all types of materials and specimens. Here, we describe the properties of synchrotron-generated X-rays and the advantages that they provide for qualitative morphological research of millimetre-sized biological organisms and biomaterials. Case studies of the anatomy of insect heads, of whole microarthropods and of the three-dimensional reconstruction of the cuticular tendons of jumping beetles, all performed at the beamline ID19 of the European Synchrotron Radiation Facility (ESRF), are presented to illustrate the techniques of phase-contrast tomography available for anatomical and structural investigations. Various sample preparation techniques are described and compared and experimental settings that we have found to be particularly successful are given. On comparing the strengths and weaknesses of the technique with traditional histological thin sectioning, we conclude that synchrotron radiation microtomography has a great potential in biological microanatomy.
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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
                8 August 2016
                : 11
                : 7
                Affiliations
                [1 ]Okinawa Institute of Science & Technology Graduate University, 1919-1 Tancha, Onna-son, Okinawa, Japan, 904-0495
                [2 ]Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI, United States of America
                CNRS, FRANCE
                Author notes

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

                Conceived and designed the experiments: GF EPE EMS. Performed the experiments: GF EPE EMS. Analyzed the data: GF EPE EMS. Contributed reagents/materials/analysis tools: EPE EMS. Wrote the paper: GF EPE EMS. Performed the microtomogaphy: GF.

                Article
                PONE-D-16-16079
                10.1371/journal.pone.0158544
                4963041
                27462877
                © 2016 Fischer 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: 11, Tables: 3, Pages: 39
                Product
                Funding
                Funded by: funder-id http://dx.doi.org/10.13039/100000001, National Science Foundation;
                Award ID: DEB-1145989
                Award Recipient : Evan Philip Economo
                Funded by: funder-id http://dx.doi.org/10.13039/100000001, National Science Foundation;
                Award ID: DEB-0425970
                Funded by: funder-id http://dx.doi.org/10.13039/100007714, Okinawa Institute of Science and Technology Promotion Corporation;
                Award Recipient : Evan Philip Economo
                This work was supported by DEB-1145989 to EPE, and by DEB-0425970 ‘Fiji Terrestrial Arthropod Survey’, both from National Science Foundation, http://www.nsf.gov/; and by subsidy funding to EPE from Okinawa Institute of Science and Technology, http://www.oist.jp/. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Anatomy
                Digestive System
                Teeth
                Medicine and Health Sciences
                Anatomy
                Digestive System
                Teeth
                Biology and Life Sciences
                Anatomy
                Head
                Jaw
                Teeth
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                Head
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                Teeth
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                Musculoskeletal System
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                Custom metadata
                Specimen images, collection and specimen data are uploaded to and available at www.antweb.org. The original X-ray microtomography scans are uploaded to Dryad ( http://datadryad.org) (DOI: http://dx.doi.org/10.5061/dryad.7d3v4).

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