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      Chemical Ecology of Cave-Dwelling Millipedes: Defensive Secretions of the Typhloiulini (Diplopoda, Julida, Julidae)

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          Abstract

          Cave animals live under highly constant ecological conditions and in permanent darkness, and many evolutionary adaptations of cave-dwellers have been triggered by their specific environment. A similar “cave effect” leading to pronounced chemical interactions under such conditions may be assumed, but the chemoecology of troglobionts is mostly unknown. We investigated the defensive chemistry of a largely cave-dwelling julid group, the controversial tribe “Typhloiulini”, and we included some cave-dwelling and some endogean representatives. While chemical defense in juliform diplopods is known to be highly uniform, and mainly based on methyl- and methoxy-substituted benzoquinones, the defensive secretions of typhloiulines contained ethyl-benzoquinones and related compounds. Interestingly, ethyl-benzoquinones were found in some, but not all cave-dwelling typhloiulines, and some non-cave dwellers also contained these compounds. On the other hand, ethyl-benzoquinones were not detected in troglobiont nor in endogean typhloiuline outgroups. In order to explain the taxonomic pattern of ethyl-benzoquinone occurrence, and to unravel whether a cave-effect triggered ethyl-benzoquinone evolution, we classed the “Typhloiulini” investigated here within a phylogenetic framework of julid taxa, and traced the evolutionary history of ethyl-benzoquinones in typhloiulines in relation to cave-dwelling. The results indicated a cave-independent evolution of ethyl-substituted benzoquinones, indicating the absence of a “cave effect” on the secretions of troglobiont Typhloiulini. Ethyl-benzoquinones probably evolved early in an epi- or endogean ancestor of a clade including several, but not all Typhloiulus (basically comprising a taxonomic entity known as “ Typhloiulus sensu stricto”) and Serboiulus. Ethyl-benzoquinones are proposed as novel and valuable chemical characters for julid systematics.

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          The online version of this article (doi:10.1007/s10886-017-0832-1) contains supplementary material, which is available to authorized users.

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          Most cited references 46

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          Selecting optimal partitioning schemes for phylogenomic datasets

          Background Partitioning involves estimating independent models of molecular evolution for different subsets of sites in a sequence alignment, and has been shown to improve phylogenetic inference. Current methods for estimating best-fit partitioning schemes, however, are only computationally feasible with datasets of fewer than 100 loci. This is a problem because datasets with thousands of loci are increasingly common in phylogenetics. Methods We develop two novel methods for estimating best-fit partitioning schemes on large phylogenomic datasets: strict and relaxed hierarchical clustering. These methods use information from the underlying data to cluster together similar subsets of sites in an alignment, and build on clustering approaches that have been proposed elsewhere. Results We compare the performance of our methods to each other, and to existing methods for selecting partitioning schemes. We demonstrate that while strict hierarchical clustering has the best computational efficiency on very large datasets, relaxed hierarchical clustering provides scalable efficiency and returns dramatically better partitioning schemes as assessed by common criteria such as AICc and BIC scores. Conclusions These two methods provide the best current approaches to inferring partitioning schemes for very large datasets. We provide free open-source implementations of the methods in the PartitionFinder software. We hope that the use of these methods will help to improve the inferences made from large phylogenomic datasets.
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            Cave Biology

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              Volatile constituents of fermented sugar baits and their attraction to lepidopteran species.

              The volatile compounds emanating from four fermented sugar baits, palm sugar, golden cane syrup, port wine, and molasses, were isolated by headspace sampling and analyzed by gas chromatography-mass spectrometry. Three classes of compounds including esters, alcohols, and aromatic compounds were identified in the headspace of the four fermented sugar baits. There was a high degree of qualitative similarity between the headspace contents of the four fermented sugar baits, although quantitatively they varied considerably. Ethyl acetate, 3-methylbutanol, ethyl hexanoate, 2-phenylethanol, ethyl octanoate, ethyl (E)-4-decenoate, ethyl decanoate, and ethyl dodecanoate were the major compounds identified in the headspace of the four fermented sugar baits. The efficacy of the four fermented sugar baits was investigated in field trapping experiments. Fermented palm sugar and golden cane syrup were superior in attracting significant numbers of moths as compared to port wine and molasses. Fermented molasses was the least attractive among the four baits. Over 90% of the insects caught were noctuids with Graphania mutans and Tmetolophota spp. being the main noctuids captured (over 55%) in the four fermented sugar baits. Male and female G. mutans were equally attracted to the four sugar baits. A number of tortricid species were also trapped.
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                Author and article information

                Contributors
                guenther.raspotnig@uni-graz.at
                Journal
                J Chem Ecol
                J. Chem. Ecol
                Journal of Chemical Ecology
                Springer US (New York )
                0098-0331
                1573-1561
                16 March 2017
                16 March 2017
                2017
                : 43
                : 4
                : 317-326
                Affiliations
                [1 ]ISNI 0000 0001 2166 9385, GRID grid.7149.b, Institute of Zoology, , University of Belgrade - Faculty of Biology, ; Studentski Trg 16, Belgrade, 11000 Serbia
                [2 ]ISNI 0000000121539003, GRID grid.5110.5, Institute of Zoology, , University of Graz, ; Universitätsplatz 2, 8010 Graz, Austria
                [3 ]ISNI 0000 0001 2166 9385, GRID grid.7149.b, Faculty of Chemistry, , University of Belgrade, ; Studentski trg 12-16, Belgrade, 11000 Serbia
                [4 ]ISNI 0000 0001 2097 3094, GRID grid.410344.6, Institute of Biodiversity and Ecosystem Research, Department of Animal Diversity and Resources, , Bulgarian Academy of Sciences, ; 2 Gagarin Street, 1113 Sofia, Bulgaria
                [5 ]ISNI 0000 0001 2192 3275, GRID grid.11355.33, Department of Zoology and Anthropology, , Sofia University “St. Kliment Ohridsky”, ; Sofia, Bulgaria
                [6 ]ISNI 0000 0001 2166 9385, GRID grid.7149.b, Institute of Chemistry, Technology and Metallurgy, , University of Belgrade, ; Studentski trg 12-16, Belgrade, 11000 Serbia
                [7 ]ISNI 0000 0000 8988 2476, GRID grid.11598.34, Research Unit of Osteology and Analytical Mass Spectrometry, , Medical University, Children’s Hospital, ; Auenbruggerplatz 30, 8036 Graz, Austria
                Article
                832
                10.1007/s10886-017-0832-1
                5399059
                28303527
                © The Author(s) 2017

                Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.

                Funding
                Funded by: FundRef http://dx.doi.org/10.13039/501100001822, Austrian Academy of Sciences;
                Award ID: 23811
                Award Recipient :
                Funded by: Ministry of Education and Science of Serbia
                Award ID: 173038; 172053
                Award Recipient :
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                © Springer Science+Business Media New York 2017

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