Abstract
Cladophorus Guérin-Méneville, 1830 are endemic Papuan net-winged beetles which take part in highly diverse Müllerian mimicry rings. Available specimens were sequenced for cox1–tRNA-Leu–cox2 mitochondrial DNA fragment and the species delimitations were based on the genetic distance, phylogenetic analysis, and morphology. Three earlier described species were identified in the recently collected material and further 10 species are described: C. pallescens sp. nov., C. bicolor sp. nov., C. craterensis sp. nov., C. motykai sp. nov., C. mindikensis sp. nov., C. kailakiensis sp. nov., C. manokwarensis sp. nov., C. haiaensis sp. nov., C. humeralis sp. nov., and C. boceki sp. nov. DNA-based identifications provided some ambiguous results and closely related species could not be robustly delimited using solely molecular data. Additionally, the species limits were based on clearly defined morphological characters and the morphological differentiation was found unlinked from the genetic divergence. Colour patterns cannot be used for identification because all species available in more specimens were polymorphic and followed various local co-mimics. The Papuan fauna of Cladophorus is very diverse and the closely related species regularly occur in limited regions. Differentiation within restricted ranges is therefore considered as the main speciation mode.
References
Bocak, L. (1998) Nomenclatural notes on taxa of the family Lycidae described by Guérin-Méneville (Insecta: Coleoptera). Annales Zoologici, 48, 245–251.
Bocak, L. (2002) Generic revision and phylogenetic analysis of Metriorrhynchinae. European Journal of Entomology, 99, 315–351.
https://doi.org/10.14411/eje.2002.043Bocak, L. (2007) A Revision of Metriorrhynchus (Coleoptera: Lycidae) from the Greater Sunda Islands and Continental Asia. Raffles Bulletin of Zoology, 55, 253–260.
Bocak, L. & Bocakova, M. (1990) Revision of the genus Hemiconderis (Coleoptera, Lycidae). Acta Entomologica Bohemoslovaca, 89, 209–220.
Bocak, L., Matsuda, K. & Yagi, T. (2006) A revision of Metriorrhynchus from the Philippines with molecular evidence of an Australian origin of the Oriental Metriorrhynchus fauna (Coleoptera: Lycidae). European Journal of Entomology, 103, 115–126.
https://doi.org/10.14411/eje.2006.017Bocak, L. & Yagi, T. (2010) Evolution of mimicry patterns in Metriorrhynchus (Coleoptera: Lycidae): the history of dispersal and speciation in Southeast Asia. Evolution, 64, 39–52.
https://doi.org/10.1111/j.1558-5646.2009.00812.xBocakova, M. (1992) Revision of the genus Calochromus Guérin-Méneville (Coleoptera, Lycidae) from New Guinea and adjacent islands. European Journal of Entomology, 89, 301–308.
Bocek, M. (2017) New species of Diatrichalus from New Guinea. Zootaxa, 4247 (5), 577–584.
https://doi.org/10.11646/zootaxa.4247.5.4Bocek, M. & Bocak, L. (2016) Where are species limits in polymorphic mimetic beetles from New Guinean mountains: a case of Eniclases net-winged beetles (Lycidae: Metriorrhynchini). ZooKeys, 593, 15–35.
https://doi.org/10.3897/zookeys.593.7728Bray, T.C. & Bocak, L. (2016) Slowly dispersing neotenic beetles can speciate on a penny coin and generate space-limited diversity in the tropical mountains. Scientific Reports, 6, 33579.
https://doi.org/10.1038/srep33579Calder, A.A. (1998) Coleoptera: Elateroidea. In: Wells, A. (Ed.), Zoological Catalogue of Australia. Vol. 296. CSIRO Publishing, Melbourne, pp. i–xiii + 1–248.
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2, more models, new heuristics and parallel computing. Nature Methods, 9 (8), 772.
https://doi.org/10.1038/nmeth.2109Dayrat, B. (2005) Towards integrative taxonomy. Biological Journal of the Linnean Society, 85, 407–415.
https://doi.org/10.1111/j.1095-8312.2005.00503.xEisner, T., Schroeder, F.C., Snyder, N., Grant, J.B., Aneshansley, D.J., Utterback, D., Meinwald, J. & Eisner, M. (2008) Defensive chemistry of lycid beetles and of mimetic cerambycid beetles that feed on them. Chemoecology, 18, 109–119. https://doi.org/10.1007/s00049-007-0398-4
Hall, R. (1998) The plate tectonics of Cenozoic SE Asia and the distribution of land and sea. In: Hall, R. & Holloway, J.D. (Eds.), Biogeography and geological evolution of SE Asia. Backhuys Publishers, Leiden, pp. 99–131.
Hebert, P.D.N., Ratnasingham, S. & deWaard, J.R. (2003) Barcoding animal life: cytochrome c oxidase subunit 1 divergences among closely related species. Proceedings of the Royal Society of London B: Biological Sciences, 270, 96–99. [S96–S99] https://doi.org/10.1098/rsbl.2003.0025
Huson, D.H., Richter, D.C., Rausch, C., Dezulian, T., Franz, M. & Rupp, R. (2007) Dendroscope: An interactive viewer for large phylogenetic trees, BMC Bioinformatics, 8, 460.
https://doi.org/10.1186/1471-2105-8-460Guérin-Méneville, F.C. (1830) Crustacés, arachnides et insectes. In: Duperrey, L.I. (Ed.), Voyage autour du monde ...sur la Corvette de Sa Majesté, La Coquille, pendant les annés 1822, 1823, 1824 et 1825 ... Zoologie. Vol. 2. Part 2. Bertrand, Paris, pl. II, fig. 9.
Guérin-Méneville, F.C. (1838) Crustacés, arachnides et insectes. In: Duperrey, L.I. (Ed.), Voyage autour du monde ...sur la Corvette de Sa Majesté, La Coquille, pendant les annés 1822, 1823, 1824 et 1825 ... Zoologie. Vol. 2. Part 2. Bertrand, Paris, pp. 24–76.
Jiruskova, A., Motyka, M. & Bocak, L. (2016) High diversity and endemism identified in Cautires (Coleoptera: Lycidae) from the Malay mountain forests. European Journal of Taxonomy, 219, 1–29.
https://doi.org/10.5852/ejt.2016.219Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780.
https://doi.org/10.1093/molbev/mst010Kleine, R. (1926) Coleoptera Lycidae. Nova Guinea. Résultats des expéditions scientifiques à la Nouvelle Guinée, 15, 91–195.
Kleine, R. (1930) Eine neue Brenthide und zwei neue Lyciden aus Neu-Guinea. Nova Guinea. Résultats des expéditions scientifiques à la Nouvelle Guinée, 15, 409–410.
Kleine, R. (1933) Lycidae. Pars 128. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. S. Schenkling, Berlin, pp. 1–145.
Kleine, R. (1935a) Fünf neue Lycidae vom Edie Creek, Britisch Neu Guinea. Nova Guinea, Leiden, 17, 151–154.
Kleine, R. (1935b) Bericht über die von Miss Cheesman in Britisch Neu-Guinea gesammelten Brenthiden und Lyciden. Nova Guinea, Leiden, 17, 303–322.
Kleine, R. (1939) Neue Brenthiden und Lyciden die Miss Cheesman 1936 in Holländisch Neu-Guinea gesammelt hat. Nova Guinea, New Series, 3, 106–112.
Kusy, D. (2017) A new species of Synchonnus (Coleoptera: Lycidae) from New Guinea, with an identification key to the Papuan species. Acta entomologica Musei Nationalis Pragae, 57, 153–160.
Kusy, D., Sklenarova, K. & Bocak, L. (2017) The effectiveness and failure of DNA-based delimitation in Synchonnus net-winged beetles (Coleoptera: Lycidae). Austral Entomology. [early view]
https://doi.org/10.1111/aen.12266)Li, Y., Gunter, N., Pang, H. & Bocak, L. (2015) DNA-based species delimitation separates highly divergent populations within morphologically coherent clades of poorly dispersing beetles. Zoological Journal of the Linnean Society, 175, 59–72. https://doi.org/10.1111/zoj.12262
Meier, R., Shiyang, K., Vaidya, G. & Ng, P.K.L. (2006) DNA barcoding and taxonomy in Diptera: a tale of high intraspecific variability and low identification success. Systematic Biology, 55, 715–728.
https://doi.org/10.1080/10635150600969864Motyka, M. & Bocak, L. (2015) The classification of Flabellochromus and Dumbrellia (Coleoptera: Lycidae) with description of two new species of Calochromus from the Sundaland. Zootaxa, 3941 (1), 125–130.
https://doi.org/10.11646/zootaxa.3941.1.7Nakane, T. (1969) Fauna Japonica. Lycidae (Insecta: Coleoptera). Academic Press of Japan, Tokyo, 224 pp.
Pic, M. (1923) Contribution à l'études des Lycides. L'Echange, 412, 33–36, hors texte.
Riedel, A., Sagata, K., Surbakti, S., Rene, T. & Balke, M. (2013) One hundred and one new species of Trigonopterus weevils from New Guinea. ZooKeys, 280, 1–150.
https://doi.org/10.3897/zookeys.280.3906
Sklenarova, K., Chesters, D. & Bocak, L. (2013) Phylogeography of poorly dispersing net-winged beetles: A role of drifting India in the origin of Afrotropical and Oriental fauna. PLoS ONE, 8, e67957.
https://doi.org/10.1371/journal.pone.0067957Sklenarova, K., Kubecek, V. & Bocak, L. (2014) Subtribal classification of Metriorrhynchini (Insecta: Coleoptera: Lycidae): an integrative approach using molecular phylogeny and morphology of adults and larvae. Arthropod Systematics & Phylogeny, 72, 37–54.
Sherratt, T.N. (2008) The evolution of Müllerian mimicry. Naturwissenschaften, 95, 681–695.
https://doi.org/10.1007/s00114-008-0403-yStamatakis, A. (2006) RAxML-VI-HPC: Maximum Likelihood-based Phylogenetic Analyses with Thousands of Taxa and Mixed Models, Bioinformatics, 22, 2688–2690.
https://doi.org/10.1093/bioinformatics/btl446Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology, 57, 758–771.
https://doi.org/10.1080/10635150802429642Strand, E. (1936) Miscellanea nomenclatorica zoologica et paleaontologica. Folia Zoologica et Hydrobiologica, 9, 167–170.
Waterhouse, C.O. (1879) Illustrations of the typical specimens of Coleoptera in the collection of the British Museum. Part I - Lycidae. British Museum, London, 93 pp.
Wilson, J.S., Jahner, J.P., Williams, K.A. & Forister, M.L. (2013) Ecological and evolutionary processes drive the origin and maintenance of imperfect mimicry. PLoS ONE, 8, e61610.
https://doi.org/10.1371/journal.pone.0061610Zhang, J., Kapli, P., Pavlidis, P. & Stamatakis, A. (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics, 29, 2869-–2876.
https://doi.org/10.1093/bioinformatics/btt499