Abstract
Three new species of the genus Himalopsyche (Trichoptera, Rhyacophilidae) from the Hengduan Mountains in China are described. Species delimitation was based on diagnostic features of genitalia, as well as molecular data from six genes analysed using the multi-species coalescent method STACEY. Formal descriptions are focused on genital morphology. Males of Himalopsyche viteceki sp. nov. are most similar to those of H. alticola and H. martynovi, and females are most similar to those of H. tibetana and H. velata. Himalopsyche immodesta sp. nov. is described based on a single male specimen and it most resembles the males of H. viteceki. Males of H. velata sp. nov. are most similar to H. tibetana, and females are most similar to those of H. maxima and H. tibetana. Diagnostic characters are found on segment IX and the superior and inferior appendages of male genitalia, and most notably on segment VIII in female genitalia. The newly discovered species underline the Hengduan Mountains as a potential source of yet undiscovered aquatic biodiversity.
References
Banks, N. (1940) Report on certain groups of neuropteroid insects from Szechwan, China. Proceedings of the United Stated National Museum, 88, 173–220.
https://doi.org/10.5479/si.00963801.88-3079.173
Banks, N. (1947) Some neuropterous insects from Szechwan, China. Fieldiana-Zoology, Chicago Natural History Museum, 31, 97–107.
https://doi.org/10.5962/bhl.title.2821
Blahnik, R.J., Holzenthal, R.W. & Prather, A. (2007) The lactic acid method for clearing Trichoptera genitalia. In: Bueno-Soria, J., Barba-Álvarez, R. & Armitage, B. (Eds.), Proceedings of the XIIth International Symposium on Trichoptera, 18–22 June, 2006. The Caddis Press, Columbus, Ohio, pp. 9–14.
Bouckaert, R. & Drummond, A. (2017) bModelTest: Bayesian phylogenetic site model averaging and model comparison. BMC Evolutionary Biology, 17, 42.
https://doi.org/10.1186/s12862-017-0890-6
Bouckaert, R., Heled, J., Kühnert, D., Vaughan, T., Wu, C.-H., Xie, D., Suchard, M.A., Rambaut, A. & Drummond, A.J. (2014) BEAST 2: A Software Platform for Bayesian Evolutionary Analysis. PLoS Computational Biology, 10 (4), e1003537.
https://doi.org/10.1371/journal.pcbi.1003537
Favre, A., Päckert, M., Pauls, S.U., Jähnig, S.C., Uhl, D., Michalak, I. & Muellner-Riehl, A.N. (2015) The role of the uplift of the Qinghai-Tibetan Plateau for the evolution of Tibetan biotas. Biological Reviews, 90, 236–253.
https://doi.org/10.1111/brv.12107
Flint, O.S. (1961) The presumed larva of Himalopsyche phryganea (Ross) (Trichoptera: Rhyacophilidae). The Pan-Pacific Entomologist, 37, 199–202.
Forsslund, H.K. (1935) Schwedisch-chinesische wissenschaftliche Expedition nach den nordwestlichen Provinzen Chinas. Arkiv för Zoologi, 1 (31), 1–21.
Fujisawa, T. & Barraclough, T.G. (2013) Delimiting species using single-locus data and the generalized mixed yule coalescent approach: A revised method and evaluation on simulated data sets. Systematic Biology, 62, 707–724.
https://doi.org/10.1093/sysbio/syt033Graf, W. & Sharma, S. (1998) The larva of Himalopsyche tibetana (Insecta: Trichoptera, Rhyacophilidae) with some ecological notes on the genus Himalopsyche Banks, 1940 from Nepal. In: Chalise, S.R. & Khanal, N.R. (Eds.), International Conference on Ecohydrology of High Mountain Areas. International Centre for Integrated Mountain Development, Kathmandu, pp. 561–571.
Hjalmarsson, A.E., Graf, W., Jähnig, S., Cai, Q., Tong, X., Li, F., Tachamo Shah, R.D., Narayan Shah, D., Sharma, S. & Pauls, S.U. (2018) Life stage association and morphological characterization of larval types of Himalopsyche (Trichoptera: Rhyacophilidae). ZooKeys, 773, 79–108.
https://doi.org/10.3897/zookeys.773.24319
Hjalmarsson, A.E., Graf, W., Vitecek, S., Jähnig, S.C., Cai, Q., Sharma, S., Tong, X., Li, F., Narayan Shah, D., Tachamo Shah, R.D. & Pauls, S.U. (2019) Molecular phylogeny of Himalopsyche (Trichoptera, Rhyacophilidae). Systematic Entomology (early view).
https://doi.org/10.1111/syen.12367
Hoorn, C., Mosbrugger, V., Mulch, A. & Antonelli, A. (2013) Biodiversity from mountain building. Nature Geoscience, 6, 154–154.
https://doi.org/10.1038/ngeo1742Hsu, L.P. (1997) A taxonomic study of Trichoptera from Taiwan (Insecta). Ph.D. Dissertation, Tokai University, Taipei 370 pp. [in Chinese with English abstract]
Jokusch, E.L., Martínez-Solano, I. & Timpe, E.K. (2014) The effects of inference method, population sampling, and gene sampling on species tree inferences: An empirical study in slender salamanders (Plethodontidae: Batrachoseps). Systematic Biology, 64, 66–83.
https://doi.org/10.1093/sysbio/syu078
Jones, G. (2017) Algorithmic improvements to species delimitation and phylogeny estimation under the multispecies coalescent. Journal of Mathematical Biology, 74, 447–467.
https://doi.org/10.1007/s00285-016-1034-0
Jones, G., Aydin, Z. & Oxelman, B. (2015) DISSECT: An assignment-free Bayesian discovery method for species delimitation under the multispecies coalescent. Bioinformatics, 31, 991–998.
https://doi.org/10.1093/bioinformatics/btu770
Katoh, K., Misawa, K., Kuma, K. & Miyata, T. (2002) MAFFT: A novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30 (14), 3059–3066.
https://doi.org/10.1093/nar/gkf436
Kawai T. & Tanida, K. (2005) Aquatic Insects of Japan: Manual with Keys and Illustrations. Tokai University Press, Kanagawa, 1342 pp. [in Japanese]
Kimmins, D.E. (1952) XL.—Indian caddis flies.—VI. New species and a new genus of the subfamily Rhyacophilinae. Annals and Magazine of Natural History, Series 12, 5 (52), 347–361.
https://doi.org/10.1080/00222935208654302
Lakhwinder, K. & Saini, M.S. (2015) A new species of the genus Himalopsyche (Trichoptera, Rhyacophilidae), with key to and catalogue of Indian species. Vestnik zoologii, 49, 3–12.
Larsson, A. (2014) AliView: A fast and lightweight alignment viewer and editor for large data sets. Bioinformatics, 30 (22), 3276–3278.
https://doi.org/10.1093/bioinformatics/btu531Lepneva, S.G. (1970) Fauna of the U.S.S.R., Trichoptera, Volume II No. 1, Larvae and Pupae of Annulipalpia. Smithsonian Institution and the National Science Foundation, Washington, D.C., 656 pp.
Li, X., Jiang, N., Cheng, R., Xue, D., Qu, Y. & Han, H. (2017) Allopatric divergence and secondary contact without genetic admixture for Arichanna perimelaina (Lepidoptera: Geometridae), an alpine moth endemic to the Hengduan Mountains. Systematic Entomology, 42, 703–713.
https://doi.org/10.1111/syen.12234
Malicky, H. (1978) Beiträge zur Kenntnis der Insektenfauna Sumatras. Teil 7: Köcherfliegen (Trichoptera) aus Sumatra und West-Neuguinea I. Rhyacophilidae, Glossosomatidae, Stenopsychidae, Goeridae. Beiträge zur naturkundlichen Forschung in Südwestdeutschland, 37, 159–173.
Malicky, H. (2000) Einige neue Köcherfliegen aus Sabah, Nepal, Indien und China (Trichoptera: Rhyacophilidae, Hydrobiosidae, Philopotamidae, Polycentropodidae, Ecnomidae, Psychomyiidae, Hydropsychidae, Brachycentridae, Odontoceridae, Molannidae). Braueria, 27, 32–39.
Malicky, H. (2011) Neue Trichopteren aus Europa und Asien. Braueria, 38, 23–43.
Martynov, A.V. (1914) Contributions a la faune des Trichoptères de la Chine. Annuaire du Musée Zoologique de l’Académie Impériale des Sciences de Saint Pétersbourg, 19, 323–339.
Martynov, A.V. (1930) On the trichopterous fauna of China and Eastern Tibet. Proceedings of the Zoological Society of London, 5, 65–112.
https://doi.org/10.1111/j.1096-3642.1930.tb00967.x
Martynov, A.B. (1935) On a collection of Trichoptera from the Indian Museum. Part I.—Annulipalpia. Records of the Indian Museum, 37, 93–306.
Rannala, B. (2005) The art and science of species delimitation Current Zoology, 61, 846–853.
https://doi.org/10.1093/czoolo/61.5.846
Ross, H.H. (1956) Evolution and Classification of the Mountain Caddisflies. University of Illinois Press, Urbana, pp.
Saini, M.S. & Kaur, L. (2011) Revised phylogenetic analysis of Indian species of genus Himalopsyche Banks (Trichoptera: Spicipalpia: Rhyacophilidae). Halteres, 3, 26–29.
Saito, T. 1965. Larvae of two fresh water insects, Himalopsyche japonica and Neohapalothrix kanii. Report of the Japanese Society of Systematic Zoology, 1, 14–16.
Schlick-Steiner, B.C., Steiner, F.M., Seifert, B., Stauffer, C., Christian, E. & Crozier, R.H. (2010) Integrative taxonomy: A multisource approach to exploring biodiversity. Annual Review of Entomology, 55 (1), 421–438.
https://doi.org/10.1146/annurev-ento-112408-085432
Schmid, F. (1963) Quelques Himalopsyche indiènnes (Trichoptères, Rhyacophilidae). Bonner Zoologische Beiträge, 14, 206–223.
Schmid, F. (1969) Sur trois Himalopsyche d’URSS (Trichoptères, Rhyacophilides). Le Naturaliste Canadien, 96, 243–346.
Schmid, F. (1970) Le genre Rhyacophila et la famille des Rhyacophilidae (Trichotera). Mémoires de la Société Entomologique du Canada, 66, 1–230.
https://doi.org/10.4039/entm10266fv
Schmid, F. & Botosaneanu, L. (1966) Le genre Himalopsyche Banks (Trichoptera, Rhyacophilidae). Annales de la Société entomologique du Québec, 11, 123–176.
Tanida, K. (1985) Trichoptera. In: Kawai, T. (Ed.), An Illustrated Book of Aquatic Insects of Japan. Tokai University Press, Tokyo, pp. 167–215.
Thamsenanupap P., Chantaramongkol, P. & Malicky, H. (2005) Description of caddis larvae (Trichoptera) from northern Thailand of the genera Himalopsyche (Rhyacophilidae), Arctopsyche (Arctopsychidae), cf. Eoneureclipsis (Psychomyiidae) and Inthanopsyche (Odontoceridae). Braueria, 32, 7–11.
Ulmer, G. (1905) Neue und wenig bekannte Trichopteren der Museen zu Brussel und Paris. Annales de la Société Entomologique de Belgique, 49, 17–42.
Vitecek, S., Kučinić, M., Previšić, A., Živić, I., Stojanović, K., Keresztes, L., Bálint, M., Hoppeler, F., Waringer, J., Graf, W. & Pauls, S.U. (2017) Integrative taxonomy by molecular species delimitation: Multi-locus data corroborate a new species of Balkan Drusinae micro-endemics. BMC Evolutionary Biology, 17, 129.
https://doi.org/10.1186/s12862-017-0972-5
Xing, Y. & Ree, R.H. (2017) Uplift-driven diversification in the Hengduan Mountains, a temperate biodiversity hotspot. Proceedings of the National Academy of Sciences, 114 (17), E3444–E3451.
https://doi.org/10.1073/pnas.1616063114
Yang, Z. & Rannala, B.H. (2010) Bayesian species delimitation using multilocus sequence data. Proceedings of the National Academy of Sciences, 107, 9264–9269.
https://doi.org/10.1073/pnas.0913022107
Zhang, J., Kapli, P., Pavlidis, P. & Stamatakis, A. (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics, 29 (12), 2869–2876.
https://doi.org/10.1093/bioinformatics/btt499