Abstract
We present the results of a survey of rhaphidophorids in Bhutan more than half a century since the first systematic exploration of orthoptera by the Natural History Museum Basel Expedition to Bhutan in 1972. Only one of two species previously described from Bhutan was resampled. A total of nine new species are described belonging to the family Rhaphidophoridae: five species in genus Diestramima Storozhenko (subfamily Aemodogryllinae), and four species in genus Rhaphidophora Serville (subfamily Rhaphidophorinae). We provide details of morphological structures for all newly described species and their known distribution and habitat. The phylogenetic relationships of the rhaphidophorids from Bhutan are inferred from mitochondrial DNA sequences (COI). New species of Diestramima are not assigned to subgenera because the shape of their paraproct is intermediate when compared to the shapes described for each subgenera. Keys to the Bhutan species based on male epiproct (Rhaphidophora) and posteromedian projection of 7th abdominal tergite (Diestramima) are provided. New species are Diestramima matermagna sp. nov., D. minjiwoonga sp. nov., D. multidenticuli sp. nov., D. samkhara sp. nov., and D. phubdorjia sp. nov.; Rhaphidophora bhutanensis sp. nov., R. bicuspis sp. nov., R. bilobata sp. nov., and R. piscicauda sp. nov..
References
- Bian, X. & Shi, F. (2016) Contribution to the Chinese subfamily Rhaphidophorinae Walker, 1869 (Orthoptera: Rhaphidophoridae; Rhaphidophorinae): New additions to the genera Eurhaphidophora and Stonychophora. Zootaxa, 4109 (1), 46–58. https://doi.org/10.11646/zootaxa.4109.1.4
- Bian, X., Zhu, Q. & Shi, F. (2017) Contribution to the Chinese Rhaphidophorinae Walker, 1869 (Orthoptera: Rhaphidophoridae: Rhaphidophorinae): New species of Rhaphidophora Serville, 1838 from China. Zootaxa, 4317 (2), 261–278. https://doi.org/10.11646/zootaxa.4317.2.4
- Bulgarella, M., Trewick, S.A., Minards, N.A., Jacobson, M.J. & Morgan-Richards, M. (2014) Shifting ranges of two tree weta species (Hemideina spp.): Competitive exclusion and changing climate. Journal of Biogeography, 41 (3), 524–535. https://doi.org/10.1111/jbi.12224
- Cigliano, M.M., Braun, H., Eades, D.C. & Otte, D. (2023) Orthoptera Species File. [http://Orthoptera.SpeciesFile.org]
- Dawwrueng, P., Gorochov, A.V., Tanomtong, A. & Suwannapoom, C. (2020) Contribution to the knowledge of Rhaphidophorinae (Orthoptera: Ensifera: Rhaphidophoridae) from Thailand: three genera Neorhaphidophora, Eurhaphidophora and Minirhaphidophora. Zootaxa, 4853 (2), 235–253. https://doi.org/10.11646/zootaxa.4853.2.5
- Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3 (5), 294–299.
- Gorochov, A.V. (1998) Material on the fauna and systematics of Stenopelmatoidea (Orthoptera) from Indochina and some other territories. I. [in Russian]. 77 (1), 73–105.
- Gorochov, A.V. (1999) Material on the fauna and systematics of Stenopelmatoidea (Orthoptera) from Indochina and some other territories. II. [in Russian]. 78 (1), 60–78.
- Gorochov, A.V. (2001) The higher classification, phylogeny and evolution of the superfamily Stenopelmatoidea. In: Field, L.H. (Ed.), The biology of wetas, king crickets and their allies. CABI Publishing, pp. 3–33. https://doi.org/10.1079/9780851994086.0003
- Gorochov, A.V. (2010) New species of the families Anostostomatidae and Rhaphidophoridae (Orthoptera: Stenopelmatoidea) from China. - Far Eastern Entomologist. 206, 1–16.
- Gorochov, A.V. (2011) Contribution to the fauna and systematics of the Stenopelmatoidea (Orthoptera) of Indochina and some other territories: IX. Entomological Review, 91 (1), 71–89. https://doi.org/10.1134/S0013873811010064
- Gorochov, A.V. (2012) Contribution to the knowledge of the Fauna and systematics of the Stenopelmatoidea (Orthoptera) of Indochina and some other territories: X. Entomological Review, 92 (7), 747–772. https://doi.org/10.1134/S0013873812070032
- Gorochov, A.V. & Storozhenko, S.Y. (1992) On the fauna of the subfamily Aemodogryllinae (Orthoptera, Rhaphidophoridae) in Vietnam. 245, 17–34.
- Gorochov, A.V. & Storozhenko, S.Y. (2015) New and little-known taxa of the tribe Diestramimini (Orthoptera: Rhaphidophoridae: Aemodogryllinae) from Southeast Asia. Part 1. Zoosystematica Rossica, 24 (1), 48–84. https://doi.org/10.31610/zsr/2015.24.1.48
- Gorochov, A.V. & Storozhenko, S.Y. (2019) New and little-known taxa of the tribe Diestramimini (Orthoptera: Rhaphidophoridae: Aemodogryllinae) from Southeast Asia. Part 2. Zoosystematica Rossica, 28 (1), 132–154. https://doi.org/10.31610/zsr/2019.28.1.132
- Hegg, D., Morgan-Richards, M. & Trewick, S.A. (2022) High alpine sorcerers: Revision of the cave wētā genus Pharmacus Pictet & de Saussure (Orthoptera: Rhaphidophoridae: Macropathinae), with the description of six new species and three new subspecies. European Journal of Taxonomy, 808. https://doi.org/10.5852/ejt.2022.808.1721
- Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: Improving the Ultrafast Bootstrap Approximation. Molecular Biology and Evolution, 35 (2), 518–522. https://doi.org/10.1093/molbev/msx281
- Ingrisch, S. (2002) Orthoptera from Bhutan, Nepal, and North India in the Natural History Museum Basel. Entomologica Basiliensia, 24, 123–159.
- Ingrisch, S. (2006) Orthoptera (Insecta) fauna of the Nepal Himalayas: Current knowledge. In Biodiversität Und Naturausstattung Im Himalaya. Erfurt, 2, 73–118.
- Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. & Drummond, A. (2012) Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28 (12), 1647–1649. https://doi.org/10.1093/bioinformatics/bts199
- Kim, D.-Y., Kim, S., Song, H. & Shin, S. (2024) Phylogeny and biogeography of the wingless orthopteran family Rhaphidophoridae. Communications Biology, 7 (1), 1–11. https://doi.org/10.1038/s42003-024-06068-x
- Liu, J., Li, Z. & Bian, X. (2021) Contribution to the Chinese subfamily Rhaphidophorinae Walker, 1869 (Orthoptera: Rhaphidophoridae: Rhaphidophorinae): One new species of Rhaphidophora. Zootaxa, 5027 (4), 597–599. https://doi.org/10.11646/zootaxa.5027.4.8
- Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Woodhams, M.D., von Haeseler, A. & Lanfear, R. (2020) IQ-TREE 2: New Models and Efficient Methods for Phylogenetic Inference in the Genomic Era. Molecular Biology and Evolution, 37 (5), 1530–1534. https://doi.org/10.1093/molbev/msaa015
- Qin, Y., Jiang, H., Liu, X. & Li, K. (2018) A new genus of Rhaphidophorinae (Orthoptera, Rhaphidophoridae) from China. Zootaxa, 4500 (2), 179–194. https://doi.org/10.11646/zootaxa.4500.2.2
- Qin, Y., Wang, H., Liu, X. & Li, K. (2016) A taxonomic study on the species of the genus Diestramima Storozhenko (Orthoptera: Rhaphidophoridae; Aemodogryllinae). Zootaxa, 4126 (4), 514–532. https://doi.org/10.11646/zootaxa.4126.4.4
- Richards, A.M. (1965) The effect of weather on Rhaphidophoridae (Orthoptera) in New Zealand and Australia, 20, 391–400.
- Royal Government of Bhutan (2022) Forest Cover Mapping Report 2022. Forest monitoring and information division, Department of forests and park services, Thimphu, Bhutan. [unknown pagination]
- Saitou, N. & Nei, M. (1987) The neighbor-joining method: A new method for reconstructing phylogenetic trees. Molecular Biology and Evolution, 4 (4), 406–425. https://doi.org/10.1093/oxfordjournals.molbev.a040454
- Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, Weighting, and Phylogenetic Utility of Mitochondrial Gene Sequences and a Compilation of Conserved Polymerase Chain Reaction Primers. Annals of the Entomological Society of America, 87 (6), 651–701. https://doi.org/10.1093/aesa/87.6.651
- Storozhenko, S.Y. & Dawwrueng, P. (2014) New genus of the tribe Diestramimini (Orthoptera: Rhaphidophoridae) from Thailand. Zootaxa, 3765 (3). https://doi.org/10.11646/zootaxa.3765.3.5
- Wang, P., Zhu, Q. & Shi, F. (2019) Supplement of the genus Diestramima Storozhenko, 1990 (Orthoptera: Rhaphidophoridae: Aemodogryllinae) from China. Zootaxa, 4615 (3), 577–584. https://doi.org/10.11646/zootaxa.4615.3.10
- Würmli, M. (1973) Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen Museums in Basel. Orthoptera: Gryllacridoidea. Verh. Naturf. Ges. Basel, 83, 337–347.
- Zhu, Q. & Shi, F.-M. (2018) Review of the genus Diestramima Storozhenko, 1990 (Orthoptera: Rhaphidophoridae: Aemodogryllinae) from China. Zootaxa, 4450 (2), 249–274. https://doi.org/10.11646/zootaxa.4450.2.5
- Zong, J.-S., Qin, Y.-Y., Li, K. & He, Z.-Q. (2021) Report of a new genus Arboramima belonging to tribe Diestramimini, with a new species A. cattusis from Guangxi, China (Orthoptera: Rhaphidophoridae: Aemodogryllinae). Zootaxa, 4981 (3), 593–600. https://doi.org/10.11646/zootaxa.4981.3.9