Abstract
Here we present updates on the taxonomy and distribution of the genera Pachyserica Brenske, 1898 and Serica MacLeay, 1819 from East Asia. Thirty eight new species are desribed from China, Myanmar, South Korea, and Vietnam: Pachyserica chenchangchini Ahrens, Zhao, Pham & Liu, new species, P. dieuthuyae Ahrens, Zhao, Pham & Liu, new species, P. hoabinhensis Ahrens, Zhao, Pham & Liu, new species, P. motuo Ahrens, Zhao, Pham & Liu, new species, P. natmatoung Ahrens, Zhao, Pham & Liu, new species, P. sanqingshanensis Ahrens, Zhao, Pham & Liu, new species, P. sunfengyii Ahrens, Zhao, Pham & Liu, new species, P. tayyentu Ahrens, Zhao, Pham & Liu, new species, P. tianxuani Ahrens, Zhao, Pham & Liu, new species, P. wangzizhaoi Ahrens, Zhao, Pham & Liu, new species, P. yaonani Ahrens, Zhao, Pham & Liu, new species, P. yinhengi Ahrens, Zhao, Pham & Liu, new species, P. zhanbaoxiangi Ahrens, Zhao, Pham & Liu, new species, Serica (s. l.) anhua Ahrens, Zhao, Pham & Liu, new species, S. (s. l.) camura Ahrens, Zhao, Pham & Liu, new species, S. (s. l.) fengxue Ahrens, Zhao, Pham & Liu, new species, S. (s. l.) nhiae Ahrens, Zhao, Pham & Liu, new species, S. (s. l.) paracallosericoides Ahrens, Zhao, Pham & Liu, new species, S. (s. l.) taythien Ahrens, Zhao, Pham & Liu, new species, S. (s. l.) wuyishan Ahrens, Zhao, Pham & Liu, new species, S. (s. l.) yuechengling Ahrens, Zhao, Pham & Liu, new species, S. (Serica) assingi Ahrens, Zhao, Pham & Liu, new species, S. (S.) bomi Ahrens, Zhao, Pham & Liu, new species, S. (S.) christophreuteri Ahrens, Zhao, Pham & Liu, Ahrens, Zhao, Pham & Liu, new species, S. (S.) cuona Ahrens, Zhao, Pham & Liu, new species, S. (S.) fansipan Ahrens, Zhao, Pham & Liu, new species, S. (S.) gongtonggouensis Ahrens, Zhao, Pham & Liu, new species, S. (S.) guangxiensis Ahrens, Zhao, Pham & Liu, new species, S. (S.) gwangjuensis Ahrens, Zhao, Pham & Liu, new species, S. (S.) hongyii Ahrens, Zhao, Pham & Liu, new species, S. (S.) jiangda Ahrens, Zhao, Pham & Liu, new species, S. (S.) jiulaoci Ahrens, Zhao, Pham & Liu, new species, S. (S.) lushui Ahrens, Zhao, Pham & Liu, new species, S. (S.) liyitengi Ahrens, Zhao, Pham & Liu, new species, S. (S.) qizhihaoi Ahrens, Zhao, Pham & Liu, new species, S. (S.) xizang Ahrens, Zhao, Pham & Liu, new species, S. (S.) zhamu Ahrens, Zhao, Pham & Liu, new species, and S. (Taiwanoserica) yexiaohani Ahrens, Zhao, Pham & Liu, new species. The lectotype of Pachyserica scalaris Arrow, 1946 is designated and its male genitalia is illustrated. Additional records of 66 other species are provided.
References
- Ahrens, D. (1996) Revision der Sericini des Himalaya und angrenzender Gebiete. Die Gattungen Lasioserica Brenske, 1896 und Gynaecoserica Brenske, 1896. Coleoptera- Schwanfelder Coleopterologische Mitteilungen, 16, 1–48.
- Ahrens, D. (1999) Revision der Gattung Serica (s. str.) MacLeay des Himalaya-Gebiets (Coleoptera, Melolonthidae). Fragmenta Entomologica, 31, 205–332.
- Ahrens, D. (2000a) Weiteres Material der Gattung Lasioserica Brenske, 1896, mit der Beschreibung acht neuer Arten (Coleoptera: Melolonthidae: Sericini). Acta Musei Moraviae, Scientiae Biologicae, 85, 1–28.
- Ahrens, D. (2000b) Synopsis der Gattung Gastroserica Brenske, 1897 des ostasiatischen Festlandes (Coleoptera: Melolonthidae: Sericini). Entomologische Abhandlungen des Staatlichen Museums für Tierkunde Dresden, 59, 73–121.
- Ahrens, D. (2000c) Serica korgei Petrovitz, 1967 aus der Türkei- ein neues Synonym von Autoserica castanea Arrow, 1913 (Coleoptera: Melolonthidae: Sericini). Reichenbachia, 33 (51), 395–298.
- Ahrens, D. (2001) Zwei neue Chrysoserica-Arten aus dem Himalaya (Coleoptera, Melolonthidae). Bolletino della Società Entomologica Italiana, 133 (2), 133–147.
- Ahrens, D. (2002) Notes on distribution and synonymy of sericid beetles from Taiwan, with descriptions of new species (Coleoptera, Scarabaeoidea, Sericini). Annales Historico-Naturales Musei Nationalis Hungarici, 94, 53–91.
- Ahrens, D. (2003a) Neue Arten aus der Gruppe der Amiserica insperata (Brenske, 1898) (Coleoptera, Scarabaeoidea, Sericini). Spixiana, 26, 65–81.
- Ahrens, D. (2003b) Zur Identität der Gattung Neoserica Brenske, 1894, nebst Beschreibung neuer Arten (Coleoptera, Melolonthidae, Sericini). Koleopterologische Rundschau, 73, 169–226.
- Ahrens, D. (2003c) Zur Identität der aus Madagaskar beschriebenen Autoserica stupida Brenske, 1900 (Coleoptera, Scarabaeidae, Sericini). Beiträge zur Entomologie, 53, 231–238. https://doi.org/10.21248/contrib.entomol.53.1.231-238
- Ahrens, D. (2003d) On the Identity of Maladera infuscata (Moser, 1915), with description of a new species from Taiwan (Coleoptera, Scarabaeidae, Sericini). Zootaxa, 392 (1), 1–8. https://doi.org/10.11646/zootaxa.392.1.1
- Ahrens, D. (2004) Monographie der Sericini des Himalaya (Coleoptera, Scarabaeidae). Dissertation.de - Verlag im Internet GmbH, Berlin, 534 pp.
- Ahrens, D. (2005a) A taxonomic review on the Serica (s. str.) MacLeay, 1819 species of Asiatic mainland (Coleoptera, Scarabaeidae, Sericini). Nova Supplementa Entomologica, 18, 1–163.
- Ahrens, D. (2005b) Taxonomic revision of the genus Anomalophylla Reitter, 1887 (Coleoptera, Scarabaeidae: Sericini). Zootaxa, 1076 (1), 1–62. https://doi.org/10.11646/zootaxa.1076.1.1
- Ahrens, D. (2006a) The phylogeny of the genus Lasioserica inferred from adult morphology – implications on the evolution of montane fauna of the South Asian orogenic belt (Coleoptera: Scarabaeidae: Sericini). Journal of Zoological Systematics and Evolutionary Research, 44 (1), 34–53. https://doi.org/10.1111/j.1439-0469.2005.00340.x
- Ahrens, D. (2006b) Sericinae. In: Löbl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Volume 3. Scarabaeoidea – Scirtoidea – Dascilloidea – Buprestoidea – Byrrhoidea. Apollo Books, Stenstrup, pp. 229–248.
- Ahrens, D. (2006c) Revision und phylogenetische Analyse der Gattung Pachyserica Brenske, 1897 (Coleoptera, Scarabaeidae, Sericini). Revue Suisse de Zoologie, 113, 487–557. https://doi.org/10.5962/bhl.part.80360
- Ahrens, D. (2007a) Revision der Serica nigroguttata Brenske, 1897 - Gruppe (Coleoptera, Scarabaeidae, Sericini). Bulletin de l’Institut Royal des Sciences naturelles de Belgique, 77, 5–37.
- Ahrens, D. (2007b) Taxonomic changes and an updated Catalogue of Palearctic Sericini (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 1504 (1), 1–51. https://doi.org/10.11646/zootaxa.1504.1.1
- Ahrens, D. (2007c) Beetle evolution in the Asian highlands: insight from a phylogeny of the scarabaeid subgenus Serica (Coleoptera, Scarabaeidae). Systematic Entomology, 32, 450–476. https://doi.org/10.1111/j.1365-3113.2006.00373.x
- Ahrens, D. (2009) A cladistic analysis reveals the occurrence of Taiwanoserica in Sichuan (China) with description of four new species (Coleoptera, Scarabaeidae). Annales de la Société Entomologique de France, New Series, 45 (3), 285–296. https://doi.org/10.1080/00379271.2009.10697612
- Ahrens, D. (2011) A revision of the genus Archeohomaloplia Nikolaev, 1982. (Coleoptera: Scarabaeidae: Sericini). Bonn Zoological Bulletin, 60, 117–138.
- Ahrens, D. (2012) New species and a morphology-based phylogeny of the genus Nepaloserica (Coleoptera, Scarabaeidae). In: Hartmann, M. & Weipert, M. (Eds.), Biodiversität und Naturausstattung im Himalaya III. Verein der Freunde und Förderer des Naturkundemuseums Erfurt e.V., Erfurt, pp. 305–316
- Ahrens, D. (2021a) Two new species of the Neoserica (sensu stricto) group from China (Coleoptera: Scarabaeidae: Melolonthinae: Sericini). Journal of Natural History, 54 (45–46), 2927–2936. https://doi.org/10.1080/00222933.2021.1884761
- Ahrens, D. (2021b) Additions to the fauna of Tetraserica Ahrens, 2004 of China and India (Coleoptera, Scarabaeidae: Sericini). Koleopterologische Rundschau, 91, 131–135.
- Ahrens, D. (2021c) New Sericini from the Zoological Institute St. Petersburg (Insecta: Coleoptera: Scarabaeidae). In: Hartmann, M., Barclay, M. & Weipert, M. (Eds.), Biodiversität und Naturausstattung im Himalaya VII. Verein der Freunde und Förderer des Naturkundemuseums Erfurt e.V., Erfurt, pp. 427–433.
- Ahrens, D. (2023a) Two new species of Archeohomaloplia Nikolajev, 1972 from China (Coleoptera, Scarabaeidae, Sericini). Bonn Zoological Bulletin, 72 (1), 55–61. https://doi.org/10.20363/BZB-2023.72.1.055
- Ahrens, D. (2023b) Updates on the Neoserica vulpes group (Coleoptera, Scarabaeidae, Melolonthinae, Sericini): new species and records. European Journal of Taxonomy, 888, 97–110. https://doi.org/10.5852/ejt.2023.888.2231
- Ahrens, D. (2024) Species diagnosis and DNA taxonomy. In: DeSalle, R. (Ed.), DNA Barcoding. Methods in Molecular Biology. Vol. 2744. Humana, New York, New York, pp. 33–52. https://doi.org/10.1007/978-1-0716-3581-0_2
- Ahrens, D. & Bezděk, A. (2016) Sericini. In: Löbl, I. & Löbl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Volume 3. Scarabaeoidea – Scirtoidea – Dascilloidea – Buprestoidea - Byrrhoidea. 2nd Edition. Revised and Updated Edition. Brill, Leiden, pp. 281–317.
- Ahrens, D. & Bezděk, A. (2024) On the publication year of Sericini genera and species described by Brenske in “Die Serica-Arten der Erde” (Coleoptera: Scarabaeidae: Sericinae). Zootaxa, 5418 (5), 442–470. https://doi.org/10.11646/zootaxa.5418.5.2
- Ahrens, D. & Bezděk, A. (2024) Catalogue of Palaearctic Sericinae (Coleoptera: Scarabaeidae) – Revised and updated 4th edition. Zootaxa, submitted.
- Ahrens, D. & Fabrizi, S. (2009a) A review on the genus Gynaecoserica Brenske, 1897 (Coleoptera, Scarabaeidae, Sericini). Journal of Natural History, 43, 1505–1584. https://doi.org/10.1080/00222930902968809
- Ahrens, D. & Fabrizi, S. (2009b) New species of Sericini from the Eastern Himalaya and Tibet (Coleoptera, Scarabaeidae). In: Hartmann, M. & Weipert, M. (Eds.), Biodiversität und Naturausstattung im Himalaya III. Verein der Freunde und Förderer des Naturkundemuseums Erfurt e.V., Erfurt, pp. 249–284.
- Ahrens, D. & Fabrizi, S. (2011) New species of Sericini from the Himalaya and adjacent mountains (Coleoptera: Scarabaeidae). Bonn Zoological Bulletin, 60, 139–164.
- Ahrens, D. & Fabrizi, S. (2016) A Monograph of the Sericini of India (Coleoptera: Scarabaeidae). Bonn Zoological Bulletin, 65, 1–355.
- Ahrens, D. & Pacholátko, P. (2003) New data on distribution of the species of Gastroserica Brenske, 1897, with description of new taxa from China and Laos (Coleoptera: Scarabaeidae: Sericini). Zootaxa, 342 (1), 1–18. https://doi.org/10.11646/zootaxa.342.1.1
- Ahrens, D. & Pacholátko, P. (2007) Two new species of Gastroserica Brenske, 1897 from China and Myanmar (Coleoptera: Scarabaeidae: Sericini). Entomologische Zeitschrift, 117, 137–139.
- Ahrens, D. & Vogler, A.P. (2008) Towards the phylogeny of chafers (Sericini): analysis of alignment-variable sequences and the evolution of segment numbers in the antennal club. Molecular Phylogenetics and Evolution, 47, 783–798. https://doi.org/10.1016/j.ympev.2008.02.010
- Ahrens, D., Liu, W.G., Fabrizi, S., Bai, M. & Yang, X.K. (2014a) A taxonomic review of the Neoserica (sensu lato) septemlamellata group (Coleoptera: Scarabaeidae: Sericini). ZooKeys, 402, 76–102. https://doi.org/10.3897/zookeys.402.7360
- Ahrens, D., Liu, W.G., Fabrizi, S., Bai, M. & Yang, X.K. (2014b) A taxonomic review of the Neoserica (sensu lato) abnormis group (Coleoptera: Scarabaeidae: Sericini). ZooKeys, 439, 28–82. https://doi.org/10.3897/zookeys.439.8055
- Ahrens, D., Liu, W.G., Fabrizi, S., Bai, M. & Yang, X.K. (2014c) A revision of the species of the Neoserica (sensu lato) vulpes group (Coleoptera: Scarabaeidae: Sericini). Journal of Natural History. 49 (17–18), 1073–1130. https://doi.org/10.1080/00222933.2014.974707
- Ahrens, D., Liu, W.G., Fabrizi, S. & Bai, M. (2021a) Taxonomic review on the Trioserica Moser, 1922 species of China (Coleoptera: Scarabaeidae: Melolonthinae: Sericini). Zootaxa, 4999 (4), 343–355. https://doi.org/10.11646/zootaxa.4999.4.4
- Ahrens, D., Liu, W.G., Pham P. & Fabrizi, S. (2021b) An overview of the genus Amiserica (Coleoptera, Scarabaeidae, Melolonthinae, Sericini). Zootaxa, 5050 (1), 1–63. https://doi.org/10.11646/zootaxa.5050.1.1
- Ahrens, D., Liu, W.G., Fabrizi, S. & Bai, M. (2022) Taxonomic revision of the species of Serica MacLeay, 1819 (sensu lato) from China and adjacent areas (Coleoptera, Scarabaeidae, Melolonthinae, Sericini), with an update on Nipponoserica Nomura, 1972. Zootaxa, 5186 (1), 1–83. https://doi.org/10.11646/zootaxa.5186.1.1
- Ahrens, D., Liu, W.G., Lukic D. & Bai, M. (2023) A taxonomic review of Microserica from continental Asia (Coleoptera, Scarabaeidae, Melolonthinae, Sericini). Zootaxa, 5241 (1), 1–115. https://doi.org/10.11646/zootaxa.5241.1.1
- Bohacz, C. & Ahrens, D. (2020) A new species of the Neoserica (sensu lato) vulpes group from China, with a corrigendum on Neoserica ailaoshanica Liu et al., 2014 (Coleoptera: Scarabaeidae: Sericini). Journal of Natural History, 53 (47–48), 2991–2997. https://doi.org/10.1080/00222933.2020.1758819
- Dietz, L., Eberle, J., Kukowka, S., Mayer, C., Bohacz, C., Baur, H., Espeland, M., Huber, B., Hutter, C., Mengual, X., Peters, R.S., Vences, M., Wesener, T., Willmott, K., Misof, B., Niehuis, O. & Ahrens, D. (2023) Standardized nuclear markers improve and homogenize species delimitation in Metazoa. Methods in Ecology and Evolution, 14, 543–555. https://doi.org/10.1111/2041-210X.14041
- Eberle, J., Warnock, R.C.M. & Ahrens, D. (2016) Bayesian species delimitation in Pleophylla chafers (Coleoptera) – the importance of prior choice and morphology. BMC Evolutionary Biology, 16, 94. https://doi.org/10.1186/s12862-016-0659-3
- Eberle, J., Fabrizi, S., Lago, P. & Ahrens, D. (2017a) A historical biogeography of megadiverse Sericini – another story “out of Africa”? Cladistics, 33, 183–197. https://doi.org/10.1111/cla.12162
- Eberle, J., Beckett, M., Özguel-Siemund, A., Shresta, J., Fabrizi, S. & Ahrens, D. (2017b) Afromontane forests hide 19 new species of ancient Pleophylla chafers (Coleoptera: Scarabaeidae): phylogeny and taxonomic revision. Zoological Journal of Linnean Society, 180, 321–353.
- Fabrizi, S., Liu, W.G., Bai, M., Yang, X.K. & Ahrens, D. (2021) A monograph of the genus Maladera of China (Coleoptera, Scarabaeidae, Sericini). Zootaxa, 4922 (1), 1–400. https://doi.org/10.11646/zootaxa.4922.1.1
- Liu, W.G. & Ahrens, D. (2014) [new taxa]. In: Liu, W.G., Bai, M., Yang, X.K. & Ahrens, D. (2014) Towards an improved knowledge of Sericini of the Tibetan highland: new species and records (Coleoptera, Scarabaeidae). Bonn Zoological Bulletin, 63, pp. 157–172.
- Liu, W.G., Ahrens, D., Bai, M. & Yang, X.K. (2011) A key to species of the genus Gastroserica Brenske of the China (Coleoptera, Scarabaeidae, Sericini), with the description of two new species and two new records for China. ZooKeys, 139, 23–44. https://doi.org/10.3897/zookeys.139.1702
- Liu, W.G., Bai, M., Yang, X.K. & Ahrens, D. (2014a) An update to the taxonomy of the genus Gastroserica Brenske (Coleoptera, Scarabaeidae, Sericini). ZooKeys, 426, 87–110. https://doi.org/10.3897/zookeys.426.7578
- Liu, W.G., Bai, M., Yang, X.K. & Ahrens, D. (2014b) Towards an improved knowledge of Sericini of the Tibetan highland: new species and records (Coleoptera, Scarabaeidae). Bonn Zoological Bulletin, 63, 157–172.
- Liu, W.G., Fabrizi, S., Bai, M., Yang, X.K. & Ahrens, D. (2014c) A taxonomic revision of the Neoserica (s.l.) pilosula group (Coleoptera, Scarabaeidae, Sericini). ZooKeys, 440, 89–113. https://doi.org/10.3897/zookeys.440.8126
- Liu, W.G., Fabrizi, S., Bai, M., Yang, X.K. & Ahrens, D. (2014d) A taxonomic revision of the Neoserica (sensu lato) calva group (Coleoptera, Scarabaeidae, Sericini). ZooKeys, 448, 47–81. https://doi.org/10.3897/zookeys.448.8368
- Liu, W.G., Fabrizi, S., Bai, M., Yang, X.K. & Ahrens, D. (2014e) A review of the Tetraserica species of China (Coleoptera, Scarabaeidae, Sericini). ZooKeys, 448, 83–121. https://doi.org/10.3897/zookeys.443.8429
- Liu, W.G., Bai, M., Yang, X.K. & Ahrens, D. (2015a) New species and records of the Neoserica (sensu stricto) group (Coleoptera, Scarabaeidae, Sericini). Journal of Natural History, 49 (39–40), 2379–2395. https://doi.org/10.1080/00222933.2015.1034208
- Liu, W.G., Bai, M., Eberle, J., Yang, X.K. & Ahrens, D. (2015b) A phylogeny of Sericini with particular reference to Chinese species using mitochondrial and ribosomal DNA (Coleoptera: Scarabaeidae). Organisms Diversity and Evolution, 15, 343–350. https://doi.org/10.1007/s13127-015-0204-z
- Liu, W.G., Fabrizi, S., Bai, M., Yang, X.K. & Ahrens, D. (2016) A taxonomic revision of Neoserica (sensu lato): the species groups N. lubrica, N. obscura, and N. silvestris (Coleoptera, Scarabaeidae, Sericini). ZooKeys, 635, 123–160. https://doi.org/10.3897/zookeys.635.9915
- Liu, W.G., Bai, M., Yang, X.K. & Ahrens, D. (2017a) New species and records of Lasioserica and Gynaecoserica from China (Coleoptera, Scarabaeidae, Sericini). Bonn Zoological Bulletin, 66, 29–36.
- Liu, W.G., Fabrizi, S., Bai, M., Yang, X.K. & Ahrens, D. (2017b) New species of Nipponoserica and Paraserica from China (Coleoptera: Scarabaeidae: Sericini). ZooKeys, 721, 65–91. https://doi.org/10.3897/zookeys.721.13918
- Liu, W.G., Fabrizi, S., Bai, M., Yang, X.K. & Ahrens, D. (2019) A taxonomic revision of Chinese Neoserica (sensu lato): final part (Coleoptera: Scarabaeidae: Sericini). Bonn Zoological Bulletin, Supplement, 64, 1–71. https://doi.org/10.20363/BZB-S-2019.64
- Pacheco, T.L., Monné, M.L. & Ahrens, D. (2023) Morphology-based phylogenetic analysis of South American Sericini chafers (Coleoptera, Scarabaeidae) contrasts patterns of morphological disparity and current classification. Zoologischer Anzeiger, 302, 43–57. https://doi.org/10.1016/j.jcz.2022.11.004
- Rana, P.S., Fabrizi, S. & Ahrens, D. (2017) New records of Sericini and a new Serica (s. str.) species from Nepal (Coleoptera: Scarabaeidae: Sericini). Entomologische Zeitschrift, 127, 144–148.
- Ranasinghe, U.G.S.L., Eberle, J., Thormann, J., Bohacz, C., Benjamin, S.P. & Ahrens, D. (2022) Multiple species delimitation approaches with COI barcodes poorly fit each other and with morphospecies – an integrative taxonomy case of Sri Lankan Sericini chafers (Coleoptera: Scarabaeidae). Ecology and Evolution, 12 (5), e8942. https://doi.org/10.1002/ece3.8942.
- Ranasinghe, U.G.S.L., Thormann, J., Benjamin, S.P., Bezděk, A., Eberle, J. & Ahrens, D. (2023) Contrasting results of multiple species delimitation approaches cause uncertainty in synecological studies: a case study on Sri Lankan chafers. Insect Conservation and Diversity, 16 (6), 870–885. https://doi.org/10.1111/icad.12684
- Sreedevi, K., Speer, J., Fabrizi, S. & Ahrens, D. (2018) New records and species of Sericini from Indian subcontinent (Coleoptera: Scarabaeidae). ZooKeys, 772, 97–128. https://doi.org/10.3897/zookeys.772.25320
- Sreedevi, K., Ranasinghe, S., Fabrizi, S. & Ahrens, D. (2019) New species and records of Sericini scarab beetles from the Indian subcontinent (Coleoptera, Scarabaeidae) II. European Journal of Taxonomy, 567, 1–26. https://doi.org/10.5852/ejt.2019.567
- Su, X., Li, W., Liu, W., Ahrens, D., Zheng, Y. & Liu, Y. (2023) An update to the taxonomy of Serica MacLeay, 1819 (sensu lato) from China (Coleoptera: Scarabaeidae: Sericinae: Sericini). ZooKeys, 1185, 181–198. https://doi.org/10.3897/zookeys.1185.109154
- Zhao, M.-Z. & Ahrens, D. (2023) New species of the tribe Sericini Kirby, 1837 from China, with further updates on their taxonomy and distribution (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 5361 (2), 181–220. https://doi.org/10.11646/zootaxa.5361.2.3