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Diplopoda, Polydesmida, Paradoxosomatidae) ZooKeys 1040: 167–185 (2021) A peer-reviewed open-access journal doi: 10.3897/zookeys.1040.66209 RESEARCH ARTICLE https://zookeys.pensoft.net Launched to accelerate biodiversity research Two new species of the millipede genus Tylopus Jeekel, 1968 from Shan State, Myanmar (Diplopoda, Polydesmida, Paradoxosomatidae) Natdanai Likhitrakarn1,2, Sergei I. Golovatch3, Ruttapon Srisonchai4, Somsak Panha5,6 1 Division of Plant Protection, Faculty of Agricultural Production, Maejo University, Chiang Mai, 50290, Thailand2 Biodiversity and Utilization Research Center of Maejo University, Maejo University, Chiang Mai, 50290, Thailand 3 Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow 119071, Russia 4 Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen, 40002, Thailand 5 Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalong- korn University, Bangkok, 10330, Thailand 6 Academy of Science, The Royal Society of Thailand, Bangkok 10300, Thailand Corresponding author: Somsak Panha ([email protected]) Academic editor: Dragan Antić | Received 20 March 2021 | Accepted 26 April 2021 | Published 28 May 2021 http://zoobank.org/910903D2-476A-439F-AC73-292D5B9A18C1 Citation: Likhitrakarn N, Golovatch SI, Srisonchai R, Panha S (2021) Two new species of the millipede genus Tylopus Jeekel, 1968 from Shan State, Myanmar (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 1040: 167–185. https://doi.org/10.3897/zookeys.1040.66209 Abstract The predominantly Indochinese to southern Chinese millipede genusTylopus presently comprises 76 described species, including two new, T. monticola sp. nov. and T. sutchariti sp. nov., both described and illustrated based on material from a limestone mountain in Taunggyi District, Shan State, Myanmar. Both new species have been found to occur syntopically near limestone caves and are assumed to be narrowly endemic to the Taunggyi Mountains, southwestern Shan State, Myanmar. A key to all six Tylopus species known to occur in Myanmar is provided, and their distributions are also mapped. Keywords Key, map, Paradoxosomatinae, Sulciferini, taxonomy, Tylopus monticola sp. nov., Tylopus sutchariti sp. nov. Copyright Natdanai Likhitrakarn et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 168 Natdanai Likhitrakarn et al. / ZooKeys 1040: 167–185 (2021) Introduction The predominantly Indochinese to southern Chinese millipede genus Tylopus Jeekel, 1968, has long been recognized as one of the most speciose and widespread not only within the family Paradoxosomatidae, but also in the entire class Diplopoda (Likhi- trakarn et al. 2010, 2016; Golovatch 2019). This genus, formerly known asAgnesia Attems, 1953, has been reviewed and rediagnosed several times (Jeekel 1965, 1968; Golovatch and Enghoff 1993; Likhitrakarn et al. 2010), but most taxonomic works have focused on adding new species descriptions, presenting a key and a distribution map to reveal the high diversity of the genus (Nguyen 2012; Liu and Luo 2013, Golo- vatch 2013, 2014, 2018, 2019, 2020; Likhitrakarn et al. 2014, 2016). At the moment, 74 species of Tylopus are known from Indochina and the adjacent parts of southern China and Myanmar (formerly Burma). Most of the known species diversity of Tylopus is encountered in Thailand and Vietnam. Myanmar forms part of the Indo-Burma biodiversity hotspot (Myers et al. 2000; Sodhi et al. 2004). It supports extremely high biodiversity and abundant natural re- sources, including millipedes (Diplopoda). At present, Myanmar’s known millipede diversity has gradually been revealed to amount to 96 species from 36 genera, 13 families and eight orders, containing 74 endemic and only five widespread synan- thropic species (Likhitrakarn et al. 2017, 2018; Pimvichai et al. 2018; Srisonchai et al. 2018a, b). Furthermore, there are 527 millipede records from Burmese amber (Bur- mite; Cretaceous, ca 100 Mya), representing 13 of the 16 extant orders. Only the orders Sphaerotheriida, Julida and Siphonocryptida have not yet been reported from Burmite. Against this background, no fossil of the family Paradoxosomatidae, one of the largest and most diverse in the entire class Diplopoda globally, has previously been recorded from Myanmar (Wesener and Moritz 2018). Four Tylopus species, all endemic, have been found in Myanmar. The first two spe- cies of Tylopus to be revealed from that country were Tylopus doriae (Pocock, 1895) and T. silvestris (Pocock, 1895), both described by Pocock (1895). It was 120+ years later that two further species were added: T. brehieri Golovatch, VandenSpiegel & Seme- nyuk, 2016 and T. punctus Likhitrakarn, Golovatch & Panha, 2016 (Golovatch et al. 2016; Likhitrakarn et al. 2016). Myanmar’s climate, geology, topography and, partly, its biota are very similar to those of the neighbouring Thailand; consequently theTylo - pus species diversity in Myanmar is surprisingly low compared to Thailand with its 31 species. This is undoubtedly due to many areas of Myanmar still being difficult to- ac cess, remaining poorly collected and often even dangerous, coupled with local natural history research being rudimentary and secluded. Hardly surprisingly, the arthropod fauna of Myanmar is poorly known and understudied. Studies on the millipede diver- sity of Myanmar have recently resumed since the British colonial times, chiefly due to the activities of the Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok, Thailand, headed by one of us (SP). The present paper puts on record two new species of Tylopus collected from a lime- stone mountain in the Taunggyi District, southwestern Shan State, Myanmar. A key Two new Tylopus from Myanmar 169 to and updated distributions of all six species of Tylopus currently known to occur in Myanmar are also provided. Materials and methods New material was collected in Myanmar, especially in limestone mountain areas, with the support of Fauna & Flora International (FFI) in 2015–2017, collaborating with the Animal Systematics Research Unit (ASRU), Chulalongkorn University. The col- lecting activities took place under the limestone conservation projects which aim to protect biodiversity in limestone habitats (Grismer et al. 2018a, b, c; Fauna & Flora International 2021). Live animals were photographed in the laboratory using a Nikon 700D digital camera with a Nikon AF-S VR 105 mm macro lens. Specimens were preserved in 75% ethanol, and morphological observations were carried out in the laboratory using an Olympus stereo microscope. Scanning electron micrographs (SEM) of gonopods coat- ed with gold were taken using a JEOL, JSM–5410 LV microscope, returned to alcohol after SEM examination. Digital images of the specimens were taken in the laboratory and assembled using the “CellD” automontage software of the Olympus Soft Imaging Solution GmbH package. In addition, line drawings of gonopod characters were also prepared. Both holotypes, as well as most of the paratypes are housed in the Museum of Zoology, Chulalongkorn University (CUMZ), Bangkok, Thailand; some paratypes are donated to the collection of the Zoological Museum, State University of Moscow (ZMUM), Russia, as indicated in the text. Collecting sites were located by GPS using the WGS84 datum. The distribution maps of all Tylopus species recorded from Myanmar were executed using QGIS 3.18.0 (QGIS Development Team 2021). Google satellite maps were downloaded via the QuickMapServices plugin. The images were enhanced and arranged in plates with Adobe Photoshop CS6 software. In the synonymy sections, D stands for the original description and/or subsequent descriptive notes, K for the appearance in a key, L for the appearance in a species list, and M for a mention. Terminology concerning gonopodal and somatic structures mostly follows Golovatch and Enghoff (1993) and Likhitrakarn et al. (2010, 2016). Abbreviations of certain gonopodal structures in the figures are explained both in the text and figure captions. Taxonomy Family Paradoxosomatidae Daday, 1889 Subfamily Paradoxosomatinae Daday, 1889 Tribe Sulciferini Attems, 1898 170 Natdanai Likhitrakarn et al. / ZooKeys 1040: 167–185 (2021) Genus Tylopus Jeekel, 1968 Tylopus brehieri Golovatch, VandenSpiegel & Semenyuk, 2016 Tylopus brehieri Golovatch, VandenSpiegel & Semenyuk, 2016: 335 (D). Record from Myanmar. Shan State, Kyauk Khaung (= Stone Cave) (Golovatch et al. 2016). Tylopus doriae (Pocock, 1895) Orthomorpha doriae Pocock, 1895: 823 (D). Orthomorpha Doriae (sic!) – Attems, 1898: 339 (L, K). Orthomorpha (Kalorthomorpha) doriae – Attems, 1936: 204 (L). Orthomorpha (Orthomorpha) doriae – Attems, 1937: 80 (D, K). Orthomorpha doriae – Weidner, 1960: 85 (L). Agnesia doriae – Jeekel, 1965: 100 (D, K). Tylopus doriae – Jeekel, 1968: 60 (M); Golovatch and Enghoff 1993: 103 (D, K); Eng- hoff 2005: 99 (R); Likhitrakarn et al. 2010: 25 (L, K); 2014: 65 (L, K); 2016: 35 (L, K); Nguyen and Sierwald 2013: 1298 (L). Records from Myanmar. Yado, 1000–1400 m; Bia-Po, 1000–1200 m, Meteleo, 900– 1200; Puepoli, 900–1200 m (Pocock 1895). Remark. Also found in Doi Suthep National Park (1400–1500 m), Chiang Mai Province, Thailand (Enghoff 2005). Tylopus punctus Likhitrakarn, Golovatch & Panha, 2016 Tylopus punctus Likhitrakarn, Golovatch & Panha, 2016:
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