Zoosyst. Evol. 100 (2) 2024, 405-423 | DOI 10.3897/zse.100.119297 yee BERLIN Taxonomic re-appraisal of Scolopocryptops quadristriatus (Verhoeff, 1934) and a description of a new species from Japan and Tatwan (Chilopoda, Scolopendromorpha, Scolopocryptopidae) Taro Jonishi!, Takafumi Nakano! 1 Department of Zoology, Graduate School of Science, Kyoto University, Kyoto 606-8502, Japan https://zoobank. org/6B07BF 7A-9A DA-4959-A654-59E747E200DE Corresponding author: Taro Jonishi (ykn347635@gmail.com) Academic editor: Martin Husemann @ Received 23 January 2024 # Accepted 14 March 2024 # Published 9 April 2024 Abstract Centipedes of the genus Scolopocryptops Newport, 1844 are blind species mostly described from the New World and East Asia. In this study, a Japanese species, S. quadristriatus (Verhoeff, 1934), which is characterised by four longitudinal keels on the tergites, is re-described, based on the likely holotype preserved in the Zoologische Staatssammlung Munchen and specimens newly collected from near the type locality. In addition, S. /ongisetosus sp. nov., a new species that bears tergal keels like S. quadristriatus, is de- scribed from the Ryukyu Islands in Japan and Taiwan. Although the presence of four keels on tergites is unique to these two species, phylogenetic analyses using nuclear and mitochondrial markers showed that S. /ongisetosus sp. nov. is not sister to S. quadristriatus. The obtained phylogeny indicates that the tergal longitudinal keels evolved in parallel within Scolopocryptops or that the presence of keels represents a plesiomorphic character of the clade containing these species. Key Words molecular phylogeny, nomenclature, plesiomorphic character, Ryukyu Islands, tergal keels Introduction Scolopocryptops Newport, 1844 is a centipede genus that mostly inhabits epigean habitats in North and South America, West Africa, East Asia and Vietnam and species have been also recorded from India, the Philippines, In- donesia, New Guinea and Fiji (Chagas-Jr et al. 2023; Le et al. 2023). The genus currently comprises 33 species and subspecies, which are classified into two groups: the Asian/North American group and the Neotropical/Afro- tropical group (Chagas-Jr 2008; Edgecombe et al. 2012: Vahtera et al. 2013; Chagas-Jr et al. 2023; Le et al. 2023; Jonishi and Nakano 2023). Recent studies showed that the Asian/North American group consists of two lineag- es: a lineage composed of S. elegans (Takakuwa, 1937) and its closest relatives (hereinafter the e/egans lineage), which is known from Far East Asia and a lineage includ- ing the rest of the Asian and North American species (hereinafter, the ex-e/egans lineage) (Jonishi and Nakano 2022 52023). Scolopocryptops centipedes are characterised by a dis- tinctive kind of collared antennal setae, the absence of eyes, 23 leg-bearing segments (except for S. sukiuyan Cha- gas-Jr, Edgecombe & Minelli, 2023, which has 25 seg- ments) and an ultimate leg prefemur with one dorso-me- dial and/or one ventral spinous process (Koch et al. 2010; Chagas-Jr et al. 2023; Le et al. 2023). Several Asian/North American species, for example, S. sexspinosus (Say, 1821) and S. spinicaudus Wood, 1862, are distinguished from others by the absence of complete paramedian sutures on tergites (Le et al. 2023). Amongst them, the Japanese S. quadristriatus (Verhoeff, 1934) bears four longitudinal Copyright Jonishi, T. & Nakano, T. 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. 406 Jonishi, T. & Nakano, T.: Scolopocryptops quadristriatus and new species from Japan and Taiwan keels on its tergites, which is a specific character that dis- tinguishes this species from all other Scolopocryptops (Verhoeff 1934; Takakuwa 1940; Shinohara 1984; Le et al. 2023). After it was described as Otocryptops sexspi- nosus quadristriatus, based on a specimen from the vi- cinity of Tokyo, S. quadristriatus has been recorded from several localities in Honshu, the Izu Islands and Kyushu in the Japanese Archipelago (Shinohara 1949; Takashima and Shinohara 1952; Miyosi 1953; Takano 1973). Phylo- genetic analyses indicated that this species belongs to the ex-elegans \ineage (Jonishi and Nakano 2022). As sum- marised by Le et al. (2023), however, the taxonomic status of S. quadristriatus remains uncertain because only a low resolution of morphological features has been reported (Verhoeff 1934; Takakuwa 1939, 1940; Shinohara 1984). In this study, a re-description of S. quadristriatus, based on the likely holotype is provided and newly-ob- tained specimens from near the type locality and adya- cent areas were also investigated. In addition, several un- identified Scolopocryptops specimens obtained from the Ryukyu Islands in southern Japan and Taiwan have been studied. These individuals are morphologically similar to S. quadristriatus, but differ in several characters. Based on both morphological examination and phylogenetic Ishigaki Isl. . Yonaguni Isl. analyses using nuclear and mitochondrial markers, these specimens are described as a new species herein. Methods Specimen collection and morphological observation A specimen labelled as “Otocryptops sexspinosus quadristriatus” deposited at the Zoologische Staatssam- mlung Munchen (ZSM A20051244), which is likely to represent the holotype of this taxon, was examined. Ad- ditional specimens of S. quadristriatus were collected from several localities in Tokyo and adjacent areas in eastern Honshu, Japan (Fig. 1). A total of 21 uniden- tified Scolopocryptops individuals were also collected from Okinawa, Ishigaki and Yonaguni Islands in the Ryukyu Islands and Hsinchu and Nantou Counties in Taiwan (Fig. 1). Specimens were placed in 30% ethanol for several minutes, fixed in 80% or 99% ethanol and then preserved in 75% ethanol. Ultimate leg-bearing segments of several specimens were dissected to exam- ine genital segments. 140°E Shikoku Figure 1. Collection localities of Scolopocryptops quadristriatus (Verhoeff, 1934) and S. /ongisetosus sp. nov. in the present study. Purple circles: S. quadristriatus; red diamonds: S. /ongisetosus sp. nov.; black diamonds: localities of the sequence data of Taiwan- ese “S. capillipedatus” (= S. longisetosus sp. nov.) obtained from INSD. Locality numbers (Q1—Q3 and L1—L6) are shown in Table 1 and Fig. 13. zse.pensoft.net Zoosyst. Evol. 100 (2) 2024, 405-423 All specimens were observed using a Leica M125C stereoscopic microscope with a drawing tube (Leica Microsystems, Wetzlar, Germany). The specimens were photographed using a Sony a6500 digital camera and a 65 mm macro lens and a Leica MC170 HD digital camera mounted on the Leica M125C. Images captured with the Leica MC170 were processed using Leica Application Suite v. 4.1.2. Specimens examined are deposited in the Zoological Collection of Kyoto University (KUZ). The terminology for external features followed Bonato et al. (2010). For dissected adults and individuals with everted genital organs, genital morphology was exam- ined following Takakuwa (1933a), Demange and Richard (1969), Iorio (2003), Siriwut et al. (2016) and Jonishi and Nakano (2022, 2023). DNA extraction, PCR and DNA sequencing Total DNA was extracted using a NucleoSpin Tissue kit (Macherey-Nagel, Duren, Germany). Following previ- ous studies (e.g. Edgecombe et al. (2012, 2019)), nucle- ar internal transcribed spacer 2 (ITS2), 28S rRNA (28S), mitochondrial cytochrome c oxidase subunit 1 (COI) and 16S rRNA (16S) markers were selected for phylo- genetic analyses. The primer sets used are as follows: the 5.8SF/28SRev (Murienne et al. 2011) for ITS2, 28Sa/28Sb (Whiting et al. 1997) for 28S, LCO1490/ HCO2198 (Folmer et al. 1994) or the reverse primer HCOoutout (Schwendinger and Giribet 2005) for COI and 16Sar (Xiong and Kocher 1991)/16Sb (Edgecombe et al. 2002) for 16S. For ITS2 and 16S, reactions were performed using a LifeECO Thermal Cycler (Bioer Technology, Hangzhou, China); 28S and COI frag- ments were amplified using a GeneAmp PCR System 407 9700 (Thermo Fisher Scientific, Waltham, USA) and a T-100 Thermal Cycler (Bio-Rad, Hercules, USA), re- spectively. The PCR mixtures were heated to 94 °C for 5 min; followed by 35 cycles at 94 °C for 10 s, differ- ent annealing temperatures for each gene (50-60 °C for ITS2 and 16S, 60 °C for 28S and 50 °C for COI) for 20 s and 72 °C for 48 s for ITS2, 30 s for 28S and 16S and 42—48 s for COI; with a final extension at 72 °C for 6 min. The cycle sequencing reactions and sequencing process followed Jonishi and Nakano (2022, 2023). All sequences obtained in this study were deposited in the International Nucleotide Sequence Databases (INSD) through the DNA Data Bank of Japan (Table 1). Molecular analyses In addition to the 42 sequences obtained in this study, 39 sequences of Japanese and Taiwanese Scolopocryptops species (Jonishi and Nakano 2022, 2023) and 31 sequenc- es of scolopocryptopid species retrieved from the INSD were included in the taxon set (Table 1). The nuclear ITS2 sequences were aligned using MAFFT L-INS-i (Katoh and Standley 2013); the nuclear 28S and the mitochondrial 16S sequences were aligned using MAFFT Q-INS-i (Kuraku et al. 2013; Katoh et al. 2019) considering the RNA secondary structure; non-conserved regions of these genes were trimmed by Gblocks v. 0.91b (Castresana 2000). The mitochondrial COI sequences showed no indels; thus, alignment was straightforward. The first four COI positions were ex- cluded from the analyses because this portion was miss- ing in most of the sequences. The concatenated sequences yielded 2510 bp of aligned positions comprising 780 bp of ITS2, 536 bp of 28S, 654 bp of COI and 540 bp of 16S. Table 1. Samples used for molecular analyses. The information on the voucher is accompanied by the collection locality and the INSD accession numbers of the DNA sequences. Locality numbers are shown in Fig. 1 and Fig.13. Acronyms: AMNH, American Museum of Natural History; KUZ; Zoological Collection of Kyoto University; MCZ, Museum of Comparative Zoology, Harvard University; and SYSU, National Sun Yat-sen University. References: 1, Jonishi and Nakano (2023); 2, Jonishi and Nakano (2022): 3, Chao et al. (unpublished); 4, Vahtera et al. (2013); 5, Vahtera et al. (2012); 6, Edgecombe et al. (2012); 7, Edgecombe and Giribet (2004); 8, Murienne et al. (2010). Sequences with an asterisk (*) were used only in the preliminary analyses. Species Voucher # Locality Locality # INSD # References ITS2 28S Col 16S Asian/North American Scolopocryptops S. quadristriatus (Verhoeff, 1934) KUZ 24083 Hachioji, Tokyo, Japan Ql - LC700508 LC700507 LC792589 This study for 16S; 2 for 28S and COl S. quadristriatus (Verhoeff, 1934) KUZ Z5098 Ome, Tokyo, Japan Q2 LC792567 LC792573 LC792581 LC792590 This study S. quadristriatus (Verhoeff, 1934) KUZ 25105 Minamitsuru, Q3 - LC792574 LC792582 LC792591 This study Yamanashi, Japan S. longisetosus sp. nov. KUZ 25108 Kunigami, Okinawa L1 LC792568 LC792575 LC792583 LC792592 This study Island, Japan S. longisetosus sp. nov. KUZ 25122 ~=Kunigami, Okinawa L2 - LC792576 1LC792584 LC792593 This study Island, Japan S. longisetosus sp. nov. KUZ 25111 Nago, Okinawa L3 LC792569 LC792577 LC792585 LC792594 This study Island, Japan S. longisetosus sp. nov. KUZ 25123 _ Ishigaki Island, Japan L4 LC792570 LC792578 LC792586 LC792595 This study S. longisetosus sp. nov. KUZ 25124 Yonaguni Island, L5 LC792571 LC792579 LC792587 LC792596 This study Japan zse.pensoft.net 408 Jonishi, T. & Nakano, T.: Scolopocryptops quadristriatus and new species from Japan and Taiwan Species Voucher # Locality Locality # INSD # References ITS2 28S Col 16S S. longisetosus sp. nov. KUZ 25127 Nantou, Ren’ai, L6 LC792572 LC792580 LC792588 LC792597 This study Taiwan “S. Capillipedatus (Takakuwa, 1938)” SYSU Yanping, Taitung, - ~ - AB617528* - 3 (= S. longisetosus sp. nov.) Chilo-044 Taiwan “S. capillipedatus (Takakuwa, 1938)” SYSU Heping, Taichung, - - - AB617529* - 3 (= S. longisetosus sp. nov.) Chilo-056 Taiwan “S. Capillipedatus (Takakuwa, 1938)” SYSU Datong, Yilan, Taiwan - - - AB617530* - 3 (= S. longisetosus sp. nov.) Chilo-061 “S. Capillipedatus (Takakuwa, 1938)” SYSU Taoyuan, Kaohsiung, - - - AB672646* - 3 (= S. longisetosus sp. nov.) Chilo-143 Taiwan S. musashiensis Shinohara, 1984 KUZ Z4085 Ichikawa, Chiba, - - LC700512 LC700511 LC792598 This study for 16S; Japan 2 for 28S and COl S. nigridius McNeil, 1887 MCZ North Carolina, USA - - HM453278 AY288744 AY288725 — 8 for 28S; 7 for DNA100807 COl and 16S S. nigridius McNeil, 1887 MCZ IZ- North Carolina, USA - - JX422594 JX422680 JX422704 6 130806 “S. nipponicus” Shinohara, 1990 MCZ IZ- Nagoya, Aichi, Japan - - JX422595 = JX422681 JX422705 6 sensu Edgecombe et al. (2012) 130804 S. ogawai Shinohara, 1984 KUZ 24395 —- Fukuroi, Shizuoka, - LC741599 LC741600 LC741601 - if Japan S. rubiginosus L. Koch, 1878 KUZ Z4082 Enoshima, Kanagawa, ~ LC741602 LC700506 LC700505 LC792599 This study for 16S; Japan 1 for ITS2; 2 for 288 and COl S. rubiginosus L. Koch, 1878 MCZ IZ- Yuanshan, Yilan, - - - JX422682 JX422706 6 130823 Taiwan S. sexspinosus (Say, 1821) MCZ IZ- North Carolina, USA - - AY288710 AY288745 AY288726 7 131450 S. spinicaudus Wood, 1862 AMNH IZC California, USA - - JX422596 JX422683 JX422707 6 00146514 S. elegans (Takakuwa, 1937) KUZ Z4062 ~Katsurahama, Kochi, - LC741566 LC700494 LC700493 LC792600 This study for 16S; Japan 1 for ITS2; 2 for 288 and COl S. elegans (Takakuwa, 1937) KUZ 24073 Higashimuro, - LC741569 LC700500 LC700499 LC792601 This study for 16S; Wakayama, Japan 1 for ITS2; 2 for 288 and COl S. elegans (Takakuwa, 1937) KUZ Z4373 Akiruno, Tokyo, Japan - LC741570 LC741571 LC741572 LC792602 This study for 16S; 1 for ITS2, 28S, and COl S. miyosii Jonishi & Nakano, 2023 KUZ 24375 Kirishima, - LC741573 =LC741574 ~LC741575 LC792603 This study for 16S; Kagoshima, Japan 1 for ITS2, 28S, and COl S. miyosii Jonishi & Nakano, 2023 KUZ Z4374 — Saeki, Oita, Japan - LC741578 LC741579 LC741580 LC792604 This study for 16S; 1 for ITS2, 28S, and COl S. miyosii Jonishi & Nakano, 2023 KUZ Z4380 Yamato-son, Amami - LC741581 LC741582 LC741583 LC792605 This study for 16S; Island, Japan 1 for ITS2, 28S, and COl S. brevisulcatus KUZ 24389 Mt. Katsuu-dake, - — LC741587 + LC741588 LC792606 This study for 16S; Jonishi & Nakano, 2023 Okinawa Island, Japan 1 for 28S and COl S. brevisulcatus KUZ 24392 Mt. Fuenchiji, Okinawa - LC741589 LC741590 LC741591 LC792607 This study for 16S; Jonishi & Nakano, 2023 Island, Japan 1 for ITS2, 28S, and COl S. curtus (Takakuwa, 1939) KUZ Z4079 Tai'an, Miaoli, Taiwan - LC741597 LC700502 LC700501 - 1 for ITS2; 2 for 28S and COl S. curtus (Takakuwa, 1939) KUZ 24081 lriomote Island, - LC741598 LC700504 LC700503 LC792608 This study for 16S; Okinawa, Japan 1 for ITS2; 2 for 28S and COl Neotropical/Afrotropical Scolopocryptops “S. macrodon’” (Kraepelin, 1903) MCZ IZ- Guyana — - - JX422675 JX422699 6 sensu Chagas-Jr (2008) and 130814 Edgecombe et al. (2012) S. miersii Newport, 1845 MCZ IZ- Brazil - - KF676364 JX422674 JX422697 6 for COl and 16S; 130729 4 for 28S S. miersii Newport, 1845 AMNH French Guiana - - - HQ402545 = JX422698 5 for COl; 6 for LP3868, 16S IZ-130730 Outgroup Newportia monticola Pocock, 1890 MCZ IZ- Parque de Cahuita, - - KF676360 KF676507 HQ402497 5 for 16S; 4 for 130777 Costa Rica 28S and COl zse.pensoft.net Zoosyst. Evol. 100 (2) 2024, 405-423 Maximum Likelihood (ML) and Bayesian Inference (BI) were applied to two separate datasets: a COI sequence dataset for preliminary analyses and a concatenated data- set of COI, 16S, ITS2 and 28S. The sequences of “S. cap- illipedatus (Takakuwa, 1938)” (see Results), of which only the COI region is available in the INSD (AB617528— AB617530, and AB672646), were used only in the pre- liminary analyses. The best-fit partition schemes and substitution models were identified, based on the Bayes- ian Information Criterion using ModelFinder (Kalyaana- moorthy et al. 2017) implemented in IQ-TREE v.1.6.12 (Nguyen et al. 2015) or PartitionFinder v.2.1.1 (Lanfear et al. 2016) with the “all” algorithm. The selected parti- tion schemes and models are shown in Suppl. material 1. The ML phylogenetic trees were reconstructed using IQ- TREE v.1.6.12 with ultrafast bootstrapping (UFBoot; Ho- ang et al. (2017)) conducted with 1000 replicates. The BI trees and Bayesian posterior probabilities (BPP) were es- timated using MrBayes v.3.2.7 (Ronquist et al. 2012). Two independent runs of four Markov chains were conducted for 10 million generations and the tree was sampled every 100 generations. Considering parameter estimates and as- sessments of convergence using Tracer v.1.7.2 (Rambaut et al. 2018), the first 2500 trees were discarded as burn-in. Uncorrected pairwise distances for COI sequences (590-654 bp) were calculated with MEGA 11 (Tamu- ra et al. 2021), with pairwise deletion of missing data (Suppl. material 2). Results Taxonomy Order Scolopendromorpha Pocock, 1895 Family Scolopocryptopidae Pocock, 1896 Genus Scolopocryptops Newport, 1844 Scolopocryptops quadristriatus (Verhoeff, 1934) Figs 2-7 Japanese name: Yosuji-akamukade Otocryptops sexspinosus quadristriatus Verhoeff, 1934: 54; Takakuwa (1939: 699), fig. 3 (as Octocryptops [sic] sexspinosus quadristria- tus), Takakuwa (1940: 73), fig. 77. Scolopocryptops quadristriatus. Shinohara (1984: 41); Shinohara et al. (2015: 880, 906); Le et al. (2023: 437, 442). Not S. guadristriatus. “Otocryptops sexspinosus quadristriatus”: Taka- kuwa (1933b: 1457, 1459) (nomen nudum). Type specimen. Ofocryptops s. quadristriatus was de- scribed, based on a single specimen from the vicinity of Tokyo, without any information on the collector, col- lecting date or deposition of the specimen (Verhoeff 1934). Most of the Verhoeff’s specimens are now kept at the Zoologische Staatssammlung Munchen (ZSM), the Museum fiir Naturkunde (MfN) and the Naturhis- torischen Museums Wien (NHMW) (Melzer et al. 2011). 409 A specimen of O. s. quadristriatus is deposited at the ZSM, but neither MfN nor NHMW have specimens labelled as O. s. quadristriatus (Moritz and Fischer 1979; Schileyko and Stag] 2004). ZSM A20051244 (Fig. 2) is, thus, the only specimen of O. s. qguadristriatus remaining within the Verhoeff’s collection and is supposed to be the holo- type of this nominal taxon, although the original label and collection data of this specimen are unavailable (Stefan Friedrich, personal communication). Its cephalic capsule, maxillae and left ultimate leg had been preserved in a sepa- rate vial (Fig. 2A); the body had been cut into two parts on leg-bearing segment 8 (Fig. 2B); its ultimate leg-bearing segment had been dissected. Additionally, its left legs 6, 8, 12, 14, 15, 17, 18, 20 and 22; right legs 3, 8, 12-18 and 20- 22; and right ultimate leg had been lost or loosened; its left leg 16 was loosened during observation by the first author. Morphological features of the likely holotype are consis- tent with the original description and specimens newly obtained in this study. Thus, a description, based on both ZSM A20051244 and our specimens, is provided below. Shinohara (1982) stated that a Japanese myriapodolo- gist, the late Dr Yoshioki Takakuwa, sent a specimen of O. s. quadristriatus to Verhoeff. Given that Verhoeff also received other chilopod specimens from Takakuwa (e.g. Verhoeff (1934, 1935, 1937)), it is plausible that the likely holotype was also dispatched from Takakuwa to Verhoeff. Material examined. Holotype (?): JAPAN * ° (approx. 33.5 mm); ZSM A20051244. Additional material. JAPAN — Tokyo « 1, 33.9 mm (KUZ Z4083); Hachioji City, Uratakao-machi; 15 Apr 2017; Taiga Kato leg. * 1, 33.2 mm (KUZ Z5091); same data as for preceding * 1, 33.0 mm (KUZ Z5092); Ha- chioji City, Mt. Takao-san; 18 Nov 2017; T. Kato leg. « 1, 28.6 mm (KUZ Z5093); Hachioji City, Uratakao-ma- chi; 35°38.69'N, 139°14.36'E; approx. 250 m alt.; 18 Oct 2021: T. Jonishi leg. * 1 3, 32.1 mm (KUZ Z5094) and 3, 33.1 mm (KUZ Z5095), 36.2 mm (KUZ Z5096), 27.7 mm (KUZ Z5097); same locality as for preceding; 35°38.72'N, 139°14.33'E; approx. 260 m alt.; 18 Oct 2021; T. Jonishi leg. ¢ 1, 38.6 mm (KUZ Z5098); Ome City, Futamatao; 35°47.87'N, 139°13.56'E; approx. 230 m alt.; 19 Oct 2021; T. Jonishi leg. « 1 4, 38.2 mm (KUZ Z5099); Ome City, Sawal; 35°48.68'N, 139°10.91'E; approx. 350 m alt.; 19 Oct 2021; T. Jonishi leg. * 2, 33.6 mm (KUZ Z5100), 29.7 mm (KUZ Z5101) and 1 juvenile, 12.8 mm (KUZ Z5102); Akiruno City, Yokosawa; 35°44.28'N, 139°14.28'E; ap- prox. 250 m alt.; 12 Apr 2022; Futaro Okuyama leg. 1, 29.6 mm (KUZ Z5103); Hachioji City, Uratakao-ma- chi; 35°38.75'N, 139°14.16'E; approx. 270 m alt.; 6 Nov 2022; T. Kato leg. — Saitama Prefecture * 1 9, 39.0 mm (KUZ Z5104); Hiki County, Hatoyama-machi; 36°0.01'N, 139°21.42'E; approx. 110 m alt.; 19 Oct 2021; T. Jonishi leg. — Yamanashi Prefecture ¢ 1, 30.5 mm (KUZ Z5105); Minamitsuru. County, Nishikatsura-cho; 35°30.82'N, 138°50.44'E; approx. 670 m alt.; 9 Apr 2022; Eitaro Mat- sushita leg. — Shizuoka Prefecture * 1, 30.4 mm (KUZ Z5106); Susono City, Suyama; 35°15.92'N, 138°47.95'E; approx. 910 m alt.; 2 June 2022; T. Nakano leg. zse.pensoft.net 410 Jonishi, T. & Nakano, T.: Scolopocryptops quadristriatus and new species from Japan and Taiwan Figure 2. Scolopocryptops quadristriatus (Verhoeff, 1934), the likely holotype, 2 (ZSM A20051244). A. Cephalic capsule, maxillae, legs and left ultimate leg; B. Forcipular segment, leg-bearing segments and legs. Scale bars: 5 mm. ws “aa - , fi ; a4 SS SS et yay ag en aes wsas=* Figure 3. Scolopocryptops quadristriatus (Verhoeff, 1934), collected from near the type locality, 2 (KUZ Z5104) and the habitat near the type locality. A. Habitat at Ome City, Tokyo; B. Live specimen, dorsal view. Scale bar: 10 mm. Figure 4. Scolopocryptops quadristriatus (Verhoeff, 1934) from near the type locality (KUZ Z5095). A. Basal articles of left anten- na, dorsal view; B. Cephalic plate, dorsal view. Scale bars: 0.5 mm (A); 1 mm (B). Diagnosis. Antenna with sparse short hairs and setae sutures, tergites 6—20 with four longitudinal keels and on dorsal surface of two basal articles, subsequent articles | median depression bordered by paramedian keels. densely covered with short setae. Cephalic plate with com- Description [variations given in square brackets]. plete lateral marginal sulci. Tergites lacking paramedian Body length approx. 27.7—-39.0 mm in 75% ethanol. zse.pensoft.net Zoosyst. Evol. 100 (2) 2024, 405-423 Tce . - All = a Figure 5. Scolopocryptops quadristriatus (Verhoeff, 1934) from near the type locality (KUZ Z5095). A. Cephalic plate and tergite 1, dorsal view; B. Distal part of article 2, article 3 and pretarsus of left second maxilla, medial view; C. Article 3 and pretarsus of left second maxilla, lateral view; D. Head, ventral view. Abbreviations: am — anterior margin of forcipular coxosternite; bs — basal suture on forcipular trochanteroprefemoral process; db — dorsal brush on article 3 of second maxilla; ds — dorsal spur on article 2 of second maxilla; pt — pretarsus of second maxilla; ptr — process of forcipular trochanteroprefemur. Scale bars: 1 mm (A, D); 0.2 mm (B, C). Colour in life yellowish-brown with dark pigment on two basal antennal articles, purplish on subsequent articles; reddish-brown on forcipules; reddish-brown with dark pigment on anterior, lateral and posterior margins of ce- phalic plate, tergites 1, 22 and 23; purplish dark brown on tergites 2—21; legs and ultimate legs brownish-yellow or orange with bluish dark pigment (Fig. 3B). Colour in ethanol slightly greenish on tergites and legs (Fig. 6A). Antennae 7.7—13.4 mm in length, approx. 0.2—0.35~x as long as body, composed of 17 articles; two basal ar- ticles with sparse short hairs and setae (sensu Bonato et al. (2010)) dorsally, subsequent articles densely covered with short setae (Fig. 4A); each seta emerging from small collar. Cephalic plate as long as wide; its surface sparsely punctate with sparse minute hairs, with complete lateral margination (Figs 4B, 5A). Second maxillae article 2 with elongated and semi-transparent [dark brown] dorsal spur distally; dorsal brush with transparent margination; pretarsus consisting of dark brown basal and semi-transparent short apical parts (Fig. 5B, C). Forcipular coxosternite and trochan- teroprefemora sparsely punctate, coxosternite with me- dian suture and transverse sutures cross median one on anterior third of coxosternite (Fig. 5D); trochanteroprefe- mur with small and blunt black process with basal suture (Fig. 5D); anterior margin of coxosternite strongly scle- rotised and slightly convex, divided into two low lobes by median diastema (Fig. 5D). Tergites sparsely punctate; tergite 1 with anterior transverse suture; anterior margin overlapped by cephal- ic plate (Figs 5A, 6A). All tergites lacking paramedian sutures; tergites 6 [S—7]—21 with longitudinal median de- pression bordered by paramedian keels, lateral keels pres- ent on tergites 6 [6—-8]-20; median depression and keels unapparent on anterior and posterior tergites [depression and keels unapparent on all tergites in adult KUZ Z5099] zse.pensoft.net 412 Jonishi, T. & Nakano, T.: Scolopocryptops quadristriatus and new species from Japan and Taiwan Se Figure 6. Scolopocryptops quadristriatus (Verhoeff, 1934) from near the type locality (KUZ Z5095). A. Cephalic plate and tergites 1-13, dorso-lateral view; B. Tergites 14-16, dorso-lateral view; C. Sternites 8-13, ventral view; D. Leg-bearing segments 3-8, lat- eral view; E. Sternite of ultimate leg-bearing segment and coxopleuron, ventral view; F. Left coxopleuron, lateral view; G. Tergite of ultimate leg-bearing segment, dorso-lateral view. Abbreviations: exp — coxopleural process; dm — dorsal margin of ultimate pleuron; mds — minute dark spine on ultimate pleuron; sp — spiracle; tlk — tergal lateral keel; tm — tergal margination; tpk — tergal paramedian keel. Scale bars: 5 mm (A); 2 mm (B—D); 0.5 mm (E-G). (Fig. 6A, B); tergites 6 [5—7]-21 [22] with complete or nearly complete lateral marginations (Fig. 6A, B); 2 or 3 short longitudinal sulci present on posterior margin of several tergites. Sternites sparsely punctate, lacking paramedian su- tures (Fig. 6C). Sides of sternite of ultimate leg-bearing segment converging posteriorly, posterior margin slightly concave [slightly convex in KUZ Z4083] (Fig. 6E). Ovoid spiracles present on leg-bearing segments 3, 5, 8, 10, 12, 14, 16, 18, 20 and 22 (Fig. 6D). Legs almost lacking setae [sparse minute setae present in several specimens]; tarsi of legs 1-21 undivided; legs 1—20 with lateral and ventral tibial spurs and tarsal spur, zse.pensoft.net leg 21 with tibial spur and tarsal spur; leg 22 with tarsal spur only; all legs with two accessory spines. Coxopleuron approx. 1.5—1.7< as long as sternite of ultimate leg-bearing segment (Fig. 6E, F). Dorsal margin of ultimate pleuron protruding from lateral side of tergite of ultimate leg-bearing segment, dorso-posterior margin with minute dark spine (Fig. 6F, G). Posterior and ventral margins of coxopleuron converging posteriorly, forming approx. 60—75° angle; coxopleural process short, tip of pro- cess pointed, slightly directed dorsally (Fig. 6F). Surface of coxopleuron without setae, covered with various-sized coxal pores (Fig. 6F). Pore-free area present on coxopleural process and dorso-posterior area of coxopleuron (Fig. 6F). Zoosyst. Evol. 100 (2) 2024, 405-423 A 413 Figure 7. Scolopocryptops quadristriatus (Verhoeff, 1934) from near the type locality. A. KUZ Z5095; B. KUZ Z5094; C. KUZ Z5104. A. Left ultimate leg, lateral view; B. Male genital segments, penis and anal valves, ventral view; C. Female genital segment and anal valves, ventral view. Abbreviations: av — anal valve; pn — penis; sgs I — sternite of genital segment 1; sgs II — sternite of genital segment 2; tg — tergite of genital segment. Scale bars: 2 mm (A); 0.5 mm (B, C). Ultimate leg 9.0-12.4 mm in length, approx. 0.3x as long as body; all articles almost lacking setae [tarsi with sparse minute setae]; prefemur with two conical and pointed spinous processes, ventral process larger than dorso-medial one; pretarsus with two accessory spines (Fig. 7A). Genital segments occupying approx. 0.7—0.8 length of sternite of ultimate leg-bearing segment; tergite of gen- ital segment covered with sparse minute setae (Fig. 7B, C). Sternite of genital segment 1 covered with sparse short setae, posterior margin weakly convex (Fig. 7B, C). Sternite of genital segment 2 well developed in male, cov- ered with sparse short setae; posterior part of genital seg- ment 2 overlapped by lamina subanalis, penis not visible in ventral view [penis visible in KUZ Z5094] (Fig. 7B); genital segment 2 not visible in female (Fig. 7C). Anal valves covered with sparse short setae (Fig. 7B, C). Distribution. This species has been recorded from Honshu and the Izu Islands and is abundant in Tokyo and adjacent areas (Takakuwa 1939, 1940; Shinohara 1949; Takashima and Shinohara 1952; Takano 1973). Miyosi (1953) recorded S. quadristriatus from Nagasaki in Ky- ushu, but this species was not obtained during the survey conducted by the first author in Nagasaki and adjacent localities (25—27 July 2023). Availability of “quadristriatus” based on Taka- kuwa’s works. The name “Otocryptops sexspino- sus quadristriatus’, which was attributed to Verhoeff, was introduced by Takakuwa’s two works (Takakuwa 1933a, b) before its formal description by Verhoeff in 1934. However, we herein decide that neither Takakuwa (1933a) nor Takakuwa (1933b) made the species-group name “quadristriatus” available. “Otocryptops sexspi- nosus quadristriatus Verh” first appeared in Takakuwa (1933a: 11), who intended to provide general anatomical features of Scolopocryptops (originally “Otocryptops’’). Nonetheless, the detailed morphological features and figures provided in this work were unambiguously based on S. rubiginosus L. Koch, 1878 (referred as “Otocryp- tops ruliginosus” [sic]). Therefore, the name quadris- triatus in the combination of Otocryptops sexspinosus quadristriatus in Takakuwa (1933a) did not satisfy the provision of Article 13.1 of the International Code of Zo- ological Nomenclature (hereinafter, Code; International Commission on Zoological Nomenclature 1999) and thus is unavailable. In a synopsis of the Japanese centipedes, Takakuwa (1933b: 1459) provided a brief taxonomic account and morphological descriptions of the subspecies referred as “Otocryptops sexspinosus quadristriatus VERHOEFF However, Takakuwa considered that “O. s. quadristria- tus” sensu Verhoeff was indistinguishable from the nomi- notypical subspecies “O. s. sexspinosus’’, thus, he did not provide any description or definition that are purported to differentiate “O. s. quadristriatus” (see Article 13.1.1 of the Code). Therefore, we conclude that the species-group name quadristriatus in the combination of Otocryptops sexspinosus quadristriatus in Takakuwa (1933b) is also unavailable and the authorship of this nominal taxon is at- tributed to Verhoeff (1934), who established O. s. guadris- triatus explicitly as a new subspecies. Moreover, ac- cording to Shinohara (1982, 1990), Takakuwa’s (1933b) description of this taxon was based on specimen(s) mis- identified as “O. sexspinosus’”, O. s. quadristriatus sensu Takakuwa (1933b) was later described as S. nipponicus Shinohara, 1990 (placed in synonymy with S. spinicau- dus by Shelley 2002). Remarks. Verhoeff (1934) established this taxon as a subspecies of the North American S. sexspinosus, based on brief taxonomic accounts. He only described the ab- sence of tergal paramedian sutures, the presence of four longitudinal keels on tergites and the colouration of head and leg-bearing segments. Shinohara (1984) elevated guadristriatus to full spe- cies status, based on the following features: 1) cephalic marginal sulci reaching from postero-lateral margin of zse.pensoft.net 414 Jonishi, T. & Nakano, T.: Scolopocryptops quadristriatus and new species from Japan and Taiwan cephalic plate to antennae; 2) tergites without paramedian sutures and with four longitudinal keels; 3) arrangement of tibial and tarsal spurs on legs 19-23; and 4) a slight- ly slender “general form” compared with other species of the genus. It is unclear whether Shinohara compared quadristriatus with the North American O. sexspinosus sexspinosus or the Japanese “O. sexspinosus” (see above) and the characters of 1), 3) and 4) cannot conclusively distinguish quadristriatus from other species. Nonethe- less, the presence of four longitudinal keels on tergites distinguishes S. quadristriatus from all other Scolopoc- ryptops (except S. /ongisetosus sp. nov.; see below). The distinctness of S. guadristriatus 1s also supported by the molecular phylogenetic analyses (Fig. 13). This species is absent from, but should be added to Chilobase 2.0 (Bonato et al. 2016). Scolopocryptops longisetosus sp. nov. https://zoobank.org/3 BCC9F07-141D-4D15-8613-3CE08877COA6 Figs 8-12 Suggested Japanese name: Kuromadara-akamukade Scolopocryptops capillipedatus: Chao and Chang (2003: 4), fig. 17; Chao (2008: 76-80), figs 82-87; Chao and Chang (2008: 14), fig. 9. Material examined. Holotype: JapAN — Okinawa Pre- fecture — Okinawa Island « <, 31.4 mm (KUZ Z5107); Kunigami-son, Uka; 26°48.45'N, 128°15.97'E; approx. 300 m alt.; 4 May 2021; F. Okuyama leg. Paratypes: JAPAN — Okinawa Prefecture — Okinawa Island « 1, 24.6 mm (KUZ Z5108); Kunigami-son, Oku; 26°48.3'N, 128°17.34'E; approx. 230 malt.; 5 May 2021; F. Okuyama leg. ° 1 4, 30.7 mm (KUZ Z5112); Nago City, Mt. Na- go-dake; 26°35.45'N, 128°0.25'E; approx. 200 m alt.; 14 June 2022; T. Jonishi leg. * 1, 29.4 mm (KUZ Z5113),; same locality as for preceding; 26°35.47'N, 128°0.27'E; approx. 210 m alt.; 14 June 2022; T. Jonishi leg. * 1 9, 28.9 mm (KUZ Z5115); same locality as for preceding; 26°35.73'N, 128°0.19'E; approx. 190 m alt.; 14 June 2022; T. Jonishi leg. « 1 3, 31.1 mm (KUZ Z5117); Kuniga- mi-son, Uka; 26°48.43'N, 128°16.04'E; approx. 330 m alt., 15 June 2022; T. Jonishi leg. * 1 4, 37.2 mm (KUZ Z5119); Kunigami-son, Benok1; 26°48.06'N, 128°16.65'E; approx. 350 m alt.; 15 June 2022: T. Jonishi leg. « 1, 30.4 mm (KUZ Z5121); Kunigami-son, Yona, Mt. Fuenchiji; 26°45.15'N, 128°14.58'E; approx. 380 m alt.; 16 June 2022; T. Kato leg. * 1, 28.6 mm (KUZ Z5122); same lo- cality as for preceding; 26°45.17'N, 128°14.58'E; approx. 380 m alt.; 16 June 2022; T. Jonishi leg. Additional material. JAPAN — Okinawa Prefecture — Okinawa Island ¢ 1 <, 30.2 mm (KUZ Z5109); Kuniga- mi-son, Uka; 26°48.30'N, 128°16.11'E; approx. 320 m alt.; A Jan 2022; F. Okuyama leg. * 2 juveniles, approx. 10 mm (KUZ Z5110), 1 9, 24.4mm(KUZ Z5111); Nago City, Mt. Nago-dake; 26°35.45'N, 128°0.25'E; approx. 200 m alt.; 14 June 2022; T. Jonishi leg. * 1, 23.4 mm (KUZ Z5114); same locality as for preceding; 26°35.47'N, 128°0.27'E; zse.pensoft.net approx. 210 m alt.; 14 June 2022; T. Kato leg. * 1 9, 29.4 mm (KUZ Z5116); same locality as for preceding; 26°35.73'N, 128°0.19'E; approx. 190 m alt.; 14 June 2022; T. Kato leg. * 1, 24.5 mm (KUZ Z5118); Kunigami-son, Uka; 26°48 .43'N, 128°16.04'E; approx. 330 m alt.; 15 June 2022; T. Jonishi leg. * 1, 27.2 mm (KUZ Z5120); Kuniga- mi-son, Benok1; 26°48.04'N, 128°16.62'E; approx. 360 m alt.; T. Jonishi leg. — Ishigaki Island « 1, 19.9 mm (KUZ Z5123); Hirakubo; 24°35.01'N, 124°20.15'E; approx. 20 m alt.; 17 Dec 2021; F. Okuyama leg. — Yonaguni Island °2 4, 27.8 mm (KUZ Z5124), 20.8 mm (KUZ Z5125); Mantabaru Forest Park; 24°27.39'N, 122°58.56'E; approx. 100 m alt.; 16 Apr 2022; Naoto Sawada leg. Tarwan ° 1, 21.5 mm (KUZ Z5126); Hsinchu County, Wufeng Town- ship, Shei-Pa National Park; 24°30.01'N, 121°4.61'E; 19 Mar 2019; T. Nakano leg. * 1, 22.2 mm (KUZ Z5127),; Nantou County, Ren’ai Township; 24°5.1'N, 121°10.73'E; 21 Mar 2019; T. Nakano leg. Type locality. Japan, Okinawa Prefecture, Okinawa Island, Kunigami-son, Uka (26°48.45'N, 128°15.97'E, approx. 300 m alt.). Diagnosis. Antenna with sparse hairs and setae of var- ious lengths dorsally on two basal articles, subsequent articles densely covered with long setae and minute se- tae. Cephalic plate with complete lateral marginal sulci. Tergites lacking paramedian sutures, tergites 5-20 with four longitudinal keels and median depression bordered by paramedian keels. Description of holotype [data from other speci- mens given in square brackets]. Body length approx. 31.4 mm [19.9-37.2 mm] in 75% ethanol. Colour in life and in ethanol yellowish-brown with dark pigment on two basal antennal articles, purplish on subsequent arti- cles; reddish-brown on forcipules; reddish-brown with dark pigment on lateral and posterior margins of cephalic plate, tergites 1, 22 and 23; brown with dark pigment on tergites 2-21; legs and ultimate legs brownish-yellow or orange with purplish dark pigment (Figs 8B, 10A, 11A). Antennae 10.1 mm in length, approx. 0.3 as long as body, composed of 17 articles; dorsal surface of two basal articles with sparse hairs and setae (sensu Bonato et al. (2010)) of various lengths, subsequent articles densely covered with long setae and minute short setae [in small specimens, most setae from article 3 shorter than those of two basal articles] (Fig. 9A); setae emerging from vari- ous-sized collars. Cephalic plate as long as wide; its sur- face sparsely punctate with minute hairs, with complete lateral margination (Figs 9B, 10A). Second maxillae article 2 with elongated and semi-transparent dorsal spur distally; dorsal brush with transparent margin; pretarsus consisting of dark brown basal and semi-transparent short apical parts (Fig. 10C, D). Forcipular coxosternite and trochanteroprefemur sparsely punctate, coxosternite with transverse sutures on anterior third of coxosternite; trochanteroprefemur with small and blunt black process and basal suture; anterior margin of coxosternite strongly sclerotised and slightly convex, di- vided into two low lobes by median diastema (Fig. 10E). Zoosyst. Evol. 100 (2) 2024, 405-423 415 Figure 8. Scolopocryptops longisetosus sp. nov., paratype, 3 (KUZ Z5119: B), non-type specimen from Yonaguni Island, 3’ (KUZ Z5124: C) and habitat near the type locality. A. Habitat (laurel tree forest) on Okinawa Island; B, C. Live specimen, dorsal view. Scale bars: 10 mm. Figure 9. Scolopocryptops longisetosus sp. nov., holotype, 3 (KUZ Z5107). A. Basal articles of left antenna, dorsal view; B. Cephalic plate, dorsal view. Scale bars: 0.5 mm (A); 1 mm (B). Tergites sparsely punctate [sparse minute setae pres- ent in small individuals]; tergite 1 with anterior trans- verse suture, anterior margin overlapped by cephalic plate (Figs 10A, 11A). All tergites lacking paramedian sutures; tergites 5 [4]—21 [20] with longitudinal medi- an depression bordered by paramedian keels, lateral keels present on tergites 5 [5—8]—20 [18-20]; median depression and keels unapparent on anterior and pos- terior tergites [depression and keels unapparent on all tergites in KUZ Z5119, Z5126 and Z5127] (Fig. 11A, B); lateral marginations complete or nearly complete on tergites 7 [5—7]—21 [22] (Fig. 11A, B); three short longitudinal sulci present on posterior margin of terg- ites 4, 6 [2 or 3 sulci present on tergites 3—21; absent in several specimens]. Sternites sparsely punctate, lacking paramedian su- tures (Fig. 11C). Sides of sternite of ultimate leg-bearing segment converging posteriorly, posterior margin almost straight [slightly concave] (Fig. 11E). Ovoid spiracles present on leg-bearing segments 3, 5, 8, 10, 12, 14, 16, 18, 20 and 22 (Fig. 11D). Legs on anterior leg-bearing segments with sparse minute setae [setae denser in small individuals], posterior legs almost lacking setae [all legs setose in KUZ Z5123]; tarsi of legs 1-21 undivided; legs 1-19 with lateral and ventral tibial spurs and tarsal spur, legs 20 and 21, re- spectively, with tibial spur and tarsal spur; leg 22 without spurs. All legs with two accessory spines. Coxopleuron approx. 1.8x [1.5—-1.8x] as long as sternite of ultimate leg-bearing segment (Fig. 11F). zse.pensoft.net 416 Jonishi, T. & Nakano, T.: Scolopocryptops quadristriatus and new species from Japan and Taiwan pt~ ai Figure 10. Scolopocryptops longisetosus sp. nov., holotype, 6 (KUZ Z5107: A, C-E) and non-type specimen from Yonaguni Is- land, 3 (KUZ Z5124: B). A, B. Cephalic plate and tergite 1, dorsal view; C. Distal part of article 2, article 3 and pretarsus of left second maxilla, medial view; D. Article 3 and pretarsus of left second maxilla, lateral view; E. Head, ventral view. Abbreviations: am — anterior margin of forcipular coxosternite; bs — basal suture on forcipular trochanteroprefemoral process; db — dorsal brush on article 3 of second maxilla; ds — dorsal spur on article 2 of second maxilla; pt — pretarsus of second maxilla; ptr — process of forcipular trochanteroprefemur. Scale bars: 1 mm (A, B, E); 0.2 mm (C, D). Dorsal margin of ultimate pleuron protruding from lat- eral margin of tergite of ultimate leg-bearing segment, dorso-posterior margin with minute dark spine (Fig. 11F, G). Posterior and ventral margins of coxopleuron con- verging posteriorly, forming approx. 60° [60—-65°] angle; coxopleural process short, tip of process pointed, slight- ly directed dorsally (Fig. 11F). Surface of coxopleuron without setae, covered with various-sized coxal pores zse.pensoft.net (Fig. 11F). Pore-free area present on coxopleural process and dorso-posterior region of coxopleuron (Fig. 11F). Ultimate leg 10.8 mm in length, approx. 0.3x [0.3— 0.37x] as long as body; prefemur, femur and tibia almost lacking setae, tarsi with sparse minute setae [tarsi almost lacking setae]; prefemur with two conical and pointed spinous processes, ventral process larger than dorso-me- dial one; pretarsus with two accessory spines (Fig. 12A). Zoosyst. Evol. 100 (2) 2024, 405-423 =e = ‘ . : i ' - mds Pe Figure 11. Scolopocryptops longisetosus sp. nov., holotype, 6 (KUZ Z5107). A. Cephalic plate and tergites 1-12, dorso-lateral view; B. Tergites 10—12, dorso-lateral view; C. Sternites 6-12, ventral view; D. Leg-bearing segments 3-8, lateral view; E. Sternite of ultimate leg-bearing segment, ventral view; F. Left coxopleuron, lateral view; G. Tergite of ultimate leg-bearing segment, dor- so-lateral view. Abbreviations: exp — coxopleural process; dm — dorsal margin of ultimate pleuron; mds — minute dark spine on ultimate pleuron; sp — spiracle; tlk — tergal lateral keel; tm — tergal margination; tpk — tergal paramedian keel. Scale bars: 5 mm (A); 2 mm (B—D); 0.5 mm (E-G). Genital segments occupying approx. 0.8 length of sternite of ultimate leg-bearing segment; tergite of genital segment sparsely setose (Fig. 12C, D). Sternite of genital segment | sparsely covered with setae, posterior margin slightly convex (Fig. 12C, D). Sternite of genital segment 2 well developed, covered with sparse setae; penis visi- ble in ventral view; anal valves covered with sparse setae [in female, genital segment 1 as described for holotype: genital segment 2 not visible; anal valves as described for holotype] (Fig. 12C—E). Variation. In specimens from the southern Ryukyus (Ishigaki and Yonaguni Islands; KUZ Z5123-Z5125) and Taiwan (KUZ Z5126, Z5127), dark pigment on ce- phalic plate absent or almost reduced (Figs 8C, 10B); ultimate leg with femur [tibia] and subsequent articles densely setose [setae relatively sparse in KUZ Z5127] (Fig. 12B). Etymology. The specific name is derived from the Latin compound adjective, “longus” (long) and “setosus” (hairy), referring to the long antennal setae of this new species. Distribution. This species is known from Okinawa, Ishigaki and Yonaguni Islands in the Ryukyu Islands, Japan and is also widespread in Taiwan. zse.pensoft.net 418 Jonishi, T. & Nakano, T.: Scolopocryptops quadristriatus and new species from Japan and Taiwan Figure 12. Scolopocryptops longisetosus sp. nov., holotype, @ (KUZ Z5107: A, C, D); non-type specimen from Yonaguni Island, 3S (KUZ Z5124: B); paratype, 9 (KUZ Z5115: E). A, B. Left ultimate leg, lateral view, C. Male genital segments, penis and anal valves, ventral view; D. Male genital segments, penis and anal valves, lateral view; E. Female genital segment and anal valves, ven- tral view. Abbreviations: av — anal valve; pn — penis; sgs I — sternite of genital segment 1; sgs II — sternite of genital segment 2; tg — tergite of genital segment. Scale bars: 2 mm (A, B); 0.5 mm (C—E). Remarks. This species resembles S. quadristria- tus, but S. longisetosus sp. nov. can be distinguished by the presence of long antennal setae (vs. setae short in S. quadristriatus, also see the Identification key provided in the Discussion). The phylogenetic analyses indicate that specimens of this species from Taiwan have been misidentified as S. capillipedatus, based on the dense setae on ultimate legs (Chao and Chang 2003, 2008; Chao 2008). However, S. longisetosus sp. nov. can be distinguished from S. cap- illipedatus by the presence of longitudinal keels and me- dian depression on tergites. Molecular phylogeny and genetic distances. The ML (In L = —14816.13; not shown) and BI (mean In L = —]14846.27; Fig. 13) trees had almost identical topol- ogies. In the Asian/North American group (UFBoot = 100%, BPP = 1.0), both the e/egans lineage (UFBoot = 100%, BPP = 1.0) and the ex-e/egans lineage (UFBoot = 98%, BPP = 1.0) were recovered as monophyletic groups. Within the latter lineage, the recovery of S. quadristriatus (UFBoot = 100%, BPP = 1.0) and S. /ongisetosus sp. nov. (UFBoot = 100%, BPP = 1.0) was strongly supported. Although the interspecific relationships remained largely undetermined, the analyses showed that S. /ongise- tosus Sp. Nov. 1s sister to a clade comprising three of the Japanese nominal species, S. ogawai Shinohara, 1984, S. musashiensis Shinohara, 1984 and “S. nipponicus” sensu zse.pensoft.net Edgecombe et al. (2012) (UFBoot = 99%, BPP = 1.0) (Fig. 13). In the preliminary analyses using the COI data- set, four sequences of “S. capillipedatus” from Taiwan (AB617528—AB617530, AB672646) were nested within S. longisetosus sp. nov. (UFBoot = 98%, BPP = 0.99; Sup- pl. material 3), but the relationships amongst populations were not resolved. In contrast, the concatenated analyses recovered two lineages within S. /ongisetosus sp. nov., which corresponded to the specimens from Okinawa Is- land (UFBoot = 100%, BPP = 1.0) and those from the southern Ryukyu Islands and Taiwan (UFBoot = 98%, BPP = 0.99). The COI pairwise distances within each lin- eage were 2.56—4.74% (Okinawa Island) and 4.89-9.17% (southern Ryukyus and Taiwan; including the sequences of Taiwanese “S. capillipedatus”). The divergence between the two lineages was 5.77—8.87% (Suppl. material 2). Discussion In the obtained phylogenetic trees, S. guadristriatus and S. longisetosus sp. nov. were strongly supported as monophy- letic lineages. The analyses indicated that S. /ongisetosus Sp. nov. comprises two lineages: Okinawa Island and the southern Ryukyus-Taiwan. These lineages differ in two ex- ternal features, 1.e. the presence/absence of dark pigment Zoosyst. Evol. 100 (2) 2024, 405-423 ex-elegans lineage Asian/North American Scolopocryptops * elegans lineage Neotropical/Afrotropical Scolopocryptops x N. monticola 98/1.0 100/0.99 419 IZ-130806 DNA100807 Ds “S. nipponicus” S. musashiensis S. ogawai KUZ Z5108 (L1) KUZ Z5111 (L3) KUZ Z5122 (L2) KUZ 25127 (L6) KUZ 25123 (L4) KUZ Z5124 (L5) IZ-130823 -——_ KUZ 24082 S. spinicaudus s7o.eoy KUZ Z4083 (Q1) * |L KUZ Z5105 (Q3) KUZ Z5098 (Q2) S. sexspinosus * KUZ 24079 KUZ 24081 KUZ 24073 KUZ 24062 ame KUZ 24373 x - KUZ 24392 KUZ 24389 KUZ 24374 / KUZ 24375 93/0.99 KUZ 24380 94/0.98 mies Se. be S. nigridius 98/0.99 99/1.0 4 85/0.50 98/0.99 93/0.99 81/0.92 * S. rubiginosus -/10.65 80/0.78 | S. curtus 65/0.90 97/1.0 97/1.0 S. elegans 70/0.89 | S. brevisulcatus * S. miyosii “S. macrodon” IZ-130730 IZ-130729 S. miersii 0.1 Figure 13. Bayesian Inference tree for 2510 bp-aligned positions of the ITS2, 28S, 16S and COI sequences. Real branch length is shown on the upper left. Numbers on nodes indicate ultrafast bootstrap values (UFBoot) and Bayesian posterior probabilities (BPP). An asterisk denotes the node with UFBoot = 100% and BPP = 1.0. Locality numbers (Q1—Q3 and L1—L6) are shown in Fig. 1 and Table 1. on the cephalic plate and the density of setae on ultimate legs. However, the pigmentation is subject to intraspecific variation in this genus and other scolopendromorph taxa and the density of the ultimate leg setae is also variable within Scolopocryptops species (e.g. Lewis (2003); Le et al. (2023)). It should be noted that the body lengths of the specimens from the southern Ryukyus and Taiwan (19.9— 27.8 mm) were generally smaller than those from Okinawa Island (23.4—37.2 mm; juveniles excluded). Nonetheless, body length difference is not considered a taxonomic char- acter here because it may be the result of different growth rates under different habitats (as speculated on Cryptops species; Lewis (2007)) or simply due to sampling bias. Additionally, because the COI genetic distances between the two lineages (5.77—8.87%) fell within the intraspecific divergence of Scolopocryptops (Garrick et al. 2018) and is smaller than the divergence amongst the sequences from the southern Ryukyus and Taiwan (4.89-9.17%), all the specimens from the Ryukyu Islands and Taiwan are con- sidered a single species, S. /ongisetosus sp. nov. It is noteworthy that four COI sequences of “S. capil- lipedatus” from Taiwan belonged to S. /ongisetosus sp. nov. (Suppl. material 3). Scolopocryptops capillipedatus, which is characterised by the dense setae on femur and subsequent articles of ultimate legs, was originally de- scribed from South Korea and has been recorded from Japan, Taiwan and Vietnam (Takakuwa 1938; Miyosi 1971; Chao and Chang 2003, 2008; Chao 2008; Le et al. 2023). Chao (2008) provided a description of Taiwanese “S. capillipedatus” and noted that there were numerous long setae densely covering the femur, tibia and tarsi of the ultimate leg. However, this nominal species is clearly distinct from S. /Jongisetosus sp. nov. because tergal keels are absent in S. capillipedatus (Takakuwa 1938, 1940). Although Chao (2008) did not mention the presence or ab- sence of tergal keels, our specimens from Taiwan (KUZ Z5126 and Z5127) exhibited the characteristic longitudinal keels on tergites. The present result thus indicates that the records of “S. capillipedatus” from Taiwan were based on misidentified specimens of S. /ongisetosus sp. nov. Chao (2008) also described an unapparent median suture on terg- ites 1-3, which was not observed in our specimens, but described in Takakuwa’s (1938) original description of S. capillipedatus. Morphological variability amongst Tai- wanese populations needs to be examined, based on further taxon sampling. Scolopocryptops longisetosus sp. nov. is quite similar to S. quadristriatus because they both have four longi- tudinal keels and median depression on tergites, as well as dark pigmentation on antennae and the dorsal sur- face of the body. Despite their phenotypic similarities, phylogenetic analyses did not support their sister rela- tionship, but united S. /ongisetosus sp. nov. with three Japanese nominal species, S. ogawai, S. musashiensis and “S. nipponicus’, which lack tergal keels (Fig. 13). The obtained phylogeny thus indicates that the longitu- dinal keels on tergites evolved in parallel within Scol- opocryptops or that the presence of keels represents a zse.pensoft.net 420 plesiomorphic character of the clade comprising these species. Within Scolopocryptops, the tergal keels and median depression are unique to these two species and S. hoanglieni Le, Schileyko & Nguyen, 2023, which bears ““drop-like’ longitudinal median depression bordered by paramedian keels” (Le et al. 2023). However, the phylo- genetic implication of the tergal keels remains uncertain because the phylogenetic position of S. hoanglieni 1s un- determined (Le et al. 2023). It is also notable that all members of the ex-e/egans lineage, except S. rubiginosus, lack complete parame- dian sutures on tergites (Shinohara 1984, 1990: Shelley Jonishi, T. & Nakano, T.: Scolopocryptops quadristriatus and new species from Japan and Taiwan 2002; Le et al. 2023), whereas all species of the elegans lineage and most of the Neotropical/Afrotropical species bear complete paramedian sutures (e.g. Chagas-Jr (2008); Chagas-Jr et al. (2023); Jonishi and Nakano (2023); Le et al. (2023)). Nonetheless, the evolutionary history of their paramedian sutures and the tergal keels remains un- clear because the analyses failed to reconstruct a robust phylogeny of the ex-e/egans lineage. Further systematic studies, for example, phylogenetic analyses using a larger taxon set including data from additional loci, need to be conducted to reveal the morphological evolution of Scol- opocryptops species. An identification key to Scolopocryptops species of East Asia Diagnostic characters and known localities mainly follow Le et al. (2023), but were updated, based on Jonishi and Nakano (2023) and the present study. wo ll NM lB KI Lesebéaring seoiment -withwelledeveloped Spiracles se. n. cnc 88 ee 55 oo eR orleans a oy A eel TS os 2 EES: PeariGeSeShneils ime MIMIOSSOITECISS sri. Tir ctsties tiles Ul eh ee ce oats Fetes tse sees d ie ehhh lee sr eats Te ieo eae ltt 3 Tergite of ultimate leg-bearing segment with median sutureS. broelemanni broelemanni Kraepelin, 1903 (eastern China) Tergite of ultimate leg-bearing segment lacking median suture .... S. broelemanni esulcatus Attems, 1938 (southern Vietnam) GepiiallGsprate-WithieOinl plore Laleral MAKING LOM: 5 1..5...<4.