Taxonomic re-appraisal of Scolopocryptops quadristriatus (Verhoeff, 1934) and a description of a new species from Japan and Taiwan (Chilopoda, Scolopendromorpha, Scolopocryptopidae) Author Jonishi, Taro https://orcid.org/0000-0002-9620-0618 Department of Zoology, Graduate School of Science, Kyoto University, Kyoto 606 - 8502, Japan ykn347635@gmail.com Author Nakano, Takafumi https://orcid.org/0000-0001-6107-2188 Department of Zoology, Graduate School of Science, Kyoto University, Kyoto 606 - 8502, Japan text Zoosystematics and Evolution 2024 2024-04-09 100 2 405 423 http://dx.doi.org/10.3897/zse.100.119297 journal article http://dx.doi.org/10.3897/zse.100.119297 1860-0743-2-405 6B07BF7A9ADA4959A65459E747E200DE EE3B0B141BBF5B9BABB2CA9EE36EC7B1 Scolopocryptops longisetosus sp. nov. Figs 8 , 9 , 10 , 11 , 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 Prefecture - 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 m alt.; 5 May 2021 ; F. Okuyama leg. • 1 ♂ , 30.7 mm (KUZ Z5112); Nago City, Mt. Nago-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 ♀ , 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 ♂ , 31.1 mm (KUZ Z5117); Kunigami-son, Uka; 26°48.43'N , 128°16.04'E ; approx. 330 m alt.; 15 June 2022 ; T. Jonishi leg. • 1 ♂ , 37.2 mm (KUZ Z5119); Kunigami-son , Benoki ; 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 locality 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); Kunigami-son , Uka ; 26°48.30'N , 128°16.11'E ; approx. 320 m alt.; 4 Jan 2022 ; F. Okuyama leg. • 2 juveniles , approx. 10 mm (KUZ Z5110), 1 ♀ , 24.4 mm (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 ; approx. 210 m alt.; 14 June 2022 ; T. Kato leg. • 1 ♀ , 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); Kunigami-son , Benoki ; 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 ♂ , 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. Taiwan • 1, 21.5 mm (KUZ Z5126); Hsinchu County , Wufeng Township , 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 various lengths dorsally on two basal articles, subsequent articles densely covered with long setae and minute setae. 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 specimens 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 articles; 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 ). Figure 8. Scolopocryptops longisetosus sp. nov., paratype, ♂ (KUZ Z5119: B ), non-type specimen from Yonaguni Island, ♂ (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. Antennae 10.1 mm in length, approx. 0.3 x 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 various-sized collars. Cephalic plate as long as wide; its surface sparsely punctate with minute hairs, with complete lateral margination (Figs 9B , 10A ). Figure 9. Scolopocryptops longisetosus sp. nov., holotype, ♂ (KUZ Z5107). A. Basal articles of left antenna, dorsal view; B. Cephalic plate, dorsal view. Scale bars: 0.5 mm ( A ); 1 mm ( B ). Figure 10. Scolopocryptops longisetosus sp. nov., holotype, ♂ (KUZ Z5107: A, C-E ) and non-type specimen from Yonaguni Island, ♂ (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 ). 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, divided into two low lobes by median diastema (Fig. 10E ). Tergites sparsely punctate [sparse minute setae present in small individuals]; tergite 1 with anterior transverse suture, anterior margin overlapped by cephalic plate (Figs 10A , 11A ). All tergites lacking paramedian sutures; tergites 5 [4]-21 [20] with longitudinal median depression bordered by paramedian keels, lateral keels present on tergites 5 [5-8]-20 [18-20]; median depression and keels unapparent on anterior and posterior 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 tergites 4, 6 [2 or 3 sulci present on tergites 3-21; absent in several specimens]. Figure 11. Scolopocryptops longisetosus sp. nov., holotype, ♂ (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, dorso-lateral view. Abbreviations: cxp - 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 ). Sternites sparsely punctate, lacking paramedian sutures (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, respectively, with tibial spur and tarsal spur; leg 22 without spurs. All legs with two accessory spines. Coxopleuron approx. 1.8 x [1.5-1.8 x ] as long as sternite of ultimate leg-bearing segment (Fig. 11F ). Dorsal margin of ultimate pleuron protruding from lateral margin of tergite of ultimate leg-bearing segment, dorso-posterior margin with minute dark spine (Fig. 11F, G ). Posterior and ventral margins of coxopleuron converging posteriorly, forming approx. 60° [60-65°] angle; coxopleural process short, tip of process pointed, slightly directed dorsally (Fig. 11F ). Surface of coxopleuron without setae, covered with various-sized coxal pores (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. 0.3 x [0.3-0.37 x ] 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-medial one; pretarsus with two accessory spines (Fig. 12A ). Figure 12. Scolopocryptops longisetosus sp. nov., holotype, ♂ (KUZ Z5107: A, C, D ); non-type specimen from Yonaguni Island, ♂ (KUZ Z5124: B ); paratype, ♀ (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, 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, B ); 0.5 mm ( C-E ). Genital segments occupying approx. 0.8 x length of sternite of ultimate leg-bearing segment; tergite of genital segment sparsely setose (Fig. 12C, D ). Sternite of genital segment 1 sparsely covered with setae, posterior margin slightly convex (Fig. 12C, D ). Sternite of genital segment 2 well developed, covered with sparse setae; penis visible 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 cephalic 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. Remarks. This species resembles S. quadristriatus , 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. capillipedatus by the presence of longitudinal keels and median depression on tergites. Molecular phylogeny and genetic distances. The ML (ln L = -14816.13; not shown) and BI (mean ln L = -14846.27; Fig. 13 ) trees had almost identical topologies. In the Asian/North American group (UFBoot = 100%, BPP = 1.0), both the Scolopocryptops elegans lineage (UFBoot = 100%, BPP = 1.0) and the ex- Scolopocryptops elegans 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. longisetosus sp. nov. (UFBoot = 100%, BPP = 1.0) was strongly supported. 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 . Although the interspecific relationships remained largely undetermined, the analyses showed that S. longisetosus sp. nov. is sister to a clade comprising three of the Japanese nominal species, S. ogawai Shinohara, 1984, S. musashiensis Shinohara, 1984 and " S. nipponicus " sensu Edgecombe et al. (2012) (UFBoot = 99%, BPP = 1.0) (Fig. 13 ). In the preliminary analyses using the COI dataset, four sequences of " S. capillipedatus " from Taiwan (AB617528-AB617530, AB672646) were nested within S. longisetosus sp. nov. (UFBoot = 98%, BPP = 0.99; Suppl. material 3), but the relationships amongst populations were not resolved. In contrast, the concatenated analyses recovered two lineages within S. longisetosus sp. nov., which corresponded to the specimens from Okinawa Island (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 lineage 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).