Scolopocryptops quadristriatus (Verhoeff, 1934)

Jonishi, Taro & Nakano, Takafumi, 2024, Taxonomic re-appraisal of Scolopocryptops quadristriatus (Verhoeff, 1934) and a description of a new species from Japan and Taiwan (Chilopoda, Scolopendromorpha, Scolopocryptopidae), Zoosystematics and Evolution 100 (2), pp. 405-423 : 405

Scolopocryptops quadristriatus (Verhoeff, 1934)

Figs 2 , 3 , 4 , 5 , 6 , 7 Japanese name: Yosuji-akamukade

Otocryptops sexspinosus quadristriatus Verhoeff, 1934: 54; Takakuwa (1939: 699), fig. 3 (as Octocryptops [sic] Scolopocryptops sexspinosus Scolopocryptops quadristriatus ); Takakuwa (1940: 73), fig. 77.

Scolopocryptops quadristriatus : Shinohara (1984: 41); Shinohara et al. (2015: 880, 906); Le et al. (2023: 437, 442).

Scolopocryptops quadristriatus Not S. quadristriatus . " Otocryptops sexspinosus quadristriatus ": Takakuwa (1933b: 1457, 1459) (nomen nudum).

Type specimen.

Otocryptops s. quadristriatus was described, based on a single specimen from the vicinity of Tokyo, without any information on the collector, collecting date or deposition of the specimen ( Verhoeff 1934). Most of the Verhoeff’s specimens are now kept at the Zoologische Staatssammlung München (ZSM), the Museum für Naturkunde (MfN) and the Naturhistorischen Museums Wien (NHMW) ( Melzer et al. 2011). 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 Stagl 2004). ZSM A20051244 (Fig. 2 ) is, thus, the only specimen of O. s. quadristriatus remaining within the Verhoeff’s collection and is supposed to be the holotype 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 separate 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 consistent 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 myriapodologist, 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 A 20051244.

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); Hachioji City, Mt. Takao-san ; 18 Nov 2017; T. Kato leg. • 1, 28.6 mm (KUZ Z5093); Hachioji City, Uratakao-machi ; 35°38.69'N, 139°14.36'E; approx. 250 m alt.; 18 Oct 2021; T. Jonishi leg. • 1 ♂, 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 ♂, 38.2 mm (KUZ Z5099); Ome City, Sawai ; 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; approx. 250 m alt.; 12 Apr 2022; Futaro Okuyama leg. • 1, 29.6 mm (KUZ Z5103); Hachioji City, Uratakao-machi ; 35°38.75'N, 139°14.16'E; approx. 270 m alt.; 6 Nov 2022; T. Kato leg. - Saitama Prefecture • 1 ♀, 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 Matsushita 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.

Diagnosis.

Antenna with sparse short hairs and setae on dorsal surface of two basal articles, subsequent articles densely covered with short setae. Cephalic plate with complete lateral marginal sulci. Tergites lacking paramedian sutures, tergites 6-20 with four longitudinal keels and median depression bordered by paramedian keels.

Description

[variations given in square brackets]. Body length approx. 27.7-39.0 mm in 75% ethanol. 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 cephalic 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 × as long as body, composed of 17 articles; two basal articles 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 trochanteroprefemora sparsely punctate, coxosternite with median suture and transverse sutures cross median one on anterior third of coxosternite (Fig. 5D ); trochanteroprefemur with small and blunt black process with basal suture (Fig. 5D ); anterior margin of coxosternite strongly sclerotised 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 cephalic plate (Figs 5A , 6A ). All tergites lacking paramedian sutures; tergites 6 [5-7]-21 with longitudinal median depression bordered by paramedian keels, lateral keels present 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] (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 sutures (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, 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 process 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 ).

Ultimate leg 9.0-12.4 mm in length, approx. 0.3 × 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 genital 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, covered with sparse short setae; posterior part of genital segment 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 Kyushu, 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 Takakuwa’s works.

The name " Otocryptops sexspinosus 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 sexspinosus 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 " Otocryptops ruliginosus " [sic]). Therefore, the name quadristriatus in the combination of Otocryptops sexspinosus quadristriatus in Takakuwa (1933a) did not satisfy the provision of Article 13.1 of the International Code of Zoological 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. quadristriatus " sensu Verhoeff was indistinguishable from the nominotypical 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 attributed to Verhoeff (1934), who established O. s. quadristriatus explicitly as a new subspecies. Moreover, according to Shinohara (1982, 1990), Takakuwa’s (1933b) description of this taxon was based on specimen(s) misidentified as " O. sexspinosus "; O. s. quadristriatus sensu Takakuwa (1933b) was later described as S. nipponicus Shinohara, 1990 (placed in synonymy with S. spinicaudus 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 absence of tergal paramedian sutures, the presence of four longitudinal keels on tergites and the colouration of head and leg-bearing segments.

Shinohara (1984) elevated Scolopocryptops quadristriatus to full species status, based on the following features: 1) cephalic marginal sulci reaching from postero-lateral margin of 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 slightly slender "general form" compared with other species of the genus. It is unclear whether Shinohara compared Scolopocryptops 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 Scolopocryptops quadristriatus from other species. Nonetheless, the presence of four longitudinal keels on tergites distinguishes S. quadristriatus from all other Scolopocryptops (except S. longisetosus sp. nov.; see below). The distinctness of S. quadristriatus is 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).

Figures

Figure 2. Scolopocryptops quadristriatus (Verhoeff, 1934), the likely holotype, ♀ (ZSM A 20051244). A. Cephalic capsule, maxillae, legs and left ultimate leg; B. Forcipular segment, leg-bearing segments and legs. Scale bars: 5 mm.

Figure 3. Scolopocryptops quadristriatus (Verhoeff, 1934), collected from near the type locality, ♀ (KUZ Z 5104) 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 Z 5095). A. Basal articles of left antenna, dorsal view; B. Cephalic plate, dorsal view. Scale bars: 0.5 mm (A); 1 mm (B).

Figure 5. Scolopocryptops quadristriatus (Verhoeff, 1934) from near the type locality (KUZ Z 5095). 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).

Figure 6. Scolopocryptops quadristriatus (Verhoeff, 1934) from near the type locality (KUZ Z 5095). 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, lateral 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: 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).

Figure 7. Scolopocryptops quadristriatus (Verhoeff, 1934) from near the type locality. A. KUZ Z 5095; B. KUZ Z 5094; C. KUZ Z 5104. 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).

Figure 13. Bayesian Inference tree for 2510 bp-aligned positions of the ITS 2, 28 S, 16 S 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 (Q 1 - Q 3 and L 1 - L 6) are shown in Fig. 1 and Table 1.