Krateraspis meinerti (Sseliwanoff, 1881)

Dyachkov, Yurii V. & Bonato, Lucio, 2022, Morphology and distribution of the Middle Asian centipede genus Krateraspis Lignau, 1929 (Chilopoda, Geophilomorpha, Mecistocephalidae), ZooKeys 1095, pp. 143-164 : 143

Krateraspis meinerti (Sseliwanoff, 1881)

Figures 2-9 , 10-14 , 15-17 , 18-24 , 25-28

Mecistocephalus meinerti : Sseliwanoff 1881a: 9 (nomen nudum). Sseliwanoff 1881b: 232 (original description). Sseliwanoff 1884: 73 (description). Attems 1903: 168, 210. Attems 1904: 115. Attems 1914: 21. Attems 1929: 156. Izotova 1960: 150 (misidentification).

Krateraspis meinerti : Lignau 1929a: 160, 165 (new record). Lignau 1929b: 207 (redescription); figs 7-11. Verhoeff 1930: 264. Titova 1965: 871 (new record). Titova 1969: 165. Titova 1975: 39, 45 (new records), 46 (in key); fig. 2: 2-4B. Bonato et al. 2003: 543, 545, 546, 550, 551, 577. Ilie et al. 2009: 14. Bonato 2011: 434. Volkova 2016: 675. Dyachkov 2019: 368, 371 (new records; description), 373 (in key); figs 6-10. Dyachkov 2020: 79 (new records), 85. Dyachkov and Nedoev 2021: 44 (new records), 47.

Tygarrup asiaticus : Verhoeff 1930: 260 (original description); figs 20-21. Verhoeff 1934: 31. Verhoeff 1937: 235 (in key). Verhoeff 1939: 88 (in key). Takakuwa 1940: 84. Verhoeff 1940: 31. Verhoeff 1942: 49 (in key). Shinohara 1965: 303 (in key), 304. Titova 1965: 871, 874 (in key). Titova 1983: 147, 148.

Type locality.

"Chinas, bl. Tashkenta" ( Sseliwanoff 1881a), also indicated as "Chinad [sic] bliz Tashkenta" ( Sseliwanoff 1881b) and "Mestechko Chinas, bliz Tashkenta" ( Sseliwanoff 1884) [Uzbekistan, Tashkent region, Chinaz town, ca. 40°56'N, 68°45'E].

Synonyms.

Tygarrup asiaticus Verhoeff, 1930 (synonymization since Titova 1975; see below, under Remarks).

Examined specimens.

Holotype of Mecistocephalus meinerti Sseliwanoff, 1881: ♀, from Chinas bl. Tashkenta [Uzbekistan, Tashkent region, Chinaz near Tashkent], 1878, VRlg (ZISP). Lectotype of Tygarrup asiaticus Verhoeff, 1930 (see below, under Remarks): ♂, from Tashkent (NHRS-JONI 714). Other material: 1 ♀, from Tashkent, 13.III [year unknown], NZlg (ZISP chilo-52); 1 ♀, from Ugam Mts, Sidzhak, nut [ Juglans ] forest, soil samples, 28.IV.[19]74, MGlg (ZMMU Rc 7408); 1 ♀, from Chimgan, nut forest, 07.V.[19]74, MGlg (ZMMU Rc 7413); 1 ♀, from Chimgan, Tashkent ravine, VII.[19]06, LBlg (ZISP chilo-1); 1 ♀, from Vrevskaya Station [now Almazar, ca. 40°57'N, 68°50'E], 26.IV.1932, V lg (ZISP chilo-5); 1 ♂ and 1 ♀, from Kamsay, near Khumsan, Juglandetum , 03.V.[19]74, MGlg (ZMMU Rc 7407); 1 ♂ and 5 ♀♀, from Khumsan, right bank of Ugam river, nut forest, 1.V.[19]74, MGlg (ZMMU Rc 7406); 1 ♀, from [Tajikistan, Districts of Republican Subordination, Roghun district] left side of Obi-kandak river valley (left stream tributary of Obigarm river), stony meadow with rocks, 38°43.275'N, 69°43.863'E, 1250-1540 m, 23.IV.2019, AFlg (ASU No. 261); 2 ♂♂, from [Khatlon region, Mu’minobod district], Muminabad [ Mu’minobod, now Leningradsky, ca. 38°06'N, 70°01'E], 0-10 [cm deep], 19.V.[19]62 (ZMMU Rc 8158) and 11.V.[19]65 (ZMMU Rc 8159); 2 ♂♂, 3 ♀♀ and 1 body fragment from Sharak [village, ca. 38°16'N, 70°04'E], 10-20 [cm deep], 15.VIII.[19]65 (ZMMU Rc 8136), 10-20 [cm deep], 27.V.[19]65 (ZMMU Rc 8185), 20-30 [cm deep], 19.X.[19]64 (ZMMU Rc 8148), 10-20 [cm deep], 31.V.[19]65 (ZMMU Rc 8155), grass, 0-10 [cm deep], 3.VI.[19]63 (ZMMU Rc 8139); 1 ♂ and 6 ♀♀, from [Yovon district], Yavan [Yovon, ca. 38°18'N, 69°03'E]: Triticum , 20-40 cm deep, 25.VII.[19]67 (ZMMU Rc 8151), Triticum , 20-60 [cm deep], 20.X.[19]67 (ZMMU Rc 8186), Triticum , 0-30 [cm deep], 19.X.[19]68 (ZMMU Rc 8170), Triticum , 10-20 [cm deep], 21.V.[19]68 (ZMMU Rc 8183), Hordeum , 0-10 [cm deep], 13.V.[19]67 (ZMMU Rc 8169), Avena , 0-10 [cm deep], 26.V.[19]68 (ZMMU Rc 8172); 2 ♂♂ and 4 body fragments, from [Sughd region], Matcha district [ca. 40°32'N, 69°25'E]: 10-20 [cm deep], [date unknown], D lg (ZMMU Rc 8157), 15.V.[19]65 (ZMMU Rc 8149), and 0-10 [cm deep], D lg (ZMMU Rc 8190); 1 ♀, from Mogol-Tau Mts [ca. 40°23'N, 69°31'E], under stones, [19]74 (ZMMU Rc 7409); 1 ♀, from F-bad [unknown region, Fayzobod], Triticum , 10-20 [cm deep], 6.V.[19]66 (ZMMU Rc 8174); 1 ♂, from [Kyrgyzstan, Jalal-Abad region], Sary-Chelek Nature Reserve, near Arkit Village [ca. 41°47'N, 71°57'E], forest with Juglans and Acer , 03.VII.[19]83, ARlg (ZMMU Rc 7670); 1 ♂, from near Kyttelsay stream, forest with Juglans , 04.VII.[19]83, ARlg (ZMMU Rc 7667); 5 ♂♂, 3 ♀♀ and 3 juv., from Kazakhstan, Turkistan region, 10 km SW Abay Village, Karatau Mt. Range, Karatau State Nature Reserve, cereals and tulip steppe, under stones, 43°47'04.2"N, 68°46'42.0"E, 1020 m, 06-07.V.2017, YDlg (ASU No. 214); 1 ♀, from 50 km NW Achisay Village, Kyzylkol Lake coast, in clay stones, 43°46'34.0"N, 69°30'36.4"E, 328 m, 08-09.V.2017, YDlg (ASU No. 215); 5 ♂♂, 10 ♀♀ and 5 juv., from Karatau Mt Range, Syrdarya-Turkestan Natural Park, near Terekty Village, Boralday River coast, Morus and cereals, under stones, 42°51'48.2"N, 69°51'55.0"E, 529 m, 14-15.V.2017, YDlg (ASU No. 216); 9 ♂♂, 6 ♀♀ and 3 juv., from Ugam Mt Range, Sayram-Ugam National Park, 10 km NE Tylkubas Village, Iirsu River Valley, meadow, under stones, 42°24'58.0"N, 70°21'30.08"E, 1296 m, 16-18.V.2017, YDlg (ASU No. 217).

Remarks on nomenclatural issues.

The species name Mecistocephalus meinerti was first introduced by Sseliwanoff (1881a) without description, definition, or indication, and therefore it is not available from that publication (ICZN 1999: Art. 12.1 and 12.2). The name became available since another paper published later ( Sseliwanoff 1881b), which provided a morphological description of the species, based on a specimen.

Verhoeff (1930) described Tygarrup asiaticus based on nine specimens from two localities (seven from Vreskaja, ca. 50 km SW of Tashkent, and two from Tashkent) and all these specimens should be considered as syntypes (ICZN 1999: Recommendation 73F). They are preserved in different museums: at least four in ZSM ( SysTax 2021), one in ZMB ( Moritz and Fischer 1979; pers. obs.), one in NHMUK ( Natural History Museum 2021), one in NHRS (pers. obs.), and one in NHMW ( Ilie et al. 2009). The descriptions and illustrations provided by Verhoeff and our direct examination of two syntypes (NHRS-JONI 714 and ZMB 3610) revealed that Verhoeff described T. asiaticus mainly on some syntypes that are fully consistent with Krateraspis meinerti . Other syntypes actually belonging to another species were misinterpreted by Verhoeff as juveniles of Tygarrup asiaticus . To stabilize the usage of the name, we herewith designate NHRS-JONI 714 as lectotype of T. asiaticus (ICZN 1999: Art. 74.1.1). This specimen (Fig. 18 ) is fully consistent with the original description and illustrations published by Verhoeff (1930) for the adult morphology of T. asiaticus and Krateraspis meinerti . It is an adult male 31 mm long, labeled " Tygarrup asiaticus Verh. Turkestan", acquired by NHRS in 1931 and indicated explicitly as type in the catalogue of NHRS. This specimen has been now labeled “lectotype”, whereas other previous syntypes has been now labeled “paralectotype” (ICZN 1999: Recommendation 74C).

ZMB 3610 (labeled as a syntype of T. asiaticus , from Tashkent, with 43 pairs of legs; Figs 29-32 ) actually belongs to a species of Arrup Chamberlin, 1912, as indicated by the following characters: clypeus with many setae on the lateral parts and very short paired plagulae (Fig. 30 ), first maxillae with relatively small telopodites (Fig. 30 ), forcipular tarsungulum with a relatively long denticle (Fig. 32 ), and 41 pairs of legs (erroneously reported 43 on the label on the microscopic slide; Fig. 29 ). More precisely, ZMB 3610 probably belongs to the species A. asiaticus (Titova, 1975), which is already known from Middle Asia and differs from all other known species of Arrup in the variable presence of coxal organs and pores (apparently absent in some specimens, including well grown specimens) and the branching structure of the channels of the anal organs and their broad openings (Fig. 31 ; Titova 1975; Dyachkov 2019).

Diagnosis.

A species of Krateraspis with: clypeus showing the transition between marked and weak areolation very close to the clypeal anterior margin (at ca. 0.1 of the medial length of the clypeus), so that all clypeal setae are inside the weakly areolate central part of the clypeus; some small spine-like sensilla on the lateral parts of the clypeus; second maxillary telopodites distinctly surpassing the tips of the telopodites of the first maxillae; first article of the second maxillary telopodites without a distinct distal bulge on the external side; all forcipular articles with a distinct denticle; invariably 45 pairs of legs. See also Table 2 .

Redescription of holotype.

Body stiffened, divided in two parts (Fig. 7 ); many legs missing. Total length ca. 48 mm; maximal width 1.4 mm (at ca. LBS 21-22). Color (in 70% ethanol) brown.

Head (Fig. 2 ). Cephalic plate 1.7 × as long as wide, sub-rectangular but slightly widening anteriorly, its posterior margin straight. Transverse suture distinct, with a medial forward angle. Antennae ca. 5 mm, ca. 4.5 × as long as the head maximum width.

Forcipular segment (Figs 2 , 3 ). Tergite sub-trapezoid, ca. 1.5 × as wide as long, with a mid-longitudinal distinct furrow inside an oval depression. Coxosternite as long as wide, with a pair of small anterior denticles. Trochanteroprefemur 1.4 × as long as wide; tarsungulum 2.9 × as long as wide. All forcipular articles with denticles: a large distal denticle on the trochanteroprefemur, femur and tibia each with a small denticle, tarsungulum with a basal small denticle. Inner edge of tarsungulum slightly serrated.

Leg-bearing segments (Figs 4-6 ). Tergites 2-43 with a pair of paramedian sulci. Metasternites 2-44 with a median longitudinal sulcus. 45 LBS. Legs 1 slightly smaller than following legs; pretarsi with two accessory spines.

Ultimate leg-bearing segment (Figs 8 , 9 ). Metatergite shield-shaped, 1.7 × as long as wide, and 1.2 × as wide as the pretergite. Metasternite subtriangular, 1.1 × as wide as long, its anterior margin ca. 3 × as wide as the posterior one. Ca. 50 pores on each coxopleuron, scattered on ventral and lateral sides. Legs slender, but incomplete (missing tarsus 2 of right leg, tibia and both tarsi of left leg).

Postpedal segments (Figs 8 , 9 ). Intermediate sternite and first genital sternite well-developed. Gonopods bi-articulate, triangular, and touching each other at their bases. Anal pores present.

Intraspecific variation.

Maximum body length: 71 mm in ♀♀ (n = 44; the largest specimen in the sample ZMMU Rc 7406), 58 mm in ♂♂ (n = 31). Color (in 70% ethanol) usually yellow, with cephalic plate, forcipular segment, and antennae light brown (Figs 10 , 11 ).

Head. Anterior markedly areolate part of the clypeus extending medially for 10-17% of the total clypeal length (Fig. 14 ). Clypeal setae usually 8, rarely 6. Labral mid-piece usually pointed and projecting backwards beyond the posterior margins of the labral lateral ones. Each mandible (Fig. 21 ) usually with six lamellae, with 5-9 teeth in each lamella. Second maxillae (Figs 15-17 ): 1st article invariably without a distinct distal bulge on the external side; distal parts of 2nd and 3rd articles usually with numerous setae.

Forcipular segment. Tergite usually partially covered by tergite 1 (Fig. 10 ) and forcipules usually surpassing the anterior margin of the cephalic plate (Figs 11 , 18 ). All forcipular articles with denticles (Figs 11-13 ), with the single exception of a specimen missing the denticle on the right femur (collected together with other specimens with usual morphology, in the sample ASU No. 216). Worth noting is that an analogous case of asymmetry has been detected in a specimen of K. sselivanovi , where a denticle has been recognized on one femur but not in the other femur (see below). Distal denticle on the trochanteroprefemur usually larger than all other denticles (Fig. 12 ). Denticle on the tibia slightly larger than the denticle on the femur and the basal denticle on the tarsungulum (Figs 12 , 13 ). Calyx of poison gland usually reaching the trochanteroprefemur in both sexes (Figs 12 , 13 ).

Leg-bearing segments. Invariably 45 pairs of legs. Worth noting is that K. sselivanovi has invariably 53 pairs of legs and the difference of eight pairs between the two species corresponds to a putative evolutionary change that have repeatedly occurred in the Mecistocephalidae ( Bonato et al. 2003).

Ultimate leg-bearing segment. Almost similar in both sexes, slightly thickened in male (Figs 22 , 23 , 25 , 26 ). Metasternite subtriangular, its length to width ratio varying between 0.8 and 1.1, and the anterior margin 3-5 wider than the posterior one; up to ca. 50 pores on each coxopleuron in both sexes; legs densely setose, without pretarsus in both sexes.

Postpedal segments. Densely setose in both sexes (Figs 22 , 23 , 25-28 ). Male gonopods bi-articulate, narrower and separated by a conic projection in between (Figs 25 , 27 ). Female gonopods bi-articulate, subtriangular, and touching each other at their bases (Figs 22 , 28 ).

Distribution.

Recorded from 24 localities, from Western Tian-Shan to the western offshoots of Pamir Mts (Fig. 1 ), in the following countries and administrative units: Kazakhstan (Turkistan and Jambyl regions), Uzbekistan (Tashkent region), Kyrgyzstan (Jalal-Abad region), and Tajikistan (Region of Republican Subordination, Khatlon, and Sughd regions) ( Sseliwanoff 1881a, 1881b, 1884; Lignau 1929a, 1929b; Titova 1965, 1975; Dyachkov 2019, 2020; Dyachkov and Nedoev 2021; present records).

A specimen from Tatarstan (European Russia) was assigned by Izotova (1960) to K. meinerti with doubt (see also Volkova 2016; Dyachkov 2019). The relative size of the forcipular tergite ( Izotova 1960: fig. 6) shows that this specimen does not belong to Mecistocephalidae , and the shape of the forcipular segment suggests instead a species of the geophilid genus Arctogeophilus Attems, 1909. The latter is known from European Russia and resembles Krateraspis in the elongation of the head, the shape of the forcipular coxosternite, the pattern of forcipular denticles, the number and arrangement of coxal pores, and the absence of ultimate pretarsi (see, e.g., Folkmanová and Dobroruka 1960).

Remarks on published morphological accounts.

Verhoeff (1930) indicated that Tygarrup asiaticus differs from Krateraspis meinerti in the clypeal areolation (a single long non-areolate plagula, with a short mid-longitudinal areolate strip, instead of two paired short plagulae), the shape of labrum (mid-piece not projecting backwards beyond the posterior margins of the labral lateral pieces), and the second maxillary pretarsi (absent). However, Verhoeff ignored the weak areolation on the central part of the clypeus and described an entire non-areolate plagula, even though recognizing a mid-longitudinal areolate strip. The putative difference in the labrum may be explained by artefacts. As for the second maxillary pretarsus, it was described and illustrated as missing in T. asiaticus by Verhoeff (1930), but this character was ignored in keys published later by the same author ( Verhoeff 1937, 1939, 1942). Moreover, a pretarsus is recognizable in the second maxillae of the lectotype (NHRS-JONI 714; Fig. 18 ), while it is absent in ZMB No. 3610, which is an Arrup specimen originally misinterpreted by Verhoeff (1930) as a juvenile T. asiaticus (see above, under Remarks on nomenclatural issues).

Figures

Figure 1. Distribution of Krateraspis Lignau, 1929: square, K. meinerti (Sseliwanoff, 1881); star, K. sselivanovi Titova, 1975. Green symbols indicate type localities. Some very close localities are marked by a single symbol. The question marks indicate alternative positions of the uncertain locality " Fayzabad " (see Materials and methods).

Figures 2 - 9. Holotype of Krateraspis meinerti (Sseliwanoff, 1881), from Chinaz near Tashkent (ZISP): 2, 3 head, forcipular and LBS 1 (dors., ventr.) 4 anterior LBS (ventr.) 5, 6 intermediate LBS (ventr., dors.) 7 anterior and posterior parts of the body (dors.) 8, 9 terminal part of the body (dors., ventr.). Abbreviations: 15 s, 16 s, 17 s - metasternites 15 - 17, cp - coxopleural pores, us - metasternite of the ultimate LBS, ut - metatergite of the ultimate LBS. Scale bars: 0.5 mm (2 - 6, 8 - 9); 2 mm (7).

Figures 10 - 14. Krateraspis meinerti (Sseliwanoff, 1881): 10, 11 head, forcipular and LBS 1 (dors., ventr.) 12, 13 left forcipule (ventr.) 14 clypeus and labrum (ventr.). Specimens: ♀ (10, 11, 13, 14) and ♂ (12), from Syrdarya-Turkestan Natural Park (ASU No. 216). Abbreviations: aa - anterior ala, ap - markedly areolate anterior part of clypeus, c - calyx of poison gland; cp - central part of clypeus with distinct but fainter areolation, fe - femur, ms - mid-longitudinal areolate strip, p - plagula, pa - posterior ala, ta - tarsungulum, ti - tibia, tl - transverse thickened line, tr - trochanteroprefemur, ts - transverse suture. Scale bars: 0.5 mm (10, 11); 0.1 mm (12 - 14).

Figures 15 - 17. Krateraspis meinerti (Sseliwanoff, 1881), ventr.: 15 maxillary complex 16 right telopodite and coxal projection of first maxillae 17 left telopodite of second maxillae. Specimen: ♀ from Syrdarya-Turkestan Natural Park (ASU No. 216). Scale bars: 0.1 mm (15); 0.05 mm (16, 17).

Figures 18 - 24. Krateraspis meinerti (Sseliwanoff, 1881): 18 head and forcipular segment (ventr.) 19 LBS 2 (ventr.) 20 distal end of tarsus of leg 12 (lateral view) 21 left mandible (ventr.) 22, 23 ultimate LBS and postpedal segments (ventr., dors.) 24 terminal articles of ultimate leg (ventr.). Specimens: 18 lectotype of Tygarrup asiaticus Verhoeff, 1930 from Tashkent (NHRS-JONI 714) 19 - 24 ♀ from Syrdarya-Turkestan Natural Park (ASU No. 216). Abbreviations: cp - coxopleural pores, ms - median longitudinal sulcus, ups - presternite of ultimate LBS, upt - pretergite of ultimate LBS, us - metasternite of ultimate LBS, ut - tergite of ultimate LBS. Scale bars: 0.2 mm (18, 19, 22 - 24); 0.1 mm (20); 0.02 mm (21).

Figures 25 - 28. Krateraspis meinerti (Sseliwanoff, 1881): 25, 26 ultimate LBS and postpedal segments (ventr., dors.) 27, 28 postpedal segments (♂ and ♀; ventr.). Specimens: ♂ (25, 26), from Sayram-Ugam National Park (ASU No. 217); ♂ (27) and ♀ (28), from Syrdarya-Turkestan Natural Park (ASU No. 216). Abbreviations: ups - presternite of ultimate LBS, upt - pretergite of ultimate LBS, us - metasternite of ultimate LBS, ut - tergite of ultimate LBS. Scale bars: 0.1 mm.

Figures 29 - 32. Specimen of Arrup misidentified by Verhoeff (1930) as juvenile Tygarrup asiaticus Verhoeff, 1930 (ZMB 3610), ventr.: 29 microscopic slide 30 head 31 ultimate LBS and postpedal segments; 32 forcipular segment and LBS 1. Abbreviation: ap - anal pore. Scale bars: 0.5 mm (30 - 32).