Phyxioschema raddei Simon, 1889

Schwendinger, Peter J. & Zonstein, Sergei L., 2011, A taxonomic revision of the genus Phyxioschema (Araneae, Dipluridae), II: species from Central Asia, Zootaxa 2815 (1), pp. 28-48 : 31-38

Phyxioschema raddei Simon, 1889

Figures 2–5

Phyxioschema raddei Simon, 1889: 385 (description of female); Simon 1892: 183, 185 (key to genera, diagnosis); Simon 1903: 968 (description of male); Charitonov 1948: 300, fig. 190 (illustration of habitus of female); Andreeva 1976: 16, figs. 8–9 (illustration of palp and leg II of male); Raven 1981: 226–228, figs. 1–7 [revision of male and of female (erroneously given as the holotype) in Simon's collection]; Raven & Schwendinger 1989: 55 (new diagnosis; synonymisation of Afghanothele lindbergi Roewer, 1960 and A. striatipes Roewer, 1960 ). For a complete list of taxonomic publications see Platnick 2011.

Ischnothele strandi Spassky, 1937: 361–363 , fig. 1 (description of female); Spassky 1952: 152, 153, 156, 200 (distribution); Spassky & Minenkova 1940 (relationships). Placed in synonymy by Charitonov 1948: 300. Synonymy is confirmed here.

Afghanothele lindbergi Roewer, 1960: 33–34 (description of female). Placed in synonymy by Raven & Schwendinger 1989: 55. Synonymy cannot be confirmed but is retained.

Afghanothele striatipes Roewer, 1960: 34 (description of female). Placed in synonymy by Raven & Schwendinger 1989: 55. Synonymy cannot be confirmed but is retained.

Type material. TURKMENISTAN: Reg. Transcaspica (Rudde [sic!; probably an incorrect transcription of the collector’s name, G. Radde, on the printed replacement label; the original label is kept apart from the specimen, personal communication by C. Rollard]), female holotype (AR 5009, 6686; MNHN) of P. raddei .—Serachs, Agar-Tschischme [= Akarcheshme], 6 syntypes of Ischnothele strandi [1 large female (vulva examined) and 5 smaller females and juvs. (vulva of one examined)] ( ZIN), 21.IV.1936, leg. L. Freiberg.

AFGHANISTAN: Darréh-Chakh [= Darreh-Shakh, Darreh-i-Shakh , Darreh-e-Shakh , Darreh-ye-Shakh ; probably in Faryab Province, E of Maimaneh, 35°41'N, 65°14'E, 1300–1500 m], 1 juv. holotype [no vulva visible in dissected genital area, but declared a female in the original description; in bad condition] (A 392; GNM) of Afghanothele striatipes , 30.X.1957, leg. K. Lindberg. — Kouh-Zarmast [= Kuhi-Zarmast, Zarmast Cave , a few km SW of Maimeneh, 870 m], 1 medium-sized juv. male holotype [given as a female in the original description] (A 365; GNM) and 1 large juv. male paratype [given as a female in the original description] ( SMF, 12995/30) of A. lindbergi , 19.X.1957, leg. K. Lindberg. — Maimanéh [= Maimeneh, Faryab Province, 35°45'N, 64°47'E, 850 m], 1 very small juv. paratype (A 397; GNM) of A. lindbergi , 29.X.1957 [27.X.1957 given in the original description], leg. K. Lindberg. — Tchidjan [near Cheikhabad, on the road Kabul-Ghazni , S of Kabul, 34°04'N, 68°45'E, 2050 m], 1 small juv. paratype (A 452; GNM) of A. lindbergi , 7.IV.1958, leg. K. Lindberg.

Other material. IRAN (northeast): Mazanderan Province [southeast of the Caspian Sea], near Dasht , 37°19'N, 56°04'E, 2 females (n° 7471; MHNG), leg. A. Senglet, 27.VII.1974 .— Mazanderan Province, above Shahpasand , 37°02'N, 55°17'E, sifting leaf litter, 1 female, 4 juvs. (n° 7473; MHNG), leg. A. Senglet, 28.VII.1974 .

TURKMENISTAN: Krasnovodsk [the type locality], 1 female ( MHNG), 3 juvs. ( TAU), leg. S. V. Ovtchinnikov, 26.III.1981 .— SW Kopetdagh Mts. , Damdam, 1 male ( TAU), leg. S. L. Zonstein, 11. VI.1983.— SW Kopetdagh, Kara-Kala , Parkhai, 2 juvs. ( TAU), leg. V. Fet, 13.III.1981 .— SW Kopetdagh, Mt. Sunt , 8 males, 1 female ( TAU), leg. N. S. Ustinova, 5.–9.VII.1982 .— W. Kopetdagh, Eldere Mt. Gorge, 5 males (one in MHNG, others in TAU), 2 females ( TAU), leg. V. Fet, 23. V.–3. VI.1980.— Aïn Deia [= Aidere], Mur. tuff., 1 female [erroneously re-described as the holotype of P. raddei by Raven 1981] and 1 male, AR 5007, 17883 ( MNHN).— SW Kopetdagh, Aidere, 16 + 13 males, 1 female, 3 juvs. ( AHK 63 ; TAU), leg. N. S. Ustinova, 3. VI.1982.— Ibidem , 1 male ( AHK 64 ; TAU), leg. N. S. Ustinova, 2.VII.1982 .— Ibidem , 1000– 1300 m, 1 male ( AHK 65 ; MHNG), leg. N. S. Ustinova, 3.–9.VII.1982 .— Kopetdagh , Aidere, house, 1 juv. male ( TAU), leg. V. Fet, 1.IV.1983 .— SW Kopetdagh, Aidere Mt. Gorge, 750–1300 m, 3 females, 1 juv. male ( TAU), leg. S. L. Zonstein, 26.–27.IV.1985 .— Kopetdagh , Aidere, 1 female, 1 juv. male ( MHNG), leg. V. Fet, 10.–20.IV.1980 .— SE Kara-Kum [= Karakum] Desert , Repetek, ca. 200 m, 1 male ( MHNG), leg. V. Krivokhatsky, 26. VI.1979.— SW Karakum Desert, Repetek , 2 females ( MHNG, TAU), leg. V. Krivokhatsky, 16.I.1982 .— Karakum Desert , Repetek, near cottage (38°35'N, 63°11'E), 1 male [partly decayed and fragmented] ( TAU), leg. V. Kapin, 22. VI.1984 ( TAU).— SE Karakum Desert, Repetek, Central Valley , 2 juv. ( TAU), leg. V. Krivokhatsky, 9. V.1982.— SW Karakum Desert, Repetek , 1 female ( MHNG), leg. Krivokhatsky, 16. V.1982.— Bairam-Ali [= Bayramaly], 1 female, leg. V. Smirnov, 31.III.1912 (n° 5513; PU) .— Badkhyz Plateau , Akartcheshme [= Akarcheshme], 1 large juvenile male ( AMNH), leg. S. Zonstein, 15.IV.1985 .— Akar-Cheshme [= Akarcheshme] well (35°47'N, 61°28'E), western part of Badkhyz Plateau, 850 m, 1 female ( TAU), leg. S. L. Zonstein, 15.IV.1985 .— Badkhyz , 1 female ( TAU), leg. N. Kuznetsov, IV.1978 .— Serakhs Distr. , Turkmenokhorasan Mts. , Elibuz Mt. Ridge, Akartcheshme Mt. Gorge, 800–850 m, 5 juv. males, 1 juv. female ( TAU), leg. S. L. Zonstein, 14.IV.1985 .— [Eroilanduz], Badkhyz Reservation , lake [probably a mistake] Kisil-Djar [= Kyzyl-Djar ravine, Kyzyl-Dzhar ravine, Kyzyldzhar ravine], 1 juv. male ( TAU), leg. V. Fet, 14.VII.1977 .— Morgunovka , N of Kushka, in house, 1 male ( MHNG), leg. V. Fet, 16.VII.1975 .— Badkhyz , Pogranichnyi Mt. Range, 4 juvs. ( TAU), leg. V. Fet, 28.II.1978 .

KAZAKHSTAN: Vost. Kyzyl-Kumy [eastern Kyzylkum desert], Khodzhal'-Bergen [Khodzha-Bergen well], 1 male, leg. N. Zarudnyi, 15. VI.1912 (n° 5510; PU).— Tchimkent Region , Arys, steppe, 6 females (including 3 juvs.), 5 juv. males (1 female in MHNG, others in TAU), leg. D. Logunov, 30.IV.1988 .

UZBEKISTAN: Tashkent Region , Bostandyk Distr. , Ugam Mt. Range, Sidjak Gorge, ca. 1200 m, 1 male ( TAU), leg. F. O. Khasanov, 30.IV.-2. V.1979.— Ferghana Valley , surroundings of Andizhan, 1 female, 3 juv. males ( TAU), leg. V. Bakhvalov, 20. V.1981.— Surroundings of Samarkand , 1 female ( TAU), leg. A. Zyuzin, 5. V.1990.— Samarkand Region & Distr. , Zeravshan Mt. Range , Ken-Kutan Gorge, near Agalyk, 1800 m, 1 female ( MHNG), leg. A. B. Nenilin, 8.IV.1979 .

TADZHIKISTAN: Aruktau Mt. Ridge , Gandjina, 800–1000 m, 37°58'N, 68°34'E, 1 female ( TAU), leg. S. Zonstein, 4. V.2002.— Ghazimailik Mt. Ridge, Ghandjina [= Gandzhina, Ganjina], 800 m, 1 female, 4 juvs. ( TAU), leg. S. L. Zonstein, 21.IV.1986 .— Ghazimailik Mt. Ridge, Gharavuti [= Garavuti], 450 m, 2 females, 3 juv. males ( TAU), leg. S. L. Zonstein, 23.IV.1986 .— South Tadzhikistan , Tigrovaya Balka [Tiger Gorge; approx. between 37°10'N, 68°18'E and 37°25'N, 68°33'E], sands, 2 females [one without opisthosoma] ( CJP), leg. T. Domracheva, 24.VII.1968 .— Pyandz-Karatau Mt. Ridge, 1600 m, Mt. Astana, W slopes, 37°23'N, 69°15'E, 2 females ( MHNG, TAU), leg. S. Zonstein, 23.IV.1991 .— Karatau Mt. Ridge, about 45 km NE of Kurgan-Tyube, 950 m, dry forest of Pistacea vera and Amygdalus spp. , 1 female ( TAU), leg. S. Zonstein, 21.IV.1989 .— Hissar Mt. Ridge, Varzob [= Varsob] Valley, 1200–1400 m, N of Dushanbe, 38°40'N, 68°47'E, 4 females ( TAU), leg. S. L. Zonstein, 26.IV.1986 .— Rangentau [= Rengentau, Rangontau] Mt. Ridge, 20 km SE of Dushanbe, Yavan Pass, 1300 m, 1 female ( TAU), leg. S. L. Zonstein, 28.IV.1986 .— Sanglok Mt. Ridge , Kolkot, 1500 m, 38°15'N, 69°17'E, 1 female ( TAU), leg. S. Zonstein, 5. V.1991 ( TAU).— Khozratisho Mt. Ridge , Sangdara, 1600 m, 38°22'N, 70°09'E, 1 female ( TAU), leg. S. Zonstein, 19. V.2002.— Khozretishoh Mt. Ridge, 1400 m, Sangdara Valley, 15 km NE of Khovaling, Juglans forest, 2 females ( TAU), leg. S. Ovtchinnikov, 15.X.1987 .

Remarks. There is some confusion about which specimen is the holotype of P. raddei . According to Raven (1981: 225) and Raven & Schwendinger (1989: 55) the female without opisthosoma from sample AR 5007, 17883 (given as no. 17889 in these two papers; either a lapsus calami or an incorrect transliteration on the printed replacement label) in Simon's collection is the holotype. The locality for this specimen is given as "Aïn Deia" on the museum label, and as "Ain-dor" and "Ain-Dor" in Raven (1981: 225) and Raven & Schwendinger (1989: 55), which are probably variant spellings of "Aidere" in the Kopetdagh Mountains of southwestern Turkmenistan. Aidere, however, is not the type locality. The type locality of P. raddei is "Krasnowodsk" (= Krasnovodsk) ( Simon 1889: 385), an old Russian settlement close to today's Turkmenbashy (= Turkmenbashi), at the eastern shores of the Caspian Sea. Krasnowodsk and Aidere are about 300 km apart. A possible explanation for this incongruence is that the original label of the holotype was lost and that a new specimen and a new label were subsequently placed with the holotype. Raven (1981: 225, apparently not being aware of the type locality) states: "… a male later added without further data to the holotype’s vial by Simon … ". However, we believe that there was no confusion of labels or mix-up of specimens, that both specimens in sample AR 5007, 17883 are from Aidere, and that the female specimen is not the holotype. It is too large to be the holotype. Carapace and chelicerae of female AR 5007, 17883 are 7.2 mm long; an intact adult P. raddei female of similar prosoma length has a body length of over 14 mm.

A second female (AR 5009, 6686) from Simon's collection deposited in the Museum of Paris, but not mentioned in Raven's revision, corresponds more closely to the original description and is most certainly the holotype of P. raddei . That specimen measures 11.1 mm from the anterior margin of the chelicerae to the posterior tip of the opisthosoma (11 mm length given in the original description), it is not fully grown ["haud plane adulta" (= not fully adult) given in the original description], it is damaged [lacking legs I–III on the right side and the distal articles of both PLS; "valde detritum" (= strongly damaged) given in the original description] and it still has its opisthosoma attached [the original description mentions the opisthosoma ("abdomen oblongum, ...")]. Furthermore, this specimen is paler than the female in sample AR 5007, 17883, which can be explained by stronger bleaching due to longer preservation in alcohol. Simon’s collection numbers (6686 for the presumed holotype and 17883 for the female from Aïn Deia) support this interpretation. The lower number was given by Simon to the earlier arrival in his collection (AR 5007 and AR 5009 are more recent MNHN registration numbers; C. Rollard, personal communication). The label data (Reg. Transcaspica) of the presumed holotype (AR 5009, 6686) do not directly refer to the type locality (Krasnowodsk), but to the holotype's region of origin and to the title of the publication (Arachnidae transcaspicae) in which the species was described.

All types of Afghanothele lindbergi and A. striatipes are juveniles (large males and small females without discernible copulatory organs) and most of them are in a very bad condition of preservation (all in the GNM have dried up and were then transferred back into alcohol). Therefore the synonymisation of these two nominal species with P. raddei by Raven & Schwendinger (1989) can be neither confirmed nor refuted. The holotype of A. striatipes was seemingly preserved shortly before the spider was to moult. Two layers of cuticle are discernible in the broken parts of the legs; hairs and spines, arranged in dark bands, of the new cuticle are clearly visible under the old cuticle of all limbs. Roewer (1960: 34) has obviously misinterpreted these underlying structures as a pigmentation of the leg cuticle in the form of dark longitudinal stripes (“1.–4. Femur bis Tarsus der Beine schwarz längs-gestreift”) and used this false character to distinguish A. striatipes from A. lindbergi . F. Coyle observed this too; he mentions it in the unpublished notes that he sent us. The paratype of A. lindbergi (a relatively large juvenile male with a body length of 10.3) in the SMF is still in fairly good condition (not dried up) and largely corresponds to females of P. raddei . Unlike most specimens of P. roxana sp. n., it has a sternum that is not much darker that the surrounding coxae. We consider it more likely that the types of Afghanothele belong to the widespread P. raddei than to the much more geographically restricted P. roxana sp. n., and thus we see no reason for removing A. lindbergi and A. striatipes from the synonymy of P. raddei .

Raven’s statement that the eye group of the female, which he considered to be the holotype, is twice as long as wide is incorrect. It is about twice as wide as long, as in all other specimens belonging to this genus.

Emended diagnosis. Fairly large species with considerable variation in size. Males distinct from those of all other Phyxioschema by: 1) ventral side of metatarsus I with one enlarged subproximal spine; 2) ventral spur on tibia II spatulate and strongly asymmetrical, its apex forming a relatively sharp concave edge with three more or less distinct distal lobes, the retrolateral half of the apex carrying two megaspines inclined from longitudinal axis of spur on tibia II; 3) metatarsus II with three proximoventral keels, the proventral one low, in some (mostly large) males indistinct. Females similar to those of P. roxana sp. n., distinguished by: 1) vulva with usually only one secondary receptacle per spermathecal trunk, 2) median receptacle with short and straight, or long and only slightly convoluted stalk, 3) lateral receptacle with base constricted to a distinct neck.

Characters not mentioned in previous descriptions. Intersegmental membranes of limbs in both sexes with some dark pigment. Tarsi I–IV of males pseudosegmented. One of the subproximal ventral spines on metatarsus I of males enlarged ( Figs. 2F , 3C–D ). Tibia I of males incrassate (slightly to distinctly wider than tibia IV, in larger specimens more so than in smaller ones), ventrally flattened ( Figs. 2F , 3E ), not cylindrical as in the species from Thailand. Patella I of males with a bent row of usually 5–6 curved and sigmoid spines retroventrally and retrodistally, without triangular projection on retrolateral margin ( Fig. 3F ). Third (proventral) keel on metatarsus II of males low, less pronounced than medioventral and retroventral keels ( Figs. 2D , 3L–O ), in some specimens difficult to see. Bands of hooked spinules retrodorsally on femur I ( Fig. 2A ) and proventrally on femur II ( Fig. 2B ) wide, the former one relatively short. Band of elongate spinules prolaterally on tibia II straight, slightly inclined from longitudinal axis of tibia ( Fig. 2D–E ). Vulva with a pair of fairly narrow spermathecae distinctly inclined from each other anteriorly, each spermathecal trunk usually carrying three receptacles (rarely two or four); median and secondary receptacles with completely sclerotised stalks; stalk of median receptacle short and straight or long and slightly convoluted; base of lateral receptacle constricted to a neck, partly sclerotised, its ental side with continuous sclerotisation, its ectal side with discontinuous (broken) sclerotisation; all receptacular heads bulbous and with pores ( Figs. 4A–H , 5A–H ).

Variation. The 51 males examined vary greatly in size: body length 6.7–15.1, carapace length 3.0–7.2, carapace width 2.6–6.1. Males from the eastern localities ( Repetek , Sidjak and Khodzha-Bergen ) are distinctly larger (12.0–15.1, 5.9–7.2, 5.0–6.1; n=4; largest male from Khodzha-Bergen in Kazakhstan) than males from the western localities ( Damdam , Mt. Sunt , Eldere , Aidere , Morgunovka ) (6.7–11.2, 3.0–4.5, 2.6– 3.6; n=47; smallest male from Aidere in Turkmenistan). The corresponding maximum measurements of the 47 adult females examined are: holotype 11.1, 3.8, 2.9; largest female (from Sangdara in Tajikistan) 21.1, 9.2, 7.4. Females from eastern localities are also generally larger than those from western localities .

A male from Repetek has three foveal setae, a female from Gandjina has only one, all others have two. Most specimens have a more or less round fovea, in some specimens it is triangular, in a few even longer than wide.

The four males from Repetek, Sidjak and Khodzha-Bergen are not only larger, but also darker, have stronger spines on the legs, more strongly incrassate tibiae I and II ( Fig. 2F , cf. Fig. 3E ), a proximally wider ventral spur on tibia II, and a less distinct proventral proximal keel on metatarsus II (in some specimens only discernible in distal view) than the males from the more western localities. The fragmented male from Repetek has two thickened spines next to the longer and distinctly enlarged subproximal ventral spine on metatarsus I, which makes the latter less conspicuous than in other males. The male from Morgunovka has seven sigmoid retroventral spines (usually 5–6) on patella I of one side. Despite the relatively large number of males examined (n=51), no variation in the number of megaspines on tibia II was observed.

Females from Uzbekistan and Tajikistan are also darker than those from in Iran and Turkmenistan. In the large female syntype of Ischnothele strandi no secondary receptacles could be found, even when examined from the ventral side; Fig. 4E . The only other P. raddei female that appears to lack secondary receptacles is a much smaller Ischnothele strandi syntype, the vulva of which is deformed by desiccation and thus is not illustrated. Two other females from the same locality (Akarcheshme, Turkmenistan), however, do possess secondary receptacles ( Fig. 4F–G ). These two specimens and a female from Mt. Astana ( Tajikistan; Fig. 5G ) seemingly have two secondary receptacles (not fully developed) on one of their spermathecal trunks. A duplicate of a lateral receptacle is present in the female from Tigrovaya Balka ( Fig. 5D ). All these extra receptacles occur only on one of both spermathecal trunks per vulva and are probably abnormal. The female from Varsob has a distinctly widened left spermathecal trunk ( Fig. 5H ), which is probably malformed.

Relationships. This species is most closely related to P. roxana sp. n.

Distribution: Phyxioschema raddei has the largest geographical range within the genus. This species occurs between the Caspian Sea in the west and the Tian Shan, Pamir and Karakorum mountain ranges in the east, a distance of about 1600 km (Fig. 1). Published records and material are available from: Iran (new records); Turkmenistan ( Simon 1889, Charitonov 1932, Spassky 1937, 1952, Spassky & Minenkova 1940, Charitonov 1969, Zonstein 1985, Mikhailov & Fet 1994); Afghanistan ( Roewer 1960; unconfirmed); Uzbekistan (new records); Tajikistan ( Andreeva 1968, 1976); Kazakhstan (new record); Kyrgyzstan ( Zonstein 1996, reporting on juveniles, which presumably also belong to P. raddei , from the surroundings of Batken). The type locality, Krasnowodsk, is the most western locality. The identity of specimens described under Afghanothele lindbergi and A. striatipes from Afghanistan require confirmation on the basis of new material (preferably mature males) from the corresponding type localities.

The vertical distribution of P. raddei ranges from about 200 m to 1800 m, and possibly even 2200 m. One of us ( SLZ) has also seen (but not collected) juveniles, which presumably also belong to this species, under living stones near the top of Mt. Sanglok in Tajikistan, at altitudes of 2000–2200 m .

Habitat and phenology. Phyxioschema raddei occurs in various, more or less arid habitats, from lowland sand deserts (e.g., at Repetek) and salt plains (e.g., at Krasnovodsk) to open mountain forests (e.g., at Aidere). This is quite unusual for diplurid spiders, which usually live in rather humid environments. The spiders live mainly under rocks, whereas in stone-free habitats they occupy abandoned burrows of rodents and reptiles. The males examined were collected (many of them in pitfall traps) between the beginning of May and mid- July.

Figures

FIGURE 2A–F. Phyxioschema raddei Simon, 1889, male (MNHN AR 5007, 17883) from Aïn Deia (= Aidere) (A–E) and decayed male from Repetek (F). A. Left femur I, retrodorsal view. B. Left femur II, proventral view. C. Left tibia II and proximal part of metatarsus II, retroventral view. D. Same, prolateral view. E. Left tibia II, prolateral view. F. Left tibia I showing flat ventral side, proventral view. All scale lines 1.0 mm.

FIGURE 3A–P. Phyxioschema raddei Simon, 1889, male MNHN AR 5007, 17883 from Aïn Deia (= Aidere) (A–I, L– P), male from Aidere (J), male from Damdam (K). A. Left palp, ventral view. B. Left bulb, retrolateral view. C. Left metatarsus I, ventral view. D. Same, prolateral view. E. Right tibia I and distal part of patella I, ventral view. F. Left patella I, retrolateral view. G. Left tibia II, ventral view. H. Left tibial spur, retroventral view. I. Right tibial spur, ventral view. J–K. Left tibial spur, ventral view. L. Left metatarsus II, distal view. M. Same, prolateral view. N. Same, proventral view. O. Same, ventral view. P. Same, retrolateral view. All scale lines 1.0 mm; A–B, C–D, G–P to same scale.

FIGURE 4A–H. Phyxioschema raddei Simon, 1889, vulvae of eight females, dorsal view. A. Holotype of P. raddei (AR 5009, 6686). B. Krasnovodsk (type locality of P.raddei). C. Dasht. D. Repetek. E. Largest syntype of Ischnothele strandi Spassky, 1937. F–G. Akarcheshme (type locality of I. strandi). H. Ken-Kutan. All scale lines 0.1 mm.

FIGURE 5A–H. Phyxioschema raddei Simon, 1889, vulvae of another eight females, dorsal view. A. Samarkand. B. Arys. C. Andizhan. D. Tigrovaya Balka. E. Ganjina. F–G. Mt. Astana. H. Varsob. All scale lines 0.1 mm.