Type species: Starengovia kirgizica Snegovaya, 2010: 351 (by monotypy).

Remarks: A genus of Nemastomatidae in the subfamily Nemastomatinae, presently comprising two species. Gender of genus name is female.

Emended diagnosis: Small species (up to 1.7 mm body length, females being larger than males), dorsal scutum with lines of anvil-shaped tubercles along margins of all scutal areas. Pairs of marked para-median tubercles on opisthosomal areas of dorsal scutum. Apophysis on basal cheliceral article of male well-marked but low, with a distad-directed hook, discharge area for secretion in a bowl-like excavation on medial side of apophysis. Its inner wall perforated by numerous minute pores, apparently the extrusion area for cheliceral gland secretion.

Genital morphology of males: Characterized by rather unspecialized truncus penis, moderately slender, not markedly inflated; muscle-containing inflated base of penis large, rather compact, deeply incised medially, occupying about one fourth to one third of truncus length, truncus in straight continuation of inflated base. Glans inconspicuous, small, short, not well differentiated from truncus; armament of glans simple, with few stiff spicules in a dorsally and ventrally symmetrical arrangement, stylus short, straight to slightly curved.

Distribution: Hitherto only two males of one species were known from Kyrgyzstan, a second species is here described from Pakistan. Staręga (1986) provisionally named a single female from the Tadjik Republic as “Mediostoma” pamiricum Staręga, 1986. The locality of this specimen is about half way between and approximately 360 km from Starengovia records in Kyrgyzstan and in Pakistan and thus this female may also belong to Starengovia. This assumption is backed by characters of external morphology (Fig. 6B; see Discussion).

Name: It refers to Wojciech Krzysztof Staręga (1939-2015), a distinguished Polish arachnologist who worked on systematics and taxonomy of European, Asian and African harvestmen (Żabka, 2015).

Relationships: Rows of anvil-like tubercles forming lines and encircling the dorsal scutum magnum or parts of it and all or some of the dorsal scutal areas (probably equivalents to “somites”) are present in several nemastomatine genera and are documented for Mitostoma Roewer, 1951, Carinostoma Kratochvíl, 1958 (Martens, 1978a) and Acromitostoma Roewer, 1951 (Rambla, 1983). These structures are excessively developed in all New World ortholasmatines. According to the (still incomplete) molecular-based results of Schönhofer & Martens (2012), these dorsal ornaments developed independently in at least two different lineages. Two of the relevant genera analyzed, namely Carinostoma and Mitostoma, are placed on different branches of the molecular-based tree and are not closely related.

Alternatively, this dorsal armament may have been lost at various stages of nemastomatine evolution. This assumption is backed by the fact that nearly all ortholasmatine genera and species exhibit rich and elaborate scutal ornamentations. It is reduced in the few known Asian species (Shear & Gruber, 1983; Shear, 2010; Zhang & Zhang, 2013). Thus, lines of anvil-shaped tubercles may represent a plesiomorphic character within the Nemastomatidae, an element of the nemastomatid “Bauplan”. In addition, equally shaped thorns turn up as coxal-rim denticles in most species of Nemastomatidae. If plesiomorphic, this character would not be useful to infer phylogenetic relationships.

Starengovia kirgizica Snegovaya, 2010: 352 (types not examined). – Schönhofer & Martens, 2012: 410 (discussion of phylogenetic placement, molecular-based phylogenetic tree). – Martens, 2016: 449 (discussion of phylogenetic placement). – Schönhofer, 2013: 47 (species mentioned).

Material examined: CJM 7649; 2 males; Kyrgyzstan, Fergansky Mountain Ridge, Babush Ata Mountains, above Arslanbob, at and near Yarodar Research Station, Juglans regia woodland, path to ravine, 41°19′N 72°58′E, 1400 m; H. Read leg. 17.5.1993. – CJM 6576; 4 males, 8 females; same locality, 1440-1500 m; S. Dashdamirov leg. 8.5.1990. – CJM 7650; 15 males, 14 females; same locality; W. Schawaller & J. Martens leg. 16.-18.5.1993. – CJM 7651; 6 males, 4 females; SMF; 2 males, 2 females; same locality; W. Schawaller & J. Martens leg. 15.5.1993. – ZMMU; 1 female; Kyrgyzstan, Chatkal Mt. Ridge, Sary Chelek Reserve, Suk-Bulak valley, litter, K. Mikhajlov leg. 7.7.1983. – CJM 7652; 1 male; Kyrgyzstan, Sary Chelek Nature Reserve, Tumanyak valley, open Juglans regia woodland, under stones and logs, 41°51′N 71°57′E, 1500 m; H. Read leg. 29.5.1993. – ZMMU; 2 females; same locality; K. Mikhajlov leg. 12.7.1983. – CJM 7653; 1 male, 1 female; same locality: Kodea Ata river valley, Juglans, Malus, Picea, under logs and stones, 71°57′E 41°51′N, 1500-1800 m; H. Read leg. 29.5.1993. – CJM 7654; 2 males, 5 females; MHNG; 5 males, 5 females; W. Schawaller & J. Martens leg. 29./30.5.1993. – ZMMU; 3 males, 3 females; same locality; W. Schawaller & J. Martens leg. 28.5.1993. – CJM 7655; 8 males, 3 females; same locality; W. Schawaller & J. Martens leg. 28.5.1993. – ZMMU; 1 male; same locality; D. Milko leg. 28.-31.5.1993. – ZMMU; 2 males, 8 females; same locality; S.I. Golovatch leg. 28.-31.5.1993. – ZMMU; 1 male, 1 female, same locality, Arkit, Juglans regia and Acer turkestanica, 1300 m; A.B. Ryvkin leg 3.7.1983. – ZMMU; 1 male, 1 female, same locality, Kil'tesay stream, Juglans regia, 1300 m; A.B. Ryvkin leg. 4.7.1983. – ZMMU; 1 female, same locality, Karatungun canyon, Picea schrenkiana stands, 1400 m; A.B. Ryvkin leg. 10.7.1983. – CJM 7660; 2 males, 3 females, same locality, Kil'tesay canyon, Juglans regia with Abies semenovi forest, 1300 m; A.B. Ryvkin leg. 17.9.1983. – CJM 7656; 1 male, 1 female; Kyrgyzstan, N. Sovetskoye S. Alash Mt. Ridge, Juglans regia and Crataegus woodland with grassy glades, in litter, 41°11′N 72°39′E, 1200-1550 m; H. Read leg. 26.5.1993. – CJM 7657; 2 males; same locality; W. Schawaller & J. Martens leg. 25.5.1993. – ZMMU; 1 male; same locality; S.I. Golovatch leg. 26.5.1993. – CJM 7658; 1 male; Uzbekistan, Uckargan; J. Martens leg. 27.5.1993.

Extended diagnosis: Characterized by genital morphology (form of wings of truncus penis, by size of inflated basal region of truncus), size (larger body than in S. ivanloebli sp. n.) and by armament of dorsal scutum and free opisthosomal tergites (large, conical para-median tubercles on opisthosomal areas I-V and on first free opisthosomal tergite in few specimens only).

Name: Named after the country in which the type specimens were collected.

Description (male)

Body, dorsal side (Figs 1-5): Scutum uniformly light to dark brown (depending on time past final moult), without any golden or silver markings. Anterior and lateral margins of prosomal area I of dorsal scutum with a continuous line of closely spaced anvil-shaped tubercles of different sizes, largest on front of prosoma. Additional lines of anvil-shapes tubercles across the scutum: First (anteriormost) straight, second bent forwards, third and fourth straight, fifth and sixth line bent backwards. Tubercles of fourth, fifth and sixth line smaller and more or less interrupted (only few low tubercles present) in median part only, nearly lacking laterally. Opisthosomal areas I-V each with a pair of quite large conical para-median tubercles; these tubercles posteriorly slightly further apart from each other than anteriorly. First free opisthosomal tergite in few specimens also carrying a pair of such tubercles. In addition, prosomal and opisthosomal areas with scattered low rounded tubercles.

Tu oc relativey large, low, touching front margin of dorsal scutum, covered by few strong anvil-shaped tubercles.

Body, ventral side (Figs 2, 4-5): Cx I-IV pro- and retrolaterally with a line of strong anvil-shaped tubercles, rear and front line of consecutive Cx touching each other. Cx surface coarse, covered with relatively large and closely spaced tubercles. Op gen covered by much fewer, large, low, rounded tubercles; free sternites with few tubercles at rear margins; all light brown.

Legs: Quite different in some populations. Relatively short (but variation considerable; see below: Dimensions), male Fe I and III stout and markedly spindle-shaped, in females less pronounced (Sari Chelek, Alash), or Fe I and III slender, not spindle-shaped and whole leg considerably longer (Yarodar).

In all populations Tr with several large rounded tubercles; Fe, Pt and Ti of legs unarmed except for few minute light hairs on Mt, and Ta with few scattered long hairs. Coarse surface especially on Fe to Ti. No “comb-teeth” (Kammzähnchen) as illustrated by Gruber (1976) for Mediostoma. Variable numbers of pseudo-articulations on Fe of legs II-IV.

Pedipalp (Figs 16-17): Slender and relatively long (in terms of Starengovia morphology), no article noticeably inflated; all articles except Tr bearing clavate setae, these most conspicuous on Pt, Ti and Ta; distal end of male Ti slightly curved downwards. No article with special armament in males and females.

Chelicera of male (Figs 7-15): Rather stout; basal article with strong pointed tubercles laterally and dorsally, a bulky frontad-directed Apo distinctly surpassing front margin of proximal article; Apo with a broad basis, approximately as long as high (in lateral view), upper side smoothly rounded and dorso-distally projecting into a pointed hook. In dorsal view Apo inclined anteriorly, towards longitudinal axis of article. Prolaterally Apo excavated nearly over its total length, forming a bowl-like excavation or hole. 2nd cheliceral article moderately inflated, with few long scattered bristles mostly on frontal surface.

Male genital morphology (Figs 20-26): Truncus penis (Figs 20-22) moderately slender; basis forming a large inflated part (occupying slightly less than one third of whole penis length) well differentiated from rest of truncus; inflated part compact and deeply split medially, completely filled by two penial muscles, their tendons spanning hole truncus length up to glans. Truncus parallel-sided (in do/ve view), slightly enlarged at level of wings, beyond wings narrowing toward glans; in lateral view narrowest above inflated basal part, from there slightly widening toward wings. One thin, fine hyaline and triangular wing on each lateral side in distal part of truncus, totally flattened, lateral corners slightly curved to ventral side (Figs 20-24).

Glans (Figs 25-26) only inconspicuously oultined, short, starting where two tendons are attached to inner truncus wall; stylus short, a continuation of the glans, tapering to distal asymmetrical opening of seminal duct, slightly curved (in lateral view). Short, stiff and unspecialized spinules forming armament of glans, their arrangement symmetrical in ventral view.

Female (Figs 14-15): Characters largely as in male but lacking the dorso-distal cheliceral Apo.

Measurements: Body length: Sary-Chelek and Alash: males: 1.5-1.75 (n=10), Uckargan male: 1.6. Females: 1.6- 2.2 (n=10). Leg II length, male, female in parentheses (two males from Yarodar and Uckargan, two females from Yarodar and Sari Chelek): Fe 1.1/1.5 (1.4/2.0) Pt 0.35/0.4 (0.5/0.5) Ti 0.75/1.2 (1.2/1.5) Mt 1.4/2.05 (1.75/2.4), Ta 1.1/1.6 (1.6/1.7). Pedipalp length (Sari Chelek): Fe 0.7 (0.9), Pt 0.6 (0.7), Ti 0.5 (0.6), Ta 0.3 (0.4). Penis length: 1.1.

Variation: The para-median tubercles of the dorsal scutum differ in size and shape in different populations. They are low and stout in individuals of short-legged populations (Sari Chelek, Alash), longer (up to twice as long) in long-legged populations (Yarodar). There is also variation within populations though to a generally much smaller extent. In addition, the dorsal scutum has either irregularly arranged anvil-shaped tubercles or these are partly arranged in transversal lines.

Distribution: The species is known from three localities in Kyrgyzstan and from one in Uzbekistan. The Kyrgyz specimens were collected at the Sary Chelek Biosphere Reservation about 60 km northwest of Tash-Kumyr, from the Fergansky Mountain Ridge in the Babush Ata Mountains above Arslanbob and from north of Sovetskoye in the Alash Mt. Ridge. The animals were collected in various habitats, mostly in open walnut (Juglans regia) forests, in riverine bushy stands, mostly under stones and old wood, and, mainly in the Sary Chelek area, also in various coniferous forest types. These opilionids were mostly sifted from soil litter and turned out to be quite common in all three areas. They were collected by different people independently from the same localities. Altitudinal records range from 1200 to 1800 m.

Type locality: Kyrgyzstan, Alash Mountain Ridge, Alash river valley, near Alash, 1550 m.

Remarks: Starengvoa kirgizica was originally described on the basis of two males. A more detailed description is presented here. Judging from the rich material available, S. kirgizica has considerable variation in the shape of the dorsal scutal armament and in the length of legs, which is pronounced in leg II and IV. This leg variation is not correlated with body size, which is homogeneous throughout all populations. Also genital and cheliceral morphology of males does not vary noticeably. A detailed genetic investigation should check if more than one species is involved.

Holotype: MHNG; male; Pakistan, Swat District, above Utrot, 35°30′N 72°28′E, 2500 m; C. Besuchet and I. Löbl leg. 13.5.1983.

Paratypes: MHNG; 1 male, 2 females; same locality as for holotype. – MHNG; 4 males, 17 females; Swat, Malam Jabba, 34°48′N 72°35′E, 2300 m; 9.5.1983. – MHNG; 1 male, 1 female; Swat, above Miandam, 35°03′N 72°34′E, 2300 m; 10.5.1983. – MHNG; 1 male; Swat, above Utrot, 35°30′N 72°28′E, 2500-2600 m; 14.5.1983. – MHNG; 3 males, 11 females, 1 juvenile; Swat, valley of Ushu, ascent to Kalam, 35°27′N 72°34′E, 2300 m; 15.5.1983. – MHNG; 1 male, 5 females; SMF; 2 males, 2 females; Swat, above Miandam, 2400-2500 m; 17.5.1983. – MHNG; 1 male; Swat, Malam Jabba, 2500-2600 m; 18.5.1983. – MHNG; 1 male; Chitral, Lawarai Pass, 35°19′N 71°49′E, 2600 m; 23.5.1983. – MHNG; 3 males; CJM 7659; 3 males, 1 female; Hazara, above Naran, 34°54′N 73°39′E, 2600 m; 1.6.1983. All specimens leg C. Besuchet and I. Löbl.

Diagnosis: Characterized by genital morphology (form of alae and of inflated basal part of truncus), size (species smaller than S. kirgizica) and armament of dorsal scutum (para-median tubercles peg-like, low and slender, not present on first free opisthosomal tergite).

Name: The name is given in honour of Ivan Löbl who during numerous expeditions collected large numbers of soil arthropods, especially in the Himalayas and other parts of Asia. His contributions to entomology are striking.

Description (male)

Body, dorsal side (Figs 6A, 27-30): Scutum uniformly light brown to dark brown (depending on time since final moult), without any golden or silver markings. Anterior and lateral margins of prosomal area I of dorsal scutum with a continuous line of closely spaced anvil-shaped tubercles of different sizes, largest on front of prosoma. Additional lines of anvil-shaped tubercles across the scutum: First (anteriormost) in male more or less straight (in female straight), second line bent forwards, third line mostly straight but slightly bent backwards laterally, fourth straight, fifth and sixth line slightly bent backwards. Tubercles of fourth, fifth and sixth line smaller and more or less interrupted (only few low tubercles present) in median part only, nearly lacking laterally.

Median part of opisthosomal areas I-V with two slender and low, peg-like para-median tubercles slightly inflated distally and rounded at tip; distance between paramedian tubercles slightly increasing posteriorly, all opisthosomal areas with several irregularly arranged low tubercles, very few on opisthosomal area I and on first free opithosomal tergite, absent on prosomal scutal area I around Tu oc. First free opisthosomal tergite without enlarged para-median tubercles.

Tu oc relatively small, low, touching front margin of scutum, densely covered with 10-12 anvil-shaped tubercles.

Body,ventral side (Figs 28, 30): Cx I-IV pro- and retrolaterally with a line of strong anvil-shaped tubercles, rear and front line of consecutive Cx touching each other. Op gen covered with rather large low, rounded, irregularly arranged tubercles; free sternites with few tubercles at margins; all light brown.

Legs: Rather short and slender (in terms of nemastomatid morphology); Cx with several large tubercles; Fe, Pt and Ti of legs I, III and IV slightly inflated (less so in female), Fe, Pt and Ti of leg I unarmed except for few minute light hairs, less so on Mt and Ta, there few scattered long hairs. Surface rough, no denticulation and no “comb-teeth” (Kammzähnchen) as figured by Gruber (1976) for Mediostoma. Various numbers of pseudo-articulations on femora of legs II-IV.

Pedipalp (Figs 18-19): Robust and short (in comparison with other nemastomatids), shorter than in S. kirgizica, male Ti slightly inflated proximally; all articles except Tr bearing clavate setae, most conspicuous on Pt, Ti and Ta (not indicated in Figs 18-19). In male and female no article with special armament.

Chelicera (Figs 31-37): Rather stout; basal article with a bulky frontad-directed Apo distinctly surpassing front margin of article; Apo with a broad basis, approximately as long as high (in lateral view), upper side smoothly rounded and dorso-distally projecting into pointed hook. In dorsal view Apo markedly inclined towards midline. Apo medially excavated for nearly its total length, forming a bowl-like excavation or hole. 2nd cheliceral article moderately inflated, with few long scattered bristles, situated mainly frontally.

Male genital morphology (Figs 38-43): Truncus penis (Figs 38-39) moderately slender; basis forming a large inflated part (occupying about one third of whole penis length) well differentiated from rest of truncus; inflated part compact, deeply incised in the middle and completely filled by penial muscles, their tendons spanning hole truncus length up to glans. Truncus narrowest above inflated part (in do/ve view), slightly widening toward lateral wings, beyond wings successively tapering to glans. One thin, fine and hyaline wing (Figs 38-41) on each lateral side of distal part of truncus, totally flattened, rectangular, abruptly and widely truncate, not pointed.

Glans (Figs 42-43) only inconspicuously outlined, short, starting where tendons are attached to inner truncus wall; stylus short, a continuation of glans, tapering to distal asymmetrical opening of seminal duct, slightly curved (in lateral view). Short, stiff and unspecialized spicules forming armament of glans; their arrangement symmetrical in ventral view (Fig. 42).

Female (Figs 29-30, 36-37): Largely as male but lacking dorso-distal Apo of first article of chelicera. This article armed with pointed tubercles laterally. Form of anvil-shaped lines of tubercles of dorsal side slightly different (Fig. 29 cf. Fig. 27, see also “Body, dorsal side”).

Measurements: Body length, male 1.3-1.5 (n=10), female 1.3-1.65 (n=10). Leg II length: male, female in parentheses: Fe 1.25 (1.25), Pt 0.35 (0.4), Ti 0.9 (0.9), Mt 1.5 (1.45), Ta 1.25 (1.2). Pedipalp length: male, female in parentheses: Fe 0.5 (0.5), Pt 0.45 (0.4), Ti 0.4 (0.35), Ta 0.2 (0.2). Penis length: 1.0.

Variation: One of the peg-like para-median tubercles on opisthosomal area V of the dorsal scutum may lack (Fig. 27); on the first free opisthosomal tergite (corresponding to opisthosomal tergite VI) such tubercles are absent (Fig. 6).

Distribution: Known from eight localities in northwestern Pakistan, close to the northwestern fringes of the Himalayan chain. The species was found in three major administrative units of the country, Swat and Chitral Districts and Hazara Division, all mountainous areas in the northwest of the country, just west of the upper reaches of the Indus river.

Detailed habitat descriptions by the collectors refer to soil litter in open pine (Pinus), fir (Abies), spruce (Picea) and cedar (Cedrus) forests, in moist grassy glades, under stones and rotten wood and at moist riverine localities. Near Naran animals were found on June 1, under stones close to the actual snow line at 2600 m. Altitudinal records stretch from 2100 m (Murree) to 2600 m (Utrot, Malam Jabba, Naran, Lawarai Pass) and indicate occurrence of this species in a narrow altitudinal belt of just 500 m. Most remarkably, S. ivanloebli sp. n. was collected together with two undescribed species of Biantes Simon, 1885 (Biantidae, a truly tropical family) at Malam Jabba, Naran to Kaghan, above Naran. One Kaghan series contained only five Biantes specimens, no Starengovia. The genus Biantes has undergone a tremendous radiation in the Himalayas; from Nepal alone 18 species were described (Martens, 1978b). Nearly all altitudinal zones there are inhabited by species in mostly narrow altitudinal belts stretching from the Terai lowlands to above 4000 m near the timberline. Many additional Biantidae species from Nepal and the Indian Himalayas await formal description. The Pakistani records of Biantes are the northwestern-most known within Asia. There they meet nemastomatines near their southeastern border.