Genus Crossopriza Simon, 1893

Crossopriza Simon, 1893: 476 (

type species: Artema pristina Simon, 1890).

Ceratopholcus Spassky, 1934: 361 (type species: C. maculipes Spassky, 1934). Synonymized in Huber et al. 2014a: 420.

Tibiosa González-Sponga, 2006: 10 (type species: T. caracensis González-Sponga 2006). Synonymized in Huber 2009b: 65; see also Huber & Villarreal 2020: 57.

Crossopriza – Wiehle 1933: 241. — Huber et al. 1999: 7. — Deeleman-Reinhold & van Harten 2001: 194.

Diagnosis

Long-legged pholcids with spotted leg femora and tibiae (e.g., Figs 558, 577, 709), abdomen dorsoposteriorly often angular or pointed (e.g., Figs 394, 398, 699), females with paired stridulatory apparatus between carapace and abdomen (Figs 468, 595, 660, 797). Distinguished from similar genera ( Maghreba gen. nov., Stygopholcus, Holocnemus) by combination of: male chelicerae with only one pair of modified (cone-shaped) hairs at tips of main cheliceral apophyses (e.g., Figs 598, 845, 859; in contrast to Stygopholcus, H. reini, and H. caudatus); male palpal coxa with rounded retrolateral hump, without distinct process (in contrast to Maghreba); male palpal femur without retrolateral proximal process (in contrast to Stygopholcus and Maghreba); male palpal femur without dorsal process (in contrast to Maghreba); male palpal tarsal organ capsulate (e.g., Figs 426, 602, 849, 862; in contrast to Stygopholcus); procursus usually with distinct prolateral hump set with numerous hairs (e.g., Figs 357, 376, 616, 628); procursus tip not strongly bent towards dorsal (in contrast to Maghreba); procursus usually without transparent membranous process at tip (in contrast to Maghreba; present in C. soudanensis); genital bulb without slender dorsal process (in contrast to Stygopholcus); distal (main) bulbal sclerite without deep retrolateral pocket (in contrast to Maghreba).

Description

Male

BODY. Total body length ~2.5–6.5; carapace width ~1.0–2.4. Carapace with deep central pit and pair of shallow furrows diverging from posterior side of pit toward posterior rim of carapace (cf. Figs 468, 595); ocular area slightly raised, eye triads relatively close together (distance PME–PME = 0.5–1.2 × PME diameter), each secondary eye (especially PME) accompanied by small elevation (arrows in Fig. 467; “pseudo-eyes”; cf. Huber 2009a), sometimes very distinct (i.e., reflecting light), e.g., in C. miskin sp. nov. PME oval; AME large (~70–110% of PME small diameter). Clypeus high, usually unmodified, in C. sengleti sp. nov. with rounded median process (Fig. 768), in C. johncloudsleyi Deeleman-Reinhold & van Harten, 2001 with pair of small hooked processes (Deeleman-Reinhold & van Harten 2001: fig. 14; Huber 2009a). Abdomen oval, dorsally posteriorly usually angular (Fig. 394), sometimes rounded (Fig. 476) or pointed (Fig. 699). Male gonopore with 4–6 epiandrous spigots (sometimes asymmetric: 2+3; Figs 420, 507, 606, 854), ALS with only two spigots each: one large widened spigot and one pointed spigot (Figs 424, 505, 607); PMS with two spigots each (Figs 424, 505, 607); PLS without spigots.

COLOR. In general ochre-yellow to light brown. Carapace mostly pale, with darker median mark, without lateral marks (cf. Figs 391, 481, 702); sternum light brown to dark brown, with darker radial marks. Legs without or with indistinct darker rings, with oval to short longitudinal line-shaped dark marks on femora and tibiae (cf. Figs 558, 577, 709), sometimes also a few on metatarsi, rarely on femora only. Abdomen usually with distinct dorsal and ventral patterns: dark heart-mark and further dark and whitish marks dorsally and laterally, ventral median band variably distinct, rarely absent (e.g., C. sengleti sp. nov.). Cave-dwelling species slightly lighter/paler (see C. moqal sp. nov.; C. kittan sp. nov.).

CHELICERAE. Chelicerae with one large modified (cone-shaped) hair on each main cheliceral apophysis (Figs 598, 845, 859); main cheliceral apophyses usually in lateral position, in some species moved to frontal position; in latter case sometimes with additional pair of lateral apophyses (without modified hairs; e.g., Figs 731, 774, 814). With fine to distinct stridulatory ridges (Figs 421, 500, 599, 841), distances between ridges ~3.5–8.0 µm, sometimes variable within file (distances proximally larger than distally; see C. sahtan sp. nov., C. maculipes).

PALPS. Coxa with rounded retrolateral-ventral hump, without distinct apophysis; trochanter barely modified, slightly protruding ventrally; femur distally widened, on ventral side usually protruding, without dorsal apophysis, without proximal retrolateral process, without or with indistinct transversal dark line on retrolateral side, with stridulatory pick (modified hair) on prolateral side (Fig. 842); femurpatella joints shifted toward prolateral side (arrows in Fig. 354); tibia-tarsus joints shifted toward retrolateral side (arrows in Fig. 356); palpal tarsus without dorsal macrotrichia, palpal tarsal organ capsulate (Figs 426, 602, 849, 862); procursus dorsally with straight or weakly curved hairs; procursus with prolateral process set with numerous long hairs, without ventral ‘knee’, distally usually with ventral sclerite (absent in C. miskin sp. nov.), usually without membranous transparent process (present in C. soudanensis; Fig. 358), tip of procursus not curved towards dorsal, with one or more transparent hair-like or spine-like processes on retrolateral side (Figs 417, 506, 596, 847, 857); genital bulb with basal sclerite connecting to tarsus (bs in Fig. 361), and distal (main) sclerite often with distinctive set of prolateral (slightly ventral) apophyses and ridges (e.g., Figs 406, 440, 716, 789), without retrolateral pocket; sperm duct opening in membranous area on prolateral side of distal bulbal sclerite (arrows in Figs 419, 597, 848, 858).

LEGS. Long and relatively thin, leg 1 length ~20–65, tibia 1 length ~5.5–18, tibia 2 longer than tibia 4 (~1.1–1.3 ×). Tibia 1 L/d usually ~50–80,> 80 in the slightly troglomorphic C. kittan sp. nov. Femur 1 usually thicker than other femora; femur 1 with spines ventrally in one row, in very small males rarely without spines; spines proximally gradually transforming into regular setae; spines rarely present on femur 2 ( C. sanaa sp. nov.); legs without curved hairs; with few short vertical hairs; retrolateral trichobothrium in proximal position (at 2–5% of tibia length in tibia 1), prolateral trichobothrium absent on tibia 1, present on other tibiae. Tarsal pseudosegments very indistinct except a few (~2–3) distally, proximally with indistinct irregular platelets rather than distinct rings. Tarsal organs of legs capsulate, with round or weakly undulating rim (Figs 428–431, 863).

Female

In general similar to male; chelicerae either with indistinct and small stridulatory files (with slightly larger distances between ridges than in males; 4.0–8.5 µm) or without stridulatory files (Figs 422, 469, 501, 600, 843, 861); legs slightly shorter than in male, without spines. Usually with pair of variably distinct processes posteriorly on carapace (arrows in Figs 468, 595) acting against pair of variably distinct plates on abdomen (Figs 660, 797), absent in C. johncloudsleyi . Epigynum usually consisting of large, simple anterior plate and short but wide posterior plate; anterior plate usually with pair of distinct pockets, sometimes close together on median elevated ridge, sometimes more like furrows than pockets; epigynum usually without processes, only in C. johncloudsleyi with two pairs of distinct apophyses (Deeleman-Reinhold & van Harten 2001: fig. 11; Huber 2009a: fig. 30); without bulging areas in front of anterior plate. Internal genitalia with sclerotized arc that consists of dorsal and ventral parts (da and va in Figs 388, 447) and is variably visible in uncleared specimens; uterus externus sometimes with small median ventral structure (pouch or pocket?), sometimes visible as round structure in untreated specimens (e.g., Figs 463, 551, 612, 837); pore plates large, flat, of variable shape and position, pores either homogeneously distributed or in groups.

Distribution

All species except C. lyoni are restricted to an area that ranges from Mali to India and from Kenya to Kazakhstan (Fig. 351). The type species C. lyoni has expanded around the globe (Fig. 351), apparently since approximately the second half of the 19 th century (see below).

Relationships

Together with Holocnemus, Stygopholcus, and Maghreba gen. nov., Crossopriza is an unambiguous representative of the spotted-leg clade, but beyond that the cladistic analysis provides only weak evidence for inter-generic relationships. Even the monophyly of Crossopriza is poorly supported by a single character of questionable strength (Appendix 3, char. 18). Within Crossopriza, a group of 14 species (including the type species) is reasonably well supported by two functionally related characters (medially-directed male cheliceral apophyses and medially positioned female epigynal pockets). Within this group, a further sub-group of eight species (again including the type species) share a second pair of male cheliceral apophyses. Finally, C. lyoni shares with three other species a ventral sclerite on the procursus provided with a retrolateral side branch (arrows in Figs 769, 788, 810, 828). The few remaining sister-group relationships suggested by the cladogram are either weakly supported or of little relevance, or both.

Composition

The genus now includes 24 named species, of which 18 are newly described. All named species are treated below except for C. johncloudsleyi which was redescribed in Huber (2009a). Several additional undescribed species are present in collections. The ZMMU has an undescribed species from Pakistan (N of Islamabad, 33.75° N, 73.06° E), which is not formally described because the only available male is in very poor condition. The ZFMK (Ar 22391) has two females from Iran (Hormozgan, Siyahu; 27.75° N, 56.34° E) that resemble C. miskin sp. nov. in habitus, size, and epigynum, but with the epigynal pockets wider apart. For further undescribed species that are very similar to species (re)described herein, see under C. dhofar sp. nov., C. khayyami sp. nov., and C. maculipes .

Natural history

Most species have been collected in sheltered spaces under rocks, in small cavities of the ground, and in the twilight area of caves. A few species build their webs in more exposed habitats, among rocks and on plants (e.g., C. tiwi sp. nov.). Others are occasionally or regularly found in houses (e.g., C. semicaudata, C. pristina, C. maculata, C. lyoni). Little is known about the biology of Crossopriza beyond these basic microhabitat data (see individual natural history sections below). Only the cosmopolitan C. lyoni has been studied in some detail, including development and prey capture (see below).

Identification

A key that works for both males and females proved difficult to construct.At the same time, a combination of geography (Figs 351–353) and diagnostic characters (mainly male palp, male chelicerae, female epigynal plate) usually makes identification relatively quick and easy.