Raveniola vulpina, Zonstein, 2024

Zonstein, Sergei L., 2024, A revision of the spider genus Raveniola (Araneae, Nemesiidae). II. Species from Central Asia, European Journal of Taxonomy 967, pp. 1-185 : 81-87

publication ID

https://doi.org/ 10.5852/ejt.2024.967.2699

publication LSID

lsid:zoobank.org:pub:C08B8027-50CC-417E-BCD4-5183B9FF6738

persistent identifier

https://treatment.plazi.org/id/C7957862-1670-4A16-AF15-DABDA8453319

taxon LSID

lsid:zoobank.org:act:C7957862-1670-4A16-AF15-DABDA8453319

treatment provided by

Plazi

scientific name

Raveniola vulpina
status

sp. nov.

Raveniola vulpina sp. nov.

urn:lsid:zoobank.org:act:C7957862-1670-4A16-AF15-DABDA8453319 Figs 34–35 View Figs 28–35 , 61–62 View Figs 54–62 , 76–77 View Figs 76–81 , 108 View Figs 100–108 , 134–135 View Figs 127–135 , 164–165 View Figs 160–171 , 194–195 View Figs 184–195 , 227–228 View Figs 220–228 , 254–255 View Figs 247–255 , 288–289 View Figs 282–289 , 309 View Figs 290–309 , 377–378 View Figs 364–378 , 459–465 View Figs 457–465 , 551–554 View Figs 537–554 , 616–618 View Figs 611–618 , 735–738, 760

Raveniola mikhailovi – Zonstein 2021: only figs 9–10 (♂; mismatched and erroneously used for the publication; see above).

Diagnosis

Males of Raveniola vulpina sp. nov. differ from the related male congeners by the following characters: from R. michailovi and R. virgata by a gently arcuate (vs slightly twisted) embolus, and from R. nenilini sp. nov. in having a considerably longer and thinner palpal tibia, as well as a narrower copulatory bulb ( Figs 377–378 View Figs 364–378 , 459–465 View Figs 457–465 cf. Figs 373–374, 376 View Figs 364–378 , 447 View Figs 439–447 , 458 View Figs 457–465 ). Females of R. vulpina differ from females of the related species in having the dorsal abdominal pattern lacking a clearly discernible median stripe (vs its presence; see Figs 61–62 View Figs 54–62 cf. Figs 56–58, 60 View Figs 54–62 ). Additionally, they are distinguishable due to a specific structure of the spermathecae, with broadly spaced, curved and flattened trunks and widely diverging lateral diverticula (vs differently arranged spermathecal structures in other species; Figs 551–554 View Figs 537–554 cf. Figs 538–543, 547–550 View Figs 537–554 ).

Etymology

The specific epithet ‘ vulpina ’ is the Latin adjective of ‘ vulpes ’ (= fox) meaning ‘vulpine’ and referring to the foxy ground colouration of this species.

Material examined

Holotype

KYRGYZSTAN • ♂; Chatkal Mts. (southern slope), Sary-Chelek Nature Reserve , Tumanyak Gorge, Karagailisai; 41°49.4′ N, 71°56.5′ E; 1400–1600 m a.s.l.; 5 Jul. 2000; S. Zonstein leg.; SMNH. GoogleMaps

Paratypes (18 ♂♂, 30 ♀♀)

KYRGYZSTAN • 3 ♀♀; same collection data as for preceding; SMNH 2 ♂♂; same collection data as for preceding; 41°48.7′ N, 71°55.3′ E; 1800 m a.s.l.; SMNH GoogleMaps 1 ♀; Karangitun Gorge ; 41°48′ N, 71°57′ E; 1700 m a.s.l.; 26 May 1993; S. Zonstein leg.; SMNH GoogleMaps 9 ♂♂, 15 ♀♀; Talas Mts (southern slope), Uzunahmat Canyon, Birbulak Gorge; 42°01.6′ N, 72°24.6′ E; 1400–1800 m a.s.l.; 14 Aug. 1986; S. Zonstein leg.; SMNH GoogleMaps 6 ♂♂, 11 ♀♀; Talas Mts , Tereksai Canyon; 42°10′ N, 72°21′ E; 2450– 2800 m a.s.l.; 16 Aug. 1986; S. Zonstein leg.; SMNH GoogleMaps 1 ♂; Talas Mts (northern slope), Beshtash Canyon, Kyrgolot Gorge; 42°18.3′ N, 72°20.4′ E; 2030 m a.s.l.; 18 Aug. 1986; S. Zonstein leg.; SMNH GoogleMaps .

Additional material (3 juvs)

KYRGYZSTAN • 2 juvs; Talas Mts , Otmek Valley 7 km WNW of Otmek Pass; 42°19′ N, 73°07′ E; 2800 m a.s.l.; 9 Jul. 1987; S. Zonstein and S. V. Ovchinnikov leg.; SMNH GoogleMaps 1 juv.; Kyrgyz Mts (northern slope), Kara-Balta Canyon , near the confluence of Kara-Balta and Chon-Mazar rivers; 42°24′ N, 73°46′ E; 2100–2200 m a.s.l.; 3 Jul. 1994; S. V. Ovchinnikov leg.; SMNH GoogleMaps .

Description

Male (holotype)

HABITUS. See Figs 34 View Figs 28–35 , 76. View Figs 76–81

MEASUREMENTS. TBL 12.90, CL 5.62, CW 4.82, LL 0.43, LW 0.83, SL 2.80, SW 2.38.

COLOUR. Carapace, palps and legs (except for darker brown femora) brownish orange; leg I slightly darker than other legs; eye tubercle blackish brown; chelicerae medium reddish brown; sternum, labium and maxillae light yellowish orange; abdomen yellowish brown, with almost indistinct brownish dorsal chevron-like pattern and a few small brown marks on ventral side; book-lungs and spinnerets pale yellowish brown.

CEPHALOTHORAX. Carapace and chelicerae as shown in Fig. 108 View Figs 100–108 . Clypeus and eye group as in Fig. 264 View Figs 256–264 . Eye diameters and interdistances: AME 0.14(0.20), ALE 0.27, PLE 0.18, PME 0.16; AME–AME 0.11(0.05), ALE–AME 0.09(0.06), ALE–PLE 0.07, PLE–PME 0.05, PME–PME 0.29. Anterior cheliceral edge with unmodified setae; rastellum not developed. Intercheliceral tumescence indiscernible. Each cheliceral furrow with 9 promarginal teeth and 1 mesobasal denticle. MIT indiscernible. Sternum, labium and maxillae as shown in Fig. 227 View Figs 220–228 . Maxillae with 11–12 cuspules each.

LEGS. Tibia and metatarsus I as in Figs 288 View Figs 282–289 , 309 View Figs 290–309 . Scopula: entire and distal on metatarsi I–II; entire on tarsus I; narrowly divided by setae on tarsus II; widely divided on tarsus III; vestigial on tarsus IV. Trichobothria: 2 rows of 8–9 each on tibiae, 13–15 on metatarsi, 11–13 on tarsi, 8 on cymbium. PTC I–II and PTC III–IV with 10–11 and 11–13 teeth on each margin, respectively.

SPINATION. Palp: femur d3, pd3, rd2; patella pd2; tibia d2, p3, r3, v6; cymbium d4. Leg I: femur d4, pd3, rd3; patella p1; tibia p2, pv2, r1, rv2+2M, metatarsus v2. Leg II: femur d4, pd3; patella p1; tibia p3, v8(7); metatarsus v6. Leg III: femur d4, pd3(2), rd3; patella p2; tibia d3, p3, r3, v9; metatarsus p4, r3, v10(9). Leg IV: femur d4, pd3, rd2; patella p2; tibia d4, p3, r3, v9; metatarsus p4, r4, v11. Tarsi I–IV aspinose.

PALP. Tibia, cymbium and copulatory bulb as shown in Fig. 377 View Figs 364–378 . Embolus long, tapering and slightly curved subapically ( Figs 459–462 View Figs 457–465 ).

SPINNERETS. See Fig. 616 View Figs 611–618 . PMS: length 0.25, diameter 0.11. PLS: maximal diameter 0.51; length of basal, medial and apical segments 0.87, 0.58, 0.48; total length 1.93; apical segment triangular.

Female (paratype)

HABITUS. See Figs 61 View Figs 54–62 , 77. View Figs 76–81

MEASUREMENTS. TBL 17.80, CL 6.31, CW 5.72, LL 0.59, LW 1.22, SL 3.22, SW 3.01.

COLOUR. In general as in male, with more intensely shaded dorsal abdomen and chelicerae.

CEPHALOTHORAX. Carapace and chelicerae as shown in Fig. 134 View Figs 127–135 . Clypeus and eye group as in Fig. 194 View Figs 184–195 . Eye diameters and interdistances: AME 0.13(0.21), ALE 0.25, PLE 0.20, PME 0.12;AME–AME 0.18(0.10),

ALE–AME 0.13(0.09), ALE–PLE 0.08, PLE–PME 0.08, PME–PME 0.43. Cheliceral rastellum absent. Each cheliceral furrow with 8 promarginal teeth and 1–3 mesobasal denticles. Sternum, labium and maxillae as shown in Fig. 254 View Figs 247–255 . Maxillae with 13–20 cuspules each.

LEGS. Scopula: entire and distal on metatarsi I–II; entire on palpal tarsus and tarsus I; narrowly divided by setae on tarsus II; sparse and widely divided on tarsus III; rudimentary on tarsus IV. Trichobothria: 2 rows of 8–10 each on tibiae, 14–15 on metatarsi, 12–15 on tarsi, 10 on palpal tarsus. Palpal claw with 3 promarginal teeth. PTC I–II and III–IV with 6–7 and 6 teeth on each margin, respectively.

SPINATION. Femora I–IV with one basodorsal spine and 3 dorsal bristles; palpal femur dorsally with 4 bristles; tarsi I–IV aspinose. Palp: femur pd1; patella p1(0); tibia p2, v8; tarsus v7. Leg I: femur pd1; patella p1; tibia p1(0), v7; metatarsus v7(6). Leg II: femur pd1; patella p1; tibia p2, v7; metatarsus v7. Leg III: femur pd2, rd2; patella p2, r1; tibia d1, p2, r2, v7(5); metatarsus d2, p4, r4, v12(9). Leg IV: femur pd1, rd1; patella p1, r1; tibia d1, p3, r2, v7; metatarsus d1, p4, r4, v12(11).

SPERMATHECAE. Each of paired spermathecae Y-shaped with relatively long and narrow base carrying two unevenly conformed branches: inner branch longer, wider, flattened and subapically dilated; outer branch shorter, club-like, with long, narrow and strictly sclerotised proximal neck, and with short, more or less rounded subapical section ( Fig. 552 View Figs 537–554 ).

SPINNERETS. See Figs 617–618 View Figs 611–618 . PMS: length 0.38, diameter 0.17. PLS: maximal diameter 0.69; length of basal, medial and apical segments 1.08, 0.64, 0.63; total length 2.35; apical segment triangular.

Variation

Carapace length in males (n =11) varies from 4.37 to 5.83, in females (n=14) from 4.28 to 7.29. The colouration and other key somatic characters of males and females from Talas Mts as shown in Figs 35 View Figs 28–35 , 77 View Figs 76–81 , 135 View Figs 127–135 , 165 View Figs 160–171 , 195 View Figs 184–195 , 228 View Figs 220–228 , 255 View Figs 247–255 , 289 View Figs 282–289 , 378 View Figs 364–378 . Variation in the structure of the spermathecae as shown in Figs 463–465 View Figs 457–465 , 551, 553–554 View Figs 537–554 .

Ecology

The species occurs in midland and highland montane belts, mostly in fragmentary woodlands composed of the spruce Picea schrenkiana Fisch. & C.A. Mey. and/or the fir Abies semenovii B. Fedtsch. , or in the mixed forest, with the participation of these coniferous trees and the walnut Juglans regia ( Figs 735– 738 View Figs 731–738 ). The spiders were found exclusively in their retreats under stones and logs. Along the river valleys, this species can penetrate into the subalpine and alpine zones.

Distribution

Known only from the type locality. See Fig. 760 View Figs 751–760 .

Data on natural history

Habitats

Depending on the altitude, prevailing landscape, precipitation regime and thermal conditions, all inhabited biotopes can be conditionally divided into three main zonal types. The first of these includes a harsh and periodically dry zone of foothills and low mountain ridges limited by lower and upper boundaries at altitudes of 450–500 m and 1000–1500 m, respectively. These biotopes are located mostly on loess substrate, on slopes covered with different types of xerophilic drought-resistant vegetation: from ephemeral semi-deserts to dense shrubland and sparse woodland composed of low sclerophyll trees ( Figs 619–626 View Figs 619–626 , 715 View Figs 715–722 ).

Similar to the Western Asian species of Raveniola , a higher level of species diversity in Central Asian congeners occurs in the most moist, mild and favorable midland-mountain forest zone (hosting 18 of the 29 regional species). However, compared with the western part of Asia, both the lower and the upper borders of this zone in the central part of the continent are at a higher altitude (which can be explained by the present-day continental climate of the region). Depending on the specific subregion, this zone can comprise a single and continuous area (as for example in southern Kyrgyzstan; Figs 720–721 View Figs 715–722 , 731 View Figs 731–738 ), or it can represent a mozaic of more or less isolated fragments (like those in Tajikistan and southern Uzbekistan, shown in Figs 631–632 View Figs 627–634 , 699–700 View Figs 699–706 , 708 View Figs 707–714 ). Within Central Asia, the lower boundary of this zone is at an altitude of at least 750 m (in the western Kopetdagh Mts), but usually 1000–1500 m a.s.l.; the corresponding upper limit is confined to 2000–2300 m a.s.l.

Finally, the third group of biotopes is represented by highland landscapes, with juniper and coniferous forests, subalpine and alpine meadows and meadow-steppes on the rocky slopes ( Figs 627–628 View Figs 627–634 , 651– 654 View Figs 651–658 , 677–679, 682 View Figs 675–682 , 717–718, 722 View Figs 715–722 , 738 View Figs 731–738 ). The lower edge of this zone is located at an altitude 2000–2300 m a.s.l. The highest altitudinal record for Central Asian Raveniola spp. is registered in this zone at an altitude of 3400–3700 m in Darvaz Mts, Tajikistan ( Andreeva 1975; 1976; under Brachythele sp. ).

Burrows and retreats

Sampling and direct observations on the ecology of Central Asian Raveniola spp. in the natural environment have shown that the majority of species do not dig their own burrows. Rather, these spiders exploit different natural refuges to hide themselves. In most cases, such refuges comprise cavities and hollows in the soil under stones, where the spiders thinly and scarcely line the bottom and walls with silk ( Figs 635–642 View Figs 635–642 , 647–650 View Figs 643–650 , 683–698 View Figs 683–690 View Figs 691–698 , 705–706 View Figs 699–706 , 733–734 View Figs 731–738 ). Several species, e.g., Raveniola cucullata sp. nov., are able to further deepen less suitable refuges ( Figs 633–634 View Figs 627–634 ). In contrast, some adult females belonging to R. mikhailovi , R. nenilini sp. nov., R. virgata and R. vulpina sp. nov., when inhabiting moist slopes that are devoid of suitable stones and logs (as shown in Figs 724–726 View Figs 723–730 , 731–732 View Figs 731–738 ), can settle using natural open depressions adjacent to tree trunks.

The three species of the caudata group are mostly known from wandering males, which have occasionally been found under stones, used as temporary shelter during daytime. The only known female of Raveniola redikorzevi , however, as well as all the known juvenuiles of R. caudata and R. redikorzevi , were collected from abandoned burrows of gerbils ( Rhombomys opimus (Lichtenstein, 1823) , Meriones spp. ) and tortoises ( Agrionemys horsfieldii (Grey, 1844)) . Within the virgata species group, R. ovchinnikovi has also been found to hide deeply inside abandoned burrows of small vertebrate animals. Another uncommon econiche settled by the latter species is that of the natural deep crevices in outcrops of loess substrate. Finally, all hand-collected specimens of R. fedotovi were found inhabiting screes in woodlands under the tree canopy ( Figs 708–710 View Figs 707–714 ).

Only a few Central Asian species of Raveniola are known to dig their own burrows. It is noteworthy that these burrowing species occurred closer to either the lower or the upper habitat limits of the Central Asian congeners, but are not found in the most favorable environments of the mid-mountain zone. The only species that inhabits the loess foothills of Western Tien Shan ( Fig. 715 View Figs 715–722 ), with their long dry summer period, is R. ferghanensis , known as a strictly obligate burrower. In contrast, the four highland members of the concolor group, R. afghana sp. nov., R. alajensis sp. nov., R. hirta , sp. nov. and R. karategensis sp. nov., which inhabit the subalpine juniper forests, subalpine and alpine meadows, and meadow-steppes at an altitude of 1900–3700 m, have also been collected exclusively from their burrows. Raveniola insolita sp. nov., known only from a single male, which was collected at an altitude of 3300–3400 m, can probably also be assigned to the members of the latter subgroup.

Unlike the similarly unprotected holes dug and used by members of the obligate burrowing Central Asian genus Anemesia , the burrows of a few bothrobiont Raveniola spp. appear to be arranged more simply. Compared with the burrow structure found in Anemesia spp. (see Zonstein 2018b: figs 330–367), these burrows are almost similarly deep (ca 30–40 cm in length), but are noticeably narrower in their median part and much more scarcely silk lined (except for the terminal living chamber, which is more densely lined). The open entrance mostly lacks the silk lining and resembles a hole used by other terrestrial arthropods (the bothrobiont coleopterans, e.g., Lethrus spp. , or woodlice; see Figs 657 View Figs 651–658 , 716 View Figs 715–722 ). Sometimes this entrance is partially camouflaged by the surrounding detritus or vegetation ( Figs 655–656 View Figs 651–658 , 661 View Figs 659–666 ). A more complex variant of an entrance arrangement was observed in R. karategensis sp. nov., in which the burrow mouth was visibly silk lined and provided with a low rim ( Figs 659–665 View Figs 659–666 ). The expanded living chamber always has a horizontal extension, as shown in Fig. 666 View Figs 659–666 . In all cases, no lateral chambers connecting the burrow shaft with the soil surface were observed.

Phenology

Data on the phenology of the congeners are sparse and based completely on fragmentary field observations. The collecting data indicate that in localities with a dry summer period the wandering adult Raveniola males occur in April–May (the most wet and favourable period). In some species inhabiting this type of biotope, a second appearance of wandering males, connected with the end of a dry period in October– November, can be observed (particularly, in R. cucullata sp. nov., R. ferghanensis , and R. kopetdaghensis ). Several congeners, such as R. ignobilis sp. nov., R. subornata sp. nov. and R. ovchinnikovi sp. nov., are known as species with males collected only in October. In highland members of the genus, e.g., in R. alajensis sp. nov. and R. hirta sp. nov., the peak of their activity, including the presence of wandering adult males, unsurprisingly moves into the mid-summer (July). Males of the most mesophilic species, like R. virgata , within the humid areas (such as the midlands of the Ferghana Mts in the environs of Arslanbob, with summer rainfalls) can be almost evenly found from mid-April to late October.

Females with egg sacs can be found in July ( R. alajensis sp. nov., R. cucullata sp. nov., R. dolosa sp. nov., R. karategensis sp. nov., R. mikhailovi , R. pamira sp. nov., R. sororcula sp. nov. and R. tarabaevi sp. nov.), or in July and August ( R. virgata , R. vulpina sp. nov.). The egg sac usually reaches 12–19 mm in diameter and contais 25– 45 eggs.

Feeding

Some currently incomplete data, which nevertheless appear to fit the general trends, have been obtained for only two Central Asian species of the genus: Raveniola ferghanensis and R. virgata . These are only generalized data on the composition of the prey remains; no quantitative counts were made. The corresponding data for other regional congeners are fragmentary and not comparable to each other, and thus are not considered below.

In Raveniola ferghanensis , a major part of the “kitchen leftovers” retrieved from the bottom of the investigated burrows was represented by the head capsules and other fragmented parts of foraging ants ( Hymenoptera , Formicidae ), including the wandering predatory foragers Catagliphus aenescens (Nylander, 1849) , and the marching carpophagous ants Messor muticus (Nylander, 1849) and M. aralocaspius (Ruzsky, 1902) . The minor part of the chitinous remains, observed as leftovers in most of the studied burrows, belonged to burrowing arthropods, which share these biotopes with R. ferghanensis : the xerophilic terrestrial woodlice Hemilepistus fedtschenkoi (Uljanin, 1874) ( Isopoda , Agnaridae ) and the colonial coleopterans Lethrus spp. ( Coleoptera , Geotrupidae ). The latter remains tended to belong to either L. sulcipennis Kraatz, 1883 or L. micronatus Semenov, 1894 , or both sympatric species, which inhabit the same biotopes. Finally, some of the burrows contained remains of Madotrogus sp. aff. ferganensis (Protzenko, 1962) ( Coleoptera , Scarabaeidae ), as well as of some other unidentified representatives of Coleoptera .

Regarding Raveniola virgata , there are somewhat more complete data on the composition of their prey. Similar to the above, most of the prey remains were represented by a few ant species ( Hymenoptera , Formicidae ), but in this case by ants wandering through or nesting within the forest floor: Camponotus reichardti K. Arnoldi, 1967 , Messor rufus Santschi, 1923 and Myrmica juglandeti K. Arnoldi, 1976 . Most of the other collected and studied remains belonged to the following representatives of the Coleoptera : Carabidae : Chilotomus sp. aff. uzgentensis Shauberger, 1932 , Eocarterus uzgentensis Heyden, 1884 , Harpalus sp. , Leistus sp. aff. ferganensis Semenov & Znojko, 1928, Poecilus sp. , Pterostichus sp. aff. sodalicius Heyden, 1885, Trechus sp. ; Curculionidae : Asiodonus sp. , Polydrosus sp. ; Elateridae : Selastomus sp. ; Glaphyridae : Amphicoma sp. ; Staphylinidae : Philonthus sp. ; Tenebrionidae : Laena sp. , Zophohelops tiro (Reitter, 1902) . The remains of Ectobius delicatulus Bei-Bienko, 1950 (Blattoidea: Ectobiidae ) were more frequent, while fragments of Hessebius plumatus Zalesskaya, 1978 , Monotarsobium sp. ( Chilopoda, Lithobiidae ) and remnants of Dysdera sp. aff. arnoldii Charitonov, 1956 ( Araneae , Dysderidae ) were found in a few isolated cases.

Mating

Regarding Central Asian Raveniola spp. , fairly satisfactory data are available only for representatives of R. virgata kept in captivity (for details see Zonstein 2002c). The corresponding observations were carried out on October 20–22, 1992. To imitate the natural environment, observations were made at night, under dim lighting, in large cages for each pair, allowing greater mobility of males and females. Each female was placed in its cage one day before the male, enabling the females to find or to dig a refuge (which mostly resembled an open cavity between lumps of soil). After introducing a male into each cage, the sequence of events in most cases was as follows.

Having discovered the entrance to the female’s refuge, the male began to tap with the tips of the palps on the substrate, demonstrating a short courtship stage. When the female appeared, within a few seconds the male made several pedalling movements with the palps (resembling the pedalling of a cyclist). The male then stretched forward its first pair of legs, moved towards the female and approached her (see Figs 739–741 View Figs 739–746 ). This is followed by touching the female with the tips of the male’s legs I for 3–4 seconds ( Fig. 742 View Figs 739–746 ). Concurrently, the male’s legs II were also raised off the ground surface and began to vibrate, tapping the female’s legs.

A few seconds later, the male used its megaspines to grab the trochanters or femora of the female’s palps, gripping them on their proventral side ( Figs 743–744 View Figs 739–746 ). The male then bent his first pair of legs at the tibiometatarsal joint and extended them upward. The male thereby closed the clamps, clasped the female’s palps, and lifted the female’s cephalothorax upward, so that the latter formed an angle with her abdomen, sometimes reaching almost 90° ( Figs 745–746 View Figs 739–746 ). The entire operation lasted no more than 2–5 seconds. Following attachment, one of the emboli entered the spermathecal openings.

During each recorded phase of mating, the longest of them was the copulation process itself (from the initial insertion of the embolus into the copulatory organs of the female until its final removal). This stage is considered hereafter as the duration of copulation. All other stages – approach, initiation, coupling and uncoupling of the partners – took several seconds each.

The data obtained from laboratory observations on the copulation behavior in 20 pairs of R. virgata indicate that the duration of a basic insertion/removal act in this species varies from 27 seconds to 13 min. and 20 sec., averaging 6 min. and 13.2 sec. Usually, the entire copulation process ended with this single act. However, in six pairs a continuation was observed, associated with a one-time change of the involved palp. Even more rarely (three observations), a quadruple insertion of the emboli was detected. In the latter case, the total duration of copulation, which averaged about 10 min. for the examined specimens, increased up to 14–19 minutes. During copulation, the male continued to move towards the female, turning her onto her dorsal side. Having completed copulation, the male removed the involved embolus. Then, pressing on the female palpal femora, the male knocked the female over, climbing over her before the female could roll back.

According to a few observations of captive representatives of R. cucullata sp. nov., R. ferghanensis and R. nenilini sp. nov. (one, one, and two observed but not documented encounters, respectively), the courtship and copulation in these species occured in a similar way. In the natural environment, a copulating pair of R. virgata has incidentally been observed only once (see Zonstein 1987). In addition, this took place during the daytime, which was somewhat surprising, because such behavior cannot be considered characteristic for either nemesiids, or for mygalomorph spiders in general.

Predators

All known data on the predators interacting with and feeding on the Central Asian species of Raveniola refer only to a few taxa of spider wasps ( Hymenoptera , Pompilidae ). The pompilids that hunt and prey on these members of Raveniola belong to the pompiline genera Pareiocurgus Haupt, 1962 and Pamirospila Wolf, 1970 . Pareiocurgus latigena (F. Morawitz, 1893) is known from Uzbekistan, Tajikistan and southern Kyrgyzstan. These wasps feed on several species of burrowing spiders, including R. ferghanensis and representatives of Anemesia ( Zonstein 2000b) . All five species of the endemic Central Asian genus Pamirospila (surveyed by Zonstein 2000b, 2002a, 2002b) are probably specialized predators feeding only on Raveniola spp. Pamirospila is confined to the same range as that of Raveniola in Central Asia. Additionally, at least twice, females of P. pamira (Haupt, 1930) have each been observed leaving a burrow of R. alajensis sp. nov., with the living chamber littered with soil lumps and with a paralyzed spider inside. A similar interaction has also been noted for R. hirta sp. nov. ( Zonstein 2000b).

SMNH

Department of Paleozoology, Swedish Museum of Natural History

V

Royal British Columbia Museum - Herbarium

Kingdom

Animalia

Phylum

Arthropoda

Class

Arachnida

Order

Araneae

Family

Nemesiidae

Genus

Raveniola

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