EUROPEAN ARACHNOLOGY 2003 (LOGUNOV D.V. & PENNEY D. eds.), pp. 185–200. © ARTHROPODA SELECTA (Special Issue No.1, 2004). ISSN 0136-006X (Proceedings of the 21st European Colloquium of Arachnology, St.-Petersburg, 4–9 August 2003)

Fauna and populations of the petrophilous spiders (Arachnida: Araneae) of north-east Siberia and north-west Canada

Ôàóíà è íàñåëåíèå ïåòðîôèëüíûõ ïàóêîâ (Arachnida: Araneae) ñåâåðî-âîñòîêà Ñèáèðè è ñåâåðî-çàïàäà Êàíàäû

YU.M. MARUSIK Þ.Ì. ÌÀÐÓÑÈÊ

Institute for Biological Problems of the North, Portovaya Str. 18, Magadan, 685000, Russia. email: [email protected] Èíñòèòóò áèîëîãè÷åñêèõ ïðîáëåì Ñåâåðà ÄÂÎ ÐÀÍ, Ïîðòîâàÿ 18, Ìàãàäàí 685000, Ðîññèÿ. email: [email protected]

ABSTRACT. Spiders associated with stony biotopes were studied in different parts of Magadan Area (Russia) and in Yukon Territory (Canada). Many of them are widespread in north-east Siberia and inhabit the tundra and taiga zones, as well as different altitudinal belts. Some species are restricted to the taiga zone, others to the tundra. The most common stony debris dwellers are Sibirocosa kolymensis, S. subsolana, Aculepeira carbonarioides and Lathys alberta. The most common dominants of the stony debris habitats in the forest belt of the taiga zone are Theridion sibiricum and L. alberta. Those of the mountain tundra belt are S. kolymensis and A. carbonario- ides. Some species were found in one or two sites of very small size (Hybauchenidium holmi, Theridion thaleri, Chalcoscirtus carbonarius, ‘Lepthyphantes’ epigynatus). Brief reviews of the geographic and spatial distribution of the lithobiont and lithophilous (= petrophilous) spider species are given. The species composition of spiders living on rock debris in Yukon Territory is rather similar to that in north-east Siberia, while both faunas differ greatly from those of central European scree slope ecosystems. The European and Beringian faunas have only two species in common. Values for the narrow endemics among the petrophilous and the exclusively lithobiont species in north-east Siberia reach 24% and 46%, respectively. The value for endemic species in the whole fauna is nine percent.

ÐÅÇÞÌÅ. Ïàóêè ñâÿçàííûå ñ êàìåííûìè îñûïÿìè è ðîññûïÿìè èçó÷àëèñü â ðàçíûõ ÷àñòÿõ Ìàãàäàíñêîé îáëàñòè è òåððèòîðèè Þêîí. Ìíîãèå èç íèõ øèðîêî ðàñïðîñòðàíåíû íà Ñåâåðî-Âîñòîêå è íàñåëÿþò òóíäðîâóþ è òà¸æíóþ çîíû, à òàêæå ðàçíûå âûñîòíûå ïîÿñà. ×àñòü âèäîâ ïðèóðî÷åíà ê îäíîé èç çîí è ñâÿçàíà ñ îäíèì èç âûñîòíûõ ïîÿñîâ. Íàèáîëåå ðàñïðîñòðàíåííûå è ìàññîâûå âèäû â êàìåííûõ ñîîáùåñòâàõ Sibirocosa kolymensis, S. subsolana, Aculepeira carbonarioides è Lathys alberta.  ëåñíîì ïîÿñå â çîíå òàéãè íà êàìåííûõ îñûïÿõ äîìèíèðóþò Theridion sibiricum è L. alberta. Äîìèíàíòàìè ãîðíîòóíäðîâî- ãî ïîÿñà ÿâëÿþòñÿ S. kolymensis è A. carbonarioides. Íåêîòîðûå èç âèäîâ áûëè íàéäåíû âñåãî â îäíîì–äâóõ íåáîëüøèõ áèîòîïàõ (Hybauchenidium holmi, Theridion thaleri, Chalcoscirtus carbonarius, ‘Lepthyphantes’ epigynatus). Äëÿ âñåõ îáëèãàòíûõ ëèòîáèîíòîâ è íàèáîëåå ìàññîâûõ ïåòðîôèëüíûõ âèäîâ ïðèâåäåíû âèäîâûå î÷åðêè, âêëþ÷àþùèå ãåîãðàôè÷åñêîå ðàñïðîñòðàíåíèå è áèîòîïè÷åñêóþ ïðèóðî÷åííîñòü. Âèäîâûå ñïèñêè ïàóêîâ êàìåííûõ îñû- 186 EUROPEAN ARACHNOLOGY 2003

ïåé íà ñåâåðî-çàïàäå Êàíàäû è ñåâåðî-âîñòîêå ñîäåðæàò îêîëî 50% îáùèõ âèäîâ, íî ñèëüíî îòëè÷àþòñÿ îò ïàóêîâ êàìåíèñòûõ áèîòîïîâ öåíòðàëüíîé Åâðîïû. Äîëÿ óçêèõ ýíäåìèêîâ ñðåäè ïåòðîôèëüíûõ âèäîâ è îáëèãàòíûõ ëèòîáèîíòîâ íà ñåâåðî-âîñòîêå Àçèè ñîñòàâëÿåò 24% è 46%, â òî âðåìÿ êàê îáùåå ÷èñëî ýíäåìèêîâ â ôàóíå — îêîëî äåâÿòè ïðîöåíòîâ.

KEY WORDS: Spiders, Araneae, north-east Siberia, Yukon Territory, scree, stony biotopes, petrophilous, lithobiont. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: Ïàóêè, Araneae, ñåâåðî-âîñòîê Ñèáèðè, òåððèòîðèÿ Þêîí, îñûïü, êàìåíèñòûå áèîòîïû, ïåòðîôèë, ëèòîáèîíò.

Introduction and no special preferences were given to the litho- biont spiders. After the first two years of research, Although stony ecosystems (i.e., screes, the petrophilous spiders attracted more of my atten- tion because they contained many interesting spe- stony taluses, stone debris, pebble river banks, cies. These studies were described in my Ph.D. etc.) are very common and widespread through- thesis [Marusik, 1988]. In 1988 and 1989, the petro- out Siberia, there are no publications specifi- philous spiders were investigated in Chukotka, viz., cally dealing with their spider faunas. A number in Anadyr’, Christ Bay, the Amguema and Bol’shaya of papers, nevertheless, contain data on habitat Osinovaya River basins. In 1985–1993, some inves- preferences for certain Siberian and Nearctic tigations were carried out in the northern Cisokho- species, including petrophilous spiders, partic- tia, Ola River basin (the upper and middle reaches) ularly from north [Eskov, 1985; Tanasevitch, and the upper-middle Kolyma River (Map 1). In 1985] or middle [Eskov, 1988] Siberia and 1993 limited research was performed in Yukon Ter- ritory of Canada. A list of the stony habitats and the from the mountains of south Siberia [Logunov sites studied in north-east Siberia is given below: & Marusik, 1995, 1999, 2001; Logunov et al., 1–6. The upper reaches of Kolyma River (62°N), 1998; Marusik et al., 2000; etc.]. Spiders asso- Sibit-Tyellakh River basin (left tributary of Kolyma ciated with cliffs in Québec were briefly dis- River), the environs of Aborigen Research Station cussed by Koponen [1992]. In contrast to Sibe- (A): ria and North America, the spiders of stony 1. Screes of a southern slope sometimes with debris (= scree slope) ecosystems in central mats of Cetraria lichens, 550–700 m. Europe are well studied thanks to numerous 2. The slatestone scree on an exposed north- publications by Rùžièka [1988, 1990a, 1993, facing slope, big stones covered with lichens, 550– 600 m. 1996, 2002, etc.] and his co-authors. 3. Granite screes and kurums1 on an exposed Stony ecosystems, morphologically and geo- south-facing slope near a flat plain in the mountain logically belong to many types (see Figs 1–6) tundra, 1 100–1 250 m. and are common both in north-east Asia and in 4. Granite-slatestone screes on an exposed east- north-west Canada. Both these territories make facing slope, 1 300 m. up parts of Beringia, a vast area between the 5. Slatestone stone field on a flat crest, 800 m. Kolyma River in the Russian Far East to the 6. Small and elongate (0.5–1.5 m wide) rock Mackenzie River in the north-west Territories debris on a south-facing slope within a semiclosed of Canada. The purpose of the present paper is birch grove, margins fixed by Graminaceae and Rosa bushes, with a small brook underneath, 600 m. to provide an account of the fauna and popula- 7 K. The upper reaches of Kolyma River, Kon- tions of the petrophilous spiders of Beringia. taktovy Creek basin, 61°51¢N, 147°40¢E, granite scree with lichen pillows on the south-facing slope, Material and methods 900 m (Figs 5, 6). 8 MA. Different granite screes and rocks around Investigations of the spiders inhabiting stony Magadan (Figs 1, 2). ecosystems in north-east Siberia were started in 9 O. Ola Plateau, klippe scree, down moist edge, 1983. The first and most detailed research was un- 1 200 m. dertaken around the Aborigen Biological Station in 1 Kurums in Siberia means the boulder screes, some- the upper reaches of Kolyma River (62°N). It was times with huge blocks (0.5–2 m), common in the moun- only a small part of the spatial distribution research tain tundra. Yu.M. Marusik. Petrophilous spiders of Beringia 187

Figs 1–6. Different stony habitats of north-east Siberia: 1 — the stony desert near Magadan; 2 — the stony desert and talus near Magadan; 3 — a small scree on the north exposed slope near the Kontakt Field Station; 4 — the stony desert on the Kolyma-Okhotian watershed; 5 — the stony talus in the mountain tundra belt near the Kontakt Field Station; 6 — the stony polygons in the mountain tundra belt near the Kontakt Field Station. Ðèñ. 1–6. Ðàçëè÷íûå êàìåííûå ìåñòîîáèòàíèÿ ñåâåðî-âîñòî÷íîé Ñèáèðè: 1 — êàìåííàÿ ïóñòûíÿ â îêðåñò- íîñòÿõ Ìàãàäàíà; 2 — êàìåííàÿ ïóñòûíÿ è ðîññûïü âáëèçè Ìàãàäàíà; 3 — ìàëåíüêèå êàìåííûå îñûïè íà ñåâåðíîì ñêëîíå â îêðåñòíîñòÿõ ñòàöèîíàðà «Êîíòàêò»; 4 — êàìåííàÿ ïóñòûíÿ íà Îõîòñêî-Êîëûìñêîì âîäîðàçäåëå; 5 — êàìåííûå îñûïè â ãîðíûõ òóíäðàõ âáëèçè ñòàöèîíàðà «Êîíòàêò»; 6 — êàìåííûå ïîëèãîíû â ãîðíûõ òóíäðàõ âáëèçè ñòàöèîíàðà «Êîíòàêò». 188 EUROPEAN ARACHNOLOGY 2003

Map 1. Main study areas. 1 — Magadan, 2 — Ola Plateau, 3 — Kontakt, 4 — Aborigen, 5 — Seimchan, 6 — Christ Bay, 7 — Kluane Lake, 8 — Carmacks. Êàðòà 1. Îñíîâíûå ðàéîíû èññëåäîâàíèé. 1 — Ìàãàäàí, 2 — Îëüñêîå Ïëàòî, 3 — Êîíòàêò, 4 — Àáîðèãåí, 5 — Ñåéì÷àí, 6 — çàëèâ Êðåñòà, 7 — îç. Êëóýéí, 8 — Êàðìàêñ.

10 S. Environs of Seimchan, slatestone scree in Stony biotopes are very diverse and cover a canyon, 600 m. huge areas. North-east Siberia, like other Arctic 11 E. Christ Bay, Egvekinot Town, south-facing and Subarctic territories, has landscapes known slope in canyon along a small creek. as cold goltsy deserts (fell-fields, alpine stony 12 E. same locality, kurums, and big stony de- deserts and rock deserts are their American bris on the east-facing slope. equivalents) [see Kuvayev, 1985]. In some ar- 13 CH. Cumulative data of Amguema River eas of the taiga belt, the cold stony deserts basin, Christ Bay and the upper reaches of Bol’shaya Osinovaya River. occupy more than 90% of the surface (e.g., in All specimens were collected either by hand or the Kolyma upland), and they are very common by using an aspirator. In population studies, species in the mountainous regions of Chukotka. were attributed to different dominant categories af- Other quite common stony habitats are the ter the criteria suggested by Palmgren [1930, 1972] stony slopes (scree slopes, screes or talus) with- for birds and spiders: viz., dominants or D, > 10% of in the taiga belt composed of small (less than specimens collected in a certain habitat; subdomi- 0.5 m blocks) or large stones. In many cases, nants or S, 5–10%; common (or influents) or C, 2– the screes have independent origins, but quite 5%; and others rare or R. Spiders of different preda- often they are formed by stony desert tongues. tion guilds such as wandering (Lycosidae, Gna- In continental highlands, as well as in coastal phosidae, Philodromidae Liocranidae, Salticidae), parts, the tongues of the stony deserts can reach ambush spiders (Thomisidae) and web builders were the valley bottoms or the sea coast. attributed to dominant categories independently, Besides the two most widespread kinds of because they required different collecting methods. stony ecosystems mentioned above, the follow- ing habitats are also present in the upper Koly- Stone debris ecosystems in north- ma: kurums, stone polygons and belts in moun- east Siberia tain tundra, cliff screes (mountain tundras), for- est debris, small stone taluses on low and flat While there are some similarities between mountain ridges and trains, and different kinds morphological types of the stony ecosystems in of morains. middle Europe (particularly in the Czech Re- The kurums in Siberia mean the boulder public, see Rùžièka [1990a, 1993]) and Siberia, screes, sometimes with huge blocks of 0.5–2 m the differences are more discernable, especially in diameter, mostly occurring in the mountain as far as north-east Siberia is concerned. tundra. Stone polygons (circles) and belts are Yu.M. Marusik. Petrophilous spiders of Beringia 189 characteristic for the tundra and in some places from one area to another, and recent ecosystems for the mountain tundra, where they were formed which are closely separated can have gradual because of the permafrost. species interchanging. The characteristic aspects of the goltsy deserts, which separate them from the tundra Spiders of the stony biotopes of the and nival zones, are (1) extremely strong winter upper Kolyma river winds (hurricanes) and (2) snow storms. While precipitation in the mountains where the goltsy The majority of stony slopes, except for the are situated is higher than in lowlands, winds goltsy, in the upper reaches of Kolyma are and hurricanes blow out all the snow. This loss restricted to the southerly exposed slopes, and of snow causes winter drying and wind erosion the adjacent ecosystems consist of dry meadow of the vegetation and stones. Together with or northern steppe (tundra-steppe) discontinu- extremely low winter temperatures (down to ous vegetation. Therefore, wandering spiders minus 60°C), rapid temperature and moisture (Gnaphosidae, Lycosidae, Liocranidae) can changes within one day, the lack of snow and a migrate to or from the scree habitats, and micro- short growing season creates an extremely se- cave dwellers such as the Micronetini and The- vere climate [Kuvayev, 1985]. The stony deserts, ridiidae can inhabit the neighbouring biotopes, in general, occupy flat and semi-flat surfaces of which have separate, flat stones. the mountains and uplands and mountain slopes In the environs of Aborigen Field Station, (1 200–2 000 m). stony slopes are formed from two different The other difference of European and Sibe- rocks, viz., slatestones (slatestone sheet) at 500– rian stony biotopes is connected with the pres- 1 000 m, and granite intrusions (800–2 500 m). ence of permafrost in Asia, causing the scree The presence of slatestones in the taiga belt, bottom to be permanently cold with tempera- makes conditions very favourable for spiders tures below 0°C. Therefore, lower depths of the (and for arachnologists to study them). Small Siberian screes are uninhabited. Because of the sites with slatestones on the border between permafrost, caves cannot exist in most parts of slatestones and granite belts, and within granite Siberia, and consequently there are no caverni- belts are inhabited by exotic species such as colous faunas in eastern Siberia (unlike Eu- Hybauchenidium holmi, Chalcoscirtus carbon- rope), which could enrich the species diversity arius, ‘Lepthyphantes’ epigynatus and Therid- of spiders in screes. ion thaleri. Another important difference between Si- Spiders inhabiting the stony ecosystems can berian and central European stony ecosystems be considered in the following four groups: is related to the surroundings of these habitats. 1) Obligate scree dwellers (moist and dry), The mountain and taiga belt ecosystems in east- not active hunters: Aculepeira carbonarioides, ern Siberia are open and usually include gravel, Chalcoscirtus grishkanae, C. carbonarius, pebbles, rocks, etc. Therefore, many species Flagelliphantes flagellifer, ‘Lepthyphantes’ are pre-adapted for living in stony debris, even punctulatus, ‘L.’ epigynatus, Poeciloneta pal- though they seem not to be true lithobionts or lida, P. petrophila, Theridion thaleri. petrophilous species. In Europe, stony ecosys- 2) Dwellers predominantly of scree ecosys- tems are always surrounded with shadowed for- tems (belonging to dominants): Acantholycosa ests or meadows with thick litter and therefore aborigenica, A. norvegica, Agroeca maculata, with no open gravel habitats. Clubiona propinqua, Enoplognatha serratosig- To conclude with the dissimilarities, it should nata, Incestophantes incestoides, Lathys alber- be mentioned that unlike in Europe, the stony ta, Procerocymbium sibiricum, Sibirocosa koly- ecosystems of north-east Siberia are not relics, mensis, Scotinotylus protervus, Tanasevitchia and such ecosystems are not discontinuous be- uralensis, Thanatus kolymensis, Theridion si- cause of the mountain ranges stretching from biricum, Xysticus rugosus. the south to the extreme north. Therefore, spe- 3) Spiders found both under stones and on cies can easily spread (in a historical sense) the screes: Agyneta birulai, Chalcoscirtus gla- 190 EUROPEAN ARACHNOLOGY 2003 cialis, Drassodes cupreus, D. neglectus, Erigo- Dictynidae noplus minaretifer, Hahnia glacialis, Para- Dictyna tyshchenkoi Marusik, 1988 syrisca holmi, Poeciloneta variegata. Siberian range; known from the Polar Urals, the 4) Eury- or polytopic (= eurybionts) dwell- upper Kolyma and Chukotka, including Wrangel ers: Euophrys proszynskii , Gnaphosa similis, Island. In the upper Kolyma, it is restricted to the Maso sundevalli, Microneta viaria, Pardosa eis- dry mountain tundra and low mountain ridges (800– 1 400 m). Common under separate stones, but very eni, Philodromus alascensis, Scotinotylus koly- rarely occurs on the margins of screes. mensis, Thanatus bungei, Titanoeca sibirica, T. nivalis, Wubanoides fissus, Xysticus albidus. Lathys alberta Gertsch, 1946 Species in the first group were not found East Siberio–north-west American range, reach- outside the screes. Some of them are known ing the south tundra in Chukotka. Associated mostly from a few specimens and only one or two with screes, but also common under separate stones localities in north-east Siberia (‘L’. punctulatus in dry habitats. In the taiga zone it is restricted to the (two localities) in Kolyma and Chukotka; C. forest belt and absent from the mountain tundra. In carbonarius (one locality in Siberia); T. thaleri the tundra zone, it occurs in lowlands on the south- (two localities in the world, both of which are in facing exposed slopes. In many places it is a domi- the upper Kolyma), H. holmi (a single locality nant species and occurs at high densities. in the world). Gnaphosidae Survey of the most common scree Drassodes cupreus (Blackwall, 1834) Trans-Palaearctic polyzonal (?) range. In north- dwellers west Siberia, it is restricted to the taiga zone and taiga belt, occurring on dry stony south-facing slopes Araneidae and screes, rarer than D. neglectus, and inhabiting Aculepeira carbonarioides (Keyserling, 1892) only lowland slopes up to 600–700 m. In some A widespread Siberio-American species distrib- habitats it reaches high densities. This species in- uted as far as the Arctic Ocean coast and everywhere habits screes in Tuva, south Siberia and the Kurile strictly restricted to stony ecosystems, viz., kurums, Islands [Logunov et al., 1998: sub (?) D. lapidosus; cliffs, stone circles, screes, where it builds its orb Marusik et al., 2000; etc.]. web between the stones. In Kolyma area it inhabits only mountains, whereas in coastal and tundra parts Drassodes neglectus (Keyserling, 1887) it can be found at low elevations. It is the only orb- Siberio–Nearctic polyzonal range; it is associat- weaver in the stony habitats of north-east Siberia, ed with dry habitats. In the upper Kolyma it is often i.e., a single representative of the ecological group associated with separate stones at the edges of screes ‘weavers’ and therefore, here it is always considered at elevations up to 1 100 m. In the tundra zone it was dominant in spite of its actual low densities at some found in a taiga oasis of Bol’shaya Osinovaya River sites. only. Clubionidae Gnaphosa similis Kulczyñski, 1926 Clubiona propinqua L. Koch, 1879 East Siberian range, from east Mongolia to Siberian range; the species reaches the south Chukotka. Associated with open gravely, south- tundra and is associated everywhere with screes. facing exposed slopes. Common both under sepa- In the taiga zone, it inhabits the mountains at 800– 1 400 m a.s.l. In both Chukotka and the upper rate stones and in shallow screes. Kolyma, the species belongs to the common, domi- nant and subdominant species groups. Gnaphosa orites Chamberlin, 1922 Circum-Holarctic range. In Asia, it occurs from Clubiona praematura Emerton, 1909 the highlands of Tuva [Marusik et al., 2000: sub Chukotka–Nearctic range [Mikhailov & Maru- Gnaphosa sp.1 (cf. orites)] to Wrangel Island. In the sik, 1996]; in Asia it is restricted exclusively to the upper Kolyma it occurs at elevations over 700 m. In tundra zone, occurring in several habitats but most the mountain tundra the species inhabits various common in the small screes or stony debris below biotopes including screes and gravelly places; in cliffs. screes it never attains a high density. Yu.M. Marusik. Petrophilous spiders of Beringia 191

Parasyrisca holmi Ovtsharenko, Platnick et Erigone arcticola Chamberlin et Ivie, 1947 Marusik, 1995 Siberio–Alaskan hypoarctic range, from Novaya East Siberian range, from Amur to the upper Zemlya to Chukotka Peninsula, southward to north Kolyma. Associated with separate stones in stony Cisokhotia [Eskov, 1994]. In the upper Kolyma and habitats occurring in the mountain tundras (conti- northern Cisokhotia, the species occurs on pebbly nental parts) and the dwarf pine belt at the sea coast. banks, whereas in Chukotka it inhabits scree and Very seldom occurs in screes. some other biotopes.

Hahniidae Erigonoplus minaretifer Eskov, 1986 Hahnia glacialis Sørensen, 1898 Endemic to the continental parts of north-east North-east Siberian–Nearctic range; in Asia it is Siberia. Most common in the goltsy and subgoltsy known from the continental parts of Magadan Area (an intermediate belt between the goltsy and taiga) and from Chukotka [Marusik et al., 1996]. Most belts under separate stones or in small stone fields, specimens of this species were collected from the which are free of snow during the winter. Unknown mountain tundra among gravel and under stones, from north of the taiga zone and never reaching a but it is also present in the forest belt. In some screes high density. it is a dominant species. Flagelliphantes flagellifer (Tanasevitch, Linyphiidae 1987) Agyneta birulai (Kulczyñski, 1908) Endemic of north-east Asia, was found in the Siberio–north-west Nearctic range, from the upper Kolyma, northern Cisokhotia and the north Polar Urals via Putorana Plateau and Arctic Yakutia Kuriles (Shumshu Isl.) only. Occurs in moist habi- to Wrangle Island and southward to south Trans- tats with embedded stone fields (slatestones or mo- baikalia [Eskov, 1994]. A dweller of the south- raine pebbles). Very rare. facing exposed slopes, occasionally can be collected among scree or under separate stones. Hybauchenidium holmi Marusik, 1988 Known from a single locality and habitat (see Bathyphantes eumenis (L. Koch, 1879) Table legend) in the upper Kolyma. Siberian (?) range: from Yenisei to Chukotka, south to south Transbaikalia and Manchuria [Eskov, Incestophantes incestoides (Tanasevitch et 1994]. Eurybiontic species. Eskov, 1987) North-east Siberian widespread endemic species, Centromerus pacificus Eskov et Marusik, occurring up to the southern tundra. Can be found 1992 under separate stones, but most common in elongate East Siberian range, from south-east Cisbaikalia thin scree. In some places may reach high densities. south-eastward to Maritime Province and north- eastward to the upper Kolyma [Eskov, 1994]. A Incestophantes kochiellus (Strand, 1900) dweller of several habitats, rarely occurring under Trans-Palaearctic boreo-hypoarctic range, from northern Scandinavia to Chukotka [Eskov, 1994]. stones or in scree. Inhabits scree in Chukotka only. Eborilaira alpina Eskov, 1989 Islandiana falsifica (Keyserling, 1886) North-east Siberian range, from the upper Koly- Trans-Palaearctic–north-west Nearctic range, ma westward to the upper Indigirka and north-east from north Scandinavia to Chukotka Peninsula and to Chukotka [Marusik et al., 1992]. In the upper Wrangel Island, southward to Kamchatka [Eskov, Kolyma it dwells in the stony deserts, screes and 1994]. In the upper Kolyma, and northern Cisokho- stony taluses in the mountain tundra (1 000 m and tia it lives on pebbly beaches. It also inhabits scree higher). in Chukotka. Entelecara sombra (Chamberlin et Ivie, 1947) Kolymocyba petrophila Eskov, 1989 Siberio–trans-Nearctic range, from Yenisei to North-east Siberian hypoarctic range, known the upper Kolyma, southward to Tuva and south from the upper Kolyma and Amguema River basin Sakhalin [Eskov, 1994]. It inhabits two different in Chukotka Peninsula [Marusik et al., 1992]. In- biotopes: stony (the kurums near Magadan, pebbles habits stony places on north-facing exposed slopes or stones in moist ground) and birch trees (under in the forest belt and south-facing slopes in the bark). mountain tundra up to1 400 m; rare. 192 EUROPEAN ARACHNOLOGY 2003

‘Lepthyphantes’ epigynatus Tanasevitch, 400–1 400 m in various open habitats with stones. A 1988 high density was observed on pebbly beaches. In Endemic to the upper Kolyma and known from Chukotka, it inhabits scree only. a single habitat (see Table legend). Perro tshuktshorum (Eskov et Marusik, ‘Lepthyphantes’ punctulatus Holm, 1939 1991) Trans-Palaearctic subarctic range. Besides the Endemic to Chukotka. Inhabits stony deserts. upper Kolyma, it was found in two localities in Chukotka, in all cases on open scree. A high density Porrhomma boreale Banks, 1892 was observed only in Chukotka. Siberio–Alaskan range, from Yenisei southward to the Commander Islands and eastward to Alaska Maso sundevalli (Westring, 1851) [Eskov, 1994]. Very rare. Approximately ten speci- Circum-Holarctic range. Eurybiont. Rarely oc- mens have been found in screes within the forest belt curs in scree. (600–900 m). Microneta viaria (Blackwall, 1841) Porrhomma cf. kulczynskii Starêga, 1974 Circum-Holarctic polyzonal range [Eskov, Known in the upper Kolyma and Chukotka only. 1994]. Rare species in the upper Kolyma and north- Rare. All specimens were collected in stony deserts. ern Cisokhotia. A leaf litter dweller, however it rarely occurs in shaded screes. Procerocymbium sibiricum Eskov, 1989 Oreoneta leviceps (L. Koch, 1879) East Siberian hypoarcto-montane range, occur- Siberio–Nearctic range [Eskov, 1994], restrict- ring from Transbaikalia to Chukotka. In the taiga zone of the upper Kolyma it exclusively inhabits ed to the tundra zone and occurs in different habitats scree in mountain tundra. In some sites it may attain including stony polygons and stony debris. high densities. Oreonetides kolymensis Eskov, 1991 North-east Siberian endemic, inhabiting differ- Scotinotylus alpinus (Banks, 1896) ent biotopes. A high density was found only in the Siberio–trans-Nearctic range [Eskov, 1994]. embedded and moist moraine pebble debris on Lank- Eurybiont. As a scree dweller it was found only in ovaya River. Ola Plateau.

Poeciloneta petrophila Tanasevitch, 1989 Scotinotylus kolymensis Eskov et Marusik, Beringian range; in addition to the upper Koly- 1994 ma, it was found in the northern Cisokhotia and in North-east Siberian range, known from northern two localities in Chukotka (together with ‘L’. punctu- Cisokhotia and the upper Kolyma only. A litter latus). For habitats in Kolyma area, see the table dweller, but was found once in a stony habitat. legend. Other habitats: open screes with a moist base. Scotinotylus protervus (L. Koch, 1879) Siberio–Alaskan range. A dweller of the forest Poeciloneta pallida (Kulczyñski, 1908) belt stone moraines. In mountains and in the tundra Endemic to north-east Siberia, reaches south zone it is associated with the moraines and moist tundra, everywhere quite rare. Associated with dry lower edges of the stone trains. In several habitats it stony habitats. Never reaches high density. attains high densities.

Poeciloneta tanasevitchi Marusik, 1988 Tanasevitchia uralensis (Tanasevitch, 1983) North-east Siberian endemic, know from three Siberian range; in north-east Siberia it was found localities only: one in eastern Yakutia and two around only in the taiga zone, and only in scree (with a few Magadan. In the environs of Magadan it was found exceptions). It occurs either in the scree with flat in the shaded, small and somewhat moist screes. stones, or in those with lichen carpets (most com- Very rare but can occur at a high density. mon), in the latter case it dwells both on the lichen cover and underneath the stones. In some scree it is Poeciloneta variegata (Blackwall, 1841) very abundant. In Tuva, southern Siberia, the spe- Trans-Palaearctic–west Nearctic range. In the cies was found among scree with lichen cover upper Kolyma this species lives at elevations of [Marusik et al., 2000]. Yu.M. Marusik. Petrophilous spiders of Beringia 193

Wubanoides fissus (Kulczyñski, 1926) Sibirocosa subsolana (Kulczyñski, 1907) East Siberian endemic, the most eurybiontic Endemic to Chukotka, occurring from the Koly- linyphiid species in Siberia. At a high density it was ma river mouth to eastern Chukotka and northward found only in a single alder grove. A relatively high to Wrangel Island. Exclusively restricted to scree density was observed in the north-exposed (and and stony deserts. correspondingly cold) screes. In the tundra zone, it is restricted to warm screes only. Philodromidae Philodromus alascensis Keyserling, 1884 Liocranidae Siberio–American range, from China to the Agroeca maculata (L. Koch, 1879) southern tundra zone. The species has no special Siberian boreal range, from east Kazakhstan and preference for stones, but rather to flat surfaces in Perm Area north-east to the upper Kolyma and general: tree trunks, big stones, ice fields. southward to central Mongolia [Mikhailov & Marusik, 1996]. It occurs in xeromorphic gravelly Philodromus cespitum (Walckenaer, 1902) habitats, but is most abundant in scree, also in the Circum-Holarctic range. A dweller of herba- ground layer of thin larch forest with sparse vegeta- ceous and shrub strata, occasionally found in scree. tion and in aspen stands. Thanatus bungei (Kulczyñski, 1908) Lycosidae Range is almost the same as in P. alascensis, but Acantholycosa aborigenica Zyuzin et Maru- restricted to the taiga and tundra zones, common on sik, 1988 tree trunks, Veratrum leaves, but most common in kurums and scree. Endemic to eastern Siberia, occurring from east- ern Mongolia to Sikhote-Alin’ Mt Ridge and north- Thanatus kolymensis Marusik, 1991 ward to the upper Kolyma. Associated with dry This species is known from two localities in the stony slopes, where it can be found in scree and upper Kolyma, occurring in the subgoltsy and goltsy other gravelly habitats. In many cases it is a co- belts among stony deserts. Logunov [1996] treats inhabitant with Acantholycosa norvegica, but never this species as a junior synonym of T. arcticus in the same proportion. Elevation: 600–800 m. It Thorell, 1872, although in north-east Siberia and should be noted that all species of the genus Acan- Tuva [cf. Logunov et al., 1998] the latter species tholycosa, except for A. lignaria (Clerck, 1757) are never occurs in stony biotopes. In the upper Kolyma associated with scree or stony slopes. T. arcticus inhabits the forest belt and occurs in various herbaceous or boggy biotopes. Acantholycosa norvegica (Thorell, 1872) Trans-Palaearctic range, present in the taiga belt Salticidae only, on dry southern exposed slopes. Common in Chalcoscirtus carbonarius Emerton, 1917 continental and coastal parts of north-east Siberia. North-east Siberio–north-west Nearctic range, within the Palaearctics it is known from the upper Pardosa eiseni (Thorell, 1875) Kolyma only. Rare. It was found in a relatively high Trans-Palaearctic range [Marusik et al., 2000], number in only one biotope: the slatestone scree (c. occurs in a wide range of habitats. In several scree 1 000 m) in the subgoltsy zone. slopes it reaches relatively high densities. Chalcoscirtus glacialis Caporiacco, 1935 Sibirocosa kolymensis Marusik, Azarkina Siberio–north-west Nearctic range northward to et Koponen, 2004 the southern tundra, restricted to dry and xeric stony Endemic to the upper Kolyma and northern habitats. In the taiga zone, it occurs in the taiga belt Cisokhotia, widespread and common everywhere in and subgoltsy zone. In south Siberia (Tuva), this the goltsy zone. Among the lycosids, this species is species is recorded as a typical dweller of the moun- most strongly afilliated with stone debris habitats tain steppes [cf. Logunov et al., 1998], but it seems and because of this it is very uncommon in pitfall a separate (sub)species lives there. traps in adjacent biotopes. In Kolyma area it inhabits the mountain tundra belt only (1 000 m and higher), Chalcoscirtus grishkanae Marusik, 1991 whereas in the coastal tundras near Magadan it can East Siberian range, from Chita Area to the occur in lowland stony debris. Previously, this spe- upper reaches of Kolyma River. In the upper Koly- cies was confused with S. subsolana [cf. Marusik et ma and in northern Cisokhotia, it is restricted to al., 2004]. scree only within the taiga belt. 194 EUROPEAN ARACHNOLOGY 2003

Euophrys proszynskii Logunov, Cutler et Xysticus rugosus Buckle et Redner, 1964 Marusik, 1993 Siberio–west Nearctic range. In the upper Koly- Siberian range, does not exceed the taiga zone. ma it occurs in the subgoltsy belt, where it is found Restricted to dry stony habitats in the taiga and in xeric snowless habitats, under separate stones or subgoltsy belts. Most numerous in the small, stone in small-stone fields. In Yukon Territory, the habi- fields or small sites within scree, where females tats of this species are almost identical. build their retreats and lay egg-sacs. Titanoecidae Theridiidae Titanoeca nivalis Simon, 1874 Enoplognatha serratosignata (L. Koch, Trans-Palaearctic–west Nearctic range [Marusik 1879) et al., 2000]. In the upper Kolyma and northern Trans-Palaearctic boreo-montane range [Marusik Cisokhotia it inhabits southern exposed slopes with et al., 2000], from Switzerland and Finland east- gravel or semi-shaded but warm aspen and birch ward to the Kolyma River mouth, southward to stands. Tuva, Mongolia and Gansu. In the upper Kolyma, all specimens were collected in open biotopes on Titanoeca sibirica L. Koch, 1879 slopes and plain places at elevations of 500–900 m. Trans-Siberian range Marusik et al., 2000], from The highest density of this species was observed in the South Urals north-eastward to the Kolyma River small stony taluses along mountain ridges with sparse mouth and southward to north Mongolia. In the Carex and lichen vegetation. upper Kolyma it occurs on steppe slopes, however in some places it inhabits pebbly river banks. In Theridion sibiricum Marusik, 1988 northern Cisokhotia this species forms high density Siberian boreo-nemoral range, known from Tuva populations on the seashore scree. to Kolyma and Kamchatka Peninsula, southward to Maritime Province, where it is restricted to dry Tetragnathidae stony habitats. In some open scree sites the species Zygiella dispar (Kulczyñski, 1885) attains high densities. Far East–Nearctic range. In north-east Siberia it ‘Theridion’ thaleri Marusik, 1988 occurs on seashore cliffs and rock debris around Was found in two localities in the upper Koly- them. ma, for localities see the table legend. Spider communities of the major Thymoites bellissimus (L. Koch, 1879) stone debris habitat types of north- Trans-Palaearctic boreal range. One or more species with similar epigynes occur in north-east east Siberia Siberia. There are at least two morphs, one specific for the stone hills and fields in northern Cisokhotia The stony deserts and Kolyma, as well as in Amur River and Karelia, In addition to the petrophilous taxa, the and the other is known only from Kolyma area. The edges of stony deserts are inhabited by moun- species lives in the Chosenia belt forests along small tain tundra and xerophilous species. Usually rivers. only a single true wandering spider commonly Thymoites oleatus (L. Koch, 1879) occupies such habitats, either Sibirocosa koly- Siberio–Nearctic range [Marusik et al., 2000]. mensis or S. subsolana (in Chukotka). Another In the upper Kolyma the only female was collected species, Parasyrisca holmi, is restricted to the in leaf litter of an alder stand on a north exposed goltsy but is rare. Occasionally Drassodes ne- slope (650 m), whereas in Chukotka it is common glectus (in the south) and Gnaphosa orites can among scree and stony debris. also be found. Thanatus bungei is quite com- mon. Thanatus kolymensis occasionally occurs Thomisidae at low altitudes (about 1 000 m). Aculepeira Xysticus albidus Grese, 1908 carbonarioides, the largest orb-weaver in north- Trans-Palaearctic species, restricted to moun- tain tundra scree on southern slopes and the pebbly east Siberia, builds its web between stones, river banks in the taiga zone. Eurytopic in the tundra usually in small hollows. In the tundra zone this zone. In the mountain tundra it is primarily restrict- species is primarily restricted to scree. Other ed to dry and warm meadows and separate stones. inhabitants of the goltsy are Eborilaira alpina, Yu.M. Marusik. Petrophilous spiders of Beringia 195

Porrhomma cf. kulczynskii and Procerocymbi- tus bungei; (2) the jumping spiders, Chalcoscir- um sibiricum. In two small sites of the upper tus glacialis, C. grishkanae (both rare); (3) the Kolyma very different and exotic populations weavers are Lathys alberta and Theridion si- were found: in the first, Chalcoscirtus carbon- biricum (dominants in the majority of biotopes); arius and ‘Lepthyphantes’ epigynatus predom- the influents: Incestophantes incestoides. inated, while other species were occasional; in the second site, Hybauchenidium holmi and The south-exposed scree of the tundra ‘Theridion’ thaleri predominated. In Chukot- ka, in huge stony deserts, Perro tchuktchorum, zone Clubiona propinqua and Mughiphantes sobri- The most common species are Sibirocosa us were found in addition to Sibirocosa sub- kolymensis or S. subsolana, Clubiona propin- solana. qua, Incestophantes incestoides, Lathys alber- ta, Pardosa eiseni, Scotinotylus protervus. The kurums An interesting fact regarding stone-affiliat- Surface dwellers were Sibirocosa kolymen- ed spiders was revealed in the upper Kolyma. sis or S. subsolana (in Chukotka), Thanatus Some of the lithobionts and petrophilous spi- bungei, sometimes Philodromus alascensis, ders have closely related siblings, which live Parasyrisca holmi, in addition to Aculepeira allopatrically in a very different habitat. They carbonarioides. can be considered ecological vicariants. They are: Hybauchenidium holmi – H. aquilonare The south-exposed scree of the taiga zone (L. Koch, 1879) (the litter of a closed flood Surface dwellers are as follows: (1) wander- plain forest); Theridion thaleri – T. ohlerti ing species dominants and influents (= com- Thorell, 1870 (dwarf birches, willows and mon): Acantholycosa aborigenica, A. norvegi- Vaccinium in lowlands); Parasyrisca holmi – ca, Drassodes neglectus, D. cupreus, Thana- P. tyshchenkoi Ovtsharenko, Platnick et Table 1. Spider populations in several stone debris ecosystems in north-east Siberia. Explanation of the numbers is given in ‘Material and methods’. Dominant categories: D and d = dominant, S = subdominant, C or i = common or influent (abundant), o occasional, R or r = rare. Capital letters are used for calculated abundances and small ones for generalized and subjective estimations. Occurrence: T = true (obligatory), N = most numerous in stony areas, S = stony debris and separate stones, E = eurybionts, O = occasional. Òàáëèöà 1. Íàñåëåíèå ïàóêîâ íåêîòîðûõ êàìåíèñòûõ áèîòîïîâ íà ñåâåðî-âîñòîêå Ñèáèðè. Ðàñøèôðîâêà òî÷åê ïðèâåäåíà ⠑Material and methods’. Ìàññîâîñòü: D è d = äîìèíàíòû, S = ñóáäîìèíàíòû, C èëè i = îáû÷íûå èëè èíôëþýíòû, R èëè r = ðåäêèå. Çàãëàâíûå áóêâû èñïîëüçîâàíû äëÿ ðàñ÷åòíûõ äàííûõ, ïðîïèñíûå äëÿ îöåíî÷íûõ äàííûõ. Âñòðå÷àåìîñòü: T = èñêëþ÷èòåëüíî êàìåíèñòûå áèîòîïû (îáëèãàòíûå ïåòðîáèîíòû), N = íàèáîëåå ìàññîâû â â êàìåíèñòûõ áèîòîïàõ, S = îñûïè èëè îòäåëüíûå êàìíè, E = ýâðèáèîíò, O = ñëó÷àéíûé. Taxa Occurrence 1 2 3 4 5 6 7 8 9 10 11 12 13 Aborigen Field Station K MA O S E E CH ARANEIDAE Aculepeira carbonarioides TDD d Rd CLUBIONIDAE Clubiona propinqua TCCDDd C. praematura Ei DICTYNIDAE Dictyna tyshchenkoi S Lathys alberta NT DC DDiS S d GNAPHOSIDAE Drassodes cupreus NR Si D. neglectus NRS ? Gnaphosa similis NR Ci i G. orites NCR Ri Parasyrisca holmi TN C R r 196 EUROPEAN ARACHNOLOGY 2003

Table 1 (continued). Òàáëèöà 1 (ïðîäîëæåíèå).

Taxa Occurrence 1 2 3456 78910111213 Aborigen Field Station K MA O S E E CH HAHNIIDAE Hahnia glacialis NDC i i LINYPHIIDAE Agyneta birulai SC Sr Bathyphantes eumenis ESi Centromerus pacificus ER r Eborilaira alpina TSi Entelecara sombra SC i Erigone arcticola SCi Erigonoplus minaretifer SRC Flagelliphantes flagellifer TC ‘Lepthyphantes’ epigynatus T ‘L.’ nenilini N i ‘L.’ punctulatus TRDRi Hybauchenidium holmi TC Incestophantes incestoides NT R C i i I. kochiellus Ni Islandiana falsifica ?Di Kolymocyba petrophila TR r? Maso sundevalli ER Microneta viaria ER i Mughiphantes flexilis NR M. sobrius Ti Oreoneta leviceps ?Si Oreonetides kolymensis T? i Perro tchuktchorum Ti Poeciloneta pallida TC CC i P. petrophila TDiDd P. tanasevitchi Td P. variegata NRd Porrhomma borealis T? P. cf. kulczynskii T? R R Procerocymbium sibiricum Ti Scotinotylus alpinus EC S. kolymensis ES S. protervus NDrDDSd Tanasevitchia uralensis TDD Walckenaeria fraudatrix OCi W. holmi OC Wubanoides fissus N C D D R r S C D i LIOCRANIDAE Agroeca maculata TC LYCOSIDAE Acantholycosa aborigenica TC A. norvegica TC R Sibirocosa kolymensis TDRi S. subsolana TRd Pardosa eiseni EC r i PHILODROMIDAE Philodromus alascensis EC i P. cespitum OR Thanatus bungei EN R DS i i T. kolymensis TC SALTICIDAE Chalcoscirtus carbonarius T C. glacialis SR C C. grishkanae TC i Euophrys proszynskii NR Dd Yu.M. Marusik. Petrophilous spiders of Beringia 197

Table 1 (continued). Òàáëèöà 1 (ïðîäîëæåíèå).

Taxa Occurrence 1 2 3 4 5 6 7 8 9 10 11 12 13 Aborigen Field Station K MA O S E E CH THERIDIIDAE Enoplognatha serratosignata Si Theridion sibiricum TS DDdD ‘T.’ thaleri TDR Thymoites bellissimus TR RdC T. oleatus O? i THOMISIDAE Xysticus albidus EDi X. rugosus T TITANOECIDAE Titanoeca nivalis S T. sibirica SC d TETRAGNATHIDAE Zygiella dispar Cliffs d

Marusik, 1995 (larch and birch tree trunks, dion thaleri, Chalcoscirtus carbonarius, Tha- under bark), Xysticus rugosus – X. sibiricus natus kolymensis. However, many species from Kulczyñski, 1908 (under larch bark), Thanatus the taiga belt are known in Chukotka. Among kolymensis – T. arcticus Thorell, 1872 (habitats the characteristic (common and widespread) with thick vegetation, mostly in lowlands); Chal- species of the taiga zone, only Theridion sibiri- coscirtus glacialis – C. hyperboreus (bogs in cum does not inhabit scree of the tundra zone. the mountain tundras, lichen pillows); Euophrys Table 2. proszynskii – E. flavoatra (Grube, 1861) (the Harvestman and spiders associated with the stony taiga belt, bogs). Sibirocosa kolymensis and S. debris near Kluane Lake, Yukon Territory. Expla- subsolana are geographical vicariants. nation of site numbers is given below. Òàáëèöà 2. As is evident from Table 1 and the com- Ñåíîêîñåö è ïàóêè, ñâÿçàííûå ñ êàìåíèñòûìè ments in the species distributions above, stony ñîîáùåñòâàìè â îêðåñòíîñòÿõ îç. Êëþýéí, òåð- debris spiders in the taiga zone can be assigned ðèòîðèÿ Þêîí. Íîìåðà áèîòîïîâ ðàñøèôðîâà- to two groups: mountain dwellers (mountain íû íèæå. tundra belt) and those which inhabit the taiga. Species/site 1 2 3 4 5 Species of both groups that reach the tundra Liopilio yukon 550 zone inhabit the same stony biotopes (or same Lathys alberta 120 18 2 27 10 belt). For example, in Chukotka the dwellers of Enoplognatha intrepida 10 2 7 40 the mountain tundra belt of the upper Kolyma, Theridion petraeum 268 Thymoites oleatus 1 viz., Clubiona propinqua, Aculepeira carbon- Steatoda borealis 17 3 1 arioides, Procerocymbium sibiricum, ‘Lepthy- ‘Lepthyphantes’ sp. 9 11 2 phantes’ punctulatus, Eborilaira alpina and Scotinella pugnata 316 Pardosa mackenziana 20 Dictyna tyshchenkoi are sympatric with dwell- P. albomaculata 2 ers of the taiga belt: Lathys alberta, Inces- Aculepeira carbonarioides 3 tophantes incestoides, Poeciloneta petrophila, Steatoda albomaculata 4 P. pallida, Wubanoides fissus, etc. Correspond- Drassodes neglectus 3 Chalcoscirtus glacialis 3 ingly, there is no subdivision among stone de- Terralonus mylothrus 24 bris dwellers according to altitude in the tundra Xysticus deichmanni 1 zone of Chukotka. Scotinotylus alpinus 12 S. protervus 3 Not all the species, which inhabit the moun- Poeciloneta variegata 2 tain stony debris in Kolyma Area, occur (i.e., Porrhomma sp. 2 were found) in Chukotka, namely: Sibirocosa Bathyphantes eumenis 2 Gnaphosa muscorum 6 kolymensis, Parasyrisca holmi, Hybauchenidi- G. orites 2 um holmi, ‘Lepthyphantes’ epigynatus, Theri- Erigoninae gen. sp. 1 5 198 EUROPEAN ARACHNOLOGY 2003

Spiders of the rock debris of Yukon In the moist pebble moraine scree near Car- Territory macks, a new species of Poeciloneta was found together with Lepthyphantes sp. The former spe- Spiders associated with stony debris around cies is closely related to the north-east Siberian Kluane Lake were examined (Table 2). Addi- P. tanasevitchi and occupies the same habitats. tionally, a small collection was made near Car- macks Town. Unlike north-east Siberia but sim- Discussion ilar to European stony debris, the Yukon stony The similarity between the spider faunas of debris ecosystems studied were formed in re- central European and north-east Siberian stony gions lacking permafrost. In all cases, the spi- debris habitats is very weak at species level. der species studied inhabited both surface and There are only two species in common: Acan- bottom layers. Most of the spiders were studied tholycosa norvegica (a true lithobiont) and Maso in the following five stony habitats around the sundevalli (a litter dweller, occasionally occur- south-east corner of Kluane Lake: ring in stony habitats). However, there are more 1) rock debris in a poplar grove (without common species in the two faunas (central Eu- continuous soil cover) on the bank (900 m); rope and the upper Kolyma) like Bathyphantes 2) open (unshaded) rock debris above site 1 simillimus (L. Koch, 1879), Drassodes cupreus, and under cliff (950 m); Diplocentria bidentata (Emerton, 1882), Mi- 3) cliff and cliff scree (1 000–1 300 m); croneta viaria, Philodromus cespitum and Po- 4) slatestone scree (1 900 m); eciloneta variegata. B. simillimus and D. bi- 5) klippe, klippe debris and moist scree in dentata are scree dwellers in central Europe the mountain tundra (1 850 m) only [Rùžièka, pers. comm.]. D. bidentata in- In other stony debris sites around Kluane habits only the lower (bottom) margins of scree, Lake, which are not shown in Table 2, Xysticus whereas in Siberia it is never associated with rugosus, ‘Lepthyphantes’ alpinus (Emerton, stones. 1882) and Scotinotylus majesticus (Chamber- There are at least two cases of vicariance lin et Ivie, 1947) were also collected. The most among stone dwellers between European and exciting observation in Kluane Lake happened east Siberian screes: Wubanoides fissus and W. to be the similarity between the lake-shore and uralensis (Pakhorukov, 1981) (central and west the high mountain scree fauna, and the absence Siberia, south Germany and Czech Republic) of many species in the intermediate zone. Such and Xysticus rugosus (Siberia and north-west a situation can apparently be explained by the America) and X. bonneti Denis, 1935 (south- similarity of the moisture in low and upland west Siberia, the Urals and south European stone debris; at least the ground beneath the Mts). stones was wet both in low and upland sites. The family composition of the two faunas is Rock debris of the steep south slope with steppe also different. On the one hand, the Czechian vegetation was dry and shallow. scree fauna includes more families (Amaurobi- As can be seen from Tables 1 and 2, the idae, Pholcidae, Agelenidae, Nesticidae, Tet- similarity of species composition between north- ragnathidae), whereas the diversity of the erig- east Asian and north-west American stony de- onine species is much lower. On the other hand, bris faunas is quite high (c. 50%), viz., of the 23 three families such as Titanoecidae, Philodro- Canadian species, 12 are also recorded in Ma- midae and Hahniidae, which were found in the gadan Area. It should be noted that the habitat Siberian screes, have not been recorded as scree preferences of Bathyphantes eumenis are dif- dwellers in the Czech Republic [Rùžièka, 1988, ferent in different parts of Beringia. In north- 1990b; Rùžièka et al., 1989]. east Siberia it lives in herbaceous vegetation or A comparison of the beetle fauna of Cze- the moraine pebbles in tundra, while in Yukon chian stony debris [Rùžièka et al., 1989] and Territory it occurs in the deep and moist layers that of north-east Siberia shows at least five of rock debris. common species [A.S. Ryabukhin, pers. comm.]. Yu.M. Marusik. Petrophilous spiders of Beringia 199

Taxonomic positions of some Siberian Aleo- (Edmonton, Canada) who critically read the manu- charinae (Staphylinidae) are unclear. All the script of this paper and checked the English of an common rove and carrion beetle species: Oma- earlier draft. The expedition to Yukon Territory was lium caesum, Scyodrepoides watsoni, Quedius supported by Drs D. Berman, (Magadan, Russia) limbatus, Tachinus rufipennis and Nicropho- and S. Armbruster (Fairbanks, Canada) to whom I extend my thanks. Dr. V. Rùžièka (Èeské Budejov- rus vespilloides are litter and carcass dwellers ice, Czech Republic) is thanked for providing very in Siberia and are not connected to the stony helpful critical comments on an earlier draft and debris habitats. additional information about the petrophilous spi- The stony debris fauna of north-east Sibe- ders of central Europe. I also thank Dr. D.V. Lo- ria, as it is seen from Table 1 and in V. Rùžièka’s gunov (Manchester, UK) for critical comments and papers, is more diverse (69 species in Table 1 helpful suggestions on the final manuscript. Har- [33 without occasional] and 55 [28 without vestman specimens from the Yukon Territory were occasional] in Rùžièka [1986]). It is worth men- identified by Dr. N. Tsurusaki (Tottori, Japan) and tioning that the similarity between the litho- the salticid species by Dr. D.V. Logunov (Manches- ter, UK). This project was sponsored in part by the biont spider faunas of north-east Siberia (par- Russian Foundation for Basic Research (RFBR ticularly the upper Kolyma) and the Czech Re- grants no. 01-04-48989, no. 04-04-48727) and the public (3%) is distinctly lower than that of the Far-East Branch of the RAS (no. 04-3-A-06-042). total araneofaunas: 21% (18% upper Kolyma– Czech Republic). References The faunas of Scandinavia and north-east Siberia are more similar [cf. Marusik, 1986]. In Eskov K.Yu. 1985. [The spiders of the tundra zone in the addition to D. cupreus and A. norvegica, there USSR] // Ovtsharenko V.I. (ed.), Fauna i ekologiya paukov SSSR. Trudy Zool. Inst. AN SSSR. T.139. are some more common species: Thymoites S.121–128 [in Russian]. bellissimus, ‘Lepthyphantes’ punctulatus, Gna- Eskov K.Yu. 1988. [Aranei of Central Siberia] // Rogache- phosa orites and some other common or rare va E.V. (ed.), Materialy po faune Srednei Sibiri i species. A further interesting conclusion can be prilezhashchikh rayonov Mongolii. Moscow: Minis- try of Agriculture. S.101–155 [in Russian]. drawn from reviewing the north-east Siberian Eskov K.Yu. 1994. Catalogue of the linyphiid spiders of stony debris fauna. Of the 72 petrophilous spe- northern Asia (Arachnida, Araneae, Linyphiidae). cies, 27 were described recently (from 1985– Sofia-Moscow: PENSOFT Publ. 144 p. 2003). This reflects the rather poor state of our Koponen S. 1992 (for 1990). Spiders (Araneae) on the cliffs of the Forillon National Park, Québec // Natural- former knowledge on spiders from stony debris iste Can. (Rev. Écol. Syst.). Vol.117. P.161–165. ecosystems. It is worth noting that the percent- Kuvayev V.B. 1985. [The cold goltsy deserts]. Moscow: age of narrow (i.e., confined to a very small Nauka. 78 p. [in Russian]. particular area) endemics among the petrophil- Logunov D.V. 1996. A critical review of the spider genera Apollophanes O. Pickard-Cambridge, 1898 and Tha- ous species (24%) in north-east Siberia is twice natus C.L. Koch, 1837 in North Asia (Araneae, Philo- that of the whole arachnofauna (about 9% [see dromidae) // Rev. Arachnol. T.11. Fasc.13. P.133– Marusik, 1988]). Having only counted the true 202. (exclusive) lithobionts (37 species), the per- Logunov D.V. & Marusik Yu.M. 1995. Spiders of the family Lycosidae from the Sokhondo reserve, Chita centage of endemic species restricted to north- Area, East Siberia (Arachnida: Araneae) // Beitr. east Siberia is as high as 46%. Araneol. Bd.4. P.109–122. The two lithobiont faunas of the relatively Logunov D.V. & Marusik Yu.M. 1999. A brief review of close geographical regions of north-east Asia the genus Chalcoscirtus Bertkau, 1880 in the fauna of Central Asia and the Caucasus (Araneae, Salticidae) and north-west America have more than 50% of // Arthropoda Sel. Vol.7. No.3. P.205–226. their species in common, especially if one merg- Logunov D.V. & Marusik Yu.M. 2000. Catalogue of the es sibling species. The similarity of the entire jumping spiders of northern Asia (Arachnida, Arane- araneofaunas of these two areas is approxi- ae, Salticidae). Moscow: KMK Sci. Press Ltd. 299 p. mately 52%. Logunov D.V., Marusik Yu.M. & Koponen S. 1998. A check-list of the spiders in Tuva, South Siberia with analysis of their habitat distribution // Ber. Naturwiss. ACKNOWLEDGEMENTS. I wish to express Med. Vereins Innsbruck. Bd.85. P.125–159. my deep gratitude to the late V. Roth and R. Leech Marusik Yu.M. 1988. [The fauna and population of spi- 200 EUROPEAN ARACHNOLOGY 2003

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