International Kuril Islands Project

Expedition No. 3, July 25-September 2, 1996

FIELD REPORT ON SPIDER SAMPLING:

PRELIMINARY RECONNAISSANCE OF THE SPIDER FAUNA

OF THE NORTHERN KURIL ISLANDS

by Rodney L. Crawford

INTRODUCTION

The general lack of prior knowledge about the spiders of the Kuril Islands is equally true of the northern group, as of the southern and central islands previously visited. The only older literature addressing these islands is a few Japanese papers: Saito (1932, 1933, 1935) and Nakatsudi (1937). In Saito's 3 papers, 16 species are recorded from Paramushir, Alaid, and Shumshu (?) (the first paper indicates Shumshu but the others indicate Simushir in the Central Kurils...for the same records!). Of these recorded species, about half are obvious misidentifications which cannot be placed now. Nakatsudi (1937) added one new species from Paramushir. The one modern checklist of Kuril spiders, by Marusik et al. (1992), lists 47 species from Paramushir and Shumshu, based on recent collecting by Russian non-arachnologists and including all of the prior records that they felt confident in assigning to known species. A paper by Eskov and Marusik (1993) supplemented this checklist with some new species descriptions, but none from the Northern Kurils. To summarize: all existing literature about the spiders of the Northern Kuril Islands refers only to the northernmost islands of Paramushir, Shumshu, and Alaid, and is extremely limited even for these islands. So far as known, no spiders had been recorded or even collected on the northern and north-central islands of Onekotan, Kharimkotan, Chirinkotan, Ekarma , Shiashkotan, Raikoke, and Matua prior to the IKIP-1996 expedition reported here.

The Marusik et al. (1992) checklist represents a considerable improvement over previous knowledge, but much more clearly remains to be done. This paper was based primarily on collections made by A. M. Basarukin, a vertebrate zoologist, and thus incorporates a bias toward less cryptic forms. Also, the islands were represented unequally, and no ecological information on Kuril Islands spiders has been available. The present report represents the first contribution based on field work by professional arachnologists in the northern Kuril Islands.

The collaborators on the spider project had somewhat different, and largely complementary, goals. The goal of RLC was to sample all productive habitats equally and produce as balanced a list for each locality and habitat as practicable in the limited time available. The goal of KYE was primarily to sample Linyphiidae, a very large family of mostly very small spiders found mainly in cryptic habitats, and especially productive of undescribed species.. KYE also sampled other spider groups as occasion offered. TK is a specialist on the family Gnaphosidae, of which only a few species are found in the Kuril Islands; he sampled general spider fauna, but sought out rocky habitats where gnaphosids tend to be found, thus probably adding further to the diversity of the collection (little of his material was field-identified, due to shortage of lab space on the ship). TWP directed much of his effort toward spider collecting on these islands; as a non-specialist, he concentrated on augmenting the numbers of specimens collected rather than concentrating on any one group, and was responsible for 30% of the 6,682 spider specimens brought back to the USA this year, including a number of species not taken otherwise. It should be noted that very little of the material collected by KYE and TK is reported on here, as that was taken back to Moscow and Osaka respectively and has not yet been available for study by RLC.

MATERIALS AND METHODS

Within limits imposed by time and access restrictions, an effort was made to collect in diverse vegetation communities. These northern islands are largely lacking in forest; the commonest habitats encountered include shore habitats (stony, meadow, and dune), inland meadow habitats, alder and dwarf-pine thickets, bogs and other wetlands, and (generally unreachable in the time available) stony mountainous habitats. Manmade habitats (buildings and their surroundings) were sampled when available. In the field, collecting sites were defined as 100 X 100 m squares within special habitats. Subsites were defined within sites wherever more than one distinct macrohabitat type was being sampled, and specimen lots were segregated by subsite and microhabitat (common microhabitats: thicket understory foliage, field layer foliage in meadow habitats, leaf litter, moss including Sphagnum from bogs, grass litter, dead wood on ground (especially driftwood), under stones and other objects, and ground surface active); other microhabitats are present in specific macrohabitat types. For macrohabitat definition, trees and some other conspicuous plant species were identified using prior experience in the southern Kurils and the resident expertise of expedition botanist Sarah Gage.

Main items of collecting equipment and techniques were: 1) heavy duty beating net, 45.7 cm diameter (18 inches) for vegetation sampling; 2) rigid plastic sifting box, 37 X 46 cm, with 1.27 cm (0.5 inch) screen mesh bottom, for moss and litter; 3) aspirator (used mainly by TWP); 4) trowel and cloth for rotten wood, soil surface web, and burrowing spider sampling; 5) dry vial collection (over net to minimize escapes) for under rock sampling; 6) visual searching for ground surface active specimens. KYE sorted moss and some forms of litter by hand on a plastic sheet. Due to the lack of large trees, bark and canopy sampling were not generally done. Because sites were not revisited, we were unable to use pitfall traps, a considerable loss in potential collection diversity. When weather conditions or shortness of time prevented adequate litter sampling on land, litter samples were brought back to the ship for later processing.

In data recording, macro- and microhabitats were recorded separately for each sample; geographic location was recorded in decimal latitude/longitude to the nearest 0.001 degree, based this year on the degree-minute-second readings of the Global Positioning Satellite (GPS) hand units; also recorded, date and approximate time of sampling. Samples were field labelled with full data by filling in blanks in preprinted labels with a Sakura Micron Pigma permanent ink pen. Checking of GPS-determined coordinates against maps showed that these data cannot be accepted uncritically; it is necessary to set the correct geodetic datum for the region in which one is working (of those available on our units, the Tananrive Datum proved the closest match for that used on Russian maps), and even so, the error in either latitude or longitude occasionally proved to be several times the nominal value of 100 m, especially on the island of Ekarma. Modern technology not withstanding, the ability to read a map is still of paramount importance in biological collecting.

Preliminary counts and identification of all spiders taken by American participants was done on the ship with a Wild M5 microscope and a card file of genitalia illustrations. However, due to limitations imposed by short available time, lack of full literature, and the motion of the ship, it will be necessary to re-examine most specimens for final identification. Accordingly, all identifications reported below should be considered preliminary and subject to revision. Many are incomplete.

NARRATIVE OF SAMPLING ACTIVITY

Paramushir Island

Paramushir, a large island which is probably the most faunistically diverse of the northern Kurils (from its size, proximity to Kamchatka, and topographic variety), was sampled on 2 days in 2 different areas. On 1 August 1996, an area accessible by road from the town of Severo-Kurilsk was visited; on 3 August, the expedition was landed on a low-lying, dune-derived peninsula at the south end of the island, from which the arachnological team hiked a considerable distance inland on fortunately accessible roads.

1 August: After a brief opportunity to collect from man-altered habitats at Severo-Kurilsk harbor, all participants were taken by Russian army truck (over promising-looking uplands, where we did not stop) to the valley of the Utyosnaya River, next major valley south of the city. Vegetation here established a pattern that was to become familiar over all the northern Kurils: lush (floristically dense, continuous ground cover) meadow habitats interspersed with relatively small thickets of low-growing Alnus maximowiczii. A typical thicket was perhaps 10-20 m in diameter and 1-2 m high. KYE and RLC visited a hillside where both thicket and meadow were sampled; a valley Sphagnum bog; and the shore meadow habitat at the river-mouth rendezvous for return to the ship. Microhabitats sampled included field layer vegetation, alder litter, Sphagnum, ground surface active, and under objects on sand. American participants (mainly RLC) took 269 spider specimens this day.

3 August: The expedition landed well out on the Vasil'yeva Peninsula, a low-lying southward extension of Paramushir probably originating in sand-dunes, though now well vegetated over much of its area. RLC, KYE and TK spent some time hiking on roads northward along the peninsula in order to collect in upland habitats at its base. Thanks to this effort, we considerably enlarged the Paramushir species list. (Through the rest of the expedition we seldom had another such chance to visit uplands on a large island). Macrohabitats encountered included shore and dune meadows; relatively extensive alder thickets (of which some were swampy); dry upland meadows (with many edible berries); extensive areas of Pinus pumila approximately 1 m high; lush riparian meadow along the Obryvistaya River; the rocky shore of this river; lake-shore meadow; Sphagnum patches and bogs. Microhabitats sampled included field layer vegetation, thicket understory vegetation, pine foliage, grass litter, Sphagnum (from hillside patch and valley bog, which yielded different faunas), and active on ground. This was the only day on which all three arachnologists worked together; for the remainder of the expedition, TK sought different habitats from RLC and KYE. American participants took 296 specimens, of which TWP, collecting closer to the peninsular landing site, contributed 94.

Total specimens brought to USA from Paramushir: 565.

Onekotan Island

More collecting effort (4 days) was devoted to this island than any other in the northern Kurils. The resulting collection contained many specimens, but barely more species than we obtained from Paramushir in half the time. Clearly, the latter has a richer spider fauna, but adverse weather and other conditions may have contributed to our less diverse sample from Onekotan also.

4 August. Landing was at Nyemo (Nemo, named after the Jules Verne character) Bay, a narrow inlet near the north end of the island which is the mouth of an unnamed river; upstream of some sand dunes and flats, the river ascends to the plateau through a rocky defile. RLC attempted collecting from rocky habitats at the latter site, with but little success, while KYE sifted moss in a more boggy section of the flats midway up the canyon. Others ascended a trail and crossed into the next valley to the south. Along the river were dune vegetation, at least one alder thicket, a few boggy patches, and lush riparian meadow with pure stands of various herbs including Urtica sp. Hillsides had meadows varying from lush to rocky. There was considerable bare and broken rock substrate in the "rocky defile." Sampling in some areas by some methods was rendered impractical by strong wind, and cold, cloudy weather kept the diurnally-active spiders, which otherwise might have been numerous, from appearing. Microhabitats sampled: under rocks, Sphagnum, meadow field layer, mixed moss and meadow litter. American participants took 223 specimens, of which TWP contributed 36 and B.K. Urbain, 37.

5 August. We had hardly landed on a beach near Cape Subbotyna, well down the east shore of the island, when it began to rain with increasing severity. An attempted ascent of a ravine to the plateau above was aborted when strong winds blowing across the island changed bad collecting conditions to impossible ones. Back in the ravine, where it was only raining, some spiders were collected under stones and a large moss sample was gathered for sorting back on the ship. Fortunately, the latter proved rich enough to prevent the day from being wasted. Habitats included meadow, stony ravine bottom with active stream, rocky hillside, and cobble/boulder upper beach. KYE ascended a different ravine and sampled in alder thickets. Microhabitats: under stones; sifted from moss and grass roots in sample returned to the ship. 156 specimens were taken by American participants, mainly RLC.

7 August. Landing at Mussel' Bay, on the east coast of the island opposite our previous site, we had access by road to the interior of the island and also to the adjacent river valley containing an abandoned army camp, "Onekotan village." RLC and KYE hiked about 2.5 km inland to a plateau 100 m in elevation, where dry and mesic meadows, dead wood along the road, alder thickets (especially in a ravine bottom), and a Pinus pumila clump were sampled. In the adjacent valley, KYE hunted unsuccessfully for a rumored Sphagnum bog while RLC found an amazing number of specimens under discarded lumber and sheet metal in the abandoned army camp. Rich riparian meadow was along the unnamed river here. Microhabitats sampled: meadow field layer, alder litter, pine litter, pine foliage, under and in dead wood, under manmade objects, active on ground. American participants took 323 specimens, of which 54 were contributed by B.K. Urbain.

9 August. An attempt at sampling the south end of the island did not turn out quite as expected. Apparently, morning fog prevented selection of a better landing site, and we landed in a narrow shoreline belt at the base of an extremely steep hillside near (but nearly out of reach of) the Trudnyi River. RLC sampled mainly shoreline habitats while KYE and others braved the steep climb and sampled in alder thickets high on the bluff. Habitats included a rocky headland with few plants but much broken rock habitat; lush hillside meadows blending into the shoreline meadow at the top of the beach; the beach itself; and the alder thickets high above. Tim Pearce, American malacologist, brought down a bag of litter from a high thicket for later sorting on the ship. Microhabitats included under stones, meadow field layer (this was one of very few sites in the northern Kurils where a numerous meadow sweep sample was obtained), active on rocks, and alder litter. American participants took 576 specimens, of which nearly half (245) were contributed by TWP who climbed out of the shoreline belt.

Total specimens brought to USA from Onekotan: 1278.

Kharimkotan Island

One day was spent on this island, unfortunately in the low old-sand-dune habitats of the northwest corner of the island rather than on the lava flows of the east end, where maps show many caves which were expected to have interesting fauna. It is not known what other habitats might have been found there or on the central peak.

8 August. Landed at Severgina Bay near Cape Ankuchi, the northwest point of the island. Rather than spend most of the day hiking over dunes to the interior, RLC elected to stay in this lowland peninsula. The spider fauna here proved better than expected for such a place, though with few surprises after Onekotan. Habitats visited included sandy-grassy dunes, interdune ponds and wetlands, a meadowy intermittent wetland (dry at that time) with band of Sphagnum, alder thickets, more diverse dune meadows, and extensive fields of driftwood on and between the foredunes. Microhabitats: field layer vegetation, alder litter, Sphagnum and other mosses, under drift logs and wood, active on ground. On this and later days TWP collected extensively from grass and grass litter. American participants took 574 specimens, of which nearly half (257) were contributed by TWP.

Chirinkotan and Ekarma Islands

Limited-size shore parties, not including this writer, landed on both these islands briefly on the same day. Judging from the limited collections brought back to the ship, the spider fauna of Chirinkotan may be the poorest in species of any island visited this year. The fauna of Ekarma should be richer due to more diverse habitats and presence of alder thickets, but this is not reflected in the present collection. Interesting results are expected from the collections (not seen by me) of KYE and TK. Most of the spiders brought back to the USA from the two islands were collected by TWP.

10 August. Morning: Chirinkotan proved to be a tall, roughly square rock rising almost vertically from the sea on all sides. Many of the steep slopes supported a lush meadow vegetation, fertilized (as was evident on closer inspection) by the abundant sea birds. Few shrubs were present. RLC was not a member of the shore party. TWP collected 84 specimens from grass and grass litter on the steep slope meadows, supplemented by other party members to make 111 in all taken by American participants. Afternoon: Ekarma had a somewhat gentler topography and the standard North-Kuril meadow and alder thicket habitats, but through an unfortunate communication failure, RLC did not land here either. Fortunately, both B.K. Urbain and Tim Pearce brought litter/moss samples back to the ship for me to sort. Habitats sampled for spiders were chiefly meadow vegetation (BKU), meadow litter (TWP), alder litter and moss (samples returned to ship). 323 specimens were taken by American participants, including 95 by TWP, 88 by BKU, and 117 in the samples brought to the ship.

Shiashkotan Island

We spent relatively little time on this fairly large island, the first day being cut short and the second made unproductive by rain; as far as could be judged from one area (both landings were at the same site), habitats were typical for the northern Kurils.

11 August. Landing at Zakatnaya Bay on the west shore of the "isthmus" of the island, we found fair (if moist) weather and old trails available to hike into the interior...but insufficient time to do so, since return to the ship was set for mid-afternoon. RLC collected in a ravine ascending the bluff, sampling in rocky habitats, an alder-Sorbus thicket, and grassy meadow, and in the driftwood and beach meadow below. KYE hiked far enough inland to find a Sphagnum bog, which, however, proved singularly unproductive. TWP collected around the remains of a former Japanese installation and even older Ainu village at the bluff top. Microhabitats: litter of Alnus, Sorbus, and Pteridium; Sphagnum; grass foliage and litter; under drift logs; in manmade pits; active on ground. On 12 August more time was available (albeit at the same landing site), but rain prevented effective spider collection; aquatic biologists brought back a few good spider specimens from a marathon hike into the interior. American participants took 361 specimens on Shiashkotan, including 175 by RLC and 147 by TWP.

Raikoke and Matua Islands

These two islands, about 20 km apart, contrast sharply in environment; Raikoke was the most barren and Matua the most lushly vegetated of the islands which were our main goals for this summer's work. Nevertheless, Raikoke proved to have more spider species than Chirinkotan or Ekarma, probably because of the proximity of Matua as a source of colonists.

13 August. In the confusion of difficult landings, RLC spent only a few minutes on this, probably the most barren island we visited with its cindery volcanic slopes. Birds were numerous, vegetation somewhat sparse and notably non-diverse. In about a half-day here, 198 spiders were taken by American participants, most (101) by TWP and mostly from litter habitats in patches of Angelica and other common plants.

14 August. After landing on Matua, we hiked along roads to the Russian army base here. Though the day was rainy, the availability of a dry room in which to sift leaf litter made it possible to collect a large number of spiders. On our progress southward, this was the first island encountered with continuous "forests" of alder "trees" (albeit only 3-4 m high), as opposed to small thickets of alder shrubs. This evidently makes a considerable difference for spiders, as several of the common species of the southern and central Kurils first appeared here. Since it continued raining most of the day, RLC collected primarily in the alder-forest and clearing habitats around the army base, while KYE scouted out other habitats and returned early. American participants took about 250 spiders this day, most by RLC.

15 August. Return to Matua. Weather remained warm and humid all day on the east coast of the island where we had worked on the 14th. Crossing the old airstrip to the south tip of the island to collect at a Sphagnum bog, we encountered foggy conditions together with a cold, biting wind. This surprising difference in weather was not accompanied by any noticeable topographic barrier. The bog proved productive; on the way back to the landing site, RLC took advantage of the better weather there to sample vegetation habitats, especially grassy meadows which yielded the second good "sweep" sample of spiders taken this year. Microhabitats: Sphagnum, grass field layer, under objects in clearing. In all (both days) American participants took 582 spiders on Matua, mostly by RLC.

Urup Island

After a side trip for water (noted below under Iturup), two days were spent on opposite shores of southern Urup (central Kurils), this being the first time on the island for all three 1996 arachnologists. On landing we were told by entomologist Arkady Lelej that this is the one Kuril island with abundant blackflies. He was right! Fortunately most of us had head nets, without which it would have been very difficult to work, as this blackfly species appears to be terrestrial and is not restricted to the vicinity of water. Perhaps because of this abundant prey, as well as the more varied (partly forested) habitats, spider collecting was relatively productive.

20 August. Landing at Ukromnaya Bay on the southeast shore of the south tip of Urup, most of the party proceeded up the Ukromnaya River among alder and willow woodland on the flood plain. The arachnologists climbed the hills on either side of the river mouth, RLC taking the southern and KYE the northern hill. On the hillsides were open woodland of alder and occasional other tree species, with a dense understory of Sasa (dwarf bamboo), interspersed with meadow areas of Sasa and occasional more grassy patches. The hill crests had some rocky habitat. Valley bottom forest and seashore environments were also visited. Microhabitats: Alder/Sasa litter; pure alder litter; field layer vegetation; under stones. American participants took 702 specimens, about half (364) by TWP.

21 August. We landed in initially good weather at Tyetyayeva (or Tetyaeva) Bay on the southwest shore of Urup, about opposite (across the island from) yesterday's landing. Habitats were different and roads were available to hike into the interior. RLC visited a ridge crest area with patches of meadow, alder forest, and dwarf-pine thicket. By this time strong wind prevented some collecting techniques, but litter sampling proved productive until mid-afternoon when it began to rain. Passage back to the ship proved extremely stormy. Microhabitats: Alder/Sasa litter, Pine/Sasa litter, meadow foliage. 331 spiders were taken by American participants, of which B.K. Urbain contributed 49.

Total specimens brought back from Urup this year: 1033.

Iturup Island

A need to replenish fresh water, inevitable in retrospect since the ship's tanks had not been full on starting, led us to visit Konservnaya Bay on Iturup (the standard ship watering place in these islands) in mid-expedition. After some work on Urup, another stop was made on Iturup en route to Kunashir. In each case, two days were partly spent on land, all in areas previously well-collected by RLC and KYE in 1994. Nevertheless, some additional species were taken, and the richer habitats of this southern island considerably augmented our specimen totals.

18-19 August. While the ship took on fresh water, we were allowed to revisit the familiar scenes of Konservnaya Bay. In efforts to visit subsites not previously sampled, KYE perilously climbed the steep bluffs to the plateau above while RLC used the bed of the main stream entering the head of the bay as a route through the impenetrable Sasa thickets to a higher zone of vegetation. Habitats visited by the latter included lush riparian forest of mixed tree species and Sasa; a higher belt of forest dominated by Betula ermanii, still with Sasa understory; rocky dry streambed; a well-drained sandy area amid the remains of the former cannery for which the bay is named. Microhabitats: Sasa litter; Betula-Alnus-Prunus-Sorbus litter; riparian forest understory foliage; under stones; active on ground. American collectors took 710 spiders, about half (379) by TWP.

22-23 August. En route southward, a two day stopover was made on the north side of Dobroye Nachalo Bay, where the spiders had been extremely well sampled in 1994. On the first day, RLC and KYE visited small Lake Natasha which we had not seen on the previous trip, sampling in swampy lakeside forest with maple, alder and true fir. Microhabitats included moss (on logs, roots and soil), litter, understory foliage, and under bark. On the second day, intensive collecting was neglected in favor of seeking out special habitats. RLC spent the day seeking additional caves in the lava flow (some unique spiders had been found in the one small cave found previously). Only one additional cave was found, but additional specimens of one species and our first mature specimens of two others resulted. KYE concentrated on finding synanthropic species in old military installations. American collectors took 464 specimens here this year, 162 of them by TWP.

Total specimens brought back from Iturup this year: 1174.

Kunashir Island

This island, one of the two richest in spider fauna in the entire archipelago (the other is Shikotan), merits as much additional work as can be managed. Even the one full day and one partial day of spider collecting done here in passing in 1996 were very beneficial and added several species and genera to the list.

25 August. KYE revisited the bog inland from Yuzhno-Kurilsk which had been sampled twice in 1994, in hopes of getting mature males of several species previously found as females. He found that the weather this year had caused flooding of the bog and greatly reduced spider fauna there, which was much "earlier" in seasonal status than would have been the case at this date in a normal year. Thus, this project was largely unsuccessful. RLC accompanied the ichthyologists to the trailhead for the "goby hot springs" in the foothills of Vulkan Mendeleeva. Available habitats included mixed deciduous forest, stands of Abies sachalinensis and Picea glehnii, an open Sasa-herb clearing, and roadside verge. Microhabitats sampled: field layer vegetation in clearing (the Sasa here had many spiders on it); Betula-Sasa litter; conifer foliage; forest floor moss; active on ground. Specimens collected by RLC (only American collector), 331.

26 August. RLC was the only arachnologist collecting this half-day, and was dropped off at a Sphagnum bog south of Lake Aliger on the west shore of the island, just inland from the dune belt. This bog was not flooded at the time, and looked extremely promising, but, as with KYE's experience the day before, the Sphagnum yielded few spiders, mostly immature. The spruce trees and bog meadow foliage were, fortunately, more productive. Specimens from this area: 146.

Total specimens brought back from Kunashir this year: 483.

RESULTS: PRELIMINARY SPECIES IDENTIFICATIONS, WITH NOTES

Abbreviation

Island

Specimens (to USA)

Days+Part Days

Species (USA specimens)

PA

Paramushir

565

2

57

ON

Onekotan

1278

4

58

KH

Kharimkotan

574

1

39

CR

Chirinkotan

111

0.5

14

EK

Ekarma

323

0.5

14

SA

Shiashkotan

361

2

30

RK

Raikoke

198

1

21

MA

Matua

582

2

33

Total, "new" Islands

3992

13

110

UR

Urup

1033

2

54

IT

Iturup

1174

4

73

KU

Kunashir

483

2

60

Total, "old" Islands

2690

8

129

Grand Total

6682

21

194

The species statistics given above and identifications given below are only estimates, from preliminary field ID. Many specimens have not been identified to species level yet and some other preliminary identifications will be changed. The final species numbers from this collection may be higher, and large additions are expected when the material collected by KYE and TK is added. The following discussions are not done island by island, but segregate this year's collection into two groups: the islands visited for the first time (Paramushir south to Matua) and those previously visited by IKIP-1994 or IKIP-1995 (Urup, Iturup and Kunashir).

Symbols: M = males, F = females, J = juveniles

North and North-Central Kuril Islands (Paramushir to Matua)

The following records mark the first collections by IKIP from any of these eight islands, and the first spiders known to have been collected on any of the seven islands from Onekotan to Matua. All or part of 13 days were spent collecting on these islands (mean 261 specimens per day).

Order Araneida

Family Amaurobiidae

Cybaeopsis typica, MA (FJ). This species, common on the southern Kurils, appeared for the first time on Matua on our route southward. There was no trace of amaurobiids on Raikoke northward, yet on Matua it is already common. Evidently the requirement for forest is the strongest determinant of the range of this species within the archipelago.

Family Tetragnathidae

Tetragnatha extensa, PA (MF), ON, SA. Common on all islands from Kunashir to Paramushir.

Family Araneidae

Larinioides patagiatus, PA (MJ), ON. Holarctic species previously found on Iturup.

Family Mimetidae

Ero furcata, ON (F). Second finding in archipelago (found on Iturup by IKIP-94).

Family Theridiidae

Enoplognatha tecta, PA (MJ), ON, KH, SA, RK, MA. Common on all islands from Kunashir to Paramushir.

Robertus lividus, PA (MF), ON (M), KH, CR (FJ), SA, RK, MA. Already known from Paramushir and Shumshu.

Theridion sp. (nr. tinctum), ON (J).

Family Linyphiidae

Agyneta allosubtilis, ON. Previously found in central Kurils, 1995.

Ainerigone saitoi, KH. Previously found in southern Kurils, 1994.

Baryphyma pini, PA. Previously found in southern Kurils.

Baryphyma sp. indet., possibly new, ON (MF), KH, CR, EK, SA, RK, MA.

Bathyphantes pogonias, ON, KH(F), EK, MA. Common on all islands, Kunashir to Paramushir, but more so in south Kurils.

Bathyphantes setiger, PA (F), ON, CR, RK. Previously recorded from Shumshu.

Bathyphantes sp. indet., PA (M).

Centromerus sylvaticus, PA (F), ON (F), KH, CR, EK, SA, RK, MA. This species proved to be common from Paramushir south to northern Iturup. Absent in southern Iturup and Kunashir, where it is replaced by C. terrigenus, which was found north to southern Urup.

Ceraticelus sibiricus, PA (F). New to archipelago.

Ceraticelus sp. indet. (red-brown), ON (F), KH. ON-96-RLC-017 and -020.

Ceraticelus sp. indet. (black), ON (F). ON-96-RLC-017.

Ceratinella sp. 1, ON (F), CR (M), SA. Undescribed species found in south Kurils by IKIP-94.

Ceratinella sp. 2, PA (J), ON (FJ), KH, EK, SA, RK. Undescribed species found on Iturup by IKIP-94.

Cnephalocotes obscurus, KH (M). New to archipelago. The finding of this specimen inadvertently led to the discovery of a junior synonym of this palaearctic species: the Japanese Nematogmus rutilis Oi.

Diplocentria bidentata, PA (F), KH (M). New to archipelago.

Diplocentria sp. indet., ON, MA.

Diplocephalus sp. nov., PA (M/KYE), ON (MF), EK.

Dismodicus sp., PA (F). A member of the difficult-to-identify bifrons group.

Eboria lapponica, PA. Holarctic species new to archipelago.

Erigone atra, MA. Known from Paramushir to Kunashir but more common in the southern Kurils.

Erigone capra, PA, ON. Found on Iturup by IKIP-94.

Erigone simillima, PA (M). Already known from Paramushir, but the male is undescribed.

Erigone sp. indet. (aletris group). PA, ON, KH.

Gnathonarium suppositum, PA (MF), ON, KH, CR, EK, SA, RK, MA. Common on all islands, Kunashir to Paramushir

Hilaira herniosa, PA. Previously known from Shumshu.

Hilaira leviceps, RK (M). Previously known from Shumshu.

Hilaira nubigena, PA. New to archipelago.

Hilaira sp. indet., ON, SA, RK.

Hypomma affine, PA (MF), ON, KH, SA, MA. Found (not very common) Kunashir to Paramushir.

Hypselistes basarukini, PA. New to archipelago.

Lepthyphantes bipilis, PA (F), KH, RK. New to archipelago.

Lepthyphantes nebulosus, MA (M). A synanthropic species found on the Army post; previously known from Iturup.

Lepthyphantes nigriventris, ON, MA. Already known from Paramushir to Iturup.

Lepthyphantes sp. indet., PA (M).

Lepthyphantes sp. indet. (tenuis group, brown), PA (F).

Lepthyphantes sp. indet. (tenuis group, patterned), PA (F), ON (MF), KH, EK, SA.

Lepthyphantes sp. indet., CR (J), SA (J).

Leptorhoptrum robustum, PA (F), ON, KH, CR (MF), EK, SA, MA. Common on northern Kurils, occurs south to Iturup.

Leptorhoptrum sp. nov., PA (F).

Maro sp., indet., PA (F).

Meioneta sp. indet., KH, CR, RK (M).

Micrargus herbigradus, KH. Previously known from Paramushir.

Nippononeta kurilensis, ON (F), SA, RK, MA. Previously known only from Kunashir.

"Oedothorax trilobatus" Banks, PA (M/KYE). The finding of this Nearctic species, incorrectly placed in Oedothorax, in the Kurils suggests that its true affinities lie with Palaearctic fauna. Its taxonomic status will be investigated by KYE. New to Palaearctic region.

Oreonetides vaginatus, PA(F). Determined as such by KYE, but could be distinct. Already known from Paramushir. SA (F), typical female.

Poeciloneta sp., PA (F). New to archipelago.

Porrhomma hakusanense, ON. New to archipelago.

Porrhomma pallidum, PA. Found on Kunashir and Iturup by IKIP-94.

Savignya saitoi, ON (MF), KH, CR, EK, SA, RK. Previously known only from south Kurils, this species proved to be common from Onekotan southward, but was not found on Paramushir.

Scotinotylus alienus, ON, KH. New to archipelago.

Scotinotylus sp. nov. (nr. antennatus), ON(F; M/KYE), KH, CR, RK, MA.

Sisicottus sp. nov. #1, ON (MF/KYE) (coll. independently later on PA by Y.Marusik). One of the most remarkable results of IKIP-96 is the finding of two new species in this genus, formerly thought to be of Nearctic-only distribution. These species are already under study by Jeremy Miller of George Washington University.

Sisicottus sp. nov. #2, ON (MF), KH, CR, EK, SA, MA.

Tiso aestivus, ON (MF). Previously known from Iturup.

Tmeticus sp., PA (MF), MA (M). Either T. japonicus (south Kurils) or T. tolli (new to archipelago).

Tunagyna debilis, PA, ON, KH. Already known from Paramushir and Shumshu.

Ummeliata sp. nov., PA (F), ON (F).

Ummeliata angulitubera, MA (MF). Common in southern Kurils, this is the northernmost record. See discussion under Cybaeopsis typica.

Walckenaeria clavicornis, KH. Found on Iturup by IKIP-94.

Walckenaeria cuspidata, PA (F), ON, MA. Previously found on Shumshu.

Walckenaeria golovatchi, ON. Previously known from south Kurils.

Walckenaeria karpinskii, RK, MA. New to archipelago.

Walckenaeria nudipalpis, PA, SA. Found on Zelionyi by IKIP-94.

Walckenaeria sp. indet. (nr. lepida), SA, MA (MF).

Walckenaeria sp. indet. (spiralis group), ON (M), KH(M), SA (M).

Walckenaeria sp. indet. (red), ON.

Sp. indet. (like Drepanotylus), PA (F). In PA-96-RLC-004.

Sp. indet. with minute scape, PA (F). In PA-96-RLC-010.

Sp. indet. bilobed, PA (F). In PA-96-RLC-010.

Sp. indet. (like Ainerigone), ON (F). In ON-96-RLC-020.

Sp. indet. (like Eulaira), ON (F). In ON-96-RLC-020.

Sp. indet. (dark area, round notch), ON (F). In ON-96-RLC-023.

Sp. indet. (rectangular notch), ON (F), KH (F). In ON-96-RLC-023, KH-96-TWP-015.

Sp. indet. (Meioneta-like), ON (F). In ON-96-RLC-023.

Sp. indet. (Porrhomma-like), EK (F). In EK-96-TWP-023.

Sp. indet. (large), MA (F). In MA-96-RLC-043.

Sp. indet. (small pale), MA (F). In MA-96-RLC-044, MA-96-TWP-030 (different?).

Family Lycosidae

Pirata piraticus, PA (F). Previously found by IKIP in southern Kurils.

Pardosa riparia, PA (F), KH. This species and P. palustris, atrata, prativaga, sibirica were found together in the same Sphagnum bog, first day on Paramushir.

Pardosa paramushirensis, SA. Described from Paramushir by Nakatsudi (1937), this species was recently rediscovered in Japan (Chikuni 1989a). This finding shows that the distribution is not disjunct. Apparently it is limited to habitats found on volcanic peaks, thus is not often found by largely shore-limited IKIP work.

Pardosa palustris, PA (F), ON (F), KH, SA, RK, MA. Holarctic species, common throughout Kurils.

Pardosa atrata, PA (F). New to archipelago.

Pardosa prativaga, PA . New to archipelago.

Pardosa sibirica, PA. New to archipelago.

Pardosa sp. indet. (black carapace), KH (J).

Trochosa terricola, PA (FJ), ON, KH, SA, MA. Common on all islands, Kunashir to Paramushir, but more so in the northern and central Kurils.

Family Gnaphosidae

Haplodrassus signifer (or related species), PA (J), ON (FJ), KH, SA (F). Already known from Paramushir; our specimen from there was kept alive for rearing.

Family Clubionidae (sensu lato)

Cheiracanthium erraticum, PA (FJ), ON (MFJ). Previously found on central Kurils and Iturup; immatures, probably this species, known from Kunashir and Shikotan.

Clubiona latericia, PA (M), ON. Found on Kunashir and Zelionyi by IKIP-94.

Clubiona mayumiae, ON, KH (MF), CR, EK, SA, MA. Previously found on Iturup, south to Japan.

Clubiona riparia, KH (M), SA (F). Known from Paramushir to Kunashir, but not very common.

Clubiona yagata, KH. New to archipelago.

Clubiona sp. (bilobed female), ON (F).

Clubiona sp. indet., SA (F).

Clubiona sp. indet., RK (J), MA (J).

Family Thomisidae (sensu lato)

Lysiteles sp. indet., ON (J).

Ozyptila sincera, PA (MJ) ON, SA, RK, MA. New to archipelago.

Ozyptila trux, PA (M), RK. Previously known from Shumshu; found on Kunashir and Iturup by IKIP-94.

Ozyptila sp. indet., KH (J).

Tibellus oblongus, PA (J), ON, MA. Previously known from south Kurils; holarctic species.

Xysticus sp., PA (J), ON (J), MA (J).

Xysticus cristatus, ON (F). New to archipelago.

Order Phalangida

Family Ischyropsalididae

Nipponopsalis yezoensis, ON (MFJ), CR (M), EK, SA, RK, MA. Previously known from south Kurils.

Family Phalangiidae

Indet. sp., PA (F).

Homolophus arcticus, PA (J), ON. Common on all islands, Kunashir to Paramushir, chiefly in beach habitats.

Mitopus morio, PA (MF), ON, KH, RK, MA. Moderately common on northern Kurils, somewhat more so on central Kurils south to Itururp.

Mizozatus flavidus, ON (MFJ), KH (J), CR (J), EK (MJ), SA(FJ), RK (MFJ), MA (FJ). This species, described from Paramushir by Nakatsudi (1937), has been "lost" for over 50 years, remaining unknown even to cataloguers, before being re-discovered this year by IKIP. It is a distinct species of the subfamily Opilioninae, with affinities to the genus Opilio, but it is possible that even the genus may be valid. If so, it would be the only known genus endemic to the Kuril Islands (unless and until it is found in Kamchatka).

Phalangium opilio, MA (F). A synanthropic species found on the Army base.

Family Sclerosomatidae

Leiobunum globosum, PA (FJ), previously known from south Kurils and Japan.

Urup, Iturup, and Kunashir Islands

The following records are from this year's work on these three islands which had previously been visited by IKIP in 1994 and/or 1995. In all, 8 days, including two half-days and one on which little collecting was done, were spent on these islands this year (mean 336 spiders per day). The greater number directly reflects the lusher and more varied vegetation and more clement weather found in the southern Kurils, which also can boast of a substantially greater spider species richness. Thus, the additional work here was by no means uncalled-for, though even better results would have been obtained by visiting new sites.

Order Araneida

Family Amaurobiidae

Cybaeopsis typica, UR, IT. Previously found on these islands.

Family Dictynidae

Dictyna sp. indet., KU (J).

Family Tetragnathidae

Dyschiriognatha quadrimaculata, IT (MF), KU (F). Previously found on Kunashir.

Meta sp. indet., UR (J), IT (FJ). Probably a species endemic to Japan and the Kuril Islands.

Metleucauge sp. indet., UR (J), IT (J).

Tetragnatha dearmata, IT. New island record; previously known from Kunashir. Holarctic.

Tetragnatha extensa, UR, IT. Common on all islands from Kunashir to Paramushir.

Tetragnatha pinicola, KU. Previously found on south Kurils.

Tetragnatha yesoensis, KU. Previously found on Kunashir.

Family Araneidae

Araneus sp. indet., UR (J).

Araneus marmoreus, KU (MF). Previously known from Kunashir, but this is first IKIP collection.

Araneus sp., probably nov. (near triguttatus), IT. This species was described from Japan (but unfortunately not named!) by Chikuni (1989b). Also found on Shikotan by IKIP-94.

Singa hamata, IT. Found by KYE in the original small cave at Dobroye Nachalo Bay. New to archipelago.

Larinioides sp. indet., IT (J), KU (J).

Yaginumia sia, IT (F), KU (J). New to archipelago. New genus for Russian territory.

Zilla sachalinensis, IT (F), KU. Previously known from south Kurils; not yet found in the north.

Family Theridiosomatidae

Theridiosoma epeiroides, IT (M). New island records; previously found on Kunashir.

Family Theridiidae

Achaearanea albipes, IT (F), KU (F). New island record; common on Kunashir.

Achaearanea tepidariorum, IT (F). Found by KYE at Dobroye Nachalo Bay; new island record of this synanthropic species, previously found on Kunashir and Zelionyi.

Argyrodes saganus, IT, KU. Previously known from south Kurils.

Dipoena mustelina, KU. Already known from south Kurils.

Enoplognatha tecta, UR, IT. Common throughout Kurils.

Robertus ungulatus, UR. Previously known from south Kurils, Sakhalin, Japan...and a disjunct distribution in Europe! Replaced in north Kurils by R. lividus.

Theridion bimaculatum, KU. Previously found on Kunashir and Sakhalin.

Theridion nigrolimbatum, UR, IT, KU. Common in south Kurils; Urup is its northernmost island.

Theridion sterninotatum, KU. New to archipelago.

Theridion sp. indet. (perhaps subpallens), UR. May be new to archipelago.

Family Nesticidae

Howaia brevipes, UR (MF), IT, KU. Common in south Kurils; Urup is its northernmost island.

Howaia sp., possibly different, IT (F).

Family Linyphiidae

Ainerigone saitoi, IT, KU. Common in south Kurils; this year also found on Kharimkotan.

Aprifrontalia mascula, IT. New to archipelago.

Baryphyma sp. indet., possibly new, UR.

Bathyphantes pogonias, UR, IT. Common on all islands, Kunashir to Paramushir, but more so in south Kurils.

Carorita sp. nov., UR (MF). Previously found on south Kurils by IKIP-94.

Centromerus sylvaticus, UR. Common from Iturup northward.

Centromerus terrigenus, UR, IT, KU. Common from southern Urup southward; the two only seem to be sympatric on southern Urup and Iturup.

Ceratinella sp. nr. rosea, UR. If correct, new to archipelago.

Ceratinella brevis, IT. New to archipelago.

Ceratinella sp. 2, UR, IT. Undescribed species previously found in south Kurils.

Dactylopisthes sp. nov., UR (M), IT (M). Found in northern Kurils by KYE this year also.

Diplocentria sp. indet., UR (F), IT (F).

Diplocentria, second species (truncate scape), UR (F).

Gnathonarium suppositum, UR, IT, KU. Common on all islands, Kunashir to Paramushir.

Gonatium nipponicum, UR, IT, KU . Known from south Kurils; Urup is its northernmost island.

Herbiphantes cericeus, IT, KU. Previously found in south Kurils.

Hypomma affine, IT. Found (not very common) Kunashir to Paramushir.

Lepthyphantes bipilis, UR. New to archipelago (also found this year in northern islands).

Lepthyphantes nigriventris, UR (MF). Already known from Paramushir to Iturup.

Leptorhoptrum robustum, UR, IT. Common on northern Kurils, occurs south to Iturup.

Leptorhoptrum sp. indet., possibly an atypical robustum, UR.

Micrargus apertus, UR, KU (M). New to archipelago.

Neriene angulifera, IT Previously known from south Kurils.

Neriene emphana, IT, KU. Abundant in south Kurils but not yet known from the north.

Neserigone basarukini, KU. Common in south Kurils; not yet found in the north.

Nippononeta kurilensis, UR. Previously known from south Kurils only.

Oreonetides shimizui, UR, IT. Known from south Kurils; Urup is its northernmost island.

Savignia saitoi, UR, IT, KU. Previously known only from south Kurils, this species proved to be common from Onekotan southward, but was not found on Paramushir.

Strandella fluctimaculata, KU (F). New to archipelago.

Strandella sp. indet., KU (F). Probably one of the species already known from this island.

Taranucnus nishikii, IT. Found in both small caves at Dobroye Nachalo Bay. Previously known from this area, Kunashir, and Japan.

Tmeticus japonicus, UR, IT, KU. Previously known from south Kurils.

Tmeticus sp. indet. (possibly 2 spp.), UR (M), IT (F).

Ummeliata angulitubera, UR (MF), IT. Common in south Kurils.

Ummeliata osakaensis, UR, IT. Common in south Kurils.

Walckenaeria golovatchi, UR, IT. Previously known from south Kurils; found this year also on north Kurils.

Walckenaeria karpinskii, UR. New to archipelago (also found this year in north Kurils).

Walckenaeria mayumiae, IT (F). New to archipelago, previously known only from Japan.

Wubanoides septentrionalis, IT, KU. Very common in south Kurils but not known from the north.

Indet. sp. (transverse oval plate), IT (F). In IT-96-TAP-072.

Indet. sp. (bright color, short truncate scape), IT (F), KU (F). In IT-96-TWP-033, KU-96-RLC-068 (possibly different).

Indet. sp. (rectangular notch; also in north), IT (F). In IT-96-TWP-040.

Indet. sp. (large dark oval plate with arcs), KU (F). In KU-96-RLC-065.

Indet. sp. (central atrium), KU (F). In KU-96-RLC-066.

Indet. sp. (Spirembolus-like), KU (F). In KU-96-RLC-066.

Indet. sp. (converging plate edges), KU (F). In KU-96-RLC-066.

Indet. sp. (appearance of intersecting plate edges), KU (F). In KU-96-RLC-069.

Indet. sp. (Tmeticus-like), KU (F). KU-96-RLC-069.

Family Agelenidae (sensu lato)

Cryphoeca silvicola, UR. First recorded from archipelago on Kunashir and Iturup by IKIP-94.

Cybaeus basarukini, UR (MF), IT (MF). Species previously known from Kunashir and Iturup. Genus is common in south Kurils; Urup is its northernmost island.

Cybaeus striatipes, UR. Formerly known from the original description of Bösenberg and Strand (1906), "Nord-Japan" (Hokkaido), this species was rediscovered by IKIP-94 on Shikotan. Urup is the farthest north site to date.

Family Hahniidae

Neoantistea sp. nov., UR (MJ). Previously found on Iturup and Zelionyi.

Family Lycosidae

Pardosa astrigera, KU (F). Previously known from south Kurils.

Pardosa lugubris, IT. Previously known from south Kurils; not yet found in the north.

Pardosa riparia, IT. Previously known from south Kurils; also found this year on Paramushir. One unusual specimen from Iturup may be different.

Pirata piraticus, UR, KU (J). Previously found by IKIP in southern Kurils; found this year also on Paramushir, so this Holarctic species is now known to be distributed throughout the chain.

Trochosa terricola, UR, IT. Prior to IKIP, the only spider recorded from Urup.

Xerolycosa nemoralis, IT. Previously known from south Kurils. A spider of dry, well-drained sites.

Family Anyphaenidae

Anyphaena pugil, KU (F). Previously found on Kunashir and Sakhalin.

Family Clubionidae (sensu lato)

Agroeca sp. indet., IT (J).

Cheiracanthium erraticum, UR, IT, KU. Previously found on central Kurils and Iturup; found this year in northern Kurils.

Clubiona sp., possibly amurensis, IT (F), KU (F). New to archipelago.

Clubiona ezoensis, UR, IT. Known from south Kurils; Urup is its northernmost island.

Clubiona kunashirensis, KU. Previously found on Kunashir and Iturup.

Clubiona kurilensis, UR. Previously known from south Kurils only.

Clubiona mayumiae, IT, KU. Previously found in south Kurils, this year first recorded from the north.

Clubiona propinqua, UR. May be new to archipelago, but female needs dissection for final ID.

Clubiona riparia, IT (MF). Previously known from south Kurils. Many taken in light trap; one unusual specimen may be different.

Clubiona sapporensis, IT. Abundant in south Kurils; not known north of Urup.

Family Gnaphosidae

Drassodes lapidosus, UR. Found on rocky hilltop; previously known from Kunashir.

Drassyllus sp. indet., IT (J), KU (J). A species of this genus has been recorded from Kunashir, but has not been taken by IKIP.

Haplodrassus sp. indet., UR (J). This genus has not been found on the southern Kurils.

Kishidaia albimaculata, KU (MJ). New to archipelago and new genus for Russian territory.

Micaria pulicaria, IT (F). New to archipelago; also found on Urup by KYE, who found a different Micaria species in the northern Kurils this year.

Family Thomisidae (sensu lato)

Diaea subdola, KU (M). Previously known from Kunashir.

Lysiteles maior, IT, KU. Previously known from south Kurils.

Misumenops sp. indet., KU (J).

Ozyptila sp. indet., IT (J), KU (J).

Philodromus cespitum, KU. Holarctic species previously found in south Kurils.

Philodromus sp., rufus group, KU (J). Previously found on Kunashir.

Tibellus oblongus, UR (J), KU. Holarctic species, now found in all parts of archipelago.

Xysticus sp. indet., UR (J), IT (J), KU (J).

Xysticus audax, IT (F). Previously known from Kunashir; first recorded this year in the north.

Xysticus kurilensis, IT (F). Previously known from south Kurils.

Xysticus sp. indet., KU (M).

Family Salticidae

Marpissa pomatia, KU (M). Previously found in south Kurils.

Neon reticulatus, IT, KU. Previously found in south Kurils.

Phintella linea, KU. Previously known from south Kurils.

Salticus sp. indet., KU (J). Genus new to archipelago.

Sitticus eskovi, IT (J). Previously known from south Kurils.

Yaginumaella ususudi, UR, IT, KU. Common in south Kurils; Urup is northernmost island for this species and its family.

Yaginumaella striatipes, IT. Previously found on Kunashir; new island record.

Order Phalangida

Family Ischyropsalididae

Nipponopsalis yezoensis, IT (F), KU (J). Previously found in south Kurils; this year first recorded from the north.

Sabacon sp. indet. (probably makinoi), UR (FJ), IT (J), KU (J). Genus not found on northern islands.

Family Caddidae

Caddo agilis, IT. New island record for this remarkable relict genus, known from Japan and the south Kurils; New England; and Baltic amber! An IKIP-94 record from Kunashir may be a second species, C. pepperella; both are known from Hokkaido (and New England!).

Family Phalangiidae

Homolophus arcticus, UR, IT. Common on all islands, Kunashir to Paramushir, chiefly in beach habitats.

Mitopus morio, UR (M). Moderately common on northern Kurils, somewhat more so on central Kurils south to Itururp.

Mizozatus flavidus, UR (J). See notes in list for northern islands. Urup appears to be its southernmost occurrence.

Oligolophus aspersus, IT, KU. Very common in south Kurils but not found from Urup northward.

Phalangium opilio, IT. Previously known from south Kurils; recorded this year also from Matua.

Family Sclerosomatidae

Leiobunum globosum, IT (FJ), KU (numerous). Common in south Kurils, this year first recorded from the north.

Nelima genufusca, IT (M), KU (numerous). Common in south Kurils, not yet recorded from the north.

PRELIMINARY DISCUSSION

Completeness of Data

As with previous years' collecting in other parts of the archipelago, our sample this year from the northern and north-central Kuril Islands gives a very incomplete picture of the total spider fauna.

Taking the data gathered by our expedition and that published by Marusik et al. (1992) together, RLC is reasonably sure that no one of the islands visited has as much as 50% of its spider species now known. Indeed, aside from Paramushir, the northern islands visited this year had no previous spider records at all. The most time we spent on any northern island this year was on Onekotan, 4 days. All of these days were in the first part of August (in the north temperate zone, generally the poorest time of year for spiders), and on only one day were weather and access conditions conducive to a really good day's collecting. The area of Onekotan is over 450 square km, with a patchwork of many habitats, only a few of which we had time to visit. The range of elevations is from sea level to 1019 m, but all spider samples were taken below 100 m. Most spider species have a limited season of maturity (including subnivial species found only in winter), and we have sampled only one season, and that one far from the most productive. Considering all these factors, it is possible that the 58 species herein reported from Onekotan represent as little as 10% of the total fauna of the island; I would say it could not be more than 30%. With pre-IKIP records added in, Paramushir now has more spider species recorded (65) than any other northern Kuril island, but it is also the largest and most diverse, so similar constraints apply there. There is a potential to improve our knowledge of the spiders of Paramushir in 1997, but a return to the islands from Onekotan to Matua is not planned under the present project.

Although the faunal lists generated this year are thus extremely incomplete, they do provide us with data for comparisons, since all the collections were taken in the same month under similar limitations. Also, taxonomic and single-species or pair-of-species zoogeographic work can be undertaken on species that have been found, even though most of the spider fauna remains unknown. What we will not be able to do is any work which requires reliable species absence data, species richness data, or comparisons of overall faunas. All of the data we have taken are important, since they fill in many gaps in species distributions, reveal the existence of new species, and provide knowledge of habitat (both microhabitat and macrohabitat) for most of the species collected for the first time.

Taxonomic Information

A number of undescribed species are represented in the above lists from 1996 collecting. More may be expected among the species not yet fully identified. Of the species already known or strongly suspected to be new, 7 have not been taken in previous years whereas 4 were previously taken in the south Kurils and have now had their ranges extended northward.

NEW THIS YEAR (All species new this year are Linyphiidae)

  1. Baryphyma sp. nov. ON to MA inclusive, UR
  2. Dactylopisthes sp. nov., PA to UR
  3. Diplocephalus sp. nov., PA to EK
  4. Leptorhoptrum sp. nov., PA
  5. Scotinotylus sp. nov., ON to MA
  6. Sisicottus sp. #1, PA to ON
  7. Sisicottus sp. #2, ON to MA
  8. Ummeliata sp. nov., PA, ON

NEW PREVIOUS YEARS, RANGE EXTENDED

  1. Ceratinella sp. 1 (north, central, south)
  2. Ceratinella sp. 2 (north, central, south)
  3. Carorita sp. nov. (south and now Urup)
  4. Neoantistea sp. nov. (south and now Urup)

The following species taken this year are of special taxonomic interest for reasons other than newness or simple range extension:

Erigone simillima. The male of this species, taken this year with females, has been undescribed. Description of the male will enable placement of this obscure species in proper relation to its congeners.

Leptorhoptrum sp. nov. and Carorita sp. nov.: These genera have previously been considered monotypic so the finding of new species is of special importance for both taxonomy and historical zoogeography.

Micrargus spp. Two species found this year in the north and central Kurils resemble M. herbigradus and M. apertus, members of a complex of species all of whose members have been described from Europe. This material will permit evaluation of whether the species of this complex in east Asia are actually distinct.

Neoantistea sp. nov. Range extension of this new species to Urup is timely, as its description is planned for the very near future.

"Oedothorax trilobatus" Banks. This species, like others described from North America in the genus Oedothorax, does not belong in that genus. It may very well be a new genus, but finding the species in the northern Kurils opens the possibility that it may have Asian relatives. KYE is investigating this question. The Kuril specimens are nearly identical to those in RLC's collection from Washington, USA.

Scotinotylus sp. nov. This species is related to the European S. antennatus and the North American S. eutypus. Its study should provide new clues to the relationships, evolution and historical zoogeography of this species complex.

Sisicottus spp. nov. Finding of these species is of peculiar interest since the genus has heretofore been found only in North America. These species are very closely related to the western North American S. panopeus (see Miller 1996). Indeed, J. Miller, who is now examining them, finds them so close that the relationship may be at the subspecies level, though RLC is inclined to disagree. In any case, they will be very important to Miller's conclusions about the genus, which will be submitted for publication early in 1997.

Pardosa paramushirensis. This species was originally described from Paramushir (Nakatsudi 1937) but remained "lost" for many years until rediscovered recently in Japan (Chikuni 1989a). Our (P. Oberg's) finding it on a mountaintop in Shiashkotan shows that the distribution is not wholly discontinuous, and suggests that the species occurs at similar sites on other islands, not sampled to date because of the logistical limitations of IKIP.

Haplodrassus signifer. These specimens, possibly not true H. signifer, will be of considerable importance to a forthcoming study of the group by our IKIP colleague Yuri Marusik.

Yaginumia sia and Kishidaia albimaculata. These Japanese species found this year in the southern Kurils add two additional genera to the fauna of Russian-held territory.

Mizozatus flavidus. This harvestman, described from Paramushir by Nakatsudi (1937), has remained unknown to science ever since. Indeed, the original publication was so obscure that it was omitted (I regret to state) from the world catalogue of the group by Crawford (1992). Earlier this year, RLC rediscovered it in IKIP-1995 material from Urup. It proved to be the most common phalangiid harvestman on the islands from Onekotan to Raikoke. The resulting wealth of material will make it possible to evaluate the genus Mizozatus and its relationships. If the genus proves to be valid, it would be the only known genus endemic to the Kuril Islands (unless and until it is found in Kamchatka).

Caddo spp. At least one, possibly two species of this harvestman genus whose distribution is decidedly relictual have been found on the South Kurils now, respectively on Kunashir and Iturup. The question of how they got there is the greatest unsolved puzzle to emerge to date from the arachnid results of IKIP.

Zoogeographic Relationships

In addition to species new to science, at least 34 species (almost certainly, more are among the presently unidentified material) were newly recorded for the Kuril archipelago and even more for individual islands; species recorded for Onekotan, Kharimkotan, Chirinkotan, Ekarma, Shiashkotan, Raikoke and Matua were the first spiders to be recorded from those islands.

Having now visited the entire length of the Kuril Archipelago, we can begin to identify north-south distribution patterns of species and higher taxa. It is, perhaps, a bit early to do this for many species, since the collection is so incomplete. However, a summary of the presently known north-south distribution of spider and harvestman taxa at the family level is interesting and is presented in the table below.

NORTH-SOUTH DISTRIBUTION PATTERNS

OF ARACHNID FAMILIES IN THE KURIL ARCHIPELAGO.

Notes: the KU column includes records from Shikotan and Zelionyi as well as Kunashir. Records from these islands and Iturup include literature records. S-CENT indicates the islands from Urup north to Rasshua, sampled mainly by IKIP-95. The PA column also includes literature records (see Appendix) which are from Paramushir, Shumshu, and Atlasova (Alaid). Many of the data in this table (even from the southern and central Kurils) were obtained in 1996.

KU IT S-CENT MA/RK SA EK/CR KH ON PA

SPIDERS

Amaurobiidae X X X X
Dictynidae X X
Pholcidae X X
Tetragnathidae X X X X X X
Araneidae X X X X X
Mimetidae X X X
Theridiosomatidae X X X
Mysmenidae X
Nesticidae X X X
Theridiidae X X X X X X X X X
Linyphiidae X X X X X X X X X
Pisauridae X
Lycosidae X X X X X X X X
Agelenidae (s.l.) X X X
Hahniidae X X X
Anyphaenidae X X
Clubionidae (s.l.) X X X X X X X X X
Gnaphosidae X X X X X X X
Philodrominae X X X X X X
Thomisinae X X X X X X X X
Salticidae X X X
HARVESTMEN
Nipponopsalididae X X X X X X X
Ischyropsalididae X X X
Caddidae X X
Phalangiidae X X X X X X X X X
Sclerosomatidae X X X

Among patterns apparent from this table:

  1. Only 9 families (of 25 listed here) occur throughout the archipelago. Of these, only 4 have been found on each island/group from which we have records (several being absent from the very small samples from Chirinkotan/Ekarma, Kharimkotan, or Shiashkotan).
  2. Three additional families (Mimetidae, Theridiosomatidae, Sclerosomatidae) are recorded from the southern and northern Kurils but have not been found in the central or north-central islands. We have too few data yet to be even reasonably certain this is not an artifact of incomplete collecting. However, it is possible that 1) the northern records of Theridiosomatidae, being literature citations only, may be incorrect; 2) the harvestman family Sclerosomatidae may be actually absent from those islands farther from mainland, being limited in dispersal ability.
  3. The spider families Amaurobiidae, Dictynidae, Pholcidae, Mysmenidae, Nesticidae, Agelenidae, (Hahniidae), Pisauridae, Salticidae; and the harvestman families Ischyropsalididae (genus Sabacon) and Caddidae (genus Caddo) have not been found north of various limits in the southern or south-central Kurils. Hahniidae is, however, known from Kamchatka and may be expected in the north Kurils. Most of these groups are relatively limited in dispersal power; the major exception to this statement is Salticidae, which are known worldwide as a group largely limited to warmer climates.
  4. The crab-spider subfamily Philodrominae (mistakenly considered a full family by most authors) is fairly diverse in the south Kurils but is represented in the central northern islands by only one species, the holarctic Tibellus oblongus.

As previously noted (our field report for 1994; Crawford in press), many arachnid species found in the southern Kurils and northern Japan show a closer relationship with the North American fauna than with most of Eurasia. However, the relationships are all at the genus level, the species being different, except with holarctic species that occur widely in both eastern and western hemispheres. In the northern Kurils, the situation is different: a number of species found here (and in Kamchatka, Chukotska, and some immediately adjacent parts of Siberia) are otherwise North American in distribution. Others may be related to North American fauna on a subspecific level. Those known to date are listed below:

NEARCTIC SPECIES AND GENERA FOUND IN THE NORTH KURILS BUT ABSENT FROM MOST OF EURASIA

Taxon

Palearctic Distribution

Nearctic Distribution

"Oedothorax trilobatus" N. Kurils only Rocky Mtns. Washington
Phlattothrata parva Kolyma, Kamchatka,Chukotska, N.Kurils Alaska to Washington and Rocky Mountains
Sisicottus spp. (orites group) N. Kurils only Northwestern North America

It will be noted that all belong to the family Linyphiidae. Preliminary study of the Sisicottus spp. collected this year (first palaearctic record of the genus) indicates that they are very close to the western North American Sisicottus panopeus and may be only subspecifically distinct.

By contrast, here are some linyphiid species found in North America and a somewhat wider portion of north-eastern Asia:

Taxon

Palaearctic Distribution

Nearctic Distribution

Dactylopisthes video Much of Siberia plus eastern Russian coast, Mongolia, N. Kurils Alaska, arctic Canada
Hilaira leviceps Across Siberia, Chukotska, N. Kurils Alaska to central Canada
Hypselistes semiflavus Central to east Siberia, N. Kurils Yukon
Tunagyna debilis Central to east Siberia, Kamchatka, N.Kurils New York to Alaska
Walckenaeria lepida Western Siberia to Russian Far East, N.Kurils Alaska, Rocky Mtns.

All distribution data in the above tables are from Eskov (1994). Note that although this second group of species are widespread in northern Asia, they extend only into the northern Kurils and have not been found in the south part of the archipelago. But it is the first group that are really interesting. Their presence in the northern Kurils is clearly the legacy of a former land bridge between northeast Asia and northwest North America, and shows that species from each of these areas were able to use this land bridge to invade the other; not just from Asia to America as popular scientific stereotypes imply. Finding more such species in a more detailed survey of the northernmost Kuril Islands can be confidently expected.

Another matter of interest is that these species, which have invaded Asia from North America and therefore certainly did not evolve in situ, nevertheless are found on islands which are volcanic and never had any connection with the Asian mainland, let alone any past land bridge to North America. Here is another convincing demonstration of the tremendous effectiveness of spider aerial dispersal by means of ballooning (see Crawford et al. 1995).

Ecological Relationships

The detailed habitat data taken with IKIP specimens will, in due course, allow us to correlate habitat requirements with distribution. It is already clear that, with some species whose dispersal ability is great enough to allow them to disperse between widely separated islands in the Kuril archipelago, the determining factor in which islands they occur on is habitat requirements and not zoogeographic history. Here are some examples from spiders collected by IKIP-1996:

Taxon

KU

IT

S-CENT

MA/RK

SA

EK/CR

KH

ON

PA

Cybaeopsis typica X X X X
Ummeliata angulitubera X X X X
Ummeliata osakaensis X X X
Theridion nigrolimbatum X X X
Howaia brevipes X X X
Gonatium nipponicum X X X
Neriene emphana X X
Cybaeus basarukini X X X
Yaginumaella ususudi X X X
Sabacon makinoi X X X
Mizozatus flavidus X X X X X X X
Haplodrassus signifer X X X X X
Robertus lividus X X X X X X X
Robertus ungulatus X X
Centromerus sylvaticus X X X X X X X X
C. terrigenus X X X
Oreonetides vaginatus X X
O. shimizui X X X

At the top of the above table are ten species found only at or south of Matua. All are found in and only in forest habitats, which do not exist to the north of that island. Several have not been found north of Urup, but might actually occur as far north as Matua in low numbers; after all, we only spent two days on Matua. The two species in this group recorded from Matua were extremely common there, though not found at all on unforested Raikoke, only 20 km farther north.

The next two species have been found only in UN-forested habitats, which are far more frequent in the north and central Kurils than in the the southern.

The table ends with three pairs of congeneric species, each of which has a member found from the northern end of the archipelago to a definite southern point, and one found from the southern end of the archipelago to a definite northern point. The two Robertus species overlap on southern Urup; the two Centromerus species overlap on southern Urup and northern Iturup, which have a similar type of stunted forest. The two Oreonetides species are not yet known to overlap, but might do so, as our knowledge of the Kuril distribution of O. vaginatus is obviously incomplete. Since sympatry does happen in some of these species pairs, the distributions are presumably determined by environment and not by competitive exclusion. In each case, the northern species of the pair is widely distributed in the Holarctic or Palaearctic regions, while the southern species is of more limited distribution and shows a faunal connection with Japan. More data will provide us with the basis for a very interesting study of these questions.

REFERENCES CITED

Bösenberg, W., and E. Strand. 1906. Japanische Spinnen. Abhandlungen, Senckenbergischen Naturforschenden Gesellschaft 30(1/2): 93-442, pls. 3-16.

Chikuni, Yasunosuke. 1989a. Pictorial encyclopedia of spiders in Japan. Tokyo: Kaisei-Sha Publishing Company, 306 pp.

_____. 1989b. Some interesting Japanese spiders of the families Amaurobiidae, Araneidae and Salticidae. in: Yoshiaki Nishikawa and Hirotsugu Ono, eds. Arachnological Papers Presented to Takeo Yaginuma on the Occasion of His Retirement. Osaka: Osaka Arachnologists' Group. Pp. 133-152.

Crawford, R.L. 1992. Catalogue of the genera and type species of the harvestman superfamily Phalangioidea (Arachnida). Burke Museum Contributions in Anthropology and Natural History 8: 1-60.

_____. In press. On Some Spider Species Described by Saburo Saito. Acta Arachnologica, to be published in 1997.

Crawford, R.L., J.S. Edwards, and P.M. Sugg. 1995. Spider arrival and primary establishment on terrain depopulated by volcanic eruption at Mt. St. Helens, Washington. American Midland Naturalist 133: 60-75.

Eskov, K. Yu. 1994. Catalogue of the linyphiid spiders of northern Asia (Arachnida, Araneae, Linyphiidae). Moscow: Pensoft Publishers, 142 pp.

Eskov, K. Yu., and Yu. M. Marusik 1993. New data on the taxonomy and faunistics of North Asian linyphiid spiders (Aranei Linyphiidae). Arthropoda Selecta 2(4): 41-79.

Marusik, Yu.M., K.Yu. Eskov, D.V. Logunov, and A.M. Basarukin. 1992. A check-list of spiders (Arachnida Aranei) from Sakhalin and Kurile Islands. Arthropoda Selecta 1(4): 73-85.

Miller, Jeremy A. 1996. Systematics of the erigonine spider genus Sisicottus (Araneae, Linyphiidae). M.S. Thesis, Western Carolina University, Cullowhee, North Carolina, 129 pp.

Nakatsudi, K. 1937. Notes on a new genus and two new species of Arachnida from the island of Paramushir, Northern Kuriles, Japan. Journal of Agricultural Science (Tokyo) 1(1): 22-27, pl. 1.

Saito, S. 1932. Descriptions of two new species of Araneida from the Northern Kurile Islands. Transactions of the Sapporo Natural History Society 12(2-3): 100-104.

Saito, S. 1933. Araneida from the Northern Kurile Islands, with descriptions of two new species. Bulletin of the Biogeographical Society of Japan 4(2): 122-132.

Saito, S. 1935. Spiders from the Northern Kurile Islands II. Transactions of the Sapporo Natural History Society 14(1): 55-56.

APPENDIX

Spider Species Previously Recorded from the Northern Kuril Islands

Total 47 spp. recorded by Marusik et al. (1992) plus 6 additional species whose records from the 1930s are possibly or probably correct.

PAR = Paramushir, SHU = Shumshu, ALA = Atlasova (Alaid)

Taxon

IKIP-96

(by Marusik et al., 1992)

(Japanese authors, 1932-37)

Tetragnathidae

Tetragnatha extensa PAR(?) (Saito 1935)

Araneidae

Araniella sp. PAR
Larinioides cornutus ? PAR

Theridiosomatidae

Theridiosoma epeiroides X PAR, ALA (Saito '33), SHU (Saito '35)

Theridiidae

Enoplognatha tecta X PAR, SHU
Robertus lividus X PAR,SHU
"Theridion araitense" ALA (Saito 1932), nom. dub.

Linyphiidae

Allomengea scopigera SHU
Baryphyma kulczynskii X PAR
Bathyphantes pogonias X PAR, SHU
Bathyphantes setiger X SHU
Bolyphantes alticeps SHU
Centromerus sylvaticus X SHU
Dactylopisthes video X SHU
Diplocentria bidentata X PAR, SHU
Diplocephalus subrostratus SHU
Dismodicus alticeps PAR, SHU
Erigone arctica sibirica PAR, SHU
Erigone atra X PAR
Erigone simillima X PAR, SHU
Gnathonarium dentatum PAR
Gnathonarium suppositum X PAR, SHU
Helophora insignis SHU
Hilaira herniosa X SHU
Hilaira leviceps X SHU
Hypomma affine X PAR
Hypselistes semiflavus SHU
Kaestneria pullata X SHU
Lepthyphantes dybowskii SHU
Lepthyphantes nigriventris X PAR
Leptorhoptrum robustum X PAR
Maro sibiricus ? SHU
Maso sundevallii SHU
Mecynargus paetulus SHU
Meioneta similis PAR
Micrargus herbigradus X PAR
Oreonetides vaginatus X PAR
Phlattothrata parva SHU
Stemonyphantes sibiricus SHU
Tiso aestivus X PAR, SHU
Tmeticus japonicus X PAR
Tunagyna debilis X PAR, SHU
Walckenaeria cuspidata X SHU
Walckenaeria lepida X SHU

Lycosidae

Pardosa palustris X PAR, SHU ALA, PAR (Saito 1932)
Pardosa paramushirensis X PAR (Nakatsudi 1937)
Pardosa riparia X ALA (Saito 1933, '35)
Trochosa terricola X PAR, SHU ALA (Saito 1935)

Clubionidae

Clubiona kurilensis ALA (Saito 1933)
Clubiona mayumiae X SHU
Clubiona riparia X PAR, SHU

Gnaphosidae

Haplodrassus signifer X PAR

Thomisidae

Ozyptila trux X SHU

Unplaced records by older authors:

Family Tetragnathidae

  • Metellina segmentata Saito 1935 European species not known from Asia

Family Theridiidae

  • Achaearanea formosa Saito 1932, 3 European, poss. S-Kurils, unlikely in north

Family Lycosidae

  • Arctosa cinerea Saito 1932, 3 Possible in S-Kurils, unlikely in north
  • Pirata montanus Saito 1932, 3 Nearctic species not found in Asia
  • Schizocosa avida Saito 1932, 3 Nearctic species not found in Asia

Family Clubionidae

  • Clubiona brevipes Saito 1932, 3 European species not known from Asia
  • Clubiona fruteorum Saito 1932, 3 European species not known from Asia

Family Thomisidae

  • Xysticus limbatus Saito 1932, 3 = X. emertoni, Nearctic species not found in Asia


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