HORTSCIENCE 43(5):1623 -1625. 2008. Ridge may have been connected with each other and with Hokkaido. Glaciers covered the northern and central Kurile Islands during Strawberry Speciespf Iturup the glacial maximum of the Pleistocene but probably did not extend any further south than and Sakháliui Islands central Iturup, beca,ise no trace of glaciations has been observed on Kunashir or in the Lesser Kim E. Hummer Kurile Islands (Kryvolutskaya, 1973). This USDA- -ARS, NCGR, 33447 Peoria Road, Corvallis, OR 97333-2521 suggests that parts of southern Iturup may have provided a refugia for ancient plant species. Andrey SabitovThe human population of lmrup is mainly VIR, Far East Experiment Station, Vavilov Str. 9 Vladivostok 690025, Russia comprised of military personnel and fisher- men. The island is difficult to access by boat or plane as a result of weather conditions and Additional index words. Era garia iinurnae, Fragaria nipponica, Fragaria yezoensis, Fragaria political constraints.. Only a few scientific it urupensis, genetic resources, germplasm expeditions have visited the island during the past decade. Abstract. A plant-collecting expedition to Iturup and Sakhalin Islands, Sakhalin The flora of this region is preboreal, and Territory, Russian Federation, occurred between 21 July and 12 Sept. 2003. Strawber-indigenous island genera have related spe- ries, Fragaria L., were observed and collected. Japanese and Russian flora describe two cies in Western Europe and North America Makino and the Yezoensis strawberry, F. diploid (2n = 2x = 14) species, F. iinumae (Charkevicz, 1996). Charkevic7 (1996) var.yezoensis (H. Hara) Kitam.I, as native nipponica Makino Isyn. = Fragaria nipponica described only two diploid Fragaria species = = 56) to those islands. In addition, a recent monograph described a new octoploid (2n native to the far eastern island region of Staudt, from a .Japanese herbarium specimen collected strawberry species, P. iturupensis Russia. F. iinumae and F. nipponica Both on Atsunupuri volcano near Lesozovodoskyl, Iturup, in 1929. The objectives of the 2003 of these species are native to Sakhalin and the collecting trip were to obtain wild strawberries from these islands and determine if a Kurile Islands. In the Flora qf.Japan, Ohwi F iinumae Makino population of F. ifurupensi.s still existed on the volcano. The native (1965) presents F. :inumae and separated was obtained from the Ogonki Village, Anivskyi Region, Southern Sakhalin Island, but F nipponica from F yezoernis with the key was not observed on Iturup. The Yezoensis strawberry was observed at Cape Otlivnoy, character difference of ascending versus iturupensis Staudt were collected Iturup. Two small colonies of the native octoploid F. spreading hairs. More recent Japanese floris- from mid-elevations on the east-facing slope of Atsunupuri volcano. The native tic treatments have submerged F. yezoensis was limited to this volcanic montane habitat and was not distribution of F. iturupensis within F. nipponica (Naruhashi and Iwata, found in the foggy, sandy locations of coastal elevations on Iturup. The leaf morphology 1988). This species association is under and growth habit of F. iturupensis plants were similar to that of F. virginiana subsp. F. additional botanical scrutiny (Staudt, per- glauca (S. Watson) Staudt of North America, but the fruit was different, more like sonal communication). Fragaria vesca L. is vesca L. Probable cultivated escapes were observed at two sites on the east coast of Iturup a European introduction into this region of F,. and at one site in southern Sakhalin. The 2003 expedition confirmed the existence (Ohwi,. 1965) and is not native to Sakhalin, it urupensis, the only known native Asian octoploid strawberry on Atsunupuri. Limited Hokkaido,. or the Kurile Islands. Staudt quantities of germplasm are available for research from the curator, U.S. Department of (1973 .1989) studied a Japanese herbarium Agriculture, Agricultural Research Service, National Clonal Germplasm Repository, specimen and seedlings of strawberries from Corvallis, OR, the U.S. national strawberry gene bank. Ih,,-,,n Island The. cnpcimen I---.nrnved to he octoploid(2n.= 8x = 56) and he named the Staudt (1973) determined that a plant southwest alignment. The Greater Kut•ile species F. iturupensis Staudt. This is the only originally collected by a Japanese botanist Islands extend northeastward from Hokkai do known native Asian octoploid species. from Atsunupuri volcano on Iturup Island to the Kamchatka. Peninsula. These islar ids ., The objective of this project was to obtain was an octoploid species that he named F. separate the Sea of Okhotsk from the Paci fie strawberry samples from Iturup and Sakhalin iturupensis. The distribution of this species Ocean. Islands and to determine if the strawberries was reported to be only from this locality. Sakhalin is along, narrow island ezlOOtDx growing, on . Atsunupuri volcano could he Examples of this species were unavailable 200 km with , a, north—south longitudi:nal verified as the octoploid F. iturupensis. in the 1990s and early 2000s for further alignment situated at its closest point research. Conjecture by American, Japanese, km east of the Asian ,continent. The forn Materials and Methods and Russian botanists called for confirmation tion of the Kurile Archipelago began in of the existence of this, the, only Asian late Cretaceous (i90 million years before I From 27 July to II Aug. 2003, a plant- octoploid, and determination of how this present) when the Okhotsk Terrain of collecting expedition was made to Iturup and species fits into Fragaria phylogeny. Scien- Kula Plate collided with the Siberian cot Sakhalin Islands, Russian Federation. This tists also wondered if additional distributions nent (Kimura and Tamaki, 1985) creatin; trip occurred through scientific collaboration of this species could be found on Sakhalin subduction zone along the southeast mar; between the U.S. Department of Agriculture or other nearby islands. of the Okhotsk Terrain. Iturup and or] and the N. I. Vavilov All-Union Research Iturup Island is the second island to the islands of the Greater Kurile Ridge began Institute, Russian Federation. On . 29 July northeast of Hokkaido and is the largest form during the late Oligocene and Mioce 2003, the expedition flew from Korsakovo, island of the Greater Kurile Archipelago. It and the volcanic activity that gave rise tot Sakhalin Island, to Burevestnik, Iturup is 100 km x 25 km with a northeast— Ridge is still in progress (McKenna, 198 Island. Because there are no roads, the islands The northern Kuriles are recently formedd were traversed on foot or with some assis- are developing their flora, receiving p1 tance from military vehicles. Three days of migrations from Kamchatka at their no hiking were needed to reach Atsunupuri Received for publication 31 Mar. 2008. Accepted and from Japan through the southern Kurt volcano from Burevestnik. During this expe- for publication 3 May 2008. (Miyabe, 1890). dition, more than 145 clonal and seed sam- We acknowledge funding for this project from ples of berries were ollected representing six USDA ARS CRIS 5358-21000-033-OOD and a Iturup has been above sea level since genera and 21 species. The collection local- USDA, ARS Plant Exploration Grant for 2003. early Pliocene. During the glacial sea-le To whom reprint requests should be addressed; regressions of the late Pleistocene, Itur ities and dates for Fragaria species that were e-mail Kim.Hummerars.usda.gov Kunashir, and the islands of the Lesser Ku: encountered are presented (Table 1). 1623 HORTSCIENCE VOL. 43(5) AuGUST 2008 Table I. Collection information for strawberries, Fragaria L., collected on Iturup and Sakhalin Islands, Russian Federation, in 2003. Collection date Collection no. Corvallis no. Species Ploidy Locality 27 July 36 CFRA 1837 F. xanunassa escape Octoploid Sakhalin Island, on the outskirts of Ujno-Sakhalinsk City, near the sanitarium called "Mountain air"; elevation 178 m;46°5762l N, 142°4627l" E 16 Aug. 37-OG 1842 F. iinumae Makino Diploid Sakhalin Island, Anivskii Region, 16 km from Ogonki Village by Ogonki-Gomozavodsk Road; elevation 123 m; 46°42492" N, 142°13-483" E 30 July 38-KH 1838 F. xananassa escape Octoploid Iturup Island, near the road from Shumi-Gorodok Village to the bridge over the Khvoynaya River; 44.8190o N, 147.5503°E 31 July 39-GK 1839 F. xananassa escape Octoploid Iturup Island, Eastern (Pacific) Coast, on the outskirts Goriachie kluchi Village; elevation 53 m: 45"01 934" N, 14745350 E 3 Aug. 40-OTC 1840 F. nipponica Makino Diploid Iturup Island, Eastern (Pacific) Coast on Cape Otlivnoy 15 km south of Shumi-Gorodok Village; 44.9276° N, 147.5766° E 5 Aug. 41-ATS 1843 F. iturupensis Staudt Octoploid Iturup Island, on eastern slope of Atsonupuri volcano; elevation 630 m; 45°06748" N, 147°58796" E 5 Aug. 42-ATS 1841 F. iturupensis Staudt Octoploid Iturup Island; on eastern slope of Atsonupuri volcano; elevation 650 m; 4506748" N, 14758796 E Ploidy is based on root tip chromosome counts and flow cytometry.

Strawberry runners and fruits were gath- grass and shrubs in association with Conval- from Shumi-Gorodok Village to the K.hvoy- ered from the wild in their native locations on lana keiskei Miq., Lilium dahuricum Ker- naya River, Iturup, and on the outskirts of Sakhalin and Iturup Islands. An open pop- Gawl, Lothvrus maritimus BigIl., Geranium Ujno-Sakhalinsk City near the Sanitarium ulation sample of as many fruit as were erianthum D.C., Sasa kunilensis Makino et called "Mountain Air", Sakhalin, we col- available from each strawberry colony was Shibata, Equisetum silvaticum L., Pinus lected three samples of larger fruited straw- collected. The seeds were extracted from the pumula Rgl., Sorhus sambucifolia Roem., berries, probably F. xananassa, from plants fruit at camp near the collection sites. Runner La,i.i curilensis Mayr., and Rosa rugosa having morphological characters that did not plants and seeds were brought to the Far East Thunb. match descriptions for native species. Fruit Experiment Station, Vladivostok, Russia. Fragania iturupensis. The plants grew on set was poor. Fruit were irregularly shaped Portions of the seeds of each seedlot were pumice scree of the eastern slope of Atsunu- and not fully mature. The maximum weight sent to the U.S. strawberry gene bank in puri volcano. The leaf and plant morphology was 4.4 g/berry. Corvallis, OR, where representative seed- of this species agreed with Staudt (1973) lings were grown. as similar to F. virginiana subsp. glaucu Discussion and Conclusions Chromosome counts were performed on (S. Watson) Staudt of North America. Fruit F. iturupensis samples in Corvallis on cells and plant specimens were obtained from Fragaria iinumae was found on southern from actively growing root tips. Samples two small populations at elevations of 630 Sakhalin Island and the Yezoensis strawberry were pretreated in 0.002 M 8-hydroxyquino- and 650 m (Fig. I). Only a few nonflowering was found on Iturup. These observations line, fixed in Farmers solution (absolute single plants were seen on the lower part of agree with distribution maps for these species ethanol and glacial acetic acid at 3:1 ratio), the stone slope. Plants occurred mostly on the in the Russian and Japanese flora (Charkevicz, rinsed, and stored in 70% ethanol. Samples border of the dwarfed forest edge. Associated 1996; Ohwi, 1965). were stained in acetocarmine (Smith, 1947) plants included Alnus maximowiczii Call., The Iturup strawberry, F. iturupensis, has and chromosomes were counted. Flow Salix fumosa Turcz., Artemesia borealis limited distribution on the Atsunupuri vol- cytometry was also performed on mature Pall., A. anctica Less., and Carex micropoda cano but is octoploid. Further analyses of summer leaves of each of the collected Asian C.A. Mey. the genomes of these Asian species are in species in Aug. 2007 (Plant Cytometry Serv- The fruit dimensions were Pz5l.5 x 1.6 cm progress. The origin of the North American ices, Schijndel, The Netherlands) as another with a maximum weight of 1.5 g/berry. octoploid species remains unknown, but estimate of ploidy. Chromosome counts from cells of actively Staudt (1999) postulated that "a hypothetical dividing seedling root tips indicated that the octoploid species probably arose in East Asia plants were octoploid (2n = 2x = 56), which and may have migrated via an Alaskan- Results agrees with Staudt et al. (2008). Siberian land bridge to North America". Escaped octoploid species. At Goryachie With seedlings provided from this explo- Strawberry specimens were collected Kluchi Village and on the left side of the road ration, Staudt et al. (2008) contrasted pollen from five locations on Iturup Island, three near sea level and two growing in higher elevations on Atsunupuri volcano. Two sam- ;c. B ples of strawberry were collected on Sakhalin pjra Island (Table 1). Fragaria iinumae. On Sakhalin, these I. plants were collected from the edge of a •••••, forest road, 7 km southwest of Ogonki Village. Salix spp. and Sasa kurilesis Makino et Shibata were growing nearby. The straw- Ne •0 berries were semiprostrate, perennial herbs 7 to 15 cm tall. Fragaria nipponica (Yezoensis Straw- :7 berry). Fruit from a large colony of F. nipponica was collected on Cape Otlivnoy. This colony was growing in coastal sand at Fig. 1. Fragaria irurupensis Staudt plant (A) and fruit (B) from Atsunupuri volcano, Iturup Island, Russian m elevation. The plants grew in dense Federation.

1624 HoRESCIENUL Voi. 43(5) AUGUST 2008 grain morphology, aroma, and flavor com- Agriculture Corvallis gene bank. Future Naruhashi, N. and T. Iwata. 1988. Taxonomic ponents in the Iturup strawberry fruit with exploration on Kunishir and other nearby reevaluation of Fragaria nipponica Makino those of North American octoploid species islands is planned. and allied species. J. Phytogeography and Taxonomy 36:59-64. F. virginiana and F. chiloensis. The pheno- Ohwi, J. 1965. Flora of Japan [in English]. logical relationship of the hermaphroditic Literature Cited Smithsonian Inst., Washington, DC. p. 523- Iturup strawberry seems further differenti- Charkevicz, S.S. 1996. Fragaria L. Plantae Vas- 524. ated than that of these heteroecious octoploid culares Onentis extremi Sovieiici. Tomus. Vol. Smith, L. 1947. The acetocarmine smear tech- species than originally postulated (Staudt 8. p. 163-165 [in Russian]. nique. Stain. Tech 22:17-26. et al.. 2008). The quest for an ancestral link Kimura, G. and K. Tamaki. 1985. Tectonic frame- Staudt, G. 1973, 1973. Fragaria iturupensis, eine between lower ploidy Asian strawberries work of the Kuril Arc since its initiation, P. neue Erdbeerart aus Ostasien [Fragaria itur- 641-676. In: Nasu, N., K. Kobayashi, S. upensis, a new strawberry species from East and North American octoploids continues. Uyeda, I. Kushiro. and H. Kagami (eds.). Asia]. Willdenowia 7:101-104. 2003 plant-collecting expedition con- Our Formation of active ocean margins. Terra Staudt, G. 1989. The species of Fragaria, their finned the existence of F. iturupensis, the Scientific Publishing Co., Tokyo, Japan. taxonomy and geographical distribution. Acta only known native Asian octoploid straw- Kryvolutskaya, G.O. 1973. Entomofauna of the Hort. 265:23-33. berry, on the east side of Atsunupuri volcano, Kurile Islands: Principal features and origin. Staudt, G. 1999. Systematics and geographical Iturup Island: F. iinumae on Sakhalin and the Izdatelstvo Nauka, Leningrad, Russia. distribution of the american strawberry species. Yezoensis strawberry on Iturup. In addition, McKenna, M.C. 1983. Holarctic landmass rear- University of California Press—Berkeley. Bot- rangement, cosmic events, and cenozoic ter- any. 81:139151. escaped octoploid strawberries were ob- restrial organisms. Ann. Mo. Bet. Gard. Staudt, G., S. Schneider, P. Scheewe, D. Ulrich, served on both islands. Limited quantities 70:459-489. and K. Olbricht. 2008. Fragaria iturupensi.s, a of seeds or plant germplasm of these straw- Miyabe, K. 1890. The flora of the Kurile Islands. new source for strawberry improvement? Proc. berries are available for research by request Memoirs of the Boston Society of Natural of the VI International Strawberry Symposium, to the author at the U.S. Department of History. Vol. IV, No. VII. p. 203-275. Huelva, Spain. p. 49. Abst. 887.

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