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Disjunctive Distribution of Vaccinium vitis-idaea and Thermal Condition* Woo-seok, Kong** Jong-Hwan, Lim*** 350m 1,200m 6.6 Abstract Vaccinium vitis-idaea or lingonberry, a typical arctic-alpine plant, is common on the circumpolar regions and alpine belts of the Northern Hemisphere, and also occurs on the alpine and subalpine belts of the Korean Peninsula, including Mt. Sorak and Hongchon. Vaccinium vitis-idaea at the elevation of c. 350m a.s.l. of Hongchon is found on the wind hole area with cool summer, and mild winter. Vaccinium vitis-idaea at Hongchon is regarded as the glacial relict of the Pleistocene period, and shows a disjunctive distribution, along with the alpine and subalpine belts of the northern and central Korea since the Holocene period. Present vertical range of Vaccinium vitis-idaea between Mt. Sorak and Hongchon might indicates that the temperatures during the glacial epoch was colder than today, down to -6 to -7 C Vaccinium vitis-idaea at this fragile wind hole site could be endanger if current global warming trends continues, and anthropogenic activities become serious. : Vaccinium vitis-idaea, arctic-alpine plant, Mt. Sorak, Hongchon, disjunctive distribution, wind hole, glacial relict, global warming 2007 (This work was supported by the research grant of the Kyunghee University in 2007). (Professor, Department of Geography and Institute of Basic Sciences, Kyunghee University), wskong@khu.ac.kr (Senior Researcher, Department of Forest Environment, Korea Forest Research Institute), limjh@forest.go.kr 495

4 500 relict species endemic species 4 Pleistocene refugia Vaccinium vitisidaea L. 350m 2005 http://www. ecojournal.co.kr wind hole natural history 200m 2 1 10 15cm 2 1 2m 3 3 SP-2000 Veriteq Co., Canada 2005 8 21 2007 12 20 30 2006 8 5 2007 5 11 3 US FS 2008a http://plants. usda.gov USDA ARS 2008 ZipcodeZoo 2008 Vaccinium vitis-idaea L. Kingdom Plantae Subkingdom Tracheobionta Superdivision Spermatophyta Division Magnoliophyta Class Magnoliopsida Dilleniidae Subclass Dilleniidae Order Ericales Family Ericaceae Vaccinioideae Subfamily Vaccinioideae Genus Vaccinium Vitis-Idaea Section Vitis-Idaea Vaccinium vitis-idaea L. 496

Dilleniidae 1 Vaccinium 858 450 51 92 Ericaceae http://ko.pandapedia.com/wiki) lingonberry, cowberry, foxberry, mountain cranberry, red whortleberry, lowbush cranberry, partridgeberry, redberry Oxycoccus Vaccinium subgenera 20 sections 2 Vitisidaea Vaccinium uliginosum Vaccinium http://www.nature. go.kr 13 4 3 1963 Vaccinium Oxycoccus Vaccinium Oxycoccus, Oxycoccoides Batodendron, Brachyceratium, Bracteata, Ciliata, Cinctosandra, Conchophyllum, Cyanococcus, Eococcus, Epigynium, Galeopetalum, Hemimyrtillus, Myrtillus, Neurodesia, Oarianthe, Oreades, Pachyanthum, Polycodium, Pyxothamnus, Vaccinium, Vitis-idaea Vaccinium bracteatum Vaccinium hirtum var. koreanum Vaccinium japonicum Vaccinium japonicum var. ciliaris Vaccinium oldhamii Vaccinium oldhamii for. glaucinum Vaccinium oxycoccus Vaccinium oxycoccus subsp. microcarpus Vaccinium uliginosum Vaccinium uliginosum for. depressum Vaccinium uliginosum for. ellipticum Vaccinium uliginosum var. alpinum Vaccinium vitis-idaea 497

uliginosum var. alpinum Vaccinium vitis-idaea var. genuinum 8 Hugeria japonica Oxycoccus microparpus Oxycoccus quadripetalus 1979 1997 5 1996 Vaccinium microcarpum 10 2001 Vaccinium japonicum var. japonicum Vaccinium japonicum var. ciliare Vaccinium koreanum Vaccinium oldhamii var. oldhamii 8 Flora of Korea Editorial Committee 2007 7 2001 Vaccinium Vaccinium uliginosum var. alpinum Vaccinium koreanum 7 20cm 1 3cm 5 13mm 1 6 7 2 3 6 7mm 4 10 8 10mm 8 9 2002 2006 2008 http://www.encyber.com hermaphrodite self-fertile http://www.pfaf.org Davis et al 2003 7 10 10 Baskin et al 2000 14 17 Hill & Vander Kloet 2005 Eriksson & Froborg 1996 5 10 Hall & Shay 1981 BWCA 2008 US FS, 2008b Ritchie 1955 Nekola 1999 498

Henry 2003 Ritchie 1955 Kardell 1986 Rook 2002 Holloway 1981 1982 Kuchko 1988 2 7 8 2 4 0 4 9 Jeglum 1971 Hall & Shay 1981 Holloway 1981 US FS 2008b 5 6 2 Fernquist 1977 Launert 1981 PFAF 2008 C A B B1 B2 B3 phytochemical BWCA 2008 Stang et al, 1990 http://www.eln.co.kr Tufts 2002 1997 http://www.kna.go.kr circumpolar circumboreal 2 Hulten 1970 499

Ritchie 1955 Hultén 1958 1962 Ohwi 1965 Holloway 1981 USDA ARS 2008 ZipcodeZoo 2008 Hultén 1970 Vaccinium vitis-idaea subsp. vitis-idaea 10 25mm 2 000m Vaccinium vitis-idaea subsp. minus 7 20mm 2 000 2 400m Fernald 1970 Hultén 1970 subsp. vitis-idaea subsp. minus Kurilo 3 Pliocene 2 Tirmenstein 1991 900 3 200m efloras 2008 1 100m 2 700m 2 500m 1 200m Hultén 1970 2 000m Landolt 1996 1925 1933 100 2 540m 1942 2001 1988 1998 Kolbek and Kucera 1989 900 2 025m 800 2 000m http: //ipcp.edunet4u.net 1965 1 103m 1 214m 350m 1 550 1 650m 958 1 638m 1 686 500

1 746m 1 200 1 600m 1 886 2 186m 1 000 1 300m 1 700 2 000m 894m 1 900 2 200m 900 2 025m 1 400m 3 9 1 420 1 816m 1 500 1 800m 2005 1 420 1 816m 1 485 1 805m 1 077 1 573m 1 586 1 893m 1 600m 1980 1 600m 1984 1998 3 1942 1965 1989a b 1989a b c 1 708m 1 950m 1 915m 10 Pinus pumila Taxus caespitosa Juniperus chinensis var. sargentii Thuja koraiensis Cypripedium guttatum var. koreanum Arctous ruber 501

Silene oliganthella Anemone narcissiflora Pedicularis manshurica Bupleurum euphorbioides Anemone narcissiflora Rosa koreana Rosa marretii Trientalis europaea Lilium cernuum Iris uniflora var. carina 80 4 2002 1 708m Vaccinium koreana 1997 2002 1 705m 1 676m 1 578m 1 550 1 650m 1 103m 1 214m 350m talus wind hole 2003 2004 10 Lee & Kim 1994 502

Cystopteris fragalis Athyrium crenatum Polypodium virginianum Hackelia deflexa 2006 350m 70m 30m 25m 7m 200m 2 5 Sorbaria sorbifolia var. stellipila Betula schmidtii Prunus padus Rhdododendron mucronulatum Diplazium sibiricum Rosa suavis Woodsia subcordata Clematis fusca var. coreana Calamagrostis langsdorfii Carex vaginata var. petersii Dryopteris fragrans Polytrichum commune Iokawa & Ishizawa 2003 Nekola 1999 Iokawa & Ishizawa 2003 palaeorefugia 350m 1 550 1 650m 1 200m 0 55 100m adiabatic lapse rate 6 6 209 http://www.nature.go.kr 2005 11 28 http://www.kma.go.kr 1971 2000 10 1 17 5 4 4 1 291 3mm 71 4 6 27 8 7 29 7 8 30 1 1 11 6 2 8 9 12 8 2 1 19 7mm 2 23 8mm 3 42 3mm 12 22 9mm 27mm 503

2005 9 2007 8 10 15cm 2 5 65 2006 1 1 12 31 1 2m 3 6 52 7 22 16 2 11 61 10 15cm 2 2 24 39 21 78 Kong & Watts 1993 19 8 23 7 30 5 27 2 2006 8 4 12 35 42 2006 2 4 9 30 26 58 1 1 1 2m 3 12 10 15cm 2 2 3 1 AWS 4 1 2m 3 1 1 1 3 28 1 2m 3 7 15 10 15cm 2 5 65 9 5 06 7 8 1 2m 3 1 8 18 3 2006 8 4 12 10 15cm 35 42 1 2m 24 93 1 2m 28 93 2005 2006 2007 9 10 11 12 1 2 3 4 5 6 7 8 1 5 06 4 04 2 06 14 28 15 45 14 11 8 86 2 62 1 08 1 81 3 25 2 78 3 28 2 16 43 8 54 0 80 11 54 11 51 11 61 2 02 6 93 13 36 19 01 22 16 17 28 5 65 3 16 05 8 82 2 07 10 10 6 66 5 01 0 62 7 06 13 65 18 20 20 00 21 11 7 15 17 46 9 69 2 82 9 64 5 50 4 89 0 88 7 67 15 03 19 48 21 14 23 74 8 16 1 2 10 15cm 3 1 2m AWS 504

2008 8 8 12 1 2m 34 58 10 15cm 42 81 1998 2002 2005 2006 Raatikainen & Vanninen 1988 PECC 2008 30mm 15 1 2m 10 4 1 1 550 1 650m 350m 350m 1 550 1 650m 1 200m 0 55 100m 505

6 6 9 2000 15 20cm 1 2m 1 10 15cm 2 5 65 24 39 21 78 Ritchie 1955 40 C Wikipedia 2008 Hall & Shay 1981 US FS 2008a Shevtsova et al 1997 2 5 22 Holloway 1981 in situ 506

core zone buffer zone 6 6 1 420m 1 816m 1 550 1 650m 1 1998 33 2 1 18 2000 35 2 77 87 2002 37 357 370 2005 507

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University of Minnesota. Holloway, P. S., Van Veldhuizen, R. M., Stushnoff, C., and Wildung, D. W., 1982, Vegetative growth and nutrient levels of lingonberries grown in four Alaskan substrates, Canadian. Journal of Plant Science., 62(4), 969~977. Hultén, E., 1958, The Amphi-Atlantic Plants and Their Phytogeographical Connections, Almqvist and Wiksell, Stockholm. Hultén, E., 1962, The Circumpolar Plants. I, Almqvist and Wiksell, Stockholm. Hultén, E., 1970, The Circumpolar Plants. II, Almqvist and Wiksell, Stockholm. Iokawa, Y. and Ishizawa, S., 2003, Vascular plants of wind-hole areas in Japan (1), J. Journal of Phytogeography and Taxon, 51, 13~26. Jeglum, J. K., 1971, Plant indicators of ph and water level in peatlands at Candle Lake, Saskatchewan, Canadian. Journal of Botany, 49, 1661~1676. Kardell, L., 1986, Occurrence and berry production of Rubus chamaemorus L., Vaccinium oxycoccus L., V. microcarpum Turcz. & V. vitis-idaea on Swedish peatlds, Scandinavian Journal of Forest Research, 1(1), 125~140. Kolbek, J. and Kucera, M., 1989, A Brief Survey of Selected Woody Species on North Korea (D.P.R.K), Botanical Institute, Czechoslovak Academy of Sciences, Czechoslovakia. Kong, W. S. and Watts, D., 1993, The Plant Geography of Korea, Kluwer Publ., The Netherlands. Kuchko, A. A., 1988, Bilberry and cowberry yields and the factors controlling them in the forests of Karelia, U.S.S.R., Acta Botanica, Fennica., 136, 23~25. Landolt, E., 1996, Vaccinium vitis-idaea subsp. minus, An overlooked circumpolar-arctic taxa of the Alps, Anales Jardine de Botanique, Madrid, 54, 277~284. Launert, E., 1981, Edible and Medicinal Plants, Hamlyn. Lee, K. S. and Kim, J. H., 1994, Primary succession on talus area at Mt. Kariwangsan, Korea, Korea Journal of Ecology, 21, 383~387. Nekola, J. C., 1999, Palaerefugia and neorefugia: the influence of colonization history on community pattern and process, Ecology, 80, 2459~2473. Ohwi, J., 1965, Flora of Japan, Washington, DC: Smithsonian Institution. Raatikainen, M. and Vanninen, I., 1988, The effects of the 1984~1985 cold winter on the bilberry and ligonberry yield in Finland, Acta Botanica Fennica, 136, 43~47. Ritchie, J. C., 1955, Vaccinium vitis-idaea, Journal of Ecology, 43(2), 701~708. Rook, E., 2002, Plants of the North, http://www.rook. org/earl/bwca/nature/flora.html. Shevtsova, A., Haukioja, E., and Ojala, A., 1997, Growth response of subarctic dwarf shrubs, Empetrum nigrum and Vaccinium vitis-idaea, to manipulated environmental conditions and species removal, Oikos, 78(3), 440~458. Stang, E. J., Weis, G. G., and Klueh, J., 1990, Lingonberry: Potential new fruit for the northern United States, pp. 321~323. In: J. Janick and J. E. Simon (eds.), Advances in New Crops, Timber Press, Portland, OR. Tirmenstein, D., 1991, Vaccinium vitis-idaea, In: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (2003, April). Fire Effects Information System. BWCA (Boundary Water Canoe Area), 2008, The natural history of the northwoods, http://www.rook.org /earl/bwca/nature/shrubs/vacciniumvit.html (accessed Aug. 2008). efloras. 2008, Flora of China, http://www.efloras.org/ florataxon.aspx?flora_id=2&taxon_id=200016732 (accessed Aug. 2008). PECC, 2008, Plant Ecophysiology and Climate Change Group, University of Helsinki, http://www. helsinki.fi/bioscience/pecc (accessed Aug. 2008). PFAF (Plants For A Future), 2008, http://www.pfaf.org (accessed Aug. 2008). 509

US FS (United States Forest Service), 2008a, Fire Effects Information System, http://www.fs.fed.us/ database/feis/plants/shrub/vacvit/all.html (accessed Aug. 2008). US FS(United States Forest Service), 2008b, Fire Effects Information System(FIES), http://www.fs.fed.us/ database/feis/plants/shrub/vacvit/all.html#dist RIBUTION%20AND%20OCCURRENCE (accessed Aug. 2008). USDA ARS(United States Department of Agriculture, Agricultural Research Service), 2008, Germplasm Resources Information Network(GRIN), http:// www.ars-grin.gov/cgi-bin/npgs/html/taxon. pl?41069 (accessed Aug. 2008). Wikipedia, 2008, http://en.wikipedia.org/wiki/ericaceae (accessed Aug. 2008). ZipcodeZoo, 2008, http://zipcodezoo.com/plants/v/ Vaccinium_vitis-idaea.asp (accessed Aug. 2008). http://ipcp.edunet4u.net/~baikdoo/adventure http://ko.pandapedia.com/wiki http://plants.usda.gov http://www.ecojournal.co.kr http://www.eln.co.kr http://www.encyber.com http://www.fs.fed.us/database/feis http://www.kma.go.kr http://www.kna.go.kr http://www.nature.go.kr 130-701 1 02-961-0548, wskong@khu.ac.kr Correspondence: Woo-seok, Kong, Department of Geography, Kyunghee University, Hoegi-dong, Dongdaemun-gu, 130-701, Seoul, Korea(Phone: 82-2- 961-0548, e-mail: wskong@khu.ac.kr) 510