Eco-Geographical Relations of the Bj~Rn~Yavascular Flora, Svalbard

Eco-geographical relations of the Bj~rn~yavascular flora, Svalbard

TORSTEIN ENGELSKJ0N

Engelskjen, T. 1986: Eco-geographical relations of the Bjerneya vascular flora, Svalbard. Polar Research 5n.~19- 127. A renewed survey of the flora of Bjemeya (74" 31") has yielded a total of 54 native vascular species. fi Details of their distribution are provided by local maps and vertical frequency tabulations. Soil preferences of species and various plant communities are outlined. The general climate and microthermal conditions of the prevailing vegetation place Bjemeya as a pronouncedly maritime part of the Svalbard archipelago, supporting mainly hygrophilous and chionophilous plant communitites and species. Only half of the island's vascular species persist at elevations above 200 m ad.. corresponding to a heat sum of July and August below approximately 190 Degree-Days. Ice-cored peat mounds coated by Rharomitrium lanuginosum were discovered on plateaux above 400 m elevation. The impact of the Vistula glaciation is discussed in view of Quaternary evidence from the Barents Sea area. Probably only one third of the present vascular flora could endure full-glacial conditions in possible ice-free cliffs on the coast of Bjerneya. Torstein Engelskjon. Botanical Garden and Museum, Universitjl of Oslo, Trondheimsveien 23b. 0560 Oslo 5. Norway. December I985 (revised September 1986).

1. Introduction genera, such as Draba, Dupontia, Festuca, Poten- Since the floristic survey published by Engel- tilla. and Puccinellia s. lat. I generally adopt pre- skjan & Schweitzer (1970) of the vascular flora liminary results from an ongoing cytotaxonomi- of Bjamaya (Bear Island), Svalbard, little cal study of vascular plant species from Bjamoya botanical work was done until 1983, when the and Spitsbergen (Engelskjan & Schweitzer 1970; present author joined an expedition of Tromsa Engelskjan 1979, and unpublished results). Museum to Bjamaya, 9 to 28 August. The nomenclature of bryophyte species fol- New observations of the flora and vegetation lows hell(1956) and Nyholm (1954-69), and during this stay make up the foundation of the of lichens, Santesson (1984) and Walker (1985). present treatment. An outline of the plant communities of Bjamaya is provided, and the thermal and edaphic conditions are related to the deve- 2. The climate of Bjornoya lopment of vegetation and plant distributions The meteorological station is situated on the N within the island. Notes on altitudinal belts are coast, at 74"31'N, 19"E, 15 m above sea level. included, as well as notes on the discovery of The extensive northern plain of the island undu- Rhacomitnum lanuginosum peat mounds at app- lates between 15 and 50 m a.s.1. This relief causes roximately 400 m above sea level on the southern only insignificant deviations from the prevailing mountain plateaux of the island. maritime climate observed at the station. Moun- Distribution maps are given for the 54 native tains to the SE and S reach altitudes from 410 to vascular species. 535 m ad., and temperatures there are lower Brattbakk (1984) visited the island to study the owing to the vertical temperature gradient and potential of the vegetation for reindeer grazing. orographic cloud formation (Steffensen 1969, He contributed an interesting floristic novelty, 1982). The thermal climate of Bjarnaya is shown Saxifaga foliolosa R. Br. in Table 1 and compared with other North At- The taxonomy and nomenclature of Svalbard lantic, Arctic stations. Due to inconsistency of vascular plants are still ambiguous for several the observation material, two sets of mean 80 monthly temperatures are presented. One is for the period 1946-65 and is continuous, with a few breaks of homogeneity due to moving of stations (Steffensen 1969). The other is the Stan- dard normal series (193 1-60) with interpola- tions of breaks during World War I I. Differences between these series reflect a secular decline of temperatures at the end of the 1950s, which lasted up to the late 1960s, e.g. on the island of Jan Mayen (Steffensen 1969, pp. 266-267). Complex phenomena such as secular climatic change could not be dealt with here, although an analysis would be of interest in relation to distribution and performance of North Atlantic, Arc- tic vegetation. Some derived parameters of ecological significance are listed in Table 2 for the same Arctic stations. Heat sums were calculated as Degree- Days, June-September (DD-4) and July- August (DD-2). all based on the 1946-65 period. Cloudiness and mean annual precipitation are summarized for the period up to 1965, annual sunshine hours are given when available, and wind chill and cooling power calculated according to the formulae of Siple & Passel (1945) and Vinje (1962). A special feature of Bjamaya and other Arctic islands is the foggy summer. The number of days with fog occurring at 7, 13, or 19 C.E.T. is given in Table 3A for the months June-September. We see that Hopen is most frequented by Arctic seafog. A closer study of the original tables (Stef- fensen 1969, Table 11) reveals that Bjamaya tends to have foggy and overcast days concentrated to July and August, which is also reflected in the low number of annual sunshine hours. The figures from Isfjord Radio on the W coast of Spitsbergen indicate a more favourable radiation climate. Bjamaya compared with North Norway (Table 3B) shows a much higher fog incidence than Vardo, which is also exposed to drifting fog from the Barents Sea. The thermal regimen of Bjamaya is comparable to that of Jan Mayen, but is less oceanic with regard to annual amplitude of temperature and total precipitation. Hopen is cooler than Bjarnoya throughout the year, whereas Isfjord Radio experiences slightly higher July tempera- 81

Table 2. Some basic and derived climatological parameters from Bjamoya and some other Norwegian Arctic stations. Data compiled from Steffensen (1969) and Andresen (1979). Comparable, 1946- 65 20 year means. Wind chill/cooling power computed according to Siple & Passel (1945) and Vinje (1962).

Station Precipi- Degree-Days Sunshine Clear Overcast Wind chilVcooling power tation June- July- hours, days, days, June July August mm Sept. August annual June- June- DD-4 DD-2 Sept. Sept. Bjornaya 357 405 265 613* I .7 93.3 898132 822129 835/30 Jan Mayen 698 475 3 10 1) 1.1 86.4 856129 7801'27 758126 Hopen 368 145 130 2) 3.6 88.4 875/28 822/26 807/26 Isfiord Radio 378 355 270 1066** 4 73 876/30 804/28 823/29 * 1960-65 I) No record; mean cloud cover 6.7 octas ** 1956-65 2) No record; meancloud cover6 octas

Table 3. Fog conditions on Bjamaya and some other exposure to this factor than the other Arctic sta- Norwegian Arctic and hemi-Arctic stations. 1946-65 tions considered during the summer months. means. However, Semlja Frantsa Josifa, at nearly 82"N, A) Cases of fog as number of days through June- has an even less favourable thermal climate September with fog at specified hours. Data from Stef- (Miiller 1982), as have Kong Karls Land and fensen (1969). Sjuoyane in the far north and east in the most sea Station At 07 At 13 At 19 ice frequented parts of Svalbard (Norsk Polar- institutt, unpublished observations). Bjamaya 19.3 14.6 17.9 Jan Mayen 16.8 12.7 12.2 The annual precipitation on Bjornoya is below Hopen 19.6 17.1 20.0 400 mm like in most Arctic regions. High humidi- Isfjord Radio 7.1 4.9 6.6 ty of the ground and vegetation cover is main- tained by the low temperatures and heavy cloud B) Monthly fog frequency: Percentage of days with cover, and fog precipitation is also considerable. cases of fog at Bjamaya and at Varda, East Finnmark, Accordingly, Bjornaya has a maritime Arctic North Norway. climate with low temperatures and humid con- Station June July August September ditions. At comparable latitudes in the North Atlantic this climatic oceanity is only surpassed 14 24 22 II Bjamaya by Jan Mayen. Varda 4 5 7 I

3. The vertical temperature gradient tures than Bjarnoya. Other stations in Spits- Data from radiosonde ascents between 10 and bergen are not compared in detail here, but inner 22 August 1983 were utilized to estimate the air Isfjord (Longyearbyen) has higher summer ave- temperatures at higher levels of Bjornoya. The rages and maxima than Isfjord Radio, and less upper air soundings start from the meteorologi- fog. cal station and are performed regularly at 1200 The heat sums for June-September and and 2400 hrs. G.M.T. The air temperature July-August (Table 2) place Bjamaya in a posi- decreases linearly with altitude, but stratification tion between Jan Mayen and Isfjord Radio, and inversions occur at various levels. The first whereas Hopen is much cooler than most other of these so-called significant levels is usually stations. crossed between 950 and 850 mb air pressure, The indices of wind chill and cooling power which corresponds to 500-1300 m above sea entered in the same table reflect the combined level. The temperatures t, and t2 at levels HI and cold and wind stress on animals and plants. We H2 are inherent in the simplified barometric for- note that Bjornaya has a more pronounced mula: 82

P: - P, ti + I? a reference site on the N coast (Fig. I), but I H: = 1600 . (I + 0.004 -) visited most parts of the island during altogether P: + PI 2 six weeks in 1967 and 1983, recording the main plant communitites. Cryptogams were collected (Bliithgen & Weischet 1980) from representative sites, totalling 220 samples of mosses, 35 of liverworts, 150 of lichens, and a Here. P, and tl are air pressure and temperature few of terrestrial algae. Assistance with the deter- at the station, P2 and tz at the lowest significant mination of some samples was provided by spe- altitude level as explained above. The vertical cialists mentioned in the Acknowledgments. I temperature gradient is then given by could also draw on ecological notes by Summer- hayes & Elton (1923), Hanssen & Holmboe ti - t. (1925), Bertram & Lack (1938), Ronning (1959), A= (o('/lOOm) and Brattbakk (1984). Ecological comments were 0.01 HZ also given by Berggren (1875) and Lynge (1926) in works on the bryo- and lichenflora of Bjorn- For successive days, the average A was computed oya. as 0.66"; 100 m at 1200 hrs. G.M.T., varying The available map of Bjornoya (Norsk Polar- between 0.15" and 0.93'. These two extreme institutt 1944) does not display the microtopo- values occurred in connection with a cyclone graphy, and the only photocoverage consists of passage with humcane windspeeds. oblique pictures on file in the Norwegian Polar The same value, 0.66' /I00 m, varying between Research Institute. These photograms do not 0.47- and 0.88', was obtained from night ascents show the high ground because of orographic fog at 2400 hrs. G.M.T. during the flight. Vertical aerial photos of a large In view of the relatively uniform weather con- scale will be needed for quantitative vegetation ditions on Bjornoya during the summer months, mapping of the island. 0.65' 100 m is considered a realistic estimate for the vertical temperature gradient in the growing season. This value is also the WMO standard for Bedrock and soil temperature reductions. Because the island is The geological map (Fig. 2) is reproduced from unprotected from the general air circulation over Worsley & Edwards (1976) and based on the the Barents Sea, the local summit effect (Eide mapping by Horn & Orvin (1928). The bedrock 1945) is not expected to lower air temperatures varies considerably as to chemical composition. much. Table 6 shows soil chemical parameters of The temperatures at about 450 m on Misery- weathering material derived from three main fjellet and Hambergfjell were calculated from groups of sedimentary rocks: this vertical gradient (Table 4). It is noted that the

temperatures on the mountains of Bjornoya are I. Carbonatites and marls. - 1. Older and Youn- lower than at Hopen Radio, two degrees of lati- ger Dolomite (Hecla Hoek), 2. Tetradium lime- tude to the north, but still higher than in extreme stone, 3. Landnardingsvika and Kapp Kare for- Arctic environments. e.g. Semlja Frantsa Josifa mations ( ='Ambigua limestone and marl'), 4. or Severnaja Semlja (Tables in Muller 1982). Kapp Duner formation ('Fusulina limestone and dolomite', cf. Siedlecka (1975)), 5. Hamberg- 4. Outline of the vegetation on fjellet formation ('Cora limestone'), 6. Misery- Bjplrn~~ya fjellet formation ('Spirifer limestone').

A short description of the terrestrial vegetation II. Intermediate sediments with some calcite. - of Bjarnaya is provided to relate the local eco- I. Sandstone within the Miseryfjellet formation, logical conditions, and to compare with other 2. Kapp Hanna formation ('Yellow sandstone'), Arctic areas. 3. Triassic shales, partly with limestone concre- Phytosociological analyses were only made at tions. 83

Table 4. Thermal conditions (air temperatures and heat sums) on the high ground of Bjornoya, 450 m ad. 1946-65 means. Temperature reductions based on the meteorological station 15 m a.s.1. and gradient 0.65"C/ 100 m. DD-4: Degree-Days, June-September. DD-2: Degree-Days, July- August. Parameter Altitude 10-22 Monthly means Mean of m a.s.1. August, June July August September June- I983 September Mean temperature 15 4.3 I .8 4.2 4.3 3.0 3.3 450 I .5 - 1.0 I .4 1.5 0.2 0.5

Mean maximum 15 5.4 3.1 6.5 6.2 4.1 5.3 450 2.5 0.9 3.1 3.4 I .9 2.5

Mean minimum 15 3.2 0.2 2.5 2.1 I .5 I .7 450 0.3 - 2.6 - 0.3 - 0.1 - 1.4 - 1.1

Heat sums 15 - 55 130 135 90 (DD-4= 405, DD-2 = 265) 450 - .30 45 45 5 (DD-4~65, DD-2 = 90)

Table 5. Air and vegetation temperatures and some other meteorological parameters, reference site at Herwig- hamna, N coast of Bjerneya, 5-8 m a.s.l.10-22 August 1983.

Main weather Mean Air Vegetation category and aspect Cloud Mean situation temperature Humid plain Lichen slope Terrace cover octas windspeed "C (I),2"E (31),20"SE (34), IOOSE m s-I Clear, sunny Diurnal 4.2 5.5 9.5 7 4 4 days (n = 4) Maximum 5.4 9 21.5 16 Minimum 3.0 2 I 1.5

Partly overcast Diurnal 4.0 5.5 6 5.5 5 6 days (n = 5) Maximum 4.8 8 13.5 10.5 Minimum 3.1 2 1 I .5

Heavily overcast, Diurnal 4.7 5 5.5 5 8 7 rainy or foggy Maximum 6.2 I I 7.5 days(n=4) Minimum 3.5 4 3.5 3.5 ...... Entire period Diurnal 4.3 5.5 I 6 6 6 (n= 13) Maximum 5.4 8 14 II Minimum 3.2 2.5 2 2

III. Silicic sediments poor in calcite. - 1, Raed- The soils of rock group I1 are less well repre- vika and Nordkapp formations ('Ursa sandstone' sented and may be influenced by seaspray and or 'Kulm'). 2. Sandstones within the Kapp KAre organic deposition. A sandstone within the 'Spi- formation. rifer limestone', originally a redeposited desert The carbonatites of group I yield alkaline sand (Siedlecka 1975), weathers easily. Its lithosol with a high content of Ca (Table 6). This vegetation, however, is windworn and sparse, explains why most of the eutrophic species are although pH is 7.1 (Table 6). The Triassic shales concentrated in the lime-rich southern third of on the summits of Miseryfjellet have not been the island, as well as below Miseryfjellet, and on analysed. Their barrenness may be caused by the northern coast. mechanical instability. 84

Fig. I. The reference site for vegetational and microthermal studies near Grav- odden. north coast of Bjornoya. I. 35: Square numbers of first and last end of vegetation transect, cf. Fig. 3. 9 August 1983.

Lithosol of the Nordkapp formation (‘Ursa The maritime influence on soils sandstone’) is slightly acid and very poor in lime. The intercalated shaly layers cause the sandstone Table 6 also shows that some soils are high in to break up into coarse blockfields or stone poly- Na, which is presumably caused by sea-spray. gons which support scattered moss and lichen This is the case on the coastal brink which is turfs, the so-called ‘soil cadaver’ (Keilhau I83 1). exposed to breaking waves, and where local Another sandstone within the Kapp Kare forma- winds may carry seafoam up to 170 m ad.even tion (Table 6) produces an equally acid and in calm weather (Brattbakk 1984, p. 13). lime-poor lithosol, which is more fine-grained The seabird rookeries on Bjc~mayaare among and supports some vegetation. The studies of the largest in the northern hemisphere (Norder- Elvebakk (1982) from Spitsbergen show similar haug et al. 1977). Bird manure contributes to bedrocklvegetation interrelations. enrichment of N and P, and the seabirds also 85

Fig. 2. Geological sketch map of Bjarnoya. Reproduced from Worsley & Edwards (1976), by per- mission.

drop shells and carcasses. Especially the polar thawing down to approximately 0.75 m in the gulls and skuas affect the vegetation by tramp- summer. Below this level permafrost continues ling and moss plucking. The bird influence down to 60-70 m. Fleetwood et al. (1974) indi- around the large rookeries of Fuglefjell and cate unfrozen ground below lake bottoms, but Hambergfjell is traced more than one km away the permafrost causes waterlogging in the surface from the coast. The zone of ornithogenic influ- sediments. Extensive areas on the northern plain ence is readily seen from the distribution map of of Bj~rn~yaare studded with ponds and muddy Cochlearia groenfandica (Section 7, map 38). polygon fields. The central sandstone areas have a better drainage, but solifluction slopes develop on shaly layers i.a. in Sigjorddalen W of Misery- The influence ofpermafrost fjellet. Horn & Orvin (1928, p. 12) observed ground The limestone and dolomite hills in the Table 6. Weathering soils on Bjerneya. pH values and some elemental analyses.

Main bedrock derivation Texture and vegetation pH Ca-AL CaO Mg-AL K-AL P-AL Na (cf. text) mg/ 100 g o:, mg/100g mg/lWg mg/IWg mg/l00g Spirifer limestone Coarse gravel, ahumic, with 8.0 35s 2.10 32 4.5 0.5 26 in the Station area Sarifraga opposirijolia I Spirifer limestone S of Coarse gravel, humic, below 6.9 SX 0.58 IX 4.6 3.0 - Station area mesotrophic moss turf ...... Arenite within limestone. Fine, sandy detritus with windworn 7. I 5.0 0.14 II 6. I 5.5 - Kapp Posadowsky Sfereocaulonsp., on the brink

Arenite within limestone, Fine, sandy detritus below Schisrid- 6.7 - 13 I I Kapp Posadowsky ium maririmum cushions, on the brink Arenite, Kapp Hanna Fine, sandy detritus below Trenre- 7.4 __ formation pohliaalgal mat. on the brink ......

Ursa sandstone SW of Gravel and cryoturbated loam, 6.6 4.0 0.07 12 I .7 0.4 ~ Station area sparse vegetation I11 Ursa sandstone SW of Gravel and cryoturbated loam, 6.3 2.0 0.04 I? I .8 0.3 - Station area sparse vegetation

Arenite. outlet of Pelsa Reddish sand, with Luzula arcuafa 6.8 2.0 0.03 6.2 2.1 0.1 - and Silene acaulis

- : Not analysed. 87 southern part of the island are better drained and such as Koenigia islandica, Phippsia algida, and less influenced by cryoturbation. Saxifaga rivularis. Then 15 m of stony ground A particular development of permafrost was follow with few vasculars and several chiono- discovered on the plateaux of Hambergfjell and philous lichens. The species content changes Fuglefjell, i.e. palsa-like mounds cored by ice from square 27 onwards, as is seen from the and coated by Rhacomitrium lanuginosum peat coverage of Festuca rubra, Poapratensis ssp. alpi- (Section 6). gena, Oxyria digyna, Saxifraga caespitosa, and cushions of Cetraria islandica and Cladina mitis. The end of the transect at square 35 is a wind- Karst phenomena exposed and well-drained terrace with i.a. Draba Runoff is directed by the rivers Lakselva and norvegica, Rhacomitrium lanuginosum, and Engelskelva in the north, Russelva and Jordbru- Ochrolechiafrigida. elva in the south. Limestone ground causes sub- This transect shows three main developments terranean drainage in the vicinity of Lakselva, of the Bjmn~yavegetation : Jordbruelva, and upper Ymerdalen. Contrary to 1) Poorly drained moss carpet communities. the topographical map, part of the Ymerdalen 2) Well-drained and wind-protected lichen river is subterranean and disappears in a karst heaths with several vascular plant species. hole W of Antarcticfjellet, to well up in a gorge 3) Wind-abraded summit or terrace vegetation. 1.4 km farther down. Protected and well-drained This standard profile is well developed in the S dolines are seen in the limestone areas, and they slope of Kikutkollen close to the Bj~rn~yaRadio support special plant communitites (Table 7, station, 30-50 m a.s.1. (Fig. 4). C.2). A provisional classification of the vegetation of Bjomoya Wind abrasion Extensive areas of Bjarnaya support mainly Windwom vegetation is seen on the calcareous non-vascular communities, where grasses, herbs sandstone plains of the Kapp Hanna formation, and dwarf willows are insignificant. However, in the pass between Hambergfjellet and Alfred- some vascular species are even found in extreme fjellet, and on the splintered shales covering the cryptogamic communities. upper slopes of Miseryfjellet. Winter conditions A classification of vegetation may be founded have not been studied, but snow abrasion may on a combination of physiognomical, floristic, be an important ecological factor on exposed and ecological criteria. The ecological factors of crests, whereas the river gorges and dolines are bedrock, marine influence, frost action, hydro- more protected. logy, and wind exposure must be considered in particular. Main groups of communities can be arranged A vegetationalgradient on the N coast in this way, and some sociations (Dahl 1957, p. The composition of vegetation was studied in 60) can be defined by the criterium of dominan- relation to hydrology, exposure and micro- ce. A grouping into associations is desirable by climate at a reference site near Gravodden, W of syntaxonomic methods. The present provisional the Bj~mayameteorological station (Fig. 1). classification (Table 7) is based on ecological Vegetation was analysed along a 35 m long strip features and the occurrence of Dominant and running ESE-WNW from a mossy plain 5 m characteristic species (D), Characteristic, but not ad., crossing a blockfield with protected fruticose dominant species (C) and Frequent, but ubiqui- lichen, grass and herb communitites, and ending tous species (F). Floristic similarities and diffe- on a wind-exposed terrace 8 m a.s.1. rences are often blurred out in the Bj~mcaya The first 10 m of the transect (Fig. 3) are uni- vegetation because several ubiquitous species are form Drepanocladus uncinatus-Brachyth ecium dominant, e.g. Cochlearia groenlandica, Saxijiraga glaciale carpets with scattered vascular species caespitosa, and Cetraria delisei. 88

Table 7. A provisional outline of the terrestrial vegetation of Hjornoya. D: Dominant and characteristic species, C: Characteristic but not very numerous species, F: Frequent and ubiquitous species.

A. STRONGLY MARINE INFLUENCED COMMUNITIES 1. Shore communities I .I, ~ccinelliaphr?ganodescommunity D hccinellia phvganodes C Schistidium maritimum F Festuca rubra C Verrucariaceuthocarpa F Cochleariagroenlandicc C Verrucaria maura I .2. Sfellaria humifusacornmunity C Stellaria humifusa F Saxifroga rivularis C Carer subspathacea F Drepanocladus uncinatus

2. Bird rookery communities 2. I. Cochleariagroenlandica community D Cochlearia groenlandica C Prasiola crispa D Drepanocladus uncinatus C Xanthoria sorediata 2.2. Festuca rubracommunity D Festuca rubra C Poapratensisssp. alpigena F Cochlearia groenlandica C Puccinelliaphryganodes

B. MODERATELY MARINE INFLUENCED COMMUNITIES 1. Saxifraga rivulariscommunity D Saxifraga rivularis I) Drepanocladus uncinatus F Phippsia algida C Cepha1o:iella aff. arctica F Koenigia islandica D Lecidea vernalis F Oryria digyna C Leciophysma arctophilum F Cochlearia groenlandica F Pertusaria spp. F Sayifraga caespitosa F Psoroma hypnorum D Brachythecium glaciale 2. Chtysosplenium tetrandrum community C Chgaosplenium tetrandrum F Drepanocladus uncinatus F Oxyria digma C Drepanocladus vemicosus F Cardamine nymanii C Plagiomnium ellipticum C Saxifraga caespitosa C Philonotisfontuna F Saxifraga cemua C RliJBtidiadelphussquarrosus F Saxifiaga hirculus C Tortula ruralis C Brachythecium salebrosum C Marchantia polymorpha C Cinclidium arcticum C feltigera membranacea C CIimacium dendroides 3. Sedum rosea ssp. arcticum community D Sedum rosea F Draba noniegica F Cochlearia groenlandica

C. INLAND COMMUNITIES, WELL-DRAINED I. Chionophobous communities I. 1. Papaver dahlianum community (Bedrock group I) C Papaver dahlianum F Saxifraga caespitosa D Saxifraga oppositifolia D Lecidea ramulosa C Arabir alpina c' Fulgensia bracteata C Drabaalpina C Lpcanora epibnon F Cerastium arcticum C Caloplaca cerina 1.2. Silenearauliscommunity (Bedrock group 11) D Silene acau1i.c F Saxifraga caespitosa F Festuca vivipara F Saxifraga opposit[fdia F Cerastium arcticum F Cetraria delisei F Draba norvegica F Och rolechia,frigida (continued) 89

I .3. Luzula arcuatacommunity (Bedrock group 111) D Luzula arcuatu D Alectoria nigricans D Festuca vivipara D Sphaerophorus globosus F Salk herbacea F Cetraria delisei D Rhacomitrium lanuginosum

2. Intermediately exposed communities 2. I. Salixpolariscommunity (Bedrock group I) D Salixpolaris F Saxifraga cemua C Equisetum scirpoides F Drepanocladus uncinatus C Poaalpina C Orthothecium chtyseurn C Polvgonum viviparum C Timmia austriaca C Ranunculuspygmaeus C Tomenthypnum nitens C Salix reticulata C Toninia lobulata F Oxyria digyna 2.2. Salixherbaceacommunity (Bedrock groups II and 111) D Salix herbacea F Drepanocladus uncinatus C Cerastium cerastoides D CIadina mitis F Oxyria digyna D Cladonia ecmocyna F Saxifraga cemua D Cladonia stricta D Brachythecium reflexum C Peltigera kristinsonii D KirPria glacialis C Peltigera rufescens C Conostomum tetragonum C Stereocaulon alpinum 2.3. Taraxacum cymbifolium community (Bedrock group 11) D Taraxacum cymbifolium F Festuca rubra D Oxyria digyna F Salk herbacea C Sedum rosea ssp. arcticum F Saxifraga cemua

D. INLAND COMMUNITIES, SEASONALLY FLUSHED I. Phippsia concinna community (Bedrock group I) D Phippsia concinna F Sagina inrermedia C Equisetum variegatum F Cerastium regelii C Juncus biglurnk F Ranunculus sulphureus C Alopecurus alpinus C Bymciyophilum C Saxifraga hirculus D Scorpidium turgescens C Saxifraga tenuis F Lecidea ramulosa F Koenigia islandica 2. Phippsia algidacommunity (Bedrock group I I) D Phippsia algida F Ranunculus sulphureus F Koenigia islandica F Drepanocladus uncinatus F Sagina intermedia F Hygrohvpnum spp. F Cerastium regelii 3. Deschampsia alpina community (Bedrock group 111) D Deschampsia alpina F Ranunculus sulphureus D Festuca vivipara F Polytrichum alpinum F Equisetum arvense C Solorina crocea

E. INLAND COM MU NIT1ES, WATER- LOGG ED 1. Dupontia community (Eutrophic/mesotrophic) D Dupontia psilosantha F Calliergonstramineum C Calamagrostis neglecta F Drepanocladus intermedius C Cardamine nymanii F Drepanocladus badius C Hippuris vulgaris F Meesea triquetra C Ranunculus hyperboreus F Paludella squarrosa 2. Arctophila community (Mesotrophic/oligotrophic) D Arctophilafulva D Calliergonsarmentosum F Dupontia psilosantha F Drepanocladusfluitans F Ranunculus hyperboreus F Sphagnum lindbergii D Brachvthecium glaciale F Sphagnum riparium D Calhergon giganteum (continued) 90

F. VARIOUS NON-CLASSIFIED COMMUNITIES i. Nearl) non-vascular Rhacomitrium lanuginosum community, montane D Rhacomrfrium lanicginosum D Cerrana nivalis D Aulacomnium rurgidum F Cetrana islandica F Tortula norvegica F Sphaerophorus globosus F Tortrcla ruralis C Placopsi.sgelida(on stones) ii. Nearly non-vascular Dn.panoc1adu.q exannulatus community D Drepanocladus e.rannulatus C Sphagnum lindbergii D Drepanorladurfluirans C Sphagnum squarrosum D Calliergon sarmenfo~um C Leptoglossum lobarum C Koeriigiu i.dandica C Mirrula graci1i.s iii. Shale screes on summits of Miseryfjellet D Cochlearia groenlandica F Saxifraga cernua F Poaalprna F Saxifraga riwlaris F Cera.rtrum regelii D Rhacomitriumfasciculare iv. Sandstone blockfields and crags in the interior of Bjornoya D .4ndreaea rupe.w-iss.lat. C Parmelia omphalodes D Dicranowersia cri.spula C Parnielia sa.ratilis F Rhacomitrium lanuginosum D Umbilicariaarctica C G!,mnom itrium coralloides C Umhilicaria cylindrica C Cladonia bellidlflora C Umhilicaria proboscidea F C(adoniu coccifera C L'ntbilicaria torrefacla \, Limestone boulders and crags in the interior of Bjornoya C Sa.rijraga opposirifolia C Caloplaca cerina C €nca!vpa spp. C Lecanora epibnon C H,pntcm revolutum C Phvsciu musrigetia C Preudoldeella spp. C Plar~~ntliiumasperellum C Schisridium apocarpum vi, Late thawing or temporarily flooded bryophyte turf snow-beds. nearly non-vascular C Conoctnmum terragonum C Kiaeria glacialis F Drepanocladus uncinarus C Polurichumspp. C G~~mnomitriumspp.

Regional comparisons Wellington 1940; Lid 1964), especially with regard to the dominance of Brachythecium spp. and The vegetation of Bjornoya may be compared chionophilous vascular species. with that of other Arctic and Oroarctic regions adjacent to the Barents Sea. Spitsbergen is best investigated in this respect, and Brattbakk (1983, 5. Thermal conditions of the vegeta- 1984) recognizes the Sphaerophoro-Rhacomi- tion trietum lanuginosi as similarly developed on the Vegetation temperatures were monitored from western coast of Spitsbergen. There are also eco- 10 to 22 August 1983 by hourly IT 100 thermistor geographical relations to the Cladina rniris-zone readings charted on a Grant writer at the reference as defined by Eurola (1968) from coastal tracts site W of the Bjarnaya meteorological station of Spitsbergen. Limestone plains of Bjornoya (Fig. 1). For discussion of temperature measuring support communities with Saxifraga oppositfolia methodology, see Walton (1984). The instru- and Cerran'a deli.sei,which are similar to coastal ments worked without failure throughout the communities of Prins Karls Forland (Summer- period, and the registration covers various repre- hayes & Elton 1923) and Braggerhalvaya on the sentative weather situations (Table 5). western coast of Spitsbergen (Brattbakk I98 1 ). Diurnal means of superficial vegetation tem- The bryophyte carpet communities resemble peratures as well as average maxima and minima those described from Jan Mayen (Russell & are summarized from three representative plant 91

Cerastium arcticum Cochlearia qroenlandica Draba norvegica Koenigia islandica Oxyria digyna

Ranunculus hyperboreus I Saxifraga caespitosa .... Saxifraga rivularis Festuca rubra Phippsia alqida Poa pratensis ssp. alpiqena Puccinellia phryqanodes

Blindia acuta I I I Brachythecium glaciale ID Bryum nitidulum Drepanocladus uncinatus Polytrichum alpinum Rhacomitrium lanuginosum

Cephaloriella aff. arctica I Tritomaria quinquedentata - Cetraria delisei Cetraria islandica Cladina mitis Cladonia ecmocyna Cladonia uncialis Cladonia sp., squamulose Lecidea vernalis Leciophysma arctophilum Ochrolechia frigida Pertusaria sp. Psoroma hypnorum Stereocaulon alpinum

Galerina pumila Leptoglossum lobatum Mitrula gracilis

Prasiola crispa

Eroded cryptoqamic turf Stones 2 Square no. (1 m 1 I 2 3 4 5 6 7 8 9 10 11 12 13 11 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35

Fig. 3. Vegetation transect, Bjorn~ya:North coast, between Tobiesenelva and Gravodden, 5-8 m above sea level. Coverage according to Hult-Sernander’s scale (- : I - I: 5).

communities along the transect shown in Figs. 1 Cepbaloziella aff. arctica. and with Pbippsia and 3. algida and Saxifraga rivularis as the only vascu- Square no. I is on a humid, freely exposed lar species of importance. It belongs to the Saxi- plain with carpets of Drepanocladus uncinatus, fraga rivulariscommunity(Table 7,B. 1). Brachythecium glaciale, Btyum nitidulum and Square no. 3 1 is in a SE facing slope with well- 92 developed fruticose lichen cushions (Cetraria most effective in square no. 31, which is favour- islandica and Cladina mitis). Poa prarensis ssp. ably exposed and wind-protected. The vegeta- alpigena. and Oxyria digyna. It is related to the tion temperatures on Bjornaya were recorded as Salk herbacea community (Table 7, C.2.2), but generally low and not on average exceeding the no Salixoccurred in the immediate vicinity. ambient air temperature by more than 1-2". Square no. 34 is on an exposed sandstone ter- This is ascribed to the low direct sun radiation race with scattered moss and lichen turfs, border- (Table 2) and the generally humid conditions. ing on a wind-swept Sedum rosea community. On foggy summer days, vegetation temperatures The summary of microthermal conditions are levelled, irrespective of slope, aspect, and (Table 5) shows the low diurnal temperature drainage. Recalling the summer fog frequency amplitude of square no. I, irrespective of the (Table 34B), Bjnm~yais evidently not suitable weather situation. The average temperature for for vascular species having high temperature the entire period was 5S0,and average maxima demands for growth and reproduction. Lignified did not exceed 9'. species such as Salk herbacea, S. polaris and S. Square no. 31 had a mean superficial vegeta- reticulara mainly grow in sheltered and south- tion temperature of 7" and an average maximum facing places, and only in the low-lying areas (cf. of 14'. The highest temperature recorded during Section 6). a calm and sunny period bas 27.5". However, the low temperatures may be com- Square no. 34 had lower averages and maxima, pensated for by a long growing season, but this which may be caused by a less sunward aspect seems to promote mainly the cryptogamic vege- and exposure to wind chill. tation. In this respect Bjorn~yamay be compared The thermal conditions in these plant with Jan Mayen (Lid 1964) and the SW coast of communities recall observations from other Arc- Spitsbergen (Triloff 1944; Eurola 1968). Parts of tic areas (cf. Gjessing & 0vstedal 1976). Water Bjarnnya have a slightly more favourable local saturation is a main factor controlling the super- climate, e.g. the southern slopes of Miseryfjellet, ficial temperature, and the former is dependent the valley at Kvalrossbukta, and the Ymerdalen on aspect, slope, and drainage. Sun heating is valley, judged from the local development of

Fig. 4. Generalized profile on the northern plain of Bjarnoya from the brink east of Kapp Posadowsky. passing southeast over Kikutkollen 12.3rn a.s.1.) to Lomvatnet (32.7 rn a.s.1.). 93

Table 8. Vascular species occurring at higher elevations on Bj0rnaya : Sterile specimens only.

Altitude Skuld Hambergfjell Fuglefjell mad. 454 Cerastium regelii Cochlearia groenlandica Saxifraga caespitosa 452 Phippsia algida 440 Cerastium arcticurn Phippsia algida Saxifraga cernua Ranunculus hyperboreus" Saxifraga rivularis Cochlearia groenlandica Saxifraga rivularis 430 Saxifraga caespitosa 310 Phippsia algida Cochlearia groenlandica Festuca rubra" 400 Poa alpina Oxyria digyna Poa pratensis ssp. alpigena'

380 Poa alpina 350 Ranunculus sulphureus Cerastium regelii 3 20 Saxifraga nivalis 310 Saxifraga oppositifolia 300 Draba alpina Draba norvegica 290 Phippsia concinna 250 Descharnpsia alpina Festuca vivipara Salk herbacea Sa1i.x polaris 220 Sagina intermedia Sa-rifiaga tenuis 200 Ranunculus pygmaeus vegetation. These are the habitats of plant com- cea and S. polaris, do not ascend above 250 m munities and species which are absent from the a.s.1. on Bj~m~ya.This is a considerable down- monotonous and water-logged northern plain, ward displacement from the Finnmark coast, e.g. Arabis alpina, Saxfraga aizoides, and where both species reach over 800 m ad.(Dahl Taraxacum cymbfolium. 1934), and still more from interior North Scandi- navia where they ascend to at least 1600 m (Engelskjm 1986 unpublished). The third ligni- 6. Vertical floristic belts fied species on the island, Salk reticulata. ascends to 80 m ad. Altitude limits of vascular species Table 9 relates the vertical distribution to heat Table 8 summarizes the altitudinal distribution sums, calculated as Degree-Days for June- of 25 species which were observed between 200 September or July- August. and 454 m ad. on the mountains of Bjarnaya. About half of the Bjarnaya species persist and Most probably some of them will also be found reproduce at the 200 m level, which corresponds on a higher altitude when the summits of Urd to DD-2 = 190. The remaining species are con- (535 m) and Verdandi (462 m) become explored. fined to lower hills, valleys and plains. Some of

Marking with a O notifies sterile plants only. This them are hydrophytes, e.g. Arctophila fulva and does not apply to Cardamine nymaniiand Ceras- Hippuns vulgaris, whereas others may be absent tium regelii which do not usually flower on from the summits for thermal reasons, e.g. Salk Bj~rnaya. reticulata, Cerastium cerastoides, Saxifraga aizoi- It is noted that the dwarf willows, Salix herba- des, and Taraxacum cymbifolium. 94

Table 9. Heat sums (as Degree-Days) at various eleva- making up the peaty coat of these ice mounds, tions on Bjarnaya, related to uppermost occurrences which are very similar to palsas (Svensson 1964; of vascular plant species. DD-4: Degree-Days. June- Lundquist 1969). The Hambergfjell field September. DD-2: Degree-Days, July- August. " : Sterile specimens only. measures approximately 0.4 x 0.3 km and is situated somewhat inside the margins of the sheer precipices to the SW, between 400 and Altitude DD-4 DD-2 Upper altitudinal limit 440 m ad. The largest mounds measure up to m ad. of vascular species 20 x 5 x 3 m. The peaty mantle is 0.3 to 0.5 m 535 - 5 60 Floristic content thick and covers ice lenses (Fig. 6) which rest on unknown Spirifer limestone pavements of the old plateau 450 65 90 Ceractiurn arcticurn. C. surface. No silty or clayey sediments were seen regelii. Coclileuria groen- inside the mounds. lundrca. Phippjia algida. Dark brown peat is exposed by erosion on Sarifiaga caespirosa. S. many of the larger, partly collapsed structures. cernua. S. rirularis Rhacomitriurn stems are recognizable also in the 0.y.ria digrna. Poa alp;- nu deepest peat layer. On intact surfaces the cryptogamic cover consists of Rhacomitriurn lanugino- 350 I40 I 30 Rarrunculus sulphureus sum. Drepanocladus uncinatus. Aulacomnium Draha alpinn. D. none- palustre. A. turgidum. Cetraria nivalis, and giccl, Pliippsia concinria. Sarjfrap~nidir. S. op- Sphaerophorus globosus. It is mainly non-vascu- potitjfblia lar, but did support some fungi, even the large Lepista polvgoni. Between the mounds are -_'50 220 I78 Dercliampsia alprna. water-soaked moss carpets of Calliergon sarmen- Fesrrica riripara. Salir Iierhucea. S. poluric tosum. Drepanocladus uncinatus, and Hygroh-vp- Cardaniine nJmanii. num alpestre. and of vascular species, Phippsia Sarjfraga Iiirculusc , S. algida. Saxijiaga rivularis, and Cochlearia groen- renui.~. Sa,gitrci inter- landica seedlings, as well as the hygrophilous media. Silene uc~~rrlis' fungi Galerina pumila and Leptogloswm loha- 200 260 190 .JraDis ulpina. Papaiw tum. dahliarrunr. Ranuncuhrs Proceeding to Alfredfjell (410 m ad.) we pJ'gmarus found a smaller area with similar peat mounds (Figs. 7, 8). They cover the top of the mountain, and eroded peat is exposed in terraces on its NW side. I suppose these structures are ice-cored, but 30 400 260 54 native species at Ion time was too short for excavation. levels

Ways of .formation of the Bj0rmya peat mounds Several factors contribute to create ice-cored mounds of the kind described (Seppala 1976). Formation ojpeat mounds on the high ground 1) Low annual mean temperature, ojBjmn0ya 2) a sufficiently high water saturation of the During the 1983 expedition, Mr. Sigbjorn Dun- ground, fjeld and the present author visited the summit 3) thermal isolation by peat during summer plateau of Hambergfjell. We came across an thaw, extensive area of 1-3 m high, peat covered and 4) cyclic erosion and regeneration courses. ice-cored hillocks (Figs. 5, 6). Rhacomitrium The mean annual temperature at the 450 m level lanuginosum is the main bryophyte species on Bjarnaya is about -5" (Table 4). Precipitation 95

Fig. 5. Peat mounds up to 3 m high on the summit plateau of Hambergfjell, approximately 430 m a.s.1. 16 August 1983.

Fig. 6. Detail of ice core in mound shown on Fig. 5. Scale (I) carved into ice, 20 cm high. 16 August 1983. 96

Fig. ?. Peat mounds on the summit of Fuglefjell, 410 m a.s.1.. looking north towards Miseryfjellet (535 m a.s.1.) and the northern low ground of Bjornoqa. 16 August 1983.

Fig. 8. Eroded Rhacomirrium lanuginosum peat terraces on Fuglefjell, probably ice-cored. The section is 3 m high. I6 August 1983. 97

is liable to be trapped in absorbent moss hum- 1. Equisetum antense L. (62).., mocks, the growth-of which is enhanced by the Frequent in various parts of the island, especially stability of the plateau and the bird manuring. on the N coast, reaching the altitude limit 130 m Situated above very large bird rookeries (Fig. 10, a.s.1. on the plain W of Miseryfjellet. Section 8), the plateaux are exposed to steady The species is indifferent to bedrock and even dropping of manure, feather, fish bones, and grows in Ursa sandstone blockfields, but requires shells. However, the plateaux are not breeding seasonal flooding and shelter from wind abrasi- sites, which would lead to overdunging and on. trampling. Characteristic of Cetraria delisei- Drepano- Although Rhacomitrium lanuginosum is a fru- cladus uncinatus and Oxyna-Saxijiraga cernua gal species (Jalas 1955) it grows vigorously with communities, or associating with Deschampsia the mentioned bird manuring, and a comparable alpina (Table 7, D.3), but also in the most growth at bird rookeries has not been described species-rich, eutrophic moss carpet communities. elsewhere in the North Atlantic Arctic (cf. Stein- dorsson 1945; Elvebakk 1979; Brattbakk 1983). Nor have I seen descriptions of Rhacomitrium- 2. Equisetum scirpoides Rich. (1 2). coated palsas (cf. Datkiewicz 1967; French 1971; The two records from the N coast (Engelskjan & Hambrey & Swett 1980; Vorren 1972). Peat accu- Schweitzer 1970, Map 16) must be excluded, mulations on Spitsbergen, described by LAg because my recent revision of the material (1980), may be compared with the Bjarnaya (TROM) proved these collections to be gracile occurrences. modifications of E. variegatum. Accordingly, E. Peat mounds were not observed on the low scirpoides is confined to the S half of the island, ground of Bj~rn~ya.Manuring is less in the ascending to 100 a.s.1. on the S slope of Misery- interior of the island, and the seasonal thermal m fjellet. fluctuations may be less suitable for formation It grows on well-drained, licheniferous of such structures. bryophyte turfs with a silty subsoil, often associa- It seems that the high ground of Bjamaya is ted with Salixpolans. the only place in the North Atlantic where such structures have developed at a large scale. Their time of formation is unknown, and it would be 3. Equisetum variegatum Schleich. (19). desirable to have them surveyed and dated. Localized to the N and S parts, from 10 to 130 m a.s.1. Preferentially in the carbonatite areas; on flooded or cryoturbated lithosol with a thin 7. Specific distribution patterns bryophyte turf, but also in nearly non-vascular within the island Drepanocladus uncinatus carpets in the Kulm The following record of horizontal and vertical areas. distributions of the vascular species of Bjam~ya is founded on accumulated information from field work in 1967 (cf. Engelskjan & Schweitzer 4. Juncus biglumis L. ( 16). 1970) and in 1983. Scattered and partly overlooked; most frequent Two species are new to the island: Alchemilla in the western pond areas but now also found in glomerulans and Saxijiraga foliolosa. The first three places in the interior. Its vertical range is numeral in the species headings corresponds to from 20 m on the brink to 50 m ad.at GAsvatna numbers of distribution maps (1 -54), which in Ymerdalen. also include a tabulation of vertical distribution. Confined to level, cryoturbated, silty patches The number in brackets gives the number of sta- with a short bryophyte or cyanophycean crust, tions known of the species on Bjamaya. Intro- associated with Phippsia concinna, Koenigia duced species are enumerated after the 54 sup- islandica, Sagina intermedia. and Cerastium posedly native species. regelii (Table 7, D.l). 98

2. Luzula arcuata (Wg.) Sw. ssp. arcuata(56). state, now recorded from 12 new stations on the Locally frequent in the interior, more sparsely S half of Bj@m@ya,including the uppermost one near the brink. The upper limit of only 150 m at Dipilen, 103 rn ad. a.s.1. is reached at Mefaringen W of Miseryfjellet. The species was found in several places in the Associated with sandstone lithosol and avoid- interior Kulm areas, but seems to prefer slightly ing carbonatites, but once found together with bird-influenced places on the brink. the eutrophic Silene acaulis (Table 6, bedrock Recorded from a variety of moss carpet com- Ill.?). munities ranging from Sphagnum riparium and L. arcuata is a characteristic species in the Calliergon sarmenfosum on the oligotrophic, to vegetation of polygon fields and boulder areas, Paludella squarrosa and Drepanorladus inter- often occurring on small sandstone outcrops mediuson the eutrophic side (Table 7, E). within limestone areas (Table 7, C 1.3). 10. Desrhampsia alpina (L.) R. & S. (70). 6. Alopecums alpinus Sm. (9). Frequent in the polygon areas inside the brink Restricted to the S half of the island, from 35 rn and ascending to the mountains: 250 m on Skuld at Royevatn to 150 m ad. in Ymerdalen. Five and approximately 300 m ad. between Ver- new localities were found in 1983. the total being dandi and Skuld (Brattbakk 1984). nine. The clones. carrying a few culms, grow The species appears indifferent as to bedrock, centrifugally on flooded, eutrophic moss carpet but requires flooding or cryoturbation, such as in communities with Hr’grohypnum spp., Drepano- the centre of polygon field communities, associa- dadus intermedius. Phippsia concinna. Dupontia ting with Festuca vivipara, encircled by Rhacomi- psilosantha. and Ranunculus sulphureus. trium lanuginosum turfs (Table 7, D.3).

7. Calamagrostis neglecta (Ehrh.)G. M.S. (26). 1 I. Poa alpina L. var. vivipara L. (60). By ponds and ribers, especially on the N coast, Concentrated in limestone areas at various levels, ascending to 52 m ad. at Krillvatn in the SE. ascending to 400 m on Skuld and 380 rn a.s.1. on New finds were made at Ellasjoen and in the Hambergfjell. interior. The species has definitely eutrophic demands, Characteristic of peaty banks, often associated but occurs on intermediate, pelitic shales on the with Dupontia psilosantha. Culms are sparse and summits of Miseryfjellet. do not appear until mid-August. The vegetative development is best in protected herb communities along brook trails. 8. Arc.tophila.firl~a(Trin.)Rupr. (17). 12. Poaprarensis L. ssp. alpigena (Fr.) Hiit. (33). A hydrophyte growing in brooks and lakes, up to 100 m a.s.1. in a pond E of Grautauget on the Locally common in some sections of the coast interior plateau. and other moderately bird-influnced places, to Exacting as to hydrology, it seems indepen- 230 m below the Ymerdalen kittiwake rookery dent of bedrock. Extensive stands were disco- and to 400 m a.s.1. on the Fuglefjell plateau. vered in the Kulm area around Storlona, upper Usually in luxuriant grass vegetation, but also in Engelskelv watercourse. Here it brims the ponds. eroded peat on the brink. Sterile at the altitude in open water as well as in soaked Calliergon limit, elsewhere flowering in August. sarmentosum carpets with few other vascular species. 13. hccinellia phtyganodes (Trin.) Scribn. & Culms are rare (cf. Engelskjon & Schweitzer Merr. (39). 1970). Dispersed on the lower coastal brinks. Being a principally littoral species (Table 7, A l.l), it 9. Dupontiapsilosantha Rupr. (44). ascends only to 30 m a.s.1. Previously overlooked due to its often sterile In moss carpet communities, usually as vegeta- 99 tive, stoloniferous clones, but developing tufts 18. Carexlachenalii Schkuhr(4). with few stolons and flowering culms when No new finds were made of this restricted growing on silty, well-drained brink sites. species, which has four stations below 50 m ad. The northern group of localities at Lakselva 14. Phippsia algida (Sol.) R. Br. (1 12). was unchanged since 1967. The tufts, partly One of the species having the most stations on sterile, were intact in 1983 as then. The best deve- Bjomoya, including the mountain plateaux: loped plants were flowering in mid-August in a Skuld 452 m, Hambergfjell 430-440 m, and community of Drepanocladus uncinatus. Festuca Fuglefjell410 m ad.Although occurring within rubra, Salix polaris, Polygonum viviparum. Ceras- the Kulm barrens, it has a certain eutrophic pre- tium regelii, and Ranunculus pygmaeus. ference. The high frequency of P. algida seems related 19. Carexsubspathacea Wormsk. (3). to the numerous available habitats with flooded mineral soil or moss carpets (Table 7, D.2). Discovered on one spot in 1967, two new stations were added in 1983: NW coast, river bed N of 15. Phippsia concinna (Th. Fr.) Lindeb. (32). Hira, 23 m a.s.1. in a Drepanocladus uncinatus carpet: and NE coast, Framnes, NE side of tarn Less common than the preceding Phippsia and 18.0 m ad., covering 10 x 6 m within a wet, confined to open, cryoturbated patches of littorally influenced Drepanocladus uncinatus carbonate-containing subsoil, especially the red carpet. It was still growing at Lusbekken on the Ambigua marl, the Hecla Hoek limestone, and N coast, 6-8 m a.s.1. The species seems to occur Spirifer limestone. Mainly on the 30 and 100 m only in the sterile state on Bjornoya. plains, but extending to 290 m ad. in the pass between Hambergfjell and Fuglefjell. P. concinna is numerous in eutrophic habitats, 20. Salix herbacea L. (I 17). accompanied by exacting species such as Juncus Extensively distributed and ascending to 250 m biglumisand Saxifraga hirculus(Tab1e 7, D. I). a.s.1. on Skuld, dispersed also in the barren Kulm sandstone areas, but clearly avoiding the bird- 16. Festuca rubra L. (102). influenced sites at the southern end of the island. Common in most sections of the brink, under S. herbacea also enters the limestone areas, bird cliffs, on skua hummocks in the interior, as associating even with Saxifraga oppositifolia on well as in non-manured limestone bluffs. Spirifer limestone on the northern coast, but Concentrated in low-lying areas, but ascend- tends to become replaced by S. polaris on this ing to 400 m a.s.1. on Fuglefjell. Avoiding the substratum. Kulm areas, if not promoted by bird manure. In polar gull areas it forms carpets without culms 21. Salixpolaris Wg. (80). (Table 7, A 2.2). Concentrated within the carbonatite areas and 17. Festuca vivipara (L.) Sm. (63). locally common, reaching the same altitude limit as S. herbacea. 250 m ad., on Skuld. Preferably Dispersed over the island, including the interior, in bryophyte communities, e.g. of Drepanocladus but less common to the W and S. Vertically, it uncinatus. D. intermedius, Oncophorus spp., reaches 250 m ad.on Miseryfjellet. Tomenthypnum nitens, Timmia austriaca, and Equally common in the limestone and Kulm Orthothecium chtyseum. areas, but avoids manured situations. F. vivipara is a specialist of gravelly, cryoturbated habitats, S. herbacea x polaris often associating with Luzula arcuata, Deschampsia alpina. and Cerastium regelii (Table Often met with in mixed growths of the parent 7, C I. 2,3, D. 3). species or with one of them (cf. Engelskjen & Culm development is not regular and usually Schweitzer 1970). Hybrid clones have their sepa- takes place from the end of July. rate signature on the S.polarismap (21). I00

22. Saliv reticulata L. (21). It is met with regularly at high levels, reaching A restricted species which is fairly common on 440 m a.s.1. on Skuld. A character species of dolomite and limestone dolomite around Ellasjsen ; elsewhere only as small clones on local limestone outcrops. Its barrens, it is best developed in bryophyte peat, highest occurrence is 80 m ad., in the floristical- seldom missing in skua hummocks and other ly rich area to the S of Miseryfjellet. local peat mounds. S. rericulata prefers eutrophic peat in protected A glabrescent biotype of C. arcticum was situations and is often associating with S.polaris. found S of Ellasjsen at 80 m ad.

23. Koenigia islandica L. (45). C.arcticum x regelii Several new stations were found during the late Intermediate clones which may be this hybrid summer of 1983 in the NW and central part of combination were found at Kikutkollen, 35 m, Bjsrnaya. It grows mainly in the low-lying and on the N side of Lakselva, 15 m ad., in both plains, reaching 130 m a.s.1. at Kollerskardet, W places with the supposed parent species, cf. of Miseryfjellet. Rsnning (1959, p. 24). A specialist of flooded or springy communities, Koenigia may be the only vascular plant 28. Cerastium cerastoides (L.)Britton (1 2). species in floating Calliergon-Drepanocladus This species, of very local distribution on the moss carpets by pools and lakes (Table 7, F ii). island, was found at new localities in the Spirifer limestone area on the N coast and around the W end of Ellasjsen on the coast, reaching 45 m 24. Oxyria digyna (L.) Hill (108). SW ad.on Kikutkollen. Seems restricted to a parti- Frequent all over the island, concentrated in the cular community of intermediate snow-cover, luxuriant herb communities, and ascending to accompanied by Brachythecium reflexum, Pelti- 400 m a.s.1. in the bird-influenced moss carpets gera kristinssonii, Oxyria digyna and Ranunculus on Hambergfjell. pygmaeus. C. cerastoides is also associating with Oxyria also occurs on barren polygon ground Alchemilla glomerulans in its one and only habitat and in seasonally flooded Cetraria delisei com- on Bjsmaya. munities. 29. Cerastium regelii Ostenf. (73). 25. Polygonum vivipamm L. (63). Unevenly distributed and largely avoiding the Dispersed in most parts of the island, reaching Kulm areas. It appears to be one of the hardiest 150 m ad. on the N plateau of Miseryfjellet. species, ascending to 454 m on Skuld and 350 m Preferring eutrophic ground and rare in the ad. on Hambergfjell. On the mountain tops it sandstone areas. grows branched in shale debris, or more contracted, on polygon mud. 26. Sagina intermedia Fenzl(58). Flowering specimens were found i.a. at the sheltered Carex lachenalii station by Lakselva, 15 Dispersed and locally frequent up to 220 m ad. m a.s.1. on the NW and W sides of the Miseryfjellet plateau. 30. Stellaria humifusa Rottb. (1 3). ~I Tolerating cryoturbation and bird trampling, Six new stations were added, and the species the species may be numerous in the central part appears as dispersed in brink vegetation (Table of limy detritus polygons, as well as in moss car- 7, A 1.2). It was met with also at the lake Skutilen, pet communities on the brink. 65 m a.s.1.. and 3 km from the sea. This is well above the supposed Holocene marine limit. Bird 27. Cerastium arcticum Lge. (I 18). dispersal to this site appears probable. Here it is Widespread, with a high number of recorded accompanied by Arctophila fulva in Drepano- stations, although preferring eutrophic habitats. cladus intermedius- Hygrohypnum carpets. 101

3 1. Silene acaulis L. (57). Particularly numerous on calcarenite plains (Bedrock group 11.2) to the NW; in the Spirifer limestone areas to the N, and in the limestone, dolomite and baryte areas to the SE. It was one of the few plants growing on lead-poisoned dolomite and baryte barrens on the hill Blyhatten. The vertical range of S. acaulis is moderate, attaining 220 m ad.on the southernmost station, where it is sterile.

32. Ranunculus hyperboreus Rottb. (38). Dispersed on the island, mainly along the brink and on the northern plain where ponds are numerous. Occumng also on the summit plateau of Hambergfjell, 435 m ad., sterile but vegetatively luxuriant. Although promoted by manure, R. hyperboreus occurs at oligotrophic pools with Sphag- num riparium and Calliergon sarmentosum. More commonly it is growing with Calliergon stramineum, Cardamine nymanii, Hippuris vulgaris. and Dupontia psilosantha. Fig. 9. Ranunculus sulphureus flowering in Ymerdalen, 33. Ranunculuspygmaeus Wg. (35). southern part of Bjomeya. Luxuriant meadow on Tetradium limestone, sheltered site. Of restricted local distribution, reaching 200 m 16 August 1983. ad.on the N plateau of Miseryfjellet. The species appears as demanding and prefers Miseryfjellet, at Kvalrosselva, and in parts of well vegetated sites in the carbonatite areas. Ymerdalen. P. dahlianum ranges from 12 m at the radio 34. Ranunculus sulphureus Sol. (65). station to 200 m ad. on the W side of Ymer- Widespread and locally common, ascending to dalen. Plants from wind-exposed dolomite ridges 350 m on Skuld and 240 m a.s.1. in Ymerdalen. A are contracted and often sterile, irrespective of substantial part of its stations is above 100 m a.s.1. altitude. The species frequents various habitats, i.a. blockfields and gravel polygons in the sandstone 36. Arabis alpina L. (18). areas, shale screes in the upper slopes of Misery- Confined to the southern, hilly part of the island, fjellet, and luxuriant herb communities along where it ascends to 190-210 m a.s.1. on both watercourses in the carbonatite areas (Fig. 9). sides of Ymerdalen. A usual substratum is well-drained, coarse 35. Papaver dahlianum Nordhagen (48). dolomite or limestone talus supporting very few Confined to carbonatite lithosol in the N and S cryptogams and the following vascular species: part of the island (Table 7, C l.l), apparently Draba alpina, Papaver dahlianum, Saxijiraga lacking in the NW part and in the Kulm areas. caespitosa, S. oppositifolia, and Silene acaulis Nor were there poppies in the Triassic shale (Table 7, C I. I). screes on Miseryfjellet. P. dahlianum is numer- Sterile rosettes are frequent, indicating un- ous and well-developed only on the S side of favourable conditions for flowering. I02

37. Cardamine nymanii Gand. (67). In slightly manured communities on the brink, D. norvegica appears to be locally more frequent Dispersed over the island, ascending to 230 m in than the preceding species. Ymerdalen: 220 m ad. on the N plateau of Miseryfjellet. Growing on flushed moss carpets by streams 41. Sedum rosea (L.) Scop. ssp. arcticum (Boris- sova) Engelskjsn & Schweitzer (55). and ponds, the species may also enter the Kulm sandstone areas. Previously considered restricted to the brink sec- Flowering is rare as previously stated tions with Kulm or other bedrock poor in lime, (Engelskjsn & Schweitzer 1970), but takes place two new stations were found in the interior, on in favourable habitats from mid-August cherty dolomite and hard calcarenite respective- onwards. ly. We did not find it above 70 m ad., but the locality of Brattbakk (1984) between Beinneset 38. Cochlearia groenlandica L. (approximately and Revdalen on the SE coast may be above 200). 100 m. Concentrated in bird-cliffs along the entire coast The species forms an interesting community as well as in the Ymerdalen kittiwake rookeries, of its own, see Engelskjsn & Schweitzer (1970) but also dispersed to the interior. As to number and the present Table 7B. 3. of stations and biomass, it must be the most 42. Saxifraga aizoides L. (4). important vascular species of BJ~rn~ya.It ascends to the high plateaux and summits on The original locality of Schweitzer (Engelskjsn & Miseryfjellet, 454 m, Hambergfjell, 440 m. and Schweitzer 1970) was investigated in 1983. About Fuglefjell, 410 m a.s.1. In non-manured situations 110 tufts are dispersed on a peaty, soliflucted, as on the shale screes of Miseryfjellet, specimens dolomite lithosol area to the SE of the hill 100.4 are contracted and lilac-flowered. In cliffs and m on Blasen, between 50 and 65 m ad.Another gullies of the rookeries, well-developed, pure population was found on alluvial silt 120 m a.s.1. Cochlearia stands are seen everywhere. The on the S slope of Miseryfjellet, here less numer- bird-influenced summit plateaux support a ous and associated with Equiserum scirpoides and Cochlearia- Drepanocladus uncinatus sociation Salk Polaris. A third locality was discovered by with few other vascular plant species (Table 7, A Brattbakk (1984) between Ymerdalen and Ella- 2.1 ). sj~en. The species is restricted on Bjsmsya and 39. Draba alpina L. (89). appears exacting as to subsoil and hydrology. Flowering starts in mid-August. Widespread in parts of the island; ascending to 300 m between Hambergfjell and Alfredfjell, and 43. Saxifraga caespitosa L. (108). to 240 mad.on Miseryfjellet. I now assess this to be the most ecologically ver- D. alpina is common in the carbonatite areas, satile and common saxifrage on Bj~msya(cf. and it is merely an exception to find it on sand- Engelskjan & Schweitzer 1970, p. 17). This may stone. not be fully evident from the distribution map Often contracted and with a few flowers on the wind-exposed dolomite barrens, but with a well- because some stations may have been left un- developed stature in moss carpet communities. noticed, due to its commonness. However, it has a preference for the low-lying carbonatite areas, flourishing also on the dolo- 40. Draba norvegica Gunn. (70). mite barrens. Widespread and occasionally also growing in the S. caespitosa ascends to the high plateaux of Kulm areas. It often occurs together with D. Skuld (450 m) and Hambergfjell(430 m a.s.1,). alpina, and they reach the same altitude limit: 300 m a.s.1.between Hambergfjell and Alfredfjell, 240 44. Saxifraga cemua L. (101). rn ad.at Krykkjedarnrnen, upper Ymerdalen. Formerly considered as the most widespread 103 saxifrage, this species has somewhat fewer locali- At its upper limit on Fuglefjell, 3 10 m a.s.l., ties than S. caespitosa, but is more conspicuous only well outside the manured zone and confined due to its stature and larger flowers. to an escarpment with exposed Cora limestone S. cernua seems more exacting than S. caespi- on the edge of Ymerdalen. rosa, by avoiding wind-exposed crest and brink areas. Well-developed plants are found in pro- 49. Saxifraga rivularis L. s.str. (78). tected herbaceous communities in the carbona- Numerous in bryophyte carpets along the brink, tite areas, but it is also alpine, reaching 440 m in the Ellasjaen depression, and Misery- a.s.1. on the Triassic shale screes on Miseryfjellet. fiellet. Schweitzer (pers. comm.) noted it in the The species shows a preference for moist moss southem bird-influenced valleys and mountain carpets, and is often the only vascular plant spe- slopes.

cies. Reaching 440 m a.s.1. on Hambergfjell~~ and on Skuld, it is among the hardiest species (Table 45. Saxifagafoliolosa R. Br. (I). 7, B. I). This important accession to the Bjumuya vascular flora was made by Brattbakk (1984). The 50. Saxifraga tenuis (Wg.) H. Smith (42). species grows locally on Oswaldfjell, S of Glugg- As different from S. nivalis, this species grows in dalsvatna, at an altitude of approximately 120 m flushed, eutrophic habitats, preferentially in the a.s.1. carbonatite areas away from the brink and bird rookeries. 46. Saxifraga hirculus L. (49). The highest record is in Scotpidium turges- The map shows a continuous distribution along cens- Hygrohypnumcommunities on the Spirifer rivers and lakes at the southern end of the island, limestone plateau on Skuld, 220 m ad. whereas it is rare on the northern half. Confined to carbonatite rocks and best developed in 5 1. Chrysospleniurn tetrandmm (Lund) Th. Fries eutrophic bryophyte carpets. (1 0). It was also noted on flushed, cryoturbated Sporadical on Bjurnuya but locally frequent in limestone gravel, mainly in a vegetative state, e.g. springy bryophyte communities below limestone W of Lakselva on the N coast and S of Ruyevatn. precipices at Miseryfjellet and Ymerdalen (to The species was sterile at its altitudinal limit, a 200 m a.s.l.), as well as in a bird-influenced brook mossy dolomite ridge 220 m a.s.1. on the SE coast. trail from MAketj0rn, 15 m a.s.l., at Kapp Levin on the E coast (Table 7, B.2). 41. Saxifraga nivalis L. (72). It is accompanied by bryophytes such as Bra- Dispersed, but apparently rare on some stretches chythecium salebrosum, Plagiomnium ellipticum, ofthe NW quadrant and in the interior sandstone Marchantia polymorpha, Paludella squarrosa, barrens. It prefers a broken micro-topography, Philonotis spp., Rhytidiadelphus squarrosus, fila- like around Lakselva, below Miseryfjellet, and mentous chlorophyceans, and well-developed near the kittiwake rookeries in Ymerdalen. Saxifraga cernua or S. hirculus. Found up to 400 m a.s.1. on Hambergfjell. 52. Alchemilla glomemlans Bus. (I). 48. Saxijiraga oppositifolia L. (81). Confined to a wind-protected gully eroded in a Most common, and locally dominant, in the Spirifer limestone plateau on the N coast, northern and southern parts of the island where 200-300 m S of Bjumuya Radio. Three large carbonatites prevail. The few finds in sandstone individuals were found together with Festuca areas seem related to limestone admixture in rubra, Oxyria digyna, Cerastium arcticurn, C. polygons. The species is absent from interior cerastoides, Ranunculus pygmaeus, and Saxi- Kulm areas as well as in sections of the brink fraga cernuu. Among the bryophyte carpets sur- with bedrock poor in lime. rounding them I noted Bruchythecium reflexum, 104

Drepanocladus uncinatus. Philonotis sp., and 4) According to Mr. Idar Tetlivold, former Timmia austriaca. head of Bjamoya Radio station, there is a This species assemblage gives the impression Taraxacum growth at the outlet of Ellasjoen on of being native. However, it is close to the station the SW coast. I was not aware of this find during area, and several introduced species were found our stay in 1983, and no specimens have been nearby. This casts some doubts on the sponta- secured. The record is presumably correct as to neity of A. glomerulans on Bjomoya, cf. the generic determination. section on introduced species. A singular station 5) A few Taraxacum specimens were found is also known at Jan Mayen (Lid 1964), here at Tunheim City in 1957 and 1967, but not redis- connected with fox burrows. covered in 1983. The primary occurence may be It is of interest to observe the development of located in cliffs facing the sea, or it has dis- Alchemilla glomemlans on Bjsmoya during the appeared because of the mining activities. coming years. The introduced vascular element on Bj0rn0~va 53. Hippuris vulgaris L. (23). Here is summarized the status of non-indigenous At pond and stream margins from 10 to 30 m species, found up to the present, around the ad., mainly on the N and W coasts. In 1983 actual and former settlements (Hanssen & Holm- found at Ellasjoen, where it was previously boe 1925; Ranning 1959; Engelskjon & Schweit- unknown although the place is well investigated zer 1970). (Ronning 1959). Very large numbers were seen in the lakes of N. Flakmyrvatna, where Hippuris 1. Agrostis capillaris L. appears to be promoted by some disturbance and One sterile tuft S of the Radio station, in a manuring by polar gulls and skuas of the floating protected gully 15 m a.s.1. (1983). Previously not moss carpets. recorded from Bjarnoya.

Deschampsia caespitosa PB. 54. Taraxacum cymbifolium H. Lindb. (5). 2. (L.) Previously treated by Engelskjon (1 967), some Persisting at Tunheim and in a protected gully S new details of its distribution and performance of the Radio station (l983), as well as at Krillvatn (Brattbakk 1984). Culms develop in late August, on Bjomaya are provided here. but ripe seed were not observed. 1) The main occurrence is in the brook valley S of Roedvika on the SE coast, 40-55 m a.s.1. T. 3. Poa alpina L. var. alpina. cymbifolium occupies a herb community (Table 7, C 2.3), but is also entering a more exposed Fes- A non-viviparous strain was found in the gully S tuca rubra-Oxyria community on the brink, as of Bjomaya Radio, represented by four vigorous more contracted plants. It is competed by closed tufts flowering on 22 August 1983. I suspect it to F. rubra carpets in the polar gull area at the be introduced, because all native strains appear mouth of the small valley facing the sea. The to be viviparous (cf. Engelskjon & Schweitzer number of individuals counts more than 200. 1970, pp. 6-7). A similar strain from Tunheim 2) A brook ravine N of Kvalrossbukta, dry in (Ronning 1959) appears extinct. August, 20 m ad. T. cvmbifolium occupies a similar intermediate zone between a wind- 4. Poa pratensis L. ssp. alpigena (Fr.) Hiit. exposed, barren plateau and the streambed. Tall-growing, deviant biotypes persist in one place There were about 30 individuals. at Tunheim, at Hammerfeststua in Herwighamna 3) A tributary brook ravine N of Kvalrosselva, close to the Radio station (1983), and at Krillvatn 250 m from the sea and 20-30 m a.s.1. T. cvmbi- (Brattbakk 1984). They are richly flower- folium grows in herb communities, on loose allu- ing and ripen seed. We cannot discount the pos- vial gravel, and on shale debris, partly together sibility of their being luxuriant modifications of with well-developed Papaver dahlianum, totally the native biotype (Engelskjon & Schweitzer about 200 individuals. 1970, p. 7). 105

5. Rumex acetosa L. sidered as well as the chronology indicated by Persisting at Engelskelva N of Tunheim City, Miller (1982). close to the walking route to Bjarnaya Radio The available Quaternary geological evidence (1983), but only sterile, and at Krillvatn (Bratt- can be summarized as follows: The northern part bakk 1984). of the Barents Sea region was a glaciation centre during the Late Vistulan. The ice sheet probably 6. Stellaria media (L.) Vill. passed Bjarnaya and advanced S and W to the margins of the Barents Sea shelf. But conditions Not rediscovered (cf. Engelskjan, Kramer & on land and the chronology of glaciation and Schweitzer 1972), probably extinct. deglaciation of Bjarnaya are still inconclusive. Hyvarinen (1968, 1972) describes pollen spec- 7. Ranunculus acris L. tra from two lakes on Bjarnaya, going back to Not rediscovered at any of the former settle- 8300 years B. P. His observation of Alms, Betula, ments, presumably extinct. Pinus, and even thermophilous tree pollen trans- ported to Bjarnaya is of considerable interest. 8. Alchemillafilicaulis Bus. The phytogeography of Bjarnaya was discussed Formerly at Krillvatn (Ranning 1959) but not by HadaE (1941, 1960), bnning (1959), and rediscovered up to 1983-1984 and presumably Engelskjan & Schweitzer (1970). This discussion extinct. has been impeded by the insufficient Quaternary geological knowledge of the Barents Sea region. Because the evidence is still ambiguous, nothing can be said of definite phytogeographical con- 8. Historical and ecological aspects sequence. However, an ecological approach of the Bj~rnplyaflora and vegeta- could be attempted to elucidate certain phytogeographical problems pertaining to the tion Bjornaya flora. The time and extent of Pleistocene glaciations Irrespective of the time and extent of the affecting the Barents Sea and Bjarnaya are still a Barents Sea glaciations, high-glacial conditions matter of discussion. Conflicting views have have affected Bjarnaya at times and caused an been presented also by geologists on the condi- ecological change. The shift from an ice-free to a tions during the Vistula glacial maximum, about pack ice sea around the island implies lower 18,000 years ago (for review, see Elverhai & Sol- temperatures and a different circulation pattern heim 1983). (Lamb 1972; Olausson 1972). Horn & Orvin (1928) found numerous traces Solar radiation was influenced by a higher of glaciation and indicated @. 55) a submergence atmospheric transmittivity due to less water of the northern low plain of Bjarnaya ‘in a part vapour and seafog (Kukla 1972), and we may of the last Ice-period’. imagine a climate on Bjarnoya quite different Considering Svalbard at large, Salvigsen & from the present maritime Arctic one. Nydal ( 198 I) provide ample documentation of A considerable local glaciation would restrict the Middle Vistulan having been an interstadial the plant habitats, but the possibility of ice-free with less glaciation than at present. The Late shelf areas must be considered. Vistulan glacier advance was limited on the west- The 400 m high precipices at the southern end ern part of Spitsbergen, but this ice sheet covered of Bjarnaya (Fig. 10) would be the least liable to the eastern archipelago completely (Salvigsen accumulate glacier ice, cf. the conditons on the 1981). The Early Vistulan, about 100,000 years cliff coast of Bouvetaya, South Atlantic Ocean ago, was a period of extensive glaciation with the (Orheim 1981). The sea-bird populations may possible exception of alpine nunataks on Prim have decreased during pack ice conditions so as Karls Forland. The palaeogeographical recon- to reduce their edaphic influence. struction by Liestal (1972), leaving shelf areas on The botanical problem may be as follows: NW Spitsbergen ice-free, should also be con- Which of the present-day vascular species could 106

Fig. 10. The southern end of Bjornoya seen from the Harnbergfjell plateau. 350 m high precipices below Fugle- fjell. Crevassed Drepanocladus uncinatus - Torrula ruralis peat on plateau margin. with Cochlearia groenlandicn. 16 August 1983

endure in steep cliffs throughout a glacial period, Bj~mayawas to study the amount of ecological when ground heating was chiefly caused by diversity and the potential for that biome to radiation? Judged from their present-day perfor- support more than the 52 vascular native species mance on Bjamaya, two thirds of the species known up to 1970. Two accessions to the flora could hardly persist under such conditions. The of the island were made, viz. Alchemilla glomenr- remaining third might well survive glacial condi- lans and Saxifagafoliolosa, the latter discovered tions. e.g. Poa alpina, Fesruca vivipara, by Brattbakk (1984). Deschampsia alpina, Phippsia spp., most Saxfraga In my opinion, we can expect discovery of few spp. except S. aizoides and S. hirculus, Cerastium vascular species on Bjamoya in the future. Then arcticum. C. regelii. Ranunculus sulphureus, Papa- the question arises of the ecological potential of ver dahlianum, Draba alpina. and D. norvegica. Bjarnaya to support species which are absent This is an assemblage of ubiquitous Arctic species, there, but occur in Spitsbergen and North Fenno- some of which are known to perform well also scandia. Examples are Ranunculus glacialis and in the extreme polar climate of Sevemaja Semlja Cardamine bellidfolia. both missing on Bjprm- (Korotkevitsj 1958; Semenov 1970), and Semlja aya, but thriving well in North Norway, at Jan Frantsa Josifa (Hanssen & Lid 1932; Tolmatsjev Mayen , and on Spitsbergen, R. glacialis only on & Sjukhtina 1974; Aleksandrova 1977). the southwest coast of Spitsbergen. To analyse this problem, I have listed 144 Ecological 'completeness' of the Bjomo.va well-documented species of the Svalbard flora in vegetation nine ecological groups (Table 10). Three levels An objective for the renewed investigation of of thermal demands are defined, as seen from I07

Table 10. Principal vascular species of Svalbard, tabulated as to hydrological and thermal preference. * Occurring on Bjarnaya. (-:) fraction of species group present on Bjarnnya.

Water-logged Intermediately Well-drained habitats habitats drained habitats

Low thermal demands

*Juncus biglumis Luzula arctica Ranunculus nivalis Poa abbreviata *Deschampsia alpina *L.arcuatas. lat. *R.sulphureus Festuca hyperborea Pleuropogon sabinei *Alopecurusalpinus Braya purpurascens Care.x nardina *Phippsia algida *Poa alpina var. *Cochlearia Minuartia rubella *P. concinna vivipara groenlandica *Papaverdahlianum Carex ursina P. arctica Cardamine bellidi- Draba nivalis *RanuncuIus hyper- *P.pratensis ssp. folia *D. norvegica boreus alpigena *Draba alpina *Saxifraga caespitosa *Saxifragafoliolosa *Festuca vivipara - D. bellii *S.nivalis S. svalbardensis different biotypes D. lactea *S.oppositifolia Colpodium vahlianum Potentilla h.vparctica Firccinellia angustata D. micropetala P. pulchella *Carex lachenalii D. oxycarpa *Oxyria digyna *Saxifraga cernua *Sagina intermedia S.flagellaris Stellaria crassipes S. hyperborea *Cerastium arcticum *S.rivularis *C.renelii *S.tenuis 16 (3,)

Intermediate thermal demands

*Equisetum arvense Huperzia selago *Arabis alpina *Equisetum scir- Dryas octopetala *E. variegatum Triseium spicatum *Chrysosplenium poides Cassiope retragona *Arctophilafulva *Puccinelliaphryga- tetrandrum Hierochloe alpina Mertensia maritima *Calcmagrostis neg- nodes *Saxifraga aizoides Poa glauca Pedicularis lanata lecta Carexglareosa S. hieraciifolia P. hartzii ssp. dasyantha Deschampsia C. misandra *S. hirculus Festuca baffinensis Polemonium boreale brevifolia *Salk herbacea Cassiope hypnoides *F. rubra Campanula unijlora Dupontiajkheri *S.polaris Pedicularis hirsuta Carex rupestris Erigeron unijlorus *D.psilosantha 5.reticulata Erigeron humilis Arenaria Arnica alpina Eriophorum *Polygonurn Taraxacum arcticum pseudofrigida scheuchzeri viviparum T. brachyceras Honckenya peploides E. rriste Minuartia bijlora *T.cymbfolium *Silene acaulis Carexparallela M. rossii S.furcata *C.subspathacea ‘Cerastium cerastoi- Draba arctica *Koenigia islandica des D. daurica *Cardamine nymanii *Stellaria humifusa D.fladnizensis Petasitesfrigidus Silene wahlbergella *Sedum rosea Ranunculus spits- “Ranunculuspyg- Potentilla nivea bergensis maeus Eutrema edwardsii

(continued) I08

Table 10. (continued)

High thermal demands

Junucus castaneus Tvjieldia pusilla Cvstopteris.fragilis ssp. dickieana J. triglumis Juncus arcticus Woodsia glabella Luzula wahlenbergii CartB.u maririma Ranunculus auricomus s. lat. Arctagrostis latfolia Salir glauca Empetrum herrnaphroditum Kobresia simpli- Betula nana Gentianella renella ciuscula Minuartia stricta Campanula rotundfolia s. lat. Cares amhlyo- Pnrentilla cranm rhj*ncha 'Alchemilla glome- C.saxatilis nilans Ranunculus Rubus ehamaemorus lapponicus VaccinI um micro- R.pallasii plirllum *H@puris vulgans

I (lo'

30 9 (a' (-)42

Not included in this survey are: Botnchium lunaria. Aiccinellia svalbardensis. Colpodium vacillans, Carer capillaris. C. aquarilis ssp. stans. Sagina caespirosa. Arenana humifusa. Sibbaldia protumbens. and Eupkrasia~fiigirla- because of very restricted or local occurrences, or ecological performance not well known on Spitsbergen. altitude limits in Scandinavia and Svalbard and of several chionophobous species with low tem- the performance of the species on the North and perature demands, i.a. Carex nardina, Minuartia East coasts of Spitsbergen. Three classes of rubella, Braya purpurascens, and Draba nivalis. hydrological preference may usually be estima- We also miss some chionophilous species: ted at the plants' habitats. It may be noted that a Ranunculus glacialis, R. nivalis, Cardamine seasonal hydrophile, e.g. Ranunculus nivalis. is bellidifolia, and Cassiope hypnoides. not ranked together with tundra swamp plants, Rather surprising is the lack of Papaver dahlia- e.g. R. pallasii. However, there are transitory num on the high ground of Bjamaya, such as the cases, and my classification is tentative. summits of Miseryfiellet. On Spitsbergen it A simple contingency test based on Table 10 ascends to 1 I00 m above sea level (Sunding 1966) proves an over-representantion in the vascular and is among the most cold-tolerant Arctic spe- flora of Bjamoya of species adapted to cold and cies. The limitations of Papaveron Bjamoya may water-logged conditions, compared to Svalbard be related to the humid climate and snow- as a whole. Conversely, there is a certain under- influenced vegetation, detrimental to Scapiflora representation of species demanding higher papavers, according to my observations in Nor- summer temperatures, especially of lignified way and Svalbard. species. Absent from Bjamaya but growing in A closer study of the bryo- and lichenflora of Scandinavia and on Spitsbergen are Salix glauca Bjornoya will probably reveal similar patterns of (verq' rare on Spitsbergen, cf. Schweitzer 1966), eco-geographical significance. Several arctic- Betula nana, Dnas octopetala, Cassiope tetra- alpine species are absent, e.g. Alectoria ochroleu- gona, and Empetrum herrnaphroditum. ca. 7hamnolia verrnicularis,and Cetraria cuculla- Even ubiquitous Arctic species are missing, fa. The absence of Usnea sphacelata is difficult e.g. Carex rupestris and Trisetum spicatum. to explain in climatic terms because this lichen Furthermore, we note the absence from Bjamaya is so frequent at Jan Mayen and in Iceland 109

(Lynge 1926, 1941). The lack of a suitable stony southern provenance because of its low chromo- substratum on the upper slopes and on the sum- some number 2n = 28, as against 2n = 49 on mits of Miseryfjellet may be a contributing cause. Spitsbergen (Engelskjan 1979). But the histories Chionophilous or oceanic species dominate of these two species on Bjamaya may be differ- the cryptogamic vegetation, e.g. Cetraria delisei, ent. Recalling the glacial history of the Barents Cladina mitis, Brachythecium glaciale, B. salebro- Sea (Salvigsen & Nydal 1981), the local F. vivi- sum. and Rhacomitrium lanuginosum.This shows para biotype could be a vestige of a pre-Vistulan the close relationship to the coastal vegetation flora. This possibility was also pointed out by section of Spitsbergen, named the Cladonia HadaE (194 1,1960) and Ranning ( 1959) for other mitis-zone by Eurola (1968). species, including the lichen- and bryoflora. Very little can be concluded about the disper- To avoid simplistic conclusions, more studies sal adaptations of the Bjamaya vascular plants. are needed of all aspects of the flora of Bjamaya, Only about 10% of the species could be con- and attention should be paid to advances in sidered as anemochores by definition. The three Quaternary research in the North Atlantic- arctic-littoral species may well disperse by sea Barents Sea region. currents. 15 YO reproduce mainly by vegetative propagules ; among these, Saxifraga foliolosa is Acknowledgements. - Among several persons who have contributed to the botanical exploration of Bjarn- most restricted on the island. A generalized dis- 0ya in cooperation with the present author, I extend persal (Berg 1983) must be regarded as the main special thanks to Ola Skifte of Tromse Museum who dispersal principle of the Bjamaya flora. Several organized the 1983 expedition. animal and geophysical vectors are involved. The following institutes and persons have contributed to the progress of the work, for which I am in- Also some man-introduced species perform well, debted: The Bjemeya meteorological station crew and e.g. non-viviparous Poa alpina and a biotype of Olav B. Syse, Vervarslinga for Nord-Norge, Brynhild P.pratensis ssp. alpigena. Merkved and the office and photographic departments The absence of several widespread, arctic- at Tromse Museum, Oivind Grothe at the Norwegian alpine species from Bjarnaya may lead to Meteorological Institute, Botanical Garden and Muse- um, University of Oslo, for providing working facilities, speculation whether the biome is ecologically and Herbjem Lysnes at the State Research Station, ‘complete’ or not, cf. Bertram & Lack (1938) who Holt, Tromse, as well as Olav M. Skulberg, the Nor- discussed the faunal aspects. Obviously, several wegian Institute for Water Research, for soil analyses. species might succeed if introduced to Bjamaya, Renit Jacquet, Graphic Office at the Museums of Natural History, University of Oslo, has drawn the e.g. the chionophilous Ranunculus glacialis and base map of Bjemraya. Cardamine bellidifolia, both of which are I am also grateful to the following colleagues for frequent on Jan Mayen (Lid 1964). The air and assistance with determination of cryptogams: vegetation temperatures and the hygric condi- Bodil Lange, University of Copenhagen (Sphag- tions would promote snow-bed plants and num), Jon Holtan-Hartwig (Peftigeru), Einar Timdal (Lecidea, Toninia, etc.), both at the Botanical Garden tundra swamp hygrophytes to settle down as and Museum, University of Oslo. Sigbjern Dunfjeld, members of the Bjamaya flora. Hattfjelldal, acted as an able field assistant and aided Snow-avoiding species grow on wind-exposed in preserving the material. Special thanks are extended crests and often tend to be stunted - Papaver to Liv Hillestad for assistance with the draft manu- script. dahlianum is a good example from Bjamaya. The Norwegian Research Council for Science and The steady fog and the wind chill may also con- the Humanities covered the travel expenses and tribute to nanism among the vascular plants. granted me a research scholarship (D. 70.49.085) for Places with favourable exposure and wind which I extend my thanks. protection are often affected by bird colonies which cause development of Cochleariaor Festuca References rubra communities with a few other species. Some species which are now confined to shal- Aleksandrova, V. D. 1977: The structure of plant associations of the polar deserts on the Aleksandra Land low ponds could have been introduced by birds (Franz Josef Land) (Russ.). Pp. 26-36 in Problemy during the Holocene, e.g. Hippuris vulgaris. The ekologii, geobotaniki, botanitsjeskojgeografii ifloristi- local biotype of Festuca vivipara may be of ki. Leningrad. 110

Andresen. L. 1979: Monthly and annual frequencies Gebieten von lsfjorden und Homsund in Westspitz- of concurrent wind forces and wind directions in bergen. Aquilo. Ser. Bot. 7.1 - 56. Northern Norway and the Arctic for the period Fleetwood, A., Pejler, B., Einsle, U. & Amemo, R. I96 I - 75. Climatological summaries,for Noma!.. Det 1974: Stratigraphical and biological investigations norske meteorologiske institutt. 202 pp. of some Bj~rnoyalakes. Norsk Pvlarinstirurr hbvk Arnell. S. 1956: Hepaticae. In lllustrared moss.flvra of’ 1972,63 -7 1. Fennoscandia 1. The Botanical Society of Lund. 308 French, H. M. 1971: Ice cored mounds and patterned PP. ground, southern Banks Island, Western Canadian Berg, R. Y. 1983: Plant distribution as seen from plant Arctic. Gevgr. Ann. 53 A (I), 32-38. dispersal : General principles and basic modes of Gjessing, Y. T. & 0vsteda1, D. 0. 1976: Energy budget plant dispersal. Sonderbd. natumiss. Vereins Ham- and ecology of two vegetation types in Svalbard. burg 7. 13-36. Asrarre 8.83-92. Berggren, S. 1875: Musci et Hepaticae Spetsbergenses. Hadat, E. 1941: Et bidrag ti1 historien om Bjornoyas Kgl. Svenska Verensk.-Akad. Hand. 13(7). 103 pp. flora. Naturen 65.141 - 155. Bertram, C. G. L. & Lack, D. 1938: Notes on the animal Hadat, E. 1960: The history of the flora of Spitsbergen ecology of Bear Island. J. Anim. Ecol. 7.27-52. and Bear Island and the age of some arctic plant Bliithgen, J. & Weischet, W. 1980: Allgemeine Klima- species. Preslia 32,225-253. geographie. W. de Gruyter. 887 pp. Hambrey, M. J. & Swett, K. 1980: A lensoid, moss- Brattbakk. I. I98 I : Braggerhalvoya Svalbard. Pegeta- covered ‘Needle Ice’ body, St. Jonsfjorden, Spits- sjonskart /:I0 000, Kartblad I- VIII. DKNVS bergen. Norsk Polarinstitutr Arbok 1979.71 -76. Museet, Bot. Avd.. Trondheim. Hanssen, 0. & Holmboe, J. 1915: The vascular plants Brattbakk, I. 1983: Lavrik morenevegetasjon pi of Bear Island. Nvr Mag. Natun-.62,210-235. Broggerhalvoya. Svalbard. DKNVS Museer. Rapport Hanssen. 0. & Lid, J. 1932: Flowering plants of Franz Bot. Ser. 19831 71.I I - 16. Josef Land. Nvrsk Polarinstiiutr Skn$er 39.42 pp. Brattbakk. I. 1984: Bjornoya, Svalbard, et potensielt Horn, G. & Orvin, A. K. 1928: Geology of Bear Island. reinbeiteomride. Report to Bjornoen A, S.20 pp. Norrk Polarinstitutt Skrifier 15. I52 pp.. 9 pl.. I geol. Dahl, E. 1957: Rondane. Mountain vegetation in South map. Noway and its relations to the environment. Norske Hyvarinen, H. 1968: Late-Quatemary sediments on Vidensk.-Akad.Skr. 1. Mat.-Natun. kl. 1956 131. 374 Bjornoya. Geogr. Ann. 50A, 235-245. pp.. 1 map. Hyiirinen, H. 1972: Pollen-analytic evidence for Dahl. 0. 1934: Floraen i Finnmark fylke. Nyt Mag. Flandrian climatic change in Svalbard. Pp. 225-237 Nalun. 69.430 pp., 17 pl. in Vasari, Y. et al. (eds.): Climatic changes in Arctic Datkiewicz, L. 1967: The distribution of periglacial areas during the last ten thousand years. Acta Univ. phenomena in NW-Sorkapp, Spitsbergen. Biuletw~ Ouluensis Ser. A 3 (I). Periglacjalny 16.37-83. Jalas, J. 1955: Rhacomirrium lanuginosum (Hedw.) Eide. 0. 1945: On the temperature difference between Brid. als Klimaindikator in Ostfennoskandien. Arch. mountain peak and free atmosphere at the same Soc. Bot. Vanamo 9. Suppl.,73-88. level. 11. Gaustatoppen-Kjeller. Meteorol. .4nn. 2. Keilhau, B. M. 1831: Reise i 0st- vg Vesr-Finmarken

183 - 206. samt ti1 Beeren-Eiland vg Spitsbergen. i Aarene 1827 Elvebakk. A. 1979: Planteso.riolvgi og :fenvlogi i er ark- og 1828. Christiania. 247 pp, 1 map, 3 pl. (Facsimile, tisk omrdde: Sruphallet. Braggerhalvct~a.Svalbard. Borsums forlag, Oslo 1973). Cand. real. thesis. Univ. Trondheim. 233 pp. Korotkevitsj, E. S. 1958: The vegetation of Severnaya Elvebakk, A. 1982 : Geological preferences among Zemlya (Russ.). Bvt. Zhurnal43(5).644-663. Svalbard plants. Inter-Nod 16. I 1-31. Kukla, G. J. 1972: Insolation and Glacials. Bvreas I, Elverhoi. A. & Solheim, A. 1983: The Barents Sea ice 63 -96.

sheet - a sedimentological discussion. Polar Lamb, H. H. 1972: Atmospheric circulation and Research 1.23 -42. climate in the Arctic since last Ice Age. Pp. 455-495 Engelskjon. T. 1967; Cyto-embryological studies in in Vasari, Y. et al. (eds.): Climatic change in Arctic Arctic-Alpine Taraxaca. 1. The Bear Island Taraxa- areas during the last ten thousand years. Acta Univ. cum. NwMag. Bot. /4, 125-137. Ouluensis Ser. A. 3 (I). Engelskjon, T. 1979: Chromosome numbers in vascu- Lid, J. 1964: The flora of Jan Mayen. Norsk Polar- lar plants from Norway. including Svalbard. Opera instirutr Sknyter 130. I07 pp. Bot. 52.38 pp. Liestol, 0. 1972: Sub-marine moraines off the west Engelskjon. T., Kramer, K. & Schweitzer, H.-J. 1972: coast of Spitsbergen. Nvrsk Polarinstitutt Arbvk 1970. Zur Flora des Van Mijenfjorden-Gebietes (Spits- 165-168. bergen) und Hopens. Norsk Polarinsritutt Arbok Lundqvist, J. 1962: Earth and ice mounds: A termino- I’)70.191--198. logical discussion. Pp. 203-2115 in Pewe, T. L. (ed.): Engelskjon, T. & Schweitzer. H.-J. 1970: Studies on the The periglacial environment. McGill-Queen’s flora of Bear Island (Bjornaya) I. Vascular plants. University Press. Astarte 3il). 1-36, Lynge, B. 1926: Lichens from Bear Island (Bjorn0ya). Eurola. S. 1968: Uber die Fjeldheidevegetation in den Nvrsk Pvlarinsritutt Skrifter 9.78 pp., 2 pl. 111

Lynge, B. 1941 : On Neuropogon sulphureus (Konig) Seppala, M. 1976: Season thawing of a palsa at Enonte- Elenk., a bipolar lichen. Sknifrer Norske Vidensk.- kio, Finnish Lapland, in 1974. Publ. Inst. Geogr. Akad. I. Mat.-Naturv. kl. 1940(10).35 pp. Univ. Turkuensis 79,I-24. LAg, J. 1980: Vertikal myr pa Svalbard. Forskningsnytt Siedlecka, A. 1975: The petrology of some Carbonife- 25 (I ), 23 -25. rous and Permian rocks from Bjernaya, Svalbard. Miller, G. H. 1982: Quaternary depositional episodes, Norsk Polarinstitutt Arbok 1973.53-72,4 pl. western Spitsbergen, Norway: aminostratigraphy Siple, P. A. & Passel, C. F. 1945: Measurements of dry and glacial history. Arct. Alp. Res. 14,321 -340. atmospheric cooling in subfreezing temperatures. Miiller, M. J. 1982: Selected climatic data for a global Proc. Americ. Philos. SOC.89, 177- 199. set of standard stations for vegetation science. W. Steffensen, E. 1969: The climate and its recent varia- Junk, Haag. 306 pp. tions at the Norwegian Arctic stations. Meteorol. Norderhaug, M., Bruun, E. & Uleberg Mollen, G. 1977: Ann. 5(8).347 pp. Barentshavets sjefuglressurser. Norsk Polarinstitutt Steffensen, E. 1982: The climate at Norwegian Arctic Meddelelser 104. 1 19 pp. stations. Klima 5.44 pp. Norsk Polarinstitutt 1944: The survey of Bjernoya Steindorsson, S. 1945: Studies on the vegetation of the (Bear Island) 1922- 193 I. Norsk Polarinsritutt central highland of Iceland. The Botany of Iceland 3 Sknifrer 86.82 pp., 2 pl., 1 map. (41,345-547. Nyholm, E. 1954- 1969: Musci. In Illustrated moss Summerhayes, V. S. & Elton, C. L. 1923: Contributions .flora of Fennoscandia. 11. The Botanical Society of to the ecology of Spitsbergen and Bear Island. J. Lund. 799 pp. Ecol. 11, 2 14-286. Olausson, E. 1972: The role of the Arctic ocean during Sunding, P. 1966: Plantefunn fra Vestspitsbergen cool climatic periods. Pp. 409-422 in Vasari, Y. et sommeren 1964. Norsk Polarinstitutt Arbok 1964. al. (eds.): Climatic change in Arctic areas during the 149- 154. last ten thousand years. Aria Univ. Ouluensis Ser. A Svensson, H. 1964: Structural observations in the 3f I). minerogenic core of a pals. Svensk Geogr. Arsbok Orheim, 0. I98 I : The glaciers of Bouveteya. Norsk 1964,138-142. Polarinstitutt Sknifter 17S,79-84. Tolmatsjev, A. I. & Sjukhtina, G. G. 1974: New data Russell, R. S. & Wellington, P. S. 1940: Physiological on the flora of Franz Josef Land (Russ.) Bot. Zhurnal and ecological studies on an Arctic vegetation. 1. The 59(2).275-279. vegetation of Jan Mayen Island. J. Ecol. 28, Triloff, E. G. 1944: Verbreitung und Okologie der 153-179. Gefassplanzen im Gebiete des Hornsundes. ein Bei- Ronning, 0. I. 1959: The vascular flora of Bear Island. trag zur Vegetationskunde Spitsbergens. Bot. Jahr- Acta Borealia Ser. A. 53 pp. bucher 73.259-360. Salvigsen, 0. I98 I : Radiocarbon dated raised beaches Vinje, T. E. 1962: The cooling power in Antarctica. in Kong Karls Land, Svalbard, and their consequen- Norsk Polarinstitutt Arbok 1961.7-22. ces for the glacial history of the Barents Sea area. Vorren, K.-D. 1972: Stratigraphical investigations of a Geogr. Ann. 64 A (3-4). 283-291. palsa bog in Northern Norway. Astarte 5.39-71. Salvigsen, 0. & Nydal, R. 1981: The Weichselian Walker, F. J. 1985: The lichen genus Usnea subgenus glaciation in Svalbard before 15,000 B. P. Boreas 10, Neuropogon. Bull. British Museum (Natural Histon) 433-446. Bof.Ser. I3 (I). I30 pp. Santesson, R. 1984: The lichens ofSweden and Norway. Walton, D. W. H. 1984: The terrestrial environment. Swedish Museum of Natural History, Stockholm & Pp. 1-60 in Laws, R. M. (ed.): Antarctic Ecology I. Uppsala. 333 pp. Academic Press, London. Schweitzer, H.-J. 1966: Beitrage zur Flora Svalbards. Worsley, D. & Edwards, M. B. 1976: The upper Palaeo- Norsk Polarinstitutt Arbok 1964,139- 148. zoic succession of Bj~rrneya. Norsk Polarinstiturt Semenov, 1. V. 1970: Severnaja Semlja. Pp. 391 -422 Arbok 1974. 17-34. in Sovjetskaja Arktika. lzdatelstva Nauka, Moskva.

Maps 1-54 114

2. Equiseticm scirpoides Rich.

pLu*

3. Eqirisenim \'nriegnrltrn Schlei ch. 4. Jiiticiis bigliimis L.

My4 % 115

5. Luzula arcuata (Wg.) SW. SSP. arcuata 6. Alopecurus alpinur Sm.

7. Calamagrostis neglecta (Ehrh.) G.M.S. 8. Arcfophila fulva (Trin.) Rupr. e-J+

1 I7

16. Fesrircn rirbrrr L. I I8 119

21. Sn1i.r polnris Wg. 22. Salix reticidom L.

24. O.ryri(i digyna (L.) Hill 120

P 121

32. Rmiiiticiiliis hyperhoreirs Rottb * zz I 123

37. C'ardarnitw tiyrnariii Gand. 38. Cochlearin groenlundica L.

140 20 20 211 I - ca.20(1 w-

39. Driibn cilpinn L. 40. Drnhn riorregicn Gunn.

Stations at various altitudin levels ( m a.s.1.):

oI3aoIoI

00000lLl 0000 c 0 -.N"=33- ,', 0 5 I - 2 70 124 I25

45. Saxifrugu foliolosa R. Br. 46. Suxifragu hirrirlirs L.

PAub t-hew

47. .s(/.\;/;7/y(/ //il O/i\ L 48. Saxifraga oppositifolia L. 126

49. Sa.rifrrign rii~ii1ari.sL. 50. Saxifraga teniris (Wg.) H.Srn.

t-b%

51. CIirvsospleti~irr?~terrandritrn (Lund) Th. Fries 127

53. Hippuris vulgaris L. 54. Taraxacum cymbifolium H.Lindb.