Flora and Vegetation of Greenland: Encyclopedia Arctica 6: Plant Sciences (Regional)

Author Stefansson, Vilhjalmur, 1879-1962

Flora and Vegetation of Greenland

(EA-PS. Tyge W. Böcher)

FLORA AND VEGETATION OF GREENLAND

CONTENTS

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Page
Introduction 1
The Flora 2
List of Vascular Plants 3
Climatically Conditioned Distributions 14
Edaphically Conditioned Distributions 16
Historically Conditioned Distributions 16
Distribution of Western and Eastern Species 18
Regions of Vegetation 19
Plant Communities 20 ^ -a ^
Scrub and Wood 20 ^ -a ^
Dwarf Shrub 21
Xerophilous Grasses and Grasslike Plants 23
Xerophilous Pteridophytes and Herbaceous Dicotyledons 25
Nondwarfish Hygrophytic and Mesophytic Herbs 26
Dwarfish Herbs and Woody Plants 27
Helophytes 30
Cryophilous Algae 32
Marine Algae 32
Hydrophytes 31
Bibliography 35

EA-Plant Sciences (Tyge W. Böcher)

FLORA AND VEGETATION OF GREENLAND
INTRODUCTION
The investigations of the flora of Greenland started in the eighteenth century, but systematically planned collections, particularly of vascular plants, were only begun after the beginning of the nineteenth century. The most important collections in the past century were made between 1828 and 1836 by Jens Vahl, the Danish botanist, who traveled the southern part of the east coast and the greater part of the west coast. His collections constituted the nucleus of the first comprehensive floral work, Johan Lange’s Conspectus florae Groenlandicae (17), which is still of great scientific interest. Later surveys of the flora have been written by Ostenfeld (20) on the whole of Greenland, Porsild (25) on the central part of West Greenland, Seidenfaden and Sorensen (32) on East Greenland, Polunin (22) on southwestern Greenland, and Böcher (1) on southern East Greenland.
Investigations of the flora particularly from an ecological point of view (plant communities, structure and biology of arctic flowering plants) were started in the latter part of the nineteenth century. The pioneer ^ in ^ this line of investigation was E. Warming, whose book Om Grønlands Vegetation [ On the Vegetation of Greenland ] (38) was epoch-making. The most important descrip– tions of the vegetation published during the Warming epoch deal with southern Greenland (30), Scoresby Sound (10; 11), and the Angmagssalik region (15).

EA-PS. Böcher: Flora and Vegetation of Greenland

From comparatively recent times we have, moreover, comprehensive descriptions of the vegetation of the coast south of Scoresby Sound (5), Godthaabs fjord (37), on account of the ecological conditions in the region of Søndre Strømfjord (2), and, in regard to northern East Greenland, a survey of the vegetation between latitude 74°30′ and 79° N.
Cytogenetic investigations of the flora of Greenland were started in 1927 with Hagerup’s investigations of Empetrum hermaphroditum . Since then, several papers have dealt with chromosome numbers determined on material from Greenland, a recent example being that of Böcher and Larsen (6).
THE FLORA
The approximate number of species of the Greenland flora is given in Table I. In 1870 there were 360 known species of vascular plants; within the next 56 years only 30 additional species were identified. By 1949 there were 456 species known, composed of: 31 pteridophytes, 1 gymnosperm, 148 monocotyle– dons, 173 Apetalae and Polypetalae, and 103 Sympetalae (Gamopetalae). As our knowledge particularly of fungi and freshwater algae is rather defective, the

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Table I. Greenland Flora .
Group Number of species Group Number of species
Vascular plants 456 Marine algae 200
Bryophytes 600 Freshwater algae 400
Lichens 400 Diatoms and dinoflagellates 650
Fungi 1,000

EA-PS. Böcher: Flora and Vegetation of Greenland

figures for these groups are too low. We may confidently expect an aggregate number of more than 4,000 plant species in Greenland.
A list of all the vascular plant species known to the author as occurring in Greenland is given below; for each species, information is given as to its distribution in Greenland as far as this is yet known. It has been attempted to divide the country into floral “provinces,” the delimitation of which is indicated in Figure 1. The boundary lines between the provinces have been drawn largely in accordance with the boundaries of the known areas of a great many species. Special stress has been laid on boundaries of species considered to be good indicators of climate, e.g., Alchemilla alpina , Carex supina , Cas–siope tetragona, and Ranunculus sulphureus .
List of Vascular Plants
The following is a key to the abbreviations used in the listing of vascular plants and throughout this article:

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N North Greenland CE central continental part of East Greenland
NW northern West Greenland
NW(T) Thule District SE southern East Greenland
NW(U) Upernivik District n northern part
CW central continental part of west Greenland s southern part
SW southern West Greenland intr introduced by man
S South Greenland alp alpine
NE northern West Greenland isol isolated
r rare
EQUISETACEAE (5): Equisetum arvense , N to S; E. scirpoides , CW, SW, S;
Fig 1.

EA-PS. Böcher: Flora and Vegetation of Greenland

E. sylvaticum , SW, S; E. trachyodon , S; E. variegatum , N to S (r in SE).
LYCOPODIACEA (5): Lycopodium alpinum , NW(U), SW, S, Se, CE (r in s); L. annotinum , NW(U), CW, SW, S, SE, CE (r); L. clavatum , SW (r), S; L. complanatum , SW, S; L. selago , N to S.
SELAGINELLACEAE (2): Selaginella rupestris , S (5); S. selaginoides, SW, S, SE.
ISOETACEAE (2): Isöetes echinospora , CW (r), SW, S; I. lacustris , S (r).
OPHILOGLOSSACEAE (4): Botrychium boreale , SW (s), S, SE (s); B. lanceolatum , SW, S, SE: B. lunaria , NW(U) (s), CW (isol in n), SW, S, SE, CE (isol in s), NE (r in s); B. tenebrosum , S.
POLYPODIACEAE (13): Asplenium viride , SW, SE; Athyrium alpestre , S, SE; Cystopteris fragilis , N to S; C. montana , S; Dryopteris austriaca , SW, S; D. filix-mas , SW, S; D. fragrans , NW, CW, SW (r in n), CE (isol in s); D. linnaeana , SW, S, SE; D. phegopteris , SW, S, SE; Polystichum lonchitis , SW, S, SE; Woodsia alpina (r), NW(U), CW, SW, S, SE, NE (s); W. glabella , N to S (r in S); W. ilvensis , NW(U), CW, SW, S, SE, NE (r), CE (r in s).
CUPRESSACEAE (1): Juniperus communis , CW, SW, S, SE.
SPARGANIACEAE (2): Sparganium antustifolium , CW (isol at lat. 68°40′ N.), SW, S, SE (at lat. 65°37′ N.); S. hyperboreum , CW (s), SW, S.
POTAMOGETONACEAE (6): Potamogeton alpinus subsp. tenuifolius , CW (s), SW (s), S; P. filiformis , CW, SW, S, SE (r), NE (r), CE (r); P. gramineus , CW (s), SW (s), S; P. natans , S; P. pusillus subsp. groenlandicus , CW, SW (r), S (r); Zostera marina , SW (r).
JUNCAGINACEAE (1): Triglochin palustre , NW(U) (s, r), CW, SW, S, SE (r), CE (r).
GRAMINEAE (59): Agropyron (see Roegneria ); Agrostis borealis , CW, SW, S, SE, CE (r); A. canina , CE (r in s), SW (s), S, SE; A. gigantea , S; A. stolo– nifera , SW (r in s), S; A. tenuis , intr to S; Alopecurus aequalis ,

EA-PS. Böcher: Flora and Vegetation of Greenland

NW (r in s), CW (r), SW, S, SE (r); A. alpinus , N, NW, CW (n), SW (r), NE, CE (n); A. pratensis , intr to S; Anthoxanthum odoratum (diploid type), S, SE (s); Arctagrostis latifolia , N, NW, CW (n), CE (n), NE; Arctophila fulva , SW (r); Calamagrostis hyperborea , SW, S, SE; C. langs–dorfii , CW (s), SW, S, SE (s); C. lapponica , CW; C. neglecta , NW(U), CW, SW, S, SE, NE, CE; C. purpurascens , NW, CW, SW (n), S ( [: ]^ r ^ ), NE, CE, SE (r); Catabrosa aquatic , SW (r), S (r); Danthonia spicata , S (r); Deschampsia alpina , SW, S, SE; D. brevifolia , NW(T), N, NE, CE; D. brevifolia subsp. pumila , NW(T), NW(U), Disko, NE; D. flexuosa , CW (single station at lat. 67° N.), SW, S, SE; Dupontia fisheri , NW(U), CW (n), NE to CE (basalt region); Elymus mollis , NW(U) (s), CW, SW, S, Se (s); Festuca brachy–phylla , N to S; F. rubra , NW, SW, CW, S, SE, NE, CE; F. vivipara , N to S (r or alp in S); F. vivipara subsp. hirsuta , SW, S, SE; Hierochloe alpina , N to S; H. odorata , S; Nardus stricta , S, SE (s); Phippsia algida , N to S (r in S and s of CW, SW, Se); Phleum commutatum , SW, S, SE; Pleuropogon sabinii , N, NW(T), NE, CE (r); Poa abbreviata , N, NW, CW (n), NE, CE; P. alpina , NW(U), CW, SW, S, SE, NE, CE; P. alpina subsp. vivipara, NE, SE (n); P. annua , intr to SW, S, Se; P. arctica , N, NW, CW (alp in s), SW (n), S (?), SE (r in s), CE, NE; P. flexuosa , S (alp); P. glauca , N to S; P. hartzii , NW(U) (r in s), CW (r), CE (r), NE (r); P. nemoralis , SW (s), S; P. pratensis (incl. P. alpigena and P. irrigata ), all districts except N, NW(T), and NE (n); Puccinellia andersonii , NW, N, NE; R.^ P. ^ angus–tata , NW, CW (n), N, NE, CE; P. coarctata , NW(U), CW, SW, S, SE, NE (r in s); P. deschampsioides , CW, SW (r in n), NW(U) (r in s); P. groenlandica , NW(U), CW, SW (n); P. langeana , SW (n), NW(U) (s); P. maritima , SW (s), S; P. Phryganodes , N to S; P. porsildii , isol on Disko; P. rosenkrantzii ,

EA-PS. Böcher: Flora and Vegetation of Greenland

isol in NW(U); P. vahliana , NW, CW (n), N, NE, CE; Roegneria borealis var. hyperar [: ]^ c ^ tica , NW(U), CW (n), CE (r), NE (r); R. doniana var. virescens , S; R. violacea , CW (r), S; Trisetum spicatum , N to S; Vahlodea atropurpurea , SW, S, SE (s).
CYPERACEAE (51): Carex aquatilis var. stans , N, NW, SW (n), CW, NE, CE (n); C. arctogena , NW(U), CW, SW, S, SE (r), CE (r); C. atrata , SW (s), S, SE; C. atrofusca , NW(T), NW(U) (r), CW (n), CE, NE; C. bicolor , NW(U), CW, SW (r), S (r), CE (r); C. bigelowii , all districts except N; C. brunnes–cens , SW, S, SE; C. buxbaumii , S (r); C. canescens , NW(U), CW (s), SW, S, SE (s); C. capillaris (s.l.), all districts except N; C. capillaris var. porsildiana , SW, S; C. capitata , isol at lat. 67° N. in CW; C. deflexa , SW, S, SE; C. glacialis , NW(U), CW, SW, S (r), SE (r in s), CE, NE; C. glareosa , all districts except N; C. goodenowii , S; C. gynocrates , NW(U), SW (r), CW, S; C. halleri , NW(U), CW, SW, S, SE (r in central part), CE, NE (r); C. holostoma , SW (n), CW; C. lachenalii , all districts except N, NW(T); C. lyngbei , S (r); C. mackenziei , SW (s), S (r); C. macloviana , CW, SW, S, Se, CE (s, r); C. magellanica , SW (r), S; C. maritima , all districts except N and SE; C. microglochin , CW, SW (r), S, SE, CE; C. misandra , N, NW, CW, SW (r in n), CE, NE; C. nardina , N to S; C. panicea , S; C. parallela, NE, CE; C. praticola , CW (r in s), SW, S; C. pseudola–gopina , NW, NE, CE; C. rariflora , NW(U), SW, CW, S, SE (r in n), CE, NE; C. rostrata , S; C. rotundata , NW(U), CW, SW, S, SE (The two species, C. rotundata and C. saxatilis , can hardly be kept separate); C. rufina , SW, S, SE, CE (single station in s); C. rupestris , NW, CW, SW (n), NE, CE, SE (r in n); C. salina , S; C. saxatilis , NW, CW, SW (n), CE, Ne; C. scirpoidea , all districts except N, NW(T); C. sparsiflora , isol in NE

EA-PS. Böcher: Flora and Vegetation of Greenland

(lat. 74°10′ N.); C. stylosa , SW (s), S, SE (s); C. subspathacea , NW(U), CW (r), SW, S, SE, NE, CE; C. supina , NW(U) (r), CW, SW (r in n), S (r), SE (r), CE, NE; C. ursina , NW, CW, SW (r in n), CE, NE; C. viridula , S; Elocharis acicularis , Disko, CW, SW (r in n); E. uniglumis , S; Erio–phorum angustifolium , N to S (absent from SE north of lat. 61°37′ N.); E. callithrix , NE, CE; E. scheuchzeri , N to S; Scirpus caespitosus , NW(U) (r), CW (r in s), SW, S, SE; S. pauciflorus , SW (r at lat. 67° N.), SW (r in s), S.
JUNCACEAE (19): Juncus alpinus , CW (r), SW (one station), S; J. arcticus , NW(U), CW, SW, S, CE, NE (r), SE (r); J. balticus , S; J. biglumis , N to S; J. bufonius , CW (s), S, NE (one station at lat. 71° N.); J. castaneus , NW(U), CW, SW, NE, CE, SE (isol at lat. 66°16′ N.); J. filiformis , SW (s), S, SE (s); J. squarrosus , S, SE (s); J. subtilis , CW (one station), SW (r), S (r), SE (r); J. trifidus , NW(U), CW (Disko Bay), SW, S, SE, NE to CE (s); J. triglumis , NW, CW, SW, S, NE, CE, SE (r); Luzula arctica , all districts except S, Se, and SW (s); L. confusa , N to S; L. groen–landica , CW, SW, S; L. multiflora , SW, S; L. multiflora subsp. frigida , NW(U), SW, S, SE, NE to CE (s); L. parviflora , NW (Disko), SW, S, SE (s); L. spicata , all districts except NW(T), N, NE (n); L. wahlenbergii , isol in NE.
LILIACEAE (3): Streptopus amplexifolius , SW (s); S; Tofieldia coccinea , NW, CW, CE, NE; T. pusilla , all districts except N and NW(T).
IRIDACEAE (1): Sisyrinchium montanum , CW (r in s), SW (Godthaab Fjord).
ORCHIDACEAE (5): Corallorhiza trifida , SW, S; Habenaria albida , SW, S, SE; H. Hyperborea , SW, S, SE; Listera cordata , SW, S, SE (s); Orchis rotun–difolia , CW (one station at lat. 67° N.), SW (one station at lat. 64°09′ N.), S (r).

EA-PS. Böcher: Flora and Vegetation of Greenland

SALICACEAE (5): Salix arctica , NW, N, NE, CE; S. arctophila , NW, CW, SW, S; S. glauca incl. S. callicarpaea , all districts except N; S. herbacea , all districts except N; S. uva-ursi , SW, S.
BETULACEAE (4): Alnus crispa , CW (s), SW, S (r); Betula glandulosa , SW, S, B. nana , NW(U), CW, SW, SE, NE, CE; B. pubescens (s.l.), SW (r in s), S.
POLYGONACEAE (7): Koenigia islandica , all districts except N, r in CW and C [: ]^ E ^ ; Oxyria digyna , N to S; Polygonum heterophyllum , intr to CW, SW, S, SE; P. viviparum , N to S; Rumex acetosa , SW, S; R. acetosella , CW, SW, S, SE, NE, CE; R. domesticus , intr to S.
CHENOPODIACEAE (1): Atriplex sp., S (r).
PORTULACACEAE (1): Montia lamprosperma , CW, SW, S, SE.
CARYOPHYLLACEAE (32); Arenaria humifusa , NW, CW (n), SW (Disko), S (r); A. pseudo–frigida , NE, CE; Cerastium alpinum , N to S; C. arvense, isol in CW (at lat. 67°5′ N.); C. caespitosum , SW, S; C. cerastioides , NW(U) (s), CW (r in s), SW, S, SE, NE, CE (r in s); C. regelii , NW (? Hare Island), N, NE; Honck–enya peploides , all districts except N; Melandrium affine , N, NW, CW, SW (n), NE, CE; M. apetalum , N, NW, CW, CE, NE; M. triflorum , N, NW, CW, SW (n), CE, NE; Minuartia biflora , all districts except NW(T) and N; M. groen–landica , SW, S, SE (s); M. rossii , NW(U), N, NE; M. rubella , N to S; M. stricta , NW(U) (s), CW, SW (Disko), CE, NE; Sagina caespitosa , NW(U), CW, SW, S, ?SE; S. intermedia , N to S except inland in CW (s); S. linnaei , SW, S, SE; S. nodosa , S; S. procumbens , S; Silene acaulis , N to S; Stellaria calycantha , SW, S, SE; S. humifusa , all districts except N; S. longipes group: S. ciliatosepala , NW, N, NE, CE (r); S. crassipes , NW, N, NE; S. laeta , CW, NW, N; S. laxmannii , CW (n), NW, N, NE; S. longipes , NW(U), CW, SW, S, ?CE (s); S. monantha , NW(U), CW, SW, S, SE (r); S. media,

EA-PS. Böcher: Flora and Vegetation of Greenland

probably intr NW(U), CW, SW, S, [: ]^ SE ^ ; Viscaria alpina , NW(U), CW, SW, S, SE, CE (s), NE (r and isol).
RANUNCULACEAE (16): Anemone richardsonii , SW (isol between 65° and 67° N. lat.); Coptis trifolia , SW, S, SE; Ranunculus acris , SW, S, SE (isol); R. affinis (incl. R. pedatifidus ), NW, CW, SW (Disko), NE, CE (r); R. auricomus var. glabrata , CE (r), NE (r), SE (n, isol); R. cymbalaria, CW (r), SW (n, r); R. glacialis , NE, SE (except s); R. hyperboreus , N to S; R. lapponicus , NW(U), CW, SW (n); R. nivalis , NW, CW, SW (n, r), CE (r), NE; R. pygmaeus , all districts except N; R. reptans , CW, SW, S, SE (isol at lat. 67°37′ N.); R. sabinii , NW(T), N; R. sulphureus , N, NW, CE (n), CW (n, r), SW (n), NE, CE (r); R. trichophyllus var. eradicatus , all districts except N; Thalic–trum alpinum , NW(U) (s), CW, SW, S, SE, CE (r), NE (r).
PAPAVERACEAE (1): Papaver radicatum , N to S.
CRUCIFERAE (34): Arabis alpina , NW(U), CW (n), SW, S, SE, CE (r in s), NE (s); A. arenicola , NW, CW, SW (r), S (r), NE (r); A. holboellii , CW, SW (r), S (r), SE (r), NE (isol in s), CE (s); Braya ( Torularia ) humilis , CW (Søndre Strømfjord), NE, CE; B. linearis , NW(U) (isol), CW [: ]^ (isol) ^ , CE; B. purpurascens , N, NW, CW (n), CE, NE; B. thorild-wulffii , N, NE (n); Capsella bursa-pastoris , intr to S; Cardamine bellidifolia , N to S (alp and r in S and CW (s)); C. pratensis , all districts except N; Cochlearia officinalis , N to S; Draba alpina , NW(U), CW (n), SW (n and isol on nuna–tak), NE, CE (r); D. aurea , CW, SW, S, SE, CE (s); D. bellii , CW (n), NW, N, NE, CE; D. cinerea , N, NW, CW, CE, NE; D. crassifolia , NW(U), CW (n), SW (n), NE, CE (s), SE (n); D. daurica , NW, CW, SW (n), NE, CE; D. fladnizensis , NE, CE; D. gredinii , NE; D. groenlandica , N, NW, CW, SW (n); D. incana , CW (r), SW, S, SE; D. lactea , N, NW, CW, SW (r), CE, NE;

EA-PS. Böcher: Flora and Vegetation of Greenland

D. lanceolata , CW; D. micropetala (incl. D. oblongata ), NW(T), N, NE, CE (r in n); D. nivalis , all districts except N; D. repens , isol in NE (s); D. rupestris , CW (r), SW, S, SE, NE (s), CE (s); D. subcapitata , N, NW, CW (n), CE, NE; Eutrema edwardsii , N, NW, CW (n), CE, NE; Hali–molobos mollis , NW, CW; Hesperis pallasii , N; Lesquerella arctica , N, NW, CW, CE, NE; Rorippa islandica , CW (isol at lat. 67° N.), SW (intr), S; Subularia aquatica , S, SE.
DROSERACEAE (1): Drose ^ r ^ a rotundifolis, S.
CRASSULACEAE (4): Sedum acre , SE (Angmagssalik District); S. ann ^ u ^ um, SW, S, SE; S. rosea , CW (s, r), SW, S, SE, NE, CE; S. villosum , CW, SW, S, SE.
SAXIFRAGACEAE (17): Chrysosplenium tetrandrum , NE (isol); Parnassia kotze–buei , CW (n, isol), S (r); Saxifraga aizoides , NW (s, r), CW, SW, S, SE (r), CE, NE; S. aizoon , NW(U), CW, SW, S, SE, CE (s); S. cernua , N to S; S. flagellaris , NW(T), N, NE; S. foliolosa , all districts except S and SE; S. groenlandica , N to S; S. hieracifolia , CE, NE; S. hirculus , NW(T), NE (isol); S. nathorstii , CE, NE; S. nivalis , N to S; S. oppositifolia , N to S; S. rivularis , all districts except N; S. stellaris , CW (? Nordre Strømfjord), SW, S, SE; S. tenuis , N to S (r in S and CW (s)); S. tricuspidata , N (western part), NW, CW, SW (n), NE (isol in s).
ROSACEAE (24): Alchemilla acutidens , S, SE (isol); A. alpina , SW, S, SE; A. filicaulis , SW, S, SE; A. glomerulans , CW (r), SW, S, SE, CE (single station in s); A. minor , S; Dryas integrifolia , N to S (two large gaps in SE); D. octopetala subsp. punctata , N, NE, CE; Potentilla chamissonis , N, NW, CW, SW (n), CE, NE; P. crantzii , NW(U), CW (r), SW, S, SE, CE (r),

EA-PS. Böcher: Flora and Vegetation of Greenland

NE; P. egedii (incl. var. groenlandica ), NW(U), CW, SW, S, SE; P. hyp–arctica , N, NW, CW, SW (n), NE, CE, SE (single station); P. nivea , NW, CW, SW, CE, NE; P. palustris , CW, SW, S, SE; P. pulchella , N, NW, CW, SW (r in n), NE, CE; P. ranunculus , CW (isol), S, SE; P. rubella , NE; P. rubricaulis , NW(T), NE, CE; P. stipularis , NE; P. tridentata , CW, SW, S, SE; P. vahli ^ n ^ ana , NW, CW (n), SW (isol at lat. 67° N.); Rubus Chamaemorus , SW, S; R. saxatilis , S, SE; Sibbaldia procumbens , NW(U), SW, S, SE, CE (s), NE (r); Sorbus decora , SW, S.
LEGUIMINOSAE (2): Lathyrus maritimus , S; Vicia cracca , intr to S.
GERANIACEAE (1): Geranium sylvaticum , single station in SW (s).
POLYGALACEAE (1): Polygala serpyllifolia , SE (isol).
CALLITRICHACEAE (4): Callitriche anceps , SW, S, SE; C. autumnalis , NW (Disko), CW (lat. 68°30′ to 69°13′ N.); C. hamulata , SW (r), S, SE; C. verna , CW (r at lat. 69°13′ N.), SW (n and s), S, SE, CE.
EMPETRACEAE (1): Empetrum nigrum subsp. hermaphroditum , all districts except N.
VIOLACEAE (4): Viola ericetorum var. friesiana , SW (s), S; V. labradorica , SW, S; V. palustris , SW, S, SE; V. selkirkii , SW (s).
ONAGRACEAE (7): Chamaenerion angustifolium , SW, CW (r), S, SE; C. latifolium N to S (perhaps not in extreme northern part of N): Epilobium anagalli–difoli [: ]^ u ^ m , NW (Disko), SW, S, SE; E. arcticum , NW(U), CW (r), NE; E. hor–nemannii , NW (Disko), CW (r), SW, S, SE; E. lactiflorum , NW(U) (r in s), SW, S, SE; E. palustre , CW (r), SW, S, SE (single isol station).
HALORAGIDACEAE (2): Myriophyllum alterniflorum , SW (r), S; M. exalbescens , CW.
HIPPURIDACEAE (1): Hippuris vulgaris , all districts except N and SE (n).
UMBELLIFERAE (2): Angelica archangelica , SW, CW (s), S, SE; Ligusticum scoti [: ]cum , SW, S.

EA-PS. Böcher: Flora and Vegetation of Greenland

CORNACEAE (2): Cornus canadensis , S (outer coast at Cape Farewell): C. suecica , SW, S, SE (s).
DISPENSIACEAE (1): Diapensia lapponica , NW (not on basalt), CW, SW, S, SE, CE, NE (s).
PYROLACEAE (3); Pyrola grandiflora , NW, CW, SW, S, NE, CE SE (isol at lat. 66°08′ N.); P. minor , southern Disko, SW, S, SE; P. secunda , SW (isol on southern Disko and at Holsteinsborg).
ERICACEAE (13); Andromeda polifolia (incl. subsp. glaucophylla ), SW (isol at lat. 68°47′ N. and 61°33′ to 62°20′ N.); Arctosphylos alpine , CW (n, isol), SW (Disko and isol at lat. 65° to 66° N.), NE, CE; A. uva-ursi , isol in SW to CW at lat. 66° to 67° N.); Cassiope hypnoides , NW(U), CW, (not inland in s), SW, S (r), SE, NE, CE (r in s); C. tetragona , N, NW, CW, SW (n); CE, NE, SE (n); Ledum decumbens , NW(U) (r in s), CW, SW (r in n); L. groenlandicum , CW (r), SW, S; Loiseleuria procumbens , NW(U) (absent from basalt regions), CW (not in s), SW, S, SE; Oxycoccus quardripetalus var. microphyllus , CW (r, isol at lat. 67°37′ N.), SW, S; Phyllodoce coerulea , NW(U), CW (not inland in s), SW, S, SE, CE (s), NE (r in s); Rhododendron lapponicum , all districts except N; Vaccinium uliginosum , all districts except N; V. vitis-idaea subsp. minus , NW, CW, SW, S (r).
PRIMULACEAE (2): Primula egaliksensis , S; P. stricta , NW(U) (r in s), CW (s), SW (Kugsuq in Godthaabsfjord), two isolated stations in CE and NE.
PLUMBAGINACEAE (1); [: ] Armeria vulgaris (incl. A. sibirica ), N to S (r in SE).
GENTIANACEAE (7): Gentiana aurea , NW (r), CW (r), SW (s, r), S, SE (r); G. detonsa , three separate areas in CW, S, and CE; G. nivalis , CW (r), SW, S, SE, isol in NE; G. tenella , CW, CE, NE; G. uliginosa , S;

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M Lomatogonium rotatum , CW (s), SW, S; Menyanthes trifoliate , CW, SW (s), S.
POLEMONIACEAE (1): Polemonium boreale , NE (lat. 73°42′ to 74°52′ N.).
BORAGINACEAE (1): Mertensia maritima , NW, CW, SW (r), S (r), SE (r).
LABITAE (1): Thymus arcticus, SW, S, SE.
SCHROPHULARIACEAE (15): Bartsia alpina , NW(U) (s), CW, SW, S, SE; Euphrasia arctica , NW(U) (S), CW, SW, S, SE, CE, NE; Limosella aquatica , CW (isol at lat. 71° N.), S, SE (isol); Pedicularis arctica , NW(T); P. capitata , NW(T); P. flammea , all districts except N and NW(T); P. hirsuta , all districts except S and s part of SW and SE; P. labradorica, CW; P. lanata , ?NW(T), NW(U), CW, SW (n); P. lapponica , NW(U), CW, SW, NE CE; Rhinan–thus groenlandicus , SW, S; Veronica alpina , NW, CW (not inland in s), SW, S, SE, NE (s), CE (r); V. fruticans , NW to CW (two stations in Nugssuaq Peninsula), ^ SW, S, ^ SE, NE (s), CE (single station in s); V. pumila , SW, S, SE, NE (s); V. wormskjoldii , SW, S, SE.
LENTIBULARIACEAE (4); Pinguicula vulgaris , NW(U) (s), CW, SW, S, SE, CE (r); Utricularia intermedia , CW (r); U. minor , CW, S (r); U. ochroleuca , CW (r).
PLANTAGINACEAE (1): Plantago maritima (incl. P. juncoides ), NW(U) (s), CW, SW, S, SE.
RUBIACEAE (3); Galium brandegeei , CW (s, r), SW (r), S, SE (isol at lat. 65°37′ N.): G. triflorum , SW, S, SE (r); G. uliginosum , SW (?s, intro).
CAPRIFOLIACEAE (1): Linnaea borealis , SW (r), S (r), SE (r).
CAMPANULACEAE (2): Campanula rotundifolia (coll.), NW(U), CW, SW, S, SE, CE, NE; C. uniflora , N to S, rand alp in S.
COMPOSITAE (46); Achillea millefolium , SW, S, SE (probably intr); Antennaria affinis , CW, S; A. alpina subsp. canescens , NW (s), CW (r in s), SW, S,

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SE, NE, CE; Antennaria sp. (closely related to A. compacta ), CW, S; A. ekmanniana , NW, CW, SW (r), ?SE; A.glabrata , NW(U), CW, SW (n); A. hansii SW, S, SE; A. intermedia , CW (n), SW; A porsildii, NW(U) (single station), CW (n, r), S (single station), SE (single station), CE, NE; Arnica alpina , NW, CW, SW (n), SE (r in n), NE, CE; Artemissia borealis , NW(U), CW, SW (r in n); Circium heterophyllum , SW (single station ?intr); Erigeron borealis , SW, S, SE; E. compositus , N to S (r in S to SE); E. eriocephalus , N, NW, CW, SW (r in n), NE, CE, SE (r); E. unalaschkensis , NW, CW SW (n and nunatak at lat. [: ]^ 6 ^ 2°40′ N.), CE, NE, SE (r in s); E. uniflorus , SW, S, SE, CE (r in s), NE (r in s); Gnaphalium norvegicum , SW, S, SE; G. supinum , NW (s), CW (r, absent inland in s), SW, S, SE, NE (s, r); G. uliginosum , intr to SW, S; Hieracium alpinum , SW (s), S, SE, CE (s), NE (s); H. amitsokense , S; H. angmagssalikensie , SE (isol); H. devoldii , S; H. groenlandicum , SW, S, SE; H. hyparcticum , SW, S, SE; H. ivigtutense , SW, S, SE (s); H. lividorubens , SW (s), S; H. nepiocratum , S; H. rigorosum , SW (s), S, SE (s); H. scholanderi , S; H. stelechodes , SE (isol); H. sylowii , S; Leontodeon autumnale , intr to S; Matricaria inodora , S, isol in [: ]^CE^ to NE; Taraxacum arcticum , N, NE, CE; T. brachyceras , CE NE SE (n); T. [: ]^ hyparcticum ^ N, NW(T); T. lacerum (and T. arctogenum ), N, NW, CW, SW (r in s); T. lapponicum , NW (s), CW (r), SW, S, SE, NE (s, r); T. maurostylum , S, SE; T. naevosum , S; T. phymato–carpum , N, NW, CW (n), NE, CE; T. pumilum , N, NE; T. purpuridens , SE; T. rhodolepis , SE.
Climatically Conditioned Distributions
In Greenland as elsewhere we may distinguish between climatic, edaphic,

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and historical reasons, or combinations of these reasons, for the distribution ^[]^of the various plants. The first distinction to be made is between distribu– tions due to summer heat and distributions also, or chiefly, controlled by the degree of continentality.
Distributions Due to Summer Heat . Here belong the southern species which in the north make for the warm inland regions, and northern species which in the south try to avoid (or are crowded out from) the warm, dry inland regions. Examples of southern species of this type are: Juniperus communis, Potentilla palustris , Galium brandegeei , and Menyanthes trifoliate . Northern species are: Alopecurus alpinus , Poa abbreviate , Ranunculus sulphureus , Draba subcapitata , and Saxifraga flagellaris .
Distributions Due to Degree of Continentality . Numerous southern species have the distributions SW, S, SE, i.e., they are connected with the parts of Greenland influenced by the ocean and make for the north outside the warm inland region because it is too dry there, being either too continental or having soil that is unsuitable because of the local continentality. Examples are Alchemilla alpina , A. vulgaris , Phleum commutatum , Thymus arcticus , Cassiope hypnoides , Cornus suecica , and Carex deflexa .
Other markedly arctic species in the south avoid the moist areas of the coastal regions, which have mild winters. Their main distributions are in N, NW, CW, NE, and CE. Here we may mention such species of Cassiope tetragona , Arnica alpina , Erigeron compositus , Lesquerella arctica , Melandirum triflorum, and Tofieldia coccinea .
There are also a number of low-arctic and subarctic species with markedly continental requirements as regards climate. The are found particularly in CW and CE, and sometimes sporadically in the interiors of SW, S, and SE. We

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may mention here Carex supine , Braya humilis , B. linearis , and, in West Green– land only, Artemisia borealis , Draba lanceolata , Pedicularis labradorica , and Ledum decumbens .
Edaphically Conditioned Distributions
A fair number of species are closely connected with definite conditions of soil, which again may be dependent upon the climate. Indeed, it is often impossible to explain whether climate or soil is decisive. Clearly edaphically conditioned distributions, on the other hand, appear by connection with, or evasion of, the basalt areas in the central parts of West and East Greenland. Connection with basalt in one or both areas is shown, for example, by Dupontia fisheri , Stellaria crassipes , Saxifraga hiriculus , Polemonium boreale , and Arenaria humifusa ; as regards the last-mentioned species the connection, how– ever, is less pronounced. Evasion is evident in acidophilous species such as Loiseleuria procumbens , Vaccinium vitis-idaea , Diapensia lapponica , Potentilla tridentata , Carex arctogena , and, as regards West Greenland, Agrostis borealis .
Historically Conditioned Distributions
Two main points of view have been advanced to explain the present-day distributions of plants: the history of the immigration of the species (whether they have come from the west or from the east), and the persistence of the species during the last glacial epoch in various ice-free [: ] refugia along the coast of Greenland (cf. 8).
There are a fair number of eastern (chiefly Eurasian) species which occur especially or exclusively in East Greenland. Thus Ranunculus glacialis , Draba repens , and Sedum acre , for example, have been found here only, while Viola palustris and Hieracium alpinum have their widest distribution in East Greenland.

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Similarly there are many western (chiefly American) species which have been found in West Greenland only: Halimo ^ lo ^ bos mollis , Carex gynocrates , Potentilla vahliana , Anemone richardsonii , and others. Some, particularly western, species are widely distributed in West Greenland but found in only a very limited area in East Greenland ( Saxifraga tricuspidata , Arabis molboellii , Dryopteris fragrans ), a fact which may be due to immigration from the west and to the different conditions in East and West Greenland during the glacial epochs.
In the southern part of Greenland the boundaries of the areas of several species occur in regions where for topographical reasons the inland ice during a greater glaciations than the present one must be supposed to have spread and formed obstacles to further dispersal. Hence these species may be supposed during the last glacial epoch to have lived in so-called refugia situated between the large tongues of the inland ice, wh ^ e ^ nce they were not able later to spread to all the places where ecological conditions might lead one to ex– pect them to live.
In West Greenland there is, at latitude 66° N., a considerable break due to the inland ice. North of this we have the southern limits of Ranunculus affinis , Carex holostoma , C. misandra , Luzula arctica , Pedicularis lanata , and other plants. It is true that they are continental species, but their southern limit is no doubt conditioned partly by historical considerations, and partly by the fact that suitable stations south of the tongue of ice are few and far between. At latitude 62°30′ N. there is another ice tongue, and immediately south of this we have the northern limit of, for example, Hieracium alpinum and Carex atrata . Still farther south many species may, in the Julianehaab District, have been cut off by ice advancing in a southward direction between

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longitude 46° and 48° W.; Viola selkirkii has its southern limit about this area. Immediately south of Angmagssalik in East Greenland there is also an area which no doubt formerly was highly glaciated, and north of this area we find the southern limits of such species as Ranunculus glacialis , Pedicularis hirsuta , and, so far as their local areas are concerned, of Betula nana and Dryas integrifolia .
Many species are extremely rare, occurring in peculiar small areas which are best explained through an assumption that nowadays they like in the near the refugia where they persisted during the glacial epoch. Good examples of such species are Polemonium boreale , Eriophorum callithrix , and Ranunculus suricomus in northern East Greenland, Sedum acre in southern East Greenland, Selaginella repestris in South Greenland, Arctostaphylos uva-ursi and Anemone richardsonii in the central part of West Greenland, and Pedicularis capitata and P. arctica in North Greenland.
There are a number of supposedly endemic species in Greenland. These, too, suggest that the flora is of a great age in Greenland itself. We may mention Melandrium triflorum with a wide distribution in Greenland (it may also occur to the west), Braya thorild-wulfii , and the amphidiploids Saxifraga nathorstii , as well as several apomictic species within the genus Hieracium . There are also certain conditions indicative of diploid races of Campanula rotundifolia being endemic to Greenland.
Distribution of Western and Eastern Species
Among the western species (with their main distribution west of Green– land), few are associated with regions of mainly oceanic character (SE, S, SW). We may, however, mention Minuartia groenlandica , Ledum groenlandicum, Coptis trifolia , Cornus canadensis , Carex stylosa , and C. deflexa . On the

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Other hand, there are a good number of western species in the continental areas, examples being: Saxifraga tricuspidata , Draba lanceolata , Antennaria Compacta , Tofieldia occinea , and Erigeron compositus .
Inversely, there are a large number of eastern (Eurasian) species asso– ciated with the oceanic regions, e.g., Alchemilla alpina , Sedum annum, Saxifraga stellaris , Juncus squarrosus , and Carex rufina. There are very few eastern species in the continental regions, although we here mention Draba repens , Carex parallela , and perhaps Gentiana tanella. Betula nana has a con– tinental tendency and is extremely rare in eastern America. Hence it has probably reached Greenland from the east.
Among both the western and eastern species in Greenland, there are several whose distribution is little influenced by the degree of continentality, e.g., Carex scirpoides and Potentilla tridentata (western), and Angelica archangelica (eastern).
As might be expected, southern East Greenland has the greatest share of eastern species, while West Greenland predominates as regards western species. This especially applies to the northern and continental portions.
REGIONS OF VEGETATION
In considering the vegetation of Greenland, Warming and Rosenvinge used three divisions: a birch region (inner fjords in S), a willow scrub region (in West Greenland extending as far north as about the Nugssuak Peninsula), and a distinctly arctic region with heaths and barrens. Later, the geograph– ical division of Greenland vegetation came to be worked out in various ways.
Subarctic Region . The mean temperature of July is about 10°C. or a little more. Warm valleys occur in the interior of S, SW, and the southern– most CW. The luxuriant freshwater vegetation, rich in species, is characteristic

EA-PS. Böcher: Flora and Vegetation of Greenland

of this region. In S the region is characterized by tall birch scrub (Fig. 2), in SW by alder scrub (Fig. 3) and tall, luxuriant willow scrub, and in CW by steppe-like communities (Fig. 7).
Low-Arctic Region . The mean temperature of July is about 6° [: ]^ to ^ 9°C. The region includes the greater parts of CW, SW, outer areas of S, the greater part of SE, and the interior of Scoresby Sound and the southernmost part of King Christian X Land in CE. The region is particularly characterized by willow scrub, in tracts near the ocean, also by meadows with high perennials (Angelica archangelica) and luxuriant herb fields and various heaths, in con– tinental areas by, among other features, a type of Kobresia myosuroides . “steppe”.
High-Arctic Region . The mean temperature of July is about 2° to 6°C. in this region, which includes N, NW, NE, northern part of CE, and high-alpine areas in south. Characteristic features are certain heaths ( Cassiope tetragona ) and dry cyperaceous communities ( Carex nardina and C. rupestris ), several types of snow-patch vegetation and of grassy meadows ( Arctagrostis and Alope–curus alpina ) and swamps.
Within these three regions various subregions may be distinguished, gen– erally separated by differences in degree of continentality. The same three main regions may be found in the mountains of Scandinavia, and in other parts of the circumpolar or circumboreal area.
Distributions of Species in the Vegetation Regions . Many species are completely or chiefly associated with one or other of the regions and hence may be characterized as subarctic, low-arctic, or high-arctic. Subarctic (relatively temperate) species are, for example, Potamogeton natans , Carex buxbaumii , Dryopteris fillix-mas ; low-arctic species include Bartsia alpina , Sibbaldia procumbens, Gnaphalium norvegicum ; and high-arctic species are

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Salix arctica , Draba subcapitata , Deschampsia brevifolia , and Pleuropogon sabinii .
Many species are widely distributed in the arctic regions: Saxifraga cernua , S. oppositifolia , Oxyria digyna , Polygonum viviparum , and others. The species may be grouped biologically in types of distribution, e.g., widely distributed arctic, high-arctic-continental, low-arctic-oceanic (see details in 1).
PLANT COMMUNITIES
The following grouping is made on the basis of life form and type of distribution of the species. The main groups are distinguished by life form, the subgroups largely by types of distribution.
Scrub and Wood
There are plant communities showing several strata, the upper one con– sisting of low trees or shrubs. The understratum consists mostly of herbs and mosses, in rarer cases of drawf shrubs or junipers.
Low birch woods (Fig. 2) consist of open woodland or tall scrub of Betula pubescens s.l., 4 to 8 meters high, with understrata of grasses ( Deschampsia flexuosa , Anthoxanthum ) or ferns ( Dryopteris linnaeana , D. phegopteris ), rich in mosses. Warm valleys are found in S on rather dry soil that is poor in nutrients.
Alder scrub (Fig. 3) contains vegetation dominated by Alnus crispa shrubs 1 1/2 to 2 1/2 meters high, with understrata of mosses and herbs. They are found in the interior of SW; particularly along running water, often as a kind of low gallery forest. The soil is fairly rich in nutrients.

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Willow scrub contains vegetation dominated by Salix glauca (incl. S. cal–licarpaea ) 1/2 to 3 meters high. The understrata are of two main types.
(1) Oceanic type: understratum rich in mosses and herbs, including num– erous species in common with the her fields; found in SW, S, and SE; on well– drained, moist soil, in the north chiefly on slopes having a southern exposure; in winter protected by a thick snow cover.
(2) Continental type (Fig. 7): Pyrola grandiflora , Calamagrostis spp., and Stellaria longipes (and S. monantha ) frequent; CW (southern part), CE (Scoresby Sound); on rather dry, warm soil, with the snow cover presumably moderate, but constant in winter.
Dwarf Shrub
There are communities of ericaceous, more or less evergreen, drawf shrubs and low willows rising over an understratum consisting particularly of mosses and lichens. The communities may be closed, and then generally are called heaths; or they may be open, with gravel and bare ground between the plants, and then are termed “fell fields” (barrens).
Oceanic Communities . There are found particularly on acid soils in regions with heavy precipitation (SW, S, SE). A number of species with oceanic distribu– tion are included.
(1) Salix uva-ursi type: open vegetation in places exposed to gales and black frost, often rich in lichens; the soil is gravelly, slightly acid; occurs in S and SW.
(2) Loiseleuria-Diapensia type: often rich in lichens, occurring in highly wind-swept places more or less bare of snow; soil acid; appears both as heath and fell field.
(3) Empetrum hermaphroditum type: often rich in Vaccinium uliginosum ;

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the heath type of the skerries (Fig. 4), occurring in wind-swept terrain often as more or less open fell field; communities in more protected terrain are rich in mosses.
( 4 ) Phyllodoce-Deschampsia flexuosa type: rich in mosses, occurring in places with a constant snow cover in winter and slight drying-up in summer; characteristic species are Cornus suecica , Lycopodium annotinum , and Ledum groenlandicum .
Continental Communities . These occur on slightly acid, neutral, or basis soils particularly in CW, NW, N, NE CE, and include several species with a continental type of distribution. The following eight types may be recognized.
( 1 ) Dryas integrifolia-Carex nardina type: found on very wind-swept ground, being often bare of snow, and occurs mostly as fell field; found in alpine situations in subarctic areas of CW; soil is acid to neutral; other Dryas communities are ecologically more closely related to steppe-like communities.
( 2 ) Vaccinium uliginosum-Diapensia type: rich in lichens, and occurs on dry, rather wind-swept places, both as heath and fell field; soil is fairly acid; found in alpine situations in subarctic areas.
( 3 ) Arctostaphylos uva-ursi type: develop on sunny slopes on dry, neutral soil, and forms a transition to the steppe-like communities; this type of heath is of a subarctic character in Greenland, where it occurs only in the southern part of CW.
( 4 ) Betula nana (or glandulosa ) type (Fig. 5): a low-arctic-subcontinental type of heath, in SW and SE often rich in Empetrum ; soil is acid and the snow covering may be considerable; Betula nana heath rich in lichens and Hierochloe alpina is developed particularly in the interior of SW and S (in the latter instance B. glandulosa in the dominant).

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( 5 ) Betula nana-Ledum decumbens-Aulacomnium turgidum type: characteristic of northern exposures in the lowland in CW; a typical associate is Pedicularis labradorica ; snow covered in winter.
( 6 ) Cassiope tetragona type: characteristic high-arctic-continental but occurs also in heath, alpine situations in the south in CW, and in northern parts of SE and SW; often rich in mosses; soil is acid or nearly neutral; develop in places protected by snow in winter and having a medium-long period of snow covering into the summer; many subtypes say be recognized, e.g., the Cassiope-Vaccinium uliglnosum type (less protected, frequently rich in such lichens as species of Stereocaulon and Catraria ), and the Cassiope-Dryas type occurring on basic soil,
( 7 ) Vaccinium uliginosum-Drayas integrifolia type (with scattered Rhodo–dendron lapponicum and Arctostaphylos alpina : more or less snow-protected, occurring on rather dry and basic soil in NW(U).
( 8 ) Rhododendron lapponicum type: developed on moist circumneutral soil, often with water oozing through, and is mostly rich in mosses; particularly fertile soil it way be rich in Dryas .
Bogs . In the southern part of Greenland, tussocky bog communities are observed that are dominated by various dwarf shrubs ( Vaccinium uliginosum , Empetrun , low phases of Salix glauca s.l.). These communities have not been investigated in detail, but an important example is the Salix arctophila type, which is particularly widely distributed in S, SW, and CW. In oceanic regions the soil is acid and the tussocks rich in Sphagnum . In continental regions there are other types (e.g., with Tomenthypnum nitens and Ranunculus lapponi–cus ) developed on neutral soil.
Xerophilous Grasses and Grasslike Plants
The vegetation consists of xerophilous Gramineae, Cyperaceae, and Juncaceae,

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and in certain cases of Dryas as well. As in the preceding types, the vege– tation may be closed or consist of open fell field, and the ground stratum way vary between being rich is lichens and being rich in mosses.
Oceanic Communities . The soil is acid, and the distribution SW, S, and SE. Three types are found.
( 1 ) Juncus trifidus types comprises open vegetation on wind-swept gravelly soil, but is mostly rich in lichens; related to the Loiseleuria– Diapensia type of dwarf shrub community.
( 2 ) Luzula spicata-Agrostis borealis type: found on lees exposed soil and is often rich in lichens; snow covering is moderates a subtype with Carex rigida dominant has been described from SE; here, among other plants, was much T ^ h ^ ymus arcticus (Fig. 6).
( 3 ) Deschampsia flexuosa type: rich in Cladonia , or in mosses on areas having a snow-patch character; forms closed communities on acid, protected soil.
Continental Communities (steppe-like and related types). The soil is neutral to basic, sad the distribution particularly In CW (NW), N, and CE (Ne). No less than five types may be distinguished.
( 1 ) Carex nardina-Poa abbreviata type: developed on wind-swept terrain, In high-arctic or alpine areas; often rich in Dryas and constitutes a transition to the [: ]^ Dryas ^ type mentioned above. Typical associates are, for example, Les–querella arctica and Arenaria pseudofri gida.
( 2 ) Carex rupestris type: often comprises a sense vegetation cover; developed on soil [: ]s that are slightly acid to neutral high-arctic, or alpine in the south; sometimes very rich in lichens; and is some places a Dryas-Carex rupestris vegetation covers rather large areas (fig. 9); typical associated species are Potentilla nivea , Pedicularis lanata , Lesquerella arctica and

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Campanula uniflora .
( 3 ) Kobresia myosuroides type (“ Elyna heath”): a steppe-like vegetation which may cover fairly large areas is CW end CE on dry, circumenutral, often loess soils; e special subtype developed on soil rich in salts occurs near salt lakes In CW(southern pert) end contains halophilous species.
( 4 ) Carex supina type (Fig. 7): occurs on dry loses soil, being often half open, with a sparse xerophytic lichen-moss flora among the higher plants; comprises a kind of subarctic steppe developing on southern exposures In CW soil is neutral.
( 5 ) Calamagrostis purpurascens types occurs in similar places to type (4), but on more alkaline soils occasionally it is developed on salt soil (subtype with Gentiana detonsa in CW); characteristic associated species ere Artemisia borealis and Potentilla chamissonis .
Psammophilous Communities . Most characteristic is ^ the ^ Elymus mollis type on sandy seashores and on inland dunes. Honckenya peploides and Festuca rubra are typical of this community.
Xerophilous Pteridophytes and Herbaceous Dicotyledons
These communities have not been investigated sufficiently for detailed description. Most of them are found on rocky terrain in which the rock is covered by shallow soil, or in rock crevices. From S a vegetation character– ized by Selaginella rupestris has been described, and types of vegetation dom– inated by Dryopteris fragrans or Woodsia ilvensls have been found in CW. Important and typical species occurring on dry ledges are, moreover, Saxifraga tricuspidata [: ]^ and ^ S. aizoon and sometimes Potentilla tridentata .

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Nondwarfish Hygrophytic and Mesophytic Herbs
These include, among others, the communities which were formerly termed herb fields. Their colorful profusion of flowers makes them the most beauti– ful types of vegetation of Greenland.
High Perennials . These are most frequent in the subarctic parts and in particularly favorable places is the low-arctic regions. They occur is S (and SW) on moist south-facing slopes that are snow-covered is winter. Examples are tall growths of Dryopteris falix-mas or Streptopus amplexifolius , associated with Angelica archangelica , Chamaenerion angustifolium , and Alchemilla glomer–ulans . Near springs in continental CW occur growths of Archangelica (often associated with Calamagrostiz langsdorfii ) which may be classified here.
Rosette Plants ( Herb Field ). In the north these occur only on south-facing soil protected by snow, but in the south they also occur along rivers, though always where they are protected by snow in winter. They are absent in the very northernmost parts of Greenland, six types may be distinguished.
( 1 ) Alchemilla “vulg aris” type (sometimes rich is Sedum rosea ) Fig. 10.: the type most closely related to the last preceding community, and is found in SW, S, SE, and Gasseland is Scoresby Sound.
(2) Taraxacum lapponicum-Phelum commutatum type, generally with Polygonum viviparum as its dominant: important or characteristic associated species include Poa alpina , Thalictrum alpinum , Veronica fruticans , Erigeron uniflorus , and Botrychium lanceolatum ; this type is found in SE, S, and SW.
( 3 ) Alchemilla alpina type: forms transitions to the Sibbaldia type of dwarfish herbs or to the Deschampsia flexuosa type; this is developed is SW, SE, and S, and is alpine in subarctic parts of S.
(4) Potentilla crantzii-Antennaria alpina (subsp. canescens ) type, often

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with Carastium alpinum as its dominant: typical associated species are Erigeron unalaschensis , Draba nivalis , etc.; this type is known from the southern part of NE; related types occur in NW(U).
( 5 ) Taraxacum arcticum type: characterized among other species by Ranunculus nivalis ^ Ranunculus nivalis ^ , R. sulphureus ^ R. sulphureus ^ , and Melandrium apetalum; ^ Melandrium apetalum; ^ this occurs in NE.
( 6 ) Arnica alpina type: forms a transition to the steppe-like communi– ties; this is a continental arctic type that is such more xerophilous than the others, and with slight snow covering; characteristic species are Erig–eron eriocephalus , Minuartia rubella , Campanula uniflora , and in CW, also Draba aurea .
Protshemicryptophytes (plant with elongated leaf-bearing aerial shoots ). Here, particularly, belong the communities dominated by Chamaenerion lati –folium occurring on open sand (Fig. 8) and gravel and in river beds.
Dwarfish Herbs and Woody Plants
These are the snow-cover communities proper, or “snow patches” — i.e., the vegetation characterized by a snow covering of very long duration, the period of vegetation accordingly being very short. The vegetation is rich in mosses and particularly rich in hepatics (species of Anthelia , Junger–mannia, etc.), and the vascular plants rise little or not at all above the moss carpet. Three types may be noted.
( 1 ) Sibbaldia procumbens type; closely related to the herb fields, as the snow covering is not of extremely long duration and the exposure is often favorable; Vernica alpina is frequent, and Draba crassifolia has been observed in several places.
( 2 ) Salix herbacea type: comprises a group of communities, some of which

EA-PS. Böcher: Flora and Vegetation of Greenland

are rich in lichens or Caasiope hypnoides , others in Oxyria , Ranunculus pyg–maeus (Fig. 13), or Equisetum arvense , etc.; further, there are many differ– ent kinds of subdominants, such as species of Anthelia , Dicranum , Polytrichum , or Timmia ; characteristic higher plant species include Gnaphalium supinum and Carex lachenalii ; this type of vegetation is extremely important in regions of Greenland rich in snow, hut rare and limited to al l ^ p ^ ine areas, for example, in the southern part of CW.
(3) Phippsia algida-Cerastium cerastoides types occurs on soil with a particularly long-lasting snow covering, and on summer-wet soil close to melt– ing snowdrifts; in East Greenland Ranunculus glacialis may be frequent in this community; other species to be mentioned are Saxifraga rivularis , S. cernua , and Carex lachenalii .
Transitional types between this kind of snow-patch vegetation and a boggy or vet meadow-like vegetation are frequent, particularly in high-arctic areas (cf. Fig, 12). There also may occur a number of related communities, for exam– ple, vegetation dominated by Luzula arctica and Potentilla hyparctica (NW), or by Draba micropet ala and Saxifraga foliolosa (NE).
Mesophilous and Hygrophilous Grasses and Grasslike Plants
These are generally meadow-like types of vegetation rich in mosses, and occur in places that are influenced by subsoil water or by water oozing out. In the north they also occur on so-called earth glaciers and vet solifluction soil. The group is not sufficiently known for detailed treatment, the several types mentioned being provisional.
Nonhalophtic Communities . ( 1 ) Arctagrostis-Alopecurus alpinus type: comprises a high-arctic grassy meadow with Poa arctica , Carex misandra , and other associates and occurs in CE, NE, NW, and N; calciphilous subtypes with Carex atrofusca also

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occur; closely related communities with Ranunculus a ffinis and Saxifraga flagellaris are found on vet solifluction clay.
( 2 ) Calamagrostis neglecta type: a low-arctic meadow vegetation described from SE (is an oceanic variant with Deschampsia alpina ) and found frequently in NW(U) and CW, where, in the southern part, summer-dry C. neglecta meadows with abundant Poa pratensis and Ranunculus affinis may develop.
( 3 ) Calamagrostis lengsdorfii type: a southern tall-grass meadow asso– ciated with willow and birch scrubs, and is found in CW mostly along running water.
( 4 ) Carex rariflora type: this often tussocky vegetation rich in mosses is greatly reminiscent of raised bogs in more southerly regions; especially frequent in the southern part of Greenland, it shows all stages of transition to the Salix arctophila type; it occurs mostly on oligotrophic soil.
( 5 ) Carex holostoma type: this has been observed to s ^ u ^ rround alpine marshes and pools of acid water in CW.
( 6 ) Scirpus caespitosus type: this develops near ozzing or slowly running water, often on shallow soil over rocks, and occurs in the southern parts of Greenland having an oceanic character; in southern CW it is alpine and rare.
( 7 ) Kobresia caricina-Juncus castaneus type: calciphilous marsh vegetation developed where water oozes or runs out, and is particularly frequent in con– tinental regions; closely related to this type, but perhaps independent, are communities in which Carex microglochin is dominant.
( 8 ) Juncus arcticus type: this is often found on sand or clay near banks of alkaline lakes or in river beds; it is most frequent in continental regions.
( 9 ) Carex aquatilis var. stans type (locally with abundant Equisetum varie–gatum ): this occurs on level areas with water oozing through the soil, and is

EA-PS. Böcher: Flora and Vegetation of Greenland

found Inland in the Nugssuaq Peninsula,
(D) Carex incurva type: this is developed in holes in Rhododendron heaths, on sandy banks of rivers and lakes, or in salt lake basins; most fre– quent in continental regions, but in general is widely distributed.
( 11 ) Scirpus pauciflorus type: this is found on banks of alkaline or salt lakes in CW.
Halophytic Communities . (1) Puccinellia phryg [: ]^ a ^ nodes type (sometimes with Carex subspathacea or, in northern Greenland, C. ursina ): this is the widely distributed salt-marsh vegetation developed on lower levels which are covered frequently by spring tides; in South Greenland Puccinellia mari–tima occurs in a similar type of vegetation.
( 2 ) Carex subspathacea-C. aquatilis var. stans type: this occurs about lagoons and on sandy or clayey soil on seashores in northernmost Greenland; farther south, related types with Stellaria humifusa and Carex glareosa are found on salt flats which are covered only by the higher spring tides.
( 3 ) Dupontia fisheri type: this is local in NW and NE occurring mainly at higher levels than the preceding type.
( 4 ) Puccinellia deschampsioides-Plantago maritima type (often associated with Triglochin palustre ): found on salt lakes in southern CW, in one place with associated Ranunculus cymbalaria ; transitions to steppe-like communities are seen when there are associated species of Braya , Primula stricta, and Gentiana detonsa .
Helophytes
This vegetation group is very sparsely developed in Greenland. It may be divided into four types.

EA-PS. Böcher: Flora and Vegetation of Greenland

( 1 ) Carex saxatilis-rotundata type: this is developed particularly as marginal vegetation around lakes, forming narrow meadow-like communities, and, as such, belonging to the preceding groups, although also found In the later as marsh vegetation; widely distributed, but occurs particularly in continental regions.
( 2 ) Carex rostrata type: this is developed in wet bogs and at the edge of lakes; it is found only in S.
( 3 ) Eriophorum scheuchzeri type: a widely distributed marsh [: ] plant com– munity about the margin of fresh water, the main species also occurring in many of the other types; the present type comprises the numerous cases in which Eriophorum scheuchzeri is the only significant species.
( 4 ) Menyanthes trifoliata type (Fig. 15); this occurs in shallow water about the margins of lakes with water poor in nutrients, and is often rich in Potentilla palustris and Sphagna ; it is found in CW, SW, and S.
Hydrophytes
Vegetation of genuine hydrophytes occurs particularly in the southern parts of Greenland, Hippuris vulgaris , the most northerly of all hydrophytes, reaches as far north as about latitude 77° N. in both West and East Greenland. Five types may be distinguished,
( 1 ) Hippuris-Pleuropogon sabinii type; Ranunculus hyperboreus may enter as a constituent of this high-arctic aquatic vegetation.
( 2 ) Potamogeton filiformis type: ^ this is ^ widely distributed, and includes in CW the vegetation of vascular plants occurring in salt lakes with the water not too saline; in the highly saline lakes only plankton was found.

EA-PS. Böcher: Flora and Vegetation of Greenland

( 3 ) Potamogeton alpinus-Sparganium hyperboreum type; this is distributed in CW, SW and S, occurring in fairly acid small lakes with the water rich in humus.
( 4 ) Sparganium angustifolium type: this inhabits oligotrophic pools in oceanic regions, and has been described from Angmagssalik in SE.
( 5 ) Subularia aquatica type: this oceanic type was described from a pond at Angmagssalik, and contained, among other associated species, Juncus bulbosus .
Besides the types mentioned above, small ponds with abundant Utricularia (in CW) and pools mainly with freshwater algae ( Stigonemataceae , Tribonema , etc.) have been observed. So far, investigations of freshwater planton have been made to only a small extent.
Cryophilous Algae
Here, particularly, belong the communities of microscopic algae forming “red snow” ( Chlamydomonas nivalis , etc.), and the communities of diatoms oc– curring in cryoconite cavities in the inland ice.
Marine Algae
A distinction it here made between littoral and sublittoral communities. The littoral region is the area between the upper limit of algal vegetation and the lowest tidemerk, below which we have the sublittoral region, which stretches down to the lower limit of algal vegetation at a depth of about 40 to 45 meters. Several types can be identified.
[: ] (1) Fucus vesiculosus-Ascophyllum type: this is developed in the littoral region in NW(U), CW, SW, S, and SE, particularly in protected places; F ucus vesicolosus and Ascophyllum nodosum are the most important species, and support various epiphytes.
(2) Fucus inflatus type: this is much as in the preceding case, but

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occupies a zone below it and sometimes occurs in less protected places.
( 3 ) Th Monostroma-Urospora-Bania : this is found in the littoral region in the southern parts of Greenland, particularly in exposed places and at such a height that the vegetation at low tide may be dried up; important species are Monostroma groenlandicum , Ulothrix flacca , Urospora mirabilis , and Bangia fuscopurpurea .
( 4 ) Chordaria-Dictyosiphon type: thin inhabits the lower part of the littoral region and rock crevices where there can be no serious drying up at low tide.
( 5 ) Laminaria type: this is found in the upper sublittoral region, at a depth of 9 to 20 meters, and constitutes large submerged “woods” of Laminaria nigripes and L. longicruris (of extensive dimensions), and in S, also L. groenlandica ; associated may be Alaria pylaii and Saccorhiza dermatodea .
( 6 ) Agarum turneri type: this occurs at a depth of 18 to 30 meters and is widely distributed on both coasts; typical associated species include Laminaria solidungula and Euthora cristata .
( 7 ) Desmarestia culeata type: this is widely distributed and, ecolog– ically speaking, closely related to the preceding type; it is developed par– ticularly in protected bays and natural harbors, the algae often lying partly loose on the bottom; among typical associated species may be mentioned Caetomorpha melagonium .
(8) Ptilota-Lithoderma type: this is widely distributed at depths of about 20 to 40 meters, and is characterized by upright Florideae including crustose species of Lithoderma and Lithothamnion; sometimes the crustose species are dominant.
( [: ]^ 9 ^ ) Lithothamnion type: this is found where there is a gravelly or

EA-PS. Böcher: Flora and Vegetation of Greenland

stony, often comparatively flat bottom at a depth of 20 to 40 meters; char– acteristic associates are the upright branched species of Lithothamnion , L. glaciale , L. flabellatum , L. fruticulosum , etc.).
For details concerning the marine plankton vegetation of the waters of Greenland (communities of diatoms such as Chaetoceras , Thalassiosira , and Coscinodiscus and various communities of Peridineae, etc.), reference should be made to the article on “Phytoplankton.”
Investigations of communities of lichens and bryophytes on rocks (cf. Fig. 16.) have recently been started in Greenland, but the results are not yet available (1950).

EA- II ^ PS. ^ Böcher: Flora and Vegetation of Greenland

BIBLIOGRAPHY

1. Böcher, T.W. “Biological distributional types in the flora of Greenland,” Medd.Grønland vol.106, no.2, 1938. With a comprehensive list of references.

2. ----. “Climate, soil, and lakes in continental West Greenland in relation to plant life,” Medd.Grønland vol.147, no.2, 1949.

3. ----. “Contributions to the flora and plant geography of West Greenland I-II,” Ibid . vol.147, no.3 and 7, 1948, 1950.

4. ----. “Phylogeographical studies of the Greenland flora,” Ibid . vol. 104, no.3, 1933.

5. ----. “Studies on the vegetation of the east coast of Greenland,” Ibid . vol.104, no.4, 1933.

6. ----, and Larsen, Kai. “Chromosome numbers of some arctic or boreal flowering plants,” Ibid . vol.147, no.6, 1950.

7. Devold, F., and Scolander, P.F. “Flowering plants and ferns of South East Greenland,” Norsk Polar institutt, Skrifter no.56, 1933.

8. Gelting, P. “Studies on the vascular plants of East Greenland between Frantz Joseph Fjord and Dove Bay,” Medd.Grønland vol.101, no.2, 1934.

9. Hagerup, O. “Empetrum hermaphroditum,” Dansk Bot.Arkiv vol.5, no.2, 1927.

10. Hartz, N. “Fanerogamer og Karkryptogamer fra Nordøst-Grønland…og Angmagsalik,” Medd.Grønland vol.18,pp.315-931, 1895.

11. ----. “Østgrønlands vegetationsforhold.” Ibid . vol.18, pp. [: ] 105-314, 1895.

12. ----, and Kruuse, C. “The [: ] vegetation of Northeast Greenland,” Ibid . vol.30, pp.333-431, 1911.

13. Iversen, J. :Moorgeologische Untersuchungen auf Grönland,” Dansk Geol.Foren. Medd . Vol.8, p.341, 1934.

14. Kruuse, Chr. “List of Phanerogams and vascular Cryptogams found in the Angmagsalik District,” Medd.Grønland vol.30, pp. ^ 2 ^ 09-88, 1906.

15. ----. “Rejser og botaniske undersøgelser i [: ] østgrønland samt Angmagsalikegnens vegetation,” Ibid . vol.49, 1912.

EA- II ^ PS ^ . Böcher. Flora and Vegetation of Greenland

16. ----. “Vegetationen i Egedesminde Skjaergaard,” Ibid . vol.14, pp.348– 400, 1897.

17. Lange, J. “Conspectus florae Groenlandicae,” Ibid . vol.3, 1880.

18. Nathorst, A.G. “Kritiska anmärkningar om den grönlandska vegetationens historia,” Svenska Vetenskapsakad. Bihang Handl . vol.16, pt.3, no.6, 1890.

19. Omang, S.O.F. “Übersicht über die Hieraciumflora Ost-Grönlands,” Norsk Polarinstitutt, Skrifter no.55, 1933.

20. Ostenfeld, C.H. “The flora of Greenland and its origin,” Danske Vidensk.Selsk.Biologiske, Medd . Vol.6, no.3, 1926.

21. ----. “The vegetation of the north-coast of Greenland,” Medd.Grønland vol.64, pp.221-68, 1923.

22. Polunin, N. “Contributions to the flora and phytogeography of south– western Greenland,” Linnean Soc. J.(Bot.) vol.52, p.349, 1942.

23. Porsild, A.E. “Contributions to the flora of West Greenland,” Medd . Grønland vol.58, pp.157-96, 1926.

24. Porsild, M. P. “Alien plants and apophytes of Greenland,” Ibid . vol.92, no.1, 1932.

25. ----. “The flora of Disko Island [: ] and the adjacent coast of West Greenland,” Ibid . vol.58, pp.1-156, 1920.

26. ----. “Skildring of vegetationen paa Øen Disko,” Ibid . vol.25, pp.91-240, 1902.

27. ----. “Stray contributions of the flora of Greenland,” Ibid . vol.77, pp.1-44, 1930; vol.93, no.3, 1935.

28. Rosenvinge, L.K. “Grønlands Havalger,” Ibid . vol.3, pp.763-974, 1893.

29. ----. “Om algevegetationen ved Grønlands kyster,” Ibid . vol.20, pp.129-244, 1899.

30. ----. “Det sydligate ^ G ^ rønlands vegetation,” Ibid . vol.15, pp.73-250, 1896.

31. Seidenfaden, G. “The vascular plants of ^ S ^ outh East Greenland,” Ibid . vol.106, no.3, 1933.

32. ----, and Sørensen, Th. “The vascular plants of Northeast Greenland from 74° 30′ to 79° 00′ N.Lat.,” Ibid . vol.101, no.4, 1937.

EA- II ^ PS ^ . Böcher. Flora and Vegetation of Greenland

33. Sørensen, Th. “The flora of Melville Bugt,” Ibid . vol.124, no.5, 1943.

34. ----. “Summary of the botanical investigations in N.E. Greenland,” Ibid . vol.144, no.3, 1945. With bibliography.

35. ----. “Temperature relations and phenology of the Northeast Greenland flowering plants.” Ibid . vol.125, no.9, 1941.

36. ----. “The vascular plants of East Greenland from 71° to 73° 30′ N.Lat.,” Ibid . vol.101, no.3, 1933.

37. Trapnell, C.G. “Vegetation types in Godthaab Fjord,” J.Ecol . vol.21, p.294, 1933.

38. Warming, Eugenius. “Om Grønlands vegetation,” [: ]^ Medd.Grønland ^ vol.12, 1888.

39. ----. “The vegetation of Greenland,” Commission for the Direction of the Geological and Geographical Investigations in Greenland. Greenland . Editors: M.Vahl, G.C. Amdrup, L. Bobe, Ad.S. Jensen. Vol. I. The Discovery of Greenland, Explora–tion and Nature of the Country . Copenhagen, Reitzel; London, Milford, 1928, pp.291-317.

Tyge W. Böcher
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