ACTA PHYTOGEOGRAPHICA SUECICA
EDIDI'l' SVENSKA V AXTGEOGRAFISKA SA.LLSKAPET
27
FLORISTIC PHYTOGEOGRAPHY OF
SOUTH-WESTERN LULE LAPPMARK
(SWEDISH LAPLAND)
I
BY
STEN SELANDER
UPPSALA 1950
ALMQVIST & WIKSELLS BOKTRYCKERI AB SVENSKA VAXTGEOGRAFISKA SALLSKAPET
(SOCIETAS PHYTOGEOGRAPHICA SUECANA)
Adress: Uppsala Universitets Vaxtbiologiska Institution, Uppsala o, Sverige
Styrelse: Ordf. Prof. G. EINAR Du RIETZ, v. ordf. Prof. Huoo OsvALD, sekr. Fil. lie. NILs QUENNERSTEDT, skattm. Fil. lie� 0LOF RUNE, red. Fil.lic.MATs W1ERN, klubbm. Fil.lic.MA. GNUS FRIEs, i5vr.: Prof. ERic HULTEN,
Doe. FREDRIK HARD AV SEGERSTAD, Prof. JoHN A.xEL NANNFELDT, Fil.lie. GusTAF SANDBERG, Fil. lie. RoLF SANTESSON, Prof. SVEN TlruNMARK.
Sallskapet, som utgor en fortsattning av Svenska skyddet samt att arbeta for den vaxtgeografiska Vaxtsociologiska Sallskapet, )>ar en foreningslank mel forskningens utnyttjande i vArt lands naringsliv.» lan Sveriges vaxtgeografer och ovriga for vaxtgeo Intrade vinnes genom inval efter anmalan hos grafisk forskning ·intresserade personer; dess andamAI sekreteraren under ovannamnda adress. Arsavgift 10 ar att vacka, underhalla och framja intresset for kronor; standig medlemsavgift 200 kronor, dock att vaxtgeografien i vidstracktaste mening, sarskilt utfors arligen betalande medlem, som tillliort Sallskapet i kandet av svensk vegetation och flora, samt att havda. minst 15 ar, kan bli standig medlem mot erlaggande vaxtgeografiens stallning inom svensk naturforsk av 100 kronor. ning>>.- »For detta andamal skall Sallskapet verka Acta Phytogeographica Suecica utsandas till med� hi. a. genom att anordna sammankomster och e:xkur lemmarna niot giropostforskott pa arsavgift + porto. sioner, att utgiva en publikationsserie Acta Phytogeo Da flera band av publikationsserien utgivas under graphica Suecica, vilken utkommer med ett eller flera ett ar, ma styrelsen uttaga en tillaggsavgift av sadan band arligen, att framja. det vaxtgeografiska natur- storlek, att kostnaderna tackas.
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Publikationerna kunna aven erhallas genom The Acta Phytogeographica Suecica may be byte efter t>verenskommelse med Uppsala Uni obtained by exchange on application to Uppsala versitets Vaxtbiologiska Institution. Universitets Vaxtbiologiska Institution.
Die Acta Phytogeographica Suecica kt>nnen nach "Obereinkommen mit der ,Uppsala Uni· versitets Vaxtbiologiska Institution" auch im Austauschswege erhalten werden. FLORISTIC PHYTOGEOGRAPHY OF
SOUTH .. WESTERN LULE LAPPMARI(
(SWEDISH LAPLAND)
I
INAUGURAL DISSERTATION
BY STEN SELANDER
FIL. LIC., STOCKH.
By due permission of the Philosophical Faculty, Natural Science Section, of the University of Uppsala to be publicly discussed in Lecture Room X, on May 24th, 1950, at 10 o'clock, for the degree of Doctor of Philosophy. In addition to this volume, "Karlvaxtfloran i sydvastra Lule Lappmark", printed in the Acta Phytogeographica Suecica No. 28, as a second part of Floristic Phytogeography of South-Western Lule Lappmark, will be publicly discussed.
UPPSALA 1950
ALMQVIST & WIKSELLS BOKTRYCKERI AB
ACTA PHYTOGEOGRAPHICA SUECICA 27
FLORISTIC PHYTOGEOGRAPHY OF SOUTH-WESTERN LULE LAPPMARK
(SWEDISH LAPLAND)
I
BY
STEN SELANDER
UPPSALA 1950 .ALMQVIST & WIKSELLS BOKTRYCKERI .AB PRINTE D WITH CO NTRIBUTION FROM
LA NGMANSKA KULT URFO NDE N CONT ENTS
Page Page
Preface...... 5 Bicentric species . . � ...... 86 I. HISTORY OF BOTANICAL EXPLORATION 7 Scandinavian plants in Spitzbergen, Iceland and Greenland ...... 89 II. SURVEY OF THE REGION INVESTI- Remaining, probable or possible, sur- GATED ...... 18 vivors ...... 90 District I...... 18 Postglacial eastern immigrants.. . . . 93 District II ...... 21 . . . Postglacial southern immigrants . . . 96 District III ...... � ...... 23 . Summary ..·...... 97 District IV...... 25 . . . VI. MIGRATORY TRACTS ...... District V...... 27 99 District VI...... 29 Glacial survivors ...... 99 District VII ...... 30 . . . . . Lowland plants west of the moun- District VIII ...... 31 tain range...... 100 Lowland plants east of the moun- Ill. HUMAN INFLUENCE ...... 34 tain range ...... 102 Settlers and settlements ...... 34 Lapps and reindeer ...... 35 VII. DISTRIBUTIONS WITHIN THE REGION 108 Lumbering of the Lapps ...... 39 Old and new findings of certain spe- cies ...... 108 IV. .ALTIDUDINAL BELTS ...... 45 Glacial survivors as apophytes . .. . . 111 >>Regio alpina descensa>>...... 45 Stability of plant communities ... . 114 Regio alpina inferior and regio alpina The flora on unstable ground . . .. . 119 media ...... 52 The vegetation on >mew soil» ...... 123 V. ORIGINS OF THE FLORA ...... 57 >>Fossil>> ranges ...... 128 Fossil finds ...... 57. . . Bedrock...... 133
The Fennoscandian flora and the flora Topography ...... 140
of the Alps...... 58 Climate ...... 158 Endemics ...... 63 VIII. CLIMATE OF THE ICE-AGE REFUGIA 162 The amphi-Atlantic element ...... 63 Discussion of the amphi-Atlantic spe- IX . BASOPHILY WITHIN THE ALPINE FLO- cies...... 71 RA OF FENNOSCANDIA ...... 168
The Alpine-North European element 80 Literature Cited ...... 181 Areas with a gap east of the Scandes 81 Index to Phanerogams and Pteridophytes 192
PREFACE
In 1844, Professor P. F. WAHLBERG wrote: and some lowland plants, e.g., Corydalis faba >>In a tract which has been visited by several of cea; for the rest, the expansion gaps here are our Land's most outstanding botanists for many probably real. The conifer region to the east, years, as is the case with Lulea Lappmark, par on the other hand, certainly supports several ticularly the surroundings of Qvickjock, there na other species than those now encountered: Poten tural(v remains very little that is new to be ob tilla rnultifida, Viola Riviniana, and Galium tri served among the plants>> (translated from the florum, especially, should occur somewhere on Swedish text). Even at that time, the moun the south-bluffs. In addition, there are species tains of south-west Lule Lappmark were, appa concerning the presence of which there are older rently, considered to be botanically very well and unconfirmed, though not improbable, re known. However, in the course of a trip from ports. The picture of the flora as a whole, how Kvikkjokk to Vaisaluokta made by Dr NILS ever, is scarcely likely to be altered by such ad DAHLBECK and myself in August, 1939, we found ditions. to our astonishment that large parts of the area Less satisfactory by far is our knowledge of were entirely unexplored. I therefore conceived the frequency of the various species. The v:ast the idea of attempting to make a botanical in uninhabited and trackless areas that can be tra ventory to fill the gap between the areas inves versed on foot only, so that much valuable time tigated by TENGWALL and BJORKl\fAN within the must be sacrificed on transport, have rendered National Parks of Sarek and Stora Sjofallet, on it impossible to make more than a somewhat the one side, and those investigated by ARWIDS cursory inspection, mainly restricted to the more SON and WISTRAND in Pite Lappmark, on the interesting parts. Particularly in the primeval other. forests of the conifer belt, only the surroundings My field research had to be done in my holi of the few paths could be investigated. Our know days, from the 1st of July to the 15th of August, ledge of the flora is relatively complete only with each year from 1941 to 1944 and in 1946, and regard to about ten, mostly rather small moun in shorter periods during the two subsequent tains to the south and south-east of Lake Viri years. In 1945, I suffered an accident which haure. At least two or three more sun1mers would compelled me to abandon the expedition I had be needed to fulfil the task I had set myself. just commenced. The consequences of this acci During my field work, though partly not until dent obliged me to discontinue my investigations it was almost completed, my attention was direc although certain tracts are as yet partially or ted to certain phytogeographical problems with wholly unexplored. a wider application than the purely local. When The flora at least of the western mountain their real nature became apparent to me, it was, tracts should, however, be rather exhaustively however, to late to change the plan of n1y in known. One may further expect to discover only vestigation. The discussion of these problems in Antennaria Porsildii and, possibly, Poa laxa ssp. Part I of this work is, therefore, chiefly foun flexuosa, Papaver radicatum, Sedum villosum ded on observations made for entirely different 6 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
purposes. Though fully conscious of these draw AND.ERSSON of Sagudden, and PER PmAK of the backs, I have, unfortunately, been unable to I.Japp village of Tuorpon. supply the deficiencies. Above all, I am indebted to Dr NrLS DAHL Part II, which deals with the local plant topo BECK, my highly esteemed companion during the graphy and is, for practical reasons, published as summers of 1939, 1943 and 1945 and, for a short a separate volume, has been written in Swedish, time, in 1946. That the consequences of my acci largely because of the difficulty of finding ade dent in 1945 did not become more serious was quate English words for the specific types of entirely due to his endurance and presence of ground in Lapland that are mentioned in the aut mind. In addition, he has generously provided ecological notes. However, this section can hard me with the notes he himself made during his ly be of particular interest to non-Scandinavian journeys in these parts. readers. The distribution of the various plants Various annotations have been kindly placed should be · clear from the maps; taxonomical and at my disposal by Dr ARVID FRISENDAHL, Dr nomenclatural observations that may be of a SVEN G:soN BLOMQVIST, Mr GuNNAR BRODAL general interest are given in the English sum� of Trondheim, and Mr OLOF RUNE. Mr �SKAR mary. LONNQVIST of Overtornea has been good enough
* to allow me the use of his unpublished investi gations of the flora of N orrbotten. Further, 1\'Ir GUNNAR KAUTSKY has obligingly informed me That this work is not still more incomplete is of certain results, likewise unpublished, from his largely due to my fellow-travellers. Many of the geological survey of the Vastenjaure-Virihaure most interesting discoveries were made by them, basin. I offer my sincere thanks to them all for and I regret that lack of space has made it im their kind generosity. possible to record these facts in the list of plant Professor ERIC HULTEN not only placed a room stations. at the Riksmuseum at my disposal, but also kind I had the great privilege of Professor 0ARL ly permitted me to study his as yet unpublished, SKOTTSBERG's company during the summer of cartographical account of the distribution of vas 1942 when, after an absence of nwre than 40 cular plants in Fennoscandia, thus sparing me years, he returned to the neighbourhood of Sarek many errors. where, as a young student, he had collaborated Dr ERIK ASPLUND has assisted me in deter with AxEL HAMBERG. In 1944, I was accom mining several critical plants. Taraxaca are deter panied by Mr OLOF RuNE, and in 1946, by mined by Dr GUSTAF HAGLUND, Hieracia mainly Mr OLLE MARTENSSO . Professor HuGo Os by Mr THORILD FOLIN, Sltlices largely by Mr VALD, Professor BERTIL LINDQUIST, Dr GuNNAR STEN GRAPENGIESSER and Mr 0ARL BLOM, some DEGELIUS, Dr WILHELM RASCJH, and Mr GUSTAF Poae by Professor J. A. NANNFELDT, certain Al VON HOFSTEN have also accompanied me for chemillae by Mr M. ENGSTED'.r, and several Betu shorter periods. To them all, I wish to convey lae by Professor BERTIL LINDQUIST. My perusal my thanks for their generous help and good fel of the herbarium at the University of Upsala lowship under often rather trying conditions. I has been supplemented by Mr OLOF RuNE. l\ir also wish to thank Mr GosTA LUNDQUIST, Mr MATS W..ERN, . editor of the Acta Phytogeogra KARL-AAGE SCHWARZKOPF, Mr BENGT ROTH phica, has spared no pains to give this book an STEIN and n1y son, TOM SELANDER, whose attractive get-up; and Mr MAGNUS LUNDQVIST, support, though less authoritative botanically, Mr SVEN ERIKSSON and Mr REINHOLD IVARS has not on that account been less appreciated. SON have taken much trouble about the maps. Nor do I want to forget my assistants and, if I greatly appreciate their kind interest. I may say so, my friends, those excellent moun Among those with whom I have had the pri taineers BORJE DANIELSSON of Kvikkjokk, ERIK vilege to discuss my work, I first wish to men- Preface 7 tion my old friend Professor G. ErNAR Du RrETZ, generous grants for field research and for the ex without whose initiative and encouragement it penses of the printing of maps and illustrations. would scarcely have come into being in its pre sent form, and whose stimulating, positive cri * ticisms have been invaluable to me. The other botanical friends whom I have troubled with my problems are too nun1erous to enumerate. In With a few exceptions, the nomenclature of addition to Dr NILS DAHLBECK I must, how Fennoscandian vascular plants is in accordance ever, mention Professor BERTIL LINDQUIST, who with HYLANDER 1941 and 1945; for specific Ice has perused my manuscript and given me his landic species, and also in some other cases, I views, and Dr AsKELL LovE of Reykjavik, who have followed A. and D. LOVE 1948. For further with inexhaustible patience has replied to my details, see Part II. For the bryophytes I questions concerning the flora of Iceland. Dr have followed JENSEN 1939. The phytogeogra GosTA LUNDQVIST has gone over the descrip phical terminology is essentially in agreement tion of the area of exploration in Chapter II, with CAIN 1944. and Dr ERNST .1\fANKER has corrected the .As far as possible, data concerning the extra Fennoscandian ranges of the plants have been map fig. 21. While thanking them, I remain, of course, alone responsible for any mistakes that taken from HULTEN's different works, as these have been overlooked. seem to me to occupy a class of their own among My former teachers in Botany, first and fore such publications, frornthe point of view of com most my revered and unforgettable friend Pro pleteness and reliability. I regret that except in fessor RUTGER SERNANDER, are, I regret to say, a few cases, time has not permitted me to pro beyond the reach of any words of mine. So is cure my material of such data from monographs and other special inv�stigations. also Dr T'noRSTEN .ARWIDSSON, who unfailingly helped me in word and deed. Stockholm, February 5th, 1950. Finally, it is my pleasant duty to express my gratitude to Langmanska Kulturfonden for their Sten S elander.
PLATE 1
Primeval pine forest on the till plains between Lake Saggat and Lake Peuraure. Ground vegetation consisting mainly of Vaccinium Myrtillus and Ledum palustre. Photo Olof Rune July 1944. PLATE 2
Primeval spruce forest with Sorbus aucuparia on the southern :;:lope of Mt Tarfek. Photo Olof Rune July 1944. PLATE 3
The "glint" on Mt Tariek. The clay-schists o£ the eastern cambro-siiurian crop out in the precipice. On the talus slope in the foreground one of the northernmost Swedish stations of Carex ornithopoda and 'l'U?-ritis glabm. Photo Olof Rune July 1944. PLATE 4
South-bluff o£ Mt Tjalta. The northernmost Swedish station of Stachys silvatica and Dryopteris spinulosa, both of which occur abundantly together with, i.a., Aconitum septentrionale, Valeriana sambttcifolia and Daphne Mezereum. Photo Nils Dahlbeck 5.8. 1946. PLATE 5
Primeval birch forest in the regio subalpina of the valley of Tjuoltavuopme. Photo Nils Dahlbeck August 1943. PLATE 6
Birch wood in the Tarra valley. In the background Mt Staika with a small glacier between its two peaks. Photo Nils Dahlbeck August 1939. PLATE 7
Sloping Scirpus caespitosus- fen on Mt Katnjunjes. In the background the Tarra valley. Photo Olof Rune July 1944. PLATE 8
Trollius europaetts- meadow in the lower regio alpina, north-eastern slope of Mt Katnjunjes. Photo Olof Rune July 1944. PLATE 9
The uppermost birch wood on the mountains north of the Rittak valley. Photo Sten Selander 6.8.1946. PLATE 10
Padjelanta is a tegion of low mountains. View to the east from Mt Allak. In the foreground the river M:ellatno. Photo Nils Dahlbeck August 1939. PLATE 11
" Recently formed nunatakk" on the glacier of Almallojekna, seen from Mt Kasakpuolta. On the raw gravel in the foreground no phanerogam vegetation but Saxijraga rivularis. The "nunatakks" were completely devoid of phanero gam vegetation. Photo Sten Selander 26.7 .1946, PLATE 12
Mt Sulitelma in the height of the summer. View from Mt Jeknaffo, c. 1400 m s.m. In the foreground habitat of Potentilla hyparctica. Photo K.-A. Schwarzkopf 29.7. 1944. I. HISTORY OF BOTANICAL EXPLORATION
The mountains around K vikkjokk and Lake IWRT is cited tHARTMAN op. c. p. 51), we may Virihaure form a classical realm of Swedish bo safely infer, however, that they were made by tany. During the 18th and the first half of the RUDBECK himself. 19th century, most leading men among our flor RUDBECK's visit to Lapland was mainly im ists, from 0LOF RUDBECK THE YOUNGER and portant because it stimulated LINNAEUS to un LINN.AEUS to WAHLENBERG, LAESTADIUS and N. dertake his famous journey. On July 12th (N.S.) J. ANDERSON, took part in the investigation of 1732, LINNAEUS crossed the boundary of our re this region. RUDBECK was a n1ember of a sci gion at Tjamotis and visited Mt Kiuri on the entific expedition to Lapland, sent out by CHAR 14th, reaching Kvikkjokk on the 16th (LINNAEUS LES XI in 1696. His collections and notes were, 1913). Next day he ascended the eastern part however, destroyed in the disastrous conflagra of Mt Vallivare (by which name the earlier tion of Upsala in 1702, so that he could not authors did not mean only the present Mt Valli� publish anything but a bare list of the plants vare, but all the Tarrekaise massif), from there observe.d (RUDBECK 1720); and because he did proceeding westwards, no doubt through the val� not visit only Kvikkjokk, but the mountains on ley of Ruonasvagge. A detailed account of his Lake Torne Trask too, this list is of little use as further route is hardly possible to give. In his a contribution to the flora of Lule Lappmark. diary LINNAEUS nwntioned only three places on Yet, the monumental figure of RuDBECK does the Swedish side of the frontier, viz., Lake Viri not play merely a decorative role in the botani haure, Mt Kaitsanjunje, and Lake Kaitum; n10st cal history of these parts. Besides the list of certainly, however, he did not visit Kaitumjaure. plant names, he left behind him a series of ex Yet for divers reasons, set forth elsewhere by cellent plant drawings, made during his trip to the present author (SELANDER 1948), it seems Lapland and now kept in a private collection reasonably sure that he went from Ruonasvagge (cf. HARTMAN 1841). From notes on a few of down to Lake Tarreluoppal and from there, over these drawings, we know them to originate from Mt Karranis and south of lVIt Silpatjakko, our region. This applies to Sedum Rosea (1;Sponte through the valley of Vejevagge to Lake Viri crescit in Wallawari monte altissimo Laponiae haure. By way of Mt Kaitsanjunje, west of I....�ake Lulensis prope Quickjock1;, cited by HAI<.TMAN, Virihaure, and the glacier of Flatkj0len, he then op. c. p. 69), Gentiana nivalis (1;in alpibus Lap. descended to T0rfjord on the Norwegian coast. Lulensis1;, ibid. p. 54), Pedicularis hirsuta (1;in His way back is impossible to reconstruet. We Lapponia Lulensh, ibid. p. 69), and Petasites fri know that he went by boat over Lake Virihaure; gidus (1;in nivato Vallavare Lap. Lul.1;, ibid. p. then he apparently erossed the table-land of 54). We can, probably, also refer Pedicularis Padjelanta. The lake mentioned in his diary, lapponica to this category (1;in Lapp. Lul.1;, ibid. under the name of Lake Kaitum, may have been p. 69). True, the descriptions of HARTMAN do I.1ake Alkajaure; and it seems rather eertain that not plainly state from whose hand the annota he went baek to Kvikkjokk, where he arrived tions derive; since no author later than TOURNE- on July 31th, by way of Ruonasvagge.
2- 496149 Sten Selander I 10 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
The question where LINNAEUS was on a given Saxijraga oppositifolia Loiseleuria procumbens day is not as insignificant as it may appear. S. stellaris Phyllodoce coerulea S. foliolosa Arctostaphylos alpina H� noted in his diary when he found a new plant. S. aizoides Primula st1·icta (veris.) Therefore, it depends on the answer to this ques S. nivalis Trientalis europaea tion where we have to look for the type localities Rubus Ohamaemorus Gentiana nivalis (at least if we do not take this concept in too Sibbaldia procumbens Veronica alpina Alchemilla alpina Pedicularis Sceptrum�ca- formal a sense) of a number of Linnean speciel-3, Dryas octopetala rolinum e. g., Juncus triglumis, Melandrium apetalum, Ra Astragalus alpinus P. lapponica nunculus pygmaeus, R. nivalis, Draba alpina, Car Geranium silvaticum Pinguicula alpina damine bellidifolia, Saxifraga cernua, S. rivularis, Viola biflora Solidago Virgaurea Rhododendron lapponicum, Cassiope tetragona, Ohamaenerion angustifo- Erigeron politum (veris.) Q hypnoides, Diapensia lapponica, Pedicularis lium E. �miflorum Angelica Archangelica Lactuca alpina hirsuta, P. flammea, Carnpanula uniflora, and Arnica alpina. This applies also to the first Swed However, we may infer, with reasonable cer ish stations of many other alpine plants. Now tainty, that LINNAEUS further noticed a consi we can define these localities only regarding spe derable number of plants not explicitly repor LINNAEUS cies found by during the early stages ted by him from our region. After leaving K vikk of his journey, for instance, on Mt Vallivare or jokk, he ascended nowhere above the conifer belt; Mt Kiuri. In regard to the remaining species, and no botanist visited the Lapland mountains we can only say that they were found some during the years between 1732 and the publish LINNAEUS where within our region, which cer ing, in 1737, of his Flora Lapponica. Therefore, tainly did not leave during his stay in the Swe it is evident that the alpine plants mentioned dish mountains. there were observed by LINNAEUS himself in the In addition to the species enumerated above, Kvikkjokk region. The same conclusion may he LINNAEUS reports a great number of plants, arrived at regarding the species reported to occur part new, part described by somebody else or 1>in alpibUS>>, >>per totam Lapponiam>>, &c. Thi:::; found by himself outsid� our region, as occur applies to the following species: ring in the Kvikkjokk district. They appear usu ally under unfamiliar names, which cannot Lycopodium annotinum Eriophorum vaginatum L. alpinum E. angustifolium always be identified to a certainty; the inter Selaginella selaginoides Scirpus Hudsonianus pretations of TH. �1. FRIES in his edition of Equisetum arvense Se. caespitosus the Iter lapponicum (LINNAEUS 1913) are often E. silvaticum Oarex paucijlora erroneous. These plants are the following: E.. limosum 0. rostrata Athyrium Filix-femina Juncus f'iliformis Lycopodium Selago Polygonum viviparum Oystopteris fragilis Luzula pilosa Poa alpina v. vivipara Stellaria nemorum Lastrea Phegopteris Maianthemum bijolium Oarex atrata Oerastium trigynum (ve- L. Dryopteris Salix hastata Juncus arcticus ris.) H ierochloe odorata Alnus incana J. . trijidus C. alpinwrn Phleum commutatum Rttmex Acetosa Luzula m·cuata (veris.) 0. glabratum Deschampsia alpina (first 0altha palustris L, spicata M inuartia biflora mentioned in LINNJEUS Trollius europaeus Tofieldia pusilla Viscaria alpina 1745) Ranunc�tlus acris Salix reticulata Silene acaulis D. caespitosa R. repens S. her·bacea M elandrium rubrum D. jlexuosa Cardamine pratensis S. glauca (veris.) Thalictrum alpinum Tr�setum spicatum Parnassia palustris S. phylicijolia (veris.) Aconitum septentrionale M olinia coerulea Filipendula Ulmaria S.lanata Ranunculus glacialis Poa pratensis Prunus Padus S. lapponum Draba norvegica ( veris.) P. alpina Rubus arcticus Betula nana Arabis alpina (veris.) Festuca ovina R. saxatilis Oxyria digyna Sedum Rosea Nardus stricta R. idaeus History of Botanical Exploration 11
Potentilla palustris Pinguicula vulgaris was, nevertheless, a mistake, since the list of P. Crantzii Linnaea borealis plants manifestly shows that the mountain re Callitriche verna Valeriana sambucijolia ferred to was Arasvare at the mouth of the river Co1·nus suecica Campanula rotundijolia Ledum palustre A ntennaria dioeca Mellatno. How Sw ARTZ managed to get there Andromeda polijolia A. alpina I do not know; it is only certain that he started Arctostaphyles Uva-ursi Petasites frigidus from Kvikkjokk, and was back late in the sum Calluna vulgaris Saussurea alpina mer. New to our region among his discoveries Empetrum hermaphrodi- Cirsium heterophyllum were Carex norvegica, Chamorchis alpina, Leu tum Hieracium alpinum Bartsia alpina chorchis albida and the nondescript Minuartia �tricta {SWARTZ 1799, 1821). In the year 17 49, LINNAEUS sent one of his In 1807, G. W AHLENBERG performed the first >>apostles>>, L. J. MONTIN, to thes� parts. MaN investigation of the flora of Lule Lappmark of a TIN's diary, now at the British Museum and not modern phytogeographical character. Thanks to accessible to me, was never published, and his his diary, now in the Library of the University collections, now partly in the Herbarium Hol of Upsala, and to the routes marked on his sketch miense, usually lack specifications as to the sta map {WAHLENBERG 1808 tab. 1), we are able to tions of the plants. For these reasons, we know follow his journey day by day. On his way to for certai� only that 1\'IONTIN visited K vikkjokk Norway, Wahlenberg crossed the mountain range and Virihaure; it seems probable, however, that already in the beginning of May; but he did not he took the same course as his teacher. His con return to Lule Lappmark until July 25th, when tributions to the flora of our region consist, as he passed the Swedish frontier at the place of far as we know, of Festuca vivipara, ,]uncus bi the present >>riksroset>> (boundary-cair-n) No. 2ln .. glumis, Salix myrsinites, Saxifraga Cotyledon and He made then for the Lapp camp, situated at S. g'roenlandica, most of them new to Science as that time below Mt Sirkavare (>>Stor-Thokin>>), well as to the Swedish flora. where he had his head-quarters until August Around 1750 J. HoLLSTEN, >>pastor Quickjock 13th. During these weeks, he explored the moun sensis>>, botanized within his parish, sending some tains of Kaitsanjunje (>>Kaisatj>>), Sirkavare, Tuki plants to :MoNTIN, i.a., Pinguicula villosa, which (>>Lill-Thokin>>) and Kuobberi (>>Kobrinoiwi>>) and had not earlier been noted within the region. their surroundings. Here he discovered Carex Concerning the Lapland journey of 0. SWARTZ glacialis, Arenaria humifusa, Saxifraga tenuis and in 1780, I have not been able to hunt out any Oxytropis lapponica, all undescribed, and Cystop definite data. In his biography (SWARTZ 1829 teris montana, Kobresia myosuroides, Carex micr() p. xxxn), WIKSTROM only writes as follows: glochin, Draba lactea, Astragalus norvegicus and >> t.Jber diese Reise, welche sich bis zum Arras Gentianella tenella, which were not previously Fjall und zum Wirihjaur hinauf erstreckte, fii.hrte known in Sweden. SwARTZ ein Tagebuch, worin er die Gegenden und On July 29th, he rowed over Virihaure to Mt dessen Produkte so wie die Lebensart der Lapp Arasvaretjakko in order to search for Chamor Htnder beschrieben hat.>> This diary is unpublished chis alpina, which he believed SWARTZ had found but for a >>Flora for tragten kring berget Arras>> there; and on August 13th, he travelled, like (Flora of the Neighbourhood of Mount Arras), wise by boat, to Staloluokta on the sout.h-eas appended to the commemoration speech that 1\'I. tern shore of Virihaure. From there, he made AF PoNTIN made in 1820 to the members of the an excursion to Lake Sarjasjaure and the gla Academy of Sciences (SWARTZ 1821). WAHLEN cier of Almallojekna, following the western and BERG, who had obtained private information northern bank of the river Stalok; besides, he from Sw ARTZ, believed the name >>Arras>> to mean explored Mt Jalle on Vejejokk and, evidently Mt Arasvaretjakko on the southern shore of Lake less carefully, Mt Kerkevare and the northern Vastenjaure (cf. the diary of WAHLENBERG). This slope of Mt Jeknaffo. During these excursions, he 12 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
found the undescribed Luzula Wahlenbergii and eluded in the notion of >>Lapponia>>). I am re- Poa arctica and Primula stricta in their first £erring to: definitely established, Swedish stations. Lycopodium clavatum Carex magellanica On his journey to Kvikkjokk, started on Au Equisetum pratense C.limosa gust 18th, he passed >>Juobba fjaJh> �Mt Junkar E. palustre C. capilla1·is tjakko) and went through the valley of Ruonas Sparganium angustifoli C, rotundata um (sub Sp. natans) C. la.
In addition to the species mentioned above, -1845, is the first and hitherto only local lnora LJESTADIUS found the following ones, not earlier of the Kvikkjokk mountains. It is founded on reported from these parts (cf. LJESTADIUS 1822, investigations, made during the summer of 1843 1824, 1839): by ANDERSON, in collaboration with the eminent zoologists, Professor P. F. WAHLBERG, Professor Agrostis canina Luzula par-viflora C. H. BOHEMAN, and C. G. LOWENHIELM, B. A. Deschampsia atropurpu- Oorallor-hiza trifida rea Salix polaris This exploring was, however, limited to the im Poa glauca Silene rupestris mediate neighbourhood of Kvikkjokk, ANDER P. nemoralis Nymphaea candida SON hardly visiting more mountains than Snje P. palustris Ranunculus lapponicus rak, Vallivare, Kaskaivo, Njunjesvare, Tarre Oar-ex tenuiflora Epilobium collinum (sub kaise, Nammatj and Lastak. He returned in 1845, Juncus balticus E. montanum) J. alpinus Oircaea alpina accompanied by W A�BERG and LOWENHIELM J. stygius Er-iger-on boreale (cf. ANDERSON 1846). This time he came from t,he south, having traversed the mountains of The elder brother of LJESTADIUS, 0ARL ERIC, Pite Lappmark, crossed Lake Peuraure on July who was vicar of Kvikkjokk, found in 1818 Koe 6th, and went on over Mt Lastak to Kvikkjokk, nigia islandica in its first Swedish locality sonle where he stayed until the 23rd. On this date, where near Kappaluoppal (Svensk Botanik 1819). he started for Virihaure in the company of Lo In the eighteen-twenties and eighteen-thirties, WENHIELM. Their route lay through Ruonas J. UI"LENrus, vicar of Jokkmokk, botanized in vagge, along the river Vassjajokk to Mt Alka these parts, from where he supplied LJESTADIUS vare, through the valley of :1\fellatno to Virihaure and N. J. ANDERSON with >>adnotationibus co and by boat to Sirkaluokta. After excursions piosissirnis et egregiis>> (ANDERSON 1844 p. 6). to the adja,cent mountains, viz., Kaitsanjunje, Only exceptionally, his reports are confirmed by Tuki and Sirkavare, they returned by way of specimens, which have found their way to our Staloluokta, where they ascended the neighbou public collections via the herbarium of LJESTA ring Mt Kerkevare. During his last visit, in 1864, DIUS. This is regrettable, i.a. because Mentha ANDERSON made excursions to the Virihaure re arvensis, reported by him, has not later been gion, visiting i.a. Mt J eknaffo and the glacier found within the region. Definite finds by ULLE of Almallojekna, and to Aktse by way of Parek NIUS are Phippsia algida, Car ex ornithopoda, Vi (ANDERSON 1866). ola rnirabilis, V. montana, and Platanthera bifolia, Among the species discovered by ANDERSON which has not been found again in the station within our region, Eriophorum medium was first (Mt Lastak) reported by him , though recently described by him, and Car-ex ru.fina, Cerastium noticed on the adjacent Mt l!'arforita. arcticum and Sagina intermedia were not pre In order to collect specimens for the Herba viously known in Sweden. His other new reports Tium Nor-male of FRIES, J. ANGSTROM travelled were rather numerous, since he was the first to in 1837 through Ruonasvagge to Virihaure. On collect and note down methodically also >>corn a detour to Mt Sakok, he found Bray a lineaTis; man>> species. Of course, quite a number of these and in the mountains to the south of Virihaure, plants had already been observed here by other he discovered Carex nardina, then undescribed, botanists, though we have no proofs of this. Any and the first Swedish station of ArenaTia norve how, the first indubitable reports or specimens gica, that he mistook, however, for A. hurnifusa. of a lot of species originate fron1 ANDERSON, viz.: His other principal finds were Botrychium Lu naria, Stellaria crassifolia, and Tussilago farfara Isoetes echinospora Woodsia ilvensis Equisetum hiemale Dryopteris Filix-mas (ANGSTROM 1839). Botrychittm bor-eale Potamogeton pusillus N. J. ANDERSON's >>Plantae vascula.res circa Asplenium viride P. alpinus Quickjoek Lapponiae Lulensis>>, published in 1844 Athyrium alpestre P. perfoliatus 14 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Phragmites communis Drosera rotundifolia tanizing in Lule Lappmark during the later half M elica nutans Dr. anglica of the century. Nearly all of them followed the Roegneria scandica Viola epipsila Eriophorum gracile V. palustris standard route through Ruonasvagge and _past Carex macloviana Epilobium davuricum Lake Festajaure and lVI t Kmkevare to Virihaure, C. brunnescens E. anagallidifolium where the journey's end was invariably l\H Tuki C. loliacea E. lactiflorum (>>Lill-Tokin>>); occasionally, they made digres C. disperma E. Ho1·nemanni sions to Mt Sakok or Lake Laidaure. It seems C. adelostoma A nthriscus silvestris therefore unnecessary to deal here with their tra C. Oederi Pyrola grandiflora C. flava Ramischia secunda vels, although some of them, besides being skilled C. vesicaria Myosotis silvatica collectors, proved their plant-geographical sense Paris quadrifolia Primula scandinavica by noticing >>common>> southern species as well Listera coTdata Prunella vulgaris as rare alpine plants. Among these botanists, Salix borealis Veronica pumila the following ones found plants not earlier known S. caprea V. serpyllifolia S. Starkeana V. tenella in the region: S. myrtilloides V. scutellata A. DAHLBERG (apparently a backwoodsman, S. pentandra V. officinalis collecting for N. J. ANDERSON during the years Stellaria graminea Utricularia intet·media 1850-1859) Carex caespitosa, Sagina Norman'i St. longifolia Galium trifidum ana; E. SANDSTROlVI Draba nemorosa; E. C. St. calycantha G. palustre Nuphat· luteum x pumilum Erigeron unalaschkense J. CEDERSTRAHLE (1856) Lycopodium compla Rosa majalis Leontodon autumnalis natum, Arabis hirsuta, Turritis glabra; A. HEIN Potentilla er·ecta Orepis paludosa RICI �1859) Potentilla. argentea; E. AlmLING & A nthyllis V ulneraria Hieracium prenanthoides M. G. BRANDELIUS (1859) Eriophorum brachy WICHURA, who made the usual trip to Viri antherum, Elatine Hydropiper, Hieracium lappo haure in 1856 and found, i.a., Montia rivularis nicum, probably Blechnum Spicant; A. FREDRICS and Actma erythrocarpa (new to Sweden), was the SON (1864) Carex pallescens, C. panicea, Urtica first and, up to the present, only foreign bota dioeca ssp. gracilis, Rumex tenuifolius, Epilobium nist to visit these mountains (WICHURA 1859). alsinifolium; and INDEBETOU & JUHLIN (1868} After him, no professional botanists came here Galium boreale. for nearly half a century, the exception being Other principal collectors of this period, cited V. :F. HoLM, who set out in 1873 on a journey in the list of plant stations in Part II, are E. from Kvikkjokk past Mt Staika to Pite Lapp J. vVIDMARK (1867 and 1870), who made exten mark; fron1 here he went by way of Lake Rave sive journeys, i.a., to the unexplored neighbour jaure, Vejevagge and Staloluokta to the moun-. hood of the river Vuojatatno, E. A. G. KLEEN tains beyond l.Jake Vastenjaure and further (1868), A. Go:Es (1872), and ALlVI & REUTER northwards, visiting on the way, i.a., the penin SKIOLD (1879). sula. of Titernjarka (cf HoLM 1875). The reports In the eighties, these collecting expeditions by HoLM are, however, often unprecise and at can1e to an end. Later visitors usually pursued times apparently erroneous; that is why I have purely scientific aims. Thus G. HAK.ANSSON thought it safest, as a rule, not to use them in (1882 ) studied the Salix flora around Kvikkjokk the list of plant stations in Part II of this_ work" and in the lower Tarra valley. In 1891 and 1893, unless confirmed by herbarium specimens. His E. NYM.AN investigated the ecology of the vege contributions to the flora of the region, viz., tation chiefly of the conifer belt arount Lake. Polygonatum verticillatum and Lotus corniculatus, Saggat (NYM.AN 1895). ASTRID 0LEVE (1896) car are rather insignificant too. The reputed find ried out ecological investigations around Lake. of Hierochloe alpina on Lake Ravejaure is, un Festajaure and on Mt Junkartjakko; however, doubtedly, a mistake. her paper (CLEVE 1901) also contains numerous All the more numerous were the amateurs bo- reports on phtnt stations. And in 1901, N. SYL- History of Botanical Exploration 15
VEN and G. BAGENHOLM travelled in the eastern flora of Lule Lappmark since the days of N. J. parts of the region, mainly in order to study the ANDERSON. TENGWALL devoted his time mostly anthropochores (SYLVEN & BAGENHOLM 1902); to the exploration of Sarek proper; however, in on that occasion, they found Limosella aquatica 1913, 1915 and 1916, he occupied himself also {BIRGER 1909). with our region (cf. TENGWALL 1914 and 1924). .Round the turn of the century, these parts During the first-mentioned year, he investiga were now and then visited by two forestry offi ted the mountains between Lake Vastenjaure and .eers interested in botany, viz. 0. VESTERLUND Lake Sallohaure, up to then completely unex (1891, 1898 etc.) and N. KR. BERLIN (at least plored, where he found, i.a., Agrostis tenuis, Poa 1904, 1906 and 1908), who gave valuable reports herjedalica, Epipactis atror'ubens, Alchemilla fili .especially from the woodland (cf. VESTERLUND caulis, A juga pyramidalis, Euphrasia lapponica 1892 and 1924, SrMMONS 1907b). During the same and Erigeron acre (as indigenous). In 1915, when period, TH. WoLF botanized in the vicinity of he noted, i.a., Sparganium hypeTboreum and Fes K vikkjokk; he furnished ANDERSSON & BIRGER tuca rubra v. mutica he took stock of the flora of {1912) with a list of the plants observed by him Padjelanta, especially on the mountains of Unna on Mt Nammatj. R. WESTLING collected round Rissavare, Stuor Rissavare, Iltom, Unna Titer, Kvikkjokk and in the lower Tarra valley in 1905. Stuor Titer, Lulep Alatjakko and Mattaive, and In 1892, H. V. RosENDAHL and, in 1906, H. made brief digressions to the mountains about SIMMONS made the traditional trip Kvikkjokk the lower course of the river Stalok. In the fol Virihaure and back. All of them gave accounts of lowing summer, he explored, i.a., the mountains their journeys tROSENDAHL 1892, WESTLING close to Kvikkjokk, among them Mt Sakok, and 1906, SIMMONS 1907a and 1907b), although they northernmost Padjelanta. The work of TENG hardly had any news to tell except WESTLING, WALL was performed with such a thoroughness who reported the finding of Listera ovata. SIM and penetration, that I have thought it unneces MONS, by the way, made some very queer state sary, once more to go over the parts where he ments, for instance, that Nardus stricta, Pyrola stayed for some time. Besides WAHLENBERG, minor and Campanula rot·undifolia should be lac LJESTADIUS, and ANDERSON, his name is the one king above the tree limit; that Pinguicula vul most frequently met with in my list of plant ga1·is should not >>exceed the birch region>>, above stations. which only P. alpina should occur; that Euphra ELISABETH EKMAN visited the mountains be sia tenuis should be >>common in the woodland>>; tween Kvikkjokk and Virihaure in 1912, mainly .and so on. TH� 0RTENBLAD established the oc in order to study their Drabae. During the sum ·Currence of Betula verrucosa (ORTENBLAD 1902); mers of 1921 and 1923, HJ. J\i oLLER stayed at and during a short visit in 1908, M. SONDEN Lake Tarraure for bryological studies; at the found Euph1·asia brevipila. same time, however, he brought together a large In connexion with the exploration of the Sarek and valuable material of vascular plants from mountains, organized by AXEL HAMBERG, the the adjacent mountains, up to then completely neighbourhood of Kvikkjokk and parts of Padje unexplored. In the twenties three salicologists, lanta were visited by C. SKOTTSBERG and T� viz. S. GRAPENGIESSER (1923), :M. ENGSTEDT VESTERGREN in 1900 and in 1901 by the latter (1926), and B. FLODERUS (1924 and 1928), wor alone, and the mountains around the river Rissa ked in the vicinity of Kvikkjokk, the last-men jokk, by the bryologists H. W. ARNELL and C. tioned extending one of his journeys to Virihaure. .JENSEN during the summer of 1902 (VESTER Especially GRAPENGIESSER interested himself al ·GREN 1902 , ARNELL & JENSEN 1907 1910). so in other plants than Salices, discovering, i.a., ' T. A. TENGW ALL was also among the collabo Asplenium 11richomanes; ENGSTEDT contributed rators of HAMBERG. In the years 1913-1918, Salix livida x xerophila and the previously over .he made the first metodical investigation of the looked S. coaetanea. In 1924, TH. ARWIDSSON 16 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
investigated the hydrophytes of the eastern lakes, Kables massif, the valley of Tjuoltavuopme and finding i.a. Potamogeton praelongus, and crossed its neighbourhood, and the mountains south and the southernmost part of the mountain range on south-east of Virihaure. The following summer, his way to Pite Lappmark (ARWIDSSON 1926 b, I worked together with 0. RUNE and, part of 1927). And C. G. ALM, who in 1926 and 1927 the time, with B. LINDQUIST in the cliffs north botanized around Lake Peuraure and in the of Lake Saggat, the delta of Anok, and the wood eastern woodlands, visiting, i.a., Mt Njanja, land between Lake Saggat and Lake Peuraure. where no other botanist has set foot, discovered Afterwards completory investigations were car Isoetes lacustris, Scheuchzeria palustris, Carex li ried out in the company of RUNE west of the vida and Lysimachia thyrsiflora. upper Tarra valley, and in parts of the areas vi In connexion with his investigation of the flora sited in the two preceding years. The excursion of the National Park of Stora SjOfallet and the of 1945 was broken off already after a few days surrounding region, G. BJORKMAN was, in 1929, spent near J...�ake Kutjaure. In 1946, I visited the first botanist to explore the mountains north the nwuntains north of J.ake Sallohaure and of Lake Sallohaure, where he found, i.a., Gentia Lake Vastenjaure, the peninsula of Kollomoloki, nella aurea in its first native Swedish station, Mt Kasakpuolta and the surrounding mountains, Scirpus pauciflorus, Sedum acre and Viola ru and parts of the plateau of Padjelanta in the pestris (BJORKMAN 1939). company of 0. M.ARTENSSON, and the Rittak val J. A. N ANNFELDT went by the standard route ley, the valley of Blackalven, and the region to Virihaure and back in 1936 in order to study around Tjamotis together with N. DAHLBECK. the Poa arctica races of the region, discovering In 194 7, I stayed for a few days at Mt Farfo on his way Roegneria m�ttabilis, Sagina caespitosa, rita and afterwards devoted some time to the and Minuartia rubella (NANNFELDT 1940, 1941). surroundings of Lake Saggat. And next sum During the same summer, A. FRISENDAHL made mer, my field research was wound up by a short an excursion, i.a., to the mountains around Mt visit to the woodlands west of J.. ake Karatsj. Silpatjakko and the valley of Vejevagge; among Every year except 1945 and 1948, 1 spent a week his finds was Eriophorum latifolium. The bryo or more in the neighbourhood of Kvikkjokk and logist A. HijLPHERS visited Lake Ravejaure in the Tarra valley. The new contributions of the 1937, also paying attention to the vascular present author and his companions to the flora , plants. And in the years 1935-1939, TH. FoLIN of the region consist of the following plants: carefully investigated the Archieracia of the Asplenium septentrionale Potentilla hyparctica neighbourhood of Kvikkjokk (FOLIN 1942). Dryopteris spinulosa Alchemilla vestita In the years 1939 and 1941 to 1948, the pre Lastrea Robertiana A. norvegica sent author took stock of the vascular plants Sparganium minimum Callitriche polymorpha of the region, as a rule staying for 5 to 6 weeks Sp. simplex Epilobium montanum Potamogeton filiformis M yriophyllum spicatum annually in these parts. The first summer I Agrostis stolonifera Pyrola chlorantha devoted to a general reconnaissance in the corn Poa irrigata M yosotis stricta pany of N. DAHLBECIL In 1941, I investigated Carex globularis Stachys silvatica the mountains to the west of the Tarra valley C. Buxbaumii Utricularia vulgaTis and around Staloluokta, the river Stalok and the Convallaria majalis U. minor Sagina nodosa Galium uliginosum westernmost coves of Virihaure, partly together . D1�aba crassifolia A ntennaria lapponica with H. OsvALD, G. DEGELIUS and G. VON Cardamine pratensis ssp . H ieracium Pilosella HOFSTEN. 0ARL SKOTTSBERG was my companion angustifolia in 1942, when we explored the region round Lake Ravejaure and the valley of Staddavagge, and Among these plants, Antennaria lapponica was the rrwuntains south of Lake Sarjasjaure. In undescribed, and Potentilla hyparctica new to 1943, N. DAHLBECK and I visited parts of the Fennoscandia. Nor had Oardamine pratensitt History of Botanical Exploration 17
• Uninvestigated areas 0 Camp for one night
• Camp for two nights ® Camp for three or more nights '
Fig. l. Cumps of the present author. The flora is fairly exhaustively investigated only near the camps for two or more nights. ssp. angustifolia earlier been noticed in Scandi lichen flora especially in the mountains south navia (SELANDER 1942, 1945 a). of Virihaure (DEGELIUS 1943), and in 1946, 0. N. DAHLllECK also has made journeys within MARTENSSON studied the moss flora around Sal the region by himself, namely in 1940 from Kvikk lohaure and Virihaure. Shorter excursions, re jokk by way of Lake Vaimok and Lake Rave ports from which are included in Part II of the jaure to Virihaure, where .Arenaria humifusa was present book, were made, i.a., in 1943 by W. found again after more than a hundred years, RASCH (Kvikkjokk-Virihaure), and in 1946 by and in 1942 , i.a., in northern Padjelanta and the S. G:soN BLOMQVIST (to Mt Tjargesvare). In mountains south-west of Lake Peuraure. He 1946 and 1948, B. ABERG studied the altitude found i.a. Botrychium multifid'ltm (DAHLBECK limits of the vascular plants (ABERG 1949), and 1940). B. LINDQUIST, having discovered the non in 1949, 0. RuNE investigated the flora of the descript PifJea .Abies var. arctica and the forgot serpentine mountain Ruopsok. Ultimately, stray ten Betula callosa within the region in 1944, revi reports in the list of plant stations in Part II sited K vikkjokk the following summer for stu come from other botanists, whose names are dies on Betula (LINDQUIST 1945, 194 7 a and specified on each occasion. 1948). In 1941, G. DEGELIUS investigated the
3-496149 Sten Selander I Il . SU RVEY OF THE RE G ION INVESTI GATED
The region investigated is located on the Nor The boundary line then follows the rivers Tau wegian border just north of the Polar circle, be rejatno and Blackalven to Tjamotis and from tween 66°45' and 67°35' N and 15°06' and 18°36' there goes due south to l.Jake Karatsj, where it E (cf. fig. 2, 3); on the ordnance map, it is found turns off westwards along the lakes of Karatsj , on the sheets Kvikkjokk, Staika, Stora Sjofal Lillselet, Storselet, and Peuraure to the valley let, Sulitalma, and Akka. According to a com of Parkajokk; from there, the boundaries of the putation, based upon the ordnance map and region coincide with those of Pite Lappmark made by A. B. Kartografiska Institutet, Stock and Norway. holm, it comprises 4079 km2, the lakes inclu The uninvestigated mountains between the eas ded; consequently, it is about three times as tern part of our region and Pite Lappmark should large as the island of Oeland, considerably lar properly have been explored: however, circum ger than the island of Gotland or the province stances beyond my control prevented my remo of Blekinge, and somewhat smaller than the state ving this white patch from the botanical map of Delaware. It is practically uninhabited and of Lapland. roadless (cf. Chapter Ill). The region is situated between those inves tigated by TENGWALL (1914, 1920, 1924, 1925), Districi I BJORKMAN (1937, 1939), ARWIDSSON (1926 a, 1943), VESTERLUND (1924), and WISTRAND (1934, For the sake of convenience the region was 1946). To the north, it does not quite reach the divided into eight districts (cf. fig. 4). District I area from which reports have been published in comprises the woodland between Lake Kara,tsj BJORKMAN 1939, the northern boundary being and l..Jake Peuraure on one side and Lake Tja a line from Lake Slappehaure due west to the motisjaure and Lake Saggat on the other; the river Sepirjokk and along this river to Lake Kut western boundary coincides with the tourist path jaure; the mountains immediately north of that of >>Kungsleden>>. line were explored by BJORKMAN in the years The eastern half of the district, which lies in after 1939, though his results are not yet pub the region of primary rock, is most insufficiently lished. investigated. However, considering its extremely From Lake Kutjaure to Lake Sjabbtjakk the poor and monotonous flora, this is hardly a se western boundary of the region coincides with rious shortcoming. In the neighbourhood of Lill that of the Sarek National Park, except in two selct, for instance, there are at places no plants places: Mt Sakok is included, the reports of the to be seen for n1Ues on end but pine, Betula co numerous botanists who visited this mountain riacea, B. verrucosa, Vaccinium Myrtillus, V. in the 19th century not being given in TENG Vitis-idaea, Empetrum, Deschampsia flexuosa and, WALL 1924; and the table-land of Parek is ex locally, Calluna, Lycopodium complanatum and cluded, since it was so thoroughly investigated single specimens of Chamaenerium; during a stay by TENGWALL (op. c.) that I did not go there. of four days in this area of barren boulder-land, Survey of the Region Investigated 19
Fig. 2. Situation of the region investigated. meagre pine heaths, and oligotrophic fens, only about 100 native vascular plants were observed, all told. The crests of Mt Ailates, Mt Fatforita, &c. which reach, alone in this district, above the timber line were not visited. More closely inves tigated were, besides the area just mentioned, the surprisingly rich and interesting south-bluff of Mt Farforita (fig. 5), the vast, though trivial Fig. 3. The investigated region (black) is situated be fen-land of Luovvajegge, and the north-bluff of tween those investigated by TENGWALL (T), BJORKMAN Mt Ailates. Both the shape and the flora of the (B), and ARWIDSSON (A). south-bluff of Mt Farforita indicate the occur rence of rocks that are less hard and acidic than chia thyrsiflora, and that the Spargania are re those of the surrounding, barren area. gularly found in flower is obviously due to the That the lakes, Karatsj and Peuraure, al fact that the feeder streams come mainly from though situated about 150 rr1 higher than Lake the woodland and, consequently, the water is Saggat, harbour a few plants not found else warmer than that of the other big lakes with where, e.g. , Spa·rganium minimum and Lysima- their inflow of glacier-water. On the other hand 20 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Fig. 4. Boundaries of the districts. the absence of all Potamogetones is caused by the per alpine range and would all seem to be shaped low nutrient content of the water. in the way described by HOLMQUIST (1900 p. 85) The eastern part of the central lowland be regarding Mt Tarfek. They rise slowly in the tween Mt Ailates and 1ft Ramek is also oligo general direction of the folding, from the north trophic in character, the drift hills and sandy west to the south-east, and end abruptly in a plains being occupied chiefly by pine heaths with row of steep bluffs, the >>glint>> (PI. 3). Rocks an undergrowth of, i.a., Ledum palustre. Here belonging to the eastern facies of the cambro one finds the only big Sphagnum bog of the re silurian formation, generally clay -schists out of gion, Stuorjegge. Farther to the west the swamps the >>hyolithus series>>, are overlapped by a crust become much nwre varied and interesting. In of harder seve schists; on the westernmost part the area of mires and small lakes around Lake of Mt Lastak hyolithus schists also overlie the Tjuskaure, rnostmire types are to be found, from seve schists (HOLMQUIST Le.). lush carrs to >>flark>> mires, which here reach their The presence of silurian roeks is as usually ap western limit. Unfortunately, these fen lands parent from the increased richness of the vege could not be investigated so carefully as they tation. Especially the cliffs, where the clay merit owing to the hellish mosquito plague, that schists crop out, display an interesting flora rich is scarcely surpassed anywhere in Lapland and both in alpine and southern species (PI. 3). Al renders a prolonged stay in these parts quite im so the lower slopes of the mountains and the possible. neighbouring lowland bear witness to the ferti The mountains farther west, i.e. Mt Sappek lity of the soil by eutrophic fens, luxuriant spruce vare, Mt Tarfek, Mt Ramek and Mt Lastak, forests rich in herbs, &c. The overlapping seve constitute the easternmost border of the pro- schists appear, on the contrary, to be rather Survey of the Region Investigated 21
Fig. 5. The southern steep of Mt Fa.rforita. Below the precipice a. fringe of vegetation, where the >>south-bluff>> speeies occur. Further down a slope of naked boulders. A lynx wa.s met with just below the white patch on the precipice. Photo Sten Selander 22.7. 194 7. barren, at least as far as they are investi Kamajokk and Njatsosjokk. It is likewise very gated. unevenly explored. The surroundings of K vikk These parts of the district lie entirely below jokk, including Mt Snjerak, were visited by all the conifer limit with the exception of the crest botanists ever since the days of LrNNlEUS or of Mt Lastak; on Mt Ramek spruce bushes grow MoNTIN, whereas the northern and north-eastern right up to the summit. parts of the Kables massif, which fills all the Thanks to the excursions of LJESTADrus, AN centre of the district, are completely unexplored. DERSON, ALM and the author the flora of Mt The south-facing bluffs of the >>glint>> near Lake Lastak and the adjoining mountains is known Saggat, the gigantic eastern precipice of Mt Kas rather exhaustively; however, the valley of Par savare, and the delta of .Anok have been fairly kajokk and the delta of the river Tselekjokk thoroughly investigated by the present author were just touched upon by DAHLBECK. jointly with DAHLBECK and RUNE. Mt Njanja with its enormous south-facing bluff, which rea ches high up into the alpine belt, is not visited District 11 by anybody but ALM, and on Mt Kiuri no bo tanist has set foot after W AHLENBERG more than District II extends north of Lake Tjamotis 140 years ago. DAHLBECK and the present author jaure and Lake Saggat and east of the rivers just wandered through the valley of Blackalven 22 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark and the western part of the Rittak valley; only getation of the valley has a mesotrophic char Mt Tjalta and the environs of Tjamotis were acter without any distinctive marks. somewhat closely investigated. In the valley of Kamajokk, one's interest is Geologically the undulating table-land of Kab captured above all by ..Anok, a delta several les seems rather like the mountains south of Sag miles wide extending south of the confluence of gat. The cambro-silurian rocks, mostly sand the rivers Kamajokk and Njatsosjokk, about 10 stones underlying clay-schists (cf. SVENONIUS km to the north of Kvikkjokk. It is rather like 1900 a p. 293), are overlapped by a harder crust, the well-known Kvikkjokk delta, though larger according to HoLMQUIST (op. c. p. 166) consis and far less influenced by Man. The river Kama ting of >>green -gray, dense seve schists>>, though jokk meanders through ..Anok in countless, partly according to LUNDQVIST (1944 p. 42) including almost circular turns� its perpendicular banks, also syenites &c. From a botanical point of view about one m high, are overgrown by dense wil this difference is, however, rather unimportant, low thickets, where Salix glandulifera often pre since the vegetation of the table-land appears dominates. Inside the banks a great number of just as poor and trivial everywhere but in parts exceedingly shallow, oligotrophic gyttja lakes of Mt Snjerak. The interesting plants are almost extend with thin stands of Equisetum limosum exclusively concentrated in the precipices, where and Carex aquatilis and, especially in Lake the cambro-silurian rocks crop out. The char ..Anujaure, elodeid communities comparatively acteristic >>south-bluff flora>> is as well developed rich in species; the lakes are usually b9rdered on Mt Tavvevare, Mt Snalkok and Mt Kassavare by isoetid communities (>>iivjebroddsformationer>> ). as on the mountains south of Lake Saggat. In These lakes communicate with the river through the east precipice of Mt Kassavare quite a num narrow, meandering channels, almost stagnant ber of basophilous alpine plants have an isolated and one or two m deep, the bottoms of which outpost where they descend far int.o the birch are covered patchwise by.Hippuris vulgaris, ste belt or even the conifer belt. Thus Woodsia gla rile Sparganium spc. and sterile Ranunculus pel bella has there its most abundant stand in our tatus. The humid parts of the interjacent flat, region and perhaps in all Sweden; and such queer alluvial terraces are occupied by impenetrable meetings as that between Salix herbacea and willow thickets, here and there interspersed with Oxalis Acetosella bear witness to the peculiar con birches; on the drier plots birch woods (of Be ditions. The mires in the forest east of Mt Kassa tula coriacea) grow, either park-like with a her vare are pronouncedly eutrophic. baceous undergrowth, which is luxuriant and The conifer forests on the south slopes of the highly coloured though rather poor in species, Kables massif consist chiefly of spruce. But in or choked-up by dense shrubberies of willows the region of primary rock, i.e. in the valley (mostly Salix borealis), Ribes spicatum ssp. lap of Blackalven, around Tjamotis, and on the lobe ponicum, and Prunus Padus ssp. borcalis. · The of gneiss-granite intruding along Lake Saggat undergrowth is like t�at of the subalpine birch past Mt A.rrevare, pine forests of a trivial heath woods except that it does not comprise such type predominate . .Around Savvonselet in the plants, e.g., Athyrium Filix-femina, Dryopteris north-eastern corner of our region, there is a del austriaca, Actaea erythrocarpa, Rubus idaeus, ta, in the main of the same type as that of ..Anok, Daphne Mezereum, Angelica Archangelica ssp. dealt with below. norvegica, and Lactuca alpina, as in these parts The Rittak valley between the massif of Kab demand mobile subsoil water. les and that of Parte lies also in the conifer belt. Vegetation of the same general character re North of it there is a well-developed birch belt; curs at Selet and round the mouth of the river south of it, on the barren north slopes of Kables, Njakajokk; for the rest the valley of Kamajokk the pine ascends up to the timber line. The ve- is occupied by rather trivial conifer forests. Survey of the Region Investigated 23
District Ill
District Ill embraces chiefly the lower Tarra valley and the adjacent mountains and valleys, i.e. the valley of Tjuoltavuopme, the Tarrekaise massif and its offshoots, the undulating table land south of the river Tarreatno with the high massifs of Mt Staika, �It Vuoka, and Mt Sjella puolta rising above it, and the Tselekjokk valley. On the north and the west the district is boun ded by the valley of Vaimokvagge, the river Kurajokk, the ravine of Paturkarso, a line west of Mt Tarrekaise and J\it Ruonas to the junc tion of the rivers Kamajokk and Tjuoltajokk, and the river Tjuoltajokk. Mt Tarrekaise and Mt Staika reach nearly 1800 m s. m.; the alti tude of the forementioned table-land is about 800 m s. m. Between the summits of Mt Staika there is a glacier facing east, which is not on the ordnance map; on the other hand, the glacier on Mt Tarrekaise does not exist except on the map. The far largest part of the district lies above the wood limit; only the lowest parts of the three forementioned big valleys and all the valley between Mt Nammatj and Mt Vuoka be long to the conifer belt, and farther up in the Tarra valley and the valley of Tjuoltavuopme there are the largest subalpine birch forests · of Fig. 6. Vegetation of high-grown herbs below the sou the region. thern steep of Mt Nammatj . Photo Gunnar Beskow August 1946. Mt N ammatj and the easternmost mountain north of the river Tarreatno, Mt Vallivare, would seem in the main to be built up in the same way south side and, possibly, in some of the preci as Mt Lastak and the Kables massif. Mt Nam pices facing north, these parts possessing a much matj, which is entirely situated in the conifer richer vegetation than the proper tableland. Bo belt, is composed of the same seve schists as Mt tanically Mt Nammatj and Mt Vallivare have Lastak; cambro-silurian rocks are not established, been exhaustively explored by most botanists but doubtless underlie the seve schists, since who visited our region. such rocks are found on the island of Storhol During the last century the parts of the lower men a few km to the east (HOLMQUIST op.c. p. Tarra valley which lie north of the river have 82). The rich flora of the mountain is concentra also been scoured by many botanists. The delta ted in the south bluff. In Mt Valliva.re the seve of the river Tarreatno and the area of small pools schists are mostly granulitic quartzites (ibid. p. around Lake Mertikjaure resemble the delta of 248); the mountain forms a level, dry table .Anok in all essentials. They differ only in the land, some 10· km long and about 900 m high, larger number of dead arms of the river, which with steep sides and a rather poor flora. How have often changed into willow carrs, and the ever, soft hyolithus schists crop out on the eas much stronger influence of cultivation; for in tern slope (ibid. p. 83) and probably also on the stance, the haymaking preserves the wet mea- 24 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
a narrow strip from Mt Pavevare to about 10 km eastwards. The area south of the river Tarreatno has not been visited by me. The low mountains south of Lake Tarraure have been investigated espe cially by MoLLER, and Mt Vuoka by, among others, WAHL ENBERG; DAHLBECK has paid a short visit to Mt Valle and Mt Unna Valle south of the river Tselekjokk; and RuNE has explored the flora of Mt Ruopsok. Since all reports seemed to confirm that the flora of, at any rate, the al pine belt is rather uninteresting, I thought it well-advised to centre my attention on other parts of the region. It must be admitted, how ever, that it is a deficiency that, therefore, my investigation does not cover the virgin conifer forests in and round the valley of Tselekjokk . .
Fig. 7. Luxuriant birch wood in the Tarra valley below The Tarrekaise massif is the southernmost off Mt Njunjesvare wit.h, i.a., Aconitu.m septent1·ionale. shoot of a huge amphibolite field, which extends Photo Nils Dahlbeck August 1939. to the north in the direction of the interior of Sarek; eastwards it reaches Mt Kaskaivo and Mt dows round the lakes, which would change into Vallispiken. It is partly characterized by the oc willow thickets if it was discontinued. The rich currence of extensive boulder-lands, which are est limnic and telmatic flora of the region occurs at places, especially in the interior of the Tarre in Lake Mertikjaure. kaise massif, almost destitute of vegetation and The conifer region of the Tarra valley, which which I have hardly investigated at all. How westwards reaches Njunjes and the river Karso- ever, the flora along the standard route of the . jokk, has the same trivial character as that of earlier botanists through the valley of Ruonas the Kamajokk valley. The subalpine birch fo vagge and farther to the north-west is fairly rest, following the northern bank of the river well known, at least partly. On the south-wes in a narrow strip, is mostly of an extremely luxu tern slope of the Tarrekaise massif lenses of mag riant meadow type; south of the river the thin, nesite and dolomite crop out (cf. also A. HoGBOM somewhat sporadic birch woods belong to meagre 1930 p. 419) which are noticeable through their heath types except in the delta at the western richer flora. end of Lake Tarraure, which is a poorer equiva The flora of Mt Njunjesvare, situated between lent to the deltas described above. Mt Karrekaise and Mt Kaskaivo, is much richer The bed-rock in the mountains south of the than that of the surrounding mountains. Al river Tarreatno consists of bard, largely quartz though I have not found any statement to this itic, seve schists, which are capped by amphi effect in the topical literature, it seems as though bolites in Mt Staika and in a belt from there the koli schists, that underlie the amphibolite eastwards to the upper parts of Mt Vuoka. Mt of Mt Tarrekaise, are exposed on Mt Njunjesvare
Ruopsok west of Mt Vuoka is made up of ser and perhaps on the adjacent Mt · Hilto. This is pentine (SVENONIUS 1884 p. 202 ff., T. Du indicated also by the fact that the birch woods, RIETZ 1935 p. 142 ff.). According to HOLM which are mostly poor and heathy below Mt QUIST (op.c. p. 81) softer koli schists extend in Tarrekaise, display the same splendid luxuriance Survey of the Region Investigated 25 below Mt Njunjesvare, &c, as farther up in the valley, on Mt Sevvatjakko and Mt Slitavare, which consist of calcareous koli schists. The mountain-sides to the north of the valley from Mt Njunjesvare to Mt Sammartjakko have the shape of steeps facing south or south-west, with the same >>south-bluff flora>> as the cliffs north of Lake Saggat; on Mt Njunjesvare the compo sition of this flora is influenced by the a bun dant moisture trickling from the rock-faces. Further west the >>South-bluff species>> gradually diminish in number and dissappear altogether west of Mt Sammartjakko . .According to SVENONIUS (1900 a, map p. 322) the upper half of Tjuoltavuopme, i.e. the valley further north running parallel with the Tarra valley, lies entirely within the forementioned field of amphibolite. Its flora is comparatively poor; softer and more nutritious rocks seem to occur only in Vuoksapakte, a south-facing pre cipice near the eastern end of Mt Tjuolta, and probably elsewhere in separate places (the south western offset of Mt Tjuolta, the north-preci pice of Mt Habres, &c.). North of the river there is a subalpine birch forest, mostly of a luxuri ant meadow type and completely untouched by Man for all its expanse from Mt Tjarrok to the river Tjuoltajokk, which is certainly the largest and finest primeval birch wood in all Fenno Fig. 8. The biggest pine of the region near Lake Vallatsj scandia. No botanist had explored this valley jauratsj . Photo Nils Dahlbeck August 1943. before the visit of DAHLBECK and the author; the crest of Mt Tjuolta and the valley north of it are as yet uninvestigated. District IV In the north-eastern corner of the district, i.e. in the lower part of Tjuoltavuopme and in the District IV embraces the mountains on both low mountains of Tjarrok, Ruotivare, and Val sides of the upper Tarra valley. It is bounded latsj, there is an area of primary rock, anortite by the Sarek National Park, District III, the &c., the geology of which is most complicated river Hadditjokk, Lake Ravejaure, and the river (cf. GAVELIN 1915). However, there is no occa Kartejokk down to Lake Nedre Skalo; from there sion for entering upon this subject here, since the boundary line runs north of Mt Kartevare the different rocks influence the vegetation in tjakko to the river Maddakajokk and then along the same unfavourable way. Especially in the the river Tarrejokk to the boundary of the Na barren boulder-lands, thinly covered by prime tional Park at Lake Tarreluoppal. .Apart from val woods of short, twisted, but often very stout the Tarra valley, the narrow groove of which pines (fig. 8), the flora is about as meagre as divides the district into two halves, this district in the oligotrophic area near Lake Karatsj (cf. is situated entirely above the tree limit. _ above p. 18). "\Ve have now entered upon >>the rich area>>, 26 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
above it the >>south-bluff flora>> has its last ou.t post in that valley. The hard seve schists extend from District Ill to Mt Skaitatstjakko, Mt Tjaurak and Mt Unna Tjaurak, in the south-western corner of District IV. .A sweep of such schists is also met with on the other side of the valley of Katnjunjes jokk; it runs from Mt Tsakkok eastwards to Mt Perro and Mt Kartevaretjakko. Prom there it turns off into District V, describing a curve to tlw north and north-west over Mt Silpatjakko, Mt Rigaive, and the adjacent mountains, to Mt Junkartjakko and the eastern crests of Mt Kerke vare (cf. HOLMQUIST, op.c., map p. 104, and. SVENONIUS, op. c., map p. 322). This sweep with its trivial flora has been less closely investigated by me than the surrounding Fig. 9. The boulder-land on Mt Sammartjakko. Mark the tracts, and the mountains between Lake Vai man in the middle of the picture! mok and Lake Tjaurakjauratsj are completely 1943. Photo Nils Dahlbeck July unexplore(l. Por the rest this district, except the northern and eastern slopes of Mt Velkis i.e. those tracts which get their geological char puolta, has been rather well investigated, chiefly acter from the western facies of the cambro by the present author with the assistance of silurian rocks, the >>koli>> schists; these predo SKOTTSBERG, DAHLBECK, and RUNE. minate in all the rest of our region. However, .A sweep of soft, highly calcareous koli schists in the eastern parts of District IV there are still extends southwards on both sides of the Tarra some fair-sized amphibolite massifs, viz., Mt valley, east of the river to Mt Sevvatjakko and Kuratjakko, Mt Sammartjakko, and Mt Velkis the north-western part of Mt Sammartjakko, and puolt.a. These are not as botanically poor as, west of it to Mt Rivkonjunjes and the north for instance, Mt Kaskaivo. In Mt Kuratjakko eastern part of 1\It Kuratjakko. This fact gives the amphibolite encloses large quantities of lime their eharacter to the flora and vegetation of stone and dolomite (HoLMQUiST op.c. p. 235); these tracts. The birch forests of the Tarra val and koli schists seem to occur in the western parts ley consist mainly of meadow woods rich in of Mt Sammartjakko as well as under the amphi herbs. True, on the valley bottom woods of that bolite further east in the same massif, where the type stop abruptly at the river Slitajokk east rock is very soft and easily weathered. In the of the river Tarrejokk, and occur only below Mt Velkispuolta massif there are considerable lenses Rivkonjunjes west of it. The occasion for this, of dolomite and magnesite (cf. HoGBOM op.c. however, is that the valley bottom north of Sli map p. 418, which, however, omits the largest tajokk and west of Tarrejokk is occupied largely deposit of this kind, that on the south-slope of by mires; lumbering of old date may perhaps Mt Velkispuolta); further, at least the western also have had its effect (cf. below p. 39). On the slopes of this mountain consist of koli schists. mountain-sides the forests are everywhere chiefly On the south-western slope of Mt Sammartjakko of a rather luxuriant meadow type, and heathy above the Tarra valley there is a huge boulder woods occur in the main only on the gravel ter land, several square km in size and practically races and other glacifluvial deposits extending devoid of vegetation, where many boulders are west of the rive1 below the nwuths of the collate as large as good-sized houses (fig. 9); in the steep ral valleys. The mires occupying good-s ized Survey of the Region Investigated 27 areas also in the meadow woods are largely slo the districts have been explored by the author, ping hill-mires (>>backmyrar>>) of a rich-fen type, who has alone visited Mt Staddatjakko and else which for instance on Mt Rivkonjunjes and Mt has been accompanied, on different occasions, Katnjunjes extend right up to the tree limit (Pl. by SKOTTSBERG, DAHLBECK, and RUNE. One or 7). Especially on the mountain-sides around the other of us has climbed all the nwuntains, Lake Tarreluoppal there is a great number of except Mt Silpatjakko, Mt Kaiseketstjakko, and >>blekekarn (highly calcareous fens), partly affec Mt Junkartjakko, either to the top or to the ted by solifluxion and possessing a scanty, though upper limit of the vegetation. Several attempts :rnost characteristic vegetation of Triglochin pa to ascend Mt Kaiseketstjakko and Mt J unkar lustre, Juncus triglumis, Koenigia islandica, Saxi tjakko failed owing to gales or snowfall; for the jraga aizoides and Epilobium dav·uricum. same reason the valley between Mt Sulitelma Koli schists occur also round Lake Ravejaure, and Mt Kaiseketstjakko is unexplored. Nor have Dn the other side of the forementioned sweep we visited Mt Hammaren west of Mt Stadda of seve schists (HOLMQUIST op.c. p. 158). The tjakko, because of the risk, unintentionally to flora, however, is less rich than might be expec pass the Norwegian border during the war. Only ted, partly because the large snow-beds do not part of the niveous and uninviting east-precipice become free from snow until very late in the of Mt Jeknaffo has been investigated. The large summer, partly because Kallovaratjeh, i.e. the mountain situated, according to the ordnance area south of Lake Ravejaure, is a dead-ice land map, between Mt Jeknaffo and the valley of scape with numberless small lakes and n1eres, Vejevagge does not exist in reality. A broad and with hills and drums mostly consisting of strip along the Norwegian border is, by the way, hard, infertile morainic material. The flora of the neither on the Swedish nor on the Norwegian two adjacent, small mountains of Jallotsj and map. It should perhaps also be mentioned that Tjahevare, where the ground has another char the glaeiers do not actually cover by far as large acter, is among the richest in all the region. areas as indicated by the ordnance map (cf. be low p. 122). This district is the most high -alpine within our District V region. Mt Sulitelma (Pl. 12) reaches 1914 m s. District V lies entirely within the large, wes m. and Mt Jeknaffo 1853 m s. m., and the alti tern field of koli schists. It comprises the three tude of the passes is 979 m s. m. in the valley huge massifs of Junkartjakko-Kerkevare, Jek of Staddavagge and about 1050 m s. m. in that naffo and Kaiseketstjakko-Staddatjakko to of Vejevagge. 'l'he whole district is situated gether with the surrounding, smaller mountains above the tree limit, except the northernmost eor and the two interjacent valleys, Vejevagge and ner, where there are some small clumps of bir Staddavagge (Kailavagge). On the south and ches round the mouth of the river Stalok. Nor west it borders upon District IV, Pite Lappmark are large willow-thickets very frequent. They and Norway; and on the north and north-east occur chiefly at the northern foot of Mt Kerke it is bounded by Lake Sarjasjaure, the rivers vare, in the Stalok valley up to the river Stadda Stalok, Katjaurejokk and Puolejokk, Lake Fes jokk, and in places on what is called here the tajaure (Katpurosjaure), and the river Tarre Sulitelma plain, i.e. the table-land round the river jokk. Hadditjokk between Mt Sulitelma and Mt Tsak The district has been rather exhaustively in kok, where the many brooks from the surroun vestigated. Mt Junkartjakko, Mt Kerkevare, Mt ding mountains form a veritable maze of mean J alle and the northernmost parts of Mt J eknaffo der bends and islanded arms. On the alluvia of have been visited by several earlier investigators, these brooks hybrids of Salix herbacea are :rnore viz., WAHL ENBERG, LJESTADIUS, ANDERSON, plentiful than i� any other place visited by me.
CLEVE, and TENGWALL. The remaining parts of Apart from the forementioned sweep of seve · 28 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
schists (cf. p. 26), the bed-rock consists almost the regio alpina sterilis, by plant communities exclusively of rocks belonging to the western fa belonging to the meadow series. cies of the cambro-silurian. Mt Karranis, which On Mt Staddatjakko one further observes the is situated east of the sweep of seve schists, is calcareous wind -heaths of the eastern offset; such also made up of can1bro-silurian rocks. These heaths are rare within the district and hardly consist in the first place of calcareous phyllites occur anywhere else but in -places oil the hills and limestones and only to a lesser extent of between Mt Kaiseketstjakko and Mt Staddatjak soft, sandstone-like quartzites or sandy mica ko and on those at the lower course of the river schists (HOLMQUIST op. c. p. 153 ff. ). The Stalok. Mt Jeknaffo, whose crest is more than olivine-gabbro of Mt Sulitelma and the moun 10 km long and throughout more than 1700 m tains farther to the north does hardly occur high and descends almost perpendicularly to the within the area investigated. Nor has the sweep east, is the mightiest massif falling entirely with of gabbro, amphibolites, &c., which according in our region. Its top ridge and east-precipice to VoGT (1927 pp. 80 ff.) extends north of consist of hard amphibolites (ace. verbal infor Mt Sulitelma, in the western parts of Mt Kai mation by Mr G. KAUTSKY not quartzites, as re seketstjakko northwards to the westernmost ported by HOLMQUIST, op.c. p. 154). This fact ex slope of Mt Staddatjakko, been investigated, plains both the abrupt change in the flora a little except for the last-mentioned, rather barren above the 1400 m level and the striking floristic area. poverty of the east side in comparison .to the The flora is relatively poor only on the sweep west side. In the valley between Mt Jeknaffo of seve schists (cf. above p. 26) and in the large and l'tft Jalle on Vejejokk, at an altitude of a depression north of Lake Ravejaure, where a little more than 900 m, there are large plateau hundred square km or more are occupied almost mires with numerous peat hummocks about one exclusively by perennial snowfields and mighty n1 high; here, and on the adjacent Lake Jalle snowbeds. But immediately after leaving these jaure, there are, i.a., the easternmost stations tracts, which permanently are more or less snowed within our region of the western Carex rufina. up, the character of the vegetation becomes The valley of Vejevagge with its relatively bar transformed. For instance, the slopes towards ren drift hills is of little botanical interest. 1\'I t the Sulitelma plain of the adjacent Mt Haddit Junkartjakko has been very unevenly investi and Kallovaratjeh are occupied by alpine ex gated; rather well explored are only the eastern treme-rich-fens with a mass vegetation of Pin offset (Passealke), the westernmost offset, a.nd guimtla alpina and Salix 1·eticulata. And anum the lowest parts of the north-slope, the latter ber of subalpine species have isolated outposts having been exhaustively investigated by ASTRm on the southern steeps of l'tit Kaiseketstjakko. OLEVE; the tremendous north-precipice, above Solifluxion is very intense in this district with the mighty accumulations of boulders in >>the its high precipitation and easily weathered rocks Valley of Death>>, is completely inaccessible. On (cf. below p. 155). This applies in the first place Lake Puolaure there is the most interesting alpine to Mt Jeknaffo, at the north-eastern foot of delta of our region, known among botanists as which there is a series of solifluxion terraces of the first Swedish station of Catex bicolor; in the mighty proportions, and Mt Staddatjakko, not vicinity is a mire with an isolated station of least its western parts, which until quite recently Eriophorum medium. The northern slopes of Mt were covered by a gigantic perennial snowfield Kerkevare are largely covered by wet solifluxion now melted away (cf. below pp. 122 f.). Solifluxion soil; the western parts of this mountain are bo certainly is the main reason why scrub heaths tanically among the richest areas of our region. are almost completely absent from large parts The flora of Mt Jalle on Vejejokk resembles that of the district. Instead the mo�ntain-sides are of Mt Kerkevare; at its western foot, in the Sta largely covered, right up to the boulder-lands of lok valley, there is a remarkable glacifluvial Survey of the Region Investigated 29
Fig. 10. Lapps' tents in the pasture land of Padjelanta. Photo Nils Dahlbeck July 1942. landscape with cones, terraces, funnels and rid Lake Kutjaure. Mt Arasvaretjakko south of ges consisting of oligotrophic gravel, the vege Lake Vastenjaure has been visited only by WAH tation of which is, consequently, rather trivial. LENBERG, and J\'It Lemakvare and Mt Allep Ala tjakko, only by the present author. Unexplored,
District VI or aln10st so, are Mt Mola, �I t Huornatj, Mt Ra pok, and Mt Spetjau. District VI comprises Padjelanta, the large To judge from the only geological map access pasture-land for reindeer between Sarek and ible to me, that by SVENONIUS (1. C.) which is, Lake Virihaure (fig. 10). On the south it bor however, most sketchy, the bedrock of Padji ders upon District V; on the east it reaches the lanta consists almost exclusively of koli schists. Sarek National Park and on the west, Lake Viri An offshoot from the amphibolite field of Sarek haure, the river Jaurekaska, Lake Vastenjaure, extends to the highest part of Mt Lulep Alatjakko the river Vuojatatno, and Lake Kutjaure. This and Mt Allep Alatjakko. The mountains north district is also rather well explored, largely thanks of the lower course of the river Mellatno, viz., to the investigations of TENGWALL who has, i.a., Mt Huornatj, Mt Mola, the eastern part of Mt alone visited �It Lulep Alatjakko, Mt Mattaive, Allak and Mt Arasvaretjakko consist likewise of and Mt Latatj. I have received a good many amphibolites (verbal information by Mr G. KA informations also from DAHLBECK who, for in UTSKY); there are, however, large intrusions of stance, is the only botanist to have investiga limestone in the western part of J\It Allak. Oli ted the northernmost mountain, Mt Njerek on gotrophic glacifluvial deposits occupy conside- 30 SELANDER: Floristic Phytogeography of South-Western Lule Lappma_rk
rable areas especially north of the river Puole of Lake Virihaure, are rather scarce and ,gener jokk and in the Mellatno valley; a boulder ridge ally occupy but small areas. between the river Mellatno and Mt Mattaive is very conspicuous. Padjelanta is a region of low mountains (see District VII Pl. 10). Only Mt Lulep Alatjakko reaches above 1500 m s. m., Mt Mattaive above 1400 m, District VII embraces the tracts immediately and Mt Allep Alatjakko and Mt Allak above south and west of Lake Virihaure, and Kollomo 1300 m; four more mountains pass an altitude loki, the western peninsula between Lake Viri of 1200 m, and the remaining ones are still lower. haure and Lake Vastenjaure. On the south and Consequently, high-alpine phenomena such as east it borders upon District V; its northern boulder-lands, solifluxion, polygon ground, &c., boundary, in the direction of district VIII, are comparatively rare. Wind-erosion, however, coincides with the river Kuotelisjokk and Lake is rather active; for instance, the forementioned Vastenjaure. glacifluvial deposits are partly intensely wind Ever since the time of WAII LENBERG some of eroded, and wind-heaths occur in many places. the mountains within this district, especially Mt Birch groves are found on the southern and Tuke (>>Lill-Thokim>), have attracted the atten western sides of Mt Unna l'riter, on the peninsula tion of botanists. Wholly unknown is only the of Titernjarka, on the west-slope of Mt Arasvare, flora of the nameless rnountain north-west of and round the mouth of the river Laddejokk. Lake Kasakj aure and that of the area south of On Mt Unna Titer and Mt Arasvare the wood Lake Raddejaure; the low mountains north of is of a meadow type and partly very rich in Alasluokta are also insufficient1y investigated. herbs. Birches, single or in small clumps, grow W AHLENBERG alone has visited the mountains a1so, e.g., on the south-steep of Mt Stuor Titer round the boundary-cairn (>>r,iksroset>>) No. 241 and on the steeps on the northern shore of Lake (>>Vajryggsroset>>) and those on the northern shore Virihaure between Arasluokta and the river Jau of Lake Sarjasjaure below the glacier of Almallo rekaska. Large willow-thickets occur above all jekna; TENGWALL was the only botanist to ex in the Mellatno valley. plore Mt Rakoktjakko and the hills around Ras On Mt Stuor Titer and below Mt Allak there tanjarka; lVIt Kasakpuolta and the hills on the are south-steeps with some characteristic plants, peninsula of Njerek have been investigated by e.g., Erysimum hieraciifolittm and Hackelia de the present author alone; and KoJlomoloki, the flexa, of great interest as they are situated chiefly hills of Luokivaratjeh, Mt Kappa, and Mt Stalo in the alpine belt. tjakko, by DAHLBECK and the author. In the Heath communities are much more important papers of W AHLENBERG and LlESTADIUSthe name than in District V. Thus Dryas-heaths rich in of >>Kappa>> or >>Kappis>> does not signify the pre species are frequent, and even Rhododend-ron lap sent Mt Kappa, but Mt Stalotjakko (cf. W AHLEN ponicum-heaths, which are very scarce elsewhere RERG 1808, nmp Tab. 1). It is evident, how in our region, occur in places; round the upper ever, that these botanists never ascended Mt Sta course of the river Mellatno, large areas are occu lotjakko, but just to�ched upon its lowest slopes pied by a Carex Bigelowii-heath (TENGWALL 1920 to the south and west. p. 390), and a Kobresia myosuroides-heath covers My statements regarding the geology of this pa.rts of the boulder ridge at Mt Mattaive (ibid. district and district VIII are almost entirely p. 385). Loiseleur:ia procurnbens�Diapensia lap based upon verbal information by Mr G. KAUT ponica-heaths characterize the glacif1uvial depo SKY, the sketchy and rather inaccurate map in sits. On the other hand >>Snow-beds rich in herbs>> SVENONIUS 1900a being the only pub1ication and similar meadow communities, which are dealing with the geology of these parts. The bed characteristic of the solifluxion mountains south rock is chiefly calcareous koli schists, nearly Survey of the Region Investigated 31
everywhere with large deposits of pure limestone. tains within this district, especially Mt Tuki, Mt Mt Rakoktjakko, situated west of Lake Viri Kuobberi, · Mt Kappa, and Mt Stalotjakko, are haure on the Norwegian frontier, is composed among those riehest in rare arctic plants in ou.r of granite and, according to SVENONIUS, in its region and in all Sweden. This is rather pecu southern part of greenstones. A sweep of green liar, since none of these mountains reach any stones also extends from the western part of considerable heights. Only Mt Stalotjakko rea Mt Stalotjakko northwards to Mt Metjer. Fur ches above the 1300 m level; Mt Kappa is about thermore, rock's of a similar kind form the sum 1200 m high, l\H Kubberi 1110 m, and Mt Tuki n1it of Mt Tuki, i.e. the high ridge west of the only 890 m. Mt Stalotjakko and Mt J eknaffo little lake, and the western part of the small · are the only stations of Potentilla hyparctica on peninsula of Pellonjarka. Also the greenstones the European mainland, except Kanin N oss in are more or less calcareous. Schists and phyllites Arctic Russia. Mt Tuki is the only Swedish sta of a hard, sandy kind extend round Alasluokta, tion of .Arenaria humifusa. On Mt Kubberi and the westernmost cove of Lake Virihaure. Ser Mt Tuki there are, i.a., steep screes of dry, highly pentines occur in severUJl places, particularly in calcareous soil where, e.g., Arenaria norvegica, Luokivaratjeh, i.e. the hills south of the cove Minuartia rubella, and Braya linearis are to be of Tukiluokta. Glacifluvial formations are found found. on Stalonjarka, the point west of the mouth of The westernmost outposts of this rich flora the river Stalok; further west no such deposits are met with on Mt Kaitsanjunje. The low, worth mention were observed by me. snowy mountains further west, round the cove No real birch woods grow within the district. of Alasluokta, are characterized, as far as my Only small clumps or single trees are to be found experience goe.s, in the first place by wide Cas in scattered places, e.g., the northern slope of Mt siope hypnoides-heaths and swampy snow-beds Piete; Jalkok, i.e. the table-land north of Mt Sta with Carex rufina &c.; they would seeni to har lotjakko; the north-side of Mt Metjer; the island bour no arctic or arctic-montane plants except of Stuorsuolo; Mt Njerek near the brook from the more or less ubiquitous ones. lt goes with Mt Kuobberi; the northern slope of Mt Sirka out saying that this applies also to the granite vare; and round the small lake on Kollomoloki. mountain of Rakoktjakko. However, some cen The two last-mentioned localities stand for the tric basophytes recur on Kollomoloki, where the western limit of the birch within our region. 'l'he northernmost outposts of the characteristic >>Viri largest willow-thickets of the region occur round haure flora>> occur on steeps facing north; in the brook west of Sirkaluokta, where many addition a few thermophilous species, e.g., Sedum square miles are covered with an impenetrable acre, are met with on the southern slopes. willow-jungle.
Wind-heaths are quite frequent. The largest District VIII such heaths within our region extend on the dry, undulating table-land of l\it Piete where, i.a., Finally, district VIII comprises the area north Carex na,rdina and C. glacialis occur in masses. of La�e Vastenjaure and west of the river Vuojat Other instances are the plateaus on the tops of atno. The mountains between Lake Vasten Mt Kappa and Mt Jalle on Sarjasjaure. The lat jaure and Lake Sallohaure have been thoroughly ter mountain is remarkable also because of its investigated by TENGWAI"L; he alone has visited Routh-steep, observed already by WAHLENBERG, Mt Kuotelis, Mt Hurrevare, l\H Rakamvare and where a nun1ber of subalpine and low-alpine parts of Mt Svartivare, and later investigations plants have their last outposts in the direction have only slightly increased the number of spe of the high mountains. The only large Cassiope cies known in these parts. Among the n1oun tetrago na-heaths of our region extend on the nor taius north of Lake Sallobaure, Mt N oijevare and thern side of l\1tKasakpuolta. Some of the moun- Mt Apmelvaratj have been visited only by BJORK- 32 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Fig. 11. The southern steep of Mt Tjargesvare. Iri the background Lake Sallohaure and the westernrnost mountains of Sarek. On this steep, Gentianella aurea is found in its only Swedish station; further remarkable plants occurring there are, e.g., Antenna1·ia lapponica, Braya linearis, Primula scandina-cica, Ajuga pyramidalis, Epipactis atrorubens and Viola 1'upestris. Photo Sten Selander 12.7. 1946.
MAN, and l\It Unna Tjargesvare, l\ti t Stupirvare, tenjaure basin. To a non-expert, the limestones and the nameless mountain north of Mt Tjar appear harder and less _easily weathered than gesvare, only by the present author. Mt Paikas in the Virihaure region, mostly possessing a de and Mt Katoktjakko are uninvestigated. finitely crystalline structure. The bedrock consists almost solely of cambro Especially characteristic of this region are the silurian schists with mighty layers of limestone. abruptly descending, south-facing steeps, most Only Mt Kuotelis is composed of granite. Am of them with big talus slopes below the preci phibolites hardly occur anywhere but in Mt Ra pices, which are found on practically the whole kamvare, where there are also big deposits of ser northern shore of Lake Vastenjaure, and on �n pentine. Mr G. KAUTSKY has told me that no Tjargesvare and the mountains west of it. Above layers of quartzites worth mention occur with the steeps wide areas are occupied by undulating in this region. However, I got the impression table-lands, which are characterized, between that the schists are somewhat quartzitic at places Lake Vastenjaure and Lake Sallohaure, by ex on the northern shore of Lake Vastenjaure as tensive Gassiope hypnoides-heaths (TENGWALL well as near the river �1alkomjokk, which would 1920 p. 379). These tracts are still more low explain the occurrence of the only real My rtil alpine than Padjelanta; none of the investigated lus-heaths observed by me in the Virihaure-Vas- mountains reach above a level of 1000 m. Survey of the Region Investigated 33
The flora of the south-cliffs includes an appre bluff of Mt Tjargesvare; elsewhere there are ciable number of thermophilous lowland species, · merely small, scattered clumps, e.g., north of several of which have their only stations with Lake Sallohaure and the river Malkomjokk and in our region on these mountains. This charac on Lake Kutjaure near the mouth of the river ter of the flora is all the more striking because Sepirjokk. On the whole, the flora of these steeps the steeps · are situated chiefly in the alpine belt. is very characteristic and differs definitely, both The only birch wood within the district is the positively and negatively, from that of the rest one occurring below the eastern part of the south- of our region (cf. below pp. 140 ff.).
4-496149 Sten Selander I Ill. HUMAN INF L UENC E
Settlers and settlements the cultured ground is made up of grassy fields, where hay from >>natural nleadows>> is On the whole, the region discussed here harvested. belongs to the parts of Scandinavia least in At or around the farmsteads, a number of fluenced by civilization. It is situated entirely weeds have been introduced, among which above the Cultivation Lin1it (>>odlingsgransen>> ), about 30 would seem to be more or less constant: and stationary settlements are met with only only about ten, viz., Poa annua, Jun(;USbufonius, in the conifer belt. The first colonizers arrived Luzula multiflora ssp. occidentalis, Cm·astium in Kvikkjokk in 1661, when mining was started holosteoides, Sagina procumbens, Trifolium repens, at Mt Silpatjakko. However, the mine was Carum Carvi, Euphrasia brevipila, Achillea Mil abandoned already about 1700, and in later lefolium, Chrysanthemum Leucanthemum and, in times, Kvikkjokk was nothing but an insignif one place, possibly Elytrigia repens, are natu icant church village, inhabited by a few families. ralized, showing a more or less pronounced In 1732, when LINNAEUS visited these parts, tendency to spread as neophytes into the na there were no settlements but Tjamotis and tive vegetation. Possibly Agrostis canina, Fes Kvikkjokk, and in 1755, only >>a few>> colonizers tuca rubra s. str., and Montia rivularis also faU lived in all Lule Lappmark (CAMPBELL 1948 under this category. Agrostis tenuis, Luzula p. 66). Njunjes dates back to the years around pallescens, Rumex tenuifolius, Stellaria graminea, 1820, and the remaining settlements are still Ranunculus acris, R. rep ens, Veronica serpylli more recent. :For the time being, there are folia, Campanula rotundifolia, and Erigeron acre three hamlets within the region, viz., Kvikk would seem to be represented by indigenous jokk, Tjamotis and Njavve, and the settler's populations as well as by introduced ones. farmsteads number ten, viz., Luovvaluokta Not a few alpine plants, otherwise rare or and I.Jillselet on Lake Karatsj, Levek, Peuraure lacking in the conifer belt, occur as apophytes gard, and Skaite on or near Lake Peuraure, on the small fields round the farmsteads, for Sagudden on Lake Tjamotisjaure, Arrenjarka instance Anthoxanthum alpinum, Phleum commu and Malenjarka on Lake Saggat, and Backen tatum, Alopecurus pratensis ssp. alpestris, Agros and Njunjes in the lower Tarra valley. The tis borealis, Poa alpina, Carex macloviana, C. colonization thus is restricted to the lake district brunnesaens, Alchemilla M urbeckiana, A. Wichu of the south-east. The stationary population rae, A. glomerulans, Astragalus alpinus, Rhi amounts to about 240 persons, i.e. about one nanthus minor ssp. groenlandicus, and Euphrasia inhabitant to each 17 square kilometres. frigida, and more rarely Minuartia biflora, Draba The inconsiderable plots of arable land, norvegica, and Gentiana nivalis (cf. TENGWALL always situated close to the houses, consist only 1925 p. 746). of small potatoe-patches, and now and then The vegetation of the conifer belt is influenced of tilted pieces of ground, where barley and also by the haymaking in the deltas of K vikk oats are sown for green forage. For the rest, jokk and Anok and, nowadays only excep- Human Influence 35
Fig. 12. The delta of the river Tarrea.tno. The meadows, which give a park-like aspect to parts of the deltas, owe their existence to the hay-making. Photo Nils Dahlbeck August 1939. tionally, on certain grassy mires. The meadows, Lapps and reindeer which give a park -like aspect to parts of the deltas, owe their existence to the haymaking; Broadly speaking, only La.ppish civilisation if this ceased, they would grow together into has left its marks above the conifer line. Excep willow-thickets (fig. 12). tions are only the establishments of Svenska .At least as significant is the timber-cutting. Turistforeningen (The Swedish Tourists .Associa Up till 194 7, woods had been cut chiefly on tion), viz., the cabins on Lake Sjabbtjakk the lands in private possession, e.g., on the wood (Partestugan), Lake Tarraure (Tarrekaisestu allotments of the Kvikkjokkers in the lower gan), the river Tarrejokk (Tarraalvshyddan), Kamajokk valley and south of Lake Saggat; and Lake Sarjasjaure, and the huts (kiltorna) the Crown Forests west of the river .Arrejokk at Sammarlapa, Tarreluoppal, Staloluokta, .Alas and a line between .Anamusluokta and .Anamus luokta, and Vastenjaureluspen. luokta were mostly untouched. Therefore, wide The Lapps, nomadizing within the region, tracts are still covered by primeval forests, belong to the three >>Lapp villages>> (lappbyarna) particularly on Mt Ramek and Mt Lastak, of Tuorpon, J akkakaska, and Sirkas. .Among where the Commissioners of Land Revenues the Tuorpon Lapps, whose caravan track have set apart a forest reservation, in the (flyttningsviig) lies north of Lake Karatsj and Tselekjokk valley, in the low-laying area be Lake Peuraure and south of the river Tarreatno, tween Mt Nammatj and Mt Vuoka, and in the one >>sita>> (tribe) has its summer camp (sommar valley of Njatsosvagge above Mt Ruotivare. viste) at Staloluokta on Lake Virihaure and its The Lapps, passing here only in spring and camp for spring and autumn at Mt Lastak or, autumn, have but occasionally cut down lichen in the case of two families, at Lillsclet; another grown trees for food for their reindeer, a thing sita has a camp for the herdsmen at Staloluokta, often met with in the forests of the lowland where the young men stay from Midsummer further east. till the beginning of .August; and a third one 36 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
has summer camps for the herdsmen at Mt the tourist path as far as to the vicinity of Pa Kartevare and Mt Katnjunjes, the families turkarso. The anthropochorous flora at Parte belonging to the two last-mentioned sitas stugan, e.g., the lowland race of Alopecurus mostly staying from spring till autumn in c�mps p'ratensis, and an introduced race of Festuoa outside the region. rubra noticed only at the .Alasluokta hut, have The Lapp village of Jakkakaska consists also followed on the heels of the tourist traffic only of two sitas, which both have their summer or the transports at the time of the building camps at the lower course of the river Mell of the huts. Usually, the weeds show no tendency atno, where they stay from the beginning of to become neophytes; thus, the anthropochorous July till the beginning of .August. The caravan race of Poa pratensis ssp. alpigena, though track of one sita lies mainly outside the region. often remaining at derelict Lapp camps, seldom The other sita, which travels by way of Ruonas or never appears outside the ruins of the huts. vagge, Tarreluoppal and Festajaure, has its .An exception is, perhaps, Poa irrigata; though camp for spring and autumn in the highland whether it be introduced or indigenous in its of Kables, some of the women and the old only station, the north slope of Mt Sirkavare, men staying during the summers in a stationary seems hardly possible to decide. camp at Njammilt in the Tarra valley. More importance certainly must be attached to the grazing of the reindeer. The exact num .Among the five sitas of the Lapp village of ber of reindeer ordinarily feeding within our Sirkas, only two sojourn for certain periods region at the height of summer cannot be within our region, one sita having their summer stated. However, in 1945, the latest year from camp at Tjakkotievva and the other theirs which figures are available, it exceeded 10000. on the river Laddejokk; in both places, the The reindeer of the Tuorpon Lapps then num Lapps stay from the beginning of July till bered 5275, those of the Jakkakaska Lapps the beginning of .August, and their caravan 1804, and those belonging to the two Sirkas tracks lie mainly outside the region. .Among sitas nomadizing here about 3000; further all these camps, only the one at Staloluokta more, 3000 to 4000 reindeer from Vaisaluokta is a >>big camp>> (storviste). Th� number of Lapps, are accustomed to stroll into the region (cf. soj ourning during the summer within the region, MA].'>;KER l. c.). amounts to about 215. For further details, However, the grazing of the reindeer probably cf. MANKER 194 7 pp. 459 ff. does not influence the vegetation to such an The weeds, introduced by the Lapps and now extent as these high figures inight suggest. occurring on Lapp yards (>>lappvallar>>) &c, The herds of reindeer are not as stationary are of little importance in the composition of by far as a herd of cattle; instead, they ramble the flora. They are but few in number: Poa far and wide, i.a., in order to escape mos annua, Stellaria media, Achillea Millefolium, quitos, gadflies, and other insects. Neither do introduced after 1940 to its only station of this the reindeer graze the ground as closely as kind (at Staloluokta), a probably non-indigenous cattle or sheep; for instance, when feeding on race of Poa pratensis ssp. alpigena, and possibly Angelica Archangelica, they eat only the leaves Poa irrigata. Some alpine plants, e.g. Deschamp and never bite off the stems. Their grazing sia atropurpurea, Poa arctica, and Sagina Lin therefore has not such a marked effect on a naei, occur on the Lapp yards as apophytes. given piece of ground as that of cattle (cf. The most common among the weeds, Poa Du RIETZ 1924 p. 95); its relative unimportance annua, spreads chiefly along the tourist paths, is evident, i.a., from the fact that there is no not along the Lapp tracks; in 1941, it was not discernible difference between the rather heavily noted farther up the Tarra valley than at Tarre grazed land east of Lake Kasakjaure and the kaisestugan, but in 1943, it had spread along river Tukijokk and the region west of this Human Influence 37 watercourse where nowadays, broadly speak year to year, except in certain places where ing, the reindeer do not go at all. the passing is difficult; e.g., in steeps, in dense The effect of the reindeer's grazing is lessened willow-thickets, between swamps, &c. Here also by the varying of their menu with the time the tracks, sharply marked and with all vegeta of the year (cf. LONNBERG 1909 pp. 144 ff.). tion worn away, extend for some tens or hund In spring and early summer, they show a special reds of yards, then branching out and disappear liking for Eriophorum, young shoots of Salix ing when the ground becomes less difficult. and Betula and the radical leaves of Deschampsia All the more frequent are occasional >>paths>> flexuosa, Festuca ovina, divers Poa species, from the passage of a large herd. By way of and the more tender Oarices, e.g., 0. brunnescens illustration, some observations may be cited, and 0. Lachenalii. Later in the summer, their made on the 22th of July, 1946, just above favourite food is about the same as that of the the tree line on the north-western slope of Mt bear, i.e. various kinds of juicy herbs such as Kuobberi. A herd of about 800 head went M enyanthes trijoliata, Lactuca alpina, Angelica, down a slope covered by Betula nana-Empe Rumex Acetosa, Oxyria digyna, Ranunculus trum hermaphroditum-heath and smaller patches nivalis, R. glacialis, &c.; aecording to TH. FRIES of Cassiope tetragona-heath, alternating with and E. BERGSTROM (in LONNBERG, op.c. p. 147), streaks of Ranunculus acris-meadow. After they prefer the Alchemillae to every other kind the passing of the reindeer, Oassiope was partly of food. When fleeing from the mosquitos to trampled to pieces; in the places most roughly the snow-beds, they also seem to eat much used, it would certainly not recover for several Salix herbacea, though I have not seen this years. The other scrubs were also severely reported in· the literature. On the other hand, mishandled. Here and there, the dwarf-birches they reject several plants, e.g., all ferns, all were stripped of their leaves; and most of the Equiseta but E. limosum, coarse graminids crowberry specimens, though seemingly not such as Deschampsia caespitosa and Carex so badly mauled, the day after were brown aquatilis, &c., apart from a not inconsiderable and withered. It seems doobtful whether densely number of alpine plants, e.g., the species of closed scrub heaths can arise or endure in places Saxifraga, Pedicularis, and Potentilla, which that from time to time are treated in this manner I have never seen snapped off by reindeer. (cf. Du RrETZ op.c. pp. 96 ff. ). Herbs and TH. FRIES (1913 p. 252) describes how wind grasses escaped more lightly; only where many erosion turns lichenous alpine heaths into des filed animals had trampled, the Ranunculus erts without vegetation, when grazing reindeer meadow also looked worn with bare furrows. have rooted out the lichens. Nothing of the No doubt, the effect of the trampling has been sort was, however, observed within our region. much intensified during the past decades of In these parts, there are few sandy plains of extensive reindeer breeding, the animals now the kind where grazing ·especially has this being allowed to flock in their summer pastures effect, and when arriving in the mountains, into herds of a thousand head or more. the reindeer have mostly left the lichens, turn An illustrative example of the colonization ing to grass and herbs. In addition, the reindeer of ground, laid bare by trampling, was a pen lichen occurs but sparingly within the region (renskillningshage), used for the sorting out of (cf. DEGELIUS 1943 p. 84). reindeers belonging to different owners and More important by far is the trampling by situated in the uppermost birch belt at Stalo many thousands of reindeer hoofs, a fact that njarka, on the south-eastern cove of Lake Viri can be verified by any one who has seen their haure. When such a pen is taken into use, paths furrowing a mountain slope. the ground is so badly trampled as to leave Apart from the few caravan tracks, there practically no vegetation whatever. The soil are no fixed tracks, used by the reindeer from becomes as bare as though turned up with a 38 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark harrow (cf. TH. FRIES 1913 p. 246). When I In the south-western parts of this territory, · visited this pen in 1943, it had not been used Cassiope tetragona occurs almost exclusively for five or six years. It was then overgrown patchwise, in greater number chiefly on cliffs by Poa arctica, P. pratensis ssp. alpigena and, facing north . .At first, I supposed this behaviour above all, Phippsia algida, which was so plenti to be due to the fact, that our species here is ful and luxuriant that I never saw the like of approximating its southern limit, and that it: all over an area of at least an acre, it grew it was therefore thinning, i.e. becoming less as abundantly as a fresh crop of corn. In the frequent, as usually happens in such instances. surrounding meadows and scrub heaths, a However, in 1946, I found Cassiope forming high-alpine species, like Phippsia restricted synusiae over large areas on the northern slope to bare soil, would of course not have the slight of Mt Kasakpuo1ta, the relatively narrow est chance of holding its own. The fold was crest of which is situated between Lake Kasak used again in 1946; it would be most interesting jaure and the glacier of Almallojekna. Here to know whether Phippsia survived it or not. our species occurs at a distanee of only 60 to The fact that Cassiope tetragona is very sen 65 km from its southernmost Scandinavian sitive to trampling may conceivably be one stations in Saltdalen and on Lake Ikesjaure. of the causes of its peculiar, crescent-shaped It was therefore most surprising to find pure distribution within Lule Lappmark. .As we Cassiope tetragona - heaths without the element know, one of the differences between the vege of rare arctic plants, which almost invariably tation of Torne Lappmark and that of Lule characterizes this community elsewhere in south Lappmark is that the vast Cassiope tetragona western Lnle Lappmark. At least partly, the heaths of the former territory have no counter explanation may be that Mt Kasakpuolta offers parts farther to the south. This is, of course, the reindeer but a rather poor pasture, and that principally due to climatical, edaphical and, its situation, between the lake and the glacier, probably, plant-historical factors. renders the mountain difficult of access to the However, not unlikely, the reindeer may also reindeer herds; from their usual grazing grounds play a part. Only a comparatively small num on Mt Kuobberi, Mt Kappa and Mt Stalotjakko, ber of reindem stay in Torne Lappmark during they but occasionally come to other parts of the summer; this is true especially of the region the mountain than the east slope, where Cassiope north of Lake Torne trask and the river Torne tetragona is lacking. The only mountain of alv, and of the time prior to 1919, when the Pite Lappmark, where Cassiope tetragona occurs Swedish-Norwegian Convention concerning the >>beinahe regionbildend>> (ARWIDSSON 1943 p. reindeer grazing was concluded. The Kare 229), Mt Rattjatjakko, occupies a similar, suando Lapps mostly have their >>summer lands>> isolated situation between Lake Peskehaure on the N orwegia.n side of the frontier and had and Lake Laddejaure: it would therefore seem so still more in ancient times; the Lule Lapps, that also that mountain is only incidentally on the contrary, always stayed mainly on visited by the reindeer. On the other hand, the Swedish ground. Within the northern parts large area in Lu1e Lappmark between the lakes of Lule Lappmark, Cassiope tetragona -heaths of Virihaure, Festajaure and Akajaure, where occur north of the river Stora Lule alv on the Cassiope is absent, is occupied by the heavily northern slope of the Kallaktjakko massif exploited pasture-lands of Padjelanta and Vaisa. (BJORKMAN 1939 p. 46) and south of that river Nor are there any north-facing bluffs, where in the valley of Kukkesvagge and on the .Apar our species could escape the trampling. tjakko massif in north-easternmost Sarek (TENG The effects of the trampling by reindeer WALL 1920 p. 383, 1924 p. 667, BJORKMAN may be summarized as follows. .At least in 1937 pp. 8 ff.), i.e. in areas not at all, or but some degree, it counteracts the formation of occasionally, used as pasture-land for reindeer. scrub heaths. It may possibly also reduce Human Influence 39 the formation of acidic raw humus by stirring woods everywhere show traces of cuttings, the soil and bringing less leached strata to the mostly made by the Lapps. Even in the part surface. At the same time, it creates transient of. Tjuoltavuopme situated south of the river, patches of >mew soil», where species of little where no caravan track has run since the be competitive power can settle; this effect is ginning of the century, numerous signs of old enhanced by the fact that the trodden places cuttings are to be seen. The high-grown mea form points of application for wind erosion dows, which here and there form clearings in {cf. SERNANDER 1905 p. 85, TH. FRIES 1913 the luxuriant birch forest of the Tarra valley, p. 251). Trampling thus would seem rather are natural plant communities only when they to favour the rare alpine plants, in about the are due to avalanches or landslips. Else they same way as, for instance, occasional ploughing mark the places of derelict Lapp yards, where favours Carex ligerica in Oeland (STERNER the forest reappears but very slowly, because 1921 p. 203) or road-building &c. favours the herbs choke the birch seedlings. The clear Gypsophila fastigiata in Dalecarlia (SAMUELS ing where the Tarrekaise cabin is situated SON 1910a p. 48). arose partly in connexion with the now discon Moreover, it seems doubtful whether the tinued mining of magnesite on Mt Tarrekaise; reindeer breeding of the Lapps should be regard and since the Lapps have established a new, ed as a factor foreign to the natural environ stationary summer camp at Njammilt, a hitherto ment of the alpine flora. When, probably untouched part of the birch forest is going to about the beginning of our era, and in any be destroyed. case before the 9th century (cf. e.g. WrKLUND The fact that the rich birch forests of the Tarra 1947 pp. 51 f.), the Lapps took up nomadizing valley end abruptly at the river Slitajokk and with reindeer and penetrated into the mountains, the upper parts of the valley are largely either they came across a stock of wild reindeer, in treeless or poorly wooded with birch heaths, all likelihood not much smaller than the present may be due partly to the cuttings made here breed of half-domesticated animals. In this in the sixties and seventies of the 17th century country, grazing and trampling by reindeer on behalf of the mining corporation of Lulea probably are contemporary with the alpine (Lulea silververk). At that time, mining was flora. To characterize these factors as condi carried out by >}tillmakning>}, at which proceeding tioned by civilization only because the reindeer huge log-fires were kindled on or near the rock have become more or less domesticated may that was to be quarried, a method requiring be justifiable formally, but it has hardly any enormous quantities of · wood. In a verdict foundation in realities. on the new silver mine at Mt Kiedkevare (i.e. Mt Silpatjakko), delivered in 1660 by JESPER KRUSEBJORN, Member of the Cameral Board, Lumbering of the Lapps he stated, that no wood was accessible nearer The lumbering of the Lapps in the subalpine the mine than >}at Paggevare on the river Darra birch forests constitutes an interference in the jokk>}, at a distance of about six miles (one natural vegetation, the extent of which is diffi Swedish n1ile). In this >}dell>}, birch woods grew, cult to state. Within the parts of our region · at first >}Somewhat poor and thin, but increasing visited by me, primeval birch wood, evidently steadily for twelve miles down the valley and quite untouched, was met with only in the above then becoming quite abundant>}, until at a dis mentioned part of the valley of Tjuoltavuopme. tance of 24 miles, >}both birch and spruce and Yet, primeval birch forest probably is to be some pine woodS>} were found (i borjan >magot found also in Tjarrokvagge, i.e., the valley to ringa och tunn, men okar sig utfore pa 2 mijhl the north of Tjuoltavuopme, where people ymnig och ligger utfore till 4 mijhl alt mehr och seldom or never set foot. Otherwise, the birch mehr>}, till dess >}bade biOrk och gran och nagot 40 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
tallskog begiynner>>; cited by LAGERGREN 1922 completely treeless at least south of Lake p. 8). By >>Darrajokk>>, KRUSEBJORN obviously Sallohaure (cf. fig. 13), is thus registered as means Tarrejokk; and >>Paggevare>> probably birch wood on the ordnance map of 1877. True, refers to Puokkikvare or . Puokkik, which is this does not mean much, the map probably situated exactly 10 km from Mt · Silpatjakko being as unreliable in this respect as in most and about 40 km above the conifer line. If other things; however, old Lapps have also told this be true, KRUSEBJORN's description indi me that there were woods in the valley when cates that the upper Tarra valley was much they were young. On the northern shore of better wooded in the 17th century than nowa Lake Sallohaure, there are nowadays no woods days. When the mining was discontinued about but the groves in the south-facing bluff of the year 1680, the valley had probably been Mt Tjargesvare and a few insignilicant copses cleaned of woods down to Slitajokk. In view of in some smaller steeps; yet an islet, where the extremely slow regeneration of the birch the Lapps evidently have not cared to go for immediately below the tree line, it is far from firewood, is well-wooded, although the slopes inconceivable that a thorough devastation of round the cove are almost bare. In addition, the woods would have such lasting results. here and there solitary birches of an age of up Around the great lakes west of the mountain to twenty or thirty years are to be found, which range, the birch woods seem to be more roughly do not resemble the wind-blown cripples usu used. Several facts indicate, that they formerly ally seen above the wood-limit, being straight occupied considerably larger areas than at stemmed and seemingly thriving quite well . present. Generally, the contemporary climatic On the completely treeless hills at the southern amelioration would seem to have raised the draw-lock on Kollomoloki, there are old flues ·limits of trees into higher positions. For instance, for smoking of fish, which the Lapps would this level on · the almost untouched Mt Prins never have built in a place without a supply kullen near Kvikkjokk was about 650 m s. m. of firewood. HAMBERG (1900 pp. 486 f. and iil 1807 .AHLENBERG (W 1808 p. 42), 705-710 1901 pp. 45 ff.) found birch trunks in a small m S. m. in 1909 (GA VELIN 1910 p. 7) and 715 m peaty depression on the south-slope of Mt S. m. in 1916 (TENGW.ALL 1920 p. 316); on the Mattaive, more than five kilometres east of similar Mt Snjerak, the corresponding figures the present birch line in the valley of Mellatno; were about 715 m in 1807 (W AHLENBERG 1. c.) in his opinion, the birch had vanished from and 767-782 m in 1916 (TENGWA.LL 1. c.); in this tract because of the doings of the Lapps. 1942, scattered (and possibly ephemere) young And on the south shore of Lake Virihaure, spruces were growing nearly up to the summit there are sparse, heathery birch copses on the of >>Turisttoppen>>, 804 m s. m. However, too dry, flat isle of Stuorsuolo, whereas grounds much importance should not be. attached to these on the mainland much more fit for the growth figures, since we cannot judge ·of the exactness of timber are practically treeless. The resem of W.AHLENBERG 's measurings. More signilicant blance to the area north of Lake Sallohaure is are, perhaps, the many vigorous young birches increased by the occurrence of solitary, vigorous that have, during the past few decades, germin birches (see fig. 14). ated above the present wood limit, in our region In 1698, the mine-inspector SADLIN men as well as in other parts of the Scandes (cf. e.g. tioned the Lapp village of >>Sirkeslocht>> (letter · S.ANDBERG 1940 p. 172). Apparently, climatic quoted by LAGERGREN 1922 p. 19). By Sirkes conditions cannot explain the state of things in locht he certainly meant Sirkaluokta on the the Virihaure-Vastenjaure basin, where a retro south-western shore of Lake Virihaure. It gression . of the birch woods seems mostly to appears from the diary of W.AHLENBERG, that have happened. a big summer camp was still situated here as All the valley of Vuojatatno, which is now late as in 1807. How long this camp existed Human Influence 41
Fig. 13. Reindeer pen in the valley of the river Vuojatatno. This valley, which is now, in a.U its southern half, completely treeless, is registered as birch wood on the ordnanc� map of 1877. The forests were cut largely to supply wood for the pens. Photo Sten Selander 13.7. 1946. is uncertain; it would seem to have been removed tain side, that the cause would rather have to to some other place when .ANDERSON visited be looked for in the dissappearance of the Lapps. Sirkaluokta in 1845 (.ANDERSON 1846 p. 20). Now the ·question arises, how treelessness .At any rate, there was a Lapp camp here for over several hundreds of square miles could more than a hundred years. Now WAHLENBERG be occasioned by a population, so small in (1808 p. 33) expressly declares, that no >>birch number as the Lapps, who moreover spend shrubs>> were growing at Sirkaluokta, and in but a few sumn1er weeks within the region. 1845, the slopes still were treeless (.ANDERSON Furthermore, TH. FRIES (1913 p. 158) empha I.e.). Since then, however, a few well-shaped tically denies, that the cuttings of the Lapps birches have appeared in the willow-thickets should have influenced the level of the wood on the north slope of Mt Sirkavare just above limit in Torne Lappmark to any degree worth the ancient Lapp camp. This may of course mentioning. However, conditions in Torne be a result of the recent climatic amelioration; Lappmark are not to be compared with those but the birches grow so high up on the moun- in Lule Lappmark. .As mentioned .above, 42 SELANDER: Floristic Phytogeography of South- Western Lule Lappmark
Fig. 14. Vigorous young birch above the wood limit on Mt Njerek, south of Lake Virihaure. The mountain slope, which is now occupied by low willow-thickets consisting mainly of Salix glauca and S. lanata, has probably been deforested by the Lapps. Photo Sten Selander 24.7. 1946. the Karesuando Lapps sojourn during the sum timber for reindeer pens (cf. TH. FRIES op.c. mer chiefly in the coastal land of Norway and p. 157 and pp. 245 f.). It is true that, according did so formerly to a still greater extent. The to MANKER (1944 p. 208), most large pens in Jokkmokk Lapps, on the other hand, always these parts originate from the time after the have had their pasture-lands mainly on the immigration of Karesuando Lapps at the be Swedish side of the frontier. That is why their ginning of this century. But the ancient folds wood cutting has had far greater effects than required more timber than the recent ones, that of the Karesuando Lapps. where the fencing partly consists of wire (see One reason why this lumbering has a larger fig. 12); further, the pens used for milking scope than probable a priori is the need of have become superfluous in our days, when Human Influence 43 the reindeers are milked but occasionally. In felling of trees on the arrival of the Lapps olden times, moreover, the pens for separating in spring, when the snow in the wood is still the reindeer herds (skillningshagarna) were one metre deep or more; although the stumps seldom used from year to year; the Lapps are often of a man's height, they are left to rot. often found it more convenient to cut down During the past few decades, the clean -cuts a birch copse and drag together the felled trees round the summer camps have grown much into fences than to repair an old pen. On more destructive than earlier. The present account of midges and other insects, the pens >>big camps>> (storvistena), where numerous fa used in summer must be situated at the very milies live during the whole, or best part, of tree line where, owing to the poor reproduction the summer, did not arise until the immigrated of the birch, a clean-cut may seem to last for Karesuando Lapps introduced the extensive ages. (The folds mentioned by FRIES, op.c. reindeer breeding round about the year of 1920 pp. 245 f., are used in .April, when no insects (cf. MANKER 1944 pp. 79 ff. ). It goes without are about, and they can, therefore, be located saying that these stationary camps occasion further down in the birch belt, where reforesta a far more intense cutting of the surrounding tion takes much shorter time, according to woods than the ancient, mobile tent camps FRIES only 30 to 40 years.) Nor is it possible that for long periods were located to the region althogether to ignore the probability of the above the tree limit. Moreover, since the reindeer reindeer's impeding the regeneration of woods are no longer milked, the Lapps are keeping on the clean-cuts by cropping the ground-shoots goats for that purpose; consequently, the re in spring. True, no observations of this kind growth of the woods around the camps is ren were made. But a comparison with injuries dered impossible by the voracious grazing of to South-Swedish forests, caused by far less the goats. numerous elks and ,roes, indicates that the There are several instances of the swift anni grazing of the reindeer must have such an hilation of the woods around the Lapp camps, effect. e.g., the huge clean-cut at Staloluokta which, However, the cutting of firewood round the cancer-like, frets away the fine birch forest Lapp camps surely is of still greater signi on the south slope of Mt Unna Titer, and the ficance. The fire on the open hearths of the surroundings of Vaisaluokta and Ritsemjokk huts must be kept burning day and night on just outside our region, where the landscape account of the mosquHos; characteristically has completely changed since the >>big camps>> enough, >>Smoke>> is the Lappish word for house arose about 30 years ago. This devastation hold. No matter how skilfully the Lapps may of the woods is all the more harmful because, economize with the fuel, to keep the fires on after a decade or two, the lack of fuel forces the open hearths continuously going involves the Lapps to move their huts, wherupon still a heavy drain on the supply of fuel. For the another part of the forest is doomed. For in sake of convenience, the firewood is procured stance, the treeless valley at Staloluokta which, by clean-cutting round the huts, not by thinning in 1807, before the arrival of the Lapps, was the forest, which would cause far less damage, forest-clad (WAHLENBERG 1808 p. 41), is dotted i.a., because the birch seedlings would not be with the ruins of abandoned huts, while the deprived of the necessary wind-break. (Concern present settlement is located to the mountain ing the influence of the wind on the subalpine side. birch forests, cf., e.g. , SYLVEN 1904 pp. 3 ff., There is hardly any doubt that the picture TH. FRIES 1913 pp. 153 ff., SMITH 1920 pp. 101 of the retrogression of the subalpine birch ff.; cf. also KIHLMAN 1890 pp. 63 ff. regarding woods given above is correct at least in its the wind and the .Arctic timber line). To this principal features. But for human interference, must be added the considerable waste from the the wood limit would be considerably higher 44 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark in the basin of Virihaure and Vastenjaure. tion was completely destroyed. If the birch Already SERNANDER (1900 p. 486), speaking were allowed to grow in peace, it would doubt of the very region of Virihaure, declared that, less, little by little, reappear, but since the young on account of the wood-cutting by the Lapps shoots are constantly annihilated, these valleys close to the tree limit, )>heaths arise in the place and mountain slopes, though forme.rly well of the forests to be simply incorporated by those wooded, will, in all likelihood, remain treeless.)> of the alpine region)> (translated from the A. more striking parallel to the conditions Swedish text). Attention has been drawn to of the Virihaure basin, as described above, the artificial lowering of the timber line also could hardly be asked for. elsewhere. ARWIDSSON (1943 p. 44) declared, Similar conditions are met with also at the that the tree limit in Pite Lappmark )> diirfte ... Arctic tree line in the direction of the tundra. in vielen Fallen nicht auf klimatische Faktoren Thus, AARIO (1943 p. 95) writes about the Pet sondern auf menschlichen Eingriff zuriickzufiih samo district as follows: ren seim>, and as early as in 1869, BLYTT described )>In allerletzter Zeit hat der Mensch die Ent similar results of the wood -cutting around the wicklung der Waldgrenzen nachteilig beein shielings (siitrarna, )>les chalets)>) of Sogn, i.e., flusst. Gegen Ende des vorigen J ahrhunderts in places where people live only during the wurden die Birkenbestande von Vaitolahti summer just as in the Lapp camps. He wrote Kervanto verwiistet, wie auch hauptsachlich (1869 pp. 112 f., translated from the Norwegian auf Pummanginniemi, wo sie jedoch nun schon text) as follows: im Aufleben begriffen sind.)> )>According to the peasants, birch woods used According to PoHLE (1903, 1917), a marked to grow in this region>> (i.e. at Horungerne in regress of the woods on the timber line of the south-western Norway) )>high up on the moun North-Russian tundras is entirely due to the tain sides, now nearly treeless, almost as far Russian settlers. )>Die A�t ist es, die Lichtun as to Oscarsvarden, about 3500 foot above the gen in Waldstreifen und -inseln schlagt. Der sea-level. At this level, ruins can be · seen of Mensch ist der auslOsende Faktor, der den For shielings that the owners were forced to move .mationen der Tundra das Eindringen in den further down for lack of fuel, when the woods Wald ermoglicht>> (POHLE 1917 p. 20). had been cut. In the upper parts of the Utle PoRSILD (1932 pp. 58 f.) reports the disappea valley (at the shielings of Guridalen, 2900 foot rance of birch woods occurring, according to above the sea-level) there were, according to old the priest IVAR BARDAR SON, in Greenland round persons, fine birch woods only 50 years ago. the year 1350 and later felled by the Norsemen. - - - I could easily enumerate more examp And with regard to the birch woods of Iceland, les of the same kind. If this matter were care which island, as evident from the Sagas, was fully gone into everywhere, the conditions much better wooded in ancient times than described above would probably be found to nowadays, STEFANSSON (1924 p. 85) makes be the rule. The principal cause of this de the following comments: )>Ever since the time plorable state is doubtless the reckless con of the first colonization, people have done all sumption of firewood at the shielings. During they could to destroy them)> (translated from a long succession of years, the farmers con the Icelandic text). tinued to exploit the woods growing at the The artificial retrogression of the subarctic highest altitudes, i.e., exactly where they most woods would thus seem to be a phenomenon needed protection, owing to the harshness of common to all inhabited parts of the high lati the climate, so that, in the end, all tree vegeta- tudes. IV . ALTITUDINAL BELTS
In the present work, the author has applied Stora Lule alv. Such outposts of alpine plants the division of the subalpine and alpine vege occur also on the great lakes of southwestern tation current among Scandinavian phytogeo Lule Lappmark. There is, however, a differ graphers, as follows: ence between our region and BJ6RKMAN's. A. The conifer belt, regio coniferina, up to In the valley of Stora Lule alv, the part of the upper limit of the natural reproduction of the the row of lakes whose littoral flora BJORK coniferous trees. Single pines and/or spruces MAN dealt with is situated in the regio subalpina may occur above this belt, ·although without and the uppermost regio coniferina, its altitude being able to reproduce themselves. The most above sea-level prior to the damming up at common name of the belt, viz., regio silvatica, Suorva being 425 m. Within our region, the is inappropriate, also the birch belt being a lakes belong to two different groups, the one, >>regio silvatica>>. viz. Virihaure and Vastenjaure, situated west B. The birch belt, regio subalpina, from the of the mountain range in the regio alpina and upper limit of the conifer belt to the wood line. the uppermost regio subalpina at an altitude Its proper name should be regio betulina, some of 580 and 548 m s. m. respectively, . and the conifer woods being just as subalpine, at any other, viz. Tjamotisjaure, Saggat, Karatsj and rate in those places where they ascend up to the Peuraure, situated east of the mountain range wood line. However, since terminological alter in the regio coniferina at an altitude of 297, ations should not be made unless absolutely 303, 414 and 443 m s. m. respectively. Because necessary, I have kept the customary term. of the eastern situation of Lake Karatsj and C. The alpine belt, regio alpina, usually sub Lake Peuraure and the lowering of the altitude divided into a regio alpina inferior, a regio alpina belts to the west, the levels of these two lakes media and a regio alpina superior. Regarding correspond, in the relevant respect, to a con the reasons why I subdivide it only into a regio siderably lower altitude than that of the lakes alpina fertilis, up to the more or less sterile, west of Suorva. Furthermore, the . difference high-alpine boulder-lands, and a regio alpina between high-water and low-water in the lakes sterilis (or superior), see below p. 52. Cf. also of Tjamotisjaure-Saggat and Karatsj-Peurare, TENGWALL 1920 pp. 285 f. whose feeding areas partly consist of woodland instead of alps, is not as great as in the source lakes of Stora Lule alv, and therefore the in »Regio alpina descensa» undation zone is not as strongly developed, nor is its flora as rich in species or individuals as BJORKMAN (19"39 p. 5 and p. 194 ff.) proposed that of the valley of Stora Lule alv. a special >>regio alpina descensa>> as a designa West of the mountain range, the vegetation tion of the >>regional alpine outposts on a lower of the inundation zone has been investigated level>> (op.c. p. 5; transl.) in and close to the more in detail only on Lake Virihaure. To inundation zone of the source-lakes of the river speak of a >>regio alpina descensa>> there would 46 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark be inappropriate, since the alpine heath extends The mosses were not set down. Sagina inter down to the shore round the greater part of media occurred in a similar habitat a few 100 m of the lake. On the whole, however, the flora away. has the same character as that of the lakes at Here the number of species generally not a lower altitude. For instance, the following occurring in the surroundings is considerably species were noted at N jerek on the south larger than at Njerek, including i.a. Poa herje shore of Lake Virihaure (24.7 1946; see fig. 15): dalica, Juncus biglumis, Sagina intermedia, Ranunculus nivalis, Cardamine bellidifolia and Equisetum arvense f. ri Saxifraga tenuis parium S. aizoides Saxifraga foliolosa in addition to the plants E. scirpoides Epilobium anagallidifo · characteristic of inundated ground. If the Agrostis stolonifera var. lium shelving shore were longer, this locality would arctica Pinguicula vulgaris be a perfect counterpart of the >>regio alpina Phippsia algida Philonotis tomentella descensa>> of BJORKMAN. Carex Lachenalii Bryum pseudotriquetrum C. fusca Schistidium angustum The vegetation of the inundation zone of C. rufina Onchophorus virens the big lakes east of the mountain range differs Salix herbacea Drepanocladus revolvens even more from that of the surroundings. S. polaris Cratoneurum sp. When dealing with its flora, I shall say a few S. hastata (about one dm Campylium stellatum worlds also concerning certain other sylvan high) Hygrohypnum spc. (the Polygonum viviparum mosses determined by habitats of alpine plants. Cerastium trigynum 0. MARTENSSON). The inundation zone, in the wide sense of Ranunculus reptans BJORKMAN (op.c. p. 195), corresponds to the Cardamine pratensis coil. Dominant : Agrostis, eulitoral of NAUMAN {1920) and LUNDQVIST (seedlings) patchwise Saxifraga (1927) and the Amphibiontenstufe of Du RIETZ Saxifraga foliolosa aizoides and Philonotis (1940b). However, the difference between the Except Equisetum arvense f. riparium, Ra vegetation below the sharply marked line, nunculus reptans, Carex rufina, which is rather indicating the water-level at the breaking up frequent at the mouths of brooks on western of the ice, and that above it (>>the lower horizon>> Virihaure, and Agrostis stolonifera var. arctica, and >>the upper horizon>> of BJORKMAN I.e.) is which is characteristic of such habitats in parts so great that a fusion of these two zones into of the region, all these plants are quite common one seems rather inappropriate. The line just on the surrounding low-alpine heath, only mentioned is rendered conspicuous, inter alia, by Phippsia algida and Salix polaris constituting the sudden disappearance of the willow thickets, probably mainly due to ice-pressure, and, r1 weak high -alpine element. broadly speaking, of a closed plant cover; At Staloloukta, i.e. the south-eastern cove · of Lake Virihaure, the following plants were very often it is also marked by a >>Shore barri noted in the inundation zone, which is bere cade>>- (cf. THUNMARK 1931 pp. 11 ff.). Below it, situated in the regio subalpina (1.8 1946): in the inundation zone proper, the ground is above water throughout the winter; frost Equisetum arvense f. ri Ranunculus reptans heaving is, therefore, sometimes rather intense. parium Cardamine pratensis coli. However, it is generally covered with snow, Agrostis stolonijera var. (seedlings) arctica C. bellidifolia which largely mitigates the effect of the frost. Poa herfedalica Saxifraga foliolosa At the spring flood it becomes submerged even Oarex fusca S. tenuis before the commencement of the growing season Juncus arcticus Parnassia palustris and does not emerge out of the water until the J. alpinus var. alpestris Primula stricta middle of July or, in the case of the lowest J. biglumis Pinguicula vulgaris Salix herbacea P. alpina parts of the zone, not until August; the ensuing Ranunculus nivalis Leontodon autumnalis zonation (cf. BJORKMAN op. c. pp. 194 f.) cannot Altitudinal Belts 47
Fig. 15. The inundation zone on Lake Virihaure below Mt Njerek. To t.he right the sterile 1>shore barricade1>. The vegetation to the left, still partly submerged, consists chiefly of Agrostis stolonifera var. arctica, with patches of Saxi fraga aizoides and Philonotis tomentella. Photo Sten Selander 24.7. 1946. be dealt with here. The flora of this zone is ra means as pronounced as those of the inundation ther peculiar. It comprises some isoetids, above zone proper; it differs from the surrounding syl all, Scirpus adcularis, Ranunculus reptans, van vegetation chiefly by the absence of some Subularia aquatica, and Callitriche veTna; not plants, e.g., Vaccinium Myrtillus and a few a few terrestrial lowland plants, most of which, other dwarf scrubs and mosses (cf. BJORKMAN e.g., Equisetum arvense var. ripaTiurn, De op.c. p. 197) and by the preponderance of the schampsia caespitosa var. glauca, Juncus alpinus, meadow communities over those belonging to Prunella vulgaris, Veronica serpyllijolia, and the heath series. The plant cover is closed, as Galium palustre, in our region are more or less distinct from that of the proper inundation zone. exclusively restricted to habitats of this kind, The alpine species occurring here, e.g., Selaginella wheras some others, e.g., Agrostis stolonijeTa, selaginoides, Equisetum variegatum, Tojieldia A. tenuis, Alopecurus aequalis, Carex Oederi, and pusilla, Salix lanata, Thalictrum alpinum, Saxi the all-regional Polygonum viviparum, occur also jraga stellaris, S. aizoides, Bartsia alpina, Peta in other habitats; and quite a number of alpine sites jrigidus, and Saussurea alpina, are, as species, the majority of which are alien to the a rule, at least as abundant in the eutrophic neighbouring vegetation. Most of these plants fens further up in the woods; the majority of occur but in single specimens or sparse patches, them are quite common also elsewhere in the and the main part of the ground is devoid of regio coniferina. vegetation. Besides these species (except Salix lanata, BJORKMA.N's >>Upper horizon>> is flooded only Petasites jrigidus, and Saussurea alpina), a for a week or so in the early summer. Conse total of 31 alpine species were observed in the quently, its characteristics are not by any inundation zone proper. Many of them recur 48 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark in similar habitats elsewhere in the conifer Sedum Rosea Gentiana nivalis (accord belt, e.g., on river banks, alluvia on the brooks, Saxifraga oppositijolia ing to WESTLING 1906 Saxijraga cernua also on footpaths) &c., where the alpine element of the vegetation S. rivularis Veronica alpina (also on is often as strong as on the lake shores (cf. Primula stricta ( charac footpaths, ibid.) SERNANDER 1899 pp. 18 ff.). On alluvium teristic of the zone) Antennaria alpina among rocks near a cataract, for instance, the following alpine plants were noted (at the .An even greater number of alpine plants river Kamajokk below the Tourist Station of were noted only on bluffs and talus slopes, Kvikkjokk, 3.8 1946): always or nearly always facing south. This applies to: Equisetum scirpoides Ranunculus nivalis Poa alpina var. vivipara Saxijraga foliolosa W oodsia alpina Draba nivalis Salix herbacea S. tenuis W. glabella Saxifraga nivalis Oerastium arcticum Polystichum Lonchitis S. groenlandica Poa glauca Potentilla nivea .Among the alpine plants of the inundation Oarex rupestris P. Ohamissonis zone, the following were found, within the coni 0. atrofusca Rhododendron lapponicum Viscaria alpina Veronica fruticans fer belt of our region, exclusively in this zone Silene acaulis Erigeron politum (and occasionally in such habitats as the fore Draba norvegica E. uniflorum mentioned one): Dr. hirta Deschampsia alpina Oardamine bellidifolia BJORKMAN ( op.c. pp. 194 ff.) assumes that Oarex Lachenalii Saxifraga foliolosa 0. Bigelowii (probably S. tenuis the occurrence of alpine species in the inunda overlooked in the Oassiope hypnoides (also tion zone should be connected with the tempe woods) once seen on the bank rature inversion in highland valleys. However, Salix polaris of a brook in a wood the mere topography of our region seems to be Oerastium trigynum land mire) imcompatible with this hypothesis; since appre 0. arcticum Veronica tenella ciable temperature inversions hardly occur In the alpine belt, these species invariably in basins as shallow as those in question here. belong to the flora of the snow-beds. .A few (In eastern Torne Lappmark, where the country plants of the same general character were noti is as flat as a pancake, many alpine plants are ced, apart from their stations in the inundation found on stony river banks far down in the zone, also on footpaths, namely: woodland, e.g. Sedum Rosea, Gentiana nivalis, Salix herbacea (once seen Sagina Linnaei Astragalus alpinus ssp. arcticus, A. frigidus, on damp forest ground Loiseleuria procumbens Saussurea, &c. at the river Vittangi alv near too) Gnaphalium supinum Vittangi, 248 m s. m. li'or other instances of Below the pine limit, J uncus biglumis, Mi the occurrence of alpine plants at low altitudes nuartia bijlora, and Sibbaldia procumbens were in level country, cf. SERNANDER 1899 pp. 18 ff. observed only_ on footpaths; Phyllodoce coe and p. 35.) rulea is much more frequent on paths than in Furthermore, BJORKMAN's argumentation im plies that alpine plants should always be favoured other places in the conifer belt (cf. BERGSTROM. 1910 pp. 221 f.). by cold. We know too little, however, about Furthermore, the inundation zone has a the cardinal temperatures in arctic and alpine considerable number of alpine species in corn plants to make such an assumption. Certain mon with bluffs and screes facing south, viz.: observations seem to indicate that the maximal temperature, at least in some such plants, is Equisetum scirpoides Juncus trifidus higher than we are inclined to believe. This is Agrostis borealis Luzula spicata Trisetum spicatum Oxyria digyna illustrated by the above-mentioned occurrence Oarex atrata Oerastium alpinum of Phippsia algida in an old reindeer pen in Altitudinal Belts 49
Fig. 16. Convallaria majalis just above the inundation zone on Lake Karatsj , below Mt Farforita. Photo Sten Selander 22.7. 194 7. the birch belt, where it luxuriated and mul in the lowland should prove that their rare tiplied to an extent unparalleled in its normal occurrence in the conifer belt of Lapland cannot habitats. In Greenland, it >>quite paradoxically always be attributed only to climatic conditions. is a very common weed in the cold frames Nor does the temperature inversion affect under glass, where it develops to 'gigantic' the south bluffs where, on the contrary, the flaccid specimens>> (PORSILD 1932 p. 25); Stel temperature is considerably higher than in the laria longipes ( coll.) also >>creeps into the cold neighbourhood (cf. ANDERSSON & BIRGER 1912 frames and grows very well under glass>> (ibid. pp. 65 ff., FRODIN 1915 pp. 200 ff.); in spite of p. 29). However, the most striking example is, this, the number of alpine plants is greater there perhaps, Koenigia islandica in Iceland: >>Even than anywhere else within the conifer belt. on the banks of brooks with water from the hot It should, further, be noticed that the flora springs I have observed it, in places where the of the inundation zone also includes several temperature of the running water was about southern species, which here have their last 70° 0.>> (GRONTVED 1942 p. 120.) This instance outposts towar.ds the mountains, viz.: is especially interesting, because Koenigia is Convallaria majalis Limosella aquatica annual and thus, evidently, able not only to Sagina nodosa Veronica serpyllifolia exist under these unusual circumstances, but Elatine Hydropiper Galium uliginosum Lysirnachia thyrsiflora G. palustre also to reproduce itself. Moreover, the great number of high-alpine species easily cultivated Convallaria and Galium uliginosum do not
5-496149 Sten Selander I 50 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark grow in the inundation zone proper, but in of highly acid litter and raw humus. The small thickets on its upper border (cf. fig. 16). Most minority that are more or less tolerant to acidic of the spontaneous appearances of Agrostis raw humus, e.g., Pedicularis lapponica, Phyl tenuis also occur in the inundation zone. Just lodoce coerulea, certain Hieracia, &c., are by outside our region, some more southern species . no means rare in the conifer belt in places where are encountered in these habitats, viz. Alisma the soil profile is comparatively mild. Several
Plantago-aquatica, Seirpus palustris, Myosotis · alpine plants, which, within our region of pre laxa ssp. caespitosa (probably reported from dominant spruce forests, occur but scarcely in our region in ANDERSON. 1844 p. 15 sub M. the conifer belt, are known to occur regularly palustris), Scutellaria galericulata, and Mentha in the woodland of other districts where light, arvensis. , dry pine forests pr�dominate and, consequently, The resemblance of ·the flora of the inunda the layers of acid raw humus are less thick tion zone to that of �he south-facing talus slopes and compact. This applies, for instance, to is enhanced by the fact that these habitats Arctostaphylos alpina, Loiseleuria procumbens, also have some lowla1�d species in common, viz.: Juncus ·trifidus, Carex Bigelowii and Lycopodium alpinum (cf., e.g., SERNANDER 1899 pp. 5 ff., Carex ornithopoda (also in Juncus alpinus a few eutrophic fens) Alchemilla filicaulis BERmlTROM 1910 p. 224, TENGWALL 1925 pp. C. Oederi Prunella vulgaris 745 f.). The rest are restricted to localities of various kinds where raw-humus formation is in Carex panicea also belongs here, though most significant or non-existent, and in most cases, of its localities are eutrophic fens, and, likewise, where bare mineral soil is available. Poa palustris which occurs, however, chiefly on Furthermore, most, if not all, alpine plants water-meadows. Rosa majalis is found exclusiv are evidently obli�ative heliophytes and thus ely on screes and lake shores, though not in the confined to habitats lacking shadowing trees inundation zone proper. and shrubs. Both these conditions are mani These facts decidedly go against BJORK fested in the habitats of the conifer belt where MAN's hypothesis. In particular the occurrence alpine plants are chiefly met with, i.e. fens, of a pronouncedly southern element in the flora talus slopes, precipices, rock fissures, footpaths, of the inundation zone and the similarities of alluvia, river banks, brook ravines, and the this flora to that of the south -slopes would be inundation zone. (No alpine plants have been quite incomprehensible if the inundation zone observed in shaded places on rivers and brooks owed its floristic character to temperature or on the wooded parts of the tal us slopes.) inversion. Another limiting factor is nutrient deficiency However, it would seem that an explanation of the soil, even when not combined with raw of the behaviour of the alpine plants in the humus formation. Among the fens, only the woodland is obtainable without resorting to a eutrophic ones support a number of alpine special >>regio alpina descensa>>. Most of the species worth n1entioning. On the whole, oli relevant ecologic factors have already been gotrophic habitats, e.g., south-bluffs built up pointed out by BJORKMAN himself (Le.). of barren rocks, hardly present arty alpine The pedology of the Scandes is still so obscure plants (cf. below pp. 176 f.), except where they that the utmost caution is needed when speaking are moistened by more or less aged water, of the soil factor. Yet as a pure hypothesis, e.g., water trickling from the rock-faces (>>sipper without any claim to definiteness, the following vatten>>, >>bergsega>>), or the water of lakes and exposition may be defensible: rivers temporarily flooding the ground. On the Most alpine plants, even those which are other hand, deposits of limestone and other indifferent to the Ca content of the soil, cannot basic rocks often harbour rather a rich alpine or stand leached podsol soils with surface layers arctic flora, even though situated hundreds of Altitudinal Belts 51 kilometers from the mountains or the tundra In other words, for such plants as cannot fulfill {cf., e.g. , POHLE 1903 pp. 8 ff., SAMZELIUS 1890 all their functions under water, the growing pp. 186 f, SERNANDER 1899 pp. 13 f., BERG season has a length of only about two months. STROM 1910 pp. 219 f., TENGWALL 1925 pp. In this connection it should be remembered . 744 f.). that in the regio alpina many alpine plants of .As to the flora of the inundation zone, the the inundation zone belong to the vegetation of significance of the nutrient content of the the snow-beds (cf. above p. 48), i.e. to plant water is evident from the difference in the com communities stamped above all by the abnormal position of this flora on the different lake-sy shortness of the growing season; according to stems. .All the rivers feeding Lake Peuraure CLEVE (1901 p. 83), in our region the whole and Lake Karatsj rise in oligotrophic areas yearly life cycle of typical snow-bed plants, of seve schists or primary rock; in the inundation such as Ranunculus nivalis, R. pygmaeus, and zone of these lakes, the alpine plants are very Salix herbacea, from the melting of the snow few in number, as regards both species and to the ripening of the seeds, is passed through specimens� J.J ake Saggat and Lake Tjamotis in 38, 49, and 52 days respectively. jaure are situated on a much lower level; in However, the shortened growing season per se spite of this, their inundation zones are consi cannot, except perhaps in rare instances, be derably richer in alpine plants, apparently be the direct explanation of the occurrence of cause the source-streams of these lakes have · alpine plants in the inundation zone� For their arise in areas partially built up of less alpine species, partly the same as on the lake barren rocks. On the lakes of the valley of shores, also appear, and in still greater number, the river Stora Lule alv, the feeders of which on south-facing steeps and talus slopes where drain, i.a., the eutrophic Virihaure-Vastenjaure the growing season is longer than anywhere basin, the inundation zone presents a flora else within our region (cf . .ANDERSSON & BIRGER which is by far th(3 richest in alpine species. I.e.). It should be noticed that the supply of dia Therefore, the abbreviation of the growing spores cannot be decisive, since practically all season in the inundation zone must be princi alpine plants at home in the inundation zone pally of indirect significance, by eliminating oecur also in the areas of seve schists. competing plant species. The comn1ensals of However, the decisive fact.or is, probably, the sylvan communities cannot master the competition, which BJORKMAN (I.e.) mentions conditions of the inundation zone proper, only incidentally (cf. SERNANDER op. c. p. 14). namely, apart from the short growing s.eason, In many cases, high-alpine plants wol.lld seem chiefly the intense frost-heaving, the ice-pres to be restricted to the high altitudes mainly sure, and the high ground-water table. .Among through their lack of competitive power. They the lowland plants, mainly the noncompetitive can hold their own only on the summits, where, isoetids and a few specialized graminids, gro i. a., the. growing season is so short that all wing in thin patches, thrive there. This gives plants are eliminated whose life cycle is not the alpine species a chance. From this point so extremely swift as that of the high-alpine of view, it is understandable that the inundation species. This point of view is far from new; zone harbours a number of southern species, too, but it should not be disregarded when we try which here, on the outskirts of . their areas, to explain the presence of alpine plants in the have a very small capacity of holding their inundation zone. For a shortening of the growing own. season is manifest also in these localities: the It should, moreover, be observed that com habitats do not emerge out of the water until petition is reduced also in other sylvan habitats late in the summer, and the autumn frosts of alpine plants, e.g., talus slopes, river banks, come on earlier than on the mountain slopes. footpaths, &c.; even in the fens, the alpine 52 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
species occur only in places where the cover of mountains within our region nowhere come up vascular plants is not closed. The footpaths, to the middle-alpine belt. Round Lake Viri for instance, differ from the adjacent ground haure, V accinium M yrtillus occurs only very . chiefly in the presence of bare soil and the sparsely, usually in small, thin patches, and is absence of litter and competing plants. practically missing on many mountains, as is Thus, the distribution of alpine plants within the case with Deschampsia flexuosa and most the conifer belt would seem not to be determined other indicator species of the Myrtillion alpinum only by temperature, but as much or more by (Du RrETZ 1942a). Hierochloe alpina-heaths edaphic factors, above all, such as decrease are entirely non-existant; except for Oarex or abolish the competition from sylvan species. Lachenalii- Oarex Bigelowii-communities which, The inundation zone, moreover, obviously has however, often descend to low altitudes (cf. close counterparts among the shore zones of TENGWALL 1920 pp. 391 f.), other grass heaths, certain lowland lakes. To retain the term of especially those belonging to the J uncion tri >>regio alpina descensa>>, invented by BJORKMAN fidi (Du RIETZ op.c), are of very rare occurrence; in order to designate an essentially local pheno and a Oassiope tetragona-heath is developed menon, would therefore be an unnecessary, over large expanses in only one locality. On not to say inappropriate, terminological burden. the other hand, Oassiope hypnoides-heaths are common even on plane ground, e.g., in the · low-alpine district round the western coves of Regio alpina inferior and regio alpina Lake Virihaure, where they extend down to media the lake, 580 m s. m., into an unquestionable A few words may be justified here to explain regio alpina inferior. Especially on mountains, why I do not use the terms regio alpina inferior such as J eknaffo, which are strongly marked by and regio alpina media. These altitude belts were solifluxion, meadow communities come up to first proposed, though not named, by VESTER the lower limit of the high-alpine belt (regio
GREN (1902 pp. 264 f.) on the basis of observa alpina sterilis) without · showing any pro tions made in the amphibolite region of Sarek. nounced change except for a gradual floristic Later, a similar division, although less sharply depauperation; on the whole, meadow communi defined, was set forth by HEINTZE (1913). The tics are characteristic of >>wet>> mountains such as middle alpine belt (regio alpina media BJORK those here in question (TH. FRIES 1917 p. 36). By MAN 1939) was invented by Du RIETZ who, this I do not imply that there should be no dif among Swedish phytogeographers, has bestowed ference between the low-alpine belt and the midd most interest on these questions and studied le-alpine one, the whole of the vegetation often them in a thoroughgoing way, mainly in Torne modifying its character at an ascent of 200 or Lappmark and central Scandinavia (Du RIETZ 300 m. I merely wish to state that I am unable 1928, 193oa; 1930b, 1942a, 1942b, 1943, &c.; to master the difficulties connected with the cf. also TH. FRIES 1917). establishing of a definite limit. However, neither in the tracts investigated by In order to illustrate the gradual manner in Du RIETZ and HEINTZE nor in the amphibolite which the vegetation changes with increasing area of Sarek conditions tally with those pre altitude, the altitude limits of the vascular plants vailing in the western, extremely calcareous part along two lines will be given below, one line of our region. running from the middle summit of Mt J eknaffo In these parts, most of the plant commu north-westwards to the upper limit of the nities constituting the difference between the willow thickets at Staddajokk, and the other two belts are either lacking, or rare. This is running from the highest summit of Mt Kerke true, e.g., of the bilberry heath, except to the vare south-westwards to the river Vejej okk. north of Lake Vastenjaure where, however, the The altitudes were determined with an aneroid. .Altitudinal Belts 53
.At the start from Staddajokk ' (28. 7 1944), 1575 m s. m. on Mt Jeknaffo is due to the fact the aneroid showed 800 m, and at the return, that the line crossed a large snow-field there. 795 m; when immediately .controlled on Lake The lower limit of the high-alpine boulder terrain Kappaluoppal (altitude ace. the ordnance map on this mountain is situated about 1440 to 699 m s. m.), it showed 700 m. On Mt Kerke 1450 m s. m. Only patches of such terrain are
vare (30.7 1944), the aneroid showed 830 m to be found on l\H Kerkevare; that is why com , both at the start from Vejejokk and at the paratively many species occur on the summit return, and 1560 m on the summit (altitude ace. proper of this mountain. The lower limit of the the ordnance map 1563 m s. m.). That no high-alpine belt can here be put at about 1415 species were registered between 1675 m and to 1425 m s. m.
Mt Jeknaffo m Mt Kerkevare The summit: 0 1700 Poa spc. (cfr laxa ssp. jlex1wsa.; sterile), Ranuncu- 1695 lus glacialis Lttztda confusa 1690 Deschamps'ia alpina. 1680 Phippsia algida, Saa.�ifraga r·ivularis. 1675 Lycopodium Selago. 1575 1563 Lycopodium Selago, Deschampsia alpina, Poa alpina var. 'Vivipara, Phippsia algida, Luzula confusa, Cerastium arcticum, Ranunculus glacialis, R_ pyg maeus, Saxifraga opposit·ifolia, 8. cernua. 1555 Saxifraga tenuis. Ranunculus pygmaeus, Cardamine bellidifolia. 1550 Sagina intermedia Sagina intermedia. 1530 Cardamine bellidifolia. , 1520 Festuca vivipam. 1490 Salix herbacea x polaris, Saxifmga groenlandica_ Ceras.tium arcticum, Saxifraga oppositifolia. 1480 Silene acaulis. Festuca vivipara, Silene acaulis. 1460 Draba lactea. Salix polaris, Saxifraga groenlandica. 1450 Salix herbacea. Poa herjedalica, Draba lactea, Dr. nivalis, Saxifraga 1440 nivalis, Potentilla hyparctica, Antennaria carpa thica. Carex Lachenalii, Sagin(l, caespitosa, Ranunculus 1430 Oxyria d'igyna. nivalis, Draba alpina, Saxifraga cernua, Pedicu laris lapponica. 1425 Carex Lachenalii. Eriophorum Scheuchzeri, Viola biflora. 1420 1415 Saxifraga foliolosa. Carex rupestris, Salix herbacea x polaris, D1·aba nor 1410 Polygonum viviparum. vegica, Sedum Rosea. Poa alpina var. vivipara, Festuca ovina, Carex Bi 1405 gelowii, Oxyria digyna, Sax·ifmga foliolosa, Saus surea alpina. Trisetum spicatum, Juncus biglumis, Cerastium tri 1400 Pedicularis hirsuta. gynum, Minuartia biflora, Saxifraga tenuis, Cas siope hypnoides. 1395 Carex Bigelowii, Draba alpina, Cassiope hypnoides. 1390 Ranunculus nivalis. Pedicularis hirsuta. 1385 Polygonum viviparum. 1375 1360 -.'Yinuart·ia biflom, Sa'ussurea alpina. 54 SELAKDER: Floristic Phytogeography of South-Western Lule Lappmark
Mt Jeknaffo m Mt Kerkevare Gnaphalium supinum. 1350 Equisetum m·vense, Eriophorurn angustifolium, Salix 1345 herbacea, Cardamine pmtensis ssp. angustifolia, Petasites frigidus. Luztda Wahlenbergii, Cerastium alpinum. 1340 1320 Erigeron unalaschkense. Campanula uniflora. 1310 Viola biflora. Empetrum hermaphroditum 1300 Trisetum spicatum, Luzula fVahlenbergii, Cerastium trigynum, Cardamine pmtensis ssp. angustifolia, Antennaria carpathica, Gnaphalium supinum. Tofieldia pusilla, Potentilla Crantzii, Dryas octo 1295 petala, Pedicularis flammea. Carex saxatilis. 1275 1260 Festuca. ovina, Salix polaris. 1255 Diapensia lapponica. 1250 Empet1·um hermaphroditum. 1230 Vaccinium uliginosum Astragalus alpinus. 1225 Pedicularis lapponica. M elandrium apetalurn. 1220 Carex rupestris, Sedum Rosea, Dryas octopetala, Campanula uniflora. Salix reticu,lata. 1215 1210 Tofieldia pusilla, Cerastium alpinurn, Thalictrurn alpinum, Potentilla Crantzii, Sibbaldia procum bens. 1205 Saxifraga nivalis. Poa alpina, Sagina Linnaei, 'Phalictrum alpinum. 1200 (Here a small precipice): Luzula spicata, Salix reti ctt.lata, 8. lanata, Sagina Linnaei, Draba nivalis, Arabis alpina, Vaccinium Vitis-idaea, Campa nula rotundifolia, Erigeron uniflorum, Hieracium alpinum. Poa glauca, Arabis alpina, Gentiana. nivalis, Erige 1190 Draba norvegica, Astragalus alpinus. ron uniflorum var. eriocephalum, . E. unalasch kense. Sibbaldia procumbens, Veronica pumila. 1170 Equisetum scirpoides. 1160 Antennaria alpina. 1150 Calamagrostis lapponica, Care.x ·vaginata, Cerastium glabratum. Carex capillaris, Ranunculus acr·is, Erigeron uni- 1140 florum, Antennaria dioeca, A. alpina, Taraxacum c·roceuru (coli.). Juncus trifidus. 1130. Cerastium fontanum ssp. scandicum. 1120 Bartsia alpina. 1115 Pedicularis flammea. 1110 Equisetum variegatum, Kob1·esia myosuroides, Carex parallela, C. atrata. Vaccinium uliginosum. 1105 1095 Ca1·ex capillaris. 1090 Poa pratensis ssp. alpigeria. Phyllodoce coerulea, Diapensia lapponica. 1085 Vaccinium M yrtillus. 1080 Saxifraga aizoides. Calamagrostis lapponica. 1075 Pyrola grandiflora. Veronica alpina. 1070 1060 Care;-c nort•egica, C. atrofusca·, Chamorchis alpina, Ran·uncul·us acris, Rhododendron lapponicum. Altitudinal Belts 55
Mt Jeknaffo m M t Kerkevare Luzula arctica. 1050 Equisetum arvense, Anthoxanthum alpinum, Poa herf edalica. 1040 M elandrium apetalum. 1035 Luzula multiflora ssp. frigida. Ohamorchis alpina. 1030 (Here a small precipice): Phleum comrntttatum, Salix hastata, Rumex Acetosa ssp. arifolius, Melandrium rubrurn, Parnassia palustris, Angelica Archange lica ssp. norvegica, Gentiana nivalis, Veronica al pina, Gnaphalium norvegicum, Taraxacum cro ceum ( coll.). 1025 Selaginella 8elaginoides, Juncu:s triglurnis, Pingui cula alpina. Anthoxanthum alpinurn, Poa pratensis ssp. alpigena, · 1020 Salix glauca, Stellaria nemorum ssp. montana. Du.zula rnultiflora ssp. frigida, Oerast-iurn glabra- turn. . 1015 Luzula parviflora, Minuartia stricta, 'Prollius euro paeus, Draba hirta. Agrostis boralis, Oarex vaginata. 1005 Koenigia islandica. 1000 Equisetum pratense, Hier·ochloe odorata, Pyrola mi nor. 995 Ooeloglossum viride ssp. islandicum. Oarex atrata, Salix arbuscula, Rumex Acetosa ssp. 990 Athyrium alpestre, Alchemilla Wichurae. arifolius. 985 Alchemilla M'lt.rbeckiana. Hierochloe odorata, Oarex Bigelowii x fttsca, Oera- 970 Luzula sudetica. stiurn fontanum ssp. sca·ndicurn. Salix lanata. 965 Oarex atrofusca. 960 Equisetum pratense, Oarex norvegica, Parnassia pa- 955 Eriophorum angustijolium, Oarex saxatilis. lustris, Bartsia alpina. Pinguicula vulgaris. 950 Euphrasia frigida 945 Alchemilla M urbeckiana. 940 Vaccinium Myrtillt ls, A ntennaria dioeca. 935 Alchernilla glomerulans. Oarex dioeca. 930 Equisetum scirpoides, Minuartia stricta, Hiera.cium 920 alpinum. 910 Lycopodium alpinum, Aconitum septentrionale. Salix glauca. 900 Phleum comrnutatum. 890 Cystopteris montana, Stellaria calycantha. 885 Oarex fusca. 870 Oarex fusca. Luzula parviflora,, Salix myrsinites. 865 Equisetum variegatum, Leuchorchis albida, Alche- 860 milla glomerulans, Pyrola minor. 855 Lycopodium annotinum, Poa arctica, Betula nana, Solidago Virgau rea. Betula nana. 840 Salix myrsinites, Trientalis europaea. Loiseleuria procumbens. 835 Ar·ctostaphylos alpina. 830 Vejejokk. Lycopodium alpinum, 8alix lapponum, Rubus 825 Ohamaemorus. Kobresia rnyosuroides, L'ltzula sudetica. 815 Oalamagrostis neglecta, Saxijraga stellaris. 805 Upper limit of willow thickets. 795 56 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
.Among the species observed on the western Stellaria calycantha Loiseleuria procumbens slope of Mt Jeknaffo, the following were not Sagina caespitosa Phyllodoce coerulea Minuartia rubella Oassiope tetragona touched by the section: Viscaria alpina Arctostaphylos alpina Lycopodium annotinum Saxifraga aizoides Rubus saxatilis Primula stricta Selaginella selaginoides Potentilla nivea Potentilla nivea (coil.) Gentianella tenella Equisetum limosurn P. Ohamissonis Astragalu.s norvegicus Euphrasia frigida Athyrium alpestre Alchemilla Wichurae Oxytropis lapponica E. lapponica Poa arctica Epilobiurn anagallidifoli- Viola epipsila Petasites frigidus Oarex parallela um Epilobium anagallidifo- Arnica alpina 0. aquatilis Pyrola grandiflora lium . Juncus m·cticus Rhododendron lapponi- J. triglumis cum .A main feature of the alpine vegetation to Luzula spica.ta Oassiope tetragona the south and south-east of Lake Virihaure Ooeloylossurn viride ssp. Vaccinium V itis-ida,ea is that several among the dwarf scrubs charac islandicum Oampanula rotundifolia teristic of Myrtillion and Empetrion (Du RIETZ Salix hastata Hieracium nigrescens Viscaria alpina (coil.) 1942b) either belong to the species, generally of rare occurrence, that were not touched by The same is the case with the following spe the sections, or, on Mt J'eknaffo, were met with cies found on the western slope of Mt Kerke only in the lowest pa!'t of the section, at· an vare: altitude of about 800 m s. m., where some hills
Oystopteris fragilis Oarex glacialis of leached morainic material were crossed. It 0. montana Juncus arcticus deserves to be mentioned here that one of the Woodsia glabella J. biglumis species separating Myrtillion from Empetrion, Agrostis borealis Luz·ula arctica viz. Deschampsia flexuosa, was not observed Poa glauca Leuchorchis albida on these carefully investigated slopes. It may, Eriophorum Scheuchzeri Salix arbuscula Oarex microglochin Koenigia islandica of course, have been overlooked. However, it 0. nardina Stellaria longifolia can scarcely be plentiful. V. OR IGINS OF THE FLORA
Fossil finds V. cfr Oxycoccus coli., Empetrum nigrum coli., and M enyanthes trifoliata ). Of course, the fact That a plant has survived at least the latest that a plant lived in Scandinavia during an in glaciation within Fennoscandia can, of course, terglacial is, on the whole, no proof of its being be proved with certainty only by findings of fos a glacial survivor, since it may have died out sils. The DTyas flora found by NATHORST and here during the subsequent glaciation. However, others in Danish and South-Swedish late-glacial if for other reasons it can be supposed to be a deposits (cf., e.g., ANDERSSON 1896 pp. 15 ff.) >>hibernator>>, this assumption becomes more pro is, however, of no immediate significance with bable, if the species is also found in interglacial regard to the problems discussed here, .since the deposits. complete lack of such fossils in deposits north The first finding of late-glacial plant fossils of the Mid-Swedish moraine sweep makes it very west of the ice-divide was made in the early questionable whether the South-Scandinavian nineties by ToLF (1893 p. 26), who found leaves Dryas flora had any connexion with the popula of Populus tremula, Betula nana, Dryas octopetala tions of the Fennoscandian mountains. More im and Phyllodoce coerulea in the bottom layers of portant in this context are the fossil finds in a bog at Ede in Brunflo (central Jamtland), interglacial layers of Pilgri!llstad and other pla in strata which did not contain any pine nee ces in Jamtland, reviewed by KULLING (1945 pp. dles. Even in 1906, A. G. HoGBOM (1906 p. 191, 29 ff. and p. 51). They include macrofossils of footnote) interpreted these fossils as originating·
Salix polaris, S. herbacea x polaris, Betula nana, from a flora immigrated from the west close on Dryas octopetala and, probably, Salix glauca, and the receding inland ice. According to present pollen or spores (ibid. p. 38) of, i.a., Lycopodium day ideas, they prove those plants to have lived Selago, L. annotinum, L. clavatum, Selaginella in the Norwegian ice-age refugia. selaginoides, Polygonurn viviparum, Thalictrum Still more conviction carry the fossils found alpinum, and Filipendula Ulmaria. The findings by SMITH (1920 pp. 130 ff.) in late-glacial depo of interglacial fossils at Bollnas, Harno, and sits of south-western Jamtland and north-wes Langsele (cf. ERIKSSON 1912, HALDEN 1915, tern Harjedalen, likewise west of the ice-divide. MUNTHE 1910 and 1946, SUNDIUS & SANDEGREN On the basis of those findings, SMITH classifies 1948) mostly consist of southern species irrele the following species as indubitable glacial sur vant to the problems discussed here; only a mi vivors: nor part of the plants, several of which could Asplenium viride Saxifraga oppositifolia not be determined with certainty, fall under the Scirpus caespitosus Dryas octopetala category of probable or possible survivors (Car ex Salix reticulata Empetrum nigrum coil. cfr microglochin, C. cfr norvegica, Rumex Aceta S. herbacea Primula stricta S. polaris sella coli. , Caltha palustris, Ranunculus repens, Rubus idaeus, Potentilla palustris, Viola cfr bi With the utmost probability, this is true: flora, Vaccinium cfr ....Myrtillus, V. uliginosum, also of the following plants (SMITH 1. c.): .58 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Selaginella selaginoides Viola spc. go eastwards, i.e. to the alpine floras of Central Equisetum spc. Loiseleuria procumbens Asia, and not northwards, to the mountain flora .Juniperus communis Andromeda Polijolia of Fennoscandia. In other words, he refers the Oarex Lachenalii Arctostaphylos alpina 0. cfr rufina Vaccinium Vitis-idaea Fennoscandian alpine flora to the Arctic branch 0. cfr Bigelowii V. uliginosurn of the total area of the Arctic-montane biota, 0. magellanica V. M yrtillus and the flora of the Alps to the Eur�siatic-mon .Salix arbuscula V. Oxycoccos coli. tane branch; according to him, these branches Betula nana Diapensia lapponica have not met west of Central Asia, at least not Thalictrum alpinum since the Great interglacial. Unfortunately, the finding by MUNTHE (1946 This may perhaps be an exaggeration. It -pp. 12 ff.) of fossil Pedicularis flammea at Porsi would seem conceivable that the Alps have re .in eastern Lule Lappmark does not tell us very ceived part of their biota from Scandinavia at much, since the fossils do not date further back a later time, possibly during the Riss glaciation than to the Ancylus time and, moreover, were (ef. KALELA 1944 pp. 145 ff. and the literature -probably not found in an unmoved position. cited there). However, there is no reason to The findings by H. LINDBERG (1911) at Ivalojoki dwell on this question here. in Lapponia Inarensis that, besides, cannot be As to a migration of plants in an opposite di said to prove much, since the fossils probably rection , all Scandinavian botanists of to-day ·originate from a flora immigrated post-glacially would seem to agree in regarding it as improb from the east (cf. NANNFELDT 1935 p. 83), able or insignificant (cf. Du RIETZ 1935 p. 227, as well as those by HEINTZE (1908) and TH. FRIES HOLMBOE 1936 p. 24, B.JORKMAN 1939 p. 220 and, (1911) on Arpojaure in Torne Lappmark, do not above all, NANNFELDT 1935 pp. 76 ff. and KA comprise any species not found by SMITH. How LELA op.c. pp. 120 ff.). It is unnecessary to re ever, as regards Betula nana, which was also peat the well-known arguments against such a :met with in the deposits of Brunflo and Pilgrim migration, all the more so since the present au sta.d, these findings increase the probability of thor shares this opinion, though partly for other its being a glacial survivor into a virtual cer reasons than those usually adduced. tainty. This also holds good of Loiseleuria pro There is, first of all, the conception that the .cumbens, which TENGWALL (1925 p. 760) found ground laid bare at the retreat of the inland ice on Andeya (Vesteraalen) in deposits that, in all was immediately occupied by pine forests im likelihood, were late-glacial. passable to alpine plants. However, this would The relatively numerous fossil finds in the coas seem largely to be a misconception, caused by tal region of Serlandet and Vestlandet in Nor an erroneous interpretation of the pollen profiles way are beyond my competence to judge; most current among all scientists prior to the inves ·of them can, apparently, not be dated with such a tigations of AARIO (1940). By means of recent ·degree of certainty as to render them altogether pollen spectra from the tundras of the· Petsamo unquestionable as evidence. district, AARIO proved that the accumulations of Pinus pollen in late-glacial deposits, anterior to the )>great subarctic Betula maximum)> of SAu 'The Fennoscandian flora and the flora of RAMo, were due to long-distance flight on open the Alps tundras and, consequently, indicate cool periods, The most important recent contribution to the instead of the reverse. The most significant history of arctic and boreal biota during the South-Swedish pollen profiles interpreted in ac ·Tertiary and Quaternary is, no doubt, HULTEN's (�ordance with the theory of AARIO are doubtless theory of equiformal progressive areas (HULTEN those from the region of the river Viskan, drawn 1937a). A consequence of his general views is that up by VON PosT. Unfortunately, they have been the historical connexions of the flora of the Alps published but in a most summary form (vo� Origins of the Flora 59
PosT 1947 pp. 206 ff. ); for our present purpose, subalpine plants should have immigrated into they are, however, sufficiently detailed. the mountains from the South (cf. M. FRIES 1949 VON PosT summarizes the course of events as pp. 54 ff. ). Nor can we be sure, that a thin fringe follows (op.c. p. 210): of alpine plants did not extend along the border The most ancient layers bear evidence of a of the receding inland ice. As suggested in a period of birch forests, occurring during Goti preceding chapter, the greatest difficulties of al glacial time and, in all probability, correspon pine plants below the conifer limit are hardly ding to the .Allerod phase in Denmark and ad due to the climate, but to the leached raw humus jacent parts. At the end of the Goti-glacial time, soils and the competition of sylvan plant spe a sharp reversion to open tundras followed, re cies. Immediately below receding glaciers in gistered by a Pinus maximum and correspon south-western Norway, even high-alpine plants, ding to the >>later Dryas time>> in Denmark &c. e.g., Deschampsia alpina, Poa laxa ssp. flexuosa, The succeeding development towards a tempe Ranttnculus glacialis, Saxifraga rivularis, &c., rate climate was interrupted by another, though occur in the conifer belt almost down to the sea less marked, reversion, occurring a good way in level and in the company or close vicinity of so to fini-glacial time. The tundra epoch then came thermophilous species as, e.g., Ribes Uva-crispa, to a definite end with a second Betula maximum Geranium Robertianum, Hypericum maculatum, (the >>great subarctic Betula maximum>>), which Digitalis purpurea, Knautia arvensis, &e., al was betokened by a culmination of Salix pollen. though the high-alpine element disappears as Almost simultaneously with this Betula maxi soon as the plant cover becomes closed (FAEGRI mum, Corylus appeared, whereas the pine did 1933). A similar occurrence of alpine plants be not become predominant until later. The sub neath the inland ice is conceivable, of course, sequent development is irrelevant in this context. provided the plants stood at the ice border when A synchronous development on exactly the the melting of the ice began. same lines can be traced at the ancient river The chances that such an ephemeral vegeta >>Svea alv>> (situated in Mid-Sweden), and in tion should lea�e traces in the form of macrofos .Angermanland (North-Sweden ), where VON PosT sils or wefts in the pollen spectra are certainly succeeded in synchronizing the Viskan dia very small. Yet from the investigations of ERDT grams with geo-chronologically dated local dia MAN (1943 pp. 57 ff.) we know that this actually grams. Thus >>the great subarctic Betula maxi happened. In the bottom layers of a bog at mum>> could be dated to the years -100 to -200 Langvattnet (near Stensele in southern Lapland) in the time-scale of DE GEER, i.e. to about 7000 410 m s. m., ERDTMAN found pollen of Hippo B. C. (v. PosT op.c. p. 211). In other words, phae rhamnoides and an Artemisia species. These at the time of the bipartition of the inland ice two plants, now completely absent from Swedish in Jamtland, which marked the transition from Lapland, must have been among the pioneers fini-glacial to postglacial time, Sweden was prob nomadizing on the bare soil hard by the retrea ably still covered mainly with subarctic birch ting ice, since they occur but in open plant com forests. (Cf. also the finding, by HALLE, of a munities; the only localities of Fennoscandia postglacial ·tufaceous limestone at Raftkalen, in where Hippophae and an Artemisia species (A. the woodland of Jamtland east of the ice-divide, bottnica) occur together are the shores of the Gulf the oldest stratum of which contained Hippo of Bothnia, �here land-elevation continually cre phae [abundantly], Dryas octopetala, Betula ado ates >>new soil». Nor could the pollen, especially _ rata coli. and Salix cfr nigricans, but no pine in the case of a herb like Artemisia, have been [HALLE 1915 pp. 20 f.]. The absence of the pine, brought to Langvattnet by long-distance flight. which occurred in younger strata, may, as a Evidently, there is a possibility, however slight, matter of course, be accidental.) that low-alpine species still living in Lapland It is, thus, quite possible that, at any rate, may have reached the mountains in the same 60 SELANDER: Floristic Phytogeography of South-TVestern Lule Lappmark way (cf. ALMQUIST 1948 pp. 82 ff.). High-alpine belt or the upper conifer belt of the Fennoscan species were excluded, i.a., because they could dian mountains, viz., Potamogeton alpinus, P. not have lived on the tundras of Central Europe praelongus, Carex rostrata, Betula pubescens, B. (cf. below p. 61) and, likewise, basophytes nana, Dry as octopetala, Myriophyllum spicatum, which could not find suitable habitats in the M. alterniflorurn, and Arctostaphylos Uva-ursi Mid-Swedish region of acidic primary rocks. (NILSSON op.c. p. 4 79). The answer to the question whether or not The evidence of the bogs of Oeland, as accoun the Fennoscandian alpine flora was partly re ted for by LUNDQVIST (1928) and ERDTM:AN cruited from the South thus largely depends (1946), leaves a similar impression, although the on what species were present immediately south material is too slender to admit of definite con of the ice. In this connexion, the conditions in clusions (LUNDQVIST op.c. p. 144). We find in Central Europe, during the far largest part of dications in the late-glacial deposits of light Be the Wiirm glaciation, are evidently not relevant, tula forests with a lot of Hippophae (up to 7 %, except indirectly. What interests us is, above ibid. pp. 28 ff. , p. 36, and pp. 48 ff. ); in addi all, conditions in South-Sweden during the Goti tion, there is pollen of, i.a., Artemisia tup to 6 % glacial time, when the plants arrived there which of NAP, ERDTMAN op.c. p. 298), Helianthemum were to occupy Mid-Sweden during the fini-gla (ibid. p. 300) and Oxytropis campestris (ibid.). cial period. According to HEMMENDORFF (1897), the macro As mentioned above, the Goti-glacial clima fossils include, e.g., Dryas octopetala, Salix reti tic amelioration, succeeding the earlier Scano culata, and S. polaris (determination possibly Danian Dryas time, and registered, i.a., by .the wrong, perhaps to be corrected to S. herba standard Aller0d section, resulted in forests, cea, Du RIETZ in NANNFELDT 1935 p. 76, Du mainly light woods of birch, willows and aspen, RIETZ 1935 p. 228); the macrofossils may, how which covered most of Scano-Dania; in the far ever, partly originate from earlier Dryas time. thest north-west, i.e. in northern Jutland, birch Especially interesting are the late-glacial de groves were to be found only in warm, shel posits of )>Morka on Mt Omberg, since they tered localities (cf., e.g., IVERSEN 1947 p. 76). Hah were laid down on the Swedish mainland close Tundra plants, unable to endure such conditions, to the ice border. pollen was evidently could not survive; nor could the later Helianthemum found also in this place: the Helianthemum spe Dryas time occasion a reinforcement of fresh cies belongs to the italicum group tundra species from the Alps, the whole of Cen (ERDTMAN pp. f.; cf. the occurrence of a Helianthe tral Europe being covered by forests. 1949 52 mum species, possibly H. alpestre (Jacq. ) DC., The pollen spectra dating from Aller0d time in the Pinega district of subarctic Russia, and later Dryas time, i.e. belonging to the zones POHLE p. p. The Omberg XI and X in the system of VON PosT, may give 1903 9, STERNER 1936 430). deposits are, further, characterized by abundant us an idea of the South-Swedish flora of those pollen, the major part of which is far-off times, provided we discount all the pol Artemisia of the vulgaris-type; yet some grains perfectly len that probably is secondary (cf. IVERSEN 1936) match those of A. rupestris op.c. pp. or, like the pine pollen , was transported by long (ERDTMAN ff.). Other interesting species present there distance flight. NILSSON (1935 pp. 480 ff.) char 49 are cf. acterizes the vegetation of Scania during those Astragalus alpinus, Filipendula, Ulmaria, cf. and cf. periods as a parallel to )>der subarktischen Bir Petasites frigidus, Saxifraga oppositi (ibid. pp. 51 ff. ). kenregiow>, divergent above all through the occur folia rence of Hippophae and, probably, of sporadic To judge from the incomplete data published Corylus. The macrofossils found, without doubt, by voN PosT ( op.c. ), the vegetation on the West in the zones XI and X of Scanian bogs consist Coast also had the same general character. Pol of plants, all at present occurring in the birch len of Hippophae and Artemisia is reported also Origins of the Flora 61 from these parts (cf. ERDTMAN 1944 p. 415). count, when discussing the provenance of the The postglacial history of Hippophae in Fenno Fennoscandian alpine flora. scandia, with a survey of all findings of fossil In the case of the subalpine plants, the posi remains, is exhaustively dealt with by SANDE tion is somewhat different. The possibility of GREN (1943), and that of Helianthemum in their having immigrated exclusively from the Denmark, by IVERSEN (1945). South can hardly be altogether left out, at least · To a botanist, the palynological evidence of in regard to some species. Many such plants the South-Swedish late-glacial deposits seems to thrive at present just as well below the conifer indicate conditions, rather like those that at pre limit as above it tcf. e.g. M. FRIES 1949 p. 61), sent prevail in the birch belt of the low moun provided the soil of the habitats has a mild mull tains (ldgfjallen), where low-alpine plants occur profile. Now such profiles were probably more ·chiefly on local, edaphically conditioned patches common, at least near the mountain range, in -of alpine heath. In addition, the flora evidently the present podsol soils prior to the great cli ·embraced a southern, continental steppe element, matic change at about 1000-500 B. C. and the perhaps owing to the stronger insolation in lower coincident immigration of the spruce; and luxu latitudes together with the continentalizing in riant herb societies, of the kind where our fluence of the inland ice south-east of the ice plants chiefly belong, were seemingly quite fre border. The alpine species were hardly numerous, quent even outside the mountains (ERDTMAN and the occurrence of truly high-alpine plants 1943 pp. 57 ff.). This would evidently facilitate would seem to be out of the question, also for the migration of subalpine species. the reason that the glacial Mid-European tundra Of course, a southern provenance is concei was, during the Wiirm glaciation, hardly ever vable especially in the case of plants whose total high-arctic in character, possibly even admitting distribution is restricted to Europe. Such a plant of the occurrence of local subarctic woods in is Myosotis silvatica ssp. frigida. It is of indubi the region between the northern inland ice and table South-European origin, all other races of the Alps (FIRBAS 1939 pp. 84 f., 87; cf. also M. silvatica but the Oentral-European ssp. eu BROCKMANN -JEROSCH 1907 pp. 396 ff. ). silvatica being Mediterranean (cf. VESTERGREN Even though Dryas and the dwarf Salices have 1938 pp. 10 ff.). Outside Scandinavia, it does not ;Small, coriaceous leaves which are easily fossilized, occur anywhere but in the South-European moun so that these plants probably are over-represen tains (VESTERGREN op.cit. pp. 7 f.). And it would ted among the macrofossils, the findings, never have been quite able to live in birch forests such theless, indicate that they were quite frequent. as the late-glacial Swedish ones. Other instances ·Their frequency may be explained by the bet are Lactuca alpina and (less probably with a view, ter soil conditions prevailing during that epoch, i.a., to its isolated area in the Samoyede Land) when leaching and podsolization had been active Saussurea alpina. This does not imply any de -but for a comparatively short time and no raw nial of the possibility, or even the probability, humus formation worth mentioning had taken of their being glacial survivors in Fennoscandia; place. Otherwise, the comparatively flat ter M. FRIES, who has made a close study of Lactu -rain would not seem to have admitted of the ca alpina, emphatically declares it to be a hiber occurrence of ecologically specialized alpine nator (1949 p. 56). _plants. Moreover, the fact that an arctic-alpine plant
However, all truly alpine species doubtless be is found either in a fossil sta-te in late-glacial, longing to the late-glacial flora of South-Swe South-Scandinavian deposits, or growing in relic den were established as glacial survivors by the stations of southern Sweden, does not of neces fossil findings of SMITH (cf. above pp. 57 f.). Also sity imply that these South-Scandinavian, ex for this reason, we need not take the South tinct or recent, populations originate from the .Swedish fossil findings of alpine plants into ac- Alps. 62 SELANDER: Floristic Phytogeography of So1-tth- Western Lule Lappmark
The contemporary stations on the Swedish mary theories; at the very best, we can hope to West Coast of Cryptogramma crispa, Poa alpina, catch a dim reflex of the truth. Visca,ria alpina, Sedum Rosea and Alchemilla Still, in exceptional cases, we can attain to alpina may be (and probably are) just outposts a reasonable certainty. There is, e.g., one de from their South-Norwegian areas (cf., e.g., cisive argument against a comprehensive post NANNFELDT 1935 p. 78, ERLANDSSON 1942a pp. glacial emigration of alpine plants from the Alps 127 ff., ST�RNER in H. FRIES 1945 pp. 70 ff. ). The to Fennoscandia, viz., the fact that these two alpine relics of south-eastern Sweden, e.g., Asple regions have very few microspecies within cri nium viride, Poa alpina, Viscaria alpina, Bartsia tical genera in common, at least not in identical alpina, and Pinguicula alpina, may, as tenta� forms, exceptions being only, e.g., Myosotis silva tively suggested by ERLANDSSON (op.c. pp. 129 tica ssp. frigida, dealt with above, and Alche ff.), originate from northern Finland, from where milla glomerulans, which is obviously of a sub they would have migrated by way of Karelia arctic provenanee (cf. SAMUELSSON 1943 pp. 85 and Balticum, or more probably, as suggested f.). This argument beeomes all the more con by STERNER (1944 p. 222) in the case of Viscaria vincing in view of the great number of micro alpina v. oelandica, from �omewhere in Central species and minor races that Fennoseandia has Europe east of the ice border. Here they may in common with Iceland, Greenland, and north have associated with some of the endemics of eastern North America (cf. below p. 72). Oeland, e.g., Helianthemum oelandicum and Ga If a eonnexion between the biota of the Alps lium pumilum ssp. oelandicum, and the Siberian and that of Fennoseandia had existed during, steppe elem.ent in the flora of Oeland and Got or after, the 'Viirm glaeiation, at least a few of land, viz., Carex obtusata, Potentilla j1·uticosa, Ar the many rather young endemies of the flora of temisia rupestris, and A. laciniata (STERNER 1922 the .Alps should be likely to have found their p. 329, 1944 pp. 221 ff.), which apparently be way into the mountains of Seandinavia. As eve longed to the pioneer Rpecies of the late-glacial rybody knows, this has not happened. In all tundra of South-Sweden (ERDTMAN 1946). likelihood, praetically every species oeeurring in This hypothesis may, to a certain degree, find both regions is an aneient unit. Here is a chanee support in a strange fact, pointed out by GAMS to determine the latest time when sueh a eon (1933 p. 4 70). The two leading species among nexion could have been in existence. GAMS (op.c. the exotic fossils of late-glacial, South-Scandi pp. 475 ff.) considers most Alpine endemies to navian deposits, viz., Dryas octopetala and Hip have arisen during the 1\Iid-Pleistocene, i.e., pophae rhamnoides, do not seem to have ap above all, during the Great interglaeial; aecor peared in the Alps before the Wiirn1 glaciation. ding to him, >>die seltenen nordischen Arten>> in At least, they, and most species of a Siberian the flora of the Alps also immigrated during >>den provenance, have not been found there in depos mittleren Eiszeiten>>. He does not class but a its anterior to that time, although Dryas occurs few races among the neo-endemies, e.g., De in 1\'lid-Pleistocene deposits of the Carpathians. schampsia rhenana Gremli, Saxifraga oppositi Considering how well the leaves of these two spe folia ssp. amphibia Siinderm., Myosotis caespi cies keep in a subfossil state, this fact may in titia Kern., and the seasonal dimorphic.races with dicate that they actually immigrated into the in Gentianella, Rhinanthus, &c., many of whieh Alps and into southern Scandinavia from the have eounterparts among the endemics, or near East at about the same time. endemies, of the Fennoseandian flora. This faet All this may seem more than a little tenta doubtless strengthens the probability of his da ti ve and vague. But like everything else in N a tings, postglacial endemism thus obtaining about tu re, the hibernations and wanderings of the al the same extent in the Alps and around the Bal pine plants were, of course, far too varied and tic.. As G.A.Ms, moreover, is very eautious in his eomplicated to be entirely covered by our sum- estimations of the age of raees and events, we Origins of the Flora 63· may safely suppose the vast majority of Alpine nunculus peltatus var. septentrionalis Lindb. fiL endemics to date back at least to the Riss gla may also be endemic in North-Fennoscandia and ciation, a fact which would seem to indicate that the adjacent parts of Russia (cf. HULTEN 1949a. after that time, at any rate, no alpine plants pp. 399 ff. and p. 390). However, as they do not emigrated to Fennoscandia from the Alps. ascend above the conifer belt, at any rate within In the following, I shall therefore disregard our region, they are, apparently, of a post the possibility of true alpine plants having rea glacial origin. ched Seandinavia from the South after the Riss With the exception of a number of Hieracia. glaciation, nor shall I deal in a different man and Taraxaca, the remaining endemic element ner with species appearing, and those missing, in the flora of our region seems probably to in the Alps. consist of epibiotics. It .is very small in number. Only Picea Abies var. arctica (LINDQUIST 1948, p. 308), Poa herjedalica 1920 p. 160), Roeg Endemics (SMITH neria scandica (LovE & LovE 1948 p. 105), Alnus Among definite glacial survivors are, first, the incana var. virescens (HuL'rEN 1949 a p. 389),. endemic species and races unless, N.B., they have Ribes spicatum ssp. lapponicum (HYLANDER 1945. arisen after the ice-age. This is probably the p. 196), Prunus Padus var. borealis (HULTEN case with Antennaria lapponica (cf. Part II, pp. 1949a pp. 390 f.), Sorbus aucuparia var. glabrata 16 f.), which seems to be a neo-endemic arisen (ibid. p. 391), Anthyllis Vulneraria var. lappo in postglacial time out of the hybrid A. alpina x nica (HYLANDER op.c. p. 226), Primula scandi dioeca. As regards Calamagrostis purpurea (see navica (BRUUN 1938 pp. 249 ff. ), Euphrasia lap NYGREN 1946 p. 348, cf. HUIJTEN 1927 p. 103, ponica (TH. FRIES 1921 p. 12), and Arnica alpina and 1942 p. 163), NYGREN (op.c. p. 207) S. str. (LAGERBERG & HOLMBOE 1940 p. 192, believes it probably to be polytopic, having M.AGUIRE 1943 ace. HULTEN 1949a) can, with arisen partly from C. oanescens through chromo any degree of certainty, be referred to this some doubling, partly through hybridization category. between C. canescens and 0. arundinacea, epi However, the same is probably true of a few geios, and neglecta. Furthermore, he thinks it more or less critical races, the distribution of >>probable that certain 0. purpurea forms are which is insufficiently known, but of which there glacial overwinterers ...while others arose after is at present, apparently, no mention outside the late glacial period>> (op.c. p. 211). For the rest Fennoscandia (and the adjacent parts of nor nobody has ascribed such a derivation to any thern Russia), viz., the following: Agrostis sto of the types concerned except certain Hieracia lonifera var. arctica (cf. Part II I>p. 8 f.), Carex and Taraxaca, not dealt with here, and possibly adelostoma (stray stations eastwards to the Urals, one or two agamospecies within the genus of C.AJ.ANDER 1935 pp. 99 f., HULTEN 1949b p. 98), Alchemilla. SAMUELSSON (1943 p. 27) thinks it Arabis hirsuta var. subalpestris (cf. Part II p. 13), most improbable that any of our Alchemillae Alchemilla borealis (SAMUELSSON op.c. p. 14), A. >mach der letzten Vereisung entstanden sei>>; but, norveg·ica (ibid.), and Erigeron politum (SIMMONS he continues, >>ich lasse. es vorH1ufig dahingestellt, 1907a p. 35, LAGERBERG 1940 p. 81). ob eine oder die andere der hier iibergangenen Arten der Acutidens-Gruppe>> (e.g., A. borealis The amphi-Atlantic element and A. norvegica) >>jiingeren Datums sein konnte>>. He does not, however, give any reason for his The >>west-arctic>> species, i.e. American species suggestion; and the chances· are, doubtless, that with isolated areas in Fennoscandia and, in a these species are of the same age as our remai few cases, northern Russia andjor Nova Zembla, ning Alchemillae, i.e. probably Old-Quaternary. can also be considered to have survived one or Pedicularis palustris ssp. borealis and Ra- more glaciations here. The theory of glacial sur- 64 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
vival was propounded, i.a., in order to explain Minnesota, Michigan, and New Yersey, N.W. Cali their peculiar distribution (cf., e.g., BLYTT 1893, fornia, Iceland, Fennoscandia, N.W. Russia, reported from single station in W. Siberia (HULTEN op.c. p. 372, SERNANDER 1896, HANSEN 1904, FRIES 1913, LOVE & LOVE 1948 p. Ill); NORDHAGEN 1933, 1935, 1937, NANNFELDT 1940), Juncus balticus, the >> main form>>: California to and it is hardly necessary to repeat here the ar Alaska, Iceland, Scotland, N. England, N. Germany, guments of these well-known works. It should Balticum, Fennoscandia (HULTEN 1943 p. 421), the · be pointed out, however, that the survival of whole European area within the boundaries of the Riss glaciation; these species remains unassailable whether we Urtica dioeca ssp. g·racilis : Alaska, S. Hudson Bay, explain their present distribution as a result of and Newfoundland to Oregon, New Mexico, Great a reduction of earlier circumpolar areas or by Lakes, N. New York, and Connecticut, Fennoscandia a migration over Transatlantic land bridges that (HULTEN 1944 p. 592, SELANDER 1947 p. 266); Cerastium arcticum : easternmost Arctic Asia, Alaska have ceased to exist. This is true also of the · and probably eastwards over the Arctic Archipelago amphi-Atlantic species with isolated areas on to N.W. Greenland, Iceland, Faroes, Spitzbergen, either side of the Atlantic, which cannot be Fennoscandia (HULTEN op.c. p. 663, LovE & LoVE referred to the >>west-arctic>> group because of op.c. p. 49, AsPLUND 1918 pp. 27 f.); their European areas being as large as, or larger Cerastium fontanum ssp. scandicum : Greenland, than, their American areas. The following plants Iceland, Faroes, Fennoscandia, Archangelsk (GART of our region representing these types of distri NER 1939 p. 69); 8agina caespitosa: area difficult to establish, owing bution do not, on the European mainland, occur to confusion with 8. intermedia; ace. to NoRDHAGEN east of the boundaries of the Wiirm glaciation: (1935 p. 55) Colorado to Alaska and Labrador, W. Deschampsia atropurpurea s. str ., excl. ssp. para Greenland, Iceland, Scandinavia; however, not in mushirensis (Kudo) and ssp. magellanica (Hook): Alaska (HULTEN op.c. p. 671) nor elsewhere W. of N. Labrador and N.W. Newfoundland to New Hamp Hudson bay (HULTEN 1949b); occurs, on the other shire, S.W. Greenland, Fennoscandia, W. Dvina hand, in the Arctic Archipelago, though not W. of Petchora distr. in Russia (HULTEN 1942 p. 182); also Baffin Land and Melville Penins. (SIMMONS 1913 in E. Greenland, though absent from S. Greenland p. 79, sub. 8. Pumilio [R. Br.] Simm.), and on Jan (BOCHER 1938 p. 220); Mayen and in S.E. Greenland (BocHER op.c. p. 81); Festuca vivipara : Hudson Bay to Newfoundland, Arenaria humifusa : Arctic and subarctic N. Ame Greenland, Iceland, Faroes, Scotland, Jan Mayen, rica, stray stations from W. Alaska to Ellesmereland, Spitzbergen, Fennoscandia (FERNALD 1926 p. 151, Baffin Land, N. Labrador, Gaspe and Newfoundland, POLUNIN 1940 p. 89, NANNFELDT 1940 p. 41); in Rocky Mts S. to Alberta, W. and SW. Greenland, Carex parallela : E. Greenland, Spitzbergen, Fenno Spitzbergen, N. Fennoscandia (PoLUNIN 1943, map scandia, Jugor Shar, Waigatch, Nova Zembla (SA p. 4.54; not in E. Greenland or Iceland); MUELSSON 1921 pp. 226 ff., NANNFELDT l.c.; SAMUELS· A renaria norvegica : Iceland (common), Shetland soN's report from W. Siberia seems to be erroneous); Is. (one station), Scotland (one station), Fennoscandia Carex nardina : Arakamtchatch I., Alaska, British (NORDHAGEN op.c. pp. 41 ff.); Columbia to Colorado, N.W. part of Hudson Bay, Draba norvegica: N.E. North America, Greenland, Ellesmereland to N. Labrador, Gaspe, W. and N.E. Iceland, Faroes, Scotland, Spitzbergen, Fennoscandia, Greenland, Iceland, Scandinavia (HULTEN op.c. pp. Nova Zembla (NANNFELDT op.c. p. 41); 299 f.); Draba crassijolia : Rocky Mts from Colorado and Carex arctogena : Tierra del Fuego, Patagonia, New Utah to British Columbia, Alaska, Richardson I., Hampshire, Labrador, W. and E. Greenland, Fenno Keewatin, Lake Agnes, S.E. Baffin Land to N. La scandia (SMITH 1940 pp. 196 ff.); brador, W. Greenland, Scandinavia (HULTEN 1945 a Carex macloviana s. lat.: Falkland Is., Patagonia to pp. 850 f.); Bolivia and Equador, Mexico, Hawaii, N. Kuriles·, Braya linearis s. str.: E. Greenland, Scandinavia Kamtchatka, Alaska and Mackenzie to California, (ScHuLz 1924 pp. 229 f.). If taken in a wider sense
Idaho and Wyoming, Labrador, Gaspe, Greenland, ( = Br. glabella Richards.), the species still belongs Iceland, Fennoscandia (HULTEN op. c. p. 319); to this category. (area Mackenzie distr., W. and E. Carex rujina : Greenland, Iceland, Scandinavia Greenland, Scandinavia, probably two stations on the (BOCHER op.c. p. 233, map p. 234); lower Lena R., Huvr.EN op.c. p. 887); Carex livida : Corea, Yezo to Kamtchatka, Alaska, Potentilla Chamissonis : Labrador, Greenland, Spitz Labrador and Newfoundland to Washington, Idaho, bergen, Fennoscandia (HULTEN 1945 b, map p. 144); Origins of the Flora 65
Alchemilla Wichurae : S.E. Greenland, Iceland, Spitzbergen, Fennoscandia and Nova Zembla (Boo Faroes, Scotland, N. England, Fennoscandia (SA HER op.c. p. 209) plus a few, rather dubious stations MUELSSON 1943 p. 16, map p. 17); in Siberia and arctic America (HuLTEN 1949b). Epilobium lactiflorum : Alaska, S. Dakota, Great Lakes and Labrador to California, Utah, Colorado, Somewhat doubtful is likewise, though for and New Hampshire, Greenland, Iceland, Fenno- · other reasons, the status of the amphi-Atlantic scandia (HuLTEN 1947 a p. 1151); species also occurring in Russia, viz., the follow Rhododendron lapponicum : from Jana R., Vilju R., and upper Yenisei R. to Chukch Peninsula, Ochotsk, ing: Saghalin, and Penshina; Alaska, Yukon, Coronation Stellaria calycantha : Ochotsk distr. and S. E. Gulf, and Ellesmereland to British Columbia, Church Kamtchatka to Saghalin and Kuriles, Alaska, Great ill, and Gaspe, Maine, New York; Greenland; Scandi Slave L., and N. Labrador to California, N. Dakota, navia (HULTEN 1948 p. 1225); and W. Massachusetts, Greenland, Iceland, Fenno Gentianella aurea: Greenland, Iceland, Fennoscandia scandia, N. Russia (HULTEN 1944 p. 649); to the White Sea (BocHER 1938 pp. 165 f.; the reports Epilobium Hornemanni : area insufficiently known, cited there from Central Asia and Arctic America, reliable reports from Ochotsk distr., Kamtchatka, incl. Alaska, are erroneous, owing to confusion with N. North America (W. area from Alaska to Minne related species); sota, southwards to California, Utah and Colorado, Euphrasia frigida : Newfoundland to Labrador, E. area from Labrador to New Hampshire), S. half Greenland, Iceland, Faroes, Fennoscandia (PuGSLEY of Greenland, Iceland, Fennoscandia, Kanin Penins., 1933 p. 308); Urals (HULTEN 194 7 a p. 1150); Rhinanthus minor ssp. groenlandicus : Newfound Cornus suecica : Chukch Penins. to S. Ussuri and land, St. Lawrence reg., S. E. Labrador, S. Greenland, N. Yezo, Alaska, Vancouver, Labrador to Gulf of Iceland, Faroes, Fennoscandia (LovE 1950, map p. St Lawrence, Magdalen Is. and Newfoundland, · s. 51); Greenland, Iceland, Faroes, Shetland, Scotland, N. Pedicularis flammea : Alaska, Canadian Rocky Mts., England, N. Germany, Denmark, Fennoscandia, Labrador, v\T. and E. Greenland, Iceland, Scandi Esthonia, N. Russia (HuLTEN op.c. p. 1188), the navia (NORDHAGEN op.c. p. 55); whole European area within the boundaries of the Campanula unijl01·a : Anadyr, Chukch Penins., Riss glaciation (map in TROLL 1925 p. 317); Alaska and Labrador to Colorado, Utah, Gaspe Pen Cassiope hypnoides : Maine to Baffin Land, W. ins. and W. Newfoundland, Arc.tic Archipelago, W. and E. Greenland, Iceland, Beeren I., Spitzbergen, and E. Greenland, Iceland, Spitzbergen, Scandinavia, Franz J oseph Land, Fennoscandia, rare along the Waigatch, Nova Zembla (HULTEN 1949c p. 1464). Russian coast to N. Urals and Nova Zembla (BocHER Erigeron boreale: Gaspe, Newfoundland, S. Labra op.c. p. 154, map in TOLMATCHEV 1930 p. 194); dor, Greenland, Iceland, Scotland, Scandinavia, E. Diapensia lapponica sens. str., excl. ssp. obovata (Fr. to Varanger, Kanin Penins. (BocHER op.c. p. 189, Schm.) Hult: N. E. North America, W. to W. coast HULTEN 1949 bp. 430); of Hudson Bay, S. to mts. of New England, Green Erigeron unalaschkense : Chukch Penins., Kam land, N. Iceland, Fennoscandia, Kanin Penins., Kol tchatka, arctic North America, W. and E. Greenland, gujev, Obi Bay, lower Yenissei (HuLTEN 1948 p. 1266); Iceland, Spitzbergen, Scandinavia, E. to Kautokeino Primula stricta: Alaska to Labrador, S. to Great (DAHL 1934 p. 402), Waigatch, Taimyr R. (HuLTEN Slave L. and S. Hudson Bay, E. Greenland (not W. 1930 p. 161); Greenland, cf. GELTING 1934 p. 153), Iceland, Fenno Antennaria alpina s. str.: Greenland, Iceland, scandia, N. Russia, Nova Zembla, probably not in Fennoscandia, possibly Kanin Penins. and Kolguj ev Asia (HULTEN op.c. p. 1275); (MALTE 1934 a p. 102, BocHER op.c. p. 186, HULTEN Taraxacum croceum coll. : Arctic America, Green 1949 b p. 433; reports as to the relevant Antennarice land, J an Mayen, Iceland, Spitzbergen, Fennoscandia of arctic Russia contradictory); (BocHER op.c. p. 198), arctic Russia (HULTEN 1949b Taraxacum rhod_olepis : E. Greenland, Fennoscandia p. 462). (BOCHER op.c. p. 198; oral confirmation by Dr. G. HAGLUND); These species may, possibly, have stayed east Taraxacum brachyceras Dahlst.: Greenland, Spitz of the ice-border during the Wurm glaciation, bergen, Fennoscandia (oral information by Dr G. which did not extend beyond the Kanin Penin HAGLUND). sula, migrating from there into Fennoscandia in Deschampsia alpina may also belong here, its postglacial time. All of them are, however� oce range comprising N. Labrador (ABBE 1936 p. 140), Greenland, Iceland, the Faroes, Scotland , Beeren I., anic rather than continental. During the glacia-
6- 496149 Sten Selander I 66 SELANDER: Flor.istic Phytogeography of South-Western Lule Lappmark tions. the continental type of the climate of .Arc Beeren I., south point of Spitzbergen, Faroes, British tic Russia became more pronounced �cf., e.g. , Is., Fennoscandia, Pyrenees, mts of Central 'Europe and Central Asia (B6CHER op.c. p. 66); HYYPPA 1936), which scarcely makes these parts Cemstium t-rigynum : American Atlantic coast from a suitable ice-age refugium for oceanic plants. 62° to 49°, Greenland, Iceland, Faroes, · Scotland, The chances are therefore that they too belong Fennoscandia, mts of S. and Central Europe, Grand to the >>hibernating�> category of Scandinavian Atlas in Morocco, large Central-Asiatic area (ibid. plants. p. 77); .Attached to the amphi-.Atlantic group is an Cemstium alpinum : conflicting reports of the distribution, owing to the lack of a· sorely needed >>amphi-.Atlantic-.Alpine>> one. These plants occur revision of this critical group; however, ace. to GRON in Greenland (and, in some cases, North .Ame- . TVED (1942 p. 219) the species seems to occur at rica), Iceland, Fennoscandia, and the mountains least in the N.E. of North America, Greenland, Ice of Central Europe (and, in some cases, Central land, N. England, mts of S. and Central Europe, .Asia), but are lacking -in all Russia, or nearly Fennoscandia; cf. also HULTEN 1949b; Viscaria alpina : Labrador to ' White Mts, Green so, and northern Siberia. Having eliminated land, Iceland, Scotland, N. England, Pyrenees, Appen the southern migratory route into Fennoscandia, nines, Alps, Fennoscandia, Caucasus, mts. of Central we may suppose this group to be glacial survi Asia (LID & ZACHAU 1928, map p. 91; not in the vors in our peninsula almost as safely as the Faroes and Nova Zembla, BocHER op.c. p. 86); >>west-arctic>> one. In the flora of our region, the Ranunculus glacialis : E. Greenland, Iceland, Jan Mayen, Faroes, Spitzbergen, Fennoscandia, Spain, following amphi-.Atlantic-.Alpine species possess Pyrenees, Alps, Carpathians, Transsylvania (ibid., areas on the North-European mainland not ex map p. 92); tending east of the boundaries of the Wiirm gla Sedum annuum : S. Greenland, Iceland, mts of S. ciation: and Central Europe, Fennoscandia (ibid. p. 110, cf. HULTEN 1949 b p. 243); Spa-rganium angustifolium: Kamtchatka, from Saxifmga stella-ris : Labrador, Greenland, Iceland, Alaska to Labrador, S. to California and Pennsylva Faroes, British Is. (common mountain plant in Ire nia, S. Greenland, Iceland, Faroes, British Is., Hol land), Fennoscandia, mts of S. and Central Europe land, N. Germany, Fennoscandia, Livonia, Pskov, from Pyrenees to Balkan (BocHER op.c. p. 120); Nichnegorodsk, Portugal , Pyrenees, Alps (HuLTEN Saxif-raga adscendens : Rocky Mts from British 1941 p. 93), the whole of the N.-European area within · Columbia to Colorado, mts of S. Europe from Pyrenees the boundaries of the Riss glaciation; to Sicily, Balkan and Carpathians, Caucasus, Estho Ca-rex mic-ruglochin : North America, scattered nia, Scandinavia, S. Finland (LAGERBERG 1940 p. 124, stations from Alaska and British Columbia to Labra HULTEN 1945a p. 904, 1949b p. 245); reports from dor and Newfoundland, N. Baffin Land, Greenland, Newfoundland and Hudson Bay distr. not confirmed; Iceland, Scotland, Fennoscandia, NW. Russia, S. to Potentilla Crantzii : Newfoundland (FERNALD 1933 Wilna, East Prussia (single station), mts of Central p. 121), Labrador, Greenland, Iceland, Spitzbergen, Europe, Carpathians, Caucasus, scattered stations in Faroes ( ?), British Is., Fennoscandia, E. to E. Kola Central Asia (HULTEN 1942 p. 310, KuKENTHAL 1909 Penins. and L. Ladoga, Balticum, Pyrenees, Alps, p. 109, STEFFEN 1931 p. 318); Carpathians, Caucasus, Asia Minor, Armenia, Nova Ca-rex no-rvegica s. str.: S.W. Greenland, Iceland, Zembla, N. Urals, N.'\V.-most Siberia, Tian Shan Scotland, Alps, Fennoscandia, eastwards to Kanin (GRONTVED 1942 p. 276, FERNALD 1925 p. 271, BocHER Penins. (KALELA 1944 pp. 13 ff.); op.c. p. 132, HuLTEN 1949 b p. 261); Ca-rex at-rata: N. North America, Greenland, Ice Alchemilla alpina : Greenland, Iceland, Faroes, land, Scotland, Wales, Pyrenees, mts of Central British Is., Fennoscandia, Sierra Nevada, Pyrenees, Europe, Fennoscandia; in Asia the closely related Appennines, Alps, Black Forest (BocHER op.c., map C. pe-rfusca Krecz., in N. America mostly ssp. at-ro p. 122); in S.E. Europe and the Caucasus probably squamea (Mack.) Hult., the )>European type)> in Green other, related races; land, Rocky mts and Alaska (HULTEN 1942 p. 354). Gentiana nivalis : Labrador, Greenland, Iceland, Juncus alpinus var. alpest-ris : Beringia, St. Law Scotland, Fennoscandia, mts of S. and Central rence reg., Greenland, Iceland, Fennoscanclia, mts of Europe from Pyrenees to Jura and Balkan, Asia S. Europe (LINDQUIST 1940 p. 123); Minor (ibid. p. 166);
Salix he-rbacea: N.E. North America, S. to 44°, Ve-ronica f-ruticans : Greenland, Iceland, · Faroes, W. to W. shore of Hudson Bay, Greenland, Iceland, Scotland, Fennoscandia, mts of S. and Central Europe Origins of the Flora 67 from S. Spain to Black Forest, Bosnia, and Carpa scandia, E. to E. Kola Penins., mts of S. and Central thians (ibid. p. 178); Europe, Caucasus, Urals, Central Asia (BocHER op.c. Veronica alpina s. str.: Greenland, Jan Mayen, p. 193), also Kolgujev (HULTEN 1949 b p. 434); Iceland, Faroes, Scotland, Fennoscandia, Alps (ibid. Gnaphalium norvegicum : Newfoundland, Gaspe p. 176); Penins., Quebec, Baffin Land (FERNALD 1926 pp. Veronica pumila: E. Greenland, Iceland, Scotland, 116 f.), Greenland, Iceland, Scotland, Fennoscandia, Scandinavia, Pyrenees, Cevennes, Alps (ibid.); Kanin Penins., mts of S. and Central Europe, Cau Hieraci�tm alpinum coli.: Greenland, Iceland, Scot casus, isolated in Central Urals, Central Asia (BocHER land, Fennoscandia, mts of Central Europe (ibid. 1. c., HuLTEN 1. c.). p. 194). Since the Fennoscandian areas of these plants Similar ranges, though including also isolated are completely isolated, it is obvious that their areas in the Urals, characterize the following areas in the Urals cannot affect their status in plants: Fennoscandia. Then there are some species whose ranges com AthyTium alpestTe, Eurasiatic type (var. typicum): prise also Arctic Russia and, in a few cases, west Newfoundland, Iceland, Fennoscandia, Kanin Pen ernmost Siberia, viz., the following: ins., mts of S. and Central Europe, isolated in Central Urals (Ufa prov.), Caucasus, Central Asia from S. Selaginella selaginoides : from Aleutians, L. Atha Tomsk prov. to L. Baikal, Kamtchatka to Honshu; basca and Labrador, S. to Colorado, Michigan and in W. America var. americanum Butt., on Gaspe Nova Scotia, S. Greenland, Iceland, Fennoscandia, Penins. var. gaspense Fern. (HuLTEN 1941 p. 34, N. Russia to Urals (also Siberian side), mts of S. FERNALD 1928); and Central Europe, Caucasus, three isolated stations Cystopteris montana: Americ'a , several separate in Central Asia, S. Kamtchatka to Honshu (Huvr:EN areas from Alaska to Labrador, S. Greenland, Scot 1941 p. 75); land, Fennoscandia, E. to E. Kola Penins. and L. Phleum alpinum L. coil. (incl. Phl. commututum, Onega, mts of S. and Central Europe from Pyrenees excl. var. americanum Fourn.): E. North America, to Transsylvania and Caucasus, single station in N. Iceland, Scotland, N. England, mts of S. and Central Russia, Urals, in Central and E. Asia three or four Europe from Pyrenees to Caucasus, Fennoscandia, separate areas (ibid. pp. 18 f.); Arctic Russia, Urals, large Central-Asiatic area from W oodsia alpina: W. American area in Alaska and Altai to Afghanistan, Sikkim and China; in ""T . North Yukon S. to 53°, E. area from N. Labrador S. along America, Kamtchatka and probably Japan var. the coast to Newfoundland and New York, Green americanum (HuLTEN 1942 p. 138); land, Iceland, mts of England, Fennoscandia, E. Agrostis boTealis : N. America, W. area from Alaska to W. Kola Peninsula and L. Onega, mts of Central to British Columbia and Great Slave L., E. area from Europe from Pyrenees to Carpathians and S. Russia, E. Baffin Land and Labrador to W. Virginia, isolated Central Urals, few stations in S. Siberia, chiefly in in N. Carolina, Greenland, Fennoscandia and Arctic Altai, Corea (HuLTEN 1941 p. 13, 1949 b p. 13); Russia to Urals, W. Alps, E. Siberia (Jakutsk and Polystichum Lonchitis: North America from Alaska Vilju distr.), Kamtchatka to Kiushiu (ibid. p. 153); and Alberta to Labrador and Michigan (area not Juncus trijidus : N.E. North America, S. Greenland, continuous), Greenland, Iceland, Faroes, Fenno Iceland, Faroes, Scotland, Fennoscandia, Arctic Rus scandia, E. to Hibina, mts of S. and Central Europe sia, Urals, mts of S. and Central Europe, then only (to Sicily and Crete), Crimea, Caucasus, Asia Minor, one station at L. Baikal (FERNALD 1925, map 70); isolated in Central Urals, Central Asia round Altai, Luzula spicata : W. American area from S. Alaska N.W. Himalaya, Kamtchatka to Saghalin and Kuriles to New Mexico, E. area from S. Baffin Land and E. (ibid. p. 33); coast of Hudson Bay to New Hampshire, Greenland, Phyllodoce coerulea: E. America from S. Baffin Iceland, Faroes, Scotland, Wales, Fennoscandia and Land and N. Labrador to New York, S. Greenland, Arctic Russia to N. Urals, isolated in S. Urals, mts Iceland, Scotland, Fennoscandia to Kanin Penins., of S. and Central Europe (to Corse, Italy, Greece Pyrenees, Urals, large E.-Asiatic area from L. Baikal and Balkan), Caucasus, Asia Minor, Central-Asiatic and Chukch Penins. to Burej a Mts, Saghalin and mts from Altai and Sajan mts to Himalaya (HULTEN Yezo, N. Corea, W. Alaska (HuLTEN 1937 a, map p. 1943 pp. 442 f.); 95, 1948 p. 1233; not in N. Italy ace. verbal informa Silene acaulis : from Alaska to Baffin Land, S. to tion from Prof. E. Du RrETz); Washington and White Mts, Greenland, Iceland, J an Gnaphalium supinum : E. North America between Mayen, Spitzbergen, Franz Josef land, Faroes, N. Ire 44 ° and 58°, Greenland, Iceland, Faroes, Fenno- land, Scotland, Wales, Fennoscandia, Arctic Russia 68 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark to Nova Zembla and Urals, isolated in S. Urals, mts cumbens, and Bartsia alpina (cf. SAMUELSSON of S. and Central Europe from Pyrenees to Bulgaria, 1943 pp. 79 ff., p. 115). E. Asia from Chukch Penins. and Anadyr to Kamt The remaining species which, with the excep chatka and Commander Is. (HULTEN 1944 p. 696, GRONTVED 1942 p. 232); tion of Silene acaulis, possess large Central-Asiatic Ambis alpina : Gaspe Penins., Newfoundland, areas have probably survived the Wiirm glacia Labrador (FERNALD 1924, map p. 560), Greenland, tion also east of the ice border, the gap between Iceland, Spitzbergen, Scotland, Fennoscandia, Russia their western and eastern areas having been fil (few stations along Arctic Ocean), Nova Zembla, led up in postglacial times; in the case of Luzula N. Urals, Alps (BocHER op.c. p. lOO); Saxifraga aizoides : Rocky Mts, N. North America spicata and Silene acaulis, which are isolated in except Alaska, Greenland, Iceland, Spitzbergen, Brit the S. Urals, this is virtually certain. ish Is., Fennoscandia, Arctic Russia to Urals and Nova Ranges of an archaic, split-up type with , i.a., Zembla, mts of S. and Central Europe (BocHER op.c. large gaps in Siberia, that render >>hibernation>> p. 114), Karsk distr. of Siberia (Flora URSS 1942 in or near their present areas more likely than p. 186); Alchemilla glomerulans : Labrador, Greenland, Ice extensive postglacial migrations, characterize al land, Scotland, Fennoscandia, Balticum, Arctic Rus so the following species: (SAMUELSSON 1943 sia to Urals, Pyrenees, Jura, Alps Lycopodium alpinum : E.-Asiatic-W.-American pp . 14 f., map 1); area from Kuznetski Alatau, Altai, middle Lena R., Sibbaldia procumbens : Chukch Penins., Kamt Chukch Penins, and Alaska S. to N. Mongolia, Da chatka, Aleutians, W. American area from Alaska huria, N. Corea, N. Saghalin, N. Kuriles, Aleutians and W. Alberta to Califomia, isolated stations at the and British Columbia, isolated on Gaspe Penins., mouth of Mackenzie R. and Great Bear L., E. area N. Labrador over Baffin Land to Greenland, Iceland, from S. Baffin Land and Hudson Bay to White Mts, Faroes, British Is., N -European-N.W.-Asiatic area S. Greenland, Iceland, J an Mayen, Faroes, Shetland from Fennoscandia E. to lower Yenisei R., then isol Is., Scotland, N.W. England, Fennoscandia, Arctic ated at Zlatoust (S. Siberia), Mid-European area Russia to Urals and Obi Bay, Sierra Nevada, Pyre from S. Germany to Pyrenees and Apennines, isolated nees, Alps, Apennines, Carpathians, Balkan; in Central area in Caucasus, Armenia and N. Asia Minor (HULTEN ULTEN Asia and Himalaya closely related types (H 1941 p. 63, GRONTVED op.c. p. 96); 1946 1034): p. Spa1·ganium hyperboreum : Kamtchatka to Honshu, Viola montana : (BbcHER S.W. Greenland op.c. Alaska to Labrador, S. to Idaho and Cape Breton I., 140), (GRONTVED 294), p. Iceland op.c. p. mts of W. Greenland, Iceland, Fennoscandia, Arctic Russia, Central Europe, Fennoscandia, N. Russia at least E. to N. Urals and the Kara tundra, single station (HULTEN 1949 to Urals b); area insufficiently known in Tyrol, mouth of Yenisei R., single station at Tju and the phytogeographical position of the species lyma R., few scattered stations in Jakutsk prov. and very uncertain; Dahuria (HULTEN 1941 p. 94); comes rather close to Loiseleuria procumbens : N. North America, Green the amphi-Atlantic (-Alpine) type; land, Iceland, Faroes, Scotland, Fennoscandia, Arctic Poa alpina : N. America (several isolated areas) Russia E. to Urals and lower Obi R., mts of S. and from Alaska to Baffin Land and W. Labrador, S. to Central Europe from Pyrenees to Carpathians, E. Vancouver I., Colorado, Utah, Lake Superior and Siberia from L. Baikal to Chukch Penins., S. to Sagha Cape Briton I., Greenland, Iceland, Jan Mayen, (HULTEN 1948 1229); lin and Yezo p. Spitzbergen, Faroes, Ireland, Scotland, N. England , Bartsia alpina : Arctic E. America, Greenland, Ice Fennoscandia, Esthonia, Arctic Russia E. to Urals, land; Faroes, Scotland, N. England, Fennoscandia, mts of S. and Central Europe from Pyrenees to Cau N. Russia, mts of S. Europe from Portugal and Pyre casus, mts of S. Siberia and Central Asia, Arctic coast nees to Carpathians, Black Forest, Vosges, Sudetes of Asia (probably only few separated areas), Chukch {BOCHER op.c. pp. 170 f.). Penins., reported from Corea and Honshu (Hur.TEN 1942 pp. 199 f., BbCHER op.c. p. 215, GRONTVED op.c. The argumentation concerning the amphi-At p. 145); lantic species with a similar distribution in Arc Viola biflora : arctic-montane Eurasia from Fenno tic Europe can be applied also to the more oce scandia eastwards, but lacking in Siberia N of the mts except along Yenisei R. and Lena R., isolated, anic among these species, viz., Selaginella selagi e.g., in Corsica and the central Urals; in America noides, Agrostis borealis, Juncus trifidus, Arabis only Alaska and (isolated) Colorado (HULTEN 1947 a alpina, Alchemilla glomm·ulans, Sibbaldia pro- p. 1132). Quite likely, identical with the Viola spc. Origins of the Flora 69
reported by SMITH (1920 p. 135) as found subfossil at that time only from the region of the Norse on Mt Helagsfjallet. Indirectly, one can arrive, with settlements in South-Greenland, as having been reasonable certainty, at the conclusion that Viola introduced by the Norsemen. According to him, biflora must be a glacial survivor. The larva of the butterfly Brenthis thore Hb. scandinavica Rygge lives these archeophytes would be no fewer than about exclusively on this plant (WAHLGREN 1941, II p. 17); 50, i.e. 13 % of the Greenland flora. Yet PoRSILD since the insect is a glacial survivor (the species occurs (1932), having subjected those presumed anthro in the Alps, Fennoscandia and North-Russia, the pochores to a careful scrutiny, reduced the num race is endemic, WAHLGREN op.c. I p. 54, map fig. ber of probable >>Old Norse>> plants to 6, viz. 57e), also Viola biflora should belong to this category. Anthoxanthum odoratum, Oarex Lyngbyei, 0. rost
Finally, attention should be called to the Eura rata. Rumex domesticus, Vicia Oracca, and Leon- _ siatic species extending across the Atlantic to todon autumnalis. Since LovE & LOVE (1948 p. Greenland and/or N.E. North America N. of the 105) have shown the Greenlandic Anthoxanthum ancient ice-border. W. of Greenland, they cen to be actually the Arctic type A. alpinum, this ter upon the ice-age refugia in Newfoundland species can, however, safely be regarded as na and round the Gulf of St Lawrence; their New tive (cf. also B6CHER op.c. p. 204). The same World areas extend over larger or lesser parts probably applies also to Leontodon autumnalis, of the region E. of the 80. longitude, which they represented by var. asperior (PORSILD op.c. p. practically never eross. The majority of these 76), which is doubtless indigenous in the Scan plants are absent from. the rest of America; a dinavian mountains and hardly occurs in settled few recur in Alaska as outposts of their .Asia country. Nor seem the arguments by PORSILD tic areas. (op.c. p. 76) concerning Oarex rostrata convin To decide which plants should be included in cing to me; with BocHER (op.c. p. 233), I think this group meets with some difficulties, since it just as probable that >>like many other southern American taxonomists at times, like their Rus species it found its own way to South-Greenland sian colleagues, are a little lavish in creating new and, like the Norsemen themselves, sought out species. A foreign botanist who has not specia the warmest and most subarctic regions>>. lized in the American flora is, therefore, often Among the plants occurring in our region and unable to decide whether an actual race diffe not previously dealt with in detail, the following rence exists or not. When a European species, ones occupy isolated areas in Greenland and/or for instance Oarex vaginata or M inuartia stricta, north-eastern North America, while missing in is reported only from Greenland, while another, the rest of America except, in some cases, in closely related type, in these cases Oarex saltu Alaska and Yukon (and, in two cases, N. Rocky erisis or Minuartia dawsonensis, is supposed to Mts): occur on the American continent, the explana Isoetes echinospora : southernmost Greenland, isol tion may simply be that the Greenlandic flora ated station in W. Greenland 68°33'; in America the was worked up by Danes, using a European no closely related I. Braunii Durieu (BooHER op.c. p. 61); menclature, while the botanists who dealt with Botrychium multifidum: Newfoundland and round the flora of the mainland stuck to the nomen the Gulf of St Lawrence; in other parts of America clature current among their American fellow the related B. silaifolium Presl. (HULTEN 1941 p. 49); Anthoxanthum alpinum: W. and E. Greenland, S. countrymen. In such dubious cases, I have as parts (BooHER op.c. p. 204; LovE & LOVE Le.); far as possible followed the species delimitation A. odoratum introduced to the American continent; jn HULTEN 1941-1949, or else included a spe Agrostis canina : W. and E. Greenland, N. to about cies only when a racial difference between the 67° (BOOHER op.c. p. 203), Newfoundland (FERNALD Greenlandic-East-An1erican population and the 1926 p. 161); introduced to other parts of America; Deschampsia flexuosa : W. and E. Greenland, N. to West-.American one has seen1ed beyond doubt. about 66° (BOOHER op.c. pp. 209 f.), from Ontario It should, perhaps, also be remembered that to Newfoundland, S. to Tennessee and N. Carolina OSTENFELD (1926) considered all plants, known (HYLANDER in LAGERBERG 1937 p. 142); 70 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Nardus stricta : W. and E. Greenland, N. to 60°30' land, N. to 69° and 75° resp. (BocHER op.c., map p. and 63°40' resp. (BOCHER op.c. p. 213), S. E. New 112), S.E. Baffin Land over Ne�ioundland, Mignan foundland (FERN'ALD 1926 p. 81); native in Greenland Is. and S. Maine to E. Pennsylvania; westernmost (PORSILD 1932) and probably in Newfoundland, else Alaska, one station; elsewhere in America ssp. integri where in America introduced; folium (Raf.) Hult. (HULTEN 1945 a p. 898); Carex panicea: S. Greenland, four stations 60°30'- Sedum acre : E. Greenland, 65°30'-66° (BocHER 61° (BocHER op.c. p. 231), southernmost station near op.c. p. 110); introduced to America; hot spring (in Greenland not the typical plant, LAN'GE Rubus saxatilis : S. Greenland 60°15', E. Greenland 1880 p. 149), Nova Scotia, St Pierre, Miquelon (PoR 63°28' (only two stations, ibid. p. 137); SILD 1932 p. 52); Rubus idaeus, Eurasiatic type: W. Newfoundland, Carex rariflora : W. and E. Greenland, N. to about N. shore of Gulf of St Lawrence; elsewhere in Ame 74° (BOCHER op.c. p. 232), Hudson Bay, · Labrador rica ssp. sachalinensis (Levl.) Focke (HULTEN 1946 and S. Baffin Land to Newfoundland and St Lawrence p. 1001); R., Mt Katahdin in Maine; Alaska, N. Mackenzie Potentilla erecta : S. E. Newfoundland (FERNALD distr. (HULTEN' 1942 p. 367); 1933 p. 273); Carex Oederi s. str.: S.W. Greenland, two stations Alchemilla vestita : southernmost Greenland (BocHER (GELERT & OsTENFELD 1902 p. 92, PoRSILD 1930 op.c. p. 127), E. Greenland 66°25', Labrador 51°-52°, p. 13), · Newfoundland and lower St Lawrence R. to Newfoundland (SAMUELSSON 1943 p. 20). Nova Scotia; elsewhere in America C. viridula Michx. Alchemilla filicaulis : W. and E. Greenland, N. to (HUL'l'EN op.c. p. 377); 66°55' and 69°18' resp. (BocHER I.e.), Labrador Carex saxatilis : N. Greenland, W. 64°-78°35', 48°-�52°, Newfoundland (SAMUELSSON I.e., cf. map E. 70°25' and 77° (BOCHER op.c. p. 236), Labrador ibid. p. 19); (HULTEN op.c. p. 382); elsewhere in America the Geranium silvaticum : W. Greenland, one station related C. physocarpa Presl. and C. mernbranacea 62°53', not introduced (PoRSILD 1932 p. 68); Hook. (ibid.); Angelica Archangelica ssp. norvegica: W. and E. Luzula multiflora ssp. frigida : "'r· and E. Green Greenland, N. to 69°47' and 66'25' resp. (BOCHER land N. to 72°31' and 70°28' resp. (BocHER op.c. op. c. p. 149); p. 249), Labrador, Newfoundland, S.E. New Bruns Call una vulgaris : New Y ersey to Newfoundland wick, E. Maine; Alaska (HULTEN 1943 p. 438); else (var atlantica Seem.; FERNALD 1926 pp. 52 f., where in America ssp. comosa (E.Meyer) Hult. (ibid.); GRONTVED op.c. p. 313); Betula tortuosa: W. and E. Greenland, .N. to about Gentianella tenella : E. and W. Greenland, Rocky 61° (BOCHER op.c. p. 71; concerning the Greenlandic Mts., Alaska (HULTEN 1948 p. 1312); and NE -American Betulae, cf. PoRSILD 1932 pp. Pedicularis hirsuta : N. Greenland, S. on the W. 55 ff.); coast to 64°10', on the E. coast to 65°35', Arctic Archipelago (BocHER op.c. p. 172); Betula· nana : W. and E. Greenland about 63°-75°, Utricularia minor: W. Greenland, 60°53' and 68°21' lacking in S. Greenland (BOCHER op.c., map p. 69), -70°59' (PORSILD 1935 p. 26); Labrador to Hudson Bay (GRONTVED 1942 p. 207); Galium palustre : S. Greenland, Newfoundland, Rumex Acetosa ssp. arifolius : S. and W. Greenland 1;\.nticosti, Gaspe and Nova Scotia to Connecticut 60°-62° (BocHER op.c. p. 73), Alaska, E. to Mackenzie and New York (FERNALD 1925, map 48, BRITTON & Delta and (isolated?) in Rocky Mts of W. British Co BROWN 1913 p. 265); lumbia and Alberta (HULTEN 1944 pp. 597 f.); ace. Erigeron uniflorum, excl. var. eriocephalum : W. PoRSILD (1932) most likely native in Greenland; and E. Greenland, N. to 67°42' and about 72° resp. Rumex tenuifolius : W. and E. Greenland, N. to (BOCHER op.c. p. 193); not in America (MALTE 1934 b 72°48' and 74°30' resp. (GELTING 1934, map p. 283, p. 191); sub R. Acetosella; LovE 1941 p. 101); R. Acetosella Leontodon autumnalis : W. Greenland 60°49'-61 o s. lat. introduced to America, chiefly ssp. angiocarpa (BOCHER op.c. p. 197). Murb. (HULTEN 1944 p. 599); Taraxacum purpuridens, reported from Greenland Sagina nodosa : S. Greenland, two stations; Ameri by SEIDENFADEN (1933 p. 76) and DEVOLD & SCHOLAN can Atlantic coast from 42° to 51°, W. to the Great DER (1933 106), Lakes (BocHER op.c. p. 83); p. probably does not belong to this Fennoscandian species, but to an endemic Green Ranunculus acris : vV. and E. Greenland, N. to landic one (verbal information by Dr G. HAGLUND ) . about 66° (ibid. p. 88; in Greenland only >>types closely allied to the main type>>, not var. pumilus nor ssp. A more thorough perusal of the floristical lite borealis, ibid.); E. Aleutians (var. frigidus Rgl, HuL TEN op.c. pp. 745 f.); introduced to America (ibid.); rature would, no doubt, reveal more instances Sedum Rosea, Eurasiatic type: �T. and E. Green- of such an >>east-arctic>> . distribution. Origins of the Flora 71
Discussion of the amphi-Atlantic species distribution, approaching the last-mentioned type or identical with it, often must be due to The significance to the problem of the corn the cause propounded by HULTEN, viz., that a position of the Fennoscandian ice-age flora of formerly circumpolar species has failed to sur these >>east-arctic>> species and their Transatlantic vive the glaciations except in a few of the un ranges is obviously determined by the general glaciated areas scattered around the earth. interpretation given to the amphi-Atlantic type However, it is, in some cases, almost impos of distribution. It has been made abundantly sible to conceive the present areas, even of very clear, above all by FERNALD (e.g., 1924, 1925, old units, as resulting from a reduction of for 1926, 1929, 1931), that plant species, which mer circumpolar ones during the glacial periods. in north-eastern North America have isolated Ranunculus glacial is, for instance, is an isolated, areas within the regions · in question, survived evidently very old species. Its distribution (cf. at least the latest glaciation in refugia at or close above p. 66) renders its European origin as good to their present stations. Likewise, the existence as certain. Being the most hardy of Scandina of unglaciated areas in Greenland cannot be vian alpine plants, it could scarcely become ex doubted (cf. e.g. GELTING 1934 p. 250 and map tinct over immense areas in consequence of one p. 252). Even those who do not share the opinion or more glaciations. Finally, it is replaced in of B6CHER (op.c. p. 312), >>that practically . an the Bering Sea region by its only close relative, Greenland plants immigrated in the last inter R. Chamissonis Schlecht. (HULTEN 1944 p. 753 glacial period or earlier and survived the last gla f.). Doubtless most arctic and arctic-montane ciation in Greenland itself>>, and that >>conditions plants radiated from that region; however, to during the last ice epoch may have been very assert that also R. glacialis did so seems far too similar to those of the 'present ice epoch'>> ( op.c. dogmatic. This would imply, either that this p. 319), will hardly oppose the more moderate plant should have reached East-Greenland, on views of GELTING (op.c. p. 264 ff.), according to the one side, and Fennoscandia and the Alps, whom at least the plants with a >>centric>> distri on the other, without undergoing the slightest bution in Greenland hibernated · as it were on change, while the population remaining in Be the spot. When not introduced by Man, the Eura ringia changed into R. Chamissonis; or that the siatic species with isolated areas in Greenland Beringia population should have remained un can be rega�ded as glacial survivors over there. changed, while the migrating populations changed If, like HULTEN, one considers all or nearly in an identical way in Greenland and in Eu all amphi-Atlantic species to have occurred ear rope.. Both assumptions are equally incredible. lier also in the remaining parts of arctic and Far simpler seems the hypothesis that the pre northern .North .America al\d to have died out sent areas of R. glacialis are remnants of a for there during the ice-age, the fact of their being mer continuous, Transatlantic· range, limited by survivors in Greenland and north-eastern North the oceanic character of the species, and that America fails to tell us anything about the past the small Beringia population migrated there of the Fennoscandian alpine flora, but that these long ago, probably already in the Tertiary, species were able to endure the hardships of a then changing into R. Chamissonis. Or else R. hibernation, at any rate in refugia as· large as glacialis s. lat. (incl. R. Chamissonis) had once the Transatlantic ones. a circumpolar distribution, · which was split, not Personally, I agree with HULTEN as to many during the ice-age, but earlier, when the present amphi-Atlantic plants, especially among the an continental climates of Asia and .America came cient taxonomic units. There seems to be no into existence. However, in that case its pre escape from the fact that a continuous series of sent range would date so far back, that specula distributional types connects the circumpolar tions as to its genesis become meaningless. The type with the amphi-Atlantic one, and that a history of other amphi-Atlantic linneons, which 72 SELANDER: Floristic Phytogeography of South-Western Lule Lapp mark are replaced by related types in the Bering Sea parts of America, and other races farther to the region, is probably similar. west. Such is the state of, for instance, Athy Comparatively young, amphi-Atlantic micro riurri alpestre, the Eurasiaticum form and var. species and races within critical genera offer just americanum Butt. (BUTTERS 1917 p. 204, FER as intricate problems. Time and again, HuLTEN NALD 1928 p. 44), Carex norvegica. s. str. and has himself emphasized the fact that circumpo other, related types, C. Oederi and C. viridula, lar macrospecies generally develop vicarious lo C. saxatilis and 0. physocarpa &c., Luzula mul cal races in different parts of their total range. tiflora ssp. frigida and ssp. comosa, Coeloglossum NOW N ANNFELDT (1940 p. 41 ff.) has pointed out viride ssp. islandicum and ssp. bracteatum, Sedum the utter improbability that such a macrospecies Rosea and its ssp. integrifolium, Rubus idaeus would be represented by exactly the same race and its ssp. sachalinensis, Potentilla Chamissonis on the two outermost borders of its range and and P. Hookeriana, the minor races within P. by other races in the interjacent areas. However, nivea s. str. (HULTEN 1945 bp. 134), Diapensia this is what the theory of HuLTEN means when lapponica and its ssp. obovata, Veronica alpina, applied to amphi-Atlantic plants of this kind. pumila and Wormskioldi, Erigeron uniflorum The original biotype compounds of the hypothet and its var. eriocephalum, Antennaria alpina and ical circumpolar populations would, of course, numerous, closely related species, &c., &c. (cf. have been depauperated towards the outskirts above pp. 69 ff.). Should such a distribution be of their ranges. This fact still more reduces the a rare phenomenon, a polytopic origin of the chances of precisely the same ecotypes reaching European types occurring in Greenland and the opposite borders: yet the same microspecies north-eastern America would, of course, be con are often distributed on both sides of Denmark ceivable; however, such cases are far too nume Street. It is, for instance, hardly conceivable rous to admit, invariably, of this explanation. that such a small unit as the ssp. caespitans with In the oceanic parts of southern Greenland, in the most complex Poa arctica should have where the climate fairly resembles that of nor spread round the whole Arctic Ocean without thern Scandinavia, the flora is dominated by undergoing appreciable morphological changes, European species, both as to numbers and fre and then have disappeared entirely from the grea quency of the species. BocHER (1938 p. 3i1) ter part of its area, particularly in view of the declares that parts of south-eastern Greenland, fact that its present distribution proves it to be especially the Angmagsalik district, >>clearly be among the most hardy of all vascular plants. Poa long to the European flora region>>; and E. DAHL arctica ssp. caespitans is by no means a unique (1946 p. 227) states, that >>the macrolichen flora case; on the contrary, the examples could easily of southern Greenland has a relationship with be multiplied (NANNFELDT 1. c.; for further in the Scandinavian ar:ctic-alpine macrolich�n flora stances cf. SAMUELSSON 1943 pp. 28 ff.). which approches identity>>. If HULTEN's theory were always applicable in These facts, added to those assembled by this particular respect, the Eurasiatic races with N .ANNFELDT and SAMUELSSON, seem to streng in the macro species would appear in western Ame then the opinion >>that a direct exchange must rica, and the American ones farther to the east. have taken place between the biota of Scandi Such a distribution is actually often met with. navia and Arctic North-America in not too Especially a type of distribution, common above remote timeS>> (N.ANNFELDT 1. c.). all among older units, where the >>Cordilleran>> Whether this exchange is to b� explained by race recurs in the north -eastern corner of the the wegenerian drift hypothesis or by the assump Continent and sometimes in the refugia near the tion of some kind of Transatlantic land bridges, Great Lakes, conforms obviously to his theory. is a question beyond the scope of this discussion. In other cases, however, the European races are In both cases, however, at least part of the mi to be found in Greenland, andjor the adjacent grating species must have passed by way either Origins of the Flora 73 of Spitzbergen or of Iceland, the hypothetical flora. As pointed out already by N ATHORST (1883) land bridges extending over the Faroes and Ice and further particularized by ANDERSSON & HEs land to Greenland and from Nova Zembla over SELMAN (1900), a number of plants, mostly less Spitzbergen to Greenland (cf., e.g., STEFFEN 1937 high-arctic than those mentioned above, never B pp. 388 ff., NANNFELDT 1940 pp. 39 f.). There bear fruit in Spitzbergen even under present con fore, the floras of Spitzbergen and Iceland should ditions. Some of these species evidently propa probably indicate the time when such a hypo gate asexually all over the Arctic. In other cases, thetical migration took place. especially in a number of plants that are very Since most vascular plants of Spitzbergen have rare in Spitzbergen and lacking in northern a holarctic distribution, plant-geographical con Greenland, Franz J osef Land and Lutke Land nexions exist both to the west, viz., to Green north of :1\'I atotchkin Shar (or in all Greenland land, and to the east, viz., to Nova Zembla. and/or Nova Zembla; cf. BocHER 1938, HANSEN Instances of plants definitely originating from & LID 1932 and LYNGE 1923) this asexual state the west are Poa Hartzii Gdgr; Serens., P. ab would seem not to be normal. To this category brcviata R. Br., Minuartia Rossii (R. Br.) Graebn., belong the following plants: Potentilla pulchella and probably several addi Calamagrostis neglecta ssp. borealis.(ANDERSSON tional species, e.g., Carex nardina and Campa & HESSELMAN op.c. p. 77), Luzula Wahlenbergii nula uniflora. On the other hand, an eastern (ibid. p. 82), Tofieldia pusilla (ibid. p. 83), Betula provenance is evident in the cases of, e.g. , Phipp nana (ibid. p. 66), Rubus Chamaemorus (ibid. p. sia concinna, Salix polaris, Ranunculus Pal 18), Empetrum hermaphroditum (ibid. p. 32) and lasii (HULTEN 1944 p. 764), Potentilla multifida probably Cassiope hypnoides (ibid. p. 18). (HULTEN 1945a p. 1021) and Taraxacum arc The same applies probably to a few other ticum (Trautv.) Dahlst. (DAHLSTEDT 1928 p. species, e.g., Carex Lachenalii, Vaccinium uligi 59.) All plants that have clearly travelled by nosum and Campanula rotundifolia (NATHORST the >}bridge>} between Nova Zembla and Green 1883 p. 63, ASPLUND 1918 p. 36). land are of necessity so extremely high -arctic Especially interesting is Empetrum hermaphro as to render it almost impossible to say anything ditum which, according to ANDERSSON & HESSEL about their past. They may well have lived MAN (1. c.), ripened its fruits in Spitzbergen du through the entire ice-age in the unglaciated areas· ring the postglacial warm period; its sterility of N. Spitzbergen (cf. I.. YNGE 1933, 1934). The must thus be due only to the present unfavou fact that the endemics of the Spitzbergen flora rable climate. (Ranunculus Wilanderi (Nath.) Freyn and the These plants, which even now barely nmnage neoendemic R. spitzbergensis Hadac, that has evi to survive, apparently could not have endured dently arisen out of the hybrid R. lapponicus x a glaciation in Spitzbergen; moreover, at least Pallasii, cf. ANDERSSON & HESSELMAN 1.900 pp. a few among them occur only in places where de 42 ff. and DAHL & HADAC 1946 p. 11) are so very finitely no ungl;:tciated refugia were situated, e.g. few does hardly disprove its supposed high age. Salix herbacea on South Cape and Empetrum The paucity of the endemic element here and in hermaphroditum on Bell Sound. Their distribu Greenland and its complete absence from the tion, with gaps in the neighbouring Arctic re Arctic American archipelago may be caused i.a. gions, renders a Fennoscandian provenance al by the high-arctic climate, the rate of mutations most certain. Rub�ts Chamaemorus, Empetrun� decreasing directly with lowered temperature, at and Vaccinium uliginosum can, possibly, have least under experimental conditions (DAUBEN been spread by migratory birds in postglacial MIRE 1947 p. 215). time. The anemochores, viz., Calamagrostis However, another element in the flora of neglecta ssp. borealis, Betula nana and Cassiope Spitzbergen is of a more immediate interest in hypnoides, may conceivably have been carried eonnexion with the history of the Scandinavian to Spitzbergen by convective air currents. The 74 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
possibility of such a transport is proved by the It does not seem indispensable to presume the occurrence on snow-fields in N. E. Spitzbergen existence of Transatlantic connexions of a more of the dipter Syrphus ribesii L. and the aphis · recent date than the early Quaternary at the Dilachnus Piceae Panz., which must have been very latest. On the other hand, a connexion brought there by a wind-transport of at least with Fennoscandia probably existed rather late, 1300 km (ELTON 1925 pp. 291 ff.). The remaining possibly even after the maxinium of the Wiirm species, Carex Lachenalii, Luxula Wahlenbergii, glaciation. Tofieldia pusilla and Campanula rotundifolia, may The opinions among bio-geographers differ have migrated over land from Fennoscandia at greatly as to the possibilities of migrations to so recent a date as after the maximum of the last Iceland and, consequently, as to the history of glaciation; in any case, they can hardly have sur the Icelandic biota. While, e.g., LINDROTH (1931 vived the Riss glaciation in Spitzbergen and thus pp. 551 ff.) and HADAC (1948 p. 22) assume that probably belonged to the Fennoscandian ice a land bridge between Iceland and the British age flora. Isles existed as late as during the last intergla There are, perhaps, other indications of migra cial, the idea of any such connexion is emphat tions between Spitzbergen and Fennoscandia, ically repudiated, e.g., by LOVE (1948 p. 106). though in a reverse direction. A few high-arctic The geologists are unanimous in rejecting these plants common in Spitzbergen, viz., Braya pur hypothetical post-tertiary land bridges; on the purascens, Potentilla pulchella, and Cerastium other hand, they are quite willing to accept the Regelii, are found in single, isolated stations on existence of unglaciated areas, particularly in the coast of northernmost Fennoscandia ( Ceras the north-western, northern, and eastern parts of tium Regelii on the Kola Peninsula, KOTILAINEN the island (cf. THORARINSSON 1937 pp . 171 ff., 1942 p. 72; for the total area, cf. GELTING 1934 map p. 174). map p. 43). They may have migrated there from .As is well known, the Icelandic flora is char Spitzbergen, though we cannot say when. acterized by its close affinity to the Scandina Such spreading seems plausible in the first vian montane flora (cf. GR0NLUND 1884, STROM place with regard to Papaver Dahlianum. Be FELT 1884, WARMING 1888, GRONTVED 1942). ing endemic in Spitzbergen and northernmost Out of about 370 apparently indigenous, vas- . Norway (NORDHAGEN 1931 p. 44; cf., however, cular plants (Taraxaca and Hieracia excluded), the occurrence in Greenland of P. radicatum var. only a few are missing in Scandinavia and the albiflorum Lge, which type probably >>comes un British Isles. They are at most nine in all (en der P. Dahlianum>>, BOCIIER op.c. p. 99), this demics and near-endemics excepted), viz., Carex plant is quite common in Spitzbergen, whereas Lyngbyei, Platanthera hypm·borea, Leuchorchis in Norway it does not occur outside a few isola straminea (Fern.) Love (cf. FERNALD 1926 b ted stations on the northern coast of the Varanger p. 17 4 and LovE 150 pp. 36 f.), Cakile eden Peninsula. Furthermore, while .specimens with tula (LOVE 1945 p. 154, LOVE & LOVE 1947), white or yellow flowers are intermingled in Spitz Cochlearia groenlandica (LOVE & LovE 1948 p. bergen, the different flower-colours occur in se 58), Saxifraga Aizoon ssp. neogaea (ibid. p. 62), parate Norwegian stations (NORDHAGEN op.e. p. Chamaenerium latifolium, Lomatogonium rotatum 12). This is, apparently, most readily explained and possibly Galium Brandegeei (cf. PoRSILD by the hypothesis that Papaver Dahlianum arose 1930 p. 26). Cochlearia groenlandica may have in Spitzbergen during the ice-age and then spread been carried there by the Polar Current, e.g., to Norway, the uniform population in each Nor from Spitzbergen, and Cakile edentula most prob wegian station originating from plants with a ably was transferred from America by the Gulf hereditary disposition but for one flower colour. Stream in postglacial times (LovE & LOVE 194 7 The conclusions that may be drawn from the p. 17). The rest, as well as about 20 other spe flora of Spitzbergen can be summarized as follows: cies, e.g., Carex nardina, C. macloviana, Sagina Origins of the Flora 75 caespitosa, Pedicularis flammea, &c., are obvious the miocene and that it most probably exis ly old arrivals from the New World. They may ted in the Scandinavian floral area ever since well have lived in Iceland through the ice-age, then>>. If this be true, also other rather thermo all of them being rather high-arctic, and the philous plants may have >>hibernated>>, at any great majority very old units as well. rate in Iceland. Till lately, the Icelandic flora was believed to Among the southern element, probable sur Jack endemics, a fact that was considered to vivors are, in the first place, such plants as grow speak against its possessing a high age. ·Recently, exclusively or mainly at hot springs and solfa however, several endemic races have been dis taras, the flora at hot springs everywhere being eovered, viz., Ophioglossum vulgatum var. islan rather queer and often comprising exotic species .dicum (LOVE & LOVE 1948 p. 105), Roegneria is which occur very far from their nearest stations, landica (ibid.), R. Doniana var. islandica (ibid.), apparently as Tertiary relics, e.g., the tropic-sub Orchis maculata ssp. islandica (ibid. p. 106), tropic Kyllingia brevifolia Rottb. and Fimbristylis Papaver radicatum ssp. islandicum and ssp. Ste annua (All.) Roem. & Sch. in their completely janssonii (LovE 1945 p. 149), Euphrasia Davids isolated stations in Kamtchatka (cf. HULTEN sonii (PUGSLEY 1933 p. 308) and Galium pumi 1927 pp. 157 ff. and pp. 170 f.). Characteristic of lum ssp. islandicum (STERNER 1944 p. 213); be., such habitats in Iceland are (GRONTVED 1942 .sides, there are the two near-endemics Papaver pp. 46 ff.) Ophioglossum vulgatum var. islandicum, radicatum ssp. jaeroense and Alchemilla faeroensi8. Blechnum Spicant, Juncus articulatus, J. bufo In this respect, the flora of Iceland shows every nius, Polygonurn Persicaria, Gardamine hirsuta, .sign of being as old as that of Fennoscandia. Grassula aquatica, Hydrocotyle vulgaris, Veronica No fewer than about 70 indigenous Icelandic A nagallis-aquatica, Plantago lanceolata, Pl. major .species are completely lacking in the New World, f. pygmaea Stefanss., Gnaphalium uliginosum. A and at least 140 undoubtedly originate from · few of these have possibly been introduced by Europe. Especially interesting is the finding of Man; the rest may well have survived at least Poa arctica ssp. depauperata in an ice-age refu the latest glaciation in their peculiar habitats, gium of northern Iceland (LovE 1947). The oc a lowering of the mean air temperature by a ·Currence of this race, that previously was re few degrees evidently having but little effect near garded as endemic in the South-Norwegian moun a hot spring, the water of which has a tempera tains, evidently suggests a connection with Nor ture of 70° C. or more. Furthermore, as the Ice way at a time that is, geologically speaking, not landic race of Ophioglossum vulgatum consti too distant (cf. NANNFELDT 1947 p. 82). t.utes an endemic variety, its high age as a mem Obviously, many species besides those men.. . ber of the flora of Iceland is, indeed, very prob tioned above can be supposed to have hi?er able; and if one of the southern species of the nated in Iceland; this holds good perhaps of the flora at hot springs and solfat.aras is a glacial major part of the Icelandic vascular plants. survivor, others may as well be so. LOVE & LOVE (194 7 p. 17) go, in fact, as far as On the other hand, not a few southern Ice to assume, that about four fifths of the native landic plants are so strongly hemerophilous that flora have survived at least one gl�ciation on the w� may suspect them of having been introduced spot and, possibly, the whole ice-age, the only to the island, although they now seem to be na exceptions being plants transported by sea cur tive. Yet even though all such species be exclu rents. This opinion is supported by LINDQUIST's ded, there remains a residue of plants, occurring establishment of Betula callosa in Tertiary de - to judge from literature only - in such in posits of Iceland. LINDQUIST (1947b p. 354) tact plant communities and so far from human considers that >>in view of the finds at phri habitations that they must be regarded as in milsoalur it may be asserted that this type of digenous, however unlikely they would seem to birch has not changed morphologically since be able to survive a glaciation in Iceland. 76 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
A foreigner not having visited Iceland is in Spitzbergen, besides the widespread, >>west hardly entitled to express an opinion of these arctic>> T. croceum coiL, is T. brachyceras Dahlst., questions. It strikes me, nevt:rtheless, as most recently identified by HAGLUND with the Fenno unlikely that, e.g. , Rosa dumalis (R. Afzeliana scandian, bicentric T. simulum (verbal informa in GRONTVED 1942 and LOVE & LOVE 1947) tion by Dr HAGLUND). Both T. brachyceras, should belong to the hibernating element, as sug which occurs in Greenland, Spitzbergen, Fenno gested by LovE & LovE (op.c. p. 18), since this scandia and Waigatch (cf. DAHLSTEDT 1906 p. 20, species, to judge from evidence available in the BOCHER op.c. p. 198), and the other dandelion literature, even under present conditions barely of Spitzbergen, T. arcticum (Trautv.) Dahlst., survives in its single Icelandic station, never which is evidently of a Eurasiatic origin, though flowers, &c. (GRONTVED 1942 p. 277); further reaching East Greenland (cf. above p. 73), are more, it occurs within a glaciated area. Sieglingia rather isolated, seemingly very old units. None of decumbens (ibid. p. 184), Carex flacca (ibid. p. them necessitates a late Transatlantic connexion. 164), Cardamine flexuosa (LOVE 1945 p. 159), and Among the native Taraxaca of Greenland, T. Rosa pimpinellifolia (GRONTVED op.c. p. 277) are cymbifolium (BOCHER op.c. p. 189, sub T. mau other instances of definitely southern plants oc rostylum), T. naevosum (ibid.), T. rhodolepis curring in single stations in those parts of Ice (ibid.; rather doubtful, possibly an endemic spe land where probably no unglaciated refugia exis cies, ace. to verbal information by Dr HAGLUND), ted, i.e. the South and the South-west (cf., how and T. brachyceras (ibid. p. 198) occur also in ever, LINDROTH 1931 p. 266, who for zoo-geogra Scandinavia. To judge from its curiously split phical reasons places a refugium >>im aussersten distribution (Greenland, Iceland, northernmost Siiden unterhalb des Myrdalsjokulls>>; Rosa pim Norway; verbal information by Dr G. HAGLUND), pinellifolia has one or two stations also in the which is unique among amphi-Atlantic areas, T. presumably unglaciated East). It would seem cymbifolium is a ·very old unit, dating back at most unlikely that such plants should be gla the last interglacial or probably still earlier. cial survivors. They are all of a European ori Since three of its four known Greenlandic sta gin, Cardamine flexuosa occurring in America on tions are situated in East-Greenland, it cannot be ly in Newfoundland, and the rest altogether lac among the >>old Norse>> plants. This applies also king there. Their presence in Iceland is appa to T. rhodolepis, its only known station being in rently an indication of comparatively recent im East-Greenland. On the other hand, T. naevosum migrations from Scandinavia andjor the Brit that has been found only in South-Greenland ish Isles. may have been introduced by the Norsemen. On the other hand, the occurrence of so ther The age of the two last-mentioned species can mophilous glacial survivors as, e.g., Juncus squar not be fixed with any degree of certainty. rosus and Polygala serpyllifolia in Greenland Scandinavian taxonomists have arrived at the (BOCHER 1938 pp. 244 ff. and 138 f.), or Ranun conception that all the circumpolar macrospecies culus hederaceus, Potentilla sterilis, P. anglica, and within Taraxacum can be completely divided in Galium hercynicum in Newfoundland (provided to, partly vicarious, microspecies, each contain they were not introduced to their only American ing one, ·or a few, biotypes (cf. GUSTAFSSON 1947 stations, on the Avalon Peninsula; cf. FERNALD pp. 259 ff.). Since the amphi-Atlantic Taraxa 1926 p. 236, 1933 p. 273) is undeniably an argu cum microspecies do not occur in the Bering Sea ment in favour of LOVE's thesis, that practically region (cf. HAGLUND 1937, 1946, 1948, 1949), but all the native Icelandic flora has survived at least are partly replaced there, - and in the interjacent the latest glaciation on the spot. regions, by vicarious agamospecies, they cannot More illuminating as to the Transatlantic con have radiated from those parts. Their distribu nexions are the Taraxacum and Hieracium floras. tion constitutes, therefore, one of the main ar The only Fennoscandian Taraxacum occurring guments in favour of a Transatlantic land con- Origins of the Flora 77 nexion of some kind or other reaching Green southern Norway and the Sudetic Mts (ELF land. However, this connexion may well be STRAND 1927 pp. 269 f.). placed as far back as to the early Quaternary No Hieracium occurs in Spitzbergen. And ac or the late Tertiary. cording to OMANG (1933 a, 1933 b, 1937), Green As to the Icelandic Taraxaca, it is rather diffi land and Scandinavia have no I-Iieracia in com cult to obtain a clear view of their total areas, mon but the widespread, >)west-arctic>), very old since their monographer CHRISTIANSEN (1942) H. alpinum, at least not in identical forms; the seems inclined, now and then, to use a nomen Scandinavian H. (sect. Prenanthoidea) thulense, clature of his own. Anyhow, according to CHRIS reported by PoRSILD (1932 p. 78) from Green TIANSEN (op.c. ) and l-IAGLUND (in LOVE 1945) , land, is according to PORSILD (L c.) generally ::E'ennoscandia and Iceland have at least the fol regarded as preglacial. The Hieracium flora of lowing indigenous Taraxacum microspecies in Greenland is rather poor, comprising oDly a do common: T. brachyceras, T. obscurans, T. sticto zen microspecies, all told. Out of these, all are phyllum, T. spectabile, T. craspedotum, T. acido endemic but H. groenlandicum Aln1qu., that oc tum, T. repletum, T. faeroense, T. calanthum, T. curs also in north-eastern America (cf. FERNALD naevosum, T. cymbifolium, and T. Hypochoeris; 1926 p. 241); H. angmaksalikense Omang (Al a few nwre among 0HRISTIANSEN's Icelandic dan pina vera) is rather closely related to the · Scan delions will probably turn out to be identical dinavian H. Sundbergii (OMANG 1937 p. 5 ). with Scandinavian types . . Further, according to :Much closer are the relations between the CHRISTIANSEN (op.c.), Greenland and Iceland Hieracium flora of Iceland and that of Fenno have several native Taraxaca in common, viz., scandia. True, OMANG (1938 p. 4 f.) characterizes T. pleniflorum, T. atroglaucum (CHRISTIANSEN the Icelandic Hieracium flora >)hauptsachlich als in GRONTVED 1942 p. 398), T. cyclocentrum, T. eine endemische Erscheinung>>; on the other hand dilutisquameum, and T. latispinulosum, besides he asserts >)dass die I-Iauptmasse der islandischen the already mentioned T. naevosum and T. Hieraciumflora ihren Ursprung von demselben cymbifolium. Our present knowledge of the distri Grundstamme wie die skandinavische I-Iieracium bution of the Iceland Taraxaca does ·not admit flora herleitet, und dass sie als ein Setzling der of any conclusions concerning the age of the for selben aufgefasst werden muss, der durch Isolie mer Transatlantic land connexions; it simply con rung seinen unterscheidenden Oharakter bekom firms their existence. men hat>>. Some Fennoscandian Hieracia, though Among the Fennoscandian microspecies with admittedly very few, appear in Iceland in alto in the genus Hieracium, most are quite young; gether identical forms, viz.; besides H. alpinurn, many must have arisen as late as after the Lito the three Prenanthoides species thulense, furves rina maximum, i.e. during the last four or five cens and basileion and the Rigidum species stic thousand years (cf. JoHANSSON 1926), some even tophyllum. A few others occur in forms which de in the course of the last millenium or so (cf. viate so little that OMANG does not even describe SAMUELSSON 1910 b). On the other hand, a few them as formae, viz., H. nordlandicum (Nigres microspecies, morphologically not more distinct centia), H. praeflocellum (Silvatica), and H. hoi than the young units, are evidently very old; dalicum (Caesia). An additional three are re according to SAMUELSSON (1943 p. 26), their age presented by insignificant varieties, viz., H. pe 1nust >)jedenfalls bis zur altquartaren Zeit zu- tiolatum and H. unguiculiferum (Alpina vera) . riickgehen>). As examples can be quoted H. kuu and H. leuceriodes (Nigrescentia). The remaining samoense, H. subarctoum, and H. subpellucidum, microspecies are endemic except two, both be which occur, in identical forms, in Fennoscandia, longing to the West-European Cerinthoidea, and in Transbaikal or the Yenisei region (SAMU which OMANG interprets as (endemic) varieties ELSSON l. c.), H. tubulosum, occurring in northern of microspecies, being at home the one in the Norway and Riesengebirge, and H. oncodes in Faroes and the other in the British Isles. On the 78 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark whole, however, endemism is hardly much more p. 283 ff. ), regard long-distance dispersal as be widespread among the Icelandic Hieracia than ing devoid of any practical significance. among the South -Swedish, mainly postglacial It is, in fact, difficult to imagine how such microspecies of the genus. a dispersal would take place. Sea currents can The differences between the Icelandic Hiera be thought of merely in coimexion with seashore cium flora and the Fennoscandian one are not plants, with seeds or fruits especially fit for in the least surprising. JoHANSSON (1926 p. water transport, and perhaps, as rare exceptions, 323 ff.) has demonstrated, by carefully investi in the case of diaspores attached to drift-ice or gating the areas of Scandinavian and Central rafts of driftwood. As to dispersal by birds, European Archieracia, that as far as we know practically all critical observers agree upon the no species has made a leap from North or Central improbability of birds carrying seeds across the Sweden to Germany or vice versa. All Swedish oceans (cf. e.g. LovE & LOVE 1947 p. 14 f. and lowland Hieracia are, of course, either post the literature cited there). And even though glacial immigrants or have arisen, as it were, comose diaspores can be blown many hundreds on the spot, and all of them have continuous of kilometers over land and sea (cf. e.g. DAUBEN distributions within larger or smaller areas. The MIRE 1947 p. 295), a few instances being known past of the vast majority of Scandinavian Archie of their having been carried as far as about racia thus being very short, we can not expect 1000 km, i.e. a distance as large as that be to encounter many of them in such an isolated tween Scandinavia and Iceland, there are hardly and out of the way place as Iceland. It is, if any any cases observed of such dissemination having thing, rather astonishing, that the affinities of resulted in ecesis (cf. especially SETCHELL 1935 the Icelandic and the Fennoscandian H ieracium pp. 289 ff.). If long-distance dispersal by air floras have not been more obliterated by Time; currents had played an important part in the they make it abundantly clear that a connexion recruiting of the flora of, e.g., Iceland, the Ame between these two regions must have existed du rican element of the flora would have been much ring a late phase of the Quaternary age. On the stronger than actually it is, since westerly winds other hand, nor the Hieracia necessitate a con prevail absolutely in the higher air strata (cf. nexion with Greenland, more recent than the LINDROTH 1931 pp. 522 ff.). late Tertiary or the early Quaternary. The state in Iceland and Spitzbergen therefore Supposing the European element within the seems to presuppose Quaternary land connexions floras of Iceland and Spitzbergen to originate with Europe of a comparatively recent date, poss from a period, when the distribution of land and ibly as late as after the maximum of the Wiirm water in the North-Atlantic and the Polar Sea glaciation and, in any case, later than the maxi was substantially the same as now, its presence mum Riss glaciation. The >>bridges>> need not there would of necessity be due to a long-distance have been continuous. Especially in those parts dispersal on a scale which scientists otherwise of the earth, where the sea in winter becomes will never admit of. l\'Iodern phytogeographers frozen or choked with pack-ice, even rather broad who have, for instance, devoted themselves, in waters will probably not constitute any serious a thorough-going way, to the floras of isolated hindrance to the dispersal of plants by the wind. islands, mostly turn down the idea of plants On the contrary, the relatively low number, and spreading across huge expanses of water (cf. e.g. fortuitous character, of the species that seem to SKOTTSBERG 1928 p. 917 f. and 1931 p. 64 f.). have crossed the ocean by the hypothetical When discussing the problems of bipolar plant >>bridges>> becomes more easily explainable, if the distribution, Du RIETZ (1940a p. 240) also denies latter are conceived as consisting of islands, se the possibility of accidental dispersal over very parated by expanses of water, broad enough to large distances. And the textbooks of plant geo obstruct, but not to prevent, the passage of graphy, e.g. HERZOG (1933 p. 24) and CAIN (1944 plants. Origins of the Flora 79
Is it eonceivable, then, that such >>land bridges>>, tions necessitated by the plant-geographical data. or rather chains of islands, have existed as late Large parts of the submarine ridge between the as has been postulated� No outsider can, of Faroes and Iceland would rise above the surface, course, answer this question; at most he can make and the Polar Sea between Norway and Spitz a guess. It seems to me that HULTEN has poin bergen, being on an average about 400 m deep, ted the way to a solution of the problem when would be changed into land to such an extent writing (1937 a p. 125) as follows: as to render a dispersal of plants, on the scale >>The possibility of the dispersal of plants spe suggested above, quite possible. The broadest cially fit for spreading over the sea from Europe remaining arms of the sea would separate Green to E. America and vice versa is naturally not land fron1 Iceland and Spitzbergen, which tallies excluded, although that has not been the history with the fact, pointed out already by W.ARMING of most of the 'amphi-atlantic' plants. It should (1888), that the main plant-geographical divi especially be pointed out that during the glacials ding-line goes through Denmark Street and far the land area, also on the Atlantic side of the ther north to the west of Spitzbergen. globe, must have been far larger than it is at pre It should also be noted, that the sea must have sent. If we assume that the sea-level was lowered maintained a lower level than the present one 200 m, land would extend far beyond the Shet for a considerable time even after the climatic land Islands, and the Faeroe Islands be separated change at the end of the glaciations, i.e. during from the continent by only a narrow channel, parts of the millenia that the melting of the ice Rockall bank would be land and Iceland and caps lasted; and during these times, i.e. during Greenland considerably larger than they are the fini-glacial epochs, even rather termophile now. The sea-level can, however, have been lo plants may have migrated from Europe to the wered so much as 200 m only during the maxi islands of the North-Atlantic and the Polar Sea. mum glaciation. But the isostatic pressure must On the other hand, the presumed land connexions have contributed to the formation of land off would have been broken off during the intergla the northwestern part of the European conti cials, as they are nowadays. nent. It is evident, therefore, that during the Whatever the significance attributed to the glacials the possibilities of plants spreading from above hypothesis, the essential fact in regard to island to island must have been greater than now.>> the composition of the Scandinavian ice-age · The essential point of HULTEN's statement is flora is the apparent inevitability of the concep obviously the increased possibilities of Trans tion of posttertiary, and probably late-quater atlantic dispersal of plants during the gla nary, land connexions of some kind or other be ciations and not, as assumed by most authors, tween Scandinavia, on the one side, and Ice during the inte rglacials, when the exi land and Spitzbergen, on the other . . stence of any kind of land connexions seems ut A connexion between Greenland and Europe terly improbable. It appears to me that we need is another prerequisite. No phytogeographical not proceed much farther on the way indicated facts, however, would seem to contradict the by HULTEN, in order to find an explanation of assumption, that this connexion was in existence the features of tlie floras of Iceland and Spitz only in fairly remote times, i.e. at the latest du bergen discussed above. ring the Riss glaciation. In Inany cases, the As is well known, the ice-caps are assumed to distributions of American plants extending into have bound so much water that the sea-level Europe, and vice versa, may date back to Cre during the Riss glaciation was lowered to an ex taceous or early Tertiary time (cf. especially tent that ENQUIST, according to oral informa FERN.ALD 1929). The most exotic, southern spe tion, now puts at ·250 m at the very least. If cies within the flora of Greenland, e.g., Sisyrin we could put this figure at 350 m, the lowering ohium angustifolium (cf. GELTING 1934 p. 253) would be just sufficient to allow of the n1igra- and Polygala serpyllifolia, are so completely 80 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
isolated that, apparently, they must be very exchange of biota between North America and old in their Greenlandic stations; GELTING and Europe took place not later than in the early BocHER (op.c. p. 139) believe these species to Quaternary. be >>southern relicts>> that have survived at least .From this interpretation of the amphi-Atlan the last glacial epoch on the spot. tic type of distribution certain corollaries can The fact, that several amphi-Atlantic plants be deduced, as follows: are lacking both in Iceland and Spitzbergen, evi Certain critical forms, e.g., the agamospecies dently constitutes an argument in favour of the within Alchemilla and Taraxacum, the races with early date of the Transatlantic land connexions in Poa arctica and P. laxa s. lat, &c., must be (cf. NORDHAGEN 1935 p. 143 ff.). True, the bo older than generally assumed (cf. N ORDHAGEN tanical exploration of Iceland is very incomplete 1935 p. 17 4). There is, of course, no reason at (cf. LovE 1948 pp. 102 f.), and some such plants all why small morphological differences between may still be found there; or they may have lived two types should be of a recent origin. We know there earlier, only to be rooted out at the many that many types have remained completely un volcano eruptions, that covered a third of Ice changed for ages of time, and this may obviously land with postglacial lava beds and ash layers. also hold good for closely related types. Having However, the amp hi-Atlantic species displaying once become isolated, through sterility barriers a gap in Iceland and Spitzbergen are so numerous or otherwise, they may remain unchanged for that this explanation will not suffice. In addi hundreds of thousands of years, irrespective of tion to the west-arctic plants enumerated by the >>magnitude>> of the taxonomic characters. NoRDHAGEN (l. �.), viz., Deschampsia atropur This evidently applies especially to apomicts. purea, Garex scirpoidea, Arenaria humifusa, Dra The west-arctic, element of the Scandinavian ba crassifolia, Braya linearis, and Rhododendron flora must, as pointed out by N ORDHAGEN ( op.c. lapponicum (the occurrence in Beeren Island is pp. 151 ff. ), have survived at least the maximum very dubious), a few species of the same general (Riss) glaciation in Scandinavia and probably the character, which were not classified as west entire ice-age. This is true even of the sylvan arctic when NoRDHAGEN wrot� his paper, viz., species, i.e. Garex livida, Urtica dioeca ssp. gra Carex aTctogena, Urtica dioeca ssp. gracilis (not cilis, and Galium triflorum (cf. SELANDER 1947 in Iceland, LovE in litt.), Galium triflorum and pp. 273 f.). In all likelihood, the whole of the Antennaria Porsildii, are also missing in these Fennoscandian mountain flora, including seve islands, as well as several European plants ex ral all-regional and subalpine species, has lived tending ·into America or Greenland, e.g. Botry here ever since the Great interglacial and, pos chium multifidum, Rubus idaeus s. str., Polygala sibly, throughout the ice-age. serpyllifolia, Scrophularia nodosa (FERNALD 1933 p. 282), &c. Personally, I am inclined to believe The Alpine-North European element that, at any rate, the amphi-Atlantic linneons whose areas cannot be interpreted as resulting Provided that no, or very few, alpine plants from a reduction of former circumpolar ranges have reached the Scandinavian mountains from got their distribution already in the Tertiary, the south in postglacial time, we can charac when Europe was still united with North Amer terize also those species as glacial survivors that, ica (cf., e.g., RAMSAY 1931 pp. 130 ff.); the outside Fennoscandia, occur, exclusively or main forementioned gaps would be readily explained ly, in the mountains of South and Central Europe by such an assumption, since the present islands with a gap between their southern and northern are but a fraction of those ancient land areas. populations and are lacking, completely or nearly In any case, everything indicates that KuL so, east of Finland and the Kola Peninsula. In CZYNSKI (1924), HANSEN (1930) and NORDHAGEN view of facts specified above (pp. 78 ff. ), I in (1935) were right in assuming that the main clude also plants occurring in Iceland in this Origins of the Flora 81 group, although their Icelandic areas are situa Furthermore, there are a few species with a ted west of the Atlantic. Within the flora of similar total distribution, though their Fenno our region, the following species constitute this scandian areas are not completely isolated from Alpine-North European clement: their Central-European ones, interjacent stations forming connexions which are interrupted only Cryptogramma crispa: rots of S. and Central Europe from Pyrenees to Balkan, Iceland, Fennoscandia, by relatively small gaps or frequency minima. isolated in the Urals between 61o and 65° {HULTEN This applies to Polygonatum verticillatum (HUL 1941 p. 41); TEN op.c. p. 127) and Melandrium rubrum (ibid. Luzula sudetica : rots of S. and Central Europe, p. 191, cf. also M. FRIES 1949 pp. 55 f.; these Fennoscandia {HULTEN 1949 b pp. 54 and 119, GRON two species with connections to the south), Epi TVED 1942 p. 195), Iceland (ibid.); reports from Russia, N. Asia, Kamtchatka, Japan, New Zealand, lobium collinum (ibid. p. 328; connexion to the Sierra Nevada in N. America (ibid.) and W. New east), and possibly a few more. They may obvi foundland (FERNALD 1933 p. 237) not confirmed, ously be postglacial immigrants to Fennoscandia; though the last-mentioned report does not seem however, the chances seem to be that their alpine improbable; populations belong to the glacial survivors. Chamorchis alpina: Alps, Carpathians, Transsyl vania, N. Balkan, Scandinavia (HEGI II p. 362); Leuchorchis albida L. straminea s. str., excl. (Fern.) Areas with a gap east of the Scandes Love: rots of S. and Central Europe (ibid. p. 369), isolated area in S.W. Sweden, then another one in the Since arctic-montane plants only exceptionally, Scandes (HARD AF SEGERSTAD 1924 p. 140, HuLTEN _if at all, could have migrated into Fennoscandia op.c., map 134); reports from the Faroes, Iceland, Greenland and N.E. North America refer to L. by any other route than the eastern one, a suf straminea (LovE 1950 pp. 36 f.); ficiently wide gap in the area of a species east Silene rupestris : rots of S. Europe from Sierra of Scandinavia and Finland, or within Fenno Nevada to Carpathians, Fennoscandia (HuLTEN op.c. scandia east of the Scandes, must be regarded p. 190, LAGERBERG 1940 p. 146); reports from S. Urals as a definite sign of its having survived the la and Altai (ibid.) not confirmed; practically certain to be a glacial survivor, since the larva of one among the test glaciation there. >>hibernating>> macrolepidoptera, the Scandinavian It is rather astonishing that, as far as I have Alpine. Eupithecia undata Frr., lives solely on its fruits been able to make out, no botanists but N ORD (WAHLGREN 1941 I p. 54, NoRDSTROM 1941 p. 278); IIAGEN (1935, 1937) and, first and foremost, HuL Saxifraga Cotyledon : Pyrenees, Alps, Iceland, Scan TEN (1937 a, 1949 b) have laid stress upon the dinavia {GRONTVED op.c. p. 263); Epilobium alsinifolium : Apennines, Alps, Sudetes, plant-historical significance of the absence of cer Carpathians, British Is., Faroes, Iceland, Fenno tain species between the Kola Peninsula (resp. scandia, Arctic Russia (GRONTVED op.c. p. 299); Finnmark or Troms) and the Urals (resp. Sibe Myosotis silvatica ssp. frigida : Spain, Pyrenees, ria) or, in some cases, between Troms and eas Jura, Alps, Fennoscandia (VESTERGREN 1933 pp. ternmost Kola or Kanin. These gaps are most 7 f.); V aleriana sambucifolia : Pyrenees, rots of Central considerable, their width varying from from 500 Europe, N. England, Scotland, Iceland, Denmark, to 1000 km or even more. They cannot have Fennoscandia (LAGERBERG op.c. p. 89, HuLTEN op.c. been bridged over in postglacial times by means p. 405); of accidental long-distance dispersal, apart from Saussurea alpina: Pyrenees, Alps, Carpathians, rare, exceptional cases. And they recur too often Wales, Scotland, Balticum, Fennoscandia, small area in Samoyed Penins.; reports from Asia and and too regularly to be explained as mere chance America refer to other, related types (LAGERBERG & irregularities, such as are sometimes met with HoLMBOE 1940 pp. 240 f.); however, ace. to POHLE in the areas of plants. (1903 p. 49) also in the Pinega and Mesen distr. of That so many species seem to be lacking in Arctic Russia. Arctic Russia east of the Kola Peninsula may Lactuca alpina: rots of S. and Central Europe from Pyrenees to N. Balkan, Scotland, Fennoscandia partly be a delusion, owing to insufficient inves {HEGI VI p. 1100). tigations; this is possibly the case with some 7- 496149 Sten Selander I 82 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark easily overlooked, sylvan or low-alpine, plants, However, the width of the region where, pro e.g., Carex laxa and Pinguicula villosa. bably, the forests used to reach the sea is not A conceivable explanation of the absence of larger than about 400 km, which does not make other species would be that they lived here im it quite unpassable to certain plant species. It mediately after the ice-age, then p�rishing duTing should also be remembered that the cliffs of lime the climatic optimum, when the forest went fur stone and gypsum on the banks of, e.g. , the rivers ther north on the present tundra. Unfortunately, J ula, Pinega and Sotka form habitats where arc I have not been able to inquire into the move tic-montane plants (Tofieldia pusilla, Minuartia, ments of the forest line in these parts. The only verna, Cardaminopsis petraea, Dryas octopetala, modern papers, known to me, where this ques Arctostaphylos alpina, &c.) occupy interjacent tion is discussed are written in Russian and, stations in the woodland (POHLE 1903 pp. 74 ff.). therefore, inaccessible to most foreign readers. Furthermore, the postglacial climatic changes The few available facts seem to indicate, how- . cannot possibly explain the absence of many arc ever, that these movements should not be left tic plants from as comparatively high-arctic re altog�ther out of account. True, the palynolo gions as Kolgujev or the Timan tundra where, gical studies of AARIO (1943) on the development rnoreover, the land was unglaciated during the of forest and climate in the tundra district of Wiirrn glaciation and, consequently, the occur Petsamo (west of the Rybachi Peninsula) do n>Warum hat denn dieses westliche Artenele TEN (1937 b p. 24) tells us, without mentioning ment von der Kolahalbinsel aus nicht weiter his, source, that the forest, at a comparatively nach Russland fortgesetzM Die ostwarts fort late time, had advanced 1-2° to the north of schreitende Kontinentalisierung des Klimas hat the present wood-line on the East-Siberian in Betreff mancher Arten (z. B. Carex marina, tundra; yet even such a big advance would leave C. norvegica, Alchemilla alpina, Myosotis silva a broad strip of tundra intact in eastern Russia. tica ssp. frigida) dazu beitragen konnen, dass ihre However, the position is essentially different west Wanderung ins Stocken geriet. In anderen Fal of the Timan Mts. Even nowadays, the forests len, zumahl in solchen, wo die betreffende Art of the river banks reach the sea coast, e.g., near auch weiter im Osten, in · noch bedeutend kon Mesen (cf., e.g., PoHLE 1903), and the treeless tinentaleren Gebieten als es die nordrussische tundra bogs conceal remains of ancient forests: Festlandtundra ist, Vorkomnisse hat, diinkt die >>Die gauze machtig� Torfschicht war stellenwei Heranziehung der klimatischen Verhaltnisse da se von Wurzeln und diinnen Stammen unter gegen unmoglich. Durch die edaphischen Ver gangener Holzer durchsetzt; es waren ... Picea, haltnisse diirfte die Sache wohl kaum zu erkla Betula, Pyrus aucuparia und Salix-Arten ver ren sein, obwohl z. B. das fast vollige Fehlen von treten. . . Auf Siidkanin lagert iiberhaupt ein Felsenstandorten im Bereich der nordrussischen grosser Theil der Tundramoore gewiss ganz direkt Tundra gewissen Arten Scbwierigkeiten bereitet auf ehemaligem Waldbo,den>> (ibid. pp. 81 ff.). haben kann. Beziiglich der meisten Falle muss There seems to be little doubt that the present man sich, wenigstens vorlaufig, mit der Annahme tundra west of Swjatoi N oss (N. of the Timan begniigen, dass die die Kolahalbinsel von N ord rots), the northern part of the Kanin Peninsula russland trennende Meeresenge auch in ihrem excepted, was forest-clad during the postglacial schmalsten Stadium zu einem Hinderniss fiir das warm period. Vorwartsdrangen der Pflanzen geworden ist.>> , Origins of the Flora 83
KALELA is obviously quite right in dismissing also be of a historical nature. The conditions the climate as sole cause of the gaps, in Russia which the flora of these parts was up against east of the Kola Peninsula, in the ranges of the during the Wiirm glaciation were, in all likeli plants discussed here, even though their migra hood, far from propitious, at any rate to arctic tions must have been obstructed, e.g. , by the and arctic-montane plants. On his map af Europe surprisingly hot summers of the region south of during the last glaciation, ENQUIST (in GusT.AFS the Kanin Peninsula (cf. POBLE 1903 p. 19) where soN 1943 p. 189) has marked out a zone of so thermophilous plants as, e.g., Lychnis flos >>sand regions with inland dunes without vege cuculi, Anemone silvestris, .:A. pate�ts, Ranunculus tation>> east of the ice-border and farther east polyanthemos, Centaurea Scabiosa, Petasites spu wards a >>steppe region>>, which would, obviously, rius, &c., are met with (ibid. pp. 66 ff., 75 f.; render wide tracts east of the ice hihabitable to cf. also STERNER 1922 pp. 319 f.). an arctic-montane or arctic flora. It goes On the other hand, one wonders if KALELA without saying that these assumptions are does not underrate the importance of the edaphic highly speculative and that the presumed zone factors. The scarcity on the North-Russian of >>inland dunes without vegetation>> practically tundra not only of cliffs and bare rocks, but of defies proving as well as disproving, there being other alpine ground formations as well, must no evidence obtainable either from macrofossils always have constituted an effective obstacle or from pollen profiles. Furthermore, ENQUIST's to a dispersal of fastidious alpine plants that hypothesis, which was put forth on a purely are confined to such habitats; the impoverish geological basis, is at variance with certain ment of the alpine flora which can be observed botanical facts. Thus attention should be called on every low, softly undulating mountain must to the highly isolated occurrence of, e.g., Allium be still more marked on the Russian tundra with strictum, Minuartia verna, Potentilla strigosa and its flat ground. Moreover, within the whole the endemic Draba cinerea var. ladogensis in Ka region from the eastern Kola Peninsula to the relia ladogensis, and, in the neighbourhood of the Tschesskaya Bay and the crests of the Timan White Sea, of Draba cinerea, Aster sibiricus, Arte l\its and, more sporadically, in the Petchora misia rupestris, Potentilla multifida (that belongs district, the tundra is, to the far greatest to another race than the Scandinavian plant, HY extent, occupied by peat bogs, where no alpine LANDER 1945 p. 199), Taraxacum glabrum, T. leu plants can live (POI:JLE op.c. p. 80). Nor will canthum (the only North-European representa the soil profile of the remaining parts be favou tive of the group Rhodotricha, nearest stations in rable to the arctic-montane flora, since intensely Altai and Tian-shan, HULTEN 1949b p. 460), the leached, infertile planosols, with surface accu endemics Ranunculus sceleratus ssp. reptabundus mulations of · exhausted minerals and acidic (HULTEN 1947b p. 352) and Helianthemum num peaty material, predominate on the tundra mularium var. arcticum, &c� The presence of expanses. Even in regions with not too unfa these plants, all of which are of an East-European, vourable a climate, e.g., the Kola Peninsula, Siberian or Central-Asiatic origin,. doubtless the Fennoscandian arctic-montane species are indicates the existence, during the Wiirm glacia lacking on the tundra proper, occurring, mainly tion, of a flora close to the ice-border. Yet the or exclusively, in the mountains and along the general character of this flora seems to have been rivers and the coast, where the tundra topo so pronouncedly continental as to suggest a pos graphy is broken and the planosols are less sible connexion with the present >>steppe>> flora predominant (cf. REGEL 1923, pp. 172 ff., &c.). of Oeland and Gotland rather than with the As far as I know, this applies also to their alpine flora of the Scandes (cf. above p. 62; see areas further east. also HIITONEN 1946). However, the causes of the singular voids Also the paleographic data should be remem within the Russian tundra flora may, partly, bered. KALELA's hypothesis as to the important 84 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
place filled by the seastrait between North-Rus crossed by mighty rivers which would also have sia and the Kola Peninsula does not carry much constituted a hindrance to certain Inigrating conviction, considering that far broader waters, plants (HYYPPA op.c. pp. 437 f.). Furthermore, such as the Baltic and the Gulf of Bothnia, have the Karelian Isthmus and adjacent parts seem to evidently in the long run failed to prevent a have been invaded by conifer forests as early as dispersal of a great number of plants; the maps of at the Salpausselka phase of the retreat of the plant areas in, e.g., HuLTEN 1949b convey the inland ice, i.e. about 8 000 B.O. (HYYPPA 1933; in impression that the phyto-geographical import view of the facts brought out by AARIO such ance of the Baltic has been very smalJ indeed. statements should, however, be accepted only Yet one more fact should be kept in mind. As with a certain amount of caution, cf. above p. pointed out by SAURAMO (194 7 p. 79) among 58). As a matter of fact, the late-glacial Dryas others, conditions on the eastern fringe of flora established on the Isthmus did not, as far the Wiirm glaciation during the finiglacial period as we know, advance further north (HYYPPA op.c. present most intricate problems. However, in pp. 10 ff.; cf. also HIITONEN 1946). The only early Goti-glacial time a broad arm of the sea late-glacial migratory tract of real significance to seems probably to have extended from the arctic-subarctic plants of an eastern origin was, Baltic to the White Sea, ancient shore marks, in all likelihood, the Kola Peninsula (cf. KALELA combined with the occurrence of marine relics op. c. pp. 42 ff.); however, it was probably fre among the biota of the lakes and of marine dia quented largely by such eastern species as do toms in deposits of that time, constituting irre not reach further west than Ponoj or the futable evidence of a great marine transgression Murman Coast. (SAURAMO op.c. pp. 81 ff.). And when the sea The present author has no personal experience began to subside, the amelioration of the cli of tundras, save those of northernmost Sweden, mate, as registered by the pollen profiles (ibid. which in many respects differ from the Russian pp. 106 f., fig. 9, 10 and 11), had proceeded so ones. Consequently, nothing but mere conjec far that the pine forests probably raised riearly tures can be offered here. Everything seems as effective a barrier to arctic plants as the taiga to indicate, however, that the North-Russian of to-day. True, the results of SAUR.AMO were flora during and shortly after the Wiirm glaciation submitted to trenchant criticisms by several Swe was at least as poor in arctic and arctic-mon.tane dish geologists (cf. Geol. Foren. Forh., Bd 69 species as nowadays, when the vegetation of 1947 pp. 293 ff.). What is relevant in this con the tundras comprises only about 300 vascular nexion, viz., the existence and the date of the plants, many of which are, moreover, of no marine transgression, seemed, however, hardly interest in this connexion, since they are lacking to be doubted. This sea strait must, to a con west of the Kola Peninsula. We have no reason siderable degree, have obstructed the dispersal to believe that, after the Great interglacial, of hypothetical arctic immigrants, at any rate Arctic Russia has ever harboured a flora, notably temporarily. Moreover, the Karelian land east richer than the present one in such plants as of the inland ice seems, irrespective of such con could have migrated into our mountains. siderations, hardly to have been a migratory Thus all or nearly all arctic-montane or tract of any significance to plants of this kind. arctic species occurring in Fennoscandia, though Even during the late fini-glacial period, about absent from northern Russia east of the Kola 7 000-8 000 B.O., it probably formed but a Peninsula, are probably glacial survivors. Their narrow land -strip between the White Sea, then number is rather difficult to detail, since no far larger than now, and the Baltic Ice-lake or modern Flora of Russia offers specifications the Yoldia Sea (cf. HYYPPA 1936, SAURAMO1942 ; of the plant ranges. The bulk of available knowl see also the map by SAURAMO in LINDROTH 1949 edge of the plants occurring in northernmost p. 727); its northernmost part was, temporarily, Russia is still contained in the old works of Origins of the Flora 85
RUPRECHT (1845, 1856), TRAUTVETTER (1849- p. 305); ace. to Flora URSS 1935 p. 381 also one 50), KORSHINSKY (1898) and POHLE (1903, 1917). station on S. coast of White Sea; Carex tenuiflora : not between W. Kola Penins., Practically no later works have been published L. Onega, and Urals (HULTEN 1949 b p. 89); in any other language than Russian. Carex bicolor : between Porsanger in Finnmark The new Flora URSS apparently sometimes (DAHL op.c. p. 264) and Obi R. one single station, includes the Kanin Peninsula and Kolgujev in the Pinega distr. near Archangelsk (HULTEN 1942 in the general term of >>Arctic Europe>>, and its p. 332); statements as to the distribution of the species Carex atrojusca : gap between Kola Penins. and N. Urals (ibid. p. 373); are, on the whole, most summary. Floristic Luzula arctica : lacking between Troms. and U rals manuals, giving total ranges of the species, would (HULTEN 1943 p. 439); often seem to report a plant as occurring in Coeloglossum viride ssp. islandicum : not reported >>Arctic Russia>>, when its actual area is restricted between Kanin Penins. and Chukch Penins (HuLTEN to the Kola Peninsula or the Urals. With these 1943 p. 473; cf. also this work, Part II, p. 10); Salix reticulata : gap between Kola Penins. and Urals qualifications, and apart from the species (HULTEN 1943 p. 507, Flora URSS 1937 p. 32); gap whose distribution in these parts is previously not mentioned in HuLTEN 1949 b; reported, the following vascular plants, belonging Salix polaris : lacking between Hibina and Kolgujev, to the flora of our region, would seem to display gap c. 550 km; on the mainland not W. of Urals gaps in their ranges east of their Fennoscandian (HULTEN 1949 b p. 147); Salix arbuscula: gap of c. 700 km from Lakse fj ord areas: to Kanin Penins. (ibid. p. 142); Asplenium viride : missing between W. Kola Penins. Betula callosa: not known between Kola Penins. and L. Onega on one side, Mid-Russia and N. Urals and Yenisei (LINDQUIST 1945 p. 176); on the other (HuLTEN 1949 b p. ll); gap at least M inuar·tia rubella : lacking between Petsamo distr. 700 . km; and >> Arctic Russia>> (HuLTEN 1944 p. 686, cfr. 1949 b Asplenium septentrionale : not between Ladoga p. 181, map 712; not reported from the Russian distr. and S. Urals (ibid.); mainland in Flora URSS 1938 p. 506); gap at least Woodsia ilvensis : lacking between E. Kola Penins. 500 km; and Central Urals (ibid. p. 13); .1l1inuartia stricta : gap of c. 500 km from Varanger Woodsia glabella : gap between Petsamo and Kuu to Swjatoj Noss, then another gap of c. 900 km from samo distr. and Urals (HuLTEN 1941 p. 14); Ponoj to Urals (HuLTEN op.c. p. 687, 1949 bp. 182); Alopecurus pmtensis ssp. alpestris : Between Varan Thalictrum alpinum : between Hibina and Urals ger Penins. (DAHL 1934 p. 244] and Kanin Penins, N ova Zembla, 1 000 km to the E., only two stations, (ANDREEV 1931 p. 156) only Imandra (HJELT 1888 at Ponoj and on Kanin Penins.; largest gap c. 600 km p. 355); largest gap c. 450 ·km; (ibid. p. 198); Calamagrostis lapponica : gap between SW. shore Aconitum septentrionale : lacking between Lule of White Sea, L. Ladoga and Central Urals (HuLTEN Lappmark and Vasterbotten on one side, Gridina in · 1949 b p. 41); Lapponia Keretinensis (HJELT 1906 p. 249) and Kare Phippsia algida : gap of c. 350 km from E. Kola lia Ladogensis on the other (map in M. FRIES 1949 Penins. to Kolgujev, then a new gap of c. 300 km to p. 62); gap c. 500 km; Nova Zembla and Waigatch (ibid. p. 57); Ranunculus nivalis : between Rybachi Penins. and Roegneria latiglumis s. lat.: between Petsamo and Urals only on Kanin Penins. and Kolgujev (HULTEN Kuusamo distr. and N. Urals one station in E. Kola 1949 b p. 210); largest gap c. 500 km;
Penins. (ibid. p. 68); Draba lactea : lacking between K vaenangen (N ORD · Kobresia myosuroides : E. of Berlevaag in Finn HAGEN 1940 p. 256), Kilpisjarvi in Lapponia Enon mark (DAHL 1934 p. 274) and Rybachi Penins. (HuL tekensis (HJELT op.c. p. 338) and Urals (HuLTEN TEN 1949 b, map 314) not until Chatanga distr. of 1945 a p. 857, Flora URSS 1940 p. 403); Siberia (HuLT:EJN 1942 p. 296); Draba nivalis : E. of Rybachi Penins. gap of about Carex capitata : E. of Kola Penins. and Vologda 600 km to Kolgujev (HuLTEN 1949 b p. 231); then not nearer than Siberia (SMITH 1940 p. 199); ace. to very rare on W aigatch and along the Siberian coast Flora URSS (1935 p. 186) in the Urals too; gap not (HULTEN 1945 a p. 862; map, partly. incorrect, in mentioned in HuLTEN 1949 b; FERNALD 1925 p. 339); Carex rupestris : gap between Kola Penins. and N. Draba alpina : between N ordreisa (NORDHAGEN Urals, Vaigatch and Nova Zembla (HULTEN 1942 op.c. p. 256) and Kanin Penins., 900 km to the E., one 86 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark single station in N.E. Kola Penins. (HULTEN 1949 b display the following plants: Eriophorum medium p. 229); largest gap c. 700 km; (Sydvaranger-Liza, c. 300 km), Juncus arcticus Oardamine bellidijolia : gap between Hibina and Sagina Urals (HULTEN 1945 a p. 833); (Rybachi Penins.-Nokujev, c. 300 km), Saxijraga oppositijolia : gap between E. Kola intermedia (Gawrilowo-Ponoj, c. 300 km), Saxi Penins. and N. Urals, Waigatch and Nova Zembla fraga aizoides (Rybachi Penins.-Hibina and (Huv.r:EN 1949 b p. 248); Swjatoi Noss, c. 200 km), and S. foliolosa (be Potentilla nivea s. str.: gap between Hibina and S. tween Paatsjoki and Ponoj, c. 550 km, only at Urals (HULTEN 1945 b, map p. 143); Imandra; largest gap c. 400 km). The last Potentilla hypa1·ctica: missing between W. Lule Lappmark and Kanin Penins. (SELANDER 1942 p. 71); mentioned gap is probably due to historical gap c. 1.100 km; causes; the remaining gaps may be without Astragalus alpinus s. str., excl. ssp. arcticus (Bunge) plant-geographical significance, to be explained Hult.: probably lacking between Kuusamo distr. and Si as resulting from insufficient investigation. beria (HuLTEN 1947 a p. 1084, cf. 1949 b, map p. 292); Very rare in Russia E. of the Kola Peninsula Astragalus norvegicus : gap of c. 600 km between Varanger and Lapponia Ponojensis (ibid. p. 293); are, finally, Carex rariflora (HULTEN 1942 p. 367), Oxytropis lapponica : lacking between Vestfinn Koenigia islandica (HULTEN 1944 p. 595), Minu mark (DAHL 1934 p. 360), Kilpisjarvi in Lapponia artia biflora (ibid. p. 680), Saxifraga nivalis Enontekensis (HJELT 1911 p. 237), and Tianshan in (HULTEN 1945a p. 924), and Diapensia lapponica Central Asia (HULTEN 1947 a p. 1097); (HULTEN 1948 p. 1266), which are reported only Epilobium anagallidijolium: gap between Kola Penins. and Urals (HULTEN 1929 p. 142, 1947 a p. from the Kanin Penins. and Kolgujev. 1143; gap not mentioned in HuLTEN 1949 b); Oassiope .tetragona : gap of c. 1.100 km between Bicentric species Urals and Hibina, then a new gap of c. 400 km between Hibina and Porsanger (HULTEN 1948 p. 1223, 1949 b Ever since TH. FRIES (1913 p. 318 ff.) made p. 352); clear the significance of the bicentric type of Gentianella tenella : missing between Alta (DAHL op.c. p. 394) and E. Kola Penins. (HULTEN 1948 p. distribution within the Scandinavian mountain 122$, 1949 b p. 367); gap c. 700 km; flora, i.e. a distribution consisting of two separate Hackelia dejlexa : gap between Karelia Ladogensis areas, a northern one, in most cases situated and Central Russia (ibid. p. 375; cf. also HoLMBOE between Vastfinnmark or Troms and Pite 1937 p. 18); gap c. 600 km; lappmark, and a southern one, whose centre Pedicularis hirsuta : lacking between Jesjavvre in Finnmark (DAHL op.c. p. 394) and Kolgujev (HuL generally lies in the South-Norwegian mountain TEN op.c. p. 407); gap c. 950 km; from Kola Penins. ranges of Dovre and Lom, such a distribution there are, ace. to HJELT (1923 pp. 86 f.), two old re has been regarded as a definite sign of glacial ports, one of which is obviously wrong and the other )>_hibernation>>. The pronouncedly bicentric spe highly dubious; cies of our region are the following ones (areas Pinguicula villosa : not reported between Kola Penins. and Obi Bay (HuLTEN 1949 b p. 412); cf., in this section according to HULTEN 1949b ): however, above p. 82; Poa arctica Minuartia rubella Erigeron unijlorum: lacking between Varanger P. herfedalica Melandrium apetalum and Kolgujev (ibid. p. 431); gap c. 800 km; Oarex parallela Ranunculus hyperboreus Antennciria carpathica : gap of c. 700 km between 0. arctogena Draba hirta Varanger and Kolgujev (ibid. p. 432); C. bicolor D1·. lactea Hieracium kuusamoense: absent between Fenno Luzula parvijlora Potentilla nivea scandia and Transbaikal (SAMUELSSON 1943 p. 26). L. arctica Rhododendron lapponicum Equisetum variegatum seems also to be missing Salix glandulifera Euphrasia lapponica between the E. Kola Peninsula and East-Russia Oerastium arcticum Oampanula uniflora (Perm); at any rate, I have not seen any reports from Sagina caespitosa Ta1·axacum brachyceras the interjacent region. Salix glandulifera has not previously been Smaller gaps in North-Finland or on the Kola included in this group, although doubtless Peninsula (cf. the maps in HULTEN 1949b) belonging here, since it does not occur between Origins of the Flora 87
K vikkjokk and Meldal in S0r-Tnmdelag (FLo parallela, Minuartia 1·ubella, Draba nivalis, Dr. DERUS 1931 p. 128, HULTEN op.c. map 572). lactea, and Gampanula uniflora in southern Less pronouncedly bicentric, with a gap in Lapland (RUNE 1945b, 1947, and 1948) consid northern Jamtland andjor South-Lapland and erably diminish the disjunctions within their the adjacent parts of Norway, less wide than areas; closer investigation may, therefore, render in the species just mentioned, are the following the bicentric distribution less striking also in plants: other .instances. .As a rule, the occcurrenc of a gap within Deschampsia atropurpu- Koenigia islandica rea Minuartia stricta the areas of the said species can, nevertheless, Phippsia algida Ranunculus nivalis not be doubted. These gaps cannot result from Kobresia myosuroides Draba nivalis the nature of the bedrock, since many strongly Oarex microglochin Dr. alpina basophilous plants, e.g., Asplenium viride, Gysto C. rariflora Angelia Archangelica ssp. pteris montana, Garex rupestris, Dryas octopetala, 0. glacialis norvegica Juncus arcticus Primula scandinavica &c., are evenly distributed all along the n1ountain Ohamorchis alpina Gentianella tenella range. Nor can I persuade myself to believe Salix polaris Pinguicula alpina that in all, or even any considerable number of, instances the gaps are the outcome of a Part of these plants were set apart already havoc, supposed to have been brought, just by TH. FRIES (Le.) as the )>Pinguicula alpina in these parts, among hypothetic, evenly distri group)>, e.g., Draba nivalis and Pinguicula alpina; buted populations of alpine plants by the post others, viz., Deschampsia atropurp�lrea, Koenigia, glacial climatic amelioration. and Angelica Archangelica, were noticed by First of all, several markedly alpine species, · RUNE (1945a pp. 8 ff.); the rest could not be e.g., Deschampsia alpina, Poa laxa ssp. flex�wsa, distinguished until HuLTEN's invaluable carto Garex Lachenalii, G. rufina, Ranunc·ulus glacialis, graphic material was compiled. R. pygmaeus, and Gardamine bellidifolia, occur In other cases, the gaps are reduced to marked more or less frequently within the gap, though minima of frequency. Such instances are the they seldom or never descend below the tree following species: line. Why should they have escaped the effects of the warm period that, presumptively, proved Trisetum spicatum Saxifraga 1·ivularis Oarex atrofusca Astragalus norvegicus fatal to the bicentric plants? Secondly, it seems 0. saxatilis Oxytropis lapponica incomprehensible that species occurring nor Juncus biglumis Primula stricta mally in the conifer belt, e�g. , Luzula parvi Salix reticulata Veronica fruticans flora, exhaustively dealt with by BJORKMAN Sagina intermedia Pedicularis lapponica (1939 pp. 205 ff.) who rightly stresses its theo Minuartia biflora Erigeron boreale Draba norvegica E. uniflorum retical significance, Salix glandulifera, Ranun Saxifraga cernua Antennaria alpina culus hyperboreus, Myricaria germanica, &c., should have been exterminated in parts of their Especially .ARwrnssoN (1943 pp. 105 ff.) areas during .the warm period by the raising of has tried to minimize the importance of the the tree limit to a higher level and the disap bicentric type of distribution for the theory of pearance of glaciers and snow patches. Nor glacial survival, pointing out that most bicentric could the warm period have such an effect on plants are basophytes, thus shunning the areas plants that are nowadays characteristic of the of acidic magma rocks, and that many alpine limestone hills of N ordland, growing almost species may have disappeared during the post down to the sea level, e.g., Euphrasia lapponica, glacial climatic optimum from the region of low Garex glacialis and Primula scandinavica. This mountains north of Storlien and the Trondheim is true also of a bicentric littoral plant like Poly Fiord. Furthermore, RUNE's findings of Garex gonum Raii ssp. norvegicum (cf. HULTEN 1949b 88 SELANDER: Floristio Phytogeography of South-Western Lule Lappmark map 642) which is, also for other reasons, an nica, which are more or less tolerant of humic obvious glacial survivor, and of several bicentric acids and normally occur in the conifer belt. We coleoptera as established by LINDROTH (1939), must find other interpretations of the gaps e.g., Bembidion Sibkei J. Miill. (op.c., map fig. 12) within their areas �or the frequency minima, loca Barynotus squarrosus Germ. (ibid., map fig. 14), ted to the gap between the areas of the bicentric and Othiorrhynchus arotious 0. Fabr. (ibid., map species and equated by RUNE [1945 a p. 10], ap fig. 15), all of which are found regularly below parently for excellent reaBons, to the bicentricity the wood limit (ibid. p. 253). Later he pointed of other plants). out that no fewer than 12 more or less pro ARWIDSSON himself emphazises (1943 p. 105) nouncedly bicentric carabids (i.e. all such cara the fact that the bicentric type of distribution bids but N ebria nivalis Payk.) occur normally displays every phase of transition from extreme in the woodland, four of which (viz., in addition cases, such as Stellaria crassipes and Saxifraga to Bembidion Sibkei, Cicindela maritima Dej., hieraoiifolia, to barely discernible vestiges, as in Dysohirius angustatus Ahr. and D. Helleni J. the ranges of Salix retioulata or Pedioularis lappo Miill.) have never been found in the regio alpina nica. A more or less pronounced tendency to (LINDROTH 1949 pp. 440 ff., 752 ff.). Lastly, a bicentricity (or, in some cases, pluricentricity) can number of plants occur in those parts where the even be said to be a pervading characteristic of >>normal>> bicentric species are missing, but are the major part of our alpine flora. And this can absent from certain regions either further north, scarcely be due to such a local phenomenon as Juncus castaneus, Nigritella nigra, Ra the consequences of the warm period within the e.g., and , nunculus platanifolius, or further south, e.g. Are Central Scandinavian region of low mountains, naria norvegioa, Arabis alpina, Saxifraga foliolosa, since the gradual changes in the composition of and S. groenlandica. These gaps cannot have been the flora reach far outside this region. On the caused by the warm period, as they are mostly other hand, distributions similar to those actu situated in high-alpine tracts, but are probably ally existing were to be expected, supposing the to be explained from the absence, or scarcity, increased number of species within certain areas of these plants in the neighbouring ice-age to be due to the nearness of an ice-age refugium refugia. from which plants have migrated a longer or I do not by any means wish to suggest that shorter distance according to their spreading ca SMITH (1920 p. 142) was wrong in stating that pacity, adaptibility, and competitive power. the amelioration of the climate diminished the That many species are lacking just within the areas of the alpine plants, especially the baso Central Scandinavian gap may at least partly philous ones. In the case of most species, this de be due to the fact that, evidently, no ice-age crease would seem to have been even more lasting refugium was situated in these parts (cf., e.g., than he assumed. When the wood limit was again HOLMBOE 1937 pp. 27 f. and EILIF DAHL 1946 lowered, at the end of the warm period, the pp. 230 ff.), and that after the melt.ing away of changes in the soil profile brought about by the the inland ice many spreading alpine species here forest vegetation, i.e. increased acidity of the met with an impassable barrier, viz., the pro soils, effected by the acidic litter of the conifers, nouncedly oceanic local climate and the resul and probably intensified podsolization (cf. e.g. tant soil types. The occurrence in Jamtland &c. DAUEENMIRE 1947 p. 49), would remain and of oceanic plant species such as Narthecium ossi cause large areas, originally inhabited by more fragum, Pedi0ularis silvatica, and Juncus squarro fastidious alpine plants, to be occupied by sus (cf. LANGE 1938), which never appear in the trivial low-alpine scrub heaths. company of fastidious alpine plants, hint, per Yet, these changes could hardly affect the haps, at the climatic causes for the absence of distribution of species such as for instance De 1nany such plants. It is also worth noticing that schampsia atropurpurea and Pedicularis lappo- a number of not too trivial, more or less evenly Origins of the Flora 89
distributed alpine species, e.g., Cryptogramma Calamagrostis neglecta ssp. Tofieldia pusilla crispa, Asplenium viride, Poa laxa ssp. flexuosa, borealis Rubus Chamaemorus Luzula Wahlenbergii Campanula rotundifolia and Carex rufina, represent an oceanic, western type of distribution. The poorness of the alpine Furthermore, Lycopodium Selago, Poa glauca, flora within the Central Scandinavian gap would and Juncus triglumis, which are also missing in in other words be a phenomenon, parallel to the northern Greenland and Liitke Land north of floral impoverishment of the outer coastal moun J\tlatotchkin Shar (or in all Nova Zembla; cf. tains of northern Norway. B6CHER 1938 and LYNGE 1923), probably migra ted to Spitzbergen from Fennoscandia at some A causal nexus between hibernation and bi time or other. They would therefore seem to be centricity gains in probability by the fact that long to the Scandinavian survivors, although four fifths of the pronouncedly bicentric species, their relict character in Spitzbergen is not empha also for other reasons, can be supposed to be sized by sterility. glacial survivors. That bicentricity certainly is Among the European plants occurring in Ice anterior to the Postglacial climatic optimum is land, those absent from Greenland and America also suggested by the differentiation of several probably immigrated to Iceland during the ice Poa arctica, Stellaria crassi bicentric species, e.g. age; if they had arrived in Iceland already in pes, Draba hirta, and into separate races within the remote times when a land connexion with their two Scandinavian areas. At least in two Greenland was in existence, they would, in all instances, there is also paleontological evidence likelihood, occur also in Greenland or north-east that bicentric organisms are interglacial sur ern North America. Most such plants probably vivors. According to LINDROTH (in SUNDIUS & hibernated in the British Isles south of the ice SANDEGREN 1948 p. 30, LINDROTH 1949 pp. 738 border. However, this can hardly have been the Simploca ia metallica ff.), the coleopter � Sturm case with the species lacking in the British Isles. obviously used to be very frequent in northern Those species would rather seem to have belonged Sweden during interglacial time, because it has to the ice-age flora of Scandinavia and to have been found there in all the four places where migrated from there to Iceland. Within the flora insectiferous interglacial deposits have been met of Lule Lappmark, this holds good of the follow with; nowadays, it is extremely rare, living only ing plants, among others: in the mountains and showing a markedly bi Erysimum hieraciijolium (GRONTVED 1942 p. 252), centric distribution (LINDROTH 1939, map fig. Viola epipsila (HuLTEN 1947 a p. 1134; in Amer 13, 1949, map p. 739). In all essentials, the same ica only ssp. repens [Turcz.] W. Bckr.), is true of another coleopter, Elaphrus lapponicus Epilobium collinum (GRoNTVED op.c. p. 300), Gyll. (LINDROTH op.c. p. 741). Vaccinium microcarpum (HuLTEN 1948 p. 1251; lack ing in Greenland, Labrador and E. North America). Salix lanata (GR6NTVED op.c. p. 204), which is absent from the mountains of Central Europe, should in all likelihood belong to the same category, though Scandinavian plants in Spitzbergen, Ice it occurs in Scotland. Nor does Betula callosa reach land and Greenland further west than to Iceland (LINDQUIST 1945 p. 176). Those plants which, apparently, migrated to Nor does it seem improbable that a similar Spitzbergen from Fennoscandia at the end of the history in Iceland and Fennoscandia could be Wiirm glaciation or, at any rate, during a late ascribed to plants that are missing in Greenland phase of the Quaternary (cf. above pp. 73 f.), and America, are more or less frequent in the should evidently belong to the Fennoscandian Scandes, and occur mainly in the mountains of glacial survivors. Among these species, the fol southern Europe and in the northern parts of the lowing could not, in the preceding chapters, be British Isles. This is the case with, e.g., the fol characterized as hibernators: lowing species (cf. GRONTVED 1942): 90 SELANDER: Floristic Phytogeography of South-Western Li(;le Lappmark
Salix phylicifolia Trientalis europaea More doubtfull still is the history of the >>east Angelica silvestris Aiuga pyramidalis arctic>> species which occur in the mountains of Vaccinium Myrtillu,s M elampyrum silvaticum South and Central Europe and in Scotland and V. Vitis-idaea (in Amer Cirsium heterophyllum ica only ssp. minus) Crepis paludosa northern England. They may, of course, have reached their Transatlantic areas from the Alps According to A. E. PORSILD (1947 p. 184), via the British Isles and Iceland, although this also Dryas octopetala belongs to this category: seems rather improbable especially with regard in America it occurs but in Alaska and Yukon, to the alpine plants. Among species not previ E. to the Mackenzie delta, and is absent from ously established as >>hibernators>>, the following Greenland, where only Dr. punctata Juz. (ibid. belong to this category: p. 185) and Dr. integrifolia M. Vahl (ibid. p. Rumex Acetosa ssp. ari Alchemilla filicaulis 188) are present. folius Geranium silvaticum .As to the history of those >>east-arctic>> spe Sedum Rosea, the Eur Leontodon autumnale var . asiatic type asperior (probably) cies that must have migrated from Europe to Rubus saxatilis Greenland and America, probably during the Late-Tertiary or early Quaternary, we can evi Still less can be said about the >>east-arctic>> dently but exceptionally come to a valid · con plants that occur also in southern England, viz., elusion. The refugia in the neighbourhood of the the following: Gulf of St Lawrence were certainly larger and Deschampsia flexuosa Ranunculus acris more hospitable than the Scandina.vian ones. Nardus stricta Sedum acre The fact that a plant hibernated in Newfound Rumex tenuifolius land &c. is, therefore, no criterion of its being able to do so likewise in Scandinavia. Still, the On the whole, it goes without saying that all refugia of Greenland and northern Labrador conclusions as to the European history of the hardly offered better conditions than those on >> east-arctic>> plants is highly conjectural, owing, this side the Atlantic (except at the hot springs i.a., to the great changes in the conditions and of Greenland). If an >>east-arctic>> species with distributions of the plants after the early Quater · stations in Greenland and/or northern Labrador nary. is lacking in Central and South J� urope and, Finally, should the Fennoscandian area of a therefore, must have reached their Transatlan plant suggest that it has survived a glaciation tic stations directly or indirectly. from Scandi here, this assumption is strengthened by the navia, it obviously belonged to the preglacial or fact that it occurs in isolated areas in Greenland interglacial flora of Fennoscandia. In such cases, and, therefore, probably is a survivor there. Ac the fact that an alpine or all-regional plant sur cording to GELTING (1934 pp. 264 ff.), this is true, vived one or more glaciations in Greenland and for i.a., of Carex vaginata and Pedicularis lapponica. northern Labrador may, consequently, be re garded as an indication of its having hibernated Remaining, probable or possible, survi also in Scandinavia. In the flora of south-west vors ern Lule Lappmark, this holds good of the fol lowing species that in the preceding chapters Some alpine species within the flora of our have either not at all, or but for inadequate rea region have not, or only with hesitation, been sons, been designated as glacial survivors: assignable to the above groups, which, by the way, have never been supposed to be either his Anthoxanthum alpinum Luzula multiflora ssp. fri torically homogeneous or representative of cat (cf. LovE & LovE 1948 gida egories actually delimited in Nature, but were p. 105) Betula tortuosa Carex rariflora Angelica A rchangelica arranged solely for practical purposes. C. saxatilis ss p. norvegica Part of these species can, nevertheless, with Origins of the Flora 91
a certain plausibility be referred to the inter is circumpolar. It goes far north in, for instance, glacial element of our flora. In view of their Greenland (BocHER 1938 p. 55) and Alaska, and occu pies isolated areas in the Central-European mts total ranges, HULTEN (1937 a pp. 62, 91, and 98) (HULTEN 1941 p. 15). considers that this applies, i.a., to some plants Oarex ornithopoda (HULTEN 1949 b, map 370) is which I enumerate below; for the argumentation, altogether absent from the N orth-German lowland, see the work cited: Denmark, southernmost Sweden and the Finnish mainland. Its main Fennoscandian area is situated Roegneria mutabilis Salix myrsinites in the southern and central parts of the Scandes with J uncus biglumis S. hastata offshoots, e.g., in the silurian area of Jamtland. In Luzula arcuata s. str. Pedicularis lapponica addition, it occurs in small, isolated areas in the One or two among these species have already interior of South-Sweden, Oeland, Gotland (very abundantly), and Balticum. Its total range com been classified as probable survivors, though for prises South and Central Europe, Asia Minor, and conspicuously insufficient reasons. the Urals (HEGI II p. 101). The area in Fenno Some circumpola:r or widely distributed Eura scandia is completely isolated. siatic species possess more or less isolated areas Alchemilla jilicaulis (HULTEN, op.c., map 1041), in the Scandes, where they are frequent, while earlier classified as >>amphi-atlantic>>, seems also to be isolated in Fennoscandia. Apart from a few missing or rare in the neighbouring woodland, stations near Skagen and Limfj ord, it is absent from viz., the following: Jutland; in Germany it occurs in the mountains and Hierochloe odorata Stellaria nemorum ssp. otherwise only in Pommerania (SAMUELSSON 1943
Milium effusum · montana p. 20), and it is absent from Russia, occuring only in Carex capillaris Tussilago farfara Balticum and Finland and in a single station at Tambow, where it may be introduced (ibid.). In and probably Pinguicula vulgaris. Fennoscandia, it centres in JVIid-Sweden, extending In most cases this fact is evident from the maps as a common plant into south-western Finland and, in the mountains,right up to the North Cape. in HULTEN 1949 b. The map of Hierochloe (op.c. p. 35) seems rather misleading, at any rate as Isolated Fennoscandian ranges of essentially far as my experience goes. This plant being eu the same type also characterize some species trophic and somewhat calciphilous, it is probably which, owing to their extra-Fennoscandian dis rare in the oligotrophic area of primary rock of, tribution, were supposed to be glacial survivors e.g., eastern Lule and Torne Lappmark, though in Scandinavia, viz., Sedum annuum (HULTEN the map indicates its being frequent in these op.c., map 946), Saxifraga adscendens (ibid., map parts. In all likelihood, a less summary map 956), Potentilla Grantzii (ibid., map 1015), Ajuga ping would reveal its area in the Scandes as pyramidalis (ibid., map 1076) and Grepis palu being almost isolated, except, perhaps, to the dosa (ibid., map 1820). It seems most likely north-east. Tussilago hardly occurs in the wood that all those species are of an interglacial origin land, except as an apophyte. The montane area in Scandinavia, in spite of their occurring also where it is indigenous appears to be isolated or (or, in some cases, mainly) in the South-Scandi nearly so. navian lowland. Then there are a few lowland plants occurring Epipactis atroru · More dubious is the status of in the mountains whose Fennoscandian areas bens (ibid., map 543). It is missing in the North on closer inspection reveal very interesting fea German lowland and Denmark but for a few sta tures: tions in Germany on the coast of the Baltic and Woodsia ilvensis (HULTEN 1949 b, map 52) is lacking one station in Denmark, and it is altogether ab in Germany north of the Harz (HEGI I p. 13), Den sent from southern and south-western Sweden, mark, southern Scania, Balticum and Russia west of while occurring in a zone from Karelia onegen the Urals. Its isolated Fennoscandian area centres sis and Balticum through Mid-Sweden to the in South-Norway, Mid-Sweden and southern Fin land. To the north, where it occurs · mainly in the South-Norwegian mountains; further north, it mountains, it reaches Nordvaranger. Its total range is rather frequent in the coastal n1ountains from 92 SELANDER: Floristic Phytogeography of South-Western Lule Lappma.rk
Helgeland to the North Cape, and has a small, Central Asia. Therefore, its North-Fenno isolated area in the Kuusamo region of north scandian isolated populations probably ought eastern Finland. Its total range comprises Eu to be designated as glacial survivors, as sug rope, the Caucasus and northern Persia (HEGI gested by BJORKMAN (1939 pp. 218 ff.). II p. 377). The Fennoscandian area is rather si Ribes spicatum ssp. scandicum is reported only milar to that of Epilobium collinum (HULTEN from Fennoscandia (mainly the northern parts), op.c., map 1278), earlier classified as a probable Lithuania and Poland (HEDLUND 1948 p. 45). glacial survivor; even though these two species If the reports from the last-named countries may have obtained their South-Scandinavian are correct, its range is one of the most peculiar populations, partly or entirely, from the south within our flora, defying interpretation with ·east in postglacial times, the montane popula even a semblance of probability; if not, it is tions may have >>hibernated>> in Scandinavia. evidently a glacial survivor. A certain resemblance to the ranges discussed The area of Salix lapponum (as well as that above characterizes those of Asplenium sep of S. myrsinites) extends from Great Britain tentrionale (HULTEN 1949b, map 41), which is to western Siberia, its centrum being Fenno absent from Russia and Denmark and, conse scandia, where these two species may possibly quently, has an isolated Fennoscandian area have arisen. resembling that of Epipactis atrorubens, though Many all-regional species are represented by Asplenium is much rarer to the north, and morphologically distinct alpin� races in the Oarex flava (ibid., map 402), which is frequent Scandes. At closer inspection, some of them in southern Norway and central Sweden, on will, perhaps, be reduced to mere modifications; the one side, and Balticum and Russia, on the while another, and probably greater, part of <>ther, though being of very scattered occur them will, no doubt, be raised to a higher taxo rence in the broad interjacent belt. However, nomical rank than forma or varietas. It is I do not venture to express an opinion as to hardly possible, at present to obtain an ade the significance of these observations. quate idea of their distribution. However, in Also the Fennoscandian distribution of Viola most cases their areas would seem to resemble rupestris (ibid., map 1263) seems strange. It those of the arctic or arctic-montane species. occurs in southern Finland, southern and central Considering that the time passed since the latest Sweden (but not in Denmark), and south-eastern glaciation would seem to be too short for the Norway. In this area it appears to be of an differentiation of such a large number of alpine eastern, postglacial provenance. In addition, races, occurring within the majority of all it occupies two separate areas further north, one regional species, these types may be regarded in north-western Norw:ay and northernmost as possible glacial survivors. Some of them Sweden, and another in the Kuusamo district have already been mentioned in connexion with and Kola Peninsula; as well there as in South the hibernating alpine species. Other instances N orway, it ascends above the tree limit. Out are the following: side of Fennoscandia, it has a Eurasiatic, much split-up range of an obviously ancient type Lycopodium Selago (v. Carex canescens (v. robus appressum Desv.) tior) (HULTEN 1937a p. 117, STERNER 1922 p. 401); L. annotinum (v. alpestre) C. aquatilis ( v. stans Drej .) in England it occurs as a glacial relict (WILMOTT Dryopteris austriaca (un Polygonum viviparum (v. 1935 p. 220). On the whole, it cannot be char named alpine race, cf. alpinum vV g) acterized as a real lowland plant, being mar WEIMARCK 1 945 p.470) Melandrium rubrum (>>oe kedly montane in Central Europe, with the Poa nemoralis (v. mon- cotypus alpinus>>, cf. tana Gaud.) TURESSON 1925 p. 227) majority of its stations there situated in the . Roegneria canina (v. lap Calthg, palustris (v. radi pine forests of the Alps, and ascending up to ponica [Laest. ]) cans Fr., ·cf. HuLTEN 3000 m s. m. in the Alps and 4000 m s. m. in Festuca rubra (v. mutica) 1949 a pp. 389 f.) Origins of the Flora 93
Ranunculus acris (v. pu Epilobium palustre (v. an phytogeographical data. On the contrary, the milus, &c.) gustum Hartm.) hardiness and adaptability that explain their R. repens (ssp. fistulosus Trientalis europaea (f. io ubiquity in the Arctic, obviously render them Rosend.) sea Neum.) Pa'rnassia palustris (v. te Carnpanula rotundifolia particularly fit for the strain of a hibernation. m�,is W g; cf. ERLANDS (seemingly several ra The same can be said about such all-regional SON 1942b) ces, as yet not cleared plants as, in the regio alpina, nearly reach the Geum rivale (v. subalpi up) regio alpina superior, and whose circumpolar num Nemn., cf. Part II Solidago Virgaurea (v. areas include the fore-mentioned high-arctic p. 14) alpestris Blytt) Lotus corniculatus (v. bo Leontodon autumnalis (v. regions, their hardiness matching that of the realis) asperior Wg, v. Tarax alpine species, their adaptability being superior. Gemnium silvaticum (cf. aci) The following, not previously discussed, species LUNDMAN 1948 p. 156) fall within this category: With regard to a few of these plants, viz. Equisetum arvense Poa pratensis ssp. alpi Melandrium rubrum, Ranunculus acris, Gera Cystopteris fragilis gena Deschampsia caespitosa Eriophorurn angustifo nium silvati�um, and Gampanula rotundifolia, lium. either the Fennoscandian area or the extra Fennoscandian distribution suggests that the As pointed out by SAMUELSSON (1943 p. 93), whole alpine populations are glacial survivors. Alchemilla M'ltrbeckianain Scandinavia probably Differences in behaviour, such as in Linnaea includes both a surviving population and one borealis and Trientalis europaea, which both immigrated from the east. Such is, in all likeli are common on alpine heaths, though the hood, the case with many other plants possessing former is a constant companion of the spruce eastwards continuous ranges, both such as were in the lowlands, and the latter grows elsewhere classified above as probable survivors, e.g. , in shady woods, possibly point to the existence several bicentric species, and such as have not of different ecological races, one of which has been discussed previously. This applies, e.g., survived the latest glaciation in Norway. to the following species: :Finally, some among the alpine plants of our Lycopodium clavatum ssp. Sagina Linnaei region have a non-committal, circumpolar, monostachyon Trolliu8 europceus circumpola.r-montane, or Eurasiatic distribution Botrychium boreale Ranunculus confervoides without considerable gaps. This applies to the Carex brunnescens Rubus Chamaernorus C. vaginata Andromeda Polifolia following: Luzula multiflora ssp. Veronica tenella Equiseturn scirpoides Saxifraga foliolosa frigida Pedicularis Sceptrum-ca- Trisetum spicatum S. nivalis Salix glauca rolinurn Eriophorum Scheuchzeri S. tenuis Polygonum viviparum Pinguicula vulgaris Carex Bigelowii S. cernua .Juncus arcticus S. rivularis and probably many more. In most such cases, it Luzula confusa S. groenlandica is for the present impossible to arrive at a defi Oxyria digyna Pyrola gmndiflora nite opinion regarding the history of a species. Ranuncttlus pygmaeus Petasites frigidus
All these plants belong to the most hardy Postglacial eastern immigrants ·element of our flora, ascending high in the moun tains, and appearing in N orth-Greenland, Spitz .Alnong indubitable postglacial immigrants, bergen and Ltitke Land. It would obviously an eastern group, that arrived by a route north be preposterous to assume that precisely these of the Gulf of Bothnia, is easily discernible. It ,species had been unable to survive the latest comprises continental species, whose Fenno _glac�ation in Scandinavia, simply because their scandian distribution is mainly northern, con ,survival cannot be demonstrated by means of tinuous to the east. They occur chiefly east 94 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark of the Scandes; in northernmost Norway they identification>>, A. E. PoRSILD in M. PoRSILD are concentrated in the interior of Finnmark. 1930 pp. 26 f.); later, it has been definitely Their total distribution is mostly Eurasiatic or identified with the American G. Brandegei by occasionally circumpolar, their ranges in the FERNALD (LOVE 1950 p. 54). Concerning the latter case displaying a gap in Iceland and Salices, cf. FLODERUS 1930. Greenland. In Central Europe they are rare To judge from their Fennoscandian ranges, throughout, completely or almost missing in several other plants are here · of an eastern pro north-western Germany; their south-western venance though in some respect or other diver boundary is often situated in East-Prussia. The ging from the type of distribution described majority of them do not ascend into the birch above, for instance, by reaching farther to the belt, and those occurring there but very seldom west andjor south in Central Europe, occurring pass the tree line. In our region, they are re in Iceland, &c.; some of them have, probably stricted to the parts east of the watershed, save or certainly, arrived in our region not by the a few that possess single stations in the Viri northern route around the Gulf of Bothnia, but haure-Vastenjaure basin. In many cases, their from the .south-east. Of the plants occurring plant-geographical character has already been there, the following ones would seem to be made clear, in the first place by HEINTZE (1909; such eastern immigrants (cf. the maps in HULTEN cf. also DAHL 1934 pp. 107 ff. ). The group 1949b and, in several cases, STEFFEN, 1931 pp. partly corresponds to the >>subarctic>> plants of 319 ff., who deals rather fully with this group of BLYTT (1893 pp. 29 ff.), which, however, include plants): also several species that are here listed as glacial Lycopodium complanatum Salix nigricans survivors. Isoetes echinospora S. Starlceana Within the flora of our region, the following Equisetum palustre Stellaria crassifolia plants fall under this category: Potamogeton g1·amineus N uphar pumilum Scheuchzeria palustris Ranunculus peltatus Botrychium multifidum Salix xerophila Alopecurus aequalis R. reptans Pinus silvestris ssp. lap S. my1·tilloides Calamagrostis neglecta Subularia aquatica ponica Betula verrucosa var. lap ssp. australis Draba nemorosa Picea Abies (excl. v. arc ponica (LINDQUIST Poa palustris Callitriche verna tica) 1947 a p. 67) Eriophorum medium Daphne Mezereum Eriophorum brachyanthe- Stellaria longifolia E. gracile M oneses uniflora rum Nymphaea alba ssp. can- Scirpus Hudsonianus Pyrola chlorantha Carex loliacea dida Carex pauciflora Ramischia secunda C. disperma Actaea erythrocarpa C. chordorrhiza Andromeda Polifolia C. globularis Ranunculus lapponicus C. caespitosa Arctostaphylos Uva-ursi Luzula pallescens Rubus arcticus C. juncella Vaccinium microcarpum Salix borealis Rosa majalis C. limosa Galeopsis bifida S. coaetanea Ledum palustre C. rotundata Mentha arvensis v. S. glauca appendiculata Galium trifidum Juncus stygius Pedicularis palustris ssp _ Listera cordata borealis The American areas of Botrychium multifidum Goodyera re'l!ens Utricularia intermedia and Luzula pallescens indicate >>hibernation>> Corallorhiza trifida Linnaea borealis in north-eastern North America; in Fennoscan N uphar pumilum is, in our region, represented dia, however, everything indicates immigration only by the hybrid with N. luteum. Equisetum from the east. The Galium trifidum reported pratense may, in northern Scandinavia, be of an from Iceland is not identical with the Eurasiatic eastern origin, though having arrived in South species (A. LOVE in litt.; >>From Iceland only Sweden from the south. And Drosera rotundi three poor sets occur in the Copenhagen herba folia var. gracilis and Dr. anglica var. pusilla. rium. They are surely not typical G. trifidum, probably also belong to the group now dealt. but the material is too scanty for a definite with, though their extra-Scandinavian ranges. Origins of the Flora 95 are too little known to warrant any positive russland, iiberlebt.>> This may possibly hold good, conclusions. though in our region the distribution of definite Also with regard to some other of these species, glacial survivors does not in the least resemble the provenance is by no means certain. For �hat of Carex angarae, whose area is rather instance, Scheuchzeria palustris and Carex limosa quite typical of those of the eastern immigrants. may be more frequent in north-eastern Sweden Its total Fennoscandian area (KALELA, op.c., than in South-Scandinavia and Central Europe map p. 161, HULTEN 1949b, map 394) is pract simply because their habitats in Non·land have ically identical with that of Eriophorum brachyan not been destroyed through artificial draining. therum (ibid., map 282) and rather like the range Others, e.g. , Carex chordorrhiza, C. juncella, of such an indubitable eastern immigrant as Corallorhiza, Ranunculus reptans, Vaccinium Rubus arcticus (ibid., map 985), except that microcarpum, and Linnaea borealis, which more Carex angarae does not reach as far south in or less regularly ascend into the regio . alpina, Finland and Sweden. The gap in northern may be glacial survivors; as to V accinium micro Finland is certainly due to the fact that our carpum and Linnaea, see above p. 89 and p. 93� species is rather eutrophic and, therefore, rare Also Ranunculus peltatus and Pedicularis pa in the extremely barren region around the North lustris ssp. borealis may, possibly, be >>hiber Finnish watershed (cf. KALELA Le.). Personally, nators>> (cf. above p. 63). I am inclined to regard it as a postglacial, eastern Eriophorum medium, whose Fennoscandian immigrant to Fennoscandia. area resembles the ranges of certain north This may apply also to Epilobium davuricum. eastern immigrants, may nevertheless fall within It is evidently more sylvan than generally another category. It seems to be a segregate of assumed, though, being eutrophic like Carex an the hybrid E. russeol·um x Scheuchzeri, that has garae, it is rare on the till-plains of the woodland. gained a degree of independence and constancy. In our region, its distribution is very similar However, E. russeolum is, evidently, a glacial to that of the last-mentioned species, as is its survivor in Iceland and north -eastern North Fennoscandian area in its entirety (HULTEN America (cf. LovE 1948); thus it, and conse op.c., map 1279); Epilobium davuricum would, quently also E. medium, may belong to the inter however, seem to be missing in Karelia one glacial, surviving flora of Fennoscandia . . gensis and the adjacent parts. Its total area Not a few species classified above as possible seems to lack gaps, apart from those in Green survivors may, on the other hand, be eastern land and Icel::;tnd (HULTEN 1947a p. 1147). It immigrants; or both alternatives may be true. may thus perhaps belong to the Siberian ele Such is the case with, e.g., the following: ment of our flora. Oarex aquatilis Angelica Archangelica A few words should also be said o£ Carex Salix phylicijolia ssp. norvegica rotundata. It is hardly an alpine plant in the S. lapponum M elampyrum silvaticum proper sense of the word; most of its stations at Viola epipsila Oirsium heterophyllum _ any rate in Sweden are situated below the tree V. montana Petasites frigidus line and a good many of them in the conifer In this context, two species should be specially belt. During a short visit in Karesuando, I 1nentioned, viz. Carex angarae and Epilobium found it on most of the visited mires and now davuricum. As to the former, KALELA (1944 and then also on Sphagnum mosses; it is cer p. i60) declares: >>Teils hat die A.rt diese un tainly much more frequent in those parts than giinstige Periode>> (i.e. the latest glaciation) is evident from HULTEN's map (1949b p. 104), >>in den norwegischen Kiistenrefugien - offen and this is probably true also of the remaining bar in mehrer€m solchen sowohl im Siiden als parts of the woodland of northernmost Sweden, im N orden - teils ausserhalb des Vereisungs still insufficiently investigated by the botanists. gebietes, und zwar ostlich desselben, in N ord- On the Kola Peninsula it is, ace. to KIHLMAN 96 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
(1890 p. 111), >>die haufigste aller Carex-.Arten>>. tiana and Listera ovata), and must, consequently, On the other hand, the map shows the Fenno be classified as southern immigrants to our scandian area of our species to be simply a region. This holds good of the following species branch of its large North-Hussian and Siberian (cf. maps in HULTEN 1949b): area. In North America, Car ex rotundata is Botrychium Lunaria Salix pentandra certainly a survivor (cf. HULTEN 1942 p. 380); Blechnum Spicant Sagina nodosa yet such is hardly the case in Fennoscandia. Asplenium Trichomanes Actaea spicata Athyrium Filix-femina Carda·mine pratensis In all likelihood, there were three waves of ssp. Dryopteris Filix-mas typica eastern immigrants into Fennoscandia, one in Dr. austriaca TU1·ritis glabra fini-glacial time, close on the receding inland Lastrea Phegopteris Sedum acre ice, another during the early postglacial period, L. Robertiana Rubus idaeus and a third one synchronous with, or posterior Polypodium vulgare Fragaria vesca to, the great climatic deterioration and the Potamogeton filiformis Potentilla argentea Triglochin palustre Alchemilla vestita immigration of the spruce. The first one would Agrostis tenuis Lotus corniculatus have consisted of arctic plants, the second one, M elica nutans Oxalis Acetosella of such sylvan species, e.g., Carex angarae, Poa remota Viola mirabilis Stellaria longifolia, Actaea erythrocarpa and P. irrigata V. palustris Rosa majalis, as usually shun acidic raw humus Nardus stricta Epilobium montanum Roegne-ria canina Circaea alpina and occur mainly in more or less eutrophic Scirpus pauciflorus Anthriscus silvestris dells, south-bluffs, &c., and the third one, of Cm·ex pallescens Angelica silvestris the taiga species occurring on the raw humus C. panicea Prunella vulgaris soils of the spruce forests, e.g., Carex globularis, C. Oederi Stachys silvatica Ranunculus lapponicus, R'ubus arcticus and C. vesicaria Veronica officinalis Paris quadrifolia Galium boreale Ledum palustre. Orchis maculata Erigeron acre Finally, it should be mentioned that two Platanthera bifolia Hieracium Pilosella indubitable north-eastern immigrants, Sagittaria Listera ovata natans Calypso bulbosa, and almost reach the A few of these plants, which ascend to compar Calypso eastern boundary of our region; may atively high altitudes, may, however, be glacial perhaps still be found within it. survivors instead of immigrants, viz. Botrychium Lunaria, Triglochin palustre, and Viola palustris. Postglacial southern immigrants This a pplies also to some species discussed earlier in this work, viz. Dryopteris austriaca, One of the rather surprising results from a Nardus stricta, Roegneria canina, Sedum acre, phytogeographical analysis of the flora of our Lotus corniculatus and Angelica silvestris. Nor region is that the indubitable southern immi does the isolated area of Lastrea Robertiana in grants are not more numerous, but rather the Kuusamo region of Finland quite conform fewer, than the glacial survivors. to the assumption of a postglacial immigration. To begin with, the former category includes Less obvious, though mostly reasonably cer such pl3illts as have continuous southward tain, is the southern or�gin of those species areas, while reaching far to the north along the that also have continuous southward areas, o N rwegian coast and being absent, or rare, in while reaching about as far north to the east northern Finland and north-easternmost· Swe of the Scandes as to the west of them. Such a den. Several among these plants are doubtless distribution characterize the following plants: eastern immigrants to South-Scandinavia. Never Equisetum hiemale Sparganium simplex theless, they have reached northern Lapland Matte·uccia Struthiopteris Potamogeton pusillus from the south, or even from the west (for Dryopteris spinulosa Agrostis canina instance, the definitely eastern Lastrea Rober- Sparganium minimum Molinia coerulea Origins of the Flora 97
Festuca rubra s. str. Potentilla erecta lsoetes lacustris Juncus jilijormis Eriophorum latijolium Callitriche polymorpha Equiset·um limosum Potentilla palustris Scirpus acicularis Elatine Hydropiper E. silvaticum Epilobium palust1·e Carex lasioca1·pa Myriophyllum spicatum Lastrea Dryopteris Chamaenerium angustifo- Luzula pilosa �71!1 . alterniflorum Juniperus communis lium Maianthemum bifolium J.11yosotis stricta Festuca ovina Hippuris vulgaris Convallaria majalis Limosella aquatica Eriophorum vaginatum Pyrola minor Gymnadenia conopsea Veronica serpyllifolia Carex dioeca Calluna vulgar·is Salix caprea V. scutellata a. canescens Menyanthes trifoliata Stellaria graminea Utricularia V'ldgaris C. magellanica Melampyrum pratense Barbarea stricta U. minor C. rostrata Antennaria dioeca Finally, the taxonomical position of a few Also some of these plants, e.g., Scirpus aci plants, viz., Betula coriacea, Sagina Normaniana cularis and Gymnadenia conopsea, arrived, no and the alpine type of Rhinanthus minor ssp. doubt, from the east in South-Scandinavia, typicus, is so uncertain as to put surmises re migrating from there northwards into l.Japland. garding their past out of the question. Matteuccia may possibly be a glacial survivor (cf. HULTEN 1949b, map 34); however its partly Summary isolated area may rather be connected with some ecological factor, e.g., the occurrence of raw In this chapter, we have, now and then, moved mull (sensu LINDQUIST, 1938). on the outskirts of botanical knowledge, often Several further lowland plants, which evi operating with mere surmises and being unable dently migrated into northern Scandinavia to arrive at definite conclusions. The results in postglacial times,. reach so far north in Fin tentatively presented on these pages, will no land as to render it scarcely possible to decide doubt before long be revised, since in a dec whether they arrived in North-Sweden from the ade or two, our knowledge of the races and south or from the east, their areas being broadly areas of the alpine plants, within Fennoscandia continuous eastwards as well .as southwards. and outside it, is sure to become more complete. This applies to the following: Our conceptions will then, doubtless, have to be changed in more respects than one. Never Potamogeton alpinus Carex Buxbaumii theless, so many indications point in the same P. praelongus Lysimachia thyrsiflora general direction, that the principal outlines P. perfoliatus Galium uliginosum Phragmites communis Galium palustre of the history of our alpine flora can be grasped Carex fusca even now. From the above account, the following facts In addition come such probable or possible concerning the alpine plants, i.e. plants that survivors as, e.g., the Vaccinium species (excl. everywhere have their proper habitats above V. rnicrocarpum), Caltha palustris, Filipendula or north of the conifer belt, stand out as more Ulmaria, Rubus saxatilis, &c., which were or less indisputable: doubtless reinforced from the south in post Out of 187 alpine plants, occurring within glacial time; their number is certainly consi our. region, 8 can be proved by means of fossil derable, although no particulars can be given. finds to be indubitable glacial survivors, and in Also most of the plants the ranges of which the case of 7 more, fossil finds render an inter within, and outside of, Fennoscandia are too glacial origin exceedingly likely. non-committal to admit even of tentative con At least 97 more plants within this group clusions may perhaps originally have consisted occupy isolated· areas in Fennoscandia and, of a population immigrated from the south therefore, in all likelihood have survived one and/or the east as well as of a · surviving one. or more glaciations in our peninsula. About This applies, e.g., to the following species: 12 would seem to be endemic in Fennoscandia. 8-496149 Sten Selander I 98 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Out of the remaining alpine plants, 12 are foveolatus Ljungh, which doubtless )>hibernated)> bicentric in Scandinavia. in northern Norway, lives in North-Scandinavia Consequently, two thirds of the alpine plants exclusively under the bark of arboraceous of the region can very probably be regarded as birches (not on Betula nana). Similar is the glacial survivors. biotope (insect-holes under the bark of trees) of Phyllodrepa vilis Among the rest, all except possibly a few the staphylinid Er., which display indications of having survived at least according to STRAND (1946 pp. 24, 210) is an the latest glaciation, though these indications indubitable glacial survivor. - In this context, are often, admittedly, extremely faint. it should be pointed out that several glacial survivors among the insects do not, at present, On the other hand, there are no imperative ascend above the timber line or even the conifer . reasons for regarding any one of these plants as limit, e.g., the carabids Bembidion aeneum Germ. a postglacial immigrant, either from the south (LINDROTH 1945, map 83, 1949 p. 442) and or the east. Dyschirius angustatus Ahr. (LINDROTH 1945, map · Thus practically the whole alpine flora of 211, 1949, p. 445) and the lepidoptera Auto our region may consist of glacial survivors. grapha parilis Hb. (WAHLBERG 1941 I p. 54, ], urthermore, six sylvan and a good dozen NoRDSTROM 1941 p. 206) and Syngrapha diasema sylvan-subalpine plants, among them seven trees, B. (ibid.), both of which are )>west-arctic)>. Picea Abies arctica, Populus tremula, viz. v. Accordingly, at least during the Wfu·m gla Betula callosa, B. tortuosa, Alnus incana var. ciation conditions in the ice-age refugia cannot virescens, Prunus Padus borealis, Sorbus var. and have been so high-arctic as generally assumed. aucuparia glabrata, var. can very likely be They should rather have resembled present-day referred to our )>hibernating)> flora. This applies conditions in the south of Greenland. And the also to several plants, now at home chiefly in main cause of the glaciations would seem to thermic habitats, above all on talus slopes and have been an increase in precipitation rather steeps facing south. than a decrease in temperature. In this connexicon, attention should be called It is therefore very likely that also the all to some indubitable )>hibernators)> among Scan regional species, or at least most of them, have dinavian lichens either growing exclusively on survived one or more glaciations. In a few trees, e.g. , Lobaria Hallii (Tuck.) Zahlbr. (.AHL cases, this is made certain, or extremely prob NER 1948 p. 62) and Cavernularia Hultenii Deg. able, by fossil finds. However, this assumption (ibid. p. 33), or never ascending above the usually defies proving, the hypo.thetical hiber timber line, e.g., Cetraria norvegica (Lynge) DR. nating populations having become mixed up (DEGELIUS 1935 p. 51) and Pseudocyphellaria cro with populations immigrated after the ice-age. cata (L.) Vain. (ibid. pp. 145 ff., .AHLNER op.c. The number of postglacial southern immi pp. 70 ff.). grants, whose status cannot be called in question, Also certain entomological facts, brought out is surprisingly small, in fact considerably smaller by LINDROTH (1949 pp. 787 ff.), indicate the than that of the indubitable survivors. An occurrence of birch and high-g rown willows in eastern, postglacial provenance is evident mainly the Norwegian ice-age refugia. A weevil, Cur in a number of woodland species; only a few culio crux Fabr., which possesses a completely such immigrants are n?rmally found in the isolated area in northernmost Fennoscandia and, regio alpina. In all likelihood, however, many consequently, must be a glacial survivor, is surviving populations, chiefly of common, low dependant on high-grown Salices, in Norway alpine species, have been reinforced from the chiefly S. phylicifolia, where its larva lives in east in early postglacial time, though this cannot nematode galls. The heteromeran Rabocerus be proved. VI. MIGRATORY TRAC TS
Glacial Survivors South-Norwegian area; this notwithstanding, nobody has asserted that it should have arrived It is, perhaps, not entirely out of the question there from any other direction. Furthermore, that a few plants can have survived the Wiirm the mountains west of Lule Lappmark, the glaciation within the boundaries of our region. area around Sulitjelma excepted, are not better GRONLIE (1927 p. 54) states regarding the investigated botanically than the rest of the adjacent part of Norway: nwuntain range. That a plant is not reported >>During the last Ice age, the ice which flowed from these parts does not imply that it is actually over the Folia district was not thick enough missing there. For instance, Draba crassifolia to cover the land, but did only form a network and Sagina caespitosa were found there quite of glaciers between mountain parties more or recently as new to Nordland fylke (NoRDHAGEN less free from ice. Along the frontier, the ice 1935 p. 55). When DAHL and NORDHAGEN sheet may have formed an almost continuous botanized in 1920 in the mountains west of cover, only allowing a few nunatakks to rise Saltdalen, their short visit resulted in the remo above the surface . . . >> val of the western limits of several alpine species .At any rate, these hypothetical nunatakks to a position considerably nearer the coast could only harbour a few, exceptionally hardy (ibid. p. 135). During an excursion in the region vascular plants, for instance, Ranunculus gla west of Lake Vastenjaure, Mr G. BRODAL dis cialis, Luzula confusa, or Lycopodiu·m Selago. covered, in 1949, a large area very rich in rare Practicallly all the vascular plants must have alpine plants, which until then had been com immigrated from outside. pletely unknown (BRODAL in litt). I do not, If we were to accept SAMUELSSON's opinion therefore, consider it unlikely that a species, (1943 pp. 80 f.),.our region should have received though not reported from northern N ordland, a large part of its alpine flora from the north, has arrived in our region from the west. since he contends, when speaking of Luzula Only four typical alpine plants, viz. Hie parviflora, that all plants lacking in N ordland rochloe alpina, Carex misandra, Astragalus frigi must be northern immigrants to Lule Lappmark. rlus, and Cassiope tetragona, are of unquestionable However, as already pointed out by .ARWIDSSON northern provenance in Lule Lappmark; and (1943 pp. 117 f.), SAMUELSSON's argumentation among these, only Cassiope reaches our region. hardly holds true. .ARWIDSSON considers the However, it is probable that a number of mainly reason why Luzula parviflora is lacking in the continental, arctic species with a tolerably con coastal mountains to be, first and foremost, the tinuous distribution in the northern parts of climate, the present as well as, particularly, the mountain range and a local southern limit that of the postglacial warm period. This will, somewhere in Pite Lappmark and Salten, e.g., of course, apply also to other alpine plants. Rhododendron lapponicurn, Pedicularis hirsuta, .ARWIDSSON draws attention to the fact that Campanula uniflora, Antennaria carpathica, and Luzula parviflora is missing also west of its Ar·nica alpina, may also have immigrated from 100 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark the north. This is certainly the case with a few whole, the ranges of most fastidious alpine plants low-alpine to sylvan hibernators which have a would seem to display a gap, or a frequency chiefly north-eastern distribution in Fenno minimum, in the mountains around Mt Kebne scandia and are restricted (or almost so) to the kaise and Skjommen. This may be partly due parts of our region situated east of the mountain to inadequate investigation. However in addi range, for instance, Alopecurus pratensis ssp. tion, the local massifs of greenstones, granites alpestris and Salix glandulifera. and gneisses seem to have acted as a barrier The southern element among the alpine plants which in many cases hindered the plant popula is even less, consisting only of Aconitum septen tions, proceeding from the northern ice age trionale. refugia, from meeting and uniting with the The majority of the alpine plants have doubt populations proceeding from the refugium west less arrived, entirely or partly, from the west, of Mt Sulitelma. where they passed the Wtirm glaciation in the That the hibernating alpine flora of our region refugium supposed to have been situated on has chiefly immigrated from the west is to a the coast between the Svartisen Peninsula and certain degree also confirmed by the fact that Vesteraalen (NoRDHAGEN op.c p. 136). This no less than a good score of species are restricted is self-evident as regards Potentilla hyparctica, to the westernmost mountains (cf. below p. 157), which is not known to occur in Fennoscandia that a great number are far more frequent outside our region, and probably also regarding there than further east, and that these western Cardamine pratensis ssp. angustifolia, whose plants are, almost withotlt exception, the same Scandinavian area would seem to extend from as those enumerated above. Lule Lappmark to .Asele Lappmark (oral in As repeatedly pointed out, the glacial sur formation by Mr. 0. RuNE) and the adjacent vivors include >>lowland plants>> in addition to parts of Norway. Carex macloviana also has its the alpine species. However, I have found it Scandinavian centre in these parts, and there more appropriate to deal with the surviving can be little doubt about its place of survival lowland plants in the paragraphR on the post (cf . .ALM 1944 p. 592). glacial immigrants. When it is only a question Also the plants whose ranges show a gap be of the way that a certain plant has passed into tween southern Lule Lappmark and Salten, on our region, the earlier phases of its history are, the one side, and northern Torne Lappmark as a n1atter of course, irrelevant. and Troms, on the other, are clearly immi grants from the west. As far as present knowledge Lowland plants west of the mountain goes, this holds good of Carex nardina, C. bioolor, range Luzula arctica, Arenaria humifusa (NORDHAGEN op.c. p. 26 f.), Sagina caespitosa (NANNFELDT Within our region, the lowland plants occur 1941, map p. 283), Euphrasia lapponica, Pedi chiefly east of the watershed. This is self cularis flammea (TENGWALL 1913 p. 272, BJORK evident in view of the fact that almost the whole MAN 1939 p. 205), and probably Agrostis stolo western parts are situated above the tree limit. nifera var. arctica. Similar, though less pro However, the basin of Lake Virihaure and nounced disjunctions would seem to occur in Lake Vastenjaure supports not a few lowland the ranges of Cystopteris montana, Polystichum plants, both probable glacial survivors and Lonchitis, Roegneria scandica, Carex microglochin, postglacial immigrants. Some of these species, Juncus arcticus, Minuartia rubella, Arenaria which occur west of the mountin range only in norvegica, Melandrium apetalum, Draba lactea, or near the Vuojatatno valley and extend from Dr. alpina, Dr. crassifolia (ARWIDSSON 1943 there continuously downwards along the river pp. 82 f.), Braya linearis, Astragalus norvegicus, Stora Lule alv (cf. BJORKMAN 1939), give the Primula stricta, and Pr. scandinavica. On the impression of having arrived from that quarter, Migratory Tracts 101 i.e. from the north-east. No doubt, this is true tions of western lowland plants to the north of of a few plants of eastern or south-eastern pro Lake Vastenjaure >>are normal outposts and may venance, viz.: Sdrpus Hudsonianus, Maianthe very well be recruited afresh this very day>> mum bifolium, Salix coeatanea, and possibly (trans�ated from the Swedish text) is hardly J uniperus communis ssp. eu-communis and Car ex more than an unsubstantiated assumption. lasiocarpa. On the contrary, it seems most unlikely that In other cases, however, for instance, Dryo these lowland plants should be able, at present, pteris Filix-mas (cf. HULTEN 1949b, map 55), to find suitable habitats on the glacier-covered a gap in the Virihaure-Vastenjaure basin occurs Mt Riddoalge and the surrounding, similar even in the ranges of plants which are missing, mountains. Although the distance between or almost so, in the lowland east of the moun their stations on Lake Vastenjaure and those tains and, therefore, must have migrated into on the inside of Leirfjord does not exceed 20 the valley of Stora Lule alv from the west. The to 30 km, a recent dispersal from the Norwegian explanation is, of course, that this migration coast directly to their limited stations on Lake took place during the postglacial warm period. Vastenjaure seems hardly probable, for reasons It should be born in mind that the altitude specified below pp. 121 f. limits of the vegetation belts have been lowered In one case at least, that of Populus tremula, between 200 and 300 m since that time (cf., the relict character of the stations of a lowland e.g., GAVELIN 1909, SMITH 1920, and ENQUIST plant in these parts appears evident. As pointed 1933), owing partly to the sinking of the tem out by BJORKMAN (op.c. p. 111), the aspen occurs perature, and partly to the raising of the land here only in the shape of >>young plants>> about masses through land -elevation. Three or four one m high which, however, seem actually to thousand years ago, the shores of Lake Vasten be ground -shoots (cf. KIHLMAN 1890 pp. 223 f. jaure and Lake Virihaure, now mainly situated as to a similar condition of Populus in the Kola in the regio alpina, were well-wooded, partly Peninsula). Arboraceous, fertile aspens are even with conifer forests (cf. below p. 132), and not to be found above the conifer belt and the birch forest -ascended into the passes to the lowermost birch belt. The distance from Norway. Consequently, several plants, which the isolated, sterile population on Mt Unna cannot stand the conditions above the tree Titer to the nearest fertile aspens is in all limit and are, therefore, now lacking, for in quarters at least 60 km as the crow flies, partly stance, on the south-facing steeps north of Lake over high mountains. It seems very unlikely Vastenjaure, doubtless used to occur there that diaspores should have been transported during the climatic optimum. It appears very such a considerable distance and then by chance likely that the lowland plants found their way have landed in the small birch groves of Mt to the great lakes almost exclusively through Unna Titer, all the more so since the seeds of the pass between Lake Vastenjaure and Ser the aspen are known only to germinate very fjordskaret on the Norwegian coast, which is rarely. The hypothesis that the aspen has only 677 m in altitude and, thus, was situated managed, by means of vegetative reproduction, entirely below the wood line during the warm to retain its position here even since the post period. On the whole, nearly all lowland plants glacial climatic optimum seems far more prob occurring in the Virihaure-Vastenjaure basin able. Parallels are not lacking. A familiar would seem to be relicts from that time, the instance is Stratiotes aloides in Fennoscandia only exceptions probably being one or two (cf., e.g., SAMUELSSON 1934 p. 187) which has late eastern immigrants, for instance, Rubus vegetated there for even greater periods of arcticus, which have isolated stations there (cf. time. TENGWALL 1925 p. 764). The relict character and the western prove BJoRKM.AN's thesis (1939 p. 215) that the sta- nance may explain a peculiarity that charac- 102 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark terizes many plants in their stations west of watershed would be arrived at if we could assume the mountain range, viz., that they ascend into that low-grown herbids and graminids, and the regio alpina on Lake Vastenjaure and Lake phanerophytes whose organs are winter and Virihaure, though they do not exceed t�e tree summer at >>meteorological height>>, react in a limit east of the watershed and, in several cases, different way to the climatic factors registered not even the conifer limit. This holds good of by our meteorological observations, in other the following species: words, that herbs and grasses are less sensitive than trees to the characteristics of an oceanic Equisetum hiemale Stellaria longifolia · Triglochin palustre Silene rupestris climate (especially the insufficient number of Agrostis tenuis Erysimum hieraciijolium summer days with a sufficiently high maximum M olinia coerulea Saxifraga adscendens temperature) which seem to lower the tree limits Scirpus acicularis Rubus idaeus near the sea and thus to bring about the tree Eriophorum medium Potentilla erecta lessness of the coastal region of northern Norway Carex canescens Callitriche verna C. panicea Calluna vulgaris (cf. ENQUIST 1933). Since, moreover, the lowland C. flava V accinium microcarpum plants occurring in the regio apina of the pro J uncus alpinus var. al- Menyanthes trifoliata nouncedly oceanic Vastenjaure-Virihaure basin pestris Hackelia deflexa are, practically all, immigrants from the west, Polygonatum verticillatum Veronica scutellata they may, perhaps, belong to ecotypes adapted Orchis maculata Melampyrum pratense Listera cordata Galium boreale to the similar climate of the treeless maritime Rumex tenuifolius Valeriana sambucifolia zone (and, in the case of the survivors, of the ice age refugia, cf. below pp. 162 ff.). Owing to In addition come the lowland species which the westward sinking of the altitudinal belts, are lacking east of the watershed and, therefore, the absolute altitude limits of the plants are, evidently reached our region only from the further, not higher in their western, alpine, than west, all of which occur in the regio alpina. The in their eastern, sylvan, stations. The fact following belong to this category: that the wood line on Lake Virihaure and Lake
Lastrea Robertiana Viola rupestris Vastenjaure has been artificially lowered through Poa irrigata Gentianella aurea the lumberings of the Lapps (cf. above pp. 39 ff.) Scirpus paucijlorus Afuga pyramidalis should also be taken into account. Finally, a Epipactis atrorubens Erigeron acre (as indi- principal cause of the phenomenon discussed Sedum acre genous; also one sta may be that, in comparison with the eastern Lotus corniculatus var. tion in the Sarek moun borealis tains) tracts, the oceanic region west of the mountain range is characterized by less sunshine and a All these plants are either possible or probable higher humidity of the air (cf. below pp. 158 f.). glacial survivors, or southern immigrants which In other words, its lococlimate is rather like the have migrated along the Norwegian coast, microclimate of the woods. The ascension of except the non-committal Oarex canescens, sylvan plants into the regio alpina may, thus, M enyanthes trijoliata, and Melampyrum pratense, in a certain degree be compared to the familiar the dubious Vaccinium microcarpum which may occurrence of sylvan species in treeless habitats be either a survivor or an eastern immigrant, on the sea shore. and the eastern Stellaria longifolia. It is, how Stellaria ever, highly doubtful whether the Lowland plants east of the mountain longifolia occurring in the regio alpina of Mt range Kerkevare belongs to the same type as the eastern woodland population (cf. Part II p. 11). All the lowland plants occurring only east of A conceivable explanation why these plants the watershed within our region, have not ar ascend above the wood line only west of the rived from the south or the east. A few have Migratory Tracts 103 doubtless come fron1 the west, viz., the inter wood line. After the great climatic changes, the esting group which occurs within an isolated species that are now lacking in Pite Lappmark area around the lakes of Saggat, Peuraure and died out within the interjacent area, unless they Karatsj. These species, which here grow almost still possess undiscovered stations in the moun exclusively in thermic habitats such as south tains of western Pite Lappmark. facing talus- slopes and precipitous rocks, reach Besides a number of probable glacial survivors, far to the north along the N orwegia.n coast but e.g. , Polygonatum verticillatum, Silene rupestris, are lacking in the provinces of Vasterbotten . .Arabis hirsuta var. subalpestris, E1·ysimum hiera and Norrbotten and in 1\iid-Lapland. Some ciifolium, Saxifraga adscendens, and Hackelia of them occur in southernmost Lapland, their deflexa, sev�ral other southern immigrants, for stations there being outposts from their Nor instance, Dryopteris Filix-mas and, probably, wegian areas. This applies to: Blechnum Spicant, Carex Oederi, had in all likelihood about the Asplenium septentrionale (one isolated station same history in these parts, though this is now in N orrbotten, two stations in Pite Lappmark), impossible to prove. However, the main part A. Trichomanes, Listera ovata, Alchemilla vestita, of the southern lowland plants certainly reached Viola mirabilis, Epilobium montanum, Cir our region from the south-east and the south. caea alpina, Stachys . silvatica, and Hieracium Even .ANDERSSON & BIRGER (op.c. p. 159) Pilosella. declared: The areas of Oxalis Acetosella and Carex >>The vast majority of species whose distribu pallescens are also rather isolated. One alpine tion was closely investigated, indicate that the plant, Saxifraga Cotyledon, which in our region coastal regions round the Gulf of Bothnia behaves like a lowland species, has a similar were the main tracts along which our flora made
N orth-Scandinavian distribution. its way northwards. From this base, the plants · As regards the way followed by these plants have spread to a varying extent along the river to the Kvikkjokk region, the observations made valleys in the direction of the mountains. This by .ANDERSSON & BIRGER (1912 p. 75 f.) are is probably true also of those species which no doubt correct. They all grow in Junker are now dispersed all over northern Sweden>> dalsuren (DYRING 1900) or in other localities (from the original Swedish text). in Salten. .A few of them occur also in Pite This opinion still holds good in the case of Lappmark, viz. Asplenium septentrionale, Carex most southern immigrants. The number of such pallescens (several stations), Oxalis Acetosella species that followed the way indicated is too (do.), .Alchemilla vestita (Lake Iraft near .Adolf great to admit enumeration. In one respect, strom, SA.MUELSSON 1943 p. 150), and Stachys the south-eastern immigration tract would silvatica (one station far to the west on Mt seem to have been of even greater significance .Avatjakko, BIRGER 1921 p. 21). than .ANDERSSON & BIRGER were willing to These species seem to have arrived in Pite admit. These authors considered all the pro Lappmark during the postglacial warm period nouncedly southern, thermophilous species with from J unkersdalen or Evenesdalen i Norway in the mountain flora of Lapland, which occur by way of the pass on Lake Juronjaure, or that exclusively on precipitous bluffs and talus near Lake Balvatn; the altitude of these passes slopes facing south (>>sydberg>> ), to have immi has not been measured, but does not, at any rate, grated, partly or entirely, from Norway. Thus, exceed 725 m s. m. Accordingly, during the cli they state (op.c. p. 167 f.), >>that among the nlatic optimum, both passes were situated in South-Scandinavian species, apart from Betula the birch belt. From there the plants probably verrucosa in the Kvikkj okk region, only a few, migrated to Lake Peuraure and Lake Saggat if any at all, can be regarded as originating by way of the pass on the river Parkajokk, solely fron1 the Bothnian stream of invaders. which is even to day wholly situated below the The great majority, especially the thermo- 104 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
philous species, would seem to have arrived able number of such mountains with a thermo in their present mountain stations by the philous flora, investigated by ULANDER (1929), Atlantic migratory tract alone>> �transl. from the LONNQVIST (1944, 1947, 1949), HANNERZ (1945) original Swedish text). and the present author (1949), are found, i.a., This can, evidently, not be correct. The in N orrbotten, especially within the area of Laplandian Betula verrucosa is not a southern Arvidsjaure porphyry in the vicinity of .Alvsbyn. immigrant but an eastern one. This error on For the time being, detailed flora lists are avail the part of ANDERSSON & BIRGER is due to able from 17 such localities in N orrbotten. The the fact that var. lapponica, in their time, had following southern and eastern species (includ not yet been distinguished from var. saxatilis ing a few of doubtful origin), which occur in Lindq. Thus, the only south-eastern immi our region, are reported from these mountains, grant recognized by them disappears. On the the figures in brackets signifying the number other hand, four thermophilous species charac of known stations: teristic of the >>sydberg>> flora are altogether Matteuccia Struthiopteris Actaea spicata ( 2) lacking in N ordland, viz. Dryopteris spinulosa (3) Draba nemorosa ( 4) (in Norway not occurring north of Nord-Tren Asplenium septentrionale Turritis glabra ( 4) delag), Myosotis stricta (do.), Galeopsis bifida (1) Sedum acre (6) (do.), and Draba nemorosa (in Norway only as Athyrium Filix-femina (5) S. annuum (4) Woodsia ilvensis ( 10) Rubus idaeus (14) a synanthrope). Consequently, they cannot Polypodium vulgare (12) Fragaria vesca ( 15) have come to Lule Lappmark from the west. Roegneria canina ( 8) Potentilla argentea (9) The three last-mentioned species occur in the Convallaria majalis (13) Rosa majalis ( 4) interior of Norrbotten (cf. below), and Dryo Gymnadenia conopsea ( 2) Daphne MezeTeum (4) pteris spinulosa is doubtless often overlooked; Platanthera bifolia (3) Epilobium collinum (8) Goody era 1·epens ( 5) PyTola chlorantha (9) closer investigation will probably show that its Stellaria graminea (9) Myosotis stTicta ( 3) two stations in the neighbourhood of Kvikk St. longifolia ( 4) Galeopsis bifida (6) jokk, now seemingly quite isolated, are con Sagina nodosa ( 1) V eTonica officinalis ( 9) nected with its area in N orrbotten where it Actaea erythrocarpa (9) ETigeTon acTe (2) reaches Matarengj (LONNQUIST 1939). Accord ingly, there is every indication that they have In this connexion, one mountain is of especial arrived in northern Lapland from the south-east. interest, viz., Sokkosberget in the parish of The reason why ANDERSSON & BIRGER Edefors, situated within the area of gneissic underestimated the significance of the south granites about half way between Boden and eastern migratory tract certainly was that, in J okkmokk, i.e. considerably nearer our region their time, hardly any botanists had explored than the other mountains dealt with here. Its the interior of N orrbotten and Vasterbotten existence was pointed out to me by Mr AXEL and the low mountains of eastern Lapland; ELGSTRAND of IJulea. The almost perpendic the botanical investigation of these parts is still ular bluff is about 100-150 m high and faces far from completed. At present, however, south-west. At my visit there, I was only able known facts do not at all agree with their state to investigate the lower ledges, the upper and, ment (op.c. p. 39), that >>out of about 70 'syd probably, more interesting parts of the precipice berg' within the region of primary rock, con being inaccessible. However, besides a few cerning which we could gather more or less alpine plants and possible glacial survivors (Poa complete records showing them to support glauca, V iscaria alpina, Sedum annuum, Saxi fraga groenlandica, Epilobium collinum), . South-Scandinavian species, not a single one the fol is situated north of a line from the river Ore lowing southern and eastern species were noted: .A alv north-westwards to upper ngermanland>> Polypodium vulgaTe ConvallaTia majalis (from the original Swedish text). A consider- RoegneTia canina StellaTia graminea Migratory Tracts 105
Stellaria longijolia Fragaria vesca get well on the way until the immigration of Actaea spicata Daphne M ezereum the spruce. Previously, i.e. during the warm A. erythrocarpa Pyrola media period, more or less open birch forests, in south Turritis glabra Veronica ojjicinalis Rubus idaeus ern Lapland with a weft of i.a. lime, and with a luxuriant undergrowth of herbs and ferns, seem Rubus idaeus and Fragaria vesca were found to have prevailed on fertile soils, at least near also in one locality between Sokkosberget and the mountain range, in the place of the present,_ the stations of the mountain range, viz. Mt monotonous spruce forests (ERDTMAN op.c. Piertinvare between J okkmokk and Luvos. pp. 58 ff.). It goes without saying that the All the species mentioned previously, except South-Scandinavian species, prior to the great Asplenium septentrionale, whose single station climatic change, were able to live, not as now in Norrbotten would seem, in all likelihood, to adays only in natural hot-beds formed by the be an outpost of its Lapland area, and Sedum rock-faces, but in other habitats too, e.g., river acre, which is found in our region only west banks, south-slopes, &c., much as they do now of the watershed, may have migrated into Lule in Mid-Sweden. Consequently, they had much Lappmark from this quarter. This seems to be greater possibilities of migrating, both from reasonably certain as regards several species the south-east and from the Atlantic coast,_ other than the four above-mentioned ones, than is suggested by their present, scattered e.g., Fragaria vesca. On account of its almost relict stations in the woodland. continuous distribution from the mountain range However, another migratory tract exists that right down to_the coast, this plant has apparently was entirely overlooked by ANDERSSON & reached its Lapland area both from Norway BIRGER, viz., the one along the border of the · and from the Bothnian coast and not, as asserted mountain range. The >>glint>>, i.e. the outer edge of by ANDERSSON & BIRGER (op.c., map p. 365), the overlapping mountain schists, &c., forms a exclusively from the west (cf. BJ6RKMAN 1939 chain of bluffs with a thermophilous flora, which p. 216). generally face south-east and are situated at a True, the distance fron1 Mt Sokkosberget, relative distance from one another of about 20 km where some of the species in question have their at the utmost. When writing their monograph, north-westernmost station outside the rrwun ANDERSSON & BIRGER knew only of a small tain range, to the easternmost locality of this fraction of these localities, chiefly those situated kind within our region, viz., l\itFarfor ita, equals on the big river valleys. In Lule Lappmark, as much as about 120 km. However, when they possessed detailed reports from only three more closely investigated, the interjacent area such places (op.c. pp. 197 ff.; ULLENIUS' and will probably reveal more such localities. WAHLENBERG's findings on Mt Kiuri as well Even if this were not so, it should be remem as WESTLING's report from Mt Tjergo were bered that present conditions are far less pro overlooked by them) and stray items from pitious to exacting southern species than those seven others, whereas at present close on 80 prevailing during the postglacial climatic opti have been more or less thoroughly investigated,_ mum. The mean temperature of South-Scandi about 20 of them on or near the >>glint>> (cf. navia was, in atlantic and subboreal times, at map fig. 17). least 2° C. higher than now (IVERSEN 1944 At present, no appreciable dispersal of South p. 4 79; ANDERSSON [1902] arrived at a similar Scandinavian species from one bluff to the result). And leaching and podsolization had other seems to occur (cf. below p. 121). How not impoverished the soils as much as nowadays. ever, during the climatic optimum, such locali Pollen analyses of podsol soils from southern ties were more numerous, the forest having Lapland, made by ERDTMAN (1943), seem to since then invaded many talus slopes, which indicate that the formation of raw humus did not have thus become >>fossil» (cf. FRODIN 1915 106 SELANDER: Floristio Phytogeography of South-Western Lule Lappmark
e Investigated 0 Mentioned by A&B. 0 Investigated o Uninvestigated before 1912 after 1912
Fig. 17. Cliffs facing south-west, south or south-east. p. 211). Since the South-Scandinavian plants Ringle valley. Among the 13 remaining species, at that time possessed also interjacent stations which are restricted to the region in question, 10f other kinds, they spread, no doubt, not only one alone occurs in all three valleys. No more along the river valleys, as assumed by ANDERS than 16 microspecies, practically all belonging SON & BIRGER, but also from valley to valley. to the widespread category, are observed in There is even proof of sorts, that such a dis the adjacent parts of Norway (OMANG in DAHL persal took place at the time presumed, namely 1912 p. 200 ff. and 1915 p. 152 ff.); the actual the distribution of the Arohieraoium microspecies number is probably slightly larger, since the in the river valleys of Lapland. Thanks to inves species concepts held by OMANG and FoLIN tigations by l!"' OLIN (1928, 1931, 1934, 1942; are hardly likely to be identical. The surpris cf. also DAHLSTEDT 1925 and ARWIDSSON 1937), ingly high number of endemics restricted to the Arohieracium floras of the upper stretches one valley may be due, as suggested by GusTAFS of the Ume, Skellefte and Lule valleys are rather SON (1947 p. 217), to the fact that )>most of well known. Among 151 microspecies found them lack all progressive properties)>. However, in the investigated valleys, only 52 are known another factor must also be taken into account, to occur outside this region. Among the latter, i.e. the climatic change at the end of the sub 12 are widely spread over large parts of northern boreal period. Since the Hieracium flora is very Scandinavia, and 8 others occur throughout poor in the middle parts of the valleys, with the mountain range, or nearly so. No less than their barren morainic soil, and immigration from 86 microspecies seem to be endemic in one Norway as a rule seems to be out of the ques- Migratory Tracts 107 tion, the hawk-weeds that are restricted to the the next valley was not bigger than that between region discussed, and present in more valleys Umedalen and Skelleftedalen. The proportion than one, must have spread from valley to between strict endemics and more widespread valley across the watersheds, although, like types is quite in accordance with the extreme most plants of a southern provenance, the im youth of most Scandinavian Hieracium micro migrated Hieracia but rarely ascend above species. the tree limit. Still during the warm period Unless a dispersal of plants along the border so many passes were situated in the birch belt of the mountain range is assumed, no explana that the hawk-weeds could easily migrate from tion can be given of the Lapland areas of, e.g.t one valley to another . .Along with the climatic Arenaria serpyllifolia (HULTEN 1949b� map deterioration, this possibility came to an end, 721), Draba nemorosa (ibid., map 901), Turritis and the microspecies arisen after that time glabra (ibid., map 922), Potentilla argentea became isolated in the valley where they had (ibid., map 1012), and Myosotis stricta (ibid.1 come into being, even though the distance to map 1470). VII. DIST RIB U TIONS WITHIN THE RE CION
Old and new findings of certain species conspicuous, early described species are likely to have escaped our keen and wide-awake Phytogeographers often try to explain pecu botanists in times gone by, _in the places visited liarities in the distribution of a plant by an by them. assumption that the spreading of the species Such a plant is Cassiope tetragona. It is has not yet come to an end. Such a conception, generally assumed not yet to have reached however, is incompatible with the trend of its definite limits in Scandinavia; this opinion modern plant geography,' according to which was first expressed by TH. FRIES (1917 p. 30) migration is mainly a mass phenomenon; the and_ later adopted by BJORKMAN (1939 p. 209), migrating units appear not to be individual Du RIETZ (1942 b p. 48), and ARWIDSSON plant species, but a sequence of communities (1943 p. 86). Phytogeographically Cassiope in a sere or climaxes following one another in tetragona belongs to the northern unicentric a clisere (cf., e.g., GAIN 1944 pp. 88 ff.). It would, group and has doubtless survived the Wiirm in fact, seem most improbable, that one single glaciation in the refugia . of North-Norway. vascular plant, belonging to the indigenous In Lule Lappmark, it lives close to the boundary alpine flora of our region, should still be gaining of its area, its southern Swedish limit being ground outside its old range, except in new situated on Mt Kapsje in northernmost Pite habitats shaped by Man. Lappmark (cf. ARwmssoN op.c. , map p. 90). As pointed out above, our region was the If it were still spreading, one might expect first part of the Scandinavian mountain range to find, in this particular region, at least some whose flora was investigated. The fact that the locality where it had turned up during the last earlier botanists usually followed the same hundred years. This, however, does not seem route is rather an advantage when it is a ques to have happened. On the map fig. 18 all known tion of forming an idea as to the distripution stations have been registered, those discovered of a plant species a hundred years ago, since before and after 1848 being marked differently; the flora of certain mountains was, consequently, the routes chosen by the earlier botanists have more thoroughly examined. True, our know also been denoted. From this it appears, that ledge of the plants occurring on a given mountain the new finds were made almost exclusively can hardly ever become complete. During within areas not previously visited, while the this century new finds, e.g., Roegneria scandica, earlier finds are assembled round the foremen Carex bicolor, Sagina caespitosa, Minuartia tioned routes. Of new finds near the old ones rubella, and Braya linearis, have been made there are only two. One was made on Mt Jun even on Mt Tuki, the smallest and most exhaus kartjakko, where CLEVE (1901 p. 43) found a tively ransacked among these mountains. But few specimens in 1896. I have passed this the species recently found on such mountains ,spot five times without finding Cassiope tetra either belong to more or less critical genera or gona, though I particularly searched for it. It are diminutive plants easily overlooked. No is therefore no surprise that it had been over- Distributions within the Region 109
• Before 1847 0 After 1847 Only one dot on each mountain ---- Customary routes of the earlier botanists
Fig. 18. Old and new findings of Ow;;siope tetragona within the investigated region. looked by earlier botanists. The second new there. Afterwards, it was not discovered in locality is the north bluff of Mt Njerek, which any new locality in these parts, although these no botanist had visited. On the other hand, rrwuntains were scoured by all the botanists the stand on Mt Sirkavare that consists of a visiting Kvikkjokk, until 1944, when I found single patch not measuring one square meter, a single tuft on the barren boulder ground of and that on Mt Tuki, where a few small tufts the eastern slope of l\'It Kaskaivo. One knows are growing, were mentioned already by WAH how a spreading species behaves, e.g., Thlaspi LENBERG (in his diary) and by .ANDERSON (1846 alpestre, Cuscuta epithymum or, in the mountains, p. 21; >>below one single cliff>>) respectively. Achillea Millefolium and a few other neophytes. In these two localities, our plant has, conse In any case, Cassiope tetragona does not imitate quently, not spread at all for a century. Out their behaviour. of the three southeasternmost outposts of the Cassiope tetragona possesses so much vitality .species, two have been known for more than and spreading capacity as to having become a hundred years. On 1\ft Vallivare, it was found predominant over huge areas, e.g., in Torne by WAHLENBERG in 1807 (cf. the diary of WAH Lappmark; and it has had a good 9000 years LENBERG; WAHLENBERG 1812 p. 107); here at its disposal for dispersal over the Scandina it still grows in the north precipice, in the vian mountain range. It would seem, then, ;Same places as then. .ANDERSON (1844 p. 26) that it should have had time, within so compar found it in 1843 on Mt Njunjesvare; and it atively limited an area, to take possession of would seem to have gained no more fresh ground the whole of available Lebensraum, provided 110 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
• 0 Cassiope tetragona -heath e 0 Fairly plentiful occurrence • o Single tufts
Dots: Stations observed by the author Circles: Stations taken from literature, & c.
Fig. 19. Ca8siope tetragona at its southern Swedish boundary. there are no impassable barriers of any kind Parte massif and central Sarek, and that the in its way to a suitable station. In my opinion gap in the west is less striking within the it has, in fact, done so. The outposts of a northern parts of our region. On the whole, Oam spreading terrestrial plant generally consist of panula uniflora is more frequent in the region a few specimens occuring in stray places. Such than Oassiope tetragona. In the mountains is not the case with Oassiope. Contrariwise, south and south-east of Lake Virihaure, it areas of Oassiope tetragona - heath occur even occurs, though rather sparingly, about 800 hard by its southern limit (cf. the map fig. 19). to 1200 m s. m. in most Dryas heaths that are The character of its southernmost stations is not too dry and become free from snow early typical of a stable limit as distinguished from in the summer, provided the vegetation does a labile one. Though its crescent-shaped area not cover more than 20 to 30 per cent of the in Lule Lappmark strikes one as rather queer, ground. Its absence from all the Vastenjaure the oceanic 1ococlimate as well as other factors and large parts of the Virihaure basin n1ust (cf. pp. 38 and 131 f.) may explain its shape. therefore be due, i.a., to climatic causes. I J\1oreover, it is hardly unique. The area of can not be persuaded to believe that it would Oampanula uniflora presents in the main the not have had time to spread, e.g., from Mt same features (cf. the map fig. 20, that shows Kerkevare, where it abounds, the few kilometers also a distribution of old and new finds similar to J\ft Unna Titer, where it is completely lack to that of Oassiope tetragona). The differences ing, although the right kind of Dryas heath. are chiefly that Campanula occurs also in the covers considerable areas also there. Distributions within the Region 111
• Reported before 1847 0 Reported after 1847 Only one station on each mountain
+ Arasvare, report in Swartz 1821 ---- Customary routes of the earlier botanists
Fig. 20. Old and new findings of Campanula unitlora within the investigated region.
Maps of the same sort, as those of the areas Glacial survivors as apophytes of Cassiope tetragona and Campanula uniflora, have been made also of some other species, viz., The reason cannot possibly be that high Melandrium apetalum, Pedicularis hirsuta, P. alpine species, even though they may po ssess flammea, and Arnica alpina, all conspicuous only an apparently very feeble spreading capac and falling within the category that ARwrnssoN ity, would be unable, under favourable cir (Le.) considers to have )} Verbreitungsgrenzen)} cumstances swiftly to invade new habitats. The above-mentioned occurrence of Phippsia in Pite Lappmark and BJORKMAN (Le.) declares algida in a reindeer pen in the birch belt at t.o be still spreading in Lule Lappmark. These Stalonjarka may serve as an instance of such maps, not reproduced here owing to lack of dispersal. True, the distance to its nearest space, show that the whole body of plants were )mormah stations, on the summits of Mt Stalo found before 184 7 in all localities known to tjakko and Mt Kuobberi, does not exceed about have been visited by botanists prior to that 5 km. Still, even this is a creditable jump on year, except, possibly, a few stations of Arnica; the part of a plant that is so little fit for long however, this species is very often sterile and distance dissemination that its diaspores usu then easily overlooked. Apparently, during the ally seem to land only a few centimeters from past century none of these six · species has the mother specimen. The explanation is, of displayed the very slightest tendency to widen course, that disseminules have happened to its area. attach themselves to reindeer hoofs. Thus, 112 SELANDER: Floristic Phytogeography of · South-Western Lule Lappmark a seemingly far from well-equipped species 1926 p. 215; map in Du RIETZ 1940 a p. 220). possesses a spreading capacity that seems to In other words, it doubtless obtained its distri invalidate the theories of distributional limits bution, broadly speaking, as ea:dy as during within the native alpine flora conditional on the Tertiary, and in Fennoscandia it belongs time. If all goes well, alpine plants may conquer to the glacial survivors that probably lived ·even far greater distances. The invasion of there through at least two glaciations (cf. .slag heaps in Bergslagen by Viscaria alpina is NORDHAGEN 1935 pp. 152 ff.). a case in point (cf. LID & ZACHAU 1928 p. 79, Yet, it has not, at the present time, lost its ;SAMUELSSON 1943 p. 84). Probably, the possi capacity of local spreading. On the contrary, bility of ecesis is, instead, the decisive factor. it so often appears in new localities that owe When new habitats with few or no con1peting their existence to cultivation, that several plant species arise, as in the cases under dis- authors doubt its occurrence in original habitats �ussion, they will often become populated by al in Lappland (BJORKMAN 1939 p. 188, ALM pine plants. However, in untouched wildernesses op. c. pp. 594 f.). In our region, I have seen such new habitats would seem to be so rare, only two populations, both on Ranunculus at any rate during periods of stable climatic acris-meadows in :what was formerly called conditions, as to be of little or no significance the willow belt, which are to all appearances (cf. below p. 119). quite native. In addition, several among the It may be objected that nobody has ascribed stations reported by TENGWALL (1924 p. 640), limits due to insufficient spreading time to the e.g. , Jillavagge, Tsakkok, and Pulkas, are two forementioned species. But the difference situated in regions where the Lapps hardly between a faintly bicentric species, such as ever come. They may, thus, safely be regarded Phippsia algida, and a markedly bicentric as original. one, such as Campanula uniflora, is quite However, also in our region the main part arbitrary; if a larger gap is determined by time, of the stations of Carex macloviana are of quite the same would evidently apply to a smaller another kind, viz., greenswards, pasture-grounds, one. Moreover, similar observations may be farm court-yards, footpaths, roadsides, &c., made with regard to those pronouncedly centric in the conifer belt. It was found at all settlers' alpine plants that occur as apophytes in the farmsteads visited, except Njunjes. In Pite lowland. They are few in number, for the Lappmark it would seem. to have been observed obvious reason that species belonging to those only as an apophyte in the woodland (.ARwrnssoN types of distribution have suffered more than 1943 p. 170). Almost all its stations in Lycksele the rest fron1 biotypic pauperization during Lappmark also belong to this type (ALM op. c. the glaeiations and, therefore, find it very p. 585), though it has recently been found there difficult to get along under unfamiliar condi in one primary alpine station (RUNE 1948 tions. There are some exceptions, though. p. 496). A well-known instance is Carex macloviana. ALM's opinion (op. c. p. 592), that Carex It is evidently a very old species. Taken as macloviana survived the Wi.irm glaciation in a collective unit, its distribution on the northern refugia between Salten and IJofoten, can hardly hemisphere belongs to the west-Arctic type; be refuted (cf. above p. 100). No doubt, BJORK moreover, it is bipolar, with stations in the .Andes MAN and ALM are right also in assuming that from Equador southwards, in Tierra del Fuego, it was brought by nomadizing Lapps to its and in the Falkland Islands; finally, it is one stations in the woodland, from alpine stations among the five (or six) vascular plants that still partly unknown (e.g., those in western the Fennoscandian flora has in common with Pite Lappmark). The secondary stations in the venerable indigenous flora of the Hawaiian the conifer belt are very numerous, being at Islands (cf. ALM 1944 p. 582, SKOTTSBERG least as many as the primary alpine ones, and Distributions within the Region 113
e Probably spontaneous stations ® Probably secondary stations, near or on caravan tracks and foot paths
0 A pophytic stations at farmsteads and on lapp yards 6. Lapp camps.
---- Caravan tracks of the lapps • · ·· · · · Foot path
Fig. 21. Carex macloviana in Lule Lappmark. dispersed over a much larger area (cf. ALM limits as a spontaneous plant must be attributed op. c. pp. 584 ff.). However, even the most to other causes. ancient of these habitats did not come into The same applies to the bicentric Ranunculus being before the colonization of the Laplanders' hyperboreus, which BJORKMAN ( op. c. p. 208) territory began, less than 300 years ago (CAMP considers not to have had the time yet to fill BELL 1948 pp. 52 ff.). In our region, most of up the gap between its northern and southern then1 are still more recent, dating, as a rule, areas. No doubt, this species is actually spread not further back than to about 1800 at the ing, but only within the conifer belt and with utmost (cf. above p. 34). In other words, the help of J\Ian, mainly in ditches, &c. (cf. Garex macloviana has been able to take posses HEINTZE 1907 p. 235). The starting-points sion of by far the largest part of its Fenno of this new-colonization were probably the scandian area in the course of an infinitesimal stations on alluvia, &c., at the rivers of the fraction of the time at its disposal for spreading regio subalpina and the regio coniferina (cf. since the melting of the inland ice. It, then, TH. FRIES 1913 p. 225). On the other hand, in seems rather preposterous to assert that it the mountains proper still untouched by human had not >>had the time>>, during the postglacial int.erference, Ranunculus hyperboreus would seem age, to spread more than a few hundreds of to occur everywhere, at any rate in Lapland, kilometers along the mountain range. Its present only in scattered,. widely separated stations 9-496149 Sten Selande� I 114 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark without showing any tendency of beeoming untouched vegetation. We are, doubtless, in more frequent or invading new territory. clined to underrate it, owing to the unstable Another instance is Alopecurus pratensis ssp. character of the plant communities on ground alpestris. It has a most peculiar Fennoscandian influenced by cultivation predominating in distribution, and its history is obscure (cf. our daily environment. Drastic instances of above p. 84). However, in Lapland it displays the swift changes in such communities are an obvious relict character. It is also one of our given, e.g. , by JULIN (1948) from South-Swedish rarest mountain plants, its known primary cleared copses, which are becoming overgrown alpine stations numbering only two in Lule since haymaking came to an end. The moss Lapp mark , and three or four in Pite Lappmark. synusiae of the bottom layer are the most In spite of this, it manifests a great spreading unstable, in some cases completely changing capacity in its secondary sylvan stations, character in as short a time as 14 years (op.c. and has become quite common on moist hay pp. 95 ff.); but also the composition of the meadows round Adolfstrom (ARWIDSSON op. c. herb layer may assume an essentially different p. 160) and around the settlements on Lake aspect in a few decades (ibid. pp. 107 ff.). Peuraure and Lake Saggat right down to Lake On the other hand, stable climax communities Tjamotisjaure. (sensu GLEASON 1927 p. 325 and Du RIETZ 1930 Among plants lacking in our region, Chryso a pp. 346 ff., not sensu CLEMENTS e.a.) are, of splenium tetrandrum (cf., e.g., HEINTZE l. c.) course, in untouched wildernesses rather the and Polemonium acutiflorum both occupy iso rule than the exception, unlike settled tracts. lated areas in northernmost Fennoscandia By climax communities I mean such communi (HULTEN 1945 a p. 950, 1948 p. 1323) with ties as Hepresent time-phases of great stability, comparatively few primary stations. In all in which we cannot observe the action of the likelihood, they are glacial survivors; yet they successional causes and for which we cannot are pronouncedly hemerophiloLis, and most predict the future)> (GLEASON Le.), i.e. about the of their present stations are to be found on same thing as Lunr's "Dauerformationen" (LuDI ground influenced by civilisation. The Higid)>, 1930 p. 532). Their constancy is often amaz extremely rare hibernator Melandrium angusti ing. The most striking instance is probably florum occLirs as an apophyte on the railway the West-American redwood forest, which has enbankment at Abisko (LAGERBERG 1940 p. 148); remained essentially unchanged since th_e Upper and the similar Triseturn subalpestre was seen Oligocene at the very least, although it has growing abundan.tly in sandy hay meadows migrated from, at any rate, Washington and and court-yards in Nordreisa (SELANDER 1945 b Oregon to western California; during the Eocene, p. 77). it probably occurred with, in all likelihood, the Accordingly, there is no single fact indicating same main composition as to-day, still far that any centric plant of our region is still ther north, for instance in Alaska, from where spreading in its natural environment, while it migrated southwards (CHANEY 1925, 1936). numerous cases are established of such plants Better than anything else the redwood forest migrating in environments influenced by culti also illustrates the rule that the migrating units vation. Consequently, their usual stationari do not consist of single species, but of plant ness will not always be due to deficient spread communities as a whole. ing capacity. The chief exceptions are, of course, the neo phytes. In our region, however, neophytes have
Stability of plant communities been observed almost exclusively in places where equilibrium had been upset by hay The cause would rather have to be looked making, grazing, lumbering, or human inter for in the stability of plant communities in ference of some other kind; alpine species inci- Distributions within the Region 115 dentally occurring in the conifer belt are also control experiments with plants cultivated far more frequent in such loca,lities than else in garden beds in order to eliminate the clima where (cf. SERNANDER 1899 p. 8). Foreign tic factor. However, the climate indoors, elements seem hardly to stand a chance of where Primula stricta managed to live, could intruding into the climax communities. hardly be better suited to arctic plants than Apparently, most plant sociologists, more that out of doors, where it died. or less unconsciously, consider this state of Certain alpine, half-open grass heaths, e.g., things as a matter of course: if the climax a Kobresia myosuroides-heath on Mt Ja,llotsj, or communities could not be supposed to keep the patches of Festuca ovina-heath, may be free from unlooked-for intruders, plant socio mentioned as examples of plant communi logy would end up in a hopeless confusion. That ties of the kind referred to by PoRSILD. They is probably why so little research has been are extremely poor in species, although with carried out on this subject, at any event in their patches of bare ground, they should invite the Arctic. However, PoRS:iLD (1932 pp. 35 ff.) to colonization from outside, considering that tried to tranfer native Greenland plants to · competition in the usual sense of the word, suitable habitats near the Arctic Station of i.e. struggle for space, is non-existent and, Disko, where they did not grow before, among further, that regarding soil-fertility, irrigation those occurring in our region Garex canescens, and other edaphic factors they do not appear Saxifraga aizoides, 8. stellaris, and Primula to differ greatly from adjacent communities strict a. The specimens flowered and bore fruit richer in species. Of course, even in such cases in the first summer, but in a few years almost the distribution of species, and the possibility all the plants died. It is particularly interesting of a certain species' becoming predominant, to note that specimens of Primula stricta grew are in the first place decided by edaphic condi well in pots indoors, though without flowering, tions, e.g. , the chemical nature of the soil and, while those planted outdoors died away after perhaps especially, the distribution of snow the first winter. PoRSILD sums up his results in the winter and the time of its melting away. as follows: However, some additional, generally overlooked, >>Every untouched plant association may be factor must be in operation. considered to be in a state of equilibrium, This factor may, conceivably, consist in the even such arctic associations where the areas capacity of certain plants to secrete a sub of bare ground exceed those 'covered' by vege stance from their subterranean parts that tation. The bare spots, whatever they may be acts as a poison upon certain · other plants. -rocks, sand and clay - are bare for reasons; As pointed out by OsVALD (1947 p. 290), already and when the plants have succeeded in getting DE CANDOLLE (1832) considered that this a foothold, they become 'beati possidentes', explained the formation of plant communities. not willingly allowing newcomers to settle, Also, more than 40 years ago, American pedolo even if these are seemingly well fitted to -- gists of The Bureau of Soils proved that a num I hope that the enumerated experiments with ber of unproductive soils contain organic toxins native Greenland plants will show that plant (cf. literature cited in OSVALD op.c. p. 303). dispersals and migrations, so willingly a.nd so Hecently OsvALD (op.c. and 1949), by experi liberally assumed in every paper of plant mental means, investigated the properties of geography, may be quite different in nature these toxic substances. His experiments were itself and that plants embraced by th e terms occasioned by the observation that rape (Bras hemerophilous and hemerophobous may possess sica napus L. f. oleifera Metzg. and Br. rapa . properties, governing their different responses, L. f. oleifera l\:Ietzg.) germinated and developed whi eh are still unknown.>> very poorly among Elytrigia repens, although It is a pity that PORSILD did not carry out there could be no question of competition · 116 SELANDER: Floristic Phytogeography of South- Western Lule Lappmark
(shading) from the thin patches of couch-grass. tains this behaviour is characteristic not only Previous observations indicated that other of the few species belonging to the ephemeral grasses, especially Avena elatior, Da(;tylis glo- plant communities on new soil, but also of the merata, Festuca rubra, and F. trachyphylla, had _ far more numerous species which are restricted a similar effect upon clover and diverse weeds. to more or less naked mineral soil. They 'l�he experiments proved that an extract of are lacking not only in communities where the couch-grass stolons, provided the concentration plant cover is closed, but in very thinly covered is high enough, inhibits the germination in meadows and grass heaths as well. The plants rape, oats and dandelion; at moderate to rather most strictly confined to mineral soil, e.g., Sagina high concentrations, the toxic substance stimu caespitosa, Braya linearis, Arenaria humifusa, lates the growth of mould fungi, e.g., Mucor A. norvegica, Carex nardina, C. glacialis, most and Penicillium, which kill seeds not or only Drabae, Potentilla nivea, Arnica alpina and partly killed by the toxin (OSVALD 1947 pp. Oxytropis lapponica, seem able to tolerate the 292 ff. ). The experiments were, however, not presence but of very few commensals, and of above criticism. They proved only that the grasses usually not at all. In other words, at couch-grass stolons contain the toxin, but least in the case of the plants most exclusively not that it is actually exuded from them. There adapted to mineral soils, (a category embracing fore, the experiments were repeated with soil most centric species supposed to be still spread from a lawn of Festuca rubra which kept remark ing) , even open or half-open plant communi ably free from weeds. An extract of this soil ties may be regarded as being closed whenever proved to have the same effect as the extract they include grasses (or, apparently, certain of couch-grass stolons (OsvALD 1949). other graminids when they appear in great The effect of the toxic substances on ger masses). This can, possibly, be explained from mjnation is strikingly similar to that _ of phyto the effect of the root exudations of the gramin hormones and hormone derivatives. The reason eous plants. why certain species are resistant to treatment Should the theory propounded by DE CAN with hormone derivatives is probably that they noLLE and OsvALD stand the test of closer exam themselves produce similar substances in ination, it will offer a simple solution to many fairly large quantities, and, in consequence, are phytogeographical problems. Thus the presence, accustomed to them. Susceptible plants, on or absence, of a species within a certain area the other hand, probably produce such sub would often be dependent on the time of its stances only in small quantities, or such of arrival there, i.e. whether the present plant other types (OSVALD 1947 p. 299). communities were already on the spot or, at OsvALD himself calls attention to the wider any rate, whether certain species had safeguarded applicability of his observations, as follows: their dmnains by )>poisoning)> the ground to )>Many of our wild plants occur, e.g., in the later arrivals. For instance, the limits of alpine mountains and on shores, exclusively on new plants belonging to the centric groups would soil, laid bare owing to one cause or the other. be explained simply by an assumption, that But whenever any grass species invade the lo they had come thus far, but no farther, when cality, the other plants disappear, even before the closed plant communities were established the grasses' having formed a close stand. Al on the ground laid bare by the retreat of the though nothing is known about the reaction of inland ice. The fact that occasional long such species to growth substances, we have prob distance dispersal seldom has lasting results ably to do, also in such cases, with an effect of and, in consequence, the untouched plant com the root exudations of the grasses.)> (Op.c. p. munities are as stable as, in fact, they are would 300; transl. from the original Swedish text.) be satisfactorily explained. Light would also To this should be added that in the moun- be thrown on the seemingly strange fact that Distributions within the Region 117 extremely rare relicts, which are considered to getting choked up, then proceeds by way of be devitalized and >>senescent>>, thrive excellently plant communities of gradually increasing xero when transplanted into botanical gardens, where phily towards heath communities rich in lichens, . they grow alone in not >>poisoned>> soil (cf. and ends up in the regeneration of the lakes in MARIE-VICTORIN 1938 p. 553). Some conse wind-shaped erosion hollows. His results were quences of the theory will be dealt with be confirmed by the investigations of BJORKMAN low; I shall restrict myself here to emphasize & Du RIETZ (1923) and Du RIE'IZ (1921, 1925). that plant-sociological and plant-geographical To the centric plants, which hardly ever research has no more urgent task than to inves occur in swamps� these events are, however, tigate the reaction of plants to foreign root of no significance whatever, except perhaps exudations. Until such investigations have in so far as they may deprive them of a fraction been carried out, we cannot proceed any further, of their habitats. As to the communities to along the way indicated by OSVALD's experi which these species belong, FRIES stated (op.c. ments. p. 256): >>Bei den alpinen Assoziationen in der However, plant migrations, and changes Heide- und Wiesenserie haben wir bisher kein in the composition of the flora, have, as a mat einziges Beispiel einer definitiven Veranderung ter of course, taken place. But the arctic and einer Assoziation nach irgend einer bestimmten subarctic regions are no exceptions from the Rich tung hin wahrnehmen konnen>> ( cfr. also rule that migrations are mass phenomena. Du RIETZ 1930a pp. 348 ff., NORDHAGEN 1928 Thus, HADAC (1948 p. 22) calls attention to the pp. 526, 567). great importance of synergetic relations be TENGWALL ( op.c. pp. 7 46 ff.) describes the tween the members of arctic plant communities. development of the vegetation in recent deltas; Even here, phytocoenoses hardly change to these ephemeral communities are, however, any noteworthy extent, except in connexion of no interest in this connexion. On the other with a change of the habitat, resulting either hand, he discusses (ibid. pp. 7 38 ff.) a phenome from altered external conditions or from the non that doubtless produces changes of the activity of the plants themselves. Chief among kind required by FRIES, viz., the leaching of such changes are, of course, the climatic ones. the surface layers of the soil and the resulting However, the climate of Europe has hardly decrease in pH. TENGWALL exemplifies this changed at all since the 13th century (ANGSTROM course of events with observations made, on 1946 pp. 94 f.). In all likelihood, no major the one hand, on Mt Pelloreppe in Sarek, where change has occurred after the subatlantic dete the interspersion of somewhat calcareous amphi rioration, i.e. for about 2600 years (ibid. p. 92). bolites increases the fertility of a soil originating The chances are, therefore, that the climax from acid syenite rock, and on the other hand, communities of the Fennoscandian mountains in subalpine areas of calcareous solifluxion soil have remained essentially unchanged ever since near Abisko. In both localities, the develop then. ment proceeded from Dryas communities · to The first investigations into the succession wards Empetrum-heaths rich in mosses. (For of plant communities in the mountains of instances of a similar development of plant Lapland were made by TH. FRIES (1913 pp. communities in the Alps, cf. BRAUN-BLANQUET 235 ff.) and TENGWALL (1925 pp. 738 ff. and & JENNY 1926). 7 46 ff. ). The results of later investigations The phenomenon doubtless is generic, even have mostly not yet been published. }""' RIES though it characterizes mainly sueh communi proved that a transgression of the Sphagnum ties as are strongly humus forming, i.e. more or bogs occurs locally, chiefly in the conifer belt, less humid and, usually, rich in mosses. Bow while above the conifer limit a cyclic develop ever, leaching doubtless takes place even ment of the marshes begins with small lakes in dry scrub heaths where very little humus is 118 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
formed. In our region, the glaci-fluvial ridges processes acknowledged by l:Ainr (1930) to north of the river Puolejokk may serve as determine the succession of plant communi instance. They are occupied by an Empetrum ties in the Alps, viz., deflation (>>Abflachung heath rich in lichens with Betula nana, Arctosta des Bodens>>) and maturing of the solums phylos alpina, Vaccinium uliginosum, Cala (>>Bodenreifung>>, i.e. increase of acidity, humus magrostis lapponica, &c., which on the windy accumulation, formation of raw humus, &c.), crests is replaced by a Diapensia lapponica which all tend to produce trivial dwarf scrub Loiseleuria procumbens-heath with Luzula confusa heaths of a type predominant on the low, &c., i.e. by communities that definitely indicate rounded mountains of the Fennoscandian areas a soil poor in lime. However, in wind-eroded of primary rock. hollows only a few dm deep basophilous plants SMITH (1920 pp. 142 f.) appears to presume are predominant, e.g. , Roegneria scandica, Ko a reversion of those processes in the grounds bresia myosuroides, Carex rupestris, and Oxy that, in the time of the postglacial climatic tropis lapponica; thus, merely a decimeter or optimum, were covered with conifer· forests. two below the surface, the reaction of the soil As far as I know, such a development has been has changed sufficiently to satisfy the rather proved nowhere, except, perhaps, locally on high requirements for lime of these species. the crests of windy hills, where dry raw humus On the other band, the summits of a calcareous layers may have weathered away, and in a few phyllite ridge on Mt Kaiseketstjakko, charac low-alpine solifluxion areas. However, at least terized principally by >>wind-heaths>> with baso in our region such grounds would rather seem philous plants such as Carex nardina, Arnica to be occupied by scrub heath communities, alpina, Kobresia myosuroides, &c., are partly oc chiefly composed of Betula nana and Empe cupied by communities of indifferent or faintly trum hermaphroditum and definitely raw humus acidocline species, i.a.,· rather paradoxically, by forming. SMITH's thesis (Le.) that >>right from small >>Sphagnum bogs>> measuring about one the postglacial climatic revolution the spread square meter, with Rubus Chamaemorus, Andro ing of the alpine flora would seem to have meda Polifolia, &c., though no difference in the been going on with ever increasing intensity>> bedrock could be detected. (Attention should (from the original Swedish text) has hardly be called, however, to the occurrence of amphi any foundation in documented facts. And if bolite, &c., in the adjacent mountains, cf. VoGT any contemporary increase in the areas of 1927.) The examples could easily be multiplied alpine plants takes place, the spreading species many times over. Leaching, of course, obstructs doubtless all belong to the most trivial element the spreading of the centric species, if any such of our mountain flora which, moreover, occurs there be, since almost all those species are more more or less regularly also on the pine heaths or less basophilous. of the regio coniferina. This holds doubtless good also of the forma As far as a change in the character of the tion of ± acidic raw humus (>>hedtorv>>, Du alpine habitats and the composition of the RIE'rz 1945 a p. 131, 1945). The changes in the alpine plant communities can be proved, it, alpine vegetation due to the formation of raw apparently, reveals a tendency disadvantageous humus have not yet been investigated. In any to the spreading of the centric species. And case, they cannot be favourable to the dispersal the normally occurring processes favourable of such plants, e.g. , most centric species, as to those syecies, viz., frost-heaving, solifluxion, arf\ adapted to mineral soils and shun humic and wind -erosion, show no signs of increasing acids. On the other hand, this process favours either in intensity · or in frequency and exten the dwarf scrubs, thus trivializing the alpine sion; a probable exception will be dealt with vegetation. This would seem to apply to all below (pp. 122 ff.). Distributions within the Region 119
The flora on unstable ground proper, though present on talus slopes in the conifer belt, e.g., Anthyllis Vulneraria var. lappo .As long as the climate does not change, nica on Mt Farforita or Minuartia rubella on Mt new habitats suited to the centric plant species Njarkavare north of the river Stora Lule .Alv, are created only by rare catastrophes such as were of necessity brought to those talus slopes land-slips and the draining of lakes; or else such by long-distance dispersal from the high moun habitats are to be found in localities where tains after the forming of the pine-forests. edaphic conditions prevent the forming of a Certain facts, however, speak against an stable plant cover. assumption that such spreading would go on In the mountains this holds good, e.g., of to any appreciable extent nowadays. The the ever-changing deltas of the sub-glacial composition of the flora of the talus slopes streams and of alluvia on brooks, where the vege seems to be more stable than might be expected. tation may assume an entirely different charac Newcomers are probably very rare; on North ter, should the brook stabilize or change its Swedish mountain bluffs, I have observed but course (cf., e.g., TENGWALL 1925 pp. 738 ff., one likely case of new-colonization, viz., Crepis N ANNFELDT 1940 p. 35). However, in our tectorum on Mt Fallforsberget near .Alvsbyn region hardly more than one centric species (SELANDER 1949 p. 117). No opposite number occurs normally in such habitats, viz., Carex was noticed in the Kvikkj okk region. A.mong bicolor which, moreover, has its >>primary>> the indubitable anthropochores, none were stations in localities of an altogether different found there on the talus slopes; even Luzula kind. pallescens, whieh is generally a neophyte The talus slopes with their recurrent slides in its Swedish stations, would seem to be native are apparently more significant. .A compara in these localities. tively large number of centric species are more On the other hand, several species, though or less characteristic of the talus slopes, in our evidently ancient inhabitants of the region, region, above all, Woodsia glabella, Roegneria occur but on one mountain or two. This applies mutabilis, _Zlfinuartia rubella (chiefly on slopes to immigrants, as well as to probable glacial facing east), Arenaria norvegica, Draba hirta, survivors. A glance at the adjoining table Braya linearis, Potentilla Chamissonis, Oxy of the floras of some typical south-facing bluffs tropis lapponica, Primula scandinavica and and talus slopes (Table 1 ) will show that, but Antennaria lapponica. It is, of course, conceiv for a few species, e.g., Cystopteris fragilis, able that these species, and similar ones, would Cerastium alpinum, Angelica Archangelica, spread on the >mew soil», laid bare by earth and Campanula rotundifolia, the plants of these slips. localities occur only in a limited number of Evidently, such dispersal has in fact taken stations, which are often more or less casually place even after the immigration of the coni scattered about. Moreover, it should be noticed ferous trees. The talus slopes could obviously that most of the plants restricted to one or two not be formed before frost-weathering, &c., mountains, e.g., Asplenium scptentrionale, Dryo bad been active for a considerable time after ptcris spinulosa, Roegneria mutabilis, Staohys the melting away of the inland ice; and by that silvatica and Hieracium Pilosella, are rather tirne, the conifer forests had occupied the ground plentiful in these isolated stations and some at least round the eastern localities of this times even found in masses (cf. Pl. 4). Such kind that are situated just on the ice-divide, a behaviour is characteristic also of many where remnants of the inland ice remained far species that either display large gaps between into the warm .Aneylus time (cf. LUNDQVIST their stations (cf., e.g., Part II maps No. 36 1943 map 1). Consequently, at any rate such and No. 331) or appear only within strictly lirn alpine species as never occur on the precipices ited areas, e.g. Polystichum Lonchitis, Potentilla 120 SELA.NDER: Floristic Phytogeography of South-Western Lule Lappmark
Table l.
Cent1·ic Glacial survivors ...... Anthyllis vulneraria v. lapponica . . . . . + + + + + + D1·aba hirta...... + + + + + + + + + + + + Drab a norvegica v. nammatis ...... + + + + + W oodsia glabella ...... + + + + + Potentilla Chamissonis ...... + + + + + + H ieracium kuusamoense ...... + + Oxytropis lapponica ...... + + + + + + + Primula scandinavica ...... + + + + + Braya linearis ...... + + + + Roegneria mutabilis ...... + Lotus cm·niculatus v. borealis ...... + + Gentianella a urea ...... +
Ubiquitous Glacial sn1·vivo1·s. Carex o1·nithopoda ...... + + + + + Saxifraga Cotyledon ...... + + + + + + + + + W oodsia ilvensis ...... + + + + + + + + + + + Erigeron politum ...... + + + + + + + + + + + + Aconitum septentrionale ...... + + + + + + + + + + + + + + + + + Sedum annuum ...... + + + + + + + + + + + + + Myosotis silvatica ssp. frigida ...... + + + + + + + + + + + + Silene rupestris ...... + + + + + + + + + + + + + + Viola montana ...... + + + + + + + + + + + + + H ackelia deflexa ...... + + + + + + + + + + + + Epilobium collinum ...... + + + + + + + + + + + + Erysimnm hiemciifolium ...... + + + + + + + + + + + + + + + W oodsia alpina ...... + + + + + + + + + + + + + Alchemilla filicaulis ...... + + + -I + + + + Arabis hirsuta v. subalpestris ...... + + + + + + + + + + + Veronica fruticans ...... + + + + + + + + + + + + + + + Cystopteris fmgilis ...... + + + + + + + + + + + + + + + + + + + + Cerastium alpinum ...... + + + + + + + + + + + + + + + + + + + + Angelica Archangelica ssp. norvegica .. + + + + + + + + + + + + + + + + + + + + Campanula rotundifolia ...... + + + + + + + + + + + + + + + + + + + + Saxifraga nivalis ...... + + + + + + + + + + + + + + + + + + Urtica dioeca ssp. gracilis ...... + + + + + + Lactuca alpina ...... + + + + + + + + + + + + Ribes spicatum ssp. lapponicum ...... + + + + + + + + + + + + + Saxif1·aga adscendens ...... + + + + + + + + + + + + Car ex capillaris ...... + + + + + + + + + + + + + + + + + + Polystichum Lonchitis ...... + + + + + + + + + Rumex tenuifolius ...... + + + Asplenium viride ...... + + + + + + + + + + A renaria norvegica ...... + + Sedum acre ...... + + Erigeron boreale ...... + + + + + Epipactis atrorubens ...... + + Aiuga pyramidalis ...... + + Viola rupestris ...... + +
North-eastern immigrants Actaea erythroca1·pa ...... + + + + + + Rosa maialis ...... + + + + + + + + Daphne M ezereum ...... + + + + + + + + + Luzula pallescens ...... + + + + + + + Stellaria longifolia ...... + + + + + + + + + + dalix coaetanea ...... + + + + + + + + + + + +
Southern immigrants Platanthera bifolia ...... + Pyrola chlorantha...... + M yosotis stricta ...... + H ieracium Pilosella ...... + Potentilla argentea ...... + + + + + Distributions within the Region 121
Table 1 continued
Turritis glabra ...... + + + + + + + Veronica officinalis ...... + + + + + + Barbarea st1·icta ...... + + + + + Ca1·ex pallescens ...... + + + Fragaria vesca ...... + + + + + + + + + + Galeopsis bifida ...... + + + + + + + + + + Rubus idaeus ...... + + + + + + + + + + Stellaria graminea ...... + + + + + + + + + + + Carex Oederi ...... + + Circaea alpina ...... + + Viola mirabilis ...... + + + + Asplenium Trichomanes ...... + + Poa palustris ...... + + Draba nemorosa ...... ( + ) Stachys silvatica ...... + Dryopteris spinulosa ...... + + Prunella vulgaris ...... + + Polypodium vulgare ...... + + + + + + + + + + + + Luzula pilosa ...... + + + + + + + + + + + + + Dryopteris Filix-mas ...... + + + + + + + + + + + + + Valeriana sambucifolia ...... + + + + + + + + + + + + + + Roegneria canina ...... + + + + + + + + + + + + + Botrychium Lunaria ...... + + + + + + + + + + + + + Anthriscus silvestris ...... + + Lastrea Robe1·tiana ...... + Erigeron acre ...... +
Species of doubtful provenance. Asplenium septentrionale ...... + Ribes spicatum ssp. scandicU?n ...... -1- + + -1- + + + + Carex flava ...... + + + A1·ctostaphylos Uva-u1·si ...... + + + + + + + + + , - + + + + +
The figures signify: 1 Mt Farforita; 2 Mt Tarfek; 13 Mt Jalle on Sarjasjaure; 14 Mt Tuke ; 15 Mt Unna. 3 Mt Sappekvare; 4 Mt Ramek; 5 Mt Nammatj ; 6 Mt Titer and Mt Stuor Titer; 16 south steep below Mt Kassavare ; 7 Mt SniUkok and Mt Tavvevare; 8 Mt Tjalta; Allak; 17 south steeps on Kollomoloki ; 18 Mt Aranaive; 9 Mt Njunjesvare ; 10 Mt Hilto and Mt Sammartj akko; 19 Mt Tjargesvare ; 20 Mt Unna Tjargesvare and adja 11 Vuoksapakte on Mt Tjuolta; 12 Mt Luoppalvaratj ; cent mountains.
Chamissonis, A nthyllis Vulneraria var. lappo .appreciable spreading from mountain to moun nica, Lotus corniculatus var. borealis, Viola tain has taken place in recent times, seeing mirabilis and V. rupestris. that the stray stations of southern species. Nor is it, for the most part, possible to dis on talus slopes outside their normal ranges cover any edaphical differences between locali have an unmistakable relict character and the ties where a given species is found or missing. centric species generally do not possess any For instance, Sedum acre was observed only pioneer stations at all. in two places within our region, viz., Kollo As to the causes of this I cannot express. moloki and 1\it Tjargesvare. On the interjacent any definite op1n10n. The gaps under dis Mt Aranaive, where practically all its commen cussion may partly be due to the disappearance sals are met with, it is lacking, although the of some plants from former stations, when the south-bluff of this mountain, with its excep climate grew less clement and the habitats. tionally rich flora, would seem to offer excellent changed in an unfavourable way owing to habitats. The distribution of the forementioned leaching, &c., while the forest invaded the talus. species is probably determined largely by chance slopes (cf. above p. 105). Further, it shoLJld only; furthermore, as far as can be judged, no be remembered that the relative distance be- 122 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark tween the >>living>> talus slopes is at present, more (Concerning the contemporary climatic amelio often than not, ten miles or so, and that inter ration, cf., e.g., SANDBERG 1940 and, partic jacent rallying-points are usually lacking. ularly, several papers by AIILMANN, e.g., 1948.) The chances that a disseminule should be carried There is probably, within our region, not a as far as that by a wind which, moreover, gener single mountain where a close inspection would ally follows the valleys, i.e., is parallel to the not reveal patches or large areas of such >mew slopes, and happen to land on a talus slope a soih. few hundred yards long, are obviously exceeding Unfortunately, I have not studied its vege ly small, perhaps one in a hundred thousand tation by far as thoroughly as it deserves, or thereabout. Since, further, only an infini since I did not arrive at an understanding of tesimal fraction of the disseminules survive, the true nature of these grounds before my the probability that a population of a few hund field research was almost finished. My notes red or thousand individuals should leave pro are, therefore, very incomplete and rather geny on a talus slope some ten miles away unsatisfactory, i.a., because they were originally appears to be practically non-existent. made for entirely different purposes. Never Almost the on1y plants that may succeed theless, after much hesitation, I decided to set in this are those spread by birds breeding in forth a few· facts, solely in the hope to animate precipices, i.e. by ravens, which disseminate some botanist, visiting the high mountains of the berries of many species, and by eagles and Lapland, to devote himself to a thorough falcons, which doubtless disseminate Hackelia going investigation of these interesting phe deflexa. Hackelia, as well as the species bearing nomena. berries, e.g., Actaea erythrocarpa, Daphne M eze Considerable areas where the perennial snow reum, Arctostaphylos Uva-ursi, &c., occur actu had recently melted away were observed on ally on all, or almost all, talus slopes within five or six places, viz., south of the eastern of .certain climatically or edaphically conditioned the two lakes called Staddajaureh; on the west limits. ern part of the high plateau of Mt Stadda T�e data concerning· the changes in un tjakko; on the west-slope of the northern summit disturbed alpine plant communities may be of Mt Kerkevare; north of the glacier of .Almal summed up in the opinion expressed by Lti"DI lojekna; on the summit and northern slope {1930 p. 588) as follows: >>Im allgemeinen gehen of Mt Kasakpuolta; and probably round the Vegetationsveranderungen im Gebirge, soweit summit of Mt Allak. According to the ordnance .sie iiber Kleinverschiebungen, die innerhalb map, there should be three good-sized glaciers des Rahmens einer Assoziatiori bleiben, hinaus within our region in addition to Rakokjekna, wachsen, vicl langsamer vor sich, als man bei .Almallojekna and the big glaciers of Mt Suli der ersten Betrachtung der reich gegliederten telma, viz., one on Mt Kaiseketstjakko, one Gehange glauben wiirde, und die Vegetation on the western part of Mt Staddatjakko, and ist in weitgehender Harmonic mit den lokalen another on Mt Jeknaffo near its southernmost Umweltfaktoren, die durch Gelandebildung ge end. As a matter of fact, there are only a small schaffen und erhalten werden.>> glacier on the eastern side of the summit of 1\H Kaiseketstjakko and two, a little larger
'The vegetation on ))new soil» ones on the eastern and south -eastern side of Mt Jeknaffo, none of them, however, being In present times, new-colonization on a by far as large as indicated by the map; the large scale does not take place in the regio >>glacier>> on Mt Staddatjakko is non-existent . .alpina but on the >mew soil» laid bare, during The explanation is, of course, that the glaciers the past decades, at the retreat of the glaciers of the map have, for the most part, actually and the melting away of perennial snowfields. been perennial snowfields; fresh moraine was Distributions within the Region 123
Fig. 22. The crest of Mt Kasakpuolta, c. 1250 m s. m. In the beginning of the 19th century, these parts of the mountain were covered by the glacier of Almallojekna (visible in the background). The striae on the rocks may, therefore, partly be of a recent origin. The patches of gravel are sparsely colonized by Draba alpina, Dr. lactea, Ra nunculus glacialis, Cerastitmn arctic-um, Sagina intermedia, Pedicularis hi1'suta, Phippsia algida, &c. Photo Sten Selander 21.7.19 46. nowhere observed in the former places of these The new soil is everywhere of about the same )>glaciers)>. During the seventy years passed kind: water-soaked, packed, almost bare gravel. since the mapping the snowfields ha.ve disap On Lake Staddajaure, the gravel is coarser peared. Similar is the case with Almallojekna, than in the other localities and looks more )!the largest gla.cier of Sweden)>, which seems to like the surface moraine of a dead-ice. As a consist of several minor glaciers connected by matter of fact, there is probably little differ snowfields that have now partly melted away; ence between a dead-ice and a perennial snow at least two good-sized )munatakks)>, not indi drift, situated, as the present one, in a hollow cated on the map, have arisen (see Pl. 11). and more or less glaciated in its lower strata. On Mt Staddatjakko square miles of )mew Since this area of )mew soih> lies at a much soil», which is not even interrupted by any par lower level than the remaining ones and its ticularly large snowpatches, extend where the vegetation, consisting mainly of Carex Lache glacier should have existed. nalii, Ranunculus glacialis and Pedicularis hir- 124 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark suta, is not at all similar to that observed in the soil. True, such plants as usually predomi other places, I shall not dwell on it in the nate at high levels, e.g. , Lycopodium Selago, following. Luzula confusa, Ranunculus pygmaeus, Carex l;arge areas on Mt Kasakpuolta and near Lachenalii and (sterile) 0. Bigelowii, are not the glacier of Almallojekna are occupied by altogether lacking anywhere but at Almallo barren boulder-land. In the beginning of the jekna, though they never attain their usual 19th century, Mt Kasakpuolta would seem to frequency. Instead, a few other species, have been completely covered by the glacier, generally differing from one mountain to an which w:;�s then far larger than now (cf. WAH other, appear in supernormal abundance. As LENBERG 1808 pp. 32 f., pp. 35 ff., and the map is well known, this is a common phenomenon tab. 1); the boulders may, therefore, partly on new soil, not least near glaciers and snow be fresh moraine, and the striae ·on the rocks fields. Thus, HuLTEN (1937 a p. 24) reports, may partly be of a recent origin (cf. fig. 22). that the ground laid bare at the retreat of Solifluxion has large proportions, at any rate the Mendenhall glacier in Alaska was colonized in places. When the snowfields melted away only by a few species, chiefly L,upinus noot and the ground was exposed to the action of kaensis Donn. In the Virihaure mountains the atmospheriles and the fluctuations in tem it has, however, to be added that these species perature, new masses of earth evidently started arc otherwise more or less rare and usually moving down the mountain sides; in this way, show a low frequency in their stations. new habitats are likely to arise, even for some On Mt Staddatjakko Draba alpina pre of the species foreign to the areas newly freed dominated, with millions of specimens, and from the perennial snow. at the time of my visit, during the last days The vegetation on the new soil consists of of July 1942, adding a faintly yellowish tinge high-alpine snow-bed communities of an extreme to several square miles of ground. The rest character, usually of the type named by SMITH of the vegetation was relatively rich in species, (1920 p. 51) >>Snow-bed sociation rich in herbs>> though only Ranunculus glacialis, R. nivalis, (>>ortrikt snolage>> ); other types occur too, which Saxifraga foliolosa and Pedicularis hirsuta were can be referred to the communities that TH. fairly numerous. FRIES (1913 p. 119) called >>eine extrem hoch Draba alpina was likewise predominant on alpine Variante>> of his >>Ranunculus glacialis Mt Kasakpuolta in 1946, although not to the Wiese>> and >>reine Phippsia algida-Assozia,tion>> same degree as on Mt Staddatjakko. It did (op.c. p. 120). TENGWALL (1920 pp. 397 ff.) not occur, however, above a level of about mentions similar communities under the names 1260 m s.m. Besides a few more common species, of >>Saxifraga-Schneeboden>>, >>Ranunculus gla Phippsia algida, Sagina intermedia and Draba cialis-Schneeboden>> , >>Gerastiurn lapponicum lactea ascended to about the same altitude, Schneeboden>>, and >>Gatabrosa algida-Schnee growing patchwise between the boulders in . boden>>. At least on Mt Staddatjakko also not inconsiderable numbers. Stray individuals the Garex rufina-sociation described by SMITH of Phippsia, Oerastium arcticum, Sagina inter (Le.) and TENGWALL (Le.) is to be found. media, Ranunculus glacialis · and Pedicularis However, as far as I am able to judge, only hirsuta went up to about 1300 m s.m., and from the Phippsia algida -sociation and the Garex there up to the summit, · 1350 n1 s. m., only rufina-sociation appear in a quite typical form sparse stand of Ranunculus glacialis were to on the new soil. Otherwise, the communities be seen. differ from normal snow-bed communities in At Ahnallojekna in 1946, Saxifraga rivularis so far as the number of individuals is abnor was the only vascular plant found on the gravel mally small and, likewise, mostly the number ground, which evidently had become free from of species, the smaller the number the ne\ver snow only in the past few years; the >munatakks>> Distributions within the Region 125 were completely devoid of phanerogam vegeta Such edaphically favoured niches as will tion. afford a vascular plant a chance of completing The new soil on JVIt Kerkevare in 1943 was its cycle of life without succumbing at any characterized mainly by the mass occurrence juncture, are evidently available but sparsely, of Sagina caespitosa. No doubt, this plant and competition within their limited space has its largest Swedish population on this appears to be rather severe. Under normal mountain; it occurred there in thousands of conditions, species with very little compe often luxuriant specimens and, patchwise, titive power take possession of them only to an became almost predominant. Codominant was insignificant extent. Otherwise, it would be Pedicularis hirs�tta; other vascular plants, e.g. , hard to understand why species which, like Salix pola1·is, Cerastium arcticum, Sagina inter Sagina caespitosa, are usually rare and small media, Ran�tnculus glacialis, Cardamine belli in number, are to be found in masses on new clifolia and Saxifraga foliolosa, were of secondary soil and nowhere else. importance, and several species found else Of course, a plant community such as the >>Saginetum caespitosae>> where in similar localities, e.g. Phippsia algida, of l\it Kerkevare is Draba alpina, Dr. lactea and Saxifraga -rivularis, not stable, but only a swiftly transient pioneer were not at all observed. phase in a sere. If this community will not be partly obliterated through the formation of Concerning the conditions on :M:t Allak in new perennial snowfields, its character will com 1939, my notes contain only a statement that paratively soon be changed through an inva Phippsia algida was predominant; also Luzula sion of such more competitive species as, at conf�tsa, Koenigia islandica, Sagina intermedia, the time of the melting away of the snowfields, Ranunculus glacialis, Saxifraga foliosa and S. were growing at too great a distance to be able rivularis were observed. immediately to conquer the new soil. In addition to the calcareous substrate, the It is, in fact, not at all unlikely that the almost total absence of competition appears >>Sagineturn caespitosae>> has already vanished to explain why the vegetation on the new soil by now, since the plant communities on this of the Virihaure mountains consists to such a kind of new soil are of a very short duration. great extent of otherwise rare species. Such FAEGRI (1933 pp. 238 ff.) states that the pioneer p species as have little com etitive ability and, phase of the succession of plant communities consequently, are more or less rare have far on new soil below receding South-Norwegian greater possibilities here than elsewhere to glaciers may come to an end in as short a time assert themselves. as 20 years. Investigations by CoAz (1887) . It may seem that the competition factor at the Rhone glacier, BRAUN-BLANQUET & would be of no importance in the high-alpine JENNY (1926) at the Sesvenna glacier in Switz communities, where the plant cover never is erland, and CooPER (1923) in Alaska indicate closed and considerable areas of bare soil are changes occurring at, approximately, the same always at hand. But the thinness of the vege rate. True, the localities described by FAEGRI tation cannot be due to an insufficient supply and CooPER are situated in the conifer belt, of diaspores, as is evident, i.a., from the mass where the course of development is probably occurrence on new soil of species such as Draba swifter than above the timber line; however, alpina and Sagina caespitosa. If more species the difference cannot be particularly marked. than one occurred simultaneously in as great Evidently, it wa.s largely a matter of chance number of individuals as, e.g., Draba alpina what species would first colonize the areas on :M:t Staddatjakko, vegetation would get a newly freed from snow. Only in one instance, higher degree of covering than it ar.tually ever viz., on :M:t Allak, was the effect of an edaphic attains in the upper regio alpina. factor perceptible. Since the main part of this 126 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark mountain consists of amphibolite, it is easily VESTERGREN 1902 p. 257) seem to spread chiefly explained that the indifferent Phippsia algida by vegetative means. is predominant here instead of the calciphile Myrmecochory is, of course, out of the ques species. On the other hand, the fact that the tion; epizoic dispersal, mainly by reindeer, is new soil is occupied exclusively by Saxifraga probably of little importance, the reindeer -rivularis at Almallojekna, chiefly by Draba only seldom visiting the high-alpine crests. alpina on Mt Staddatjakko and J\ft Kasakpuolta Endozoic spreading by birds cannot take place and largely by Sagina caespitosa and Pedicularis except on a very small scale, since practically hirsuta on Mt Kerkevare, can hardly be accounted no plants with juicy fruits grow in this belt for but by assuming that these particular spe and the snow-bunting (Passerina nivalis), the cies happened to be present there in sufficiently only high-alpine seed-eater, has its haunts great numbers when the perennial snowfield.s chiefly on boulder-land and not on gravel melted away. ground (cf. EKMAN 1944 p. 155). More impor On such occasions distances are probably tance may, perhaps, be ascribed to dissemination more decisive to the flora of the· high moun over the snow in winter by means of tumbling tains than to that of the lowlands, since the disseminules (>>Schneelaufer>>, >>marklopare>>, cf. high-alpine plants usually lack organs for long SERNANDER 1901 p. 11 ff. and p. 355, HEIKTZE distance dissemination. The persistent gales 1914, Du RIETZ 1931); however, diaspores of practically prevent the anemochores from ob this kind also will, in all likelihood, mostly taining a foothold on the summits, the diaspores be blown away from flat expanses such as simply being blown away. Such anemochores these. Most species within the flora of the high aR possess very light diaspors, e.g. , Ericales, alpine gravel ground have heavy diaspores, often Gentianae, orchids, and ferns, belong almost consisting of germinal buds, and usually de invariably to the flora of the low-alpine belt, posed close to the mother specimens. the only important exceptions being Cassiope Consequently, the new soil must have been hypnoides and the Saxifragae. However, Cas colonized mainly step by step, at first chiefly siope, which, moreover, hardly ever grows on by plants from the immediate vicinity; the the summits proper, seems mainly to spread colonizers were, therefore, mostly hardy, though vegetatively. And among the Saxifragae char often uncompetitive and, for that reason, acteristic of the >mew soil» only S. oppositifolia more or less rare species that had been forced is a real anemochore: S. cernua and S. foliolosa to retire to habitats close to .the perennial snow propagate almost exclusively by means of bul where most competitors were unable to live. bils, and S. rivularis, in spite of its abund.ant The colonization of new soil is particularly seed-production, spreads largely through runners interesting as the only contemporary change and subterranean buds. Usually the anemo in the vegetation of the Scandinavian moun chores with comose disseminules, e.g., Dryas, tains that is easily observable and unques Epilobia, Hieracia, and Taraxaca, also come to tionably connected with a change in the climate. a halt on a lower level. In our region, the flora Unlike the other, immensely slow events in of the summits proper hardly includes more the life of the alpine flora, it is largely dependent than four anemochores of this type, viz., .Anten on time. Especially on the assumption that naria carpathica, Erigeron unalaschkense, E. uni foreign root exudations have toxic effects, florum, and Arnica alpina. Furthermore, Salix the competitive power of a species in a given polaris, S. herbacea and Gnaphalium supinum locality must to a certain degree be proportional occur in the upper regio alpina, although mostly to the number of specimens growing there. on the snowbeds of the hollows and rarely on 'l'he more numerous the individuals of a species, fresh gravel ground; moreover, the dwarf Salices the greater its chances of vindicating itself, which, at this altitude, are largely sterile (cf. even though it otherwise may possess but small Distributions within the Region 127
competitive ability. Therefore, the important soil profile vanish and the gravel ground re thing is probably often to be first on the spot covers its ancient character. (Cf. the soil pro and thus to get the start of competitors. files from low-alpine snow-beds in G.JAEREVOLL This should apply to all periods of climatic 1949 with their surface layers of raw humus.} revolutions and more especia.Uy to the last At the beginning of a climatic deterioration, phase of a glaciation, when the inland ice melted most high-arctic plants therefore pass through away. The conditions on recent >mew soil» a critical time. present a miniature of those prevailing during About the year 1750, the glaciers all over finiglacial time. Also then, the new soil must, Europe reached their greatest extent since at least here and there and now and then, have the ice-age (cf. :FAEGRI 1933 p. 12), and the real been invaded by uncompetitive species occurring regress did not begin until a hundred years in masses; and then as well as now, it must often later. In the beginning of the last century,. have been a matter of chance what species the alpine flora, thus, still felt the after-effects would first establish themselves in a habitat, of this climatic deterioration. For instance, thus taking the lead and getting better prospects according to WAHLENBERG (1808 p. 33), the of holding their own, when stronger compe ice in the western part of Lake Vastenjaure titors came crowding on. At least in the case did not break up, in 1807, until the beginning of some species, e.g., the Papaver races, 8tella of August, i.e. about a month later than now ria crassipes, the Arenaria species, some Drabae, adays. The reason why WAHLENBERG, LAESTA &c., the explanation of their present, irregular DIUS and N. J. ANDERSON overlooked, e.g., and scattered, distribution is probably to be 8agina f.Jaespitosa, Draba crassifolia, and Poten sought for in events such as those suggested tilla hyparctica was, no doubt, not only the above. partiality of the earlier botanists for the lower However, the boom for certain high-alpine mountain slopes, but also the very low fre plants during the first phases of a climatic quency of the forementioned species in those amelioration is doubtless only temporary. The days. Nor can a glorious future be predicted new soil will soon be occupied by a flora of for those plants, when the present temporary a more low-alpine character, and the high recovery comes to an end. alpine species will be driven away to gradually For the present we can, however, only express diminishing areas round the summits: the final vague suppositions on these matters. None outcome will be retrogression. This also applies the less, this much seems certain. Most ele to the more low-alpine, rare species,. having ments of the Fennoscandian alpine flora have difficulties in finding new habitats, when their long ago reached all the localities where they old ones are rendered uninhabitable by a cli can live and vindicate themselves under present matic change. conditions, provided they have not been barred Should the climate, then, again deteriorate, by particularly wide stretches without any the lowering of the region limits will produce suitable habitats. Among the centric species, practically no improvement in the conditions at any rate the high-alpine ones do probably of the high-alpine species, anyhow, not imme not widen their ranges at all. On the contrary, diately. The increase of glaciers and snow they seem to have struggled, for a long time past, fields deprives them at once of part of their simply to keep their modest positions. Our high-alpine habitats, while their former habitats rare alpine plants appear to become still rarer further down are still occupied by more or less with every change in the climate, irrespective closed plant communities, where plants bound of its character; during the intervening spells, to naked mineral soil are excluded. These com they probably remain stationary or, possibly, munities disappear but little by little, and it will undergo a slow decimation from deterioration take a still longer time till the changes in the of the habitats. 128 SELANDER: Floristic Phytogeography of South-Western Lule Lapprnark
»Fossil» ranges pronouncedly >>aggressive>>, while the Green landic is >>conservative>>. Many phytogeographical facts would appa Nor can a given species, therefore, be expected, rently be conveniently explained on the basis everywhere and always, to react in the same {)f a supposition that, in the history of every way, for instance to a climatic change, · still plant species, such spells of stationariness have less two different ones. Consequently, great alternated with spells of dispersal, coinciding caution is needed when discussing the inter with periods when, for some reason or other, relationship of plant ranges and climatic the environment underwent great, far-reaching changes or other past phenomena. Still, the changes (cf., e.g., GLEASON 1924 pp. 543 f.). future of many species was, probably, largely At present, such a change is brought about settled by the events of the final phase of the by the exploitation of the earth by :Man. One last glaciation. Such plants as Stellaria crassi particular category among the plants is, as pes, Arenaria hurnif�tsa, Potentilla hyparctica, a result, gaining ground everywhere, viz., the Papaver Laestadianurn, P. lapponicurn, Draba anthropochores. This change is unique in so crassifolia, Sagina caespitosa, &c., could scar far as its scope is world-wide and its course cely have reached the few, very scattered and everywhere proceeds in the main similarly, characteristic habitats, where they are able more or less irrespectively of climate and other to maintain themselves against competition, local conditions. However, this or that species after the time when the plant cover of the wide, is not affected in the same way, in the different interjacent areas closed up. As far as can be parts of its total area, by the changes occasioned judged, they spread almost exclusively during by human activity. Thus, in Greenl3Jnd the the period when the ground, uncovered by the plants displaying hemerophilous tendencies are wasting away of the inland ice, still remained {)nly partly the same as in Pennoscandia. more or less bare and an uncompetitive species Several species stated by PoRSILD (1932 pp. was not, when migrating, everywhere confronted 24 ff.) to be hemerophilous in Greenland, e.g., by superior competitors. Cerastiurn trigynurn, Ranunculus pygrnaeus, Saxi As established by MANNERFELT (1945, e.g. fraga cernua, S. foliolosa, and S. groenlandica, p. 10 ff., p. 116 ff., p. 163, &c.), the melting are, as far as my experience goes, definitely inland ice was, during fini-glacial and early hemerophobous even in those parts . of Fenno postglacial times, in the main a gigantic dead scandia, for instance, northernmost Norway, ice. The ice-cap, therefore, melted away first where they occur close to human habitations. where i� was thinnest, i.e. on the summits, On the other hand, some plants that are strongly which cropped up out of the ice sheet, forming hemerophilous with us, e.g. , Carex rnacloviana, nunatakks surrounded by nunatakk lakes. C. canescens, Charnaeneriurn angustifoliurn, and Later, when the high massifs were largely free Geranium silvaticurn, are explicitly described from ice, the valleys were still filled with dead by PORSILD as not favoured by human activity, ices and ice-locked lakes. In all likelihood, as far as Greenland is concerned; the same evi the high-alpine plants spread, chiefly or exclu dently applies to the Alchernillae in their Trans sively, during this stage of the wearing out atlantic stations. In some cases, such behaviour of the inland ice. This hypothesis, i.a., explains within small populations living close on the why, ceteris paribus, the big mountains and boundaries of the range of the species is doubtless the hills immediately surrounding them are due to impoverishment of the biotype com much richer in rare alpine plants adapted to pound. However, this cannot always be t.he ex mineral soil than the table-lands and the iso planation. For instance, Carex rnacloviana is of lated, lower mountains. These plants probably undoubted An1erican provenance; . in . spite of established themselves on the high mountains this, the isolated Scandinavian population is as . early as at the nunatakk stage and then Distributions within the Region 129
followed the ice on its retreat down the slopes. Yet even among the high -alpine plants there As the remaining dead-ice in the valleys and are not a few ubiquitous, comparatively aggres on the tablelands n1elted away and the ice sive, species, e.g., Lycopodium Selago, Carex
locked lakes were drained, · they were able, Lachenalii, Luzula confusa, Ranuncul'ltS glacialis, partly, to colonize the nearest hills; but being &c., which got ahead of their conservative, slow starters and poorly adapted to a climate slow-spreading commensals, whether the desti as warm as that of the early postglacial period, nation was distant areas, e.g., Sarek, or the they could not proceed very far, owing, i.a., wide extents of land nearer the starting-point to the competition from invading low-alpine that suddenly became available for colonization, species. It should be kept in mind that our e.g., at the draining of the ice-locked lakes region did not begin to become free from ice and the melting away of the huge remnants of until the end of the Y oidia time or the beginning dead-ice on the tableland west of Sarek. When of the .Ancylus time, round about the year 7000 the slow-paced species arrived there, the first B. C. (cf. A.HLMANN a.o., 1949 p. 8), i.e. but
.. Potentilla 0 Arenaria � Luzula arctica � Sagi�1a caespitosa hyparctica humifusa
- Minuartia rubella � Pedicularis flammea ...... Carex nardina
Fig. 23. Areas of some arctic plant species.
The ranges within our region of the rare ities within a larger area, where their specific arctic plants seem, largely, to be of this kind. wants were satisfied. Nor these plants did, Their limits generally suggest rings round a however, reach very far from their starting stone thrown into water, with the mountains points. north and north-east of Mt Sulitelma forming .An especially interesting instance of such the centre (cf. map fig. 23). Evidently these >>fossil>> limits may, perhaps, be found in one species managed to occupy areas of a size particular part of the region. Attention has varying with the degree of biotypic depaupera been called above (p. 110) to the strange fact tion during the glacial period. Those most that Campanula 'ltniflora is practically absent depleted of biotypes, e.g. Arenaria humifusa, from the low mountains between the river Potentilla hyparctica, and Luzula arctica, were Puolejokk and the river Mellatno, viz., Iltom, unable to advance beyond a very short distance. Lemakvare, Unna Titer, Stuor Titer, and Aras .At present they can assert themselves only vare, as well as from the area round the lower under conditions so unusual as to be found but course of the river Stalok, including Mt Piete, exceptionally, at a few points within their although suitable habitats abound and Carn potential areas. Other species with narrow panula is not uncommon in the surrounding ecological amplitudes, e.g., Carex nardina, C. mountains. The same applies to several other bicolor, Draba crassifolia, and Braya linearis, plants. Poa herjedalica, Festuca rubra v. mutica, did not lose their spreading capacity to the Cm·ex bicolor, Minuartia rubella, Arenaria humi same extent, being able to occupy most local- fusa, A. norvegica, Sagina caespitosa, Draba lac- Distributions within the Region 131 tea, Braya linearis, and Cassiope tetragona seem The absence from the forementioned low to be lacking in this area, though they occur alpine area of a few truly high-alpine plants, in the neighbourhood. Others are extremely e.g., Luzula arctica, Draba crassifolia, and scarce; thus, Asplenium viride, Salix polaris, Potentilla hyparctica, is easily explained. How and Saxifraga rivularis were observed but ever, a number of high-alpine species, e.g. , in one station each. Among the alpine species Phippsia algida, Draba alpina, Pedicularis hir that HYLANDER (1941) classifies as more or suta, and Erigeron unalaschkense, are fairly less rare (by a figure of 5 or more), and 5 other common in that area; further west all the plants species (Asplenium viride, Carex rariflora, Jun lacking in southern Padjelanta, except Draba cus arcticus, Saxifraga ri1)ularis, and Astragalus crassifolia and Potentilla hyparctica, grow on norvegicus) that are comparatively scarce in mountains, e.g., Kuobberi, Tuki and the hills our region, only 22.0 have, on an average, been of Kollomoloki, which are about as low, or noted on each of the six above-mentioned even lower. mountains, while the seven surrounding moun The discussed gaps cannot be caused by tains of Tuki, Kuobberi, Stalotjakko, Jalle the substrate, since the mountains in ques on Vejejokk, Eltivare, Allep Alatjakko, and tion are, largely, made up of limestones, highly Mattaive, and the hills of Kollomoloki, on an calcareous phyllites, &c., and abound in cal average, harbour each 37.7 of such species. ciphytes, nor by the local climate, seeing that For various reasons, this difference is not most of the plants occur in all quarters in the altogether clear from the maps that present immediate neighbourhood. The absence of the distribution of the species in question. The Gassiope tetragona may be due to trampling by area between Puolejokk and Mellatno has reindeer (cf. above p. 38); for the absence been thoroughly investigated by TENGWALL of the remaining species, other causes have and the present author, among others, while to be looked for. the mountains north of it, i.e. Mt Huornatj , Now, in early postglacial time, this particular Mt Mola, Mt Arasvaretjakko, and the hills area was evidently occupied by a huge lump round Rapok, have hardly been visited at all. of dead-ice which caused the valleys of the In those parts, as well as in central Sarek, rivers Stalok, Puolejokk and MelH1tno to be to which the same observations in this instance filled with ice-locked lakes (cf. LUNDQVIST apply, the maps, therefore, reveal gaps that 1943 map 1). Shore-lines marking the sites may not actually exist. (Seeing that the fore of ancient nunatakk lakes are to be seen even mentioned mountains consist mainly of amphi as high up as at a level of 1100 m s. m., on bolite, the gaps in those parts, if real, are prob the northern slope of :1\ft Mattaive (LUNDQVIST ably caused by ecological, and not by historical, 1948 p. 162), while the bottoms of the valleys factors�) The apparent gaps east of 1\tl t Jek are to a great extent covered with gravel terraces, naffo are, partly, due to the inaccuracies in eskers, cone-shaped gravel hills, &c., built the ordnance map that includes a non-existing up of hard, rather barren material from the mountain in these parts. Furthermore, a surrounding mountains, and obviously of a single dot, for instance the one signifying Mt glacifluvial or sub glacial origin. West of Mt Latatj, may represent a wide area with several Piete and J alkok, no such formations were stations, while on the other hand, the dots noticed. come close to each other in southern Padje The theory propounded by MANNBRFELT lanta, simply because the mountains there, (op.c.) makes this state of things perfectly though comparatively small, were kept apart comprehensible, though it forces us to turn by the investigators. Consequently, the differ our old conceptions upside down. Nowadays, ence is actually more marked than the maps when the firn line west of Lake Virihaure ap appear to indicate. proximates a level of 1300 m s. m., all the high 132 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark mountains of this neighbourhood are glacia.ted optimum forests are eertain to have grown while, at the end of the ice-age, when the praetieally all over these parts� The real prob firn line was considerably higher, the present lem in obviously how plants that do not grow centres of glaciation, e.g., Mt Almallos, Mt on the high mountains, e.g., Arenaria humifusa, Kuotelis, Mt Rakoktjakko, &c., were free from A. norvegica, Bray a linearis, and Primula scandi ice, the remaining ice-masses being located navica, managed to survive the elimatie opti to the hollows, chiefly to the present Lake mum. It is hardly probable that they went up Virihaure. Further, since the inland-ice sloped the mountains and then down again, sinee they to the west, the ice cover was thinner in the are not known to oeeur anywhere in reliet western part of the Virihaure basin than at stations above their normal present limits, and its eastern end. Thus, the western parts were since the faetors - too intense solifluxion, &c., first freed from ice. Consequently, the pioneer see below p. 151 - that now keep them away plants were able to spread from the ci-de1)ant from these mountains were, presumably, oper nunatakks, e.g., Mt Almallos, Mt Stalotjakko, ative also during the warm period. Also the &c., to lower mountains close by, such as Mt interjaeent areas of bireh woods, willow thiek Kuobberi and Mt Tuki, while the low mountains ets and dense scrub heaths should have formed east of Lake Virihaure were still ice-covered. rather an effeetive barrier. In all likelihood, When, later, the remnants of the local dead-ice they lived, all during the warm period, in or melted away and the ice-lakes were drained near their present stations on precipiees, talus and, as a result, this area became available slopes, serees of dry ealeareous soil, serpentine for colonization, a number of the high-alpine barrens and loeal areas of solifluxion soil, i.e. gravel plants were, apparently, unable to in in habitats where no tree growth was possible. vade it, either because the climate had be (The station of Arenaria humifusa on Mt Tuke come too warm for their successful ecesis or seems to be seeondary, the primary ones, as owing to the overwhelming competition from yet undiseovered, probably being situated on low-alpine plants. the serpentine hilLs of Luokivaratjeh.) The Be this as it may, the fact remains that a aretic epibioties of Newfoundland, the region number of plants are practieally or entirely of St. Lawrenee, and the eoastal region of absent from this area and, further, that for Maine &e., are a ease in point, praetieally all the most part their absence eannot be explained of them living >>exelusively at very low levels, by present eonditions. A eoneeivable explana sometimes near the sea>> (MARIE-VICTORIN 1938 tion is of course that they oceurred here before p. 527), on river gravels, serees, &e., of the the postglaeial warm period, to be ousted as woodland (ef. a�so FERNALD 1907, 1926, 1933, bireh and, here and there, pine invaded the PIERCE 1936). mountains. (Coneerning the earlier oeeurrenee The only hypothesis that seems to fit in of pine in the Virihaure basin ef. SVENONIUS with all known faets is that propounded above. 1900 b p. 485. Subfossil pine trunks were It may seem far-fetehed; yet I am unable to observed by rne only in a tarn below Mt Metjer, suggest a better one. at about 700 m s. rn.; aeeording to the Lapps, Finally, the inability of rare aretie plants sueh trunks were found also on the south slope to intrude into established eommunities may, of Mt Mattaive, · probably at approximately perhaps, be illustrated by a report by SwARTZ the same level.) Even in that instance the (1821). In 1783, he found one single speeimen ranges would, however, be >>fossil>>. And how, of Campanula uniflora on Mt Arasvare, where then, explain that, for instanee, nearly all the it certainly does not grow nowadays. This forementioned plants are to be found on Mt lonely passenger may have been a monument Tuki, and many of them on the low, isolated over an unsueeessful attempt at widening estab hills of Kollomoloki, although during the elimate lished ranges. Distributions within the Region 133
Another instance of such >>fossil ranges>> is, minerals in the soiL Nor should any single perhaps, to be found in the mountains round environmental factor be dealt with separately, Lake Sallohaure. There was a large ice-locked since the factors of the environment act col lake in those parts (cf. LUNDQVIST 1943, map 1), lectively, and the action of any one factor is though as far as I know, the distribution of qualified by the other factors, i.e. the environ glacifluvial and subglacial deposits from the ment is holocoenotic (cf., e.g. , ALLEE & PARK finiglacial time has not been thoroughly investi 1939, CAIN 1944 pp. 16 ff. ). Nevertheless, an gated. In this area, a considerable number attempt at ascertaining the degree to which of plants, mostly those lacking on the eastern the distribution of the plants of our region is end of Lake Virihaure, display gaps or marked determined by geological conditions is neces frequency minima that will be discussed below sary, i.a., in order to form an opinion of the (pp. 140 ff.), although several among them recur part played by other factors. west of the Norwegian border, on J\H Riddoalge Unfortunately, the distribution of the differ and the adjacent mountains (G. BRODAL in litt.), ent rocks of our region is far from sufficiently as well as further north in Vaisa, on Mt Stipok, investigated. No geological map of Lule La pp &c. (unpublished findings by G. BJORKMAN). mark exists, and most investigations available An explanation on the lines suggested above in print date back to the turn of the century. seems rather probable. The broad outlines are known, however, for instance, the dividing-line between mountain
Bedrock schists and acidic, primary rock which is, of course, also the most conspicuous phytogeo Climatic and historic factors determine the graphical dividing-line within the region. general trend of the distribution of plants. Even as regards the distribution of the plants But the details of the areas are largely shaped on both sides of this dividing-line, most ques by edaphic factors, in the Scandinavian moun tions could be accurately answered only by tains in the first place by the nature of the n1eans of pH determinations. Unfortunately, bedrock, i.e. the distribution of basic (or . cir it proved impossible to perform such deter cumneutral) and a.cid rocks. In Scandinavia minations. The transports in the pathless its importance to the alpine plants was pointed wilderness could not be encumbered with in out by A. BLYTT (1869, 1871). In his work struments or soil-samples. This disadvantage on the flora of Helgeland, published as early becomes still more serious with regard to the as in 1912, 0. DAHL classified most Norwegian distribution of plants within the mountain alpine species in regard to their reaction to range with its varying geological nature, partic calciferous rocks. And ever since the pub ularly in view of the fact that the differences lication of the papers by TH. FRIES (1913 are here often rather sn1all. The only massifs pp. 228 ff., 1925), most Scandinavian phyto of primary rock within our region are the glacier geographers have, in fact, concentrated their covered mountains of Rakoktjakko and Kuo attention on the significance of the chemical telis farthest to the north-west, and the acidic nature of the soil, seemingly sometimes to the syenites which are frequent in Sarek (cf. HAM exclusion of other, perhaps as important factors BERG 1910) extend into our region only in single (cf., e.g., TENGWALL 1916, FRODli'{ 1919, PESOLA places, for instance, in Mt Tjalta and the Kables 1928, BRENNER 1930, EKLUND 1931, 1933). massif. Also the seve schists and the amphi The relationship between the plants and the bolites are always more or less calciferous, nature of the bedrock is, evidently, not as though as far as the amphibolites are concerned simple as often assumed. For instance, mobil often only to a small degree, while only certain subsoil water rich in electrolytes may serve, quartzites are practically non-calciferous. Ac to a certain degree, as a substitute for basic cordingly, the soils of our region would seem 134 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
predominantly to be circumneutral or basic. vegica, A. humifusa, Gerastium glabratum, &c., Except for local deposits of non-calciferous that are characteristic of the flora on non rocks, mainly quartzites which, n1oreover, occur calcareous serpentines (cf. RUNE 1949), the often in the form of narrow strips only a few term >> b as o phy t e>> is preferable. meters wide, no habitats suitable for acidophytes In literature written in English the different would seem to exist within the larger part denominations for plants occurring, exclusively of the regi9n outside the morainic and glaci or mainly, on soils with a basic or circumneutral fluvial deposits of leached material which, how reaction (calciphytes, calciphilous plants, cal ever, are mostly also more or less, calciferous cicole plants, &c.) would seem to be invariably unless composed of, usually high-alpine, rocks treated as synonyms (cf., e.g., CAIN op.c. p. poor in basic minerals ab origine. 4 77, DAUBENMIRE 194 7 p. 50): no distinction However, the deficiency of pH-determinations is maintained between the German conceptions may, to a certain degree, be met by means of of kalkstet (kalkbunden in Swedish) and kalkhold the numerous determinations made elsewhere (kalkgynnad in Swedish). However, this sim in the Scandes (cf., e.g. , 0HRISTOPHERSEN 1925, plification is· hardly desirable. \Vith a slight NoRDHAGEN 1928, 1935, 1936, 1943, Du RrnTz alteration of the terminology proposed by FER 1942a, 1942b, 1943, 1949, RUNE 1945a, GJAERE DINANDSEN (1918), I shall distinguish between VOLL 1949, WITTING 1949). such basophilous plants as are basocole, corre These determinations have proved that, as sponding to the German kalkstet, and such regards the distribution of plants, the essen as are merely basocline, corresponding to the tial thing is the basicity of the rock, while in German kalkhold. Similarly, I shall distinguish other respects the chemical composition is more between such acidophytes (corresponding to or less insignific.ant. Limestones, dolomites, oxylophytes, calcifuge plants, &c.) as are aci magnesites, even certain gabbros are almost docole (kieselstet) and such as are only acido equivalent as a substrate for the plants, ser cline (kieselhold). Amphicline should, perhaps, pentine being the only conspicuous exception. also be a better term than indifferent (cf. FER Even a non-geologist is able to distinguish in DINANDSEN op.c.). the field between such rocks and primary rocks, A few plants occur only in the eastern area of quartzites, hard, non-calciferous amphibolites, primary rock, viz., Betula verrucosa, Botrychium &c.; also .the serpentines are easily recognized. multifidum, Sparganium minimum, Scheuohzeria That is why I have considered my observations palustris, Oarex globularis, G. livida, Juncus worth publishing in spite of the want of pH stygius, Ranunculus lapponicus, Ledum palustre, determinations. and Lysimachia thyrsiflora, besides several plants The fact that lime is not the only geological that are at home in the inundation zone of factor determining the occurrence of plants the lakes. Ledum and probably Ranunculus is obvious, since typical calciphytes occur also lapponicus have each one known station situa on dolomite and magnesite, where Ca is partly ted on the mountain schists, and Scheuchzeria replaced by Mg, and on olivine stone and ser occurs in one place within the oligotrophic pentine, where Oa may be completely lacking. area of primary rock, anortite, &c., in the lower Therefore, it appears to me inaccurate to speak, part of the valley of Tjuoltavuopme (cf. above as is often done, of the flora on calcareous soil, p. 25). The same is the case with Phragmites &c., and below I shall, . instead, use the ex communis, which is also practically restricted pression circumneutral rocks as distinct from to the region of primary rock (only three sta the acidic ones. tions outside it). In some instances (Sparga Nor seems the term >>calciphyte>> quite ade nium minimum, Juncus stygius, Phragmites, quate. In view of the >>calciphytes>>, e.g., Asple and Lysimachia thyrsiflora,) the distribution nium viride, Minuartia rubella, Arenaria nor- is in all likelihood caused by the inability of Distributions within the Region 135
• • · ...... Approximate western boundary · 2 · Approximate limit of common of the area of primary rocks occurence of ledum palustre
e ledum palustre 0 Betula verrucosa 0 Ranunculus lapponicus
• Carex globularis 8 Botrychium multifidum 0 Juncus stygius
Fig. 24. vV esterrunost stations in Lule Lappmark of some sylvan lowland species.
the plants to: ascend above a certain altitude. on the eastern cambro-silurian, i.e. on rock The remaining species would seem to be more that is more or less calciferous. At least in the or less· basophobe, at least in our region. This case of such species as are found only in a few applies especially to Betula verrucosa, Botry steeps, basophily therefore might well be merely chium multifidum, and Car ex globularis (cf. the apparent. When they occur also in the region map. fig. 24). of primary rock east of the mountain range, Little can be said about the relationship they are evidently not basophilous at all. For between the bedrock of the mountain range instance, in the mountain range W oodsict ilven and the lowland plants occurring there. The sis, Polypodium vulgare, and Potentilla argentea observations made generally confirn1 the assump give a distinct impression of being basophytes. tion that many species display an increased Thus, ARwrnssoN (1943) characterized them tendency towards basophily when approaching as kalkhold, Polypodium even as kalkstet. But the limits of their ranges (cf., e.g., BEGER since they are rather frequent on the granite 1930 pp. 488 f., ALMQUIST 1929 p. 413). Regard mountains of Norrbotten, this must obviously ing those lowland plants which occur exclu be a mistake, due to the topography of the sively or mainly on south-bluffs a qualification mountain range; accordingly, these plants are must, however, be made. By far the greatest amphicline even in the northernmost parts number of such bluffs within our region are of their areas. situated on the mountain schists, especially In other instances, however, the behaviour 136 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
of the plants in Lapland doubtless differs, while in our region it occurs also (or exclusively) in the respect discussed here, from that in South in poor-fens. Sweden (regarding their behaviour in South The reason why some lowland species behave Sweden, cf. e.g. HARD AF SEGERSTAD 1924 differently in relation to the chemical nature and STERNER 1938). Two species that south of the soil in southern Scandinavia and in the wards are pronouncedly acidophile will, within mountains, while others react similarly in both our region, be found exclusively on basic or places, has possibly to be looked for in their circumneutral rocks, viz., Asplenium Tricho history (cf. below p. 175 f.). manes and A juga pyramidalis; and there is As regards the relations of the different quite a number of plants which are acidocline alpine plant species to the chemical nature in South-Sweden though noticeably basocline of the soil I can, almost invariably, endorse within our region, e.g. Botrychium Lunaria, the statements by ARWIDSSON (1943). The Milium effusum (LINDQUIST 1931 p. 249), Mai only exceptions among the basophytes arc a anthemum bifolium, Orchis maculata, Stellaria few plants characterized by him as amphicline graminea, and Sedum annuum. Carex ra agella or · basocline (kalkhold), which were observed by nica, which is an indicator species (>>ledart>>) me only on circumneutral rocks, in spite of the of the South-Swedish poor-fens (cf. WALDHBIM fact that most of them have a considerably grea & WEIMARCK 1943 p. 9), is amphicline in Lule ter number of stations in our region than in Lappmark, occurring, for instance, also in the Pite Lappmark. This holds good of Roegneria rich-fens of the Tarra valley; and, similarly, scandica, Carex capitata, C. atrofusca, Meland Eriophorum vaginatum, which in South-Sweden rium apetalum, and Potentilla nivea; furthermore, is a differential species (>>skiljeart>>) between ARWIDSSON does not distinguish between Fest�wa bogs and poor-fens on the one hand and rich rubra v. mutica and Cardamine pratensis ssp. an fens on the other, in our region occurs on all gustifolia, which are basocole within our region, kinds of mires (cf. S.roRS 1946b p. 43 and Du and F. rubra s. str. and C. pratensis ssp. RIETZ 1949 p. 289). Among the species which typica, which are amphicline. Most alpine in northern Lapland would seem to be strictly plants lacking in Pite Lappmark but occurring basocole, several are amphicline in the south, in Lule Lappmark are strictly basocole; this e.g. Carex pallescens, Oxal�s Acetosella, Viola is true of Cystopteris fragilis ssp. Dickieana, rupestris, Galium boreale, and Campanula rotun Poa herjedalica, Luzula arctica, Sagina caespi difolia. Especially some fen plants, the eco tosa, Minuartia rubella, .Lirenaria humifusa, logical amplitude of which is rather wide in A. norvegica, Draba hirta, Braya linearis, Poten southern Sweden, would in the north seem to tilla hyparctica, P. Chamissonis, and Primula ·be confined to highly calcareous rich-fens, scandinavica. Basocline, in a different degree, e.g., 11riglochin palustre and Carex panicea are only Roegneria mutabilis, Salix glandulifera, . (cf. Du RIETZ 1942c p. 7). The only excep Saxifraga Cotyledon, and Veronica tenella. (Cf. tions among the lowland plants are Stachys also TENGWALL 1925, pp. 745 ff.). silvatica, which in the South of Sweden is .All basophytes except Anthyllis Vulneraria basocline (LINDQUIST 1938 p. 203) and confined and a few other >>south-bluff species>> occupy to eutrophic habitats, while in our region it is continuous, western areas that include the met with only on the syenite mountain of whole, or larger or smaller parts, of the >>rich Tjalta, and Carex caespitosa, which in South area>>. This . distribution field ends abruptly Sweden (Scania) is among the indicator species at a line that almost coincides with the boundary of the rich-fens with its optimum in the extreme of the Sarek National park; east of this line rich-fens (WALDHEIM & WEIMARCK op.c. p. 27), many basophytes are lacking or more or less and is absent from the oligotrophic region of rare. It is evident that this sharply marked Smaland (HARD AV SEGERSTAD 1924 pp. 62 f.), boundary line has come into being owing to Distributions within the Region 137
1111 - Koli schists Eastern Serpentine Seve schists {western cambro-silurian) cambro-silurian
S ...,. �E
Amphibolites Gneisses, Eastern and southern and gabbros gra nites, & c. boundary of "rich area"
Fig. 25. Survey of the bed-rock within the region investigated, mainly according to Holmquist 1900 and Svenonius 1900 a. the transition from koli schists to the amphi ceeded by Phippsia algida, Sagina intermedia, bolites, syenites, &c. of the high mountains. Ranunculus glacialis, R. pygmaeus, &c. A dis The western slope of Mt .Allep Alatjakko can tance of a few steps is sufficient to change the, serve as an object lesson of its nature. Up entire character of the flora. to 1225-1240 m s. m. the koli schists predom On the Tarra valley the transition from the inate and, consequently, the rich >>Virihaure >>rich area>> to the poorer region east of it is. flora>> with Sagina caespitosa, Draba alpina, far more diffuse. On the sweep of soft, highly Dr. lactea, Pedicularis flammea, .Antennaria calcareous koli schists extending south to Mt. carpathica, &c. At this altitude the amphi Kuratjakko (cf. above p. 26) a vegetation of bolite sets in with a razor-sharp contact. The pronouncedly basophilous type predominates, edge of the amphibolite cap is indicated by with fens rich in species, and partly belonging to· a precipice with a long, narrow snowfield the extreme-rich-fen type, Trollius-meadows, below; and above the precipice a regio alpina Dryas-heaths, wind-heaths rich in basophytes,. media of the type frequent in Sarek begins &c. Here the characteristic >>Virihaure flora>> at once. Here the >>Virihaure flora>> is sue- also stretches out an offshoot which gradually 138 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
thins out and comes to an end on Mt Kura do not become free from snow until late in the tjakko (cf. below p. 152). Further west the summer (cf. NORDHAGEN 1943 p. 301); and boundary line once more becomes sharp and boulder-lands do not occur within the areas follows, approximately, the valley between of soft schists (phyllites, �ica schists, &c.; Mt Pajep Kuratjakko and Mt Rivkonjunjes cf. LUNDQUIST 1948 p. 398). These conditions, and the river Katnjunjesjokk, on the southern rather than the chemical nature of the bed bank of which only parts of Mt Skaitatstjakko rock per se, may conceivably decide its distri harbour a few basophytes. On the northern bank bution. In fact, TH. FRIES (1925 p. 11) and the koli schists of Mt Katnjunjes put in a .A.RwrnssoN (1943 p. 154) deny that it is always wedge between two fields of seve schists (HoLM acidocole; on the other hand, N ORDHAGEN (Le.) QUIST 1900 p. 79; cf. above p. 26); and on Mt and GJAEREVOLL (1949 p. 80) unhesitatingly Katnjunjes the >>Virihaure flora>> is, in fact, declares it to be an acidophyte. very well-represented. Alchemilla alpina appears to be completely Further east Mt N junjesvare forms an island lacking in the >>rich area>> except for a station of calciferous rocks with a rich flora, in rather on Mt N umirvare reported by BJORKMAN barren surroundings (cf. above p. 24). Several (1939 p. 13), which I have not seen and the western basophytes recur also on the eastern nature of which I am, therefore, unable to judge. cambro-silurian, especially in the east-precipice Its stations in the Tarra . valley are all situated of Mt Kassavare (cf. above p. 22). The occurr on glacifluvial deposits poor in lime, except ence of Hyolithus schist on the crest of Mt that on Mt Njammilt; however, here the sub Lastak announces itself, i.a., through an iso stratum seems to have been formed by an lated station for Pinguicula alpina. Finally, ancient mountain-slip from the above precipice in spite of its eastern situation Mt Sakok har which, at least partly, would seem to consist bours a great many alpine basophytes, viz., of amphibolite. Personally, I am inclined to Woodsia glabella, Polystichum Lonchitis, Kobre accept Alchemilla alpina as acidocole as far .sia myosu'roides, Minuartia str;·icta, Melandrium as its stations within our region are concerned, .apetalum, Braya linearis, Oxytropis lapponica, as NORDHAGEN (1943 p. 245) does with regard Rhododendron lapponicum, Gentianella tenella , to its stations in Sikilsdalen (south-western Veronica fruticans, Pedicularis hirsuta, Cam Norway) and GJAEREVOLL (op.c. p. 86) as to panula uniflora, Eriger;·on boreale, Arnica alpina, those in Mt Oviksfjallen (Jamtland) . .&c. The explanation doubtless is that parts Carex rottmdata is the only alpine plant con of this mountain are situated on the eastern sidered by TH. FRIES (I.e.) to be acidocole every
· cambro-silurian (SVENONIDS 1900a). In Sarek where within · the mountain range. However, proper many basophytes recur on the clay Du RIETZ and WITTING haYe found it dominating .schists of Kukkesvagge (HAMBERG 1910 p. 693). in transitional rich-fens (Warnstorfio-Scorpidion, The alpine acidophytes are, of course, as cf. Du RIETZ 1949 p. 303) in Kattervagge in relatively few in number within our region as Torne Lappmark, where the pH of the water in the rest of the Scandes. In the Scandinavian is up to 5.9 (unpublished). It can thus not be flora only three alpine plants are generally classified as calciphobe. In our region, it occurs considered to fall within this cathegory, viz., also in the >>rich area>>, though its habitats Cryptogramma crispa, Carex rotundata and Alche there are glacifluvial deposits that, to judge milla alpina. The status of Cryptogramma, from the flora, are more or less oligotrophic howev�r, would seem to be a little doubtful, in character. I have, for instance, never seen although, in our region, it has been ob it grow in the company of its basocole parallel served only on non -calcareous amphibolite and species C. saxatilis, though in two stations, on quartzite. Its habitats are, without exception, 1\It Puokkik and Lake Katjaure, they occur boulder-lands in the lower alpine belt which in close vicinity of one another. NORDHAGEN Distributions within the Region 139
(1943 p. 449]. classifies it as oligotrophic and heath peat (raw humus); cf. NoRDHAGEN op.c. ac.idophilious. pp. 121 ff.; The question is whether a few other plants Gnaphalium supinum: astonishingly rare all may not rather be considered to shun basic over the rich area, probably altogether absent rocks, more or less definitely. One of them is from some mountains. Since I have not searched Betula tortuosa. The only large stand of pure systematically for it, the latter impression may, Betula tortuosa that I have seen there grows however, be erroneous. Anyhow, the species on the barren, obviously non-calcareous lit is pronouncedly acidocline and perhaps acido Tjarrok, where it descends down to the coni cole (cf. G.JAEREVOLL op.c. pp. 89 f.); fer belt; on the whole, the species seems to Gnaphalium norvegicum: about the same be fairly common only on mountains of an distribution as Gn. supinum, though less high oligotrophic character, e.g., Mt Kaskaivo and alpine, within the rich area mainly occurring Mt Tjalta. Until the taxonomy and distribution on the glacifluvial accumulations of the valleys, of the alpine birches have been cleared up, and also less acidophilous, having numerous it is, however, impossible to decide how far stations for instance in the birch belt of the these impressions correspond to actual facts. Tarra valley (cf. G.JAEREVOLL op.c. p. 90); None the less, it seems reasonably certain that H ieraciurn alpinum coli. and H. nigrescens pure Betula tortuosa, like most acidophytes, coll.: seem in the rich area to be confined to has a preponderatingly eastern distribution acidic heath peat (raw humus), practically within our region. · to the exclusion of other habitats. Large stands Further instances of acidophilous alpine spe of H ieracium alpinum or Pedicularis lapponiaa cies of our region are: announce already from a distance a >>poor>> Athyrium alpestre: frequent on the amphi locality where it is no use looking for rare baso bolites and quartzitic seve schists of the eastern phytes. mountains, being in the >>rich area>> mostly Among the lowland species normally occurring confined to the seve schists, while scarce on the in the alpine belt especially Desohampsia eutrophic mountains. The largest stands of flexuosa, Nardus stricta, Vaccinium Myrtillus, the species occur on the barren :1\ft Kartevare Trientalis europaea, and Solidago Virgaurea fall tjakko and the equally barren Mt Kaskaivo. within this category (cf. Du RIETZ 1942b p. 60). (Of. G.JAEREVOLL op.c. p. 80); It is rather striking that so many acidophytes Deschampsia atropurpurea: fairly common, at belong to the vegetation of the snowbeds� any rate locally, on the amphibolites and quartz Only comparatively few species, e.g., Antho xanthum alpinum, Poa arctica, Phippsia al ites of the eastern part of the region; in all . the Virihaure-Vastenjaure basin observed by gida, Carex Lachenalii, Juncus biglurnis, Salix me only on amphibolite and, in a few places, glauca, Oxyria digyna, Stellaria ca.lycantha, Cera Dn leached glacifluvial material; stium alpinum� Saxifraga stellaris, Sibbald·ia Juncus trifidus : surprisingly scarce in the procumbens, Viola biflora, Gassiope hypnoides, rich area, where hardly any Juncus trifidus Yeronica alpina, Euphrasia frigida, and Peta heaths occur at all, frequent only on glacifluvial sites frigidus, are definitely as frequent within deposits, for instance, at the river Puolejokk the rich area as outside of it, i.e. truly indifferent and in the Stalok valley below Mt Jalle on to the chemical nature of the soil. Also Carex Vejejokk; common on the quartzitic . schists, rufina is entirely amphicline. On the other amphibolites and syenites of the eastern moun hand, at least the following alpine plants, many tains (cf. NORDHAGEN 1943 p. 208); of which belong to the common dwarf scrubs Pedicularis lapponwa: not completely miss (cf. TENGWALL 1925 p. 742), are doubtless more ing on any mountain, but very scanty on the scarce on basic or circumneutral rocks than on eutrophic ones and mainly appearing on acidic more or less acidic ones: 140 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Lycopodium alpinum, Carex brunnescens, Salix tains close to the Norwegian border, M t Rakok herbacea, Rumex Acetosa ssp. a1·ijolius (at any tjakko and Mt K.uotelis with their poor flora, rate in the alpine belt), Epilobium lactiflo and a fe'v amphibolite mountains (cf. p. 29) rum, E. Hornemanni, Loiseleuria procumbens, immediately east of the river J aurekaska, the Phyllodoce coerulea, Arctostaphylos alpina, Dia bedrock here consists, as pointed out above pensia lapponica, and possibly Phleum commu (p. 27 ff. ) , almost exclusively of soft rocks rich tatum, Agrostis borealis, Carex Bigelotvii, and in Ca or Mg, viz., phyllites and other easily Cerastiurn trigynum. weathered mountain schists, limestone, dolo However, this type of distribution must mite, olivine stone, &c. True, north of Lake not of necessity be interpreted as a negative Sallohaure and to a . certain extent between reaction, on the part of the plants in question, Lake Sallohaure and J.1ake Virihaure the lime to the basicity of the soil. As already pointed stone appears, as already mentioned (above out, by OLSEN (1936) among others, compe p. 32), to be harder and more crystalline and tition must be less severe on acidic soil where the mountain schists to be less easily weathered the basocole plants are excluded. Consequently, than in the area south and east of Lake Viri the species which are able to live in such habitats haure. However, the difference is hard1y big find it easier to assert themselves there and enough to explain the considerable dissimilarity become dominant. of the flora south of Lake Vastenjaure and However, the reasons why many amphi north of this lake, and especially north of cline species occur mainly on more or less acidic Lake Sallohaure. soils lie outside the scope of this investigation. Among the 327 undoubtedly native species, For our present purposes it suffices simply excluding Taraxaca and Hieracia, that are to state the position and note that it is re known from these parts of our region, no less flected in the distribution of these amphi than 94 (plus 3 Hieracia) are lacking either cline plants, their stations mostly being concen north or south of some of these three lakes. trated to the eastern mountains, and much more A few hydrophytes and shore plants that have scattered within the rich . area. been noticed exclusively in or around Lake Virihaure were, however, mostly observed there
Topography only in the past few years; they are, probably, not noted within the northern tracts simply The chemical nature of the soil is, of course, because those parts were investigated less com not the only edaphic factor that should be pletely and earlier in the summer than the taken into account when trying to explain the Virihaure region. This holds true of the follow distribution of the plants of a region. Naturally, ing species: the topography is also of great importance. Potarnogeton filiformis Scir us acicularis Its influence, however, manifests itself in so p P. alpiwus J·uncus alpinus many ways and is often so intimately connected P. gramineus Oallit·riche verna with the chemical nature of the bedrock, that Agrostis stolonifera var. Hi pp·uris vulgaTis it would be n1eaningless to try to define its arctica Veronica scutellata significance with regard to the distribution of the plants within our region in its entirety. In regard to the remaining plants the gaps Only a smaller, well-investigated area which should, generally, be real. That some of these geologically is comparatively uniforn1 can give species recur in northern Sarek, the mountains an idea of the importance of topography in at the river Stora Lule alv, and central Vaisa, this respect. on the one side, and on Lake Peskehaure, the Such an area is the Virihaure-Vastenjaure Tarra valley and Lake Saggat, on the other, basin. With the exception of the granite moun- is irrelevant in the present instance. Distributions within the Region 141
The following plants occur only south of a Prunus Padus var. bore Viola montana line Virihaure-MelH1tno: alis Afuga pyramidalis Potentilla erecta Erigeron acre W oodsia glabella Ranunculus hyperboreus Lotus corniculatus Hieracium diasemum Festuca rubra v. mut1:ca Draba crassifolia Carex capitata Potentilla hyparctica Finally, south of the line Virihaure-MelH1tno C. bicolor Astragalus norvegicus the following plants are lacking: Luzu.la arctica Epilobium davuricum Rumex tenuifolius E. alsinifotium Carex flava Hackelia deflexa Sagina caespitosa Campanula uniflora Silene rupestris Valeriana sambucifolia M inuartia rubella Antennaria carpathica Sedum acre and probably A renaria hurnifusa Alchemilla filicaulis Erigeron boreale Calluna vulgaris The following are missing north of Lake Vastenj aure: However, the d_ifference between the flora north and south of Lake Vastenjaure does not Spar·ganium hype1·boreum Cassiope tetragona Ca'rex 'rnicroglochin Vaccinium rnicrocarpum stand out clearly if the dissimilarities in the 0. arctogena Pedicularis hirsuta frequency of many species be disregarded. 0. macloviana P. flammea For the purpose of finding areas, for the sake Arenaria norvegica Galium boreale of comparison, that are not of too unequal a Rubus arcticus Arnica alpina size, Padjelanta (District VI) has been excluded North of Lake Sallohaure the following species from the southern area, and only stations south are absent: and south -east of Lake Virihaure and on the peninsula of Kollomoloki are taken into account. Lycopodium clava,tum Listera cm·data Even then, the northern area is considerably Poa arctica Corallorhiza trifida P. herfedalica Salix polaris smaller than the southern; and since, further, Carex rufina Stellaria nemorum ssp. it is far less completely investigated, an equal J'uncus arcticus montana number of stations in both areas will, as a J. filiformis Aconitum septentrionale . matter of course, indicate a higher frequency Ramischia secunda Luzula par,vijlora in the northern one. Of the figures in brackets, L. Wahlenbergii the first one signifies the number of known On the other hand, some species are found stations south of Lake Vastenjaure and the only north of Lake Sallohaure, viz.: second one stations north of this lake.
Athyrium Filix-femina Viola rupestris Decidedly more common north of Lake Dryopteris Filix-mas Epilobium collinum Vastenjaure are the following plants: Gentianella aurea Polypodium vulga1·e Asplenium viride (3, 13) Bray a linearis (I, 3) Veronica tenella Scirpus pauciflorus Woodsia alpina (3, 5) Saxijraga adscendens (3, Antennaria lapponica Se. Hudsonianus Roegneria can·ina ( 1, 4) 8) Hieracium arctogenum M aianthemum bifolium 01·chis maculata (0, 2), Primula scandinavica (2, H. prenanthoides coli. Salix coaetanea Gymnadenia conopsea ( 2, 5) Alchemilla alpina 5) Veronica fruticans (6, 10) Erysimum hie'ruciifolium Erigeron politum ( 0, 5) Several other plants are observed only north (0, 2) .of Lake Vastenjallre, as follows:
Equisetum hiemale Carex lasiocarpa Keeping in mind the difference in the size Poly::;tichum Lonchitis Polygonatum verticillatum of the respective areas, and the blanks in the Lastrea Robertiana Epipactis atrorubens investigations, the following species n1ust also Agrostis tenuis Caltha palustris be considered to have a higher frequency in Ambis hirsuta var . . sub M elica nutans the north: Roegneria mutabilis alpestris Eriophorum brachyanthe- Sedum annuum Roegneria scandica (3, 4) Draba hirta (10, 9) rum Carex glacialis· (13, 13) 142 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
The number of species that are more frequent tum 'Verticillatum, Ajuga pyramidalis, and Viola south of Lake Vastenjaure is considerably larger, rupestris. Other basocole species, e.g., Asple as follows: nium viride, Carex glacialis, Draba hirta, Braya OystoptP.r·is montana ( 11,2) Draba r1,lpina (31, 4) linearis, and Primula scandinavica, a,re decidedly Poa herjedalica ( 15, 2) Gardamine prate'nsis ssp. more frequent north of Lake Vastenjaure than Garex parallela (22, 1) angustifolia (28, 4) in the rest of our region; others, e.g., Carex a. nardina (22, 2) Potentilla Ghamissonis rupestris, Juncus triglumis, Leuchorchis albida, Juncus arcticus (23, 2) (5, 1) J_ll inua.rtia stricta, Dry as octopetala, Oxytropis Gerastium glabratum (21, P. nivea s. lat. (28, 2) 4) Rhododendron lapponi lapponica, and Euphrasia lapponica, are about Sagina intermedia (23, 6) cum (30, 3) as frequent there as elsewhere; and Pinguicula M elandrium apetalum Gentianella tenella (23, 3) alpina, which is confined to highly calcareous (30, 4) Erigeron unalaschkense fens, reaches a maximum of frequency round Draba lactea (24, 2) (26, 2) Lake Sallohaure. To this has to be added that Dr. nivalis (41, 3) a few more or less amphicline species, e.g., In addition, there is a not insignificant num Car ex arctogena, C. macloviana, and Ranun ber of species, for instance Phippsia algida, culus hyperboreus, seem to be lacking north Salix polaris, Ranunculus glacialis, Cardamine of Lake Vastenjaure, while others, e.g., Poa bellidifolia, Saxifraga foliolosa, S. rivularis, arctica, Juncus arcticus, Luzula parviflora, and &c., which doubtless are much rarer north of L. "fV ahlenbergii, that occur between r.. ake Lake Vastenjaure than south of it, though Vastenjaure and Lake Sallohaure, were not no figures can be given, since TENGWA:LL (1924) found north of the latter lake. fails to state the number of stations in the moun Therefore, the causes must be searched for tains visited by him alone. elsewhere. An explanation is suggested in the Occasionally the explanation of the ranges striking affinity of the flora north of Lake, is fairly obvious. l?or instance, Aconitum Vastenjaure to that of the limestone hills on septentrionale reaches its northern Swedish limit the coast of Helgeland, investigated by 0. DAHL within this �irea. And the major part of the (1912, 1915), though these are situated just. plants occurring only north of Lake Sallohaure above the sea-level. consist in subalpine species, the stations of A number of lowland species, most of which which are outposts from their areas on the river are lacking in the rest of our region, ascend Stora Lule alv (cf. above pp. 100 f.). The gap into the alpine belt near Lake Vastenjaure. within the areas of some lowland species (Lyco All these species recur in Helgeland. Accord podium clavatum, Juncus filiformis, Listera ing to DAHL Polygonatum verticillatum, Sedurn cordata, Corallorhiza trifida, Ramischia secunda) acre, S. annuum, Lot·us corniculatus, and Erigeron north of Lake Sallohaure is probably due to acre aTe frequent there; Gentianella aurea is imperfect investigations; it will not be taken fairly frequent and Epipactis atrorubens, not. into consideration below. rare. Arabis hirsuta is said to be a )>character On the whole, however, the reasons of the plant of the schist and the limestone)> and striking dissimilarities between the floras of Aj1tga pyramidalis, to occur )>here and there)>; the two areas are not as evident as in these the same statement is made by NORDHAGEN particular cases. 'l'he differences cannot be oc (1940 p. 270) regarding Viola rupestris, which casioned only by the presumably better supply DAHL does not distinguish from V. Riviniana; of assimilable lime in the Virihaure area. Several and Lastrea Robertiana is reported from 11 species lacking round Lake Virihaure are baso stations, all told. cole, either everywhere as Arabis hirsuta var. This similarity, however, is hardly surprising,. subalpestris, Epipactis atrorubens, and Lastrea since the stations near Lake Vastenjaure of Robertiana., or in these parts as, e;g., Polygona- precisely these species must be regarded as. Distributions within the Region 143
Fig. 26. Vie>v t.o the west from Mt Tj argesvare. In the foreground Mt Unna 'l'jargesvare, that is already free from snow, while the Norwegian mountains, where the westerly winds discharge most of their moisture, are still largely snow clad. In Mt Unna Tj argesvare, the northward slanting of the rock-layers is plainly t.o be seen. Photo Sten Selander 12.7. 1946. outposts from their areas on the Norwegian hirta, which DAHL does not distinguish from coast. More significant is that the affinity Dr. norvegica, is reported from these parts, is almost as pronounced regarding the alpine i.a., by NORDHAGEN (op.c. p. 257). In Beige species. Among such plants characteristic of land, all these plants descend almost to the the mountains north of Lake Vastenjaure only sea-level. Pinguicula alpina is lacking on the coast of Out of the plants, between 50 and 60 in Helgeland, where also Minuartia stricta is number, which are characteristic of the Viri missing. According to DAHL, Oarex rupestris, haure region, no less than about 45 are decidedly Dryas octopetala, Veronica fruticans, and Eu alpine and occur mainly on the northern slopes phra,sia lapponica are >>Character plants>> of the of the mountains. Only three among them, limestone hills; Asplenium viride, Polystichum . viz., Luzula Wahlenbergii, Arenaria norvegica, Lonchitis, Juncus triglumis, Primula scandina and Potentilla nivea, appear in the coastal vica, and Erigeron boreale are rather frequent; region of Helgeland, where the number of W oodsia alpina, Roegneria scandica, Oar ex gla known stations, according to DAHL, is 3, 4 cialis, Leuchorchis albida, and Oxytropis lappo and 1, respectively. The flora north of Lake nica are said not to be rare; Bray a linearis, Vastenjaure can in its entirety be characterized which is rare everywhere in Scandinavia, as an outpost of the flora occurring on the lime occurs, however, in three plaees; and Draba stone hills of the Norwegian coast south of 144 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark our region; the >}Virihaure flora>} is mainly arctic with regard to the occurrence of southern species, and high-alpine in character. In other words, the reason being that some >}South-bluffS>} (syd the >}Va,stenjaure flora>} may be said to be large berg) on the Rapa valley, e.g., Mt Litnokvaratj ly a, relict fr01n the postglacial warm period, and Mt Tjakkeli, face north. BJORICMAN (1939 while the >}Virihaure flora>} is characterized p. 213) likewise points out the occurrence on chiefly by glacial epibiotics. the river Stora Lule alv of a few steeps facing This floristic difference would, in the first north -east with a southern element in their place, seem to be occasioned by the geotectonics vegetation, although >}the number of individuals of the Virihaure-Vastenjaure basin. As far is always considerably smaller and the speci as I know, no expert investigation of the basin, rnens show a lower degree of vitality>} there from this point of view, has been published. than on steeps facing south. However, it seems LUNDQVIST (1948 p. 123) states that the schists to me that, at all events, TENGWALL overrates round Mt Sulitelma slant towards this massif, not a little the significance of his observations. which, thus, originally formed the bottom of a On the steeps mentioned by him only very fold, and that the great valleys are the remains few lowland plants occur, none but Polypodium of the ascending parts of the folds. For the vulg�re being an indubitable southern immi rest, we must be content with the unsupported grant. It would, in fact, be very extraordinary impressions of non-expert. Especially north if his assertion were true, seeing that a slope of Lake Sallohaure, though a.lso, less pronounc of as little as 5° towards the pole reduces soil edly, between Lake Sallohalire and I;ake Vasten temperature approximately as much as 300 jaure, the rock-layers would seem to slant north miles of latitude in the same direction (DAu wards (cf. fig. 26). The consequence is that BENMIRE 1947 p. 177). the north slopes here, as wherever the surface Within our region there are also one or two of the mountains and the slanting rock-layers steeps facing north though possessing a flora run parallel, are too little weathered to support of partly southern character, e.g., the big north more than a rather poor flora, while the richer eastern steep of Mt Kaddepakte on Lake Saggat flora is concentrated in the southern steeps (cf. NYMAN 1895). These are, however, sheer with their abundant, highly weathered mineral exceptions. In the region of primary rock soil. The reverse is true of the mountains south the significance of exposition is proved by of Lake Virihaure, where the gradually slanting >}Predikstolen>}, a northern precipice on Mt south slopes are relatively poor in species, while Ailates, when compared with the neighbouring the steep northern sides are botanically very southern precipice of Mt Kanivare. The former rich. is situated within a relatively eutrophic area; As pointed out by NANNFELDT (1940 p. 37), yet on this steep no southern species are to among others, on sufficiently high latitudes be found, and no noteworthy plants at all, the northern sides of the mountains offer the except Dryopteris austriaca and Stellaria longi most favourable conditions to an arctic-alpine folia. On the contrary, the latter precipice, vegetation: there is sufficient light, even there, although situated within the extremely oligo for the needs of the plants, the ground does trophic area north-west of Lake Karatsj, and not dry up completely as is often the case on at a higher altitude than that of >}Predikstolen>}, the southern sides, and the temperature does harbours at least three >}South-bluff specieS>}, not exceed the maxima of the arctic species. viz., Woodsia ilvensis, Silene rupestris, and It goes without saying that the reverse holds Sedum annttum. good of the southern sides, as well as of the To the west of the mountain range practi southern plant species. cally no lowland species occur on steeps facing True, TENGWALL (1925 p. 768) asserts that another quarter than south and south-west. the exposition is only of a secondary importance Even in the birch belt there are nearly always Distributions within the Region 145 snow-beds below the wet northern steeps. represents, in miniature, the floristic contrast Only in one place, on Mt Njerek in the regio between the northern steeps of the Virihaure alpina 650 m s. m., a single lowland species, mountains and the southern steeps of the moun Rubus idaeus, has been observed on a northern tains north of Lake Vastenjaure. (Of. the sim steep, at the foot of which it forms thickets ilar observations in FRODIN 1915 pp. Z05 ff.) together with Salix glau0a and S. glauca x This southern steep, and still more the large phylicifolia. For some reason, perhaps that >>south-bluff>> of Mt Aran:Uve, situated in the it is partly made up of olivine stone, this steep regio alpina on the northern shore of Lake is, also in other respects, unlike all the neigh Vastenjaure, may give us an idea of the locali bouring ones: thus it supports some xerophytes, ties on the Norwegian coast where lowland viz., Kobresia myosuroides, Oarex rupestris, species are supposed to have survived the . Draba nivalis, and Potentilla Ohamissonis, which latest glaciation. We are here at a very short otherwise do not at all belong to the flora distance from two of the mightiest glaciers of the northern steeps. of Scandinavia, viz., BHl.mansisen which meas The most striking instance of the signifi ures about 150 krn square, and Flatkjelen cance of exposition is, however, to be found which is not mu.ch smaller, not to mention on Kollomoloki, the peninsula between Lake Rakokjekna, Kuotelisjekna, Hurrejekna, Veik Virihaure and Lake Vastenjaure. On the same dalsisen, Almallojekna and other large glaciers. hill, at the same altitude, about 610 m s. m., There is hardly any reason to suppose that and at a mutual distance from one another of an ecologic change worth mentioning should only about 2 km, there are a southern steep ensue, were these glaciers to fuse into one huge (fig. 27) and a northern steep (fig. 28) of an ice-sheet. Yet, not only the lowland species identical geological nature. Nevertheless, the enumerated above but, e.g., Lastrea Robertiana, differences in insolation, humidity and date Polygonatum verticillatum, Epipactis atrorubens, of the melting of the snow and, consequently, Ery�imum hieraciifolium, Lotus corniculatus, in the length of the growing season occasion Ajuga py1·amidalis, Erigeron acre, and Tussilago dissimilarities between the floras of the two farfara occur here, in close vicinity of the gla steeps that are surprisingly marked. On the ciers. And on the similar �It Tjargesvare i.a. northern steep, which at the end of July is S edum� annuum, Viola rupestris, Epilobium still irrigated from a snow-field above it, there collinum, Gentianella aurea, and H ackelia deflexa are to be found, e.g., Poa herjedalica, Dr·aba join them. The great majority of these plants lactea, Saxifraga tenuis, S. foliolosa, Pedi6ularis may be suspected of being glacial survivors; hirsuta, P. flammea, &c., several of which are and when seeing them in their stations round characteristic of the upper parts of the regio Lake Vastenjaure, this assumption appears alpina. On the other hand, clumps of birches most plausible. with Filipendula Ulmaria, Geum rivale var. In our region, the hypothetic thermophilous subalpinum, Valeriana sambucifolia, and other survivors nowadays reach an altitude of 700 m subalpine plants are growing on the southern s. m. or more. However, the ice-age refugia steep at the same level and 70 m further up; were situated mainly near the sea level. The and in the drier, more scree-like parts of this temperature in mountainous . regions decreasing steep, there are to be found, i.a., Silene rupestris, about 5,5° C. for each increase in elevation of Sedum acre, Saxifraga adscendens, Anthyllis 1000 m, irrespective of latitude, this difference Vulneraria var. lapponica, and (in scrub-heath) in altitude corresponds to a difference in tempe Listera cordata, besides some xerophilous alpine rature of about 4° C. Consequently, the lowland plants, e.g., Kobresia myosuroides, Oarex gla plants supposed to be glacial survivors would cialis, Arenaria norvegica, Draba hirta, Dryas be able to live in the refugia, even though the octopetala, and Oxytropis lapponica. This hill mean temperature during the glaciations fell
11 - 496149 Sten Selander I 146 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Fig. 27. Southern slope of a hill on Kollomoloki, the peninsula between Lake Virihaure and Lake Vastenjaure, with birch shrubs and lowland plants, e.g., Sedum acre, Listera cordata, l!'ilipendula Ulmaria, Valeriana sambucifolia, &c. Photo Sten Selander 20.7. 1946. as much as 4 o C. As far as I know, no compu of the glaciations, at approximately 4 o C. tation of the changes in the temperature during As a matter of course, this does not imply the glacial periods in Scandinavia is made. that identical changes took place in Scandina The many calculations of the hypothetical cli via; on the Norwegian west-coast, the difference mate of Central Europe during the glacial epochs between present times and the glaciations was, do not apply to Scandinavia, since they refer in all likelihood, considerably less marked (cf. to a continental region betwen two ice-caps. LINDROTH 1949 pp. 782 ff. ) . The calculation of Only an estimation of the ice-age climate of a CRANKY & MASON may, none the less, give us a coastal region would be relevant. And no such rough idea of the maximal dimensions of the estimation is made as regards Europe. changes in ternperature that · have to be taken Basing their argumentation on the composi into account. Accordingly, it appears that the tion of the fossil closed -cone pine forest flora existence of hibernating lowland plants in Scan of Santa Cruz Island CHANEY & MASON (1930), dinavia is not a matter beyond the bounds of however, estimated the lowering of the mean possibility. (For furthe.r instances of a thermo temperature of South-California, during one philous biota living hard by large glaciers, cf., Distributions within the Region 147
Fig. 28. Northern slope of this hill, situated at the same altitude and at a distance of c. 2 km. The flora consists exclusively of alpine species, e.g., Draba lactea, Saxifraga tenuis, S. foliolosa, Pedicularis hirsuta, P. flammea, &c. Photo Sten Selander 20.7. 1946. e.g., LINDROTH 1931 pp. 541 ff., Du RIETZ 1935 plants belonged to the glacial survivors. This p. 228, BOCHER 1938 pp. 318 f.) is, however, hardly conclusive. Such species Furthermore, one indubitable glacial sur were, of course, far from frequent in the ice vivor, Braya linearis, is confined to xerothermic age refugia, occurring only in limited, edaphi habitats of exactly the same kind as those of cally favoured localities. That they would the forementioned plants. It hardly ascends have left traces in the pollen profiles seems higher than the lowland species, and seems, utterly unlikely, even disregarding the fact on the whole, to demand approximately the that most of them are entomophilous plants same edaphical and thermic conditions as, for with low pollen production. One may rest assured instance, Viola rupest1·is or Ajuga pyramidalis that Epipactis atrorubens, Sedum acre, Ajuga in their North-Scandinavian stations. This, pyramidalis, &c., do not manifest themselves evidently, renders the hypothesis of hibernat in the recent pollen spectra of the Virihaure ing lowland plants still more reasonable. Vastenjaure basin which, no doubt, are domi Late-glacial pollen profiles are sometimes nated by Betula nana, Salices, &c., i.e. are brought forward as proofs that no lowland subarctic in character. There is obviously no 148 SELANDER: Floristic Phytogeography of South-Western Lule Lappmar k
reason to presume the spectra of the late is deprived of the melting-water from snow glacial deposits to have differed in this respect. fields, &c. Pollen profiles indicating subarctic or even arctic As snow receptacles, the eastern sides of the conditions do not disprove the occurrence mountains equal the northern ones, since the of southern lowland plants in the close vicinity. snow, usually coming with the west winds, The contrast between the flora north of is deposited mainly on the lee sides, i.e. the Lake Vastenjaure and that south of it is empha east slopes (ENQUIST 1916). All glaeiers within .Al sized by the fact that especially the southern. our region (Kuotelisjekna, Rakokjekna, steeps north of Lake Sallohaure, on Mt Tjar mallojekna, the small glacier on Mt Kaisekets gesvare and the adjacent mountains, are rather tjakko, Jeknaffojekna, and the glaciers on Mt dry because the rock-layers slant towards the Sulitelma and Mt Staika) face east or, in one north and, consequently, practically no water or two cases, north -east or south -east, and the oozes out of the rock-faces. Immediately north eastern sides of the mountains are throughout of Lake Vastenjaure the . rock-layers, at any characterized by the abundance of snow-beds rate here and there, would seem to be almost and perennial snowfields. In terms proposed horizontal, so that the southern precipices, by NORMAN (1895 p. V), the >>backS>> of the for instance on Mt .Aram1ive, are irrigated, mountains, where the altitude belts have been locally, by considerable quantities of trickling forced downwards and the flora of the snow water. Here one finds, in fact, mesophytes such beds and of the ground flushed by melting as Poa herjedalica, Epilobiurn lactiflorum, Tussi water predominates, do, consequently, not lago fwrfara, &c., growing in the damp ravines. inelude only the northern sides, but the eastern For the rest these steeps are largely as dry ones as well. (According to GELTING, 1934 as those on Lake Sallohaure and chiefly support p. 234, in East-Greenland the southern sides the same xerophilous flora. are the >>backs>> of the mountains owing to the northerly winds that prevail at winter-time.) Another cause of the xerophilous character The more xerothermic >>Sunny sides>> embrace of the flora north of Lake Vastenjaure is that the southern, south-western and western slopes; no rnountain tops within this area exceed the the south -eastern and north-western slopes 1000 m level to any degree worth mention. are indifferent in this respect. This is in itself sufficient to explain the absence, Mt Tuki is typical of the low mountains in or lesser frequency, of certain high -alpine spe the Virihaure region. The vegetation of its cies, e.g., Luzula arctica, Ranunculus glacialis, western slope is pronouncedly xerophilous and Draba laotea, Dr. alpina, Dr. crassifolia, Car low-alpine, while a number of rather high damine bellidifolia, Potentilla hyparctica, Erigeron alpine, partly chionophilous species occur on and unalaschkense, &c. However, with the exception below its strongly developed eastern precipice, of Draba crassifolia and Potentilla hyparctica, where the snow remains. On the adjacent low the character species of the >>Virihaure flora>> mountain of Kuobberi there is a damp northern occur, as pointed out above (p. 131), also on steep, where even so high-alpine a species as mountains that are as low as, or even lower Luzula arctica manages to exist. On .a third than, those north of Lake Vastenjaure. In similar mountain, Unna Titer, a corrie facing this respect the characteristics of the flora east fills the same place. The northern steeps are, accordingly, mainly an indirect result of on the peninsulas of N jerek and Kollomoloki the absence of high peaks. have already been mentioned. Such a result is that in normal summers, North of Lake Vastenjaure, on the contrary, the mountains become free from snow already all steeps face south. On the gently sloping at the beginning of July. During the major northern and eastern sides of the mountains, part of the growing seaso:n, the vegetation thus there are no steeps where the snow would remain Distributions within the Region 149 even at this low altitude. In consequence, snow beds occur only on level ground where, as usual in such places, they are chiefly occupied by Gassiope hypnoides- and · Salix herbacea heaths poor in species (cf. TENGW.ALL1920 p. 379). To this has to be added that generally the clouds do not pass below an altitude of 1000- 1200 m and, therefore, do not touch the low mountains in the northernmost part of our region. In the Virihaure basin the number of cloudy days during the growing season is very large; the figures given by CLEVE (1901 p. 65) are not representative, since the summer of 1896 evidently, as she points out, was un usually dry and fair. Even during the driest summer month, that of July, there may not be a single hour of really fine >>top weather>> without cloud-caps on the highest summits. So far as my experience goes, even in fine summers ten July days that can be called clear Fig. 29. The glacier on the south-eastern slope of Mt J ek are exceptional, and during the summer of naffo. As usual, the upper parts of the mountain are hidden 1942, which was not especially ill-favoured, in rain-clouds. Photo Nils Dahlbeck August 1940. there were altogether three sunny days in July. It is well known that moisture, precipitated as fog, &c., on the cloud-capped peaks, is very able in view of the fact that both Arnica important to their vegetation. This is, of course, and Gampan�tla uniflora recur on the neigh particularly true pertinent to so cloudy a region bouring, high-alpine Mt Riddoalge (ace. in as this. Especially on mountains that barely forma.tion in litt. by Mr G. BROD.AL). In this reach above the >>lower c.loud limit>>, e.g., Mt connexion atten.tion may be called to the small Kappa, Mt Mctjer and Mt Jalle on Vejejokk, >>Rubus Ghamaetnorus-bogs>>, mentioned above the flora generally is richer on the summits on p. 118, that occur on mountain crests south than further down. On the mountains round of Lake Virihaure; they have, of course, no Lake Sallohaure the flora of the tops is, on the counterparts in the northern area. contrary, remarkably poor in spite of the far Also the local climate occasioned by the from unfavourable bedrock. Among species topography of this area would seem to influ lacking there are Minuartia rubella, Gampanula ence its floral character. Exact meteorological uniflora, and Ar,nica alpina, that elsewhere data from these parts are not available: the occur Inainly in the >>cloud belt>> 1100-1300 n1 nearest meteorological station on the Swedish s. m., on mountain crests that become free from side is situated in Kvikkjokk, east of the moun snow very early in the summer or remain bare tain range. Also with regard to the climate, even during the winter. Their absence north my own, sporadic and amateurish, observations of I.ake Vastenjaure is, probably, in the first will have to serve for real facts. The west- . place due to historical causes. However, they winds prevail, also in the summer. They do possibly require the moisture from the clouds not by any means always bring rain; as a rule in order to be able to live in these dry habitats1 they are attended by fairish weather. Still, and the gap under discussion may be connected in the Virihaure basin, where they have not with this need. This seems particularly prob- confronted any high mountains on their way 150 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
from the Atlantic, their n1eeting with the high vestigation being, as yet, unpublished but for massifs occasions afternoon showers that for a short summary (SANDBERG 1938). This prob long periods on end may be almost daily, and lem will, therefore, only be touched upon here, sometimes are turned into · heavy rains lasting without any pretence to thoroughness. for a day or two. Also in the summer, the precip In our region solifluxion may act upon an itation comes chiefly from westerly winds. The entire mountain-side so intensely that hardly a Vastenjaure and Sallohaure basins are, contra square yard remains unaffected. In view of riwise, screened fron1 the sea by rather high, the factors determining the extent of solifluxion, glacier-covered massifs, e.g., Mt Kuotelis, Mt i.e., in addition to the gradient of the slopes, the Veikdalsisen and Mt Reinoksfjell, where the occurrence of easily weathered rocks yielding west winds discharge most of their moisture fine-grained soil, the amount of precipitation, (cf. fig. 26). When a westerly wind brings rain and the frequency of perennially frozen ground on Lake Virihaure, the sky is often clear together with the resulting level of the subsoil northwards. water (cf. BESKOW op.c. pp. 622 ff. , LUNDQVIST East winds are much rarer. Especially when 1948 pp. 369 ff., DAUBENMIRE 1947 p. 69), coming from the north -east they regularly bring it goes without saying that solifluxion is wide rain on the eastern side of the mountain range, spread particularly in the mountains south but less often on Lake Virihaure which, on these and south -east of Lake Virihaure. occasions, is in the rain-shadow of the Sarek When solifluxion is intense enough to trans massifs. However, when it rains here with form the ground into a semi-fluid mush, it an easterly wind the downpour will go on long becomes fatal to vegetation. In most such locali and without interruption, now and then grow ties no vascular plants at all, and in places not ing into torrents. Even then precipitation would even bryophytes, are to be found (cf., e.g., seem to be less heavy round Lake Sallohaure, GJAEREVOLL 1949 p. 72). In its less extreme where Mt Akka, Mt Kisuris, Mt Stuor Niak forms solifluxion seems rather to favour certain and the other high mountains to the east come plants confined to mineral soil, partly by nearer, forming a more effective rain-screen mixing the material and bringing lower strata · than on Lake Virihaure. A certain similarity of unleached, basic soil to the surfaee (cf. TENG to the coastal region of Helgeland may be WALL 1925 pp. 739 ff., NORDHAGEN 1928 p. 536), noticed also in . regard to the local climate. the resultant pH sometimes exceeding 9 (SAND There, as well as north of Lake Vaste.njaure, BERG op.c. p. 335), partly by giving rise to naked an otherwise pronouncedly oceanic area appears >mew soib, watery immediately below the soli . to have a considerably lower precipitation than fluxion terraces (>>flytjordsvalkarna>>), and more the adjacent mountains, e.g., only 846 mm or less dry in parts of their surfaces. in Bod0, as against nearly 2000 mm in the moun Furthermore, solifluxion reduces competition tains further inland. by keeping out most dwarf scrubs, which on Among the phenomena attending the heavy flowing (or strongly frost-heaved) soil are rains and the abundant supply of melting-water restricted to limited patches of a more stable in the Virihaure basin and, especially, in the character, for instance the upper edges of old high mountains to the south and south -east of solifluxion terraces, and, to a less degree, by the lake, the extensive and most violent soli reducing the vitality of Salix herbacea and S. fluxion is particularly important to the vege polaris. This will, evidently, favour the arctic tation. (Regarding the connexion between epibiotics, which generally possess little competi precipitation and solifluxion cf., i.a., BESKOW tive power. 1930 p. 636.) The effect of solifluxion on the Apparently, the above holds more or less flora has as yet not been satisfactorily ex good also of the effects of frost-heaving on ground amined, the results of G. SANDBERG's close in- with little or no boulders. Distributions within the Region 151
Not a few plants seem to occur, exclusively or nearly so, on ground affected by solifluxion, i.a., several among the arctic or arctic-alpine epibiotics of the Virihaure region such as Carex bicolor (in its primary, high-alpine stations), Luzula arctica, Sagina caespitosa, Draba cTassi folia, and Potentilla hypaTctica. Most other spe cies of such origin would also appear to benefit by solifluxion, e.g., Poa heTjedalica, Garex parallela, M elandrium apetalum, DTaba lactea, Dr. alpina, Pedicularis hirsuta, P. flammea, Erigeron unalaschkense, and A ntennaria carpa thica. Carex microglochi·n and C. aTctogena are especiallly characteristic of solifluxion soil that is thoroughly soaked during the melting of the snow and later in the summer dries up com pletely. These species abound, for instance, in such localities at the western foot of Mt Kerke vare. Practically all these plants are lacking or rare north of Lake Vastenjaure; Of course, soli fluxion occurs also here, for instance locally in the southern steeps. However, it seems to Fig. 30. Solifluxion soil on the southern slope of Mt Tuipe. date, mainly, rather far back (cf. LUNDQVIST Photo Nils Dahlbeck July 1942. op.c. p. 396), and over large areas of the level table-lands it is quite insignificant. Rather ex tensive solifluxion is characteristic only of Mt heaths and dry southern steeps. The following Tuip� in the south-eastern part of the district plants are lacking there: (cf. fig. 30), where, in fact, DTaba alpina, Dr. Asplenium viride Braya linearis lactea, and RhododendTon lapponicum are to be Roegneria scandica Saxifraga adscendens seen. The remaining >>Virihaure species>> ob Carex nardina Primula scandinavica served within the northern district were mostly 0. glacialis Pr. stricta M inuartia rubella Euphrasia lapponica found in small patches of solifluxion soil or Arenaria norvegica Arnica alpina polygon ground on Mt Stupirvare and/or the Draba hirta nameless mountain south of Mt Paikas. On the other hand a number of species evi Furthermore, W oodsia alpina, "OxytTopis lap dently are at a disadvantage when precipitation ponica, and Veronica fruticans, reported from and the resulting solifluxion are too abundant. Mt Jeknaffo by �rENGWALL (1924) though not Mt Jeknaffo, where solifluxion is more intense observed by me during my repeated visits than anywhere else within our region, possesses there, do not, in all likelihood, occur on the a very rich flora and is rather well-investigated, mountain proper but on the hills at its northern at least partly. In spite of this, several plants foot. were not found there, although they occur on With the exception of Minuartia rubella, the adjacent mountains. Their absence on Mt Arenaria norvegica, and Arnica alpina, all Jeknaffo can, in most cases, not be put down these plants occur in n1ore or less numerous to any other causes than too much irrigation, stations north of Lake Vastenjaure; several intense solifluxion, and lack of eroded wind- are among the character plants of this area. 152 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Accordingly, · the factors causing solifluxion calcareous koli schists and the same topography; would seen1 to be of great importance, posi especially their eastern steeps seem everywhere tively as well as negatively, with regard to the to offer the same favourable conditions to alpine distribution of the plants within the Virihaure plants with little competitive capacity. In spite Vastenjaure basin. of this the number of >>Virihaure specieS>> con In many cases it is dif.ficult or impossible to tinually diminishes from north to south. The decide whether historical factors or topograph southern mountains do not possess any species ical ones are decisive: generally their trend not recurring farther north. On the other hand, seems to be about the same. This is, however, Luzula arctica and Sagina caespitosa do not rather self-evident, since topography and the occur south of their isolated outposts on Mt ensuing local climate must partly have exerted Karranis, where Garex microglochin and 0. about the same influence at the end of the ice arctogena also come to a stop. Poa arctica, P. age as nowadays. herjedalica, Sagina intermcdia, Cardamine pra tensis ssp. angustifolia, and Veronica fruticans
Climate do not reach further south than to Mt Karte vare. Carex parallela has its local southern Edaphic and topographic factors, accordingly, limit on Mt Vuossavare; Kobresia myosuro�des, seem to have co-operated with the historical Carex nardina, Cerastium arcticum, Draba alpina, ones in creating the conspicuous line of demar Potentilla Chamissonis, Oxytropis lapponica, and cation between the rich area and the adjacent ErigeTon unalaschkense ha.ve theirs on Mt tracts. However, a ·closer study of the dividing Katnjunjes; while the southernmost station of line reveals that this explanation does not cover Carex glacialis, Chctmorchis alpina, Gentianella all the known facts. tenella, l!Juphrasia lapponica, PediC'ltlaris flam True, the sharp boundary line to the east mea, and Arnica alpina is to be found on Mt and south is sufficiently explained through the Rivkonjunjes. Only a smaller number of more change in the nature of the bedrock. And the or less rare alpine plants reach the southern topography may be decisive with regard to limit of the koli schists on Mt Kuratjakko, e.g., the thinning out of the >>Virihaure species>> to the Asplenium viride, Woodsia glabella, Min�tartia north-east, on the vast table-land of Padjelanta. rubella, M. stricta, Melandrium apetal�tm, Draba The farther to the north past Mt Allak and Mt lactea, Dr. niva.lis, Potentilla nivea, Rhodon Mattaive, the more flat-domed become the dendron lapponicum, Cassiope tetragona, Pedicu mountains. Because of the evenly rounded laris hirsuta, Ping�ticula alpina, Campanula shapes without high peaks, precipices, &c., uniflora, and Antennaria carpathica, most of where the snow could remain, nwre or less trivial, which go far to the east in Sarek. Future, closer dry scrub heaths of a Dryadion type, unsuitable investigation will, no doubt, correct the details to many >>Viri'haure species>>, predominate. This of this picture; the principal features will, how fact, however, does not explain the gradual ever, certainly remain unchanged. disappearence aJso of species which normally The same trend is manifest on the eastern belong to the Dryadion communities. Yet, boundary of the rich area, i.e. on the other side this area is too little known to admit of any of the Tarra valley. Among the western species definite conclusions. that reach the western side of the valley, some Above all, the shape of the diffuse boundary do not cross it at all, e.g., Carex microglochin, of the rich area on the south-east, in the moun J uncus arcticus, Luzula arctica, Sagina caespi tains on the Tarra valley, cannot be fully ex tosa, Minuartia rubella, and Euphrasia lappo plained either by geological or by topographi nica; others only in a place or two, e.g., Asple cal facts. All these mountains have, in the nium viride, Poa herjedalica, and Pedicularis main, the same bedrock of easily weathered, flammea. Most such plants do not reach further Distributions within the Region 153 east or south than to the north-westernmost growing season is fairly plentiful and the mountains, viz. Mt Puokkik and Mt Luoppal Rnow, while being sufficiently deep in the winter, varatj, and in some cases to the western side does not melt away too late in the spring. There of Mt Velkispuolta. The southern and eastern fore, it is rather surprising that Pedic·ularis sides of this large massif are far poorer. On its flammea suddenly disappears a good distance south slope there are, for instance, mighty within the boundaries of the rich area, being, dolomite screes of exactly the same type as, for instance, absent from practically all the forty kilometers to the west, abound in rare baso mountains east of the upper Tarra vaUey, where phytes; however, on Mt Velkispuolta they con suitable habitats abound. tain no notable plants but Kobresia myosuroides, Evidently, the facts compiled above cannot Carex nardina, and 0. glacialis. The southern be explained through the geological conditions. most mountain among those . discussed here, To this is to be added that certain amphicline the commanding Mt Sammartjakko, should, species display a western tendency contrary supposedly, offer ideal conditions on its northern to the usual type of distribution of such plants. and north-western sides for fastidious alpine This is true of the. following species (the figures plants: irrigated slopes and damp precipices within brackets signifying, the first one the are plentiful, and in many places the soft schist number of stations within the rich area, Mt is so thoroughly weathered that it crumbles Sarvestjakko, Mt Tjagnoristjakko and Mt Para when squeezed. Yet, only the following notable varatj included, the second the number of those alpine plants were observed there: Asplenit(,m within the Sarek National Park and our region vi ride, W oodsia glabella, Minuartia stricta, Me outside of the rich area): landrium apetal'ltm, Draba lactea, Saxifraga Phtppsia algida (61, 27), Carex rufina (28, O)r Cotyledon, Rhododendron lapponicum, Veronica C. rariflora (46, 9), Luzula Wahlenbergii (61, 37), fruticans, Pedicularis hirzuta, Pinguicula alpina, Koenigia islandica (73, 25), and Sagina inter Campanula ·uniflora, and Erigeron unalaschkense, media (51, 21). all of which recur further east, on Mt Njunjesvare Also Deschampsia alpina and Festuca vivipara, and the eastern cambro-silurian. which are so frequent within the rich area that This trend is illustrated also by the distribu no stations were noted, are more or less rare tion of the separate >>Virihaure species>>, for cast of it. instance Pedicularis flammea. The probable immigration history likewise It n1ay be characterized as common in the fails as an explanation. To begin with, it would mountains immediately south and east of Lake be most extraordinary, if a great number of Virihaure. And it cannot be said to be ecologi species, strikingly dissimilar in · qualifications cally specialized to a very high degree, except for effective dissemination, biotype compounds, in so far as it is strictly basocole and evidently and competitive ability, should all have come welcomes · solifluxion. As distinguished from to a stop at very much the same line. And Pedimtlaris hirsuta it shuns snow-beds and ground further, the hypothesis propounded above (p. 129)· flushed by melting-water, which explains that regarding a distribution stop early in the post it is practically absent from the snowy depres glacial time pertains exlusively to high-alpine sion round Lake Ravejaure. Furthermore, it pioneer plants confined to mineral soils, and prefers the sunny sides of the n1ountains and has no reference to comparatively low-alpine is seldom to be noted above 1200 m s. m., on species occurring also below the upper limit of the one hand, or below the upper limit of the the willow thickets, e.g., Carex microglochi·n,. willow thickets, on the other. In other respects Chamorchis alpina, Oxytropis lapponica, and it is not fastidious, occurring in meadow com Pinguicula alpina. munities of many kinds and even in scrub The shape of the discussed line of demarca heaths, provided the irrigation during the tion can apparently only be explained by presup- 154 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
Fig. 31. Cloud-bank over the western mountains. In the foreground the Kvikkjokk delta and Mt Kaskaivo. posing that a climatic boundary line by chance met with on Mt Rivkonj unjes and the northern more or less coincides with those determined part of Mt Kuratjakko west of the river and on by geological and historical factors. Three Mt Sevvatjakko east of it; aceordingly, Mt Sam additional facts seem to point in the same martjakko and the southern part of Mt Kura direction. The >>Virihaure species>> restricted tjakko lie outside both the region of steep boggy to the northern mountains on the Tarra valley slopes and that of the proper >>Virihaure flora>>. largely belong to the category distinctly favoured Thus, the climatic boundary line seems to go by solifluxion, while those occurring also in between the atlantic local climate west of the the southern mountains of this area generally mountain range and the more continental cli belong to other ecological types. All viviparous mate east of it. Besides the tract near Riks grasses, which, as pointed out by NANNFELDT gransen, the Virihaure-Vastenjaure basin is the (1940 p. 24) and others, everywhere occur region where the frontier of Sweden comes chiefly in regions with a humid climate, have a nearest to the Atlantic: the distance from Mt western distribution within our region indepen Rakoktjakko to the Norwegian sea shore at Hel dent of their reaction to the chemical nature larvik is only 16 km. As is well known, this of the soil. And the steep mire slopes (>>back basin is among he most oceanic parts of Sweden, myTarna>> ), characteristic, as they are, of oceanic with an annual precipitation of 1800-2000 mm regions, are frequent in the regio subalpina (ANGSTROM 1946, map p. 61). On the other side around Lake Virihaure and in the upper Tarra of the mountain range the climate is much valley, while they will only be found exce_i)tio drier and more continental: Kvikkjokk has an nally, at least as far as steeply sloping ground annual precipitation of only 509 mm (LUND is concerned, east and south of the line where QVIST 1948 p. 228). As already mentioned, no
the >>Virihaure flora>> disappears. The southern exact meteorologic · data regarding the area n1 ost mires, definitely belonging to this type, are west of Kvikkjokk are available. However, Distributions within the Region 155
Fig. 32. The same cloud-bank six hours later. according to information volunteered by the the massif is being pelted with rain, while the inhabitants, wholly confirmed through obser weather is fine in its rain-shadow towards l; ake vations of my own during six summers, the Tarraure. Just as striking is the difference point where the Tarra valley bends round Mt between the wet west side of Mt Velkispuolta Sammartjakko and is crossed by the foremen and the dry east side; in the valley of Tjuolta tioned phytogeographical dividing-line coin vuopme, heavy downpours can often be seen ddes with the boundary line between the region in the mountains to the west, though not a where the easterly winds and that where ehiefly drop of rain is falling in the valley. the westerly winds bring rain, i.e. between the Conditions in the valley of the river Stora region approximating the neighbourhood of Lule alv wo'uld seem to be essentially the same Kvikkjokk as regards precipitation and that as in our region. However, in that valley no which in that respect resembles the Virihaure high mountains bar the way of the .Atlantic . basin. On a summer's day with a west wind air. The climatic dividing-line is, . therefore, at Tarreluoppal, the rain-laden clouds rolling to be found further east, in the neighbourhood in from the west will' often be seen to come, as of Suorva (LUNDQVIST 1944 p. 160). Most by magic, to a standstill towards Mt Sammar western basophytes are stopped by the belt of tjakko and Mt Staika, while to the east the sky acidic syenities, &c., west of Suorva; but those will generally be clear. Similarly, on fine sum able to force this obstacle reach the climatic mer's days at Kvikkjokk a cloud-bank is usually boundary line, adjoining, here, the area of the hanging over the mountains to the west, which eastern cambro-silurian. In other words, they may remain for days on end (cf. fig. 31, 32). Many reach further east than is usual in the valley a time a rain-curtain from the west is pulled of the river Lilla Lule alv, where no equivalents <>ver the mountains on the Tarra valley down to to the rich stations of western alpine plants in Mt Sammartjakko: the north-western side of the mountains round Stora SjOfallet are known. 156 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
The true criterion of the views presented exclude the local climate as a factor determining here is evidently whether the general phyto their distribution. geographical character of the plants that seem This probably holds good als9 of some species to have a climatically conditioned eastern limit with greater claims as regards the pH of the within our region actually indicates that they soil, which, for that reason, are still rarer in the have been favoured by the oceanic climate of forementioned areas of acid rocks, viz., as follows: the Virihaure basin. In order to form an opinion Kobresia myosuroides Potentilla nivea on these matters, it will be necessary to give a Carex parallela P. Chamissonis summary sketch of the distribution of the alpine 0. rupestris Pyrola grandiflora plants within Lule Lappmark, though this will, C. atrofusca Rhododendron lapponicum to some extent, be a repetition of facts already Cerastium glabratum Cassiope tetragona Minuartia stricta Veronica fruticans set forth. JJ,1elandrium apetalum Campanula uniflora Besides such alpine species as are frequent Draba nivalis in all the Scandes, the following ones have a tolerably uniform distribution within our region Woodsia glabella, Arnica alpina, and Anten and Sarek (cf. TENGWALL 1924; in view of the naria carpathica would also seem to belong to forementioned facts (p. 155), no attention will this category, though, as far as they are con here be paid to the mountains immediately cerned, the gaps in Sarek are still larger. That
: TENGWALL (1924 p. 673) reported Arnica alpina south of the river Stora Lule alv) . from but a small number of stations in Sarek Equisetum variegatum A rabis alpina may be due to his having ignored its habit of E. scirpoides Sedum Rosea Bot1·ychium boreale Saxifraga oppositijolia occuring in a sterile condition and, conse Polystichum Lonchitis S. joliolosa quently, overlooked it. T1·isetum spicatum S. nivalis BJORKMAN's (1937) thorough investigation of Poa arctica S. tenuis the area round the magnesite deposit on }it P. glauca S. aizoides Apartjakko in easternmost Sarek, where the Oarex atrata S. cernua J uncus triglumis S. rivularis bedrock to a great extent is basic or circum J. biglumis S. groenlandica neutral, is particularly instructive. All the Luzula parvijlora Potentilla Orantzii basophytes just enumerated occur there, except Salix Teticulata Dryas octopetala Woodsia glabella, Kobresia myosuroides, Carex S. polaTis Epilobium anagallidijoli- atrata, Juncus triglumis, Saxifraga nivalis, Poten S. myrsinites urn S. lanata AngelicaArchangelica ssp. tilla nivea, P. Chamissonis, and the low-alpine Oerastium fontanum ssp. norvegica Veronica fruticans, while only seven western scandicum Gentiana nivalis species, viz. Carex arctogena, Luzula Wahlen M inuartia bijlora Pedicularis hirsuta bergii, Koenigia islandica, Cerastium wrcticum,. Erigeron politum Viscaria alpina Draba lactea, Pinguicula alpina, and Erigeron Silene acaulis E. uniflorum Draba norvegica P etasites frigid us unalaschkense, occupy more or less isolated Oardamine bellidijolia stations within the area. · In addition Ranun culus gladalis and R. nivalis were found; though The basophytes among these species have a distinctly thinning out to the east, they do not smaller absolute number of stations in Sarek seem to be completely lacking in any high-alpine and the low eastern mountains than further area of this size. west, owing to the large areas occupied by Sometimes the lower frequency in Sarek is syenites and other acidic rocks where basocole wholly or partly caused by the fact that the plants cannot live. However, in proportion to species in question are low-alpine and never, or the number of possible habitats, their relative but exceptionally, overstep the upper limit of frequency is uniform enough as practically to the willow thickets and, therefore, do not thrive Distributions within the Region 157 in high -alpine areas. This holds true, i.a., of (cf. BJORKMAN 1939 p. 117), and R. nivalis Cystopteris montana, Carex capitata, Salix ar (cf. ibid. p. 118), which are so frequent in the buscula, Draba hirta, and Myosotis silvatica ssp. western mountains that no figures of the number frigida. of their stations can be given. Also Erigeron On the other hand, a considerable number of boreale belongs, no doubt, to the western type, species are so much more frequent within the though the number of its stations cannot be rich area and in the whole western part of our stated, since the reports from Sarek by TENG region that their distribution cannot be ex WALL (op.c. p. 672) are hardly reliable, and, . plained simply by the greater number of suit likewise, Carex norvegica s. str., the distribution able habitats. This holds good particularly of of which is insufficiently known, since TENGWALL the plants exclusively, or almost exclusively, was, naturally, unable to distinguish it from restricted to the rich area, as follows: Carex angarae . .Accordingly, the number of western alpine Festuca rubra v. mutica Cardamine pratensis spp. Roegneria scandica angustifolia plants, the distribution of which cannot be Garex nardina Drabct alpina explained through their edaphic demands, is 46. G. rufina Dr. crassifolia Before entering upon the question whether G. bicolor Braya linearis their total ranges are oceanic or not, a few preli G. glacialis Potentilla hyparctica minary remarks should be made. rr o classify Luzula arctica Astragalus norvegicus Sagina caespitosa Primula scandinavica arctic, more or less circumpolar, species as Minuartia rubella Euph?'asia lapponica oceanic or continental seems to rne almost Arenaria humifusa Pedicularis flammea unfeasable and, furthermore, rather meaning A. no'rvegica Antennaria lapponica less. HULTEN writes (1937a p. 60) as follows: >>It should be remembered, that all arctic plants .Among these plants Carex glacialis has two are to be regarded as more or less oceanic, as Roegneria scandica, Carex nardina, stations and they are confined to a strip along the shore Braya linearis and one station each outside of an ocean>>. On the other hand, BociiER the rich area; the rest are entirely restricted (1938 p. 257) proclaims a contrary opinion of to this area, and none of the species occur in his >>high-arctic types>>, as follows: >>All these Sarek proper. types are relatively psychrophilous and none .A western distribution characterizes, though are outstandingly oceanic, as even the coastal less pronouncedly, also the following species parts of the high-arctic regions may be regarded (concerning the purport of the figures, cf. as being fairly continental. ... In addition, the above p. 153): high-arctic species are relatively continental,
Asplenium viride (25, 8) 0erastium arcticum ( c .l7 0, their main area being in the rainless high-arctic Poa herfedalica (34, 5) 38) region>>. If an opinion not based on personal Phippsia algida (61, 27) Sagina intermedia (51, 21) experience of the high -arctic regions may be Garex microglochin (29, 3) Draba lactea (47, 10) volunteered, the most outstandingly arctic G. arctogena (42, 9) Oxytropis lapponica (60, plants would seem to be neither continental G. rariflora (46, 9) 3) Juncus arcticus (62, 16) Primula stricta (20, 6) nor oceanic, but adapted to the peculiar arctic Luzula Wahlenbergii ( 61, Gentianella tenella (59, 7) climate. .Among the characteristics of that 37) Pinguicula. alpina ( > 200, climate, worth particular notion in this con Ghamorchis alpina ( 64, 8) 9) nexion, within the region n1ost akin to Fennoscan Leuchorchis albida (50, 6) Erigeron unalaschkense dia, viz. >>the comparatively humid East .Arctic Koenigia islandica (73, (55, 10) 25) (Spitzbergen, the Franz J oseph archipelago, Novaya Zemlya)>> (GELTING 1934 p. 204), should To the same category belong Deschampsia be remembered that the growing season of alpina, Festuca vivipara, Ranunculus glacialis the plants is characterized by constantly recur- 158 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark ring fogs and a chilly climate with low maximum even in summer largely in the form of snow (cf. temperatures. VESTERGREN 1902 p. 259, who emphasizes the Counting among the purely arctic types the absence of vascular plants in the region of circumpolar species as well as those )>west abundant precipitation and high humidity of arctic)> species that occur both in the oceanic the air). For these reasons, it would seem to be regions, e.g., Iceland, and the continental ones, of comparatively little use to the vegetation. e.g., the Arc_tic Archipelago, a considerable Ecologically and with regard to the vascular number of plants with a western distribution plants, the interior of Sarek should be appreci within Lule Lappmark belong to this category, ably drier than the low mountains to the west as follows: and the western border zone of the high moun Phippsia algida Sagina intermedia tains. Festuca rubra v. mutica M'inuartia rubella At least as regards the arctic plants, however, Carex nardina Arenaria humifusa this difference seems to be of less importance C. glacialis Ranunculus nivalis than the difference in cloudiness. As pointed C. rarijlora Draba lactea Luzula Wahlenbergii Dr. crassifolia out above (p. 154), this is very pronounced: L. arctica Potentilla hyparctica the number of clear days is far larger in the Cerastium a-rcticum Pedicularis flammea eastern than in the western mountains, and in Sagina caespitosa Erigeron unalaschkense Sarek, as in other high-alpine tracts, fog often At first sight it seems very strange that such hangs on the intermediate slopes, while the a relatively great number of arctic plants are crests are cloudless (cf. fig. 33), so that the in� concentrated to the western, comparatively low terior is exposed to the sun. According to mountains, while absent from or rare in the H. E. HAMBERG (1909), the difference in the mean really high -alpine area, i.e. Sarek. In some numbers of sunny hours during the summer cases this is certainly due to historical causes, (April-September) between K vikkjokk and the as pointed out above. But this explanation Virihaure-Vastenjaure basin is only about 100> cannot apply to those species, e.g., Luzula hours. However, there were, at that time, no W ahlenbergii, Ranunculus nivalis, and Erigeron meteorological stations between K vikkjokk and unalaschkense, which actually occur in Sarek, Mt Partefjallet, on the one hand, and Bod0,. though being much rarer there than further on the other. The figure for the Virihaure west. Research in the field leaves a general Vastenjaure basin must, therefore, have been impression, difficult to substantiate statistically, obtained through interpolation and does not. that the arctic plants go from bad to worse make allowance for the locally increased cloudi the farther they penetrate towards the interior ness which seems considerably to exceed that. of Sarek. of both Sarek and the coastal region. Nor does. The explanation can hardly be anything but the cloud caps of the mountains, which are,. the climate. The fact that the interior of Sarek apparently, of great importance to the vegeta-· has so high an annual precipitation as 1245- tion, appear from that figure. 1500 mm (LUNDQVIST 1948 pp. 228 f.) may The cloudiness of the rich area, most marked seem to be incompatible with the hypothesis at the altitude where the arctic plants chiefly that the relevant difference between the western occur, may be regarded as a sort of equivalent. and eastern parts of our region would consist, to the fogs characteristic of the Arctic. It. in the m0re abundant precipitation and the prevents the light intensity, which is always. more oceanic character of the former. How low in the Arctic (RUBEL 1930 p. 285), from ever, HAMBERG (1901) states that precipitation exceeding the tolerances of arctic plants and,. in Sarek strongly increases above a level of further, insolation , which in high-arctic lati 1000 to 1500 m s. m. Consequently, it falls mainly tudes is much feebler than further south, from on the highest, sterile parts of the massifs and causing high ground temperatures, injurious. Distributions uithin the Region 159
Fig. 33. The Sarek mountains seen from Lake Stuor Tata. In Sarek, as in other high-alpine tracts, fog often hangs on the intermediate slopes, while the crests are cloucUess. Photo Sten Selander 3.8. 1946. to arctic plants, as often as in the eastern , high However, to some extent it seems possible alpine area. That, nevertheless, the effect of to trace an oceanic and a continental element insolation is considerable even in the western even within the Arctic flora. The former con mountains is proved by figures given by CLEVE sists in such species, chiefly, of course, low (1901 pp. 64 f.). On the two occasions, at the a.rctic ones, as are confined to, or mainly occur, beginning and end of July, when measurings in the distinctly oceanic parts of the Arctic, -were made, the ground temperature exceeded i.e. the Aleutian Islands, the south-coast of that of the air by 10°, reaching +27° C. at Alaska and the adjacent parts of the Rockies, an altitude of about 1000 m s. m. on Mt Junkar where at all events the western slopes should tjakko, near the eastern border of the rich area. be regarded as fairly oceanic, Newfoundland :For the sake of comparison, a few figures and the region of St Lawrence, southern and may be given from so low-arctic a region as the south -eastern Greenland, Iceland, and the North eastern Kola Peninsula (Orlow near Ponoj), Scandinavian coastal region west of the North where KIHLMAN (1890, Beilage pp. XIII ff.) Cape. These plants thus occur on coasts south during the month of July, 1889, observed ground of the boundary of the pack-ice which, in the temperatures exceeding 17° C. only 6 times (22,0° summer, are more or less ice-free. Most of these one time, 19,7° one time). The largest difference coasts are washed be warm sea-currents. observed between the temperature at ground The continental-arctic group, on the other level and that at a height of 2 m above the hand, is made up of species avoiding the re ground was 8° (observed but one time); as a gions just mentioned and confined to, or mainly rule, the difference was only about 1 to 2°. occurring in, North-Russia, Siberia and North In view of these facts, it is understandable Greenland and on the north-coast of America that arctic plants, especially high-arctic ones from the Arctic Archipelago to northern Alaska, such as Luzula arctica or Potentilla hyparctica, i.e. on coasts with cold sea-currents and n1uch have assembled. in the low mountains of the pack-ice. Climatically a permanently frozen we'3tern area. sea may be compared to a continent; and to a 160 SELANDER: Floristic Phytogeography of South- Western Lule Lappmark certain degree this must hold true also of a The remaining species, viz. Asplenium viride sea which during, practically, the whole year (HULTEN 1941 p. 38), Carerc microglochin, Koe is choked up with pack-ice. nigia islandica, Draba alpina (HULTEN 1945a The western element of the flora of Lule p. 846), Astragalus norvegicus (HULTEN 1947a Lappmark embraces no representatives of the p. 1087), Gentianella tenella, and Pinguicula continental-arctic group. To the oceanic-arctic alpina (FAEGRI in LAGERBERG 1939 p. 91; not type belong, more or less definitely, the following in Iceland, GRONTVED 1942 p. 436), all occupy species: areas iu the interior of .Asia, in addition to
Deschampsia alpina (cf. Cardamine pratensis ssp. their Arctic or Arctic-Alpine ranges. Yet, they B6CHER 1938 p. 209) angustifolia cannot be characterized as continental. Carex Festuca vivipara Braya linearis microglochin, Koenigia islandica, and Gentia Carex arctogena Primula stricta (GRONT nella tenella are common in Iceland, probably C. rufina VED 1942 pp. 319 f.) the most oceanic of all arctic countries, where C. norvegica s. str. Erigeron boreale A renaria norvegica also Draba alpina is to be found. Furthermore, all except Asplenium viride, Koenigia islandica An Arctic-Alpine group can also be perceived, and Pinguicula alpina have an eastern limit in occuring in the mounta.ins of South and Central Fennoscandia, which evidently is not due to Europe, on the one hand, and in the maritime historical causes alone, but largely to the parts of the Arctic, and/ or :Fennoscandia, on transition from an oceanic to a more conti the other. .As these mountains can be said nental climate. In view of their total areas, mainly to form part of the suboceanic region of BocHER (op.c. pp. 48, 72) does not hesitate Europe (cf., e.g. , TROLL 1925 pp. 328 ff.), this to classify Asplenium viride and Koenigia as group corresponds to the arctic-oceanie one. It definitely oceanic. It is obvious that the present comprises the following species: ranges of these plants were shaped, first and Carex bicolor Leuchorchis albida foremost, by the redu,ction and splitting up of Juncus arcticus coil. Ranunculus glacialis thir areas during the ice-age and that their Chamorchis alpina Central-Asiatic populations have, in the course Oxytropis lapponica possibly also belongs of time, adapted themselves to climatic con here. Its range is most peculiar, embracing tbe ditions absolutely foreign to the isolated, North Alps, Scandinavia west of Porsanger, where European populations that may rather be char it is one of the most pronouncedly coast-bound acterized as oceanic. among the alpin.e plants, and a large area in The above facts may be summed up as follows. Central Asia from Altai to Himalaya. Bearing None of the forementioned 46 species are con in mind how very critical the genus Oxytropis tinental. 19 are purely arctic and would, there is in Asia, it is not out of the question that the fore, be favoured by the cloudiness and de 0. lapponica of Central Asia may be something creased insolation of the oceanic local climate. apart from our species; it must, at any rate, 14 are arctic-oceanic, or they belong to related belong to an altogether different ecotype. Also types. 5 are endemics with oceanic ranges in rnorphologically, our plant, var. marmorata Scandinavia. The remaining 8 species are unde Nordh., is slightly different from, at any rate, fined, or difficult to interpret in this respect. the race of the Alps (NORDHAGEN 1940 p. 369). For the sake of comparison, the alpine and Four species are endemic, viz. Poa herjedalica, subalpine plants with eastern tendencies in Lule Roegneria scandica, Primula scandinavica, and Lappmark may be considered. They are far E·uphrasia lapponica. All have a definitely fewer in number. Apart from such species as, oceanic distribution. To them has to be added like Carex angarae, C. rot·undata, Epilobium the neo-endemic Antennaria lapponica, not davuTicum, and Pinguicula villosa, may be sus known to appear outside our region. pected of being postglacial eastern immigrants Distributions within the Region 161 or, like Cryptograrnrna crisp a, Descharnpsia Alpine total range which is rather non -committal, atropurpurea, Alchernilla alpina, Epilobiurn . lac comprising areas both in Altai and in Ice tiflorurn, .E. Hornernanni, Gnaphaliurn s�tpinurn, land. All the remaining species are pronoun &c., are lacking or rare within the rich area, cedly continental, centring upon Central Asia owing to their being more or less acidophilous, and Siberia, while missing in Iceland and other hardly any eastern species remain but the arctic-oceanic areas. following ones: This difference in the general character W oodsia alpina, Alopecurus pratensis ssp. between the species with western ranges within alpestris, Salix xerophila (HULTEN 1928 p. 10), Lule Lappmark and those with an eastern S. hastata (ibid. p. 13, cf. BJORKMAN 1939 tendency indicates that, apart from the his p. 129), S. glandulifera (FLODERUS 1931 p. 128), torical and edaphical factors, the climatic Saxifraga Cotyledon, Potentilla rnultifida (HUL contrast between the oeeanic Virihaure-Vasten TEN 1929 pp. 60 f.), Astragal�f-S frigidus (ibid. jaure basin and the more continental eastern p. 99), and Veronica tenella. mountains determined the distribution of alpine As far as Saxifraga Cotyledon, elsewhere plants within our region. The distances involved pronouncedly oceanic, is concerned, the eastern (about 60 km, as the crow flies) also correspond distribution is solely due to the fact that the to the analogous distances established by GEL plant is sylvan-subalpine within our region. TING (1934 p. 227) in north-eastern Greenland. Among the rest, W oodsia alpina has an Arctic-
. '
12 - 496149 Sten Selander I VIII . CLIMATE OF THE ICE-AGE REFUGIA
Among plants that probably survived the age refugia. Phytogeographical data will carry latest glaciation in the refugium west of Mt us a good way further. The flora is a great re Sulitelma (cf. above p. 100.) none seem, as gistering apparatus from which the climates of previously stated, to possess a total range of a times gone by can be deduced with a degree of continental type. This indicates that, also dur accuracy that no speculations unsupported by ing the Wiirm glaciation, a maritime climate observable facts can attain. In my opinion, this prevailed in the refugia on the N orwegia.n west registering apparatus confirms that, during the coast, a conclusion that, of course, differs from glaciations, a more or less maritime climate pre the conception current among glaciologists and vailed on the Scandinavian coast west of the plant historians. According to them, the anti ic�c-cap. cyclones arising over the ice-sheet and causing As set forth by EILIF DAHL (op.c.), even dur dry, fohn-like downward winds produced a local ing glaciations with a high precipitation ice-free climate of a pronouncedly continental nature refugia with a rich flora may have existed in below the ice-border (cf., e.g., HANSEN 1930 and places where sufficiently high mountains rea NORDHAGEN 1935). EILIF DAHL (1946 p. 238, ched the sea, provided the firn-line lay at a 194 7) is almost alone in his opposite opinion. higher level than the lowest parts of the ice-free Without the slightest doubt, a highly conti area. DAHL (1947 p. 234) assumes that refugia nental climate prevailed east of the ice-cap (cf., existed on coastal mountains in 1\'l0re and Troms, e.g., HYYPPA 1936). However, as to the condi on the west-coast of Norway, while, according tions west of the ice-cap, hardly anything is to him, Finnmark comprised a >>tundra refu known, i.a., because observations regarding the gium>> which owed its existence to a continental contact of an inland ice with a warm sea cur local climate with no more winter precipitation rent like the Gulf Stream cannot be made any than the sun could melt during the summer. where in present times. That proximity to a DAHL presupposes refugia on coastal mountains land-ice does .not of necessity produce a conti in Iceland and South-Greenland and, probably, nental local climate, even though the sea cur in Scotland, the Faroes, and Labrador, and rent skirting along the coast is far from being tundra refugia in Siberia, northern and eastern as warm or strong as the Gulf Stream, is born Spitzbergen, North-Greenland, Arctic Canada, out by conditions in southern and south-eastern and possibly on the Kola Peninsula (1947 pp. Greenland, where the climate must be charac 234 f.). In addition, a refugium must have existed terized as markedly maritime (BocHER 1938 also on the coastal mountains of N ordland (Nor pp. 9 ff. ). In Greenland this is indicated also way) in order to explain the occurrence in these by the occurrence of plants as oceanic as, for parts of, i.a., Carex scirpoidea, Saxifraga .Aizoon instance, Juncus squarrosus, Polygala serpylli ssp. Laestadii, and Potentilla hyparctica (cf. folia, and Armeria maritima. TENGWALL 1913 p. 269). The refugia in New However, we need not content ourselves with foundland and Gaspe, which DAHL does not vague assumptions as to the climate of the ice- classify, were also situated on coastal mountains Climate of the Ice-Age Refugia 163
(cf. FERNALD 1925). On the other hand, to p. 116), Carex pediforrnis (HULTEN 1927 p. 200), judge from the flora, the refugia in northern C. rnisandra (HULTEN 1942 p. 374), Salix po Labrador probably belonged to the tundra type laris (LAGERBERG 1940 p. 153), S. glanduli (cf. ABBE 1936 pp. 132 ff.). fera, Stellaria crassipes (HULTEN 1943 b map p. DAHL imagines that the refugia on coastal 258), Clematis alpina ssp. sibirica (HULTEN 1928 mountains for the most part consisted of nuna p. 119), Aconitum septentrionale (cf. above p. takks, though he adds that in all likelihood 85 and M. FRIES 1949 pp. 54 ff.; has spread there were also >>ice-free areas down to the sea far northwards in postglacial time), Draba hirta, level>> (op.c. p. 236). In view of the ecological Draba fladnizensis (HULTEN 1945 a p. 853)� character of some indubitable survivors (e.g., Saxifraga hieraciifolia (ibid. p. 919), Astragalus M yricaria germanica and Galium pumilum ssp. frigidus (has spread far in postglacial time), N ormani), this assumption seems fairly inevit Pedicularis Oederi (HULTEN 1930 p. 120), Aster able. sibiricus (ibid. pp. 158 f.), and Artemisia norve Thanks to DAHL's theory, it is now possible gica (ibid. p. 180). Carex parallela and Potentilla to make a finer distinction between oceanic and nivea s. str. also, possibly, fall within this cate continental arctic species. Species that evidently gory. survived the glaciations only in refugia on coastal Predorninantly oceanic, in the sense defined mountains (and in some cases in the Alps, &c.) above, are the following non-endemic species are manifestly oceanic, while those that sur within this group: vived only in tundra refugia (and in some cases Poa laxa ssp. flexuosa (NANNFELDT 1935 pp. in Central Asia, &c.) should be regarded as more 54 ff.; has spread far to the north in postglacial or less continental. An analysis of the distri time), Carex arctogcna, C. bicolor, Juncus arcti bution of the different floral elements in the cus, Charnorchis alpina, Nigritella nigra (WI Norwegian refugia decidedly bears out DAHL's STRAND in LAGERBERG 1937 pp. 196 ff.), Koeni opinion. In the following summary the words gia islandica, Arenaria norvegica, Ranunculus oceanic and continental are used in the sense platanifolius (LAGERBERG 1940 p. 137), Carda defined above; it should be expressly stated rninopsis petraea (GRONTVED 1942 p. 24 7), Sedum that the classification of several species is in villosum (ibid. p. 259), Saxifraga Aizoon ssp. evitably rather arbitrary. The survey is very cu-Aizoon (ibid. p. 260), Primula stricta (ibid. incomplete, as it comprises only plants the p. 319), Gentiana purpurea (HULTEN 1930 p. 66, hibernation areas of which are known with a the report from Kamtchatka most probably reasonable amount of certainty and as, further, erroneous, ibid.), and Campanula barbata (HEGI all endemic plants are passed by as well as all VI: 1 p. 336). plants whose total ranges are more or less non The following relevant species possess an committal. References regarding the distri arctic area without definitely oceanic or conti bution are OJ?-lY given once for each species and nental features: altogether omitted when the total range, in Phippsia algida (HULTEN 1942 p. 144), Carcx cludjng sources, has been given earlier in this glacialis, C. rariflora, Luzula parviflora (HULTEN work. 1943a p. 441), L. arctica, Cerastium arcticum, Among plants occurring in the Central-Scan Sagina caespitosa, Minuartia rubella, Ranun dinavian mountains and, in some instances, culus hyperboreus (HULTEN 1944 p. 758), R. northernmost Scandinavia, while missing in nivalis (ibid. p. 761), Draba lactea,, Dr. nivalis Nordland and the adjacent parts of Lapland, (Greenland should be added to the area reported the following represent the continental type of by HuLTEN, 1945a pp. 862 f., cf. BoCHER distribution: 1938 p. 106), Saxifraga foliolosa (HULTEN op.c. Athyriurn crenatum (HOLMBERG 1922 p. 19), p. 917), S. groenlandica (ibid. p. 910), Rhodo Phippsia concinna (GELERT & 0STENFELD 1902 dendron lapponicum, and Campanula uniflora. •
164 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
The general phytogeographical character of their centres of survival cannot be located, viz. the plants than can be supposed to have passed the following: the Wiirm glaciation also, or exclusively, in Carex 'rufina, Salix arbuscula (FLODERUS refugia on the coast between the Trondheim 1931 p. 114), Ranunculus glacialis, Saxifraga Fjord and Troms is distinctly dissimilar. Cotyledon, Alchemilla W ichurae, and Veronica Within this group there is only one seemingly pumila. continental species, -viz. Calamagrostis obtusata .Among the plants that have, in all prob (HOLMBERG 1922 p. 155). However, according to ability, survived the latest glacation in Troms NYGREN (1946 pp. 195 ff., 254) our plant which is and/or Finnmark, the oceanic element plays endemic in Fennoscandia (Lapland, Kola Pe but an inferior part. If species with ranges ins.) and should properly be called 0. chalybaea continuous southwards be excluded, this ele (Laest.) Fr. is not identical with the closely re ment will comprise but the following spe lated 0. obtusata of Central .Asia, being, i.a., cies that, moreover, are totally absent from aporp.icticwhile the latter is amphimictic. It may East-Finnmark and \V est-Finnmark east of have arisen from the hybrid C. arundinacea x Porsanger Fjord: canescens, or from C. obtusata. In any case, we Carex bicolor, Nigritella nigra, Ramtnculus need not take it into account in the present con platanifolius, and Braya linearis. nexion. The purely arctic element is somewhat bigger The purely arctic species belonging to this in number, embracing the following species, category are rather numerous, as follows: among which, however, none but Ranunculus Carcx scirpoidea (HULTEN 1942 p. 304), C. sulphureus seem to have hibernated in East nardina, Luzula arctica, Sagina caespitosa, M i Finnmark: nuartia rubella, Arenaria humifusa, Draba lactea, Carex na1·dina, Luzula arctica, Sagina caespi Dr. crassifolia, Potentilla hyparctica, Pedicularis tosa, Arenaria humifusa, Ranunculus sulphureus flammea, and A ntennaria Porsildii. (HULTEN 194-4 p. 770), Draba crassifolia, Rho To these plants should, probably, be added dodendron lapponicum, Pedicularis flammea, some species reported by RuNE (1945b, 1947, Campanula uniflora, Erigeron unalaschkense, 1948) from J..� ycksele and .A.sele Lappmark and, Antennaria Porsildii, and A. carpathica. maybe, isolated there, viz. the following: The continental species, on the contrary, are Draba nivalis, Rhododendron lapponic·um, Cam very numerous, as follows: panula uniflora, and possibly Carex parallela. Dryopteris fragTans (HULTEN 1941 p. 25), However, the last-mentioned species probably Hierochloe alpina (not typical; occurring also has a distribution continuous northwards and in Newfoundland &c., HULTEN 1942 p. 133, and may, thus, have >>hibernated>> in northernmost in South-Greenland, BOCHER op.c. p. 212), Alo Norway. In the following it will, therefore, be pecurus pratensis ssp. alpestris, .A.Tctagrostis excluded. latifolia (HULTEN op.c. p. 148, cf. NORDHAGEN Among the species that, in all likelihood, 1935 p. 124), 11risetum subalpestre (HULTEN . hibernated in these pa1·ts, the following are more 1926 p. 176), Poa arctica ssp. caespitans (NANN or less oceanic: FELDT 1940 pp. 73 ff.), Roegneria mutabilis Carex rnacloviana, C. bicolor, Arena,ria nor (HULTEN 1927 p. 150), Scirpus pumil1�s (NORD vegica, Cardamine pratensis ssp. angustifolia, HAGEN 1935 p. 93), Carex holostoma (NYGREN Braya linearis, Sedum villosum, and Primula 1936, map p. 148), 0. misandra, Veratrum album stricta. ssp. virescens (HULTEN 1937a, map p. 112), Al · However, a few more plants, all of them defi lium Schoenoprasum ssp. sibiTicum (HULTEN 1943 nitely oceanic, have an eastern limit in Troms pp. 453 f.), Platanthera oligantha (HULTEN 1927 or West-Finnmark and, therefore, evidently p. 266), Salix xerophila (FLODERUS 1931 p. 87), hibernated on the Norwegian west coast, though S. glandulifera, Stellaria m·assipes, Arenaria Climate of the Ice-Age Refugia 165 ciUata ssp. pseudofrigida (NORDHAGEN op.c. In West-Finnmark and Troms the continental pp. 46 ff.), Melandrium angustiflorum (HuL'rEN element is slightly larger in number than the 1928 p. 92), Papaver Dahlianum (NORDHAGEN oceanic and arctic elements put together: out 1931 p. 44), P. lapponicum (ibid. p. 13), Draba of 32 non-endemic plants with isolated areas fladnizensis, Braya purpurascens (HULTEN 1945a between Porsanger and Lofoten, 14 are conti p. 888; also in Newfoundland, &c., and, ace. nental, 4 oceanic, and 7 purely arctic. BOCHER op.c. p. 102, in North-Greenland), In Nordland and the adjacent parts of Lap Saxifraga hieraciifolia, Chrysosplenium tetrand land, on the other hand, the oceanic and arctic rum (HTJLTEN op.c. p. 950), Potentilla multifida, elements predominate to about the same P. Chamissonis, Astragalus alpinus ssp. arcticus degree as the continental element in East (HULTEN 1947a p. 1083), A. frigidus, Oxytropis Finnmark. Apart from the endemics, at least deflexa (ibid. p. 1240), Cassiope tetragona, Ar 28 relevant plants can be supposed to have meria scabra ssp. sibirica (HULTEN 1948 p. 1297; hibernated somewhere in the refugia between also in Newfoundland, &c.), Polemoniurn acu Troms and the Trondheim Fjord. Out of these tiflorum s. str. (ibid. p. 1323), Pedicularis hir plants, 12 are oceanic, and 16 arctic; none are suta (BOCHER op.c. p. 172), and Crepis rnulti continental. caulis (NORDHAGEN 1935 p. 120). The continental element reappears south of It seems superfluous to deal with the hypo the Trondheim Fjord. Out of 48 relevant, non thetical refugia in or near the Kola Peninsula, endemic plants with isolated areas in the Cen the existence of which is indicated by the isola tral-Scandinavian mountains, no less than 17 are ted stations of, e.g., Calarnagrostis deschamp continental; 15 are oceanic, and 16 purely arctic. sioides, Lornatogonium rotatum, and the plants The general distribution of the different enumerated above p. 83; with a few exceptions, elements thus aceords with present climatical the survivors of these parts are pronouncedly conditions. The continental species are, largely, continental. confined to the continental regions, i.e. Finnmark As already pointed out, this survey is incom and the adjacent parts of Finland and northern plete and largely tentative. No doubt, a closer most Sweden, on the one side, and the South phytogeographical and taxonomical examination Norwegian mountains, on the other, while the of the whole alpine flora of Scandinavia would oeeanic element prevails in the maritime region render it more satisfactory. Unfortunately, I of Nordland and westernmost Lapland; the have not been in a position to undertake this north-eastern limit of several oceanic species time-absorbing task. The available data n1ay, in North Scandinavia nearly coincides with the however, suffice to give a general outline of limit of an annual precipitation of 500 mm. the distribution of the oceanic and continental Certain irregularities are apparently connected elements of the Scandinavian arctic-montane with postglacial changes of the clin1ate. Thus, flora. the effects of the warm period would have been In East-Finnmark the continental element more disastrous to psychrophilous plants in predominates. The 11 species with isolated low-lying Finnmark than in regions with higher areas in these parts which, therefore, probably mountains. This may conceivably explain why survived the last glaciation in the Varanger some species, e.g., Hierochloe alpina, Melandrium Peninsula are all continental. 12 non-endemic angustiflorum, Ranunculus glacialis, Papaver plants have isolated North-Scandinavian areas lapponicum, Saxifraga hieraciifolia, and Cas in the region between East-Finnmark and siope tetragona, are absent from East-Finnmark, Troms, although their centres of survival cannot though occurring both further west and in the be definitely established. All of these are con Kola Peninsula.. tinental except the arctic-circumpolar Ra The present position would, however, be nunculus sulphureus. incomprehensible, if a climate of a pronouncedly 166 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark continental type had prevailed during the isolated outposts south of these areas, e.g., Wiirm glaciation also in the coastal mountains Alopecurus pratensis ssp. alpestris, Salix glatpdu west of the ice-cap. The predominance of lifera, M elandrium angustiflorum, and Pole oceanic species within the regions that, in all monium acutiflorum, they are, without exception, likelihood, received the hibernating element situated on the eastern side of the mountain of their floras chiefly from refugia on the coastal range. mountains of western Norway may, seemingly, Apart from the few more or less ubiquitous be explained if these species be supposed to continental species, e.g., Carex aquatilis, Salix have survived a glaciation with a continental hastata, Viola biflora, and Pedicularis lapponica, climate in isolated stations where local con there is but one continental survivor in Helge ditions suited them particularly well and, land and South Lapland, viz. Astragal'ltS frigidus. further, to have obtained their present wider It seems to have hibernated in South Norway, distribution as the climate became more mari on the one hand, and in Troms-Finnmark, on time after the glaciation. However, why should, the other, and then to have spread north, re then, the continental element be almost absent, spectively south, without being able to fill up not only in the coastal mountains but in the the gap between its two areas; this, by the way, chief part of interior Lapland� If "the surviving indicates how continental ubiquitous plants flora west of the ice-cap had comprised conti of the type just mentioned may have obtained nental species in any number worth mentioning, their present distribution. The remarkable they could have followed the melting ice, in findings of rare alpine plants recently made by the vicinity of which, according to current RuNE (1945b, 194 7) in Asele Lappmark do not opinion, a markedly continental climate pre include any c.ontinental species; even as eastern vailed, and proceeded to stations of a continental a massif as Mt l\iurberget and Mt Graipesvare character east of the watershed, where they has a flora of an oceanic or purely arctic char must have been able to survive even after the acter, comprising, i.a., Oarex glacialis, Minuartia time when the alleged continentalizing effect rubella, Arenaria norvegica, and Draba norve from the ice-cap ceased. gica (RuNE 1947 p. 487). All the region from The actual position is, of course, quite diffe Troms to Opdal and South Jamtland harbours rent. The continental hibernators of the Lap practically no continental species which cannot land flora, a few ubiquituous species of great be supposed to have immigrated there in post spreading capacity excepted, occur almost ex glacial time. clusively in .the northernmost territories, where The continental plants of the South-Norwegian they have more or less continuous areas, only · mountains present problems of their own. Many exceptionally as far south as in Pite Lappmark. of them are highly isolated in their Central Unlike most alpine plants, they are at least as Scandinavian stations. AthyTium crenatum, Ca frequent in the eastern parts of the mountain rex pediformis ssp. rhizodes, Clematis alpina range as in the western parts, wherever the ssp. sibirica, and possibly Aster sibiricus, which number \of edaphically suitable habitats is here only occur in the conifer belt, n1ay, per equal. In several cases, they are even thinning haps, be supposed to have immigrated from out westwards. Only rarely, as in the case Finland during some postglacial period with of Hierochloe alpina, Carex misandra, Salix a continental climate (cf. EILIF DABL 1947 xerophila, Potentilla nivea, P. Chamissonis, p. 251). This supposition is supported by the Astragalus frigidus, Cassiope tetragona, Veronica occurrence of Clematis in an isolated station tenella, Pedicularis hirsuta, and Arnica alpina, on the Finnish coast of the Gulf of Bothnia are the mountains of Lule Lappmark wholly (cf. HuLTEN 1949b, map 798). As regards the or partly included in their . continuous areas. rest, such a history seems unimaginable: they When indubitable continental survivors have must be glacial survivors. However, had they Climate of the Ice-Age Refugia 167 actually survived in refugia on the coastal moun west coast of Norway would, during many tains of Mere, &c., why should they be lacking thousands of years, have been exposed to a in the vV est-Norwegian refugia farther north� severely continental climate, which must have There is no reason why the ice-age climate of favoured the continental eeotypes and elimi the coast of Mere should have been less oceanic nated the oceanic ones. than that of Nordland. Not improbably, the Below I shall, therefore, presuppose, first, continental South-Norwegian survivors hiber that the local climate of the West-Norwegian nated on inland nunatakks with a more con coastal refugia was definitely more inclement tinental local climate than that of the refugia than it is at present, though not much more on the coastal mountains. The existenc� in continental than now, even during the maxima Trollheimen of such a nunatakk with a hiber of the glaciations; and, secondly, that EILIF nating flora was recently suggested by N . .A. DAHL is right in presuming the existence of SoRENSEN (1949); however, its flora seems to tundra refugia with a low precipitation in have been preponderant1y oceanic in character. East-_Finnmark and of refugia with a high pre The nunatakks suggested in the present work cipitation on coastal mountains in western should have been situated further inland, and Norway. I shall, furthermore, assume that the more southwards. refugia of West-Finnmark and Troms had a The above data are, apparently, entirely climatic character intermediate between these incompatible with the theory that the par two types, and that in South Norway continen ticular part of the Fennoscandian rrwuntain tal refugia existed on inland nunatakks in addi flora that survived the latest glaciation on the tion to the maritime coastal ones. IX. BASOPHILY WIT HIN THE AL PINE "FLORA OF FENNOSCAN DIA
To Scandinavian botanists it is a matter of Roegneria mutabilis and R. scandica, Carex course that the rich and interesting flora is arctogena and C. capitata, Saxifraga tenuis and to be found on basic or circumneutral rocks, and S. nivalis, and Veronica alpina and V. pumila. that only very trivial plants occur on acidic Nor is it unlikely that some further amphicline magma rocks. However, this applies by no species, e.g., Astragalus alpinus (cf. TH. FRIES means to all alpine floras. In the .Alps, for 1925 p. 6), will be divided into smaller units instance, many local epibiotics and endemics differing in their reactions to the chemical are confined to areas of acidic rock. Broadly nature of the soils. However, such instances speaking, the flora on the �cidic rocks of the will certainly remain rare exceptions. Further .Alps does not seem to be much poorer than more, it should be .observed that, with the that on basic or circumneutral rocks: for in exception of Carex rotundata and possibly stance, the flora of the gneissic part of the Salix herbacea (cf. Du RIETZ 1942b p. 60, region of Trient comprises 310 species as against GJ.AEREVOLL 1949 p. 84), all the forementioned 390 species in the limestone part (J.ACC.ARD pJants prefer circumneutral soils, though to a 1930 p. 168), and a gneiss slope at Luisin con varying degree. Pairs ecologically as clearly tained 173 species as against 150 species on a distinct as the .Alpine ones are scarcely to be closely similar slope on limestone at Kiih-Dungel, found in Fennoscandia. and 107 species in a similar station on calcareous This is rather surprising. Genotypical depau schist at La Peulaz (ibid. p. 187). peration within the small populations surviving .Another well-known phenomenon,. with no in the Fennoscandian refugia has, no doubt, real equivalent i Fennoscandia, is the great. prevented differentiation on a large scale. number of vicarious, related types in the .Alps, Seeing that, nevertheless, new types actually one basophilous, the other acidophilous or arose within the genera of Papaver, Poa, Draba, amphicline. It will suffice to enumerate but &c., it is indeed st.range that practically none a few examples, where one of the parallel species of them differ in respect to their reaction to is represented also in Fennoscandia, e.g., Poa the chemical nature of the soil, as many com laxa and P. minor, Juncus trifidus and J. paratively young units within the flora of the monanthos, Sedum annuum and S. atratum, .Alps do. (An explanation may conceivably be Alchemilla alpina and A. Hoppeana, Campanula that, as is always the case within very small barbata and C. alpina, &c. populations, elimination prevailed against selec True, a few such pairs of species or races occur tion in the Fennoscandian ice-age refugia, also in Fennoscandia. Apart from the best adaption, consequently, being imperfect or known instance, Carex rotundata and 0. saxa lacking.) Furthermore, since the original bio tilis which, however, are not very closely related, type sets were hardly less complete in the pre there are Salix herbacea and S. polaris, Ce glacial or interglacial flora of the Scandes than _ 1 rastium alpinum and C. glabratum, and, possibly, in that 0f the .Alps, it is even stranger that so ex- Basophily within the Alpine Flora of Fennoscandia 169 ceedingly few acidophytes occur also among the W oodsia alpina Papaver radicatum call. old units of the Fennoscandian mountain flora. W. glabella Draba fladnizensis Kobresia myosuroides Dr. nivalis A great many of the alpine species common K. simpliciuscula Dr. alpina to Fennoscandia and the Alps also react diffe Carex scirpoidea Saxifraga Aizoon rently to the particular kinds of rock. In the C. nardina S. oppositifolia Alps not a few Fennoscandian basophytes occur C. rupestris S. groenlandica exclusively, or almost so, on acid, primary rock, C. misandra Potentilla nivea C. glacialis P. hyparctica e.g., Woodsia alpina, Kobresia simpliciuscula, C. capillaris Pyrola grandiflora Carex bicolor, Juncus triglumis, Minuartia stricta, Juncus castaneus Rhododendron lapponicum and Saxifraga Cotyledon; others are amphicline Luzula arctica Pedicularis flammea in the Alps, e.g., Selaginella selaginoides, Equi M inuartia rubella Campanula unijlora setum variegatum, Asplenium viride, Scirpus Melandrium apetalum pumilus, Kobresia myosuroides, Carex micro In these respects north-eastern Greenland, glochin, C. atrofusca, Arenaria ciliata (coli.), partly known thanks to investigations by S0REN Draba fladnizensis, Saxijraga aizoides, and Gen SEN (1933) and GELTING (1934), is less different tianella tenella; finally, many plants that are from Scandinavia, however substantial the amphicline in Fennoscandia are confined to, dissilnilarities. Thus GELTING mentions but or rare outside, the areas of primary rock in two basocole plants, viz. Rumex Acetosella top.c. the Alps, e.g., CaTex Lachenalii, C. brunnescens, p. 91; should be R. tenuifolius, A. l.JOVE in litt.) G. Bigelowii, Luzula spicata, Salix glauca, and Saxifraga Hirculus (ibid. p. 124), neither of Oxyria digyna, Cerastium trigynum, and Viola which is a basophyte in Fennoscandia. His montana. There is, thus, a general trend towards list of the remaining basophytes only comprises a decrease in basophily within the . flora of the the following Fennoscandian plants (ibid. p. 228): .Alps as compared to that of the Scandes. The Kobresia simpliciuscula, Carex parallela, C. opposite is only a rare exception, for instance, nardina 168), in the case of Arabis alpina and Solidago Vir 'talso on primary rock, ibid. p. C. misandra, C. atrofusca, C. capillarls, C. gaurea var. alpestris, which are amphieline, resp. acidophilous, with us and basophilous in saxatilis ()>to some degree)>, p. 172), C. rotundata C. saxatilis, 173, the Alps. (Data from the Alps according to (more basophilous than p. SCHROETER 1908 and HEGI 1906-31). though acidophilous in Fennoscandia), Juncus arcticus, Braya linearis The relationship of the flora to the bedrock (included by GELTING in the Arctic .Archipelago of America differs in Br. glabella), Br. purpurascens, and Saxifraga yet more from that met with in the Scandes aizoides. (cf. SIMMONS 1906, 1913, POLUNIN 1940). With According to 80RENSEN (op.c), Equisetum respect to the number of species, as well as of variegaturrn and Pedicularis flammea have to be individuals, the vegetation in the former region added. All of these plants but Equisetum variega is much richer on magma rocks than on sedi tum, Carex rotundata and Juncus arcticus are mentary rocks, particularly silurian limestone. basocole in the Scandes, while only more or less The more basic the soil, the poorer the vege basocline in N.E. Greenland. tation. On silurian limestones it becomes On the other hand the following species, mostly dwarfed, and deficient in species, and the rare amphicline in Fennoscandia, are characterized plants stick to acid primary rock almost as by GELTING (op.c. p. 228) as acidophilous in exclusively as to basic or circumneutral soil in north -eastern Greenland: Fennoscandia. At any rate, the following Hierochloe alpina, Poa arctica, P. alpina Fennoscandian basophytes are, in the Arctic (definitely basocline in Fennoscandia), Carex Archipelago, acidophilous or, in a few cases, scirpoidea (growing, in its only Scandinavian amphicline: station, on calciferous soil), Luzula arcuata 170 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
s. lat., L. spicata, Salix herbacea, Ranunculus instead, the edaphic factor principally decisive glacialis, Cassiope hypnoides (basocline in Fenno of the distribution of plants appears to be the scandia), Antennaria alpina (do.). age, and the resulting state of weathering, of The rest of the Fennoscandian plants occurring the lava rocks. in north-eastern Greenland, i.e. the majority To nly knowledge, there is only one region of our alpine flora, are evidently more or less where the position, in the respect relevant here, arilphicline there. Thus S0RENSEN (op.c. p. 170) is essentially the same as in the Scandes, viz., stresses the fact that W oodsia glabella is not Newfoundland and the adjacent parts of the calciphilous, while GELTING (op.c. p. 71) points mainland. FERNAI,D (1924 p. 563) writes of out that among the Drabae, whose alpine repre the epibiotics of Newfoundland as follows: sentatives are all basocole or basocline in Scan >>The very restricted distribution on the island dinavia, only Dr. Bellii Holm is calciphilous of the species confined to the region centering to any degree worth mentioning. The great on the Long Range is associated not only with majority of the character plants of the commu the ungla.ciated conditions of those tablelands nities that in Fennoscandia are confined to but with the fact that to a considerable extent basic or circumneutral soils are evidently they are composed of calcareous or magnesian amphicline in those parts of Greenland, e.g., rock and that the plants which center there Dryas octopetala (coll.; ibid. p. 98), Carex ru are such as usually occur- on these rocks>>. The pestris (p. 171), and Saxifraga oppositifolia forementioned plants are largely our Scandi (p. 128). navian, more or less rare, basophytes. Also in Stray notices from South Greenland subn1itted the mountains between New York and Gaspe by PORSILD (1920, 1930, 1935) and BOCHER Co. they behave mostly in the same way. Accor (1938) suggest that the relationship between ding to FERNALD (1907 pp. 158 ff., 1926, 1933), plants and bedrock, especially in the oceanic the following arctic-subarctic plants occurring region of south-eastern Greenland, rather re also in Fennoscandia. are, in the regions near the sembles that in Fennoscandia. BocHER, how Golf of St. Lawrence, confined to areas of basic ever, points out certain differences. Thus several or circumneutral rock (lirnestone, serpentine, or Fennoscandian basophytes are amphicline in in a few cases trap or slate; some plants ab these parts, e.g. , Car ex microglochin ( op.c. sent from either Newfoundland or the .American p. 228), C. rupest·ris (pp. 235 f.), C. glacial is mainland are also included):
(p. 232), C. capillaris (p. 223), and Pedicularis Selaginella selaginoides 0. glacialis flammea (p. 172). Rhododendron lapponicum Equisetum variegatum 0. capillaris is basophilous only in the outer, pronouncedly Botrychium boreale J uncus arcticus oceanic, fiord region,- but acidophilous on the Asplenium viride Tojieldia pusilla Oystopteris montana Ooeloglossum viride inner, more continental parts of the fiords Woodsia alpina Salix reticulata (ibid. p. 160). .And a few plants, e.g., Loise W. glabella Arenaria humijusa leuria procumbens (ibid. pp. 156 f.) and Diapensia Dryopteris fragrans Viscaria alpina lapponica (p. 163), that are amphicline in Fenno Polystichum Lonchitis Thalictrum alpinum scandia, are acidophilous in South Greenland. Poa alpina Draba norvegica Festuca rubra v. mutica Dr. hirta Conditions in Iceland can hardly be compared f. prolijera Dr. nivalis to those in the other arctic-subarctic countries Eriophorum brachyanthe- Dr. lactea on the North-.Atlantic and the Polar Sea, since rum Oardamine pratensis ssp. the Icelandic basalts and other n1agma rocks Kobresia simpliciuscula angustifolia everywhere appear to be basic enough to satisfy Oarex microglochin Saxijraga Aizoon 0. capitata s. lat. S. oppositijolia the wants of most arctic and arctic-montane 0. rupestris S. nivalis basophytes. As a whole, the flora of Irela.nd 0. maritima S. aizoides seems to be more or less basophilous, while, 0. bicolor S. groenlandica Basophily within the Alpine Flora of Fennoscandia 171
Potent-illa nivea Veronica pumila Finally, the following plants occur only or, P. hyparctica (on Gaspe Bartsia alpina in a few cases, mainly on quartzite and other Penins., PIERCE 1936) Pedicularis flammea silicious rocks in Newfoundland and/or on the P. Cmntzi·i Campanula unijl01·a Epilobium davuricum Erigeron boreale mainland of N. E. North America:
Pyrola grandiflora Lycopodium annotinum v. paludosa "\Vimm. & Gr.
pungens ( = spp. typica) Practically all these plants are basophilous L. alpinum Cardamine bellidifolia also in Pennoscandia, the only exceptions being Athyrium alpestre Saxijraga foliolosa the amphicline Botrychium boreale and Erio Woodsia ilvensis S. rivularis phorum brachyantherum; concerning Dryopteris Hierochloe alpina Sibbaldia procumbens Deschampsia atropurpu- Epilobium lactijlorum jragrans, see below p. 172. Characteristic of rea (also on trap) the serpentine barrens are in the first place Juncus trifidus E. H ornemanni Asplenium viride, .Arenaria humifusa, and Vis Luzttla conjus a Phyllodoce coerulea caria alpina (which is found only on serpentine); Salix herbacea (also on Cassiope hypnoides they are character plants of this rock also in our slate) Gnaphalium supinum Cardamine pratensis v. peninsula. Gn. norvegicum The following plants seem to be amphicline Part of these species are acidophilous also in both in Newfoundland and on the mainland Fennoscandia. However, many of them are (or in one of these regions if present only there) amphicline there, although not occurring on as well as in Pennoscandia (the position of limestone in the forementioned American sta some plants, however, being somewhat uncer tions. As regards the mainland, this may be tain, since PERNALD, particularly in FERNALD due to the fact that the relatively high-alpine 1907, does not always classify the rocks in the areas consist either of primary rock (Green Mts, way current among Scandinavian botanists): White Mts, Mt Katahdin, Table-top Mt) or of Lycopodium Selago v. ap Oxyria digyna serpentine (Mt Albert), while the limestone areas pressum Cerastium alpinum chiefly embrace cliffs, &c., at low altitudes. Botrychium matrica?·iijo - Sagina Linnaei Another explanation, which applies also to lium Silene acaulis Newfoundland, may be that many of these B. multifidum Ranuncttlus hyperboreus Sparganium hyperboreum R. pygmaeus species are very rare, occurring only in one or Phleum alpinum L. s. lat. A rabis alpina two stations, and, consequently, plants that Alopecurus aequalis Alchemilla ve8tita are actually amphicline have, by chance, been Agrostis borealis A. filicaulis found only in habitats of one and the same kind. Festuca vivipara Astragalus alpinus This may apply also to Botrychium boreale and E'riophorum Scheuchzeri Epilobium anagallidifo- Carex Lachenalii lium Eriophorum brachyantherum (cf. above), as well C. brunnescens Loiseleuria procumbens as to Luzula spicata, amphicline on the American C. Bigelowii ATctostaphylos alpina mainland and in Fennoscandia, but acidocole C. mriflora Vaccinium Vitis-idaea in Newfoundland; Cerastium trigynum, amphi Luzula parviflora (only ssp. minus) cline in Newfoundland and Fennoscandia, but Salix glauca Diapensia lapponica acidocole on the mainland of N. E. North Ame 0 erastium alpinum, Ranunculus hyperboreus � rica; and Juniperus communis ssp. nana and .Arabis alpina, and Astragalus alpinus prefer Hackelia deflexa, basocole in that region, though limestone in their American stations. Botry amphicline in Fennoscandia. chium multifidum may be acidocole in Fenno On the other hand, a real difference seems to scandia (see above p. 135). Silene acaulis, which exist as regards Carex livida, which is basocole Du RIETZ (1942a) counts among the character in Newfoundland, although amphicline or pos plants of the Swedish Dryadion, seems to grow sibly acidophilous in Fennoscandia, and Rhodo in N. E. North America chiefly on serpentine. dendron lapponicum, which is basocole in New- 172 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
foundland and Fennoscandia, although acidocole tion of Dryopteris fragrans on the continent of in N. E. North America and Labrador (FER Europe, provides an instance. The bedrock is NALD 1933 p. 10). here >>zum Teil Granate fuhrender Hornblen However, as is evident from the above lists, degneiss mit dunkleren amphibolitischen Partien>> the general trend is exactly the same in New (KALLIOLA 1937 p. 31); no lime could be estab foundland and N. E. North-America, on the lished, and when KALLIOLA states that the rock one hand, and Fennoscandia, on the other, and must be >>einigermassen kalkhaltig>>, he does so - the differences are hardly more pronounced than only because of the flora. Yet, this steep har those met with within the boundaries of Fenno bours such otherwise basophilous plants as scandia. Woodsia glabella, Carex rupestris, C. norvegica, At least some of the alpine basophytes of C. capillaris, Urtica dioeca ssp. gracilis, Draba Fennoscandia, e.g., .Asplenium vi'ride, Woodsia hirta, Saxifraga nivalis, S. groenlandica, Poten alpina, Draba hirta, Saxifraga nivalis, and S. tilla nivea, and Primula striata (ibid. p. 29). groenlandica, live on silicious primary rock in On the other hand, basophobe mosses (e.g., their isolated stations on the >>south-bluffs>> of Rhacomitrium lanuginosum, Grimmia oval is, and the North -Swedish woodland (cf. AN.DERSSON & H edwigia ciliata) are to be found there (ibid. BIRGER 1912 p. 103). Here, however, the p. 31). This should be decisive as to the chem discrepancy is probably more apparent than ical nature of the rock, since mosses are more real. The alpine plants always grow in the sensitive than vascular plants to such diffe precipice proper, where the trickling water will rences. Other basophytes occurring in the have absorbed enough nutritive salts, during North-Finnish region of non-calcareous primary its passage through the fissures of the rock, to rock are, e.g., .Asplenium viride, Carex glacialis, satisfy their rather modest needs of Ca, &c. CeTastium glabratum, Thalictrum alpinum (com Further, the forementioned phanerogams, which mon in Utsjoki), Draba incana (do.), SaxifTaga in the lowlands shed their seeds already early aizoides, .Astragalus frigidus (common in Uts in July, are among the very few >>Spring plants» joki), and Pinguicula alpina (HusTICH 1941 of these parts; and since the growing season pp. ·222 ff.). Further examples are to be found in the dry rock habitats is curtailed by the among DAHL's reports from Inner Finnmark summer drought, they are not much hampered (OVE DAHL 1934). The bedrock of the Kola by competition and can, therefore, live on the Peninsula is mainly primary rock, probably very limit of their tolerances. prevailingly granitoid gneisses (RAMSAY 1931 p. The South-Swedish, generally basophilous po 254), and sandstones in the south and east; Ump pulations of alpine plants, most of which are tek and Lujavr-Urt consist of nepheline syenite amphicline in the mountains, are, similarly, of (ibid. p. 296). Limestones are very rare (as solid little interest in this connexion, since their pro rock only on Kildin and the Rybachi Peninsula, venance probably differs from that of the alpine KIHLMAN 1890 p. 5 ); calciphilous lichens occur populations (cf. above p. 62) and, in most cases also, e.g., at Ponoj (NYLANDER 1882 pp. - 169 f.) at any rate, they are composed of other ecotypes where, however, soil samples showed a Ca con (cf. TURESSON 1927). tent of only 0,23-0,33 % (KIHLMAN 1. c.). The appearance of several arctic or arctic Nevertheless, several basophytes are common montane basophytes on acid rock within the also in regions where no basic or circumneutra,l north-eastern parts of Fennoscandia, i.e. in inner soils are to be expected, e.g., Tofieldia pusilla Finnmark and in the North-Finnish, oligo (>>eines von den gewohnlichsten Sumpfkrautern>>, trophic and severely continental, region of low ibid. p. 112), Salix myrsinites (very common, mountains of primary rock, constitutes a matter ibid. p. 108) and, to judge from the maps in of more importance. The steep on the river Kevu HULTEN 1949b, Salix hastata (map 570), Pyrola joki in the Utsjoki district, the only known sta- . grandiflora (map 1360), Pinguicula alpina (map Basophily within the Alpine Flora of Fennoscandia 173
1603), &c. It should be observed, however, ssp. sordida, Polemonium boreale, and Benecio that in the Kuusamo district (in northeastern integrifolius, are amphicline, and the far grea Finland) the relationship between plants and test number of their stations are on primary bedrock seems to be exactly the same as in the rock and hard schists poor in lime and mag Scandes. Thus, PESOLA {1934 pp. 14 ff.) nesia (cf. OVE DAHL 1934). This holds good also when investigating 55 cliffs in this district of a number of plants with a similar distri found but two otherwise basophilous plants, bution in these parts, though reaching farther viz., Woodsia alpina and Baxifraga nivalis, on south, e.g., Moehringia lateriflora, Silene tata non-calcareous rock, while even so faintly baso rica, Dianthus superbus , and Adoxa Moschatellina. cline species as, e.g., Actaea erythrocarpa, Baxi Out of 13 species with limited areas in North jraga cernua, and Baussurea alpina were found Norway and northern Lapland, reaching east only on calcareous rock. wards to · East-Finnmark (and, in some cases, Anyhow, some kind of connexion between to the Kola Peninsula and further east), Poten basophily and oceanic distribution seems un tilla Chamissonis, Armeria scabra ssp. sibirica, questionably to exist, at any rate in the Scandes. Arnica alpina, and probably Carex holostoma Within our region of Lule Lappmark the baso are basocole; Potentilla multifida and Astragalus phytes predominate in the western oceanic frigidus (which, however, occur also on primary tracts, to a degree that cannot be accounted rock, cf., e.g. , SJORS 1946a p. 85, HUSTICH op.c. for simply by a larger number of appropriate p. 225) are basocline; the rest is amphicline. habitats (cf. above pp. 156 f). The connexion On the other hand, no less than 22, from between · basophily an'd oceanic distribution is among 28 species centring on West-Finnmark, further borne out by a survey of the distribution Troms and North-Lapland, are basocole. Fur of the basophytes in the different parts of the ther, .Melandri,um angustiflorum (which, how northern Scandes. ever, also is found on primary rock, cf. S. 0. Among 11 plants, in northern Fennoscandia BJORKMAN 1947 p. 309) and Cassiope tetragona west of the Kola Peninsula restricted to conti are pronouncedly basocline. Amphicline are nental East-Finnmark and the adjacent terri only H ierochloe alpina, Triseturn subalpestre, tories, only two are basocole, viz., Arenaria ciliata Ranunculus platanifolius, and Myricaria ger ssp. pseudof'rigida and Papaver Dahlianum. manica. In all likelihood, this is. true also of Crepis mul Still more widespread is basophily among the ticaulis, that, in its only station on the river plants, 30 in all, that centre on the region Meskelva, lives on clay slate (TH. M. FRIES between West-Finnmark and Helgeland and 1857 p. 182). Thymus Berpyllum ssp. tanaensis the adjacent parts of Lapland. 27 alpine plants occupies a fairly large area in oligotrophic North belong to this group, all being basophilous, Finland and the neighbouring parts of Norway except Phippsia algida and Ca·rex macloviana and Arctic Russia {HULTEN 1949b, map 1515); which are amphicline. Among the three low at the river Tana elv it grows on alluvium land plants, the endemic Galium pumilum ssp. (cf., e.g., HuSTICH op.c. pp. 216 f.) that, in all Normani is basocole {OVE DAHL 1915 p. 137), likelihood, is non-calcareous. Dryopteris frag while Myricaria germanica and, probably, the rans, which in West-Greenland is found only endemic Thymus Serpyllum ssp. arcticus are on gneiss (PoRSILD 1920 p. 23), appears on a amphicline. similar rock in North-Finland, though occurring Finally, those alpine species whose areas exclusively on limestone and other basic rocks reach from the North Cape to southern Norway in eastern North America (cf. FERNALD 1907 consist of 9 basophytes and 3 amphicline species, J). 158, POLUNIN 1940 p. 32). The remaining viz. Carex rufina, Ranunculus glacialis , and 6 plants, viz. Arctagrostis latifolia, Veratrum Alchemilla Wichurae. Within the boundaries album, Thalictrum kem�nse, Oxytropis campestris of their oceanic areas, especially the three latter 174 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark species have manifested a considerable spreading less, which, moreover, almost exclusively takes capacity. place in winter in the form of snow; in summer On the whole. the connexion between amphi there is practically no rain at all (cf., e.g., cliny and spreading capacity seems to be· more GELTING 1934 p. 224). This affects both the marked than a greater number of appropriate chemical and the physical properties of the soil. habitats for amphicline plants will explain: Paramount to the plants are the chemical pro out of 96 doubtless amphicline alpine plants, perties, as· expressed in the pH values; this is no less than 83, i.e. 86,5 per cent, are ubiquitous proved, i.a., by the fact that many basophilous all over the Scandes , while only 22, i.e. 27,5 plants thrive excellently on calcareous primary per cent, of 80 unquestionably basocole species rock (>>urkalksten>>) that, physically, differs but fall within the ubiquitous category. Amphi slightly from certain acidic rocks (cf. EKLUND cline hibernators with little spreading power 1931, 1933). In the rainless Arctic, pH values occur only in the north-eastern district and, as high as up to 7 were established by BocHER in a few cases (e.g. Gentiana purpurea, Cam (1949) on acid primary rock (gneiss) in an extre panula barbata, Taraxacum dovrense), in South mely continental region of West-Greenland, the Norway. explanation evidently being the almost total This brief outline shows that conditions absence of leaching, owing to which the basic similar to those in western Scandinavia, with ions are not carried away but on a very small a rich flora on basic or circumneutral rocks and scale. In such places, the soil solutions are a poor one on acid rocks, prevail in those arctic obviously nutritious enough to satisfy the wants and subarctic regions (Newfoundland, north of practically all basophytes. On the other hand, eastern North-America, south -eastern Green in such circumstances Ca and other bases may, land) where we have reason to presuppose the in limestone areas, be supplied in solutions ex existence of ice-age refugia on coastal moun ceeding the tolerance of most plants. Purther, tains, while the opposite happens in regions the osmotic pressure of the soil solutions may be (the Arctic Archipelago, north-eastern Green raised to so high a degree as to obstruct the water land, and, less conspicuously, East-Pinnmark) absorption of the plants (cf., e.g., HARD AV SE where the refugia probably were of a tundra GERSTAD 1924 p. 68). type, this being the more marked, the more Nor should the physical components of what continental the climate. BRENNER (1930 p. 85) called the >>Kalkfaktoren No doubt, this state of things is partly brought komplex>> be altogether forgotten. The best ac about by present conditions. Thus, practi count known to me of the ecologically signi cally no raw humus forms in high-arctic, con ficant, physical properties of limestone is given tinental regions, while, in more oceanic re by an entomologist, LINDROTH (1949 pp. 177 ff.), gions, raw-humus forms on a large scale even who also carried out some very illuminating as far north as, for instance, on Mikis }., jord in experiments. Thermally, limestone will rather East-Greenland, in latitude 68° (BocHER 1938 favour the plants under arctic conditions, since it p. 157). Plants that do not tolerate acidic raw is characterized by high minima and small varia humus and, consequently, are confined to tions of temperature (op. c. pp. 184 ff.; cf. also basic or circmnneutral rocks in more humid KRAUS 1911); thus, cold snaps during the growing tracts, are, therefore, enabled to invade areas season will be less pernicious on lintestone of acid primary rock in high -arctic, continental than on primary rock. However, this advantage regions. However, the direct effects of the is outbalanced by the dryness of the limestone extremely low precipitation of these parts (LINDROTH op.c. pp. 187 f., 195) resulting partly unquestionably play a far greater part. The from its porosity and the larger evaporating Arctic Archipelago and North Greenland have surface of the particles as compared with those an annual precipitation of only 200 mm or of gneiss, &c., partly from its fissility and perm ea- Basophily within the Alpine Flora of Fennoscandia 175
bility to water: In the Arctic, the last-mentioned vegetation of the Alps makes an impression of property seriously hampers the plants which have being more arid than that of the Scandes (Du to manage, during the rainless summer, on the RIETZ 1924 pp. 92 ff.) probably depends on the supply of melting-water from the remaining snow. more intense insolation, the stronger evapora On sedimentary rocks, habitats that are not tion, and the absence of large, perennial snow irrigated from glaciers or perennial snowfields dry fields at low altitudes and permanently frozen up so completely during the summer that they soil which would occasion a higher level of the become almost inhabitable (cf. GELTING op. c. subsoil water table. However, these factors p. 224). Considering all relevant factors, silicious affect neither the Scandinavian alpine xero magma rocks are, in the extremely dry continen phytes nor the chionophilous plants of the Alps tal regions of the Arctic, evidently at least as which, none the less, differ in their reaction to favourable as basic or circumneutral, sedi the soil in the same way as the remaining flora. mentary ones, even to plants that are basophil Also for other reasons the distribution of ous elsewhere. basophytes and amphicline plants in the Scan A further fact should be observed. In Fenno des and, more particularly, the almost complete scandia, as well as in Newfoundland and the absence of acidocole species cannot be fully region of St Lawrence and Gaspe Co., the arctic explained on the basis of present conditions. plants live far outside their normal ranges. Many Supposing them to be decisive, there could plants tend to become nwre basophilous on the be no difference between the old inhabitants of outskirts of their areas. This holds good on the Scandes and the species arrived . there at the southern, as well as northern, boundaries a later date in their relationship to the chemical (cf. above p. 135). nature of the soil. However, such a difference doubtless exists, though not among the alpine It is a well-known fact that, on the whole, species, all of which are probably glacial sur the ecological amplitude of a species tends usu vivors, but among the lowland plants. ally to become narrower near the boundaries Those lowland plants and all-regional species, of its area; in most cases, plants are more re which may be supposed to have survived at stricted to eutrophic habitats there than nearer any rate one glaciation in the West-Norwegian their centres of dispersal, though also the opposite refugia, are as a rule basocole or pronouncedly may happen (cf. Carex magellanica in South basocline, in spite of the fact that the lowland Sweden, see above p. 136) . . It is, therefore, not populations are often amphicline or even acido astonishing that the arctic plants are confined philous. This holds good not only of many to nutricious, basic or circumneutral soils in the species that are represented in the mountains forementioned oceanic regions where intense by nwrphologically distinct races, e.g., Coelo leaching has increased the oligotrophy of soils glossurn viride ssp. islandicum, Urtica dioeca ssp. derived from acidic rocks. gracilis, Cardamine pratensis ssp. angustifolia, This would, perhaps, suffice to explain the Arabis hirsuta var. subalpestris, Geum rivale different behaviour, in the respect discussed var. subalpinum, Anthyllis Vulneraria var. lap here, of plants in regions of tundra refugia and ponica, Lotus cornimtlatus var. borealis, Rhi refugia on coastal nwuntains, and in the Arc nanthus minor ssp. groenlandicus, and the alpine tic proper and within the outposts in Fenno races of Campanula rotundifolia. The same scandia, Newfoundland, &c., but for the fact applies to a number of species that, though for that the flora on acid primary rock is much more or less satisfactory reasons they may be richer in the Alps than in the Scandes, even counted among the survivors, fail to display any though in the Alps many plants occur still nwrphological difference between the popula farther from their main areas and precipitation tions of the mountains and those of the lowland, is, on an average, higher. That, nevertheless, the viz., within our region, e.g., Sedum acre, Alche- 176 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
1nilla filicaulis, Viola rupestris, and Ajuga distribution of the different biotypes. Thus pyramidalis. Erigeron uniflorum and E. alpinum L. are amphi On the other hand, unquestionable post cline in the western and central Alps, and pro glacial immigrants do not generally manifest nouncedly acidophilous in the East-Alps, while · any differences in behaviour between the alpine E. neglectum Kern. is amphicline in the Tyrol, and the lowland populations, except in so far and basocole in the rest of the Alps (pp. 386 ff. ). as species that as, e.g., Carex vesicaria, Oxalis Apart from the basophytes on acid rock in Acetosella, Viola mirabilis and are everywhere East-Finnmark and northernmost Finland (see confined to more or less eutrophic habitats, in above p. 172), Scandinavian examples of such the mountains are almost exclusively to be a varying behaviour are, as far as we know, found in places with a basic or circumneutral very rare. The only instances known to me are soil, eutrophic habitats being practica11y non Alchemilla alpina, which is basocole in northern existent in the areas of acid primary rock. As Torne Lappmark (Trr. FRIES 1913 p. 231) and e to the >>south-bluff species>> the greater fr quency acidocline or even acidocole in Lule Lappmark of steeps on the mountain schists also plays a (see above p. 138) as well as in the Alps, part (see above p. 135).- The lowland species and Cassiope tetragona which is more or less that have been characterized earlier (p. 134) as amphicline in northern Torne Lappmark (ibid.), conceivable acidophytes are an eastern or, more at any rate at high altitudes (though seldom exceptionally, southern immigrants. And while occurring on granite and gneiss, Trr. FRIES 1917 the number of glacial survivors gradually in p. 30), while it is basocline at low altitudes (Du creases, from a syenite mountain such as Mt RIETZ 1942a pp. 132 f.) and basocole in Pite Tj:Uta, to a mountain composed of doubtless Lappmark (ARwrnssoN 1943 pp. 230, 238) and somewhat calciferous .primary rock such as south-western Lule Lappmark. Further, Saxi �it Farforita, and further to a mountain partly fraga Cotyledon appears to be amphicline in consisting of cambro-silurian rocks such as Mt south-western Norway, though basophilous in its Kassavare, the number of postglacial immigrants Laplandian stations (and acidocole in the Alps). remains about the same on all these mountains. In all these cases, the explanation may be that, Obviously, basophily and acidophily arf'. pro in those regions where the plants are baso perties that may vary from one biotype to an philous, they are approaching the boundaries other, whether the biotypes are morphologically of their areas, viz. Alchemilla alpina and Saxi . distinct or not. In this respect, the flora of the fraga Cotyledon their north-eastern limits, and Alps has been more closely investigated than Cassiope tetragona its southern one. the flora of the Scandes. Therefore, some in stances from the Alps, taken from ScHROETER However, also in Fennoscandia the position (1908), may first be considered. Not a few seems to have been best explained by TH. FRIES species apparently avoid the areas of calciferous in his hypothesis (1925 p. 6), that at any rate rocks in the western and central Alps, though part of the amphicline species consist of biotypes being basocole in the East-Alps, e.g., Sedum reacting differently to the chemical nature of Rosea (p. 538) and Carex rupestris (p. 520); the soils. Within populations whose history even such a >>Kallcanzeiger>> as Dryas octopetala differs, different biotypes may have been elimi (pp. 192 f.) seems to be basocole only in the nated, the result being that one and the same East-Alps, but amphicline-basocline in the west. species may be basophilous in one quarter and This would, no doubt, indicate a connexion acidophilous in another. between precipitation and the relationship of The proportion of basophytes to acidophytes, plants to the bedrock in the different parts within the alpine flora of l!., ennoscandia, is of the Alpine range, were it not for the fact hardly intelligible unless basophilous or amphi that there are instances also of an opposite cline biotypes were the only ones able to survive Basophily within the Alpine Flora of Fennoscandia 177 t.he glaciations in the refugia on the Norwegian basic and circumneutral soils; consequently, west-coast. most habitats appropriate to the alpine flora True, in all the areas on the Norwegian coast were situated on such soils, a�d the basophilous where an ice-age refugium can be reasonably or amphicline biotypes were favoured, as com assumed to have existed, both acid primary pared with the acidophilous ones. rocks and basic or circumneutral rocks come down More particularly, as pointed out especially to the sea. Prom this it would appear that by N ANNFELDT (1935 p. 75), the changes in the acidophytes, as well as basophytes, have had a climate must have immensely increased the ' ' chance of surviving. However, in view of the areas of the peat-bogs. The huge peat-forming conditions that probably prevailed during the carpets of Rhacomitrium lanuginosum, which initial phases of the glaciations, this is far from in present tintes cover vast areas in the coastal certain or even likely. regions and oceanic mountain tracts of western As already mentioned, our actual knowledge Norway (cf., e.g., Du RIETZ 1945b p. 75), must of the climate of the ice-age is rather slight. also have increased in a similar way. However, Among the explana-tions of the origin of the neither Sphagna nor Rhacomitria tolerate lime glaciations known to me, that presented by and other basic matters. Therefore, the peat MILANKOVITCH (1930) seems the most convin formation must have chiefly affected the regions cing. He suggests a causal connexion between of primary rock, most of which were, in all the four last glaciations and the intervening inter likelihood, wrapped up in a continuous carpet glacials, on the one side, and certain variations of Sphagnum and Rhacornitrium peat. in the inclination and eccentricity of the eclip Proper alpine plants do not tolerate peaty tic and the precession of the equinoctial points, .soils. Such acidophilous biotypes among the on the other; these variations apparently coin alpine plants as n1ay have existed in preglacial cided in time with oscillations of the ice-edge or interglacial tin1es could, therefore, hardly in the Alps. When these three factors co-operate find appropriate new habitats in the West 1n a. certain direction, the effect in high latitudes N orwegian refugia during the earlier phases will be periods of increased precipitation, reduced of the glaciations, when the ice had deprived difference between the seasons, and cool and them of part of their ancient stations. The wet summers, in other words, a deterioration crests of primary rock protruding from the in the climate pronounced enough to cause a n1arsh-logged, peat-covered ground were not glaciation. likely to support more than a scanty flora, MILANKOVITCH's theory implies that, at the poor in species, not unlike that which at present beginning of the glaciations, Scandinavia had submits to similar inhospitable habitats in outer a chilly, extremely rainy, oceanic climate with Lofoten. At any rate during the initial sta.ges rather n1ild winters. Apparently, these initial of the glaciations, conditions in the refugia on phases when local glaciation was intense, though the West-Norwegian coastal mountains, all no continuous ice-sheet covered the land, were situated a.t low altitudes, were probably not a of a long duration, seeing that the Scandes little suggestive of the present state on the probably obtained their present surface sculp islands of Helgeland, where the alpine flora ture essentially during this phase of the latest centres upon the limestone hills and is almost glaciation, and not in postglacial times (MAN completely absent from the areas of primary NERFELT 1945). In these circumstances, the rock with their peat-bogs and heatherclad soils of the coastal region rnust have been moors (cf. OvE DAHL 1912, 1915). subjected to an intense leaching, podsolization Above all, however, conditions in the West and raw-humus formation, i.e. to processes Norwegian refugia �re likely to have resembled adverse to practically all alpine plants. Purther, the present ones in Newfoundland. Thanks to these processes must have been less severe in PERNALD's graphic descriptions (1925, 1926,
13- 496149 Sten Selander I 178 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
1933, &c.), we are able to form a mental picture (CoLEMAN 1926). This group comprises the fol of the vegetation on different kinds of rock in lowing species (FERN.ALD 1926 p. 50, pp. 52 f., those parts. The table-lands, especially when p. 161, pp. 235 f., 1933 p. 273): consisting of silicious rock (as, e.g., on Lookout Potamogeton polygonifoli- Ranunculus hederaceus Mt and in the Blomidon and Topsail regions), us R. Flammula are >>a prairie-like expanse of peat, with innu Agrostis canina Potentilla sterilis merable shallow depressions and small pools>> Sieglingia decumbens P. erecta Glyceria fluitans P. anglica (FERNALD 1933 p. 85). In western Newfound Festuca ovina ssp. capil- Oalluna vulgaris (v. ame- land, all the >>b ogs, wet thickets, brooksides and lata ricana Seem.) meadows are characterized by southern types>>, Nardus stricta Pedicularis palustris while >>the arctic types prevail on dry rocks or Oarex leporina P. silvatica on windswept crests>> (FERNALD 1925 p. 332). Juncus conglomeratus Galium hercynicum J. bulbosus Thus, the habitats of the arctic plants are to be found in such places as have beyond doubt Part of these species, in the first place the always been free from peat, particularly on the graminids, may, of course, have been introduced. precipices and steep screes just above sea level; In the case of, e.g., the hydrophytes, the Pedi :FERN.ALD (1933 p. 125) emphasizes >>the deep cularis species, and the endemic race of Calluna, mantle of weathered debris>> and >> the aridity and this seems, however, hardly possible. FERNALD the calcareous nature of the ex posed ledges>> as evidently considers that they are native, except espe.cially characteristic of the stations of >>the Potentilla sterilis, about whose station and ' truly arctic species and their endemic relatives>>. habitat in Newfoundland nothing is known As already mentioned (above I). 170), the great (cf. FERNALD 1933 p. 273). Apart from Festuca majority of these plants are confined to the ovina ssp. capillata, all the remaining plants are areas of calcareous or magnesian rocks. The characteristic of the oligotrophic regions of flora on quartzite, &c., is about as poor as in western, oceanic Europe, where they chiefly Scandinavia. However, probably a few acidocole occur either in wet habitats of some kind or species (and biotypes) managed to survive, other or on leached, more or less damp, often owing to the fact that the unglaciated areas somewhat peaty ground (in Swedish called of Newfoundland were far larger and more fukthedar). Since at any rate the majority of varied than the small Norwegian refugia and, the species must be regarded as glacial survivors consequently, habitats appropriate to acido in Newfoundland, their presence there confirms phytes escaped, in some places, from being co the probability of a course of events on the lines vered by peat, viz. either dry, windswept crests suggested above. or ravines and steep brook furrows of the kind When the local glaciers in Scandinavia united where nowadays the few rare, acidophilous into one continuous ice-sheet, the alpine plants epibiotics seem mainly to occur (cf., e.g. , FER were, as a matter of course, deprived of their N.ALD 1933 p. 116). last stations in the mountains. They became In this connexion, a particular interest at restricted to new habitats in the refugia, the taches to a number of European lowland plants nature of which was suggested above. The that occupy completely isolated areas (in Sphagnum bogs may have decreased, as the several cases the only ones in America) on highly fully developed ice-cap lowered the temperature, silicious rock in the Avalon Peninsula of south though this is far from certain. To judge from eastern Newfoundland, which was largely free the character of several plants that ai·e likely from ice during the last (Wisconsin) glaciation, to have survived the last glaciation, for instance, although covered, like all Qf Newfoundland ex in Newfoundland, especially those belonging cept the southern part of Long Range, by an ice to the thermophilous flora of the Atlantic cap during an earlier phase of the Pleistocene coastal plain of America, but also certain Euro- Basophily within the Alpine Flora of Fennoscandia 179 pean types, e.g., Ranunculus hederaceus, Carda point of view applied here, this difference mine flexuosa, and Potentilla anglica, the cli between vascular plants and others is easily mate can hardly have been much colder during explained: boulders, &c., of primary rock, where this glaciation than it is at present; and there basophobe lichens and mosses could find refuges, is no ground for assuming that conditions in would have protruded also from the thickest of Norway differed in that respect. Even a lower peat carpets, while niches, of a size sufficient ing of the mean temperature of about 4 o C, to harbour such basofuge species of moss and as suggested above (p. 146), would scarcely have lichens as live on earth, would always have affected the peat bogs to any critical degree. been available in rock-crevices, &c. But be that as it may: provided the above argu Apart from species surviving in the tundra mentation holds good, the harm to the alpine refugium of East-Finnmark, to which this ar acidophytes had already been done, even though gumentation does not apply, the only likely, or there were no peat-bogs left during the maximum conceivable, acidophytes in the alpine flora of of the glaciation. Therefore, only biotypes Lapl::md are the species enurnerated above (pp. tolerant of calcareous or magnesian rocks can 138 f.). .Among them, Carex rotundata is one be supposed to have overcome all trials, espe of the two (or three) Fennoscandian Carices that cially if our alpine flora has survived more glacia are actually to be found on typical Sphagnum tions than one and, perhaps, all the ice-age in bogs (>>mossan>); besides, it may be a postglacial Fennoscandia. In the latter instance, the immigrant. The habitats of Ctyptogramma events outlined above would, presumably, have crispa, i.e. steep boulder-lands, could evidently been repeated at the beginning of each glaciation, not be affected by the processes suggested and the elimination of basofuge biotypes almost above. Pedicularis lapponica, H ieracium alpi total. In other words: the proportion of baso num, and H. nigrescens are not favoured by a philous and acidophilous biotypes ·within the mineral soil. They rather thrive just as well, or West-Sca ndinavi an alpine flora would, then, perhaps better, on such raw-humus soils as have become exactly such as they actually are. would be common under the forementioned It goes without saying that rare plants with conditions. This is true also of Deschampsia small populations comprising only a limited atropurpurea. The remaining species are such number of biotypes would, in the first place, extremely hardy and unpretentious ones as be affected in this way. Common species with would hold their own on the barren crests of a large ecologic amplitude would be better primary rock in any marsh-logged and peat qualified to keep large and vigorous popula covered refugium; nowadays, for instance, all tions; this may be reflected in the considerable of them appear among the scanty flora of the postglacial spreading capacity of the amp hi Lofoten islands. cline species (cf. above p. 174). Thus, the com },urthermore, it shoul9- be emphasized that mon plants would have becon1e still more such a course of events as that outlined above common, the rare ones still rarer. could hardly have taken place, within the The hypothesis propounded here appears to be Atlantic region, outside West-Norway, Scotland, borne out also by the wellknown fact that, the Faroes, Newfoundland and the region of St within the alpine flora of Fennoscandia, bryo Lawrence and, possibly, south-eastern Green phytes and thallophytes, as distinguished from land. However, in Scotland and the Faroes vascular plants, are not characterized by the vegetation is so strongly affected by culture, predominance of basophilous and amphicline particularly by intense sheep pasturing, that types. On the contrary, a very large number it is impossible to form an idea of the actual of alpine mosses and, especially, of alpine position there from literature alone. In the lichens are confined to silicate rocks; to quote .Alps, the alpine flora was not, as in Fenno instances should be superfluous. From the scandia, closed in between ice and sea. Espe- 180 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark cially on the southern side of the Alpine range, took them in the other regions where the refu where the J\iediterranean climate should have gia were of a coastal type. On the other hand, changed, during the glaciations, in a way favou an arctic, or arctic-montane, flora survived rable to alpine plants, there must have been also on acid rocks in the tundra refugia, where many places where alpine acidophytes could the continental climate precluded such processes find refuges. as those described above and, consequently, The hypothesis propounded here is admittedly also an elimination of tlie acidocole types among highly conjectural. However, I have not been the plants restricted to mineral soil. This posi able to find out any data that conflict with the tion has, later, been maintained because of above-stated suppositions. They can be summa the climatic conditions during the postglacial rized as follows: time, which have had a similar effect. Also the Each glaciation was ushered in by a very long fact that most plants become more basophilous period of a wet and chilly, extremely oceanic, near the boundaries of their areas has probably climate. This caused an intense leaching, played a part. Accordingly, in the regions of podsolization and raw-humus formation and former coastal refugia (western Scandinavia, a vast increase of Sphagnum bogs and Rhaco Newfoundland, the region of St Lawrence, rnittium carpets within all areas of primary possibly south-eastern Greenland) the epibio acid rocks on the Norwegian west-coast. Con tics consist exclusively, or almost so, of baso sequently, such acidocole arctic and arctic philous and amphicline types, while in the montane species and biotypes as were more or regions of former tundra refugia (the Arctic less confined to mineral so!l were unable to find Archipelago of America,. · North-Greenland, East appropriate habitats in the refugia on the Finnmark and, probably, some inland nunatakks ·Coastal mountains of western Norway and, in South-Norway) the epibiotics consist also of therefore, became extinct. A similar fate over- a.cidophytes.
ADDENDA ET CORRIGENDA
Investigations by LovE (1950), published when this Vitis-idrea ssp. minor (Lodd.) Hult.) (ibid. p. 47). V. book had been set up, prove that Sesleria varia (Jacq. ) Vitis-idrea sens. str. ( = ssp. maior Love) is restricted Wettst. ssp. islandica Love (op.c. pp. 27 ff.) and Silene to Eurasia, including the Faroes (ibid.). Consequently, maritima With. ssp. islandir:a Love et Love (ibid. p. 40) it should be· excluded from the list of Eurasiatic plants should be added to the Icelandic endemics enumerated reaching Iceland (above p. 90). above (p. 75). Also the American species in Iceland Further, Polypodium vulgare should be included in have proved to be more numerous than previously the list (p. 10) of the plants reported by LINNAEUS assumed; in addition to those enumerated above (p. from south-western Lule Lappmark.
74) come Elymus mollis Trin. (LovE , op.c. pp. 31 ff.), The pictures in Fig. 31 and 32 on pp. 154-155 are Salix cordijolia Pursh (ibid. pp. 37 f.), and Vaccinium photographed by Borje Mineur July 1947. LITERAT URE CI TED
AARIO, L., 1940: Waldgrenzen und subrezente Pollen skandinaviska arter. - Norrlandskt Hand spektren in Petsamo Lappland. -Ann. Acad. bib!. V. Scient. Fenn. A 54: 8. ANDERSSON, G., & HESSELMAN, H., 1901: Bidrag till 1943: Uber die Wald- und Klimaentwicklung kannedomen om Spetsbergens och Beeren an der lapplandischen Eismeerkiiste in Petsamo. Islands Karlvaxtflora. - Bih. K. Sv. Vet. - Ann. Bot. Soc. Zool. Bot. Fenn. V anamo Akad. Handl. 26, 3: 1. 19: I. ANDREEV, V., 1931: Materiaux pour la flare de la ABBE, E. C., 1936: Botanical Results of the Grenfell presqu'ile de Kanin. - Trav. du mus. Bot. de Forbes Northern Labrador Expedition 1931. l'Acad. d. Sci. de l'URSS. 1931. - Rhodora 38. ANGSTROM, A., 1946: Sveriges klimat. - Stockholm. ABERG, B., 1949: Om karlvaxternas hoj dgranser i ANGSTROM, J., 1839: Antekningar.-Bot. Not. 1839. Sarek. - Sv. Bot. Tidskr. 43. ARNELL, H. W., & JENSEN, C., 1907, 1910: Die Moose AHLMANN, H. W:soN, 1948: Den nutida klimatfluk des Sarekgebietes. - Naturw. Urtters. d. tuationen och dess utforskande. - N orsk Sarekgeb. in ·schwed. Lappl. Ill: 2, 3. Geogr. Tidskr. 11 (1947). ARWIDSSON, Th., 1926 a: Floristiska notiser fran AHLMANN, H. W:SON, GARLAND, S.-0., MANNER Pite lappmark, huvudsakligen nordvastligaste FELDT, C. M:soN, BECKMAN, B., 1949: Sverige delen. - Bot. Not. 1926. nu. AVC:s atlas over Sveriges folk, land och 1926 b: Sjoarnas vegetation i Lilla Lule alvs naringar. - Stockholm. vattenomrade. - Arkiv. f. bot. 20 A: 14. AHLNER, S., 1948: Utbredningstyper bland nordiska 1927: Vaxtgeografiska notiser fran Norrland I. barrtradslavar. - Acta Phytogeogr. Suec. 22. ----'- Bot. Not. 1927. ALLEE, W. C., & PARK, T., 1939: Concerning ecolog 1937: Om Hieracium Kjellandii Folin n. sp. - ical principles. - Science 89. Ibid. 1937. ALM, C. G., 1944: Om Carex macloviana d'Urv. och 1943: Studien uber die Gefasspflanzen in den dess utbredning i Nordeuropa. - The Svedberg Hochgebirgen der Pite Lappmark. - Acta 1884 30.8. 1944. Uppsala och Stockholm. Phytogeogr. Suec. XVIII. ALMQUIST, E., 1929: Upplands vegetation och flora. AsPLUND, E., 1918: Beitrage zur Kenntniss der - Acta Phytogeogr. Suec. I. Flora des Eisfjordsgebietes. - Arkiv f. bot. --, 1948: Nagra markliga Dalavaxter. - Sv. Bot. 15: 14. Tidskr. 42. ANDERSON, N. J., 1844-45: Plantae vasculares circa BEGER, H., 1930: Praktische Richtlinien der struk Quickjock Lapponiae Lulensis. - Diss. Upsa turellen Assoziationsforschung im Sinne der von liae. der Zurich-Montpellier-Schule geiibten Methode. 1846: Anteckningar under en resa i Umea, Pitea - Abderhalden, Handb. d. biol. Arbeitsmeth. och Lulea lappmarker 1845. -Bot. Not. 1846. XI: 5. 1866: Nya bidrag till Quickjocktraktens flora. BERGSTROM, E., 1911: En anteckning om fjallvaxter - Ibid. 1866. · i Torne Lappmarks barrskogsregion. - Sv. ANDERSSON, G., 1896: Svenska vaxtvarldens historia Bot. Tidskr. 4. i korthet framstalld. 2: a uppl. - Stockholm. BESKOW, G., 1930: Erdfliessen und Str:ukturboden --, 1902: Hasseln i Sverige fordom och nu. - Sv. der Hochgebirge im Licht der Frosthebung. Geol. Unders. C a: 3. Vorlaufige Mitteilung. - Geol. Foren. Forh. 52. ANDERSSON, G., & BIRGER, S., 1912: Den norrlandska BILLBERG, C. J.: See Svensk Botanik. florans geografiska fordelning och invandrings BIRGER, S., 1909: Vaxtlokaler fran Norrland och historia med sarskild hansyn till dess syd- Dalarna. - Sv. Bot. Tidskr. 3. 182 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark
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--, 1935: Glacial Survival of Plants in Scandinavia fasen i den svenska florans invandringshistoria. and the British Isles. - Proceed. Royal Soc. - Sv. Bot. Tidskr. 40. B 118: 808. 1949: Palynological Aspects on the Pioneer 1940 a: Problems of Bipolar Plant Distribution. Phase in the Immigration of the Swedish Flora. - Acta Phytogeogr. Suec. XIII. II. Identification of Pollen Grains in Late 1940 b: Das limnologisch-thalassologische Vege Glacial Samples from Mt Omberg, Ostrogothia. tationsstufensystem. - V erh. Intern. Verein. -Ibid. 43. f. theor. u. angew. Limnologie IX. ERIKSSON, B., 1912: En submoran fossilforande aflag 1942 a: Rishedsforband i Tornetraskomradets ring vid Bollnas i Halsingland. - Geol. Foren. lagfjallbalte. - Sv. Bot. Tidskr. 36. Forh. 34. 1942 b: De svenska fjall ens vaxtvarld. - N orr ERLANDSSON, S., 1942 a: Fjallvaxter i sodra Sverige. land, natur, befolkning och naringar. Stock - Bygd o. Natur, Arsb. 1942. holm. 1942 b: Cytologiskt-vaxtgeografiska rasstudier i 1942 c: Vaxtsamhallslarans grunder. - Sten Nordens Parnassia palustris-population. - Acta cilerat kompendium. Horti Berg. 13: 4. 1943: Tornetraskomradets hoj dgranser for Salix FAEGRI, K., 1933: Uber die Langenvariation einiger herbacea och Salix polaris. - Bot. Not. 1931. Gletscher des Jostedalsbre und die dadurch 1945 a: Om terminologien for forna och organo bedingten Pflanzensukzessionen. - Bergens gen jord samt om circumneutral hedtorv och Mus. Arb. 1933, Naturv. rekke 7. angstorv (>>Alpenhumus>>) i de svenska fjallen. FERDINANDSEN, C., 1918: Unders0gelser over danske - Geol. Foren. Forh. 67. Ukrudsformationer paa Mineraljorder. - Diss.
-- , 1945 b: Bryophyta. - NANNFELDT, J. A., och K0benhavn. Du RIETZ, G. E., Vilda vaxter i Norden. Mos FERNALD, L. M., 1907: The soil preferences of certain sor. Lavar. Svampar. Alger. Stockholm. alpine and subalpine plants. - Rhodora 9. 184 SELANDER: Floristic Phytogeography ,of South-Western Lule Lappmark
FERNALD, L. M., 1924: Isolation and Endemism in FRIES, TH. M., 1857: Nya Skandinaviska vaxter. Northeastern America. - Am. J ourn. Bot. 11. Intybus multicaulis (Led.). - Bot. Not. 1857. 1925: Persistence of Plants in Unglaciated --, 1913: See LINNAEUS 1913. Areas of Boreal America. - M em. Am er. A cad. FRonrn, J., 1915: Nagra markliga sydberg i Lule lapp Arts & Sci. XV: Ill. mark. Tva nya lokaler for Potentilla multijida. 1926: Two Summers of Botanizing in New - Sv. Bot . .Tidskr. 9. foundland. - Rhodora 28. 1919: Om forhallandet mellan berggrundens 1928: Eastern American Occurrence of Athy kalkhalt och de nordsvenska vaxternas utbred rium alpestre. - Ibid. 30. ning. - Bot. Not. 1919. -- , 1929: Some Relationships of the Floras of the GAMS, H., 1933: Das Alter des alpinen Endemismus. Northern Hemisphere. - Proc. Intern. Congr. - Ber. Schweizer Bot. Ges. 42. Plant Sci. Ithaca, N. Y., 1926 2. GARTNER, H., 1939: Zur systematischen Anordnung 1931: Specific Segregations and Identities in einiger Arten der Gattung Cerastium L. some Floras of Eastern North America and Feddes · Repertorium, Beih. CXIII 113. the Old World. - Rhodora 33. GAVELIN, A., 1909: Om tradgransens nedgang i de 1933: Recent Discoveries in the Newfoundland svenska fjalltrakterna. - Sv. SkogsvardsfOren. Flora. - Ibid. 35. Tidskr. 7. FIRBAS, F., 1939: Vegetationsentwicklung und Kli 1910: TradgransfOrskjutningarna inom Kama mawandel in · der Mitteleuropaischen Spat jokks vattenomrade (Lilla Lulealf). - Sv. und N acheiszeit. - Die N aturwissenschaften Geol. Unders. C: 227. 27: 6. --, 1915: Om den geologiska byggnaden inom FLODERUS, B., 1930: Sibiriska inslag i Fenno13kandias Ruotevare omradet. - Geol. Foren. Forh'. 37. Salix-flora. - Bot. Not. 1930. GELERT, 0., & OsTENFELD, C. H., 1902: Flora arctica. --, 1931: Salicaceae. - Skand. Flora, utg. av I. Pteridophyta, Gymnospermae and Mono 0. R. Holmberg. II: l. Stockholm. cotyledones. - Copenhagen.
- Leningrad. of East Greenland between Franz J oseph Fj ord FoLIN, T., 1928: Hieracia vulgata efter Ume alvs 6vre and Dove Bay (Lat. 73°15'-76°20' N.). - lopp. - Arkiv f. Bot. 22 A: 5. Medd. om Gmnl. 101: 2. 1931: Hieracia vulgata efter Skellefte alvs GJAEREVOLL, 0., 1949: Sn0lejevegetationen i Oviks 6vre lopp. - Ibid. 24 A: 1. fjellene. -- Acta Phytogeogr. Suec. 25. 193'4: Nya Hieracia vulgata fran Tarna socken. GLEASON, H. A., 1924: Age and Area from the view - Ibid. 26 A: 6. point of phytogeography. - Amer. Journ. 1942: Kvikkjokkstraktens Hieracia vulgata. Bot. 11. - Ibid. 30 A: 5. 1927: Further views on the succession concept. FRIES, H., 1945: Goteborgs och Bohus lans faneroga -Ecology 8. mer och ormbunkar. Forteckning med fynd GRANLUND , E., 1932: De svenska hogmossarnas geo orter. - G6teborg. logi, deras bildningsbetingelser, utvecklingshis }"' RIES, M., 1949: Den nordiska utbredningen av Lac toria och utbredning jamte sambandet mellan tuca alpina, Aconitum septentrionale, Ranun h6gmossebildning och f6rsumpning. - Sv. Geol. culus platanifolius och Polygonatum verticilla Unders. C: 373.
tum. - Acta Phytogeogr. Suec. 24. GR0NLIE, · 0. T., 1927: The Folden Fiord. Contribu FRIES, TH. C. E., 1911: Avlagringarna vid Arpojaure. tions to the quaternary geology of the Folia - Geol. Foren. Forh. 33. district. - Troms0 Mus. Skr. I: II. --, 1913: Botanische Untersuchungen im nord- GR0NLUND , CHR., 1884: Karakteristik af Plante lichsten Schweden. - Vetensk. o. prakt. unders. vaeksten paa Island, sammenlignet med Flo i. Lappl. Fl. o. Fauna 2. raen i flere andre Lande. - Naturv. Foren. 1917: Uber die regionale Gliederung der alpinen Festskr. K0benhavn . Vegetation der fennoskandischen Hochgebirge. GRONTVED, J., 1942: The Pteridophyta and Sperma -- Ibid. 4. tophyta of Iceland. -- Botany of Iceland IV: I. 1921: Die skandinavischen Formen der Euphm GuSTAFSSON, A., 1943: The Genesis of the European
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HADAC, E., 1948: On the History of the Flora of HEINTZE, A., 1914: Iakttagelser over kionokor fro Iceland. - Studia Bot. Cechosl. 9: l. spridning. - Bot. Not. 1914. HAGLUND, G., 1937: Taraxacum. In HULTEN 1937 b. HEMMENDORFF, E., 1897: Om Glands vegetation. - 1946: Contributions to the Knowledge of the Diss. Uppsala. Taraxacum Flora of Alaska and Yukon. - HERZOG, T., 1933: Pflanzengeographie. - Handb. d. Sv. Bot. Tidskr. 40. Geogr. Wissensch. Potsdam. 1948: Further Contributions to the Knowledge HnTONEN, I., 1946: Die Karelische Landenge als. of the Taraxacum Flora of Alaska and Yukon . Einwanderungsweg der Pflanzenarten im Lichte - Ibid. 42. ihrer heutigen Verbreitung. - Ann. Bot. 1949: Supplementary Notes on the Taraxacum Soc. Zool. Bot. Fenn. Vanamo 22. Flora of Alaska and Yukon. - Ibid. 43. HJELT, HJ., 1888-1926: Conspectus Florae Fennicae HALDEN, B., 1915: Det interglaciala Bollnasfyndets I-VII. - Acta Soc. Fauna Fl. Fenn. V,. stratigrafi. - Geol. Foren. For h. 37. XXI: I, 30: I, 35: I, 41: I, 51: I, 54. --, 1948: See SuNDIUS & SANDEGREN. H6GBOM, A., 1930: Om Tarrekaisses magnesit- och HALLE, T. G., 1915: Nagra jamtlandska kalktuffer jarnmalmsforekomster. - Geol. Foren. Forh. och deras flora. - Sv. Geol. Unders. C: 260. 52. HAMBERG, A., 1900: Sarjektraktens qvartara historia. H6GBOM, A. G., 1906: Norrland. Naturbeskrifning. - Geol. Foren. Forh. 22. - N orrlandskt Handbibl. 1. 1901 : Geologiska och fysiskt -geografiska under HOLM, V. F., 1875: En resa i Lappland och Norge. sokningar i Sarekfjallen. - Ymer 21. - Bot. Not. 1875. 1910: Gesteine und Tektonik des Sarekgebirges HoLMBERG, 0. R., 1922: Hartmans handbok i Skan-
etc. - Geol. · Foren. Forh. 32. dinaviens flora. Haft. 1. -- Stockholm. HAMBERG, H. E., 1909: Molnighet och solsken pa den --, 1926: Handbok i Skandinaviens flora. Haft. 2. skandinaviska halvon. - Bih. Meteor. Iaktt. - Stockholm. i. Sverige 50. HoLMBOE, J., 1936: Uber Nigritella nigra (L.) Rich.,. HANNERZ, E., 1945: Fallforsberget, ett sydberg i ihre V erbreitung und Geschichte in Skandina Norrbotten. - Sveriges Natur 36. vien. - Ber. d. Schweiz. Bot. Ges. 46. HANSEN, A. M., 1904: Hvorledes har Norge faat sit 1937: The Trondheim District as a Centre of plantedekke? - Naturen 1904. Late Glacial and Postglacial Plant Migrations .. --, 1930: Bre og biota. - Skr. utg. av D. Norske - Avh. utg. av D. Norske Vid.-Akad. I. 1936 Vid. Akad. I Mat. Naturv. Klasse 1929: 3. No. 9. HANSEN, 0., & LID, J., 1932: Flowering Plants of HOLMQUIST, P. J., 1900: En geologisk profil ofver Franz J oseph Land. Collected on the Nor fjellomradena emellan Kvikkjokk och norska wegian Scientific Expedition 1930. - Skr. om kusten. - Geol. Foren. Forh. 22. Svalbard og Ishavet 39. HuLTEN, E., 1927-30: Flora of Kamtchatka and the· HARD AV SEGERSTAD, F., 1924: Sydsvenska florans Adjacent Islands I-VI. - 1927: I. K. Sv. vaxtgeografiska huvudgrupper. (Deutsche Zu Vet.-Akad. Handl. 3: e ser. 5: l. 1928: II.. sammenfassung). - Malmo. Ibid. 5: 2. 1929: Ill. Ibid. 8: l. 1930: IV. Ibid. HARTMAN, C. J., 1841: Olof Rudbecks Lappska ve4ter. 8: 2. - Bot. Not. 1841. 1937 a: Outline of the History of Arctic and HEDLUND, T., 1948: Om Ribes vulgare och Ribes rub Boreal Biota during the Quaternary Period. - rum. - Bot. Not. 1948. Stockholm. HEGI, G., 1906-31: Illustrierte Flora von Mittel 1937 b: Flora of the Aleutian Islands and europa I-VII. - Munchen. westernmost Alaska Peninsula with notes on HEINTZE, A., 1907: Om Chrysosplenium alternifolium the flora of Commander Islands. - Stockholm. L. v. tetrandrum Lund och dess utbredning i 194 1-50: Flora of Alaska and Yukon I-IX . Skandinavien. - Bot. Not. 1907. - 1941: I. Lunds Uni\r. Arsskr. N. F. Avd. 2. 1908: Vaxttopografiska anteckri.ingar fran ett 37: I. 1942: II. Ibid. 38: I. 1943: Ill. Ibid. par farder genom Skibottendalen i Tromso 39: I. 1944: IV. Ibid. 40: l. 1945 a: V. Ibid. amt. - Arkiv f. Bot. 7: 11. 41: l. 1946: IV. Ibid. 42: I. 1947 a: VII. Ibid .. 1909: Om Rammculus lapponicus och andra 43: I. 1948: VIII. Ibid. 44: l. 1949 c: IX. granens foljevaxter i Skandinavien. - Bot. Ibid. 45: l. Not. 1909. 1943 a: Stellaria longipes Goldie and its allies .. 1913: Vaxttopografiska undersokningar i Asele - Bot. Not. 1943. Lappmarks fjalltrakter. II. - Arkiv f. Bot. 1945 b: Studies in the Potentilla nivea Group .. 13: 5. - Bot. Not. 1945. 186 SEL.ANDER: Floristic Phytogeography of South-Western Lule Lappmark
HuLTEN, E., 1947 b: Ranunculus sceleratus subsp. Studien an der Oarex-Subsektion Alpinae Kale la. reptabundus inom det Fennoskandiska florom - Ann. Bot. Soc. Zool. Bot. Fenn. Vanamo radet. - Bot. Not. 194 7. 19: 3. 1949 a: On the Races in the Scandinavian KALLIOLA, R., 1937: Dryopteris fragrans (L.) Schott, Flora. - Sv. Bot. Tidskr. 43. ein fiir Europa neuer Farn. - Ibid. 9: 4. 1949 b: Atlas over vaxternas utbredning i KIHLMAN, A. 0., 1890: Pflanzenbiologische Studien N orden. Fanerogamer och ormbunksvaxter. aus Russisch Lappland. Ein Beitrag zur - Stockholm. (Seen in the proof-stage, only Kenntnis der regionalen Gliederung an der po partly paged and dated 1949; the year of im laren Waldgrenze. - Acta Soc. Fauna Fl. pression of the published volume will probably Fenn. VI: 3. be 1950.) KoRSHINSKY, S., 1898: Tentamen Florae Rossiae ori HusTICH, I., 1941: Nagra vaxtgeografiska anteck entalis. - Mem. Acad. Imp. Sci. St.-Peters ningar under en juliresa 1940 i Utsjoki, norra bourg VIII Ser. VII: 1. Lapp land. - Mem. Soc. Fauna Fl. Fenn. 17. KoTILAINEN, M., 1942: Alpiinis-arktisen kasvis HYLANDER, N., 1941: Forteckn{ng over Skandina tonaineksen vaellusteista ja sekoittumisesta viens vaxter. I. Karlvaxter. - Lund. kvartaarikautena. - Luonnon Ystava 46. 1945: omenldatorische und systematische N KRAUS, G., 1911: Boden und Klima auf kleinstem Studien iiber nordische Gefasspflanzen. - Raum. V ersuch einer exakten Behandlung des Upps. Univ. Arsskr. 1945: 7. Standorts auf dem Wellenkalk. - J ena. HYYPPA, E., 1933: Das Klima und die Walder der KuKENTHAL, C., 1909: Cyperaceae. - Caricoideae. spatglazialen Zeit im Bereich der Karelischen - Pflanzenreich IV: 20. Landenge. - Acta Forest. Fenn. 39. KuLCZYNSKI, S., 1924: Das boreale und arktisch 1936: Uber die spatquartare Entwicklung alpine Eiement in der mittel-europaischen N ordfinnlands mit Erganzungen zur Kennt Flora. - Bull. Acad. Polon. Se. Lettr. Ser. B, nis des spatglazialen Klimas. - Compt. Rend. 1923. Soc. Geol. Finl. IX. KULLING, 0., 1945: Om fynd av mammut vid Pil lVERSEN, J., 1936: Sekundares Pollen als Fehler grimstad i Jamtland. Med en inledning av quelle. - Danm. Geol. Unders. IV: 2. PER GEIJER. - Sv. Geol. Unders. Arsb. 39: 6. 1941: Landnam i Danmarks Stenalder. En pol LAESTADIUS, L. L., 1822: Botaniska Anmarkningar, lenanalytisk Unders0gelse over det f0rste Land gjorda i Lappmarken och tillgransande Lands brugs Indvirking paa Vegetationsudviklingen. orter. - K. Sv. Vet.-Akad. Handl. 1822. - Ibid. II: 66. 1824: Beskrifning ofver nagra sallsyntare Vax 1944: Viscum, Hedera and Ilex as Climate ter fran norra delen af Sverige jemte Anmark Indicators. A Contribution to the Study of the ningar i Vaxtgeografien. Ibid. 1824. Post-Glacial Temperature Climate. - Geol. 1839: Loca parallela plantarum. - Nova Acta Foren. Forh. 66. Soc. Scient. Upsal. 11. 1945: Helianthemum som fossil Glacialplante i Danmark. - Ibid. 67. LAGERBERG, T., 1937, 1939: Vilda vaxter i Norden. 1947: Plantevaekst, Dyreliv og Klima i det Band I, 1937. Band Ill, 1939. -Stockholm. senglaciale Danmark. - Ibid. 69. 1940: Svenska fjallblommor. Andra, tillokade JACCARD, P., 1930: Die statistisch-floristische Methode uppl. - STF:s handb. om det svenska fjallet l. als Grundlage der Pflanzensoziologie. - Abder Stockholm. halden, Handb. bioi. Arbeitsmeth. 11: 5. LAGERBERG, T., & HoLMBOE, J., 1940: Vare ville JENNY, H.: See BRAUN-BLANQUET, J. planter. Bd VI. - Oslo. .JENSEN, C., Skandinaviens bladmossflora.- K0ben LAGERGREN, H., 1922: Lulea Silververk. - Norr havn. bottens lans hembygdsfor. Tidskr. 1922. . JoHANSSON, K., 1926: Vaxtgeografiska sporsmal LANGE, J., 1880: Conspectus Florae Groenlandicae . rorande den svenska Hieracium-floran. II. - Pars prima. - Medd. om Gr0nl 3. Sv. Bot. Tidskr. 20. LANGE, TH., 1938: Jamtlands karlvaxtflora. - Acta .JuLIN, E., 1948: Vessers udde. Mark och vegetation Bot. Fenn. 21. i en igenvaxande lovang vid Bjarka-Saby. - LID, J., & ZACHAU, A. R., 1928: Utbredningen av Acta Phytogeogr. Suec. 23. Viscaria alpina (L.) G. Don, Alchemilla alpina KALELA, A., 1943: Die Ostgrenze Fennoskandiens in L. och Rhodiola rosea L. i Skandinavien. pflanzengeographischer Beziehung. - Veroff. Medd. Goteb. bot. tradg. IV. Geobot. Inst. Riibel 20. LINDBERG, H., 1911: Floran och vegetationen. 2. 1944: Systematische und pflanzengeographische Karlvaxter. 3. Subfossila viixtfynd belysande Literature Cited 187
florans utveckling. - Text till Kartblad N :o Norrbottens Natur. Norrb. Bioi. Foren.' 20 i Atlas over Finland. Helsingfors. Smaskr. 1 1949. Boden. LINDQUIST, B., 1931: Den skandinaviska bokskogens LovE, A., 1941: Rumex tenuifolius nova species. - biologi. - Sv. Skogsvardsforen. Tidskr. 29. Bot. Not. 1941. 1938: Dalby Soderskog. En skansk lovskog i 1944: The dioecious forms of Rumex subgenus forntid och nutid. - Acta Phytogeogr. Suec. X. Acetosa in Scandinavia. - Ibid. 1944. 1940: Juncus alpinus Vill. var Marshallii (Pugs!.) 1945: Islenzkar jurtir. - Lund. Lindq. in Scotland and the Race Differentia 1947: Heimskautusveifgras (Poa arctica R. Br.) tion in the Species J. alpinus V ill. - Acta fundi5 a Hornstrondum. - N atturufr. 17. Phytogeogr. Suec. XIII 1948: Eriophorum russeolum Fr. in the North 1945: Betula callosa Not0, a neglected species West of Iceland. _____:_ Bot. Not. 1948. in the Scandinavian subalpine forests. - Sv. 1950: Some innovations and nomenclatural sug Bot. Tidskr. 39. gestions in the Icelandic flora. - Ibid. 1950. 1947 a: On the variation in Scandinavian Betula LovE, A., & LovE, D., 1947: Studies on the Origin verrucosa Ehrh. with some notes on the Betula of the Icelandic Flora. I. Cyto-ecological series Verrucosae Sukacz. - Ibid. 41. Investigations on Cakile. - Reykjavik Univ. 1947 b: Two species of Betula from the Iceland Inst .. Appl. Se. Dpt Agric. Rep, B: 2. Miocene. - Ibid. 1948: Chromosome Numbers of Northern 1948: The main varieties of Picea Abies (L.) Plant Species. - Ibid. B: 3. Karst. in Europe, with a contribution to the LUDI, W., 1930: Methoden der Sukzessionsforschung theory of a forest vegetation in Scandinavia in der Pflanzensoziologie. - Abderhalden during the last Pleistocene glaciation. - Acta Handb. bioi. Arbeitsmeth. XI: 5. Hort. Berg. 14: 7. LUNDMAN, B., 1948: Nagot om den regionala varia LINDROTH, C. H., 1931: Die Insektenfauna Islands tionen av blomfargen hos midsommarblomster und ihre Probleme. - Zool. Bidr. fr. Uppsala (Geranium silvaticum L.). - Sv. Bot. Tidskr. XIII. 42. 1939: Die skandinavische Kaferfauna als Ergeb LuNDQVIS'.r, G., 1927: Bodenablagerungen und Ent nis der letzten Vereisung. - VII. Intern. Kongr. wicklungstypen der Seen. - Thienemann, Die Ent. Berlin 1938 I. Weimar. Binnengewasser II. Stuttgart. 1945: Die fennoskandischen Carabidae. Eine 1928: Studier i Olands myrmarker. - Sv. tiergeographische Studie. II. Die Karten. - Geol. Unders. C: 353. K. Vet. Vitt.-Samh. Handl. F. 6 (B) 4: 2. 1943: Norrlands jordarter. - Sv. Geol. Unders. 1948: See SuNDIUS & SANDEGREN. Arsb. 37: 6. 1949: Die fennoskandischen Carabidae. Eine 1944, 1948: De svenska fjallens natur.- STF:s tiergeographische Studie. Ill. Allgemeiner handb. om det svenska fjallet 2. Stockholm Teil. - K. Vet. Vitt.-Samh. Handl. F. 6 (B) 1944. - 2:a uppl. 1948. 4: 3. LYNGE, B., 1923: Vascular Plants from Novaya LINNAEUS, C., 1737: Flora Lapponica. - Amstelo Zemlya. - Rep. Sci. Res. Norw. Exp. Nov. dami. Zemlya 1921, 13. 1745: Flora Suecica. - Stockholmiae. 1933: On Dufourea and Dactylina. Three --, 1913: Iter lapponicum. Andra upplagan. Med Arctic Lichens. - Skr. om Svalbard o. Ishavet bilagor och noter ombesorjd af TH. M. FRIES. 59. - Skr. af Carl von Linne utg. af K. Sv. Vet. 1934: Some general results of recent Norwegian Akad. V. Uppsala. research work on Arctic Lichens. - Rhodora LoNNBERG, 1909: E., Om renarne och deras lefnads 36. vanor. - Bil. Forh. infor skiljedomstolen af MAGUIRE, B., 1943: Monograph of the Genus Arnica. 1909 i renbetesfr., Afd. I. Svensk inl. n:r 3. - Brittonia 4. (Not seen.) Uppsala. MALTE , M. 0., 1934 a: Antennaria of Arctic America. LONNQVIST, 0., 1939: Nagra vaxtfynd i Overtornea - Rhodora 36. trakten. - Bot. Not. 1939. 1944: Nagot om £loran pa sydberget Pullingi 1934 b: Critical Notes on Plants of Arctic i Overtornea socken i Norrbotten. - Bioi. America. - Ibid. Foren. i Boden Smaskr. 2. Boden. MANKER, E., 1944: Lapsk kultur vid Stora Lule alvs 1947: Floran i tva sydbranter i Norrbotten. - kallsjoar. - Nord. Mus. Acta Lapp. IV. Bot. Not. 1947. 1947: De svenska fj allapparna. - STF:s handb. 1949: Pa sydberg och backdalar i N orrbotten. om det sv. fjallet 4. Stockholm. 188 SEL.ANDER: Floristic Phytogeography of South-Western Lule Lappmark
MANNERFELDT, C. M:soN, 1945: Nagra glacialmorfo NoRDHAGEN, R., 1943: Sikilsdalen og Norges fjellbei logiska formelement . och deras vittnesb6rd ter. - Bergens Mus. Skr. 22. om inlandsisens avsmaltningsmekanik i svensk NoRDSTROM, F.: 'see NoRDSTROM, WAHLGREN & och norsk fj allterrang. - Geogr. Ann. 1945: TULLGREN. 1-2. NoRDSTROM, F., WAHLGREN, E., TuLLGREN, A., 1941: MARIE -VICTORIN, FRERE, 1938: Phytogeographical Svenska fjarilar. Systematisk bearbetning av Problems of Eastern Canada. - Contrib. Labor. Sveriges storfjarilar, Macrolepidoptera. Bot. Univ. Montreal 30. Stockholm. MrLANKOVITCH, M., 1930: Matematische Klimalehre NoRMAN, J. M., 1895: Norges arktiske Flora II. - und astronomische Theorie der Klimaschwan Kristiania. kungen. - Handb. Klimatologie I: A. NYGREN, A., 1936: Carex holostoma, en for Sverige MuNTHE, H., 1910: Preliminary list of plant remains ny fanerogam, funnen i Torne Lappmark. - found in the Hernogyttja. - Geol. Foren. Sv. Bot. Tidskr. 30. Forh. 32. 1946: The Genesis of some Scandinavian Species 1946: Nya bidrag till kannedomen om Harn6 - of Calamagrostis. - Hereditas 32. gyttjan. Sv. Geol. Unders. C: 481. NYLANDER, W., 1882: Lichenes Lapponiae orientalis. NANNFELDT, J. A., 1935: Taxonomical and Plant - Not. Sallsk. Fauna Fl. Fenn. Forh. VIII. Geographical Studies in the Poa laxa Group. NYMAN, E., 1895: Vegetationsbilder fran Lappland. - Symb. Bot. Ups. 1: 5. - Bot. Not. 1895. 1940: On the Polymorphy of · Poa arctica R. OLSEN, C., 1936: Kalkplanter og kalksky Planter. __:___ Br., with Special Reference to its Scandinavian Tidskr. Land0konomie 11. Forms. - Ibid. IV: 4. 0MANG, s. 0. F., 1933 a: Ubersicht uber die Hiera 1941: Sagina caespitosa funnen i Lule Lapp ciumflora Ost-Gronlands. - Skr. om Svalbard mark. - Bot. Not. 1941. o. Ishavl3t 55. 1947: Nagra synpunkter pa den skandinaviska· 1933 b: Contributions to the Hieracium Flora fjallflorans alder. - K. Vet.- Soc. Arsb. 1947. of Greenland. - In SEIDENFADEN 1933. NATHORST, A. G., 1883: Nya bidrag till kannedomen 1937: Uber einige Hieracium-Arten aus Gran om Spetsbergens karlvaxter och dess vaxt land. - Skr. om Svalbard o. Ishavet 40. geografiska forhallanden. - K. Sv. Vet.-Akad. 193S: Monographische Bearbeitung der Hiera Handl. 20: 6. cia Islands. - Oslo. NAUMANN, E., 1920: Nagra synpunkter angaende 0RTENBLAD, Ta., 1902: Om masurbjorkens (Betula de limniska avlagringarnas terminologi. - verrucosa Ehrh.) utbredning i Skandinavien. Sv. Geol. Unders. Arsb. 14: 2. - Bot. Not. 1902. NrLSSON, T., 1935: Die pollenanalytische Zonen OsTENFELD, C. H., 1926: The flora of Greenland and gliederung der spat- und postglazialen Bil its Origin. - K. Danske Vid.-Selsk. Biol. dungen Schonens. - Geol. F6ren. Forh. 57. Medd. 6: 3. NoRDHAGEN, R., 1928: Die Vegetation und Flora OsvALD, H., 1947: Vaxternas vapen i kampen om des Sylenegebietes I, II. - Skr. utg. av D. utrymmet. - Vaxtodling. Skr. Inst. Vaxt Norske Vid.-akad. J. Mat.-naturv. klasse 1927. odlingsl. K. Lantbruksh6gsk. 2. F6rs6k ror_ 1931: Studien uber die skandinavischen Rassen kampen mot ograsen 1935- 1946. Uppsala. des Papaver radicatum Rottb. sowie einige 1949: Root Exudates and Seed Germination. mit denselben verwechselte neue Arten. - - Annals Royal Agric. Coli. Sweden 16. Bergens Mus. Arb. 1931. Naturv. rekke 1. PESOLA, V. A., 1928: Calcium carbonate as a factor 1933: De senkvartaere klimavekslinger i Nord in the distribution of plants in Finland. (Fin europa og deres betydning for kulturforsk nish). - Ann. Soc. Zool. Bot. Fenn. Vanamo ningen. - Inst. Sammenl. Kulturforskn. A: 12. 9: 1. 1935: Om Arenaria humifusa og dens betydning for utforskningen av Skandinavias eldste 1934: Uber die Felsenvegetation in NE-Kuu floraelement. - Bergens Mus. Arb. 1935, samo und SE-Kuolajarvi ..- Ibid. 5: 7. Naturv. rekke 1. PIERCE, J. H., 1936: Range Extensions of certain 1936: Versuch einer neuen Einteilung des Plants on the Gaspe Peninsula. - Rhodora subalpinen-alpinen Vegetation Norwegens. - 38. - Ibid. 1936 7. PoHLE, R., 1903: Pflanzengeographische Studien 1937: Om Norges fjellflora og dens oprinnelse. uber die Halbinsel Kanin und das angrenzende - Naturen 1937. Waldgebiet. - St. Petersburg. 1940: N orsk flora. - Oslo. 1917: Wald- und Baumgrenzen in Nord-Russ- Literature Cited 189
land. - Zeitschr. Ges. f. Erdkunde zu Berlin RuPRECHT, F. J., 1845: Flores Samojedorum cisura 1917. lensium. - St. Petersburg. PoLUNIN, N., 1940: Botany of the Canadian East --, 1856: Flora boreali-uralensis. - Ibid. Arctic. I. Pteridophyta and Spermatophyta. SAMUELSSON, G., 1910 a: RegionfOrskjutningar - N at. M us. Canada Bull. 92. Dalarna. - Sv. Bot. Tidskr. 4. 1943: Geographical distribution of Arenaria 1910 b: Uber die Verbreitung einiger ende humifusa Wahlenb., new to the flora of Spits mischer Pflanzen. - Arkiv f. Bot. 9: 12. bergen. - Nature 152. 1921: Carex dioeca-gruppen i den nordiska PoNTIN, M. AF: See SwARTZ 1821. floran. - Acta Fl. Suec. I. PoRSILD, A. E., 1947: The Genus Dryas in North 1934: Die Verbreitung der hoheren Wasser Amerba. - Canadian Field Natural. 61. pflanzen in Nordeuropa (Fennoskandia und PORSILD, M. P., 1930: Stray Contributions to the Danemark). - Acta Phytogeogr. Suec. VI. Flora of Greenland. I-V. - Medd. om Gr0nl. 1943: Die V erbreitung der Alchemilla-Arten LXXVII. aus der vulgaris-Gruppe in N ordeuropa. - 1932: Alien Plants and Apophytes of Green Ibid. XVI. land. - Ibid. 92: l. SAMZELIUS, H., 1890: Vegetationsiakttagelser inom 1935: Stray Contributions to the Flora of Pajala socken af Norrbottens lan. - Bot. Not. Greenland. VI-XII. - Ibid. 93: 3. 1890. PoRSILD, M. P. & A. E., 1920: The Flora of Disco SANDBERG, G., 1938: Redogorelse for undersokningar Island and the Adjacent Coast of West Green utforda med understod av Sallskapets stipen land from 66°-71° N. lat. - Ibid. LVII. dier. - Ymer 1938. PosT, L. voN, 1924: Ur de sydsvenska skogarnas re 1940: Den pagaende klimaMorbattringen. - gionala historia under postglacial tid. - Geol Sv. Vall- o. MosskulturfOren. Kvartalsskr. For. Forh. 46. 1940. 194 7: Diskussionsinlagg vid N ordiskt kvartar SANDEGREN, R., 194:3 : Hippophae rhamnoides L. i geologiskt mote den 5-7 november 1945. Sverige i senkvartar tid. - Sv. Bot. Tidskr. 37. Ibid. 69. SAURAMO, M., 1942: Kvartargeologiska studier i 6stra PuGSLEY, H. W., 1933: The Euphrasias of Iceland Fennoskandia. - Geol. Foren. Forh. 64. and the Faroes. - J ourn. Bot. British and 1947 : Studier over de senglaciala nivMorand Foreign LXXI. ringarna i Fennoskandia. - Ibid. 69. RAMSAY, W., 1931: Geologiens grunder. 3:e uppl. see LINDROTH 1949. II. Oversikt av den geologiska utvecklingen. ScHROETER, C., 1908: Das Pflanzenleben der Alpen. Fennoskandias geologi. - Stockholm. Eine Schilderung der Hochgebirgsflora. REGEL, K., 1923: Die Planzendecke der Halbinsel Kola. Ziirich. Lapponia Varsugae. - Mem. Faculte Sci. Univ. ScHULZ, 0. E., 1924: Cruciferae. Sisymbriaceae. Lithuanie 1922. Pflanzenreich IV: 105. RosENDAHL, H. V., 1892: Nagra anteclmingar fraq SEIDENFADEN, G., 1933: The Vascular Plants of en exkursion genom Lulea lappmark sommaren South East Greenland 60°04' to 64°30' N. lat. 1892. - Bot. Not. 1892. - Medd. om Gmnl. 101: 1. RuBEL, E., 1930: Lichtklima und Lichtgenuss: - SELANDER, S., 1942: Potentilla emargina.ta Pursh i Abderhalden, Handb. biol. Arbeitsmeth. XI: 5. Sverige. - Bot. Not. 1942. RuDBECK, 0. fil., 1720: Index Plantarum praecipua 1945 a: Ny svensk forekomst av Potentilla rum, quas in Itinere Lapponico anno 1695 emarginata Pursh. - Ibid. 1945. observavit Dn. Olaus Rudbeck, filius. - Acta 1945 b: Mitt roligaste vaxtfynd. - Sveriges Litt. Suec. IV. Natur 36. RuNE, 0., 1945 a: Karlvaxtfloran i Offerdals och 1947: Urtica gracilis Ait. in Fennoscandia. nordostra Kalls fjallomrade. - Arkiv f. Bot. - Sv. Bot. Tidskr. 41. 32 A: 11. 1948: Linne i Lule lappmark. Sv. Linne- 1945 b: Nagra anmarkningsvarda vaxtfynd i sallsk. arsskr. XXX. sodra Lapplands fjall. - Sv. Bot. Tidskr. 39. 1949: Potentilla norvegica som spontan i --, 1947: Minuartia rubella funnen i sodra Lapp Sverige. - Sv. Bot. Tidskr. 43. land. - Ibid. 41. SERNANDER, R., 1896: Nagra ord med anledning af 1948: Nya vaxtfynd i Lycksele Lappmarks GUNNAR ANDERSSON, Svenska vaxtvarldens fjallomrade. - Ibid. 42. historia. - Bot. Not. 1896. 1949: Serpentinfloran - en botanisk gata. - 1899: Studier ofver vegetationen i mellersta Norrb. Natur. Smaskr. l. Boden. Skandinaviens fjalltrakter. 2. Fjallvaxter i 190 SEL.ANDER: Floristic Phytogeography of South-Western Lule Lappmark
barrskogsregionen. - Bib. K. Sv. Vet.-Akad. STERNER, R., 1921: Carex ligerica J. Gay. En floris Handl. 24 Ill: 11. tisk och vaxtgeografisk studie. - Acta Fl. SERNANDER, R., 1900: Yttrande med anledning af A. Suec. I. Hambergs foredrag om Sarjektraktens qvartara 1922: The Continental Element in the Flora historia. - Geol. Foren. Forh. 22. of South Sweden. - Geogr. Ann. IV. 1901: Den skandinaviska vegetationens sprid 1936: Helianthemum oelandicum (L.) Willd. ningsbiologi. Zur Verbreitungsbiologie der och dess anfOrvanter pa bland. - Sv. Bot. skandinavischen Pflanzenwelt. - Berlin & Tidskr. 30. Upsala. -- , 1938: Flora der Insel Gland. Die Areale der 1905: Flytj ord i svenska fjalltrakter. - Geol. Gefasspflanzen Glands nebst bemerkungen Foren. Forh. 27. zu ihrer Oekologie und Soziologie. - Acta SETCHELL, W. A., 1935: Pacific insular floras and Phytogeogr. Suec. IX. Pacific paleography. - Amer. Naturalist 69. 1944: Galium pumilum Murr. i nordvastra SIMMONS, H., 1906: The Vascular Plants in the Flora Europa. - Medd. Goteb. bot. tradg. XV. of Ellesmereland. - Rep. Se. Norw. Arct. 1945: See H. FRIES 1945. Exp. in the >>Fram>> 1898-1902 2. STRAND, A., 1946: Nord-Norges Coleoptera. - Troms0 1907 a: Uber einige lapplandische Phanero Mus. Arsh. 67 (1944). gamen. - Arkiv f. Bot. VI: 17. STROMFELT, H. F. G., 1884: Islands karlvaxter, be 1907 b: Nagra bidrag till Lule lappmarks flora. traktade fran vaxtgeografisk och floristisk syn - Bot. Not. 1907. punkt. - Ofvers. K. Vet .-Akad. Forhandl. 8. 1913: A survey of the phytogeography of the SuNDius, N., & SANDEGREN, R., 1948: Interglacial Arctic American Archipelago with some notes fyndet vid Llmgsele. Med bidrag av T. LAGER on its exploration. - Lunds Univ. Arsskr. BERG, C. LINDROTH och H. PERSSON. Bihang. N. F. 2 9: 19. Nya data rorande det interglaciala Bollnas SJORS, H., 1946 a: Myrar i Muddus. - Sveriges Natur fyndet av B. HALDEN. - Sv. Geol. Unders. 37. Arsb. 42: 4.
--, 1946 b: Myrvegetationen i ovre Langanomradet. SwARTZ, 0., 1799: Botaniske Anmarkningar; samt - Arkiv f. Bot. 33 A: 6. beskrifning pa en Svensk och hittills okand 1948: Myrvegetation i Bergslagen. - Acta Spergula stricta. - Nova Acta Holm. XX. Phytogeogr. Suec. 21. 1821: Flora for tragten kring berget Arras. - SKOTTSBERG, C., 1926: Vascular plants from the Bib. t. Aminnelsetal ofver Kongl. Wet.-Acad. Hawaiian Islands. I. - Medd. fr. Goteborgs framl. ledamot, Herr Olof Swartz, hallit inf. Bot. Tradg. II. Kongl. Wet.-Acad. den 21 Nov. 1820; af dess 1928: Remarks on the Relative Independency Ledamot MAGNUS AF PONTIN. Stockholm. of Pacific Floras. - Proceed. Third Pan-Paci 1829: Adnotationes Botanicae quas reliquit fic Se. Congr. Tokyo. Olavus Swartz . . . Post mortem Auctoris 1931: Remarks on the Flora of the high Hawai collectae . . . a J OHANNE EM. WIKSTROM. ian Volcanoes. - Medd. Goteb. bot. tradg. 6 Holmiae. (1930). SvENONIUS, F., 1884: Nya olivinstensforekomster i SMITH, H., 1920: Vegetationen och dess utvecklings Norrland. - Geol. Foren. Forh. VII: 88. historia i det centralsvenska hogfjallsomradet. 1900 a: Ofversikt af Stora Sjofallets och angran - N orrl. Handbibl. IX. sande fjalltrakters geologi. II. Berggrunden. 1940: Carex arctogena nova species. - Acta -Ibid. 22. Phytogeogr. Suec. XIII. 1900 b: Yttrande med anledning af A. HAM S0RENSEN, N. A., 1949: Gjeilvasskammene - nuna bergs foredrag om Sarj ektraktens qvartara takker i Trollheimens mitte? - Naturen 73. historia. - Ibid. S0RENSEN, Tu., 1933: The Vascular Plants of East Svensk Botanik, 1819. 8. Bandet utg. af G. J. Greenland from 71°00' to 73°30' N. lat. - BILLBERG. - Stockholm. Medd. om Gronl. 101: 3. SYLVEN, N., 1904: Studier ofver vegetationen i Torne STEFANSSON, s., 1924: Flora Islands. II utgafa, lappmarks bjorkregion. - Arkiv f. Bot. 3: 3. aukin. - Kaupmannahofn. SYLVEN, N., & B.AGENHOLM, G., 1902: Ruderatvaxter STEFFEN, H., 1931: Vegetationskunde von Ostpreus antecknade fran Lule lappmark sommaren sen. - Jena. 1901. - Bot. Not. 1902. 1937: Gedanken zur Entwicklungsgeschichte TENGWALL, T. A., 1913: De sydliga skandinaviska der arktischen Flora. - Beih. Bot. Zentralbl. fjallvaxterna och deras invandringshistoria. 56: B, 57: B. - Sv. Bot. Tidskr. 7. Literature Cited 191
TENGWALL, T. A., 1914: Braya alpina Sternb. & WAHLGREN, E.: See NoRDSTROM, WAHLGREN & Hoppe vid V astenjaure. - Ibid. 8. TULLGREN. 1916: Uber die Bedeutung des Kalkes fiir die WALDHEIM, S., och WEIMARCK, H., 1943: Bidrag till V erbreitung einiger schwedischer Hochgebirgs Skanes Flora 18. Skanes myrtyper. - Bot. pflanzen. - Ibid. 10. Not. 1943. 1920, 1925: Die Vegetation des Sarekgebietes I, WARMING, E., 1888: Tabellarisk Oversigt over Gr0n II. - I 1920, Naturwiss. Unters. des Sarekgeb. lands, Islands og Faer0ernes Flora i 1887. - in Schwed.-Lappl. Ill, I: 4. II 1925, ibid. I: 8. Vidensk. Medd. fr. Naturhist. Foren. i K0ben 1924: Die GeHisspflanzen des Sarekgebietes. havn (1887). - Ibid. II: 7. WEIMARCK, H., 1943: See WALDHEIM & WEIMARCK. THORARINSSON, S., 1937: Vatnaj okull. Scientific 1945: Rec. av HYLANDER, Nomenklatorische Results of the Swedish-Icelandic Investiga und systematische Studien, &c. - Bot. Not. tions 1936-1937. Chapter II. The main 1943. geological and topographical features of Ice- · VESTERGREN, T., 1902: Om den olikformiga snobe land. - Geogr. Ann. XIX. tackningens inflytande pa vegetationen i Sarek THUNMARK, S., 1931: Der See Fiolen und seine Vege fjallen. - Bot. Not. 1902. tation. - Acta Phytogeogr. Suec. II. 1938: Systematische Beobachtungen uber Myo ToLF, R., 1893: GranUimningar i svenska torfmossar. sotis silvatica (Ehrh.) Hoffm. und verwandte - Bih. K. Sv. Vet.-Akad. Handl. 18: 3. Formen. - Arkiv f. Bot. 29. ToLMATCHEV, A., 1930: Sur la provenance de la flore VESTERLUND, 0., 1924: Forteckning over fanerogam du Vaigatch et de Novaja Semlja. - Trav. och karlkryptogamfloran inom Jokkmokks och Mus. Bot. Acad. Se. URSS 22. Kvikkj okks skogsregion. - Sv. Bot. Tidskr. 18. TRAUTVETTER, E. R., VON, 1849-50: Die pflanzen WESTLING, R., 1906: Pa botanistfard i Lappmarken. geographischen Verhaltnisse des europaischen - Svensk Farmaceutisk Tidskr. 1906. Russlands. - St. Petersburg. WICHURA, M., 1859: Ein Ausflug nach der Lulea TROLL, K., 1925: Ozeanische Zuge im Pflanzenkleid Lappmarken. - Flora 42. Mitteleuropas. - Freie Wege vergl. Erdk. WIKLUND , K. B., 1947: Lapparna. - Nordisk Kultur Festg. E. von Drygalsky. Munchen u. Berlin. X. Stockholm. TURESSON, G., 1925: The plant species in relation to WIKSTROM, J. E.: See SwARTZ 1829. habitat and climate. - Hereditas 6. WILMOTT, A. J., 1935: Evidence in Favour of Survival
--, 1927: Contributions to the genecology of glacial of the British Flora in Glacial Times. - Pro relics. - Ibid. 9. ceed. Royal Soc. B 118: 808. WISTRAND, G., 1934: Bidrag till kannedomen om ULANDER, A., 1929: I .sydbergens hagn. - Norrbotten. Norrb. lans Hembygdsforen. Tidskr. 1929. flora i Pite Lappmarks barrskogsregion. - Sv. Bot. Tidskr. 28. WAHLENBERG, G., 1808: Berattelse om matningar 1946: Vaxtlokaler vid Tjeggelvas. - Norr och observationer for att bestamma lappska bottens N atur. Norrb. Bioi. Foren. Smaskr_ fjallens hojd och temperatur vid 67 graders pol 3. Boden. hojd forrattade 1807. - Stockholm. WITTING, M., 1949: Kalciumhalten i nagra nord 1810: Anmarkningar om Lappska Vegetationen svenska myrvatten. - Sv. Bot. Tidskr. 43. med beskrifning om Myosotis deflexa eller ett VoGT, TH., 1927: Sulitelmafeltets geologi og petro nytt Forgat mig ej fran Lappland. - Nova 121. Acta Holm. XXXI. grafi. - Norges Geol. Unders. --, 1812: Flora lapponica. - Berolini. WAHLBERG, P. F., 1844: Vaxtformer i Lulea Lapp WAHLENBERG, G.: G. Wahlenbergs Resa i Lapp mark. - Ofvers. K. Sv. Vet.-Akad. Handl. marken ar 1807. - Manuskript. Uppsala Univ. 1844. Bibl. No. S 169 b. INDEX TO PHANEROGA MS AND PTER IDOPHYTES
Figures in italics indicate passages where data concerning the distribution and history of a plants species are given, or a species is dealt with at some length.
Achillea Millefolium 34, 36, 109 Anthriscus silvestris 14, 96, 121 Aconitum septentrionale 10, 55, 85, lOO, 120, 141, 142, 163 Anthyllis Vulneraria v. lapponica 14, 63, 119, 120, 121, Actaea erythrocarpa 14, 22, 94, 96, 104, 105, 120, 122, 173 136, 145, 175 A. spicata 96, 104, 105 Arabis alpina 10, 54, 68, 88, 120, 156, 169, 171 Acloxa Moschatellina 173 A. hirsuta v. subalpestris 14, 63, 103, 120, 141, 142, 175 Agrostis l:)orealis 12, 34, 48, 55, 56, 67, 68, 140, 171 Aretagrostis latifolia 164, 173 A. canina 13, 34, 69, 96, 178 Arctostaphylos alpina 10, 50, 55, 56, 58, 118, 140, 171 A. stolonifera v. arctica 16, 46, 47, 63, 100, 139 A. Uva-ursi 11, 60, · 94, 121, 122 A. tenuis 15, 34, 47, 50, 96, 102, 141 Arenaria ciliata coll. 169 Ajuga pyramidalis 15, 90, 91, 102, 120, 136, 141; 142, A. cil. ssp. pseudofrigida 164, 173, 145, 147 A. humifusa 11, 31, 64, 80, 100, 116, 128, 130, 132, 134, Alchemilla alpina 10, 62, 66, 82, 138, 141, 161, 168, 176 136, 141, 157, 158, 164, 170, 171 A. borealis 63 A. norvegica 13, 31, 64, 88, 100, 116, 119, 120, 130, 132, A. fa.eroensis 7 5 134, 136, 141, 143, 145, 151, 157, 160, 163, 164, 166 A. filicaulis 15, 50, 70, 90, .9 1, 120, 141, 171, 175 A. serpyllifolia 107 A. glomerulans 34, 55, 62, 68 Armeria maritima 162 A. Hoppeana 168 A. scabra ssp. sibit·ica 165, 173 A. Murbeckiana 34, 55, 93 Arnica alpina 10, 56, 63, 99, 111, 11(\, 118, 126, 138, 141, - A. norvegica 16, 63 149, 151, 152, 156, 166, 173 A. vestita 16, 70, 96, 103, 171 Artemisia bottnica 59 A. \Vichurae 34, 55, 56, 65, 164, 173 A. laciniata 62 Alisma Plantago-aquatica 50 A. norvegica 163 Allium Schoenoprasum v. sibiricum lG4 A. rupestris 60, 62, 83 A. strictum 83 A. vulgaris 60 Alnus incana v. virescens 10, 63, 98 Asplenium septentrionale 16, 85, 92, 103, 104, 105, 119, 121 Alopecurus aequalis 12, 47, 94, 171 A. Trichomanes 15, 96, 103, 121, 136 A. pratensis ssp. alpestris 12, 34, 85, lOO, 114, 161, 164, 166 A. viride 13, 57, 62, 85, 97, 89, 120, 131, 134, 141, 142, A. prat. ::>sp. praticolus 36 143, 151, 152, 153, 157, 160, 169, 170, 171, 172 Andromeda Polifolia 11, 58, 93, 118 Aster sibiricus 83, 163, 166
Anemone patens 83 Astragalus alpinus 10, 34, 54, 60, 86, · 168, 171 A. silvestris 83 A. alp. ssp. arcticus 48, 86, 165 Angelica Archangelica ssp. norvegica 10, 22, 37, 55, 70, A. frigidus 48, 99, 161, 163, 165, 166, 172, 173 87, 90, 95, 119, 120, 156 A. norvegicus 11, 56, 86, 87, 100, 131, 141, 157, 160 A. silvestris 90, 96 Athyrium alpestre 13, 55, 56, 67, 72, 139, 171 Antennaria alpina 11, 48, 54, 65, 72, 82, 87, 137, 170 A. alp. v. americanum 67, 72 A. carpatbica 12, 53, 54, 86, 99, 126, 141, 151, 152, 156, A. alp. v. gaspense 67 164 A. alp. v. typicum 67 A. dioeca 11, 54, 55, 97 A. crenatum 163, 166 A. lapponica. 16, 63, 119, 141, 157, 160 A. Filix-femina 10, 22, 96, 104, 141 A. Porsildii 5, 80, 164 A vena elatior 116 Anthoxanthum alpinum 12, 34, 55, 69, 90, 139 Barbarea stricta 12, 97, 121 A. odoratum 12, 69 Bartsia alpina 11, 47, 54, 55, 62, 68, 171 Index to Phanerogams and Pteridophytes 193
Betula callosa 17, 75, 85, 89, 98 Carex angarae 95, 96, 157, 160 B. coriacea 18, 97 C. aquatilis 12, 22, 37, 56, 92, 95, 166 B. nana 10, 37, 55, 57, 58, 60, 70, 73, 98, 118, 147 C. aq. v. stans 92 B. odorata coli. 59 C. arctogena 64, 80, 86, 141, 142, 151, 152, 156, 157, 160, B. pubescens coli. 60 163, 168 B. tortuosa 70, 90, 98, 139 C. atrata 10, 48, 54, 55, 66, 156 B. verrucosa coli. 103, 104, 134, 135 C. atr. ssp. atrosquame a 66 B. verr. v. lapponica 15, 18, 94, 104 C. atrofusca 12, 48, 54, 55, 85, 87, 136, 156, 169 B. verr. v. saxatilis 104 C. bicolor 12, 28, 85, 86, 100, 108, 130, 141, 151, 157, 160, Blechnum Spica,nt 14, 75, 96, 103 163, 164, 169, 170 Botrychium boreale 13, 93, 156, 170, 171 C. Bigelowii 12, 30, 48, 50, 52, 53, 58, 9.3, 124, 140, 169, 171 B. Lunaria 13, 96, 121, 136 C. bnmnescens 14, 34, 37, 93, 140, 169, 171 B. matricariifolium 171 C. Buxbaumii 16, 97 B. multifidum 17, 69, 80, 94, 134, 135, 171 C. caespitosa 14, 94, 136 B. silaifolium 69 C. canescens 12, 92, 97, 102, 115, 128 Brassica napus f. oleifera 115 C. can. v. robustior 92 Br. rapa f. oleifera 115 C. capillaris 12, 54, 91, 120, 169, 170, 172 Braya glabella 64, 169 C. capitata sens. lat. 12, 170 Br. linearis 13, 31, 64, 80, 100, 108, 116, 119, 120, 130, C. capitata sens. str. 85, 136, 141, 157, 168 131, 132, 136, 138, 141, 142, 143, 147, 151, 157, 160, C. chordorrhiza 12, 94, 95 164, 169 C. dioeca 12, 55, 97 Br. purpurascens 74, 165, 169 C. disperma 14, 94 Cakile edentula 74 C. flacca 76 Calamagrostis arundinacea 63, 164 C. flava 14, 92, 102, 121, 141 C. canescens 63, 164 C. fusca 46, 55, 97 C. chalybaea, 164 U. glacialis 11, 31, 56, 87, 116, 141, 142, 143, 145, 151, C. deschampsioides 165 152, 153, 157; 158, 163, 166, 169, 170, 172 C. epigeios 63 C. globularis 16, 94, 96, 134, 135 C. lapponica 12, 54, 85, 118 C. holostoma 164, 173 C. neglecta coll. 12, 55, 63 C. juncelia 94, 95 C. negl. ssp. australis 94 C. Lachenalii 12, 37, 46, 48, 52, 53, 58, 73, 74, 87, 123, C. negl. ssp. borealis 73, 89 124, 129, 139, 169, 171 C. obtusata 164 C. lasiocarpa 12, 97,- 101, 141 C. purpurea 12, 63 C. laxa 82 Callitriche polymorpha 16, 97 C. leporina 178 C. verna 11, 47, 94, 102, 140 C. ligerica 39 Calluna vulgaris 11, 18, 97, 102, 141 C. limosa 12, 94, 95 C. vulg. v. atlantica 70, 178 C. livida 16, 64, 80, 134, 171 Caltha palustris 10, 57, 92, 97, 141 C. loliacea 14, 94 Calypso bulbosa 96 C. Lyngbyei 69, 74 Campanula alpina 168 C. macloviana 14, 34, 64, 74, 100, 112, 113, 128, 141, 142, • C. barbata 163, 168, 174 164, 173 C. rotundifolia 11, 15, 34, 54, 56, 73, 74, 89, 93, 119, 120, C. magellanica 12, 58, 97, 136, 175 1�{6, 175 C. marina (syn. C. glareosa) 82 C. uniflora 10, 54, 6S, 73, 86, 87, 99, 110, 111, 112, 130, C. maritima 170 132, 138, 141, 149, 152, 153, 156, 163, 104, 169, 171 C. membranacea 70 Cardamine bellidifolia 10, 46, 48, 53, 86, 87, 125, 142, 148, C. microglochin 11, 56, 57, 66, 87, 100, 141, 151, 152, 153, 156, 171 157, 160, 169, 170 C. flexuosa 76 C. misandra 99, 163, 164, 166, 169 C. hirsuta 7 5 C. nardina 13, 31, 56, 64, 73, 74, 100, 116, 118, 130, 142, C. pratensis coli. 10, 46 151, 152, 153, 157, 158, 164, 169 C. prat. ssp. angustifolia 16, 54, 100, 136, 142, 152, 157, C. norvegica 11, 54:, 55, 57, 66, 72, 82, 157, 160, 172 160, 164, 170, 175 C. obtusata 62 C. prat. ssp. typica 96, 136, 171 C. Oederi 14, 47, 50, 70, 72, 96, 103, 121 Cardaminopsis petraea 82, 163 C. ornithopoda 13, 50, 91, 120 Carex adelostoma 14, 63 C. pallescens 14, 96, 103, 121, 136
14 - 496149 Sten Selo.nder I 194 SELANDER: Floristic Phytogeography of South- Western Lule Lappmark
Carex panicea 14, 50, 70, 96, 102, 136 Crepis tectorum 119 C. parallela 12, 54, 56, 64, 86, 87, 142, 151, 152, 156, Cryptogramma crispa 12, 62, 81, 89, 138, 161, 178 163, 164, 169 Cuscuta Epithymum 109 C. pauciflora 10, 94 Cystopteris fragilis ssp. eu-fragilis I 0, 56, 93, ll9, 120 C. pediformis ssp. rhizodes 163, 166 C. frag. ssp. Dickieana 136 C. perfusca 66 C. montana ll, 55, 56, 67, 87, 100, 142, 157, 170 C. physocarpa 70, 72 Dactylis glomerata 116 C. rariflora 70, 86, $7, 90, 131, 153, 157, 158, 163, 171 Daphne Mezereum 12, 22, 94, 104, 105, 120, 122 C. rostrata 10, 60, 69, 97 Deschampsia alpina 10, 48, 53, 59, G5, 87, 153, 157, 160 C. rotundata 12, 94, 95, 96, .138, 160, 168, 169, 179 D. atropurpurea 13, 36, 64, 80, 87, 88, 139, 161, 171, 179 C. rufina 13, 28, 31, 46, 58, 64, 87, 89, 124, 139, 141, 153, D. atr. ssp. magellanica 64 157, 160, 164, 173 D. atr. ssp. paramuschirensis 64 C. rupestris 12, 48, 53, 54, 85, 87, ll8, 142, 143, 145, 156 D. caespitosa 10, 37, 93 169, 170, 172, 176 D. ca>sp. v. glauca 47 C. saltuensis 69 D. flexuosa 10, 18, 37, 52, 56, 69, 90, 139 C. saxatilis 12, 54, 55, 70, 72, 87, 90, 168, 169 D. rhenana 62 C. scirpoidea 80, 162, 164, 169 Dianthus superbus 173 C. tenuiflora 13, 85 Diapensia lapponica 10, 30, 54, 58, 65, 72, 86, ll8, 140, C. vaginata 12, 54, 55, 69, 90, 93 170, 171 C. vesicaria 14, 96, 176 D. lapp. ssp. obovata 65, 72 C. viridula 70, 72 Digitalis purpurea 59 Carum Carvi 34 Draba alpina 10, 53, 85, 87, 100, 124, 125, 126, 131, 137, Cassiope hypnoides 10, 31, 32, 48, 52, 53, 65, 73, 126, 139, 142, 148, 151, 152, 157, 160, 169 149, 170, 171 Dr. Bellii 170 C. tetragona 10, 31, 37, 38, 52, 56, 86, 99, 108 ££., 131, Dr. cinerea 83 141, 156, 165, 166, 173, 176 Dr. cin. v. ladogensis 83 Centaurea Scabiosa 83 Dr. crassifolia 16, 64, 80, 99, 100, 127, 128, 130, 131, 141, Cerastium alpinum 10, 48, 54, 66, ll9, 120, 139, 168, 171 148, 151, 157, 158, 164 C. arcticum 13, 48, 53, G4, 86, 124, 125, 152, 156, 157, Dr. fladnizensis 163, 165, 169 158, 163 Dr. hirta 48, 55, 86, 89, ll9, 120, 136, 141, 142, 143, 145, c._ fontanum ssp. scandicum 54, 55, 64, 156 151, 157, 163, 170, 172 C. glabratum 10, 54, 55, 134, 142, 156, 168, 172 Dr. incana 172 C. holosteoides 34 Dr. lactea ll, 53, 85, 86, 87, 100, 124, 125, 130, 137, 142, C. Regelii 74 145, 148, 151, 152, 153, 156, 157' 158, 163, 164, 170 C. trigynum 10, 46, 48, 53, 54, 66, 128, 140, 169, 171 Dr. nemorosa 14, 94, 104, 107, 121 Chamaenerium angustifolium 10, 18, 97, 128 Dr. nivalis 12, 48, 53, 54, 85, 87, 142, 145, 152, 156, 163, Ch. latifolium 7 4 164, 169, 170 Chamorchis alpina ll, 54, 55, 81, 87, 152, 153, 157, 160, Dr. norvegica 10, 34, 48, 53, 54, 64, 87, 143, 156, 166, 170 163 Dr. norv. v. nammatis 120 Chrysanthemum Leucanthemum 34 Drosera anglica v. pusilla 14, 94 Chrysosplenium tetrandrum ll4, 165 Dr. rotundifolia v. gracilis 14, 94 Circaea alpina 13, 96, 103, 121 Dryas integrifolia 90 Cirsium heterophyllum ll, 90, 95 Dr. octopetala 10, 30, 54, 57, 59, 60, 61, 6,'2, 82, 84, 87, Clematis alpina ssp. sibirica 163, 166 90, ll7' 126, 142, 143, 145, 156, 170, 176 Cochlearia groenlandica 7 4 Dr. punctata 90 Coeloglossum viride coll. 12, 170 Dryopteris austriaca 12, 22, 92, 96, 144 C. vir. ssp. bracteatum 72 Dr. Filix-mas 13, 96, 101, 103, 121, 141 C. vir. ssp. islandicum 55, 56, 72, 85, 175 Dr. fragrans 164, 170, 171, 173 Convallaria majalis 16, 49, 9'7, 104 Dr. spinulosa 16, 96, 104, ll9, 121 Corallorhiza trifida 13, 94, 95, 141, 142 Elatine Hydropiper 14, 49, 97 Cornus suecica 11, 65 Elymus mollis 180 Corydalis fabacea 5 Elytrigia repens 34, ll5 Corylus Avellana 59, 60 Empetrum hermaphroditum 18, 37, 54, 73, ll7, ll8 Crassula aquatic a 7 5 E. nigrum coll. 57 Crepis multicaulis 165, 173 Epilobium alsinifolium 14, 81, 141 Cr. paludosa 14, 90, 91 E. anagallidifolium 14, 46, 56, 85, 156, 171 Index to Phanerogams and Pteridophytes 195
Epilobium collinum 13, 81, 89, 92, 104, 120, 141, 145 Galium palustre 14, 47, 49, 70, 97 E. davuricum 14, 27, 95, 141, 160, 171 G. pumilum ssp. islandicum 75 E. Hornemanni 14, 65, 140, 161, 171 Hippophae rhamnoides 59, 60, 61, 62 Luzula arcuata sens. str. 12, 91 Hippuris vulgaris 12, 22, 97, 140 L. confusa 53, 93, 99, ll8, 124, 125, 129, 171 Hydrocotyle vulgaris 7 5 L. multiflora ssp. comosa 70, 72 Hypericum maculatum 59 L. mult. ssp. frigida 55, 70, 72, 90, 93 Isoetes Braunii 69 L. mult. ssp. occidentalis 34 I. echinospora 13, 69, 94 L. pallescens 12, 34, 94, ll9, 120 I. lacustris 16, 97 L. parviflora 13, 55, 86, 87, 99, 141, 142, 156, 163, 171 Juncus alpinus v. alpestris 13, 46, 47, 50, 66, 102, 140 L. pilosa 10, 97, 121 J. arcticus 10, 46, 56, 86, 87, 93, 100, 131, 141, 142, 152, L. spicata 10, 48, 54, 56, 67, 68, 169, 170, 171 157, 160, 163, 169, 170 L. sudetica 12, 55, 81 .T. articulatus 7 5 L. Wahlenbergii 12, 54, 73, 74, 89, 141, 142, 143, 153, J. balticus 1:J, 64 156, 157, 158 J. biglumis ll, 46, 48, 53, 5o, 87, 91, 139, 156 Lychnis Flos-cuculi 83 J. bufonius 34, 7 5 Lycopodium alpinum 10, 50, 55, 68, 140, 171 J. bulbosus 178 L. annotinum 10, 55, 56, 57, 92 J. castaneus 88, 169 L. ann. v. alpestre 92 J. conglomeratus 178 L. ann. v. pungens 171 J. filiformis 10, 97, 141, 142 L. clavatum ssp. monostachyon 12, 57, 93, 141, 142 J. monanthos 168 L. complanatum ssp. anceps 14, 18, 94 J. squarrosus 76, 88, 162 L. Selago 10, 53, 57, 89, 92, 99, 124, 129 J. stygius 13, 94, 134 L. Sel. v. appressum 92, 171 J. trifidus 10, 48, 50, 54, 67, 68, 139, 168, 171 Lysimachia thyrsiflora 16, 19, 49, 97, 134 J. triglumis 10, 27, 55, 56, 89, 142, 143, 156, 169 Maianthemum bifolium 10, 97, 101, 136, 141 Juniperus communis coli. 58, 97 Matteuccia Struthiopteris 12, 96, 97, 104 J. comm. ssp. en-communis 101 Melampyrum pratense 12, 97, 102 J. comm. ssp. nana I 71 M. silvaticum 12, 90, 95 Knautia arvensis 59 Melandrium angustiflorum ll4, 165, 166, 173 Kobresia myosuroides ll, 30, 54, 55, 85, 87, ll5, ll8, M. apetalum 10, 54, 55, 86, 100, Ill, 136, 138, 142, 151, 138, 145, 152, 153, 156, 169 152, 153, 156, 169 K. simpliciuscula 169, 170 M. rubrum 10, 55, 81, 92, 93 Koenigia islandica 13, 27, 49, 55, 56, 86, 87, 125, 153, Melica nutans 14, 96, 141 156, 157, 160, 163 Mentha arvensis 13, 50, 94 Kyllingia brevifolia 7 5 Menyanthes trifoliata 13, 37, 57, 97, 102 Lactuca alpina 10, 27, 37, 61, 81, 120 Milium effusum 12, 91, 136 Lastrea Dryopteris 10, 97, 121 Minuartia biflora 10, 34, 48, 53, 86, 87, 156 L. Phegopteris 10, 96 M. dawsonensis 69 L. Robertiana 16, 96, 102, 141, 142, 145 M. Rossii 73 Ledum palustre ll, 20, 94, 96, 134 M. rubella 16, 31, 56, 85, 86, 87, 100, 108, ll9, 130, 134, Leontodon autumnalis 14, 46, 69, 70, 93 136, 141, 149, 151, 152, 157, 158, 163, 164, 166, 169 L. aut. v. asperior 69, 90, 93 M. stricta ll, 55, 59, 85, 87, 138, 142, 143, 152, 153, 156, L. aut. v. Taraxaci 93 169 Leuchorchis albida ll, 55, 56, 81, 142, 143, 157, 160 M. verna 82, 83 L. straminea 74, 81 Moehringia lateriflora 173 Limosella aquatica 15, 49, 97 Molinia coerulea 10, 96, 102 Linnaea borealis ll, 93, 94, 95 Moneses uniflora 12, 94 Listera cordata 14, 94, 102, 141, 142, 145 Montia rivularis 12, 34 L. ovata 15, 96, 103 Myosotis caespititia 62 Loiseleuria procumbens 10, 30, 48, 50, 55, 56, 58, 68, liS, M. laxa ssp. caespitosa 50 140, 170, 171 M. silvatica ssp. frigida 14, 61, 62, 81, 82, 120, 157 Lomatogonium rotatum 74, 165 M. stricta 16, 97, 104, 107, 120 Lotus corniculatus v. borealis 14, 93, 96, 102, 120, 121, Myricaria germanica 87, 163, 173 141, 142, 145, 175 Myriophyllum alterniflorum 12, 60, 97 Lupinus nootkaensis 124 M. spicatum 16, 60, 97 Luzula arctica 12, 55, 56, 85, 86, 100, 130, 131, 136, 141, Nardus stricta 10, 15, 70, 90, 96, 139, 178 148, 151, 152, 157, 158, 159, 163, 164, 169 N arthecium ossifragum 88 L. arcuata coli. 10, 169 Nigritella nigra 88, 163, 164 Index to Phanerogams and Pteridophytes 1.97 �uphar luteum x pumilum 14, 94 Poa abbreviata 73 N. pumilum 94 P. alpina 10, 34, 48, 53, 54, 62, 68, 169, 170 Nymphaea alba ssp. candida 13, 94 P. annua 34, 36 Ophioglossum vulgatum v. islandicum 7 5 P. arctica 12, 36, 38, 55, 56, 72, 80, 86, 89, 139, 141, 142, Orchis maculata 12, 96, 102, 136, 141 152, 156, 169 0. mac. ssp. islandica 75 P. arct. ssp. caespitans 72, 164 Oxalis Acetosella 12, 22, 96, 103, 136, 176 P. arct. ssp. depauperata 7 5 Oxyria digyna 10, 37, 48, 53, 93, 139, 169, 171 P. glauca 13, 48, 54, 56, 89, 104, 156 Oxytropis campestris 60 P. Hartzii 7 3 0. camp. ssp. sordida 173 P. herjedalica 15, 46, 53, 55, 63, 86, 130, 136, 141, 142, 0. deflexa 165 145, 148, 151, 152, 157, 160 0. lapponica 11, 56, 86, 87, 116, 118, 119, 120, 138, 142, P. irrigata 16, 36, 96, 102 143, 145, 151, 152, 153, 157, 160 P. laxa coli. 80, 168 Papaver Dahlianum 74, 164, 173 P. laxa ssp. flexuosa 5, 53, 59, 87, 89, 163 P. Laestadianum 128 P. minor 168 P. lapponicum 128, 164, !'65 P. nemoralis 13, 92 P. radicatum coll. 5, 169 P. nem. v. montana 92 P. rad. v. albiflorum 74 P. palustris 13, 50, 94, 121 P. rad. ssp. faeroense 75 P. pratensis ssp. alpigena 10, 36, 38, 54, 55, 93 P. rad. ssp. islandicum 7 5 P. remota 12, 96 P. rad. ssp. Stefansonii 75 Polemonium acutiflorum 114, 165, 166 Paris quadrifolia 14, 96 P. boreale 173 Parnassia palustris 10, 46, 55, 93 Polygala serpyllifolia 76, 79, 80, 162 P. pal. v. tenuis 93 Polygonatum verticillatum 14, 81, 102, 103, 141, 142, 145 Pedicularis flammea 10, 54, 58, 65, 75, lOO, Ill, 131, 137, Polygonum Persicaria 7 5 141, 145, 151, 152, 153, 157, 158, 164, 169, 170, 171 P. Raii ssp. norvegicum 87 P. hirsuta 10, 53, 70, 86, 99, 111, 123, 124, 125, 126, 138, P. viviparum 10, 46, 47, 53, 57, 92, 93 141, 145, 151, 152, 153, 156, 165, 166 P. viv. v. alpinum 92 P. lapponica 10, 50, 53, 54, 87, 88, 90, 91, 139, 166, 179 Polypodium vulgare 96, 104, 121, 135, 141, 144, 180 P. Oederi 163 Polystichum Lonchitis 12, 48, 67, 100, 119, 120, 138, 141, P. palustris coll. 12, 178 143, 156, 170 P. pal. ssp. borealis 63, 94, 95 Populus tremula 57, 98, 101 P. Sceptrum-carolinum 10, 93 Potamogeton alpinus 13, 60, 97, 140 P. silvatica 88, 178 P. filiformis 16, 96, 140 Petasites frigidus 11, 47, 54, 56, 60, 93, 95, 139, 156 P. gramineus 12, 94, 140 P. spurius 83 P. perfoliatus 13, 97 Phippsia algida 13, 38, 46, 48, 53, 85, 87, 111, 112, 124, P. polygonifolius l 7 8 125, 126, 131, 137, 139, 142, 153, 157, 158, 163, 173 P. praelongus 16, 60, 97 Ph. concinna 73, 163 P. pusillus 13, 96 Phleum alpinum coli. 67, 171 Potentilla anglica 76, 178 Phl. alp. v. americanum 67 P. argentea 14, 96, 104, 107, 120, 131 Phl. commutatum 10, 34, 55, 67, 140 P. Chamissonis 48, 56, 64, 7.2, 119, 120, 136, 142, 145, Phragmites communis 13, 97, 134 152, 156, 165, 166, 173 Phyllodoce coerulea 10, 48, 50, 54, 56, 57, 67, 171 P. Crantzii 11, 54, 66, 91, 156, 171 Picea Abies 94 P. erecta 14, 70, 97, 102, 141, 178 P. Ab. v. arctica 17, 63, 94, 98 P. fruticosa 62 Pinguicula alpina 10, 15, 28, 46, 55, 62, 87, 138, 142, 143, P. Hookeriana 7 2 152, 153, 156, 157, 160, 172 P. hyparctica 16, 31, 53, 86, lOO, 127, 128, 130, 131, P. villosa 11, 82, 86, 160 136, 141, 148, 151, 157, 158, 159, 162, 164, 169, 171 P. vulgaris 11, 15, 46, 55, 91, 93 P. multifida 5, 73, 83, 161, 165, 173 Pinus silvestris ssp. lapponica 94 P. nivea coil. 12, 56, 142, 172 Plantago lanceolata 7 5 P. nivea sens. str. 48, 56, 72, 86, 116, 136, 143, 152, 156, Pl. major f. pygmaea 75 163, 166, 169, 171 Platanthera bifolia 13, 96, 104, 120 P. palustris ll, 57, 97 Pl. hyperborea 74 P. pulchella 73, 74 PI. oligantha 164 P. sterilis 76, 178 198 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark Potamogeton strigosa 83 Rosa majalis 14, 50, 94, 96, 104, 120 ' Primula scandinavica 14, 63, 87, 100, 119, 120, 132, 136, R. pimpinellifolia 76 141, 142, 143, 151, 157, 160 Rubus arcticus 10, 94, 95, 96, 101, 141 Pr. stricta 10, 12, 48, 56, 57, 65, 87, 100, 115, 151, 157, R. Chamaemorus 10, 55, 73, 89, 93, 118, 149 160, 163, 164, 172 R. idaeus 10, 22, 57, 70, 72, so: 96, 102, 104, 105, 121, 145 Prunella vulgaris 14, 47, 50, 96, 121 R. id. ssp. sachalinensis 70, 72 Prunus Padus v. borealis 10, 22, 63, 98, 141 R. saxatilis 10, 56, 90, 97 Pyrola chlorantha 16, 94, 104, 120 Rumex Acetosa ssp. arifolius 10, 37, 55, 70, 90, 140 P. grandiflora 14, 54, 56, 93, 156, 169, 171, 172 R. Acetosella coil. 57, 70, 169 P. media 105 R. Ac. ssp. angiocarpus 70 P. minor 12, 15, 55, 97 R. domesticus 69 Ramischia secunda 14, 94, 141, 142 R. tenuifolius 14, 34, 70, 90, 102, 120, 141, 169 Ranunculus acris 10, 34, 37, 54, 70, 90, 93, 112 Sagina caespitosa 16, 53, 56, 64, 74, 86, 99, 100, 108, 116, R. acris v. frigidus 70 125, 126, 127, 128, 130, 137, 141, 151, 152, 157, 158, R. acris v. pumilus 70, 93 163, 164 R. acris ssp. borealis 70 S. intermedia 13, 46, 53, 64, 86, 87, 124, 125, 137, 142, R. Chamissonis 71 152, 153, 157, 158 R. confervoides 93 S. Linnaei 12, 36, 48, 54, 93, 171 R. Flammula 178 S. nodosa 16, 49, 70, 96, 104 R. glacialis 10, 37, 53, 59, 66, 71, 87, 99, 123, 124, 125, S. Normaniana 14, 97 129, 137, 142, 148, 156, 157, 160, 164, 165, 170, 17:3 S. procumbens 34 R. hederaceus 76, 178 S. Pumilio 64 R. hyperboreus 86, 87, 113, 114, 141, 142, 163, 171 Sagittaria natans 96 R. lapponicus 13, 73, 94, 96, 134 Salix arbuscula 12, 55, 56, 58, 85, 157, 164 R. nivalis 10, 37, 46, 48, 50, 53, 85, 87, 124, 156, 157, 158, S. borealis 14, 94 163 S. caprea 14, 97 R. Pallasii 73 S. coaetanea 15, 94, 101, 120, 141 R. peltatus 12, 22, 94, 95 S. cordifolia 180 R. pelt. v. septentrionalis 63 S. glandulifera 12, 22, 86, 87, 100, 136, 161, 163, 164, 166 R. platanifolius 88, 163, 164, 173 S. glauca 10, 55, 57, 93, 139, 145, 169, 171 R. polyanthemos 83 S. gl. v. appendiculata 94 R. pygmaeus 10, 50, 53, 87, 93, 124, 128, 137, 171 S. hastata 10, 46, 55, 56, 91, 161, 166, 172 R. repens 10, 34, 57, 93 S. herbacea 10, 22, 27, 37, 46, 48, 50, 53, 54, 57, 60, 66, R. rep. ssp. fistulosus 93 126, 140, 149, 150, 168, 170, 171 R. reptans 12, 46, 47, 94, 95 S. lanata 10, 47, 54, 55, 89, 156 R. sceleratus ssp. reptabundus 83 S. lapponum 10, 55, 92, 95 R. spitzbergensis 73 S. myrsinites 11, 55, 91, 92, 156, 172 R. sulphureus 164, 165 S. myrtilloides 14, 94 R. Wilai:J.deri 73 S. nigricans 12, 59, 94 Rhinanthus minor ssp. groenlandicus 34, 65, 175 S. pentandra 14, 96 Rh. minor ssp. typicus 97 S. phylicifolia 10, 90, 95, 98, 145 Rhododendron lapponicum 10, 30, 48, 54, 56, 65, 80, 86, S. polaris 13, 46, 48, 53, 54, 57, 60, 73, 85, 87, 125, 126, 99, 138, 142, 151, 152, 153, 156, 163, 164, 169, 170, 171 131, 141, 142, 150, 156, 163, 168 Ribes spicatum coli. 12 S. reticulata 10, 28, 54, 57, 60, 85, 87, 88, 156, 170 R. spic. ssp. lapponicum 22, 63, 120 S. Starkeana 14, 94 R. spic. ssp. scandicum 92, 121 S. xerophila 15, 94, 161, 164, 166 R. Uva-crispa 59 Saussurea alpina 11, 47, 48, 53, 61, 81, 173 Roegneria canina 12, 92, 96, 104, 121, 141 Saxifraga adscendens 12, 66, 91, 102, 103, 120, 141, 14·5, R. can. v. lapponica 92 151 R. Doniana v. islandica 7 5 S. aizoides 10, 27, 46,. 47, 54, 56, 68, 86, 115, 156, 169, 170, R. islandica 7 5 172 R. latiglumis 85 S. Aizoon coil. 169, 170 R. mutabilis 16, 91, 119, 1�0, 136, 141, 164, 168 S. Aiz. ssp. eu-Aizoon 163 R. scandica 14, 63, 100, 108, 118, 136, 141, 143, 151, 157, S. Aiz. ssp. Laestadii 162 160, 168 S. Aiz. ssp. neogaea 74 Rosa dumalis 76 S. cernua 10, 48, 53, 87, 93, 126, 128, 156, 173 Index to P hanerogams and Pteridop hytes 199 Saxifraga Cotyledon 11, 81, 103, 120, 136, 153, 161, 164, Stellaria longifolia 14, 56, 94, 96, 102, 104, 105, 120, 144 169, 176 St. longipes coll. 49 S. foliolosa 10, 46, 48, 53, 86, 88, 93, 124, 125, 126, 128, St. media 36 142, 145, 156, 163, 171 St. nemorum ssp. montana 10, 55, 91, 141 S. groenlandica 11, 48, 53, 88, 93, 104, 128, 156, 163, 169, Stratiotes aloides 101 170, 172 Subularia aquatica 12, 47, 94 S. hieraciifolia 88, 163, 165 Taraxacum acidotum 77 S. Hirculus 169 T. arcticum 73, 76 S. nivalis 10, 48, 53, 54, 86, 93, 120, 156, 168, 170, 172, 173 T. atroglaucum 77 S. oppositifolia 10, 48, 53, 57, 60, 86, 126, 156, 169, 170 T. brachyceras 65, 76, 77 S. opp. ssp. amphibia 62 T. calanthum 77 S. rivularis 10, 48, 53, 59, 87, 93, 124, 125, 126, 131, 142, T. craspedotum 77 156, 171 T. croceum coll. 54, 55, 65, 76 S. stellaris 10, 47, 55, 66, 115, 139 T. cyclocentrum 77 S. tenuis 11, 46, 48, 53, 93, 145, 156, 168 T. cymbifolium 76, 77 Scheuchzeria palustris 16, 94, 95, 134 T. dilutisquameum 77 Scirpus acicularis 47, 97, 102, 140 T. dovrense 174 Se. caespitosus ssp. austriacus 10, 57 T. faeroense 77 Se. Hudsonianus 10, 94, 101, 141 T. glabrum 83 Se. palustris 50 T. Hypochoeris 7 7 Se. pauciflorus 16, 96, 102, 141 T. latispinulosum 77 Se. pumilus 164, 169 T. leucanthum 83 Scrophularia nodosa 80 T. naevosum 76, 77 Scutellaria galericulata 50 T. o bscurans 7 7 Sedum acre 16, 31, 70, 90, 96, 102, 104, 105, 120, 141, T. pleniflorum 77 142, 145; 147, 175 T. purpuridens 70 S. annuum 12, 66, 91, 104, 120, 136, 141, 142, 144, 145, T. repletum 77 168 T. rhodolepis 65, 76 S. atratum 168 T. simulum 76 S. Rosea 10, 48, 54, 62, 70, 72, 90, 156, 176 T. spectabile 77 S. Rosea ssp. integrifolium 70, 72 T. stictophyllum 77 S. villosum 5, 163, 164 Thalictrum alpinum 10, 47, 54, 57, 58, 85, 170, 172 Selaginella selaginoides 10, 47, 55, 56, 57, 58, 67, 68, Th. kemense 173 169, 170 Thlaspi alpestre 109 Senecio integrifolius 173 Thymus Serpyllum ssp. arcticus 173 Sesleria varia ssp. islandica 180 Th. Serp. ssp. tanaensis 173 Sibbaldia procumbens 10, 48, 54, 68, 139, 171 Tofieldia pusilla 10, 47, 54, 73, 74, 82, 89, 170, 172 Sieglingia decumbens 76, 178 Trientalis europaea 10, 55, 90, 93, 139 Silene acaulis 10, 48, 53, 67, 68, 156, 171 Tr. eur. f. rosea 93 S. maritima ssp. islandica 180 Trifolium repens 34 S. rupestris 13, 81, 102, 103, 120, 141, 144, 145 Triglochin palustre 27, 96, 136 S. tatarica 173 Trisetum spicatum 10, 48, 54, 87, 93 Sisyrinchium angustifolium 79 Tr. subalpestre 114, 164, 173 Solidago Virgaurea 10, 55, 93, 139 Trollius europaeus 10, 55, 93 S. Virg. v. alpestris 93, 169 Turritis glabra 14, 96, 104, 105, 107, 121 Sorbus Aucuparia v. glabrata 63, 98 Tussilago farfara 13, 91, 145, 148 Sparganium angustifolium 12, 66 Urtica dioeca ssp. gracilis 14, 64, 80, 120, 172, 175 Sp. hyperboreum 15, 68, 141, 171 Utricularia intermedia 14, 94 Sp. minimum 16, 19, 96, 134 U. minor 16, 70, 97 Sp. simplex 16, 96 U. vulgaris 16, 97 Sp. spc. 22 Vaccinium microcarpum 89, 94, 95, 97, 102, 141 Stachys silvatica 16, 96, 103, 119, 121, 136 V. Myrtillus 18, 47, 52, 55, 57, 58, 90, 97, 139 Stellaria calycantha 14, 55, 56, 65, 139 V. Oxycoccos coll. 57, 58 St. crassifolia 13, 94 V. uliginosum 54, 57, 58, 73, 97, 118 St. crassipes 88, 89, 127, 128, 163, 164 V. Vitis-idaea 18, 54, 56, 58, 90, 97 St. graminea 14, 34, 97, 104, 121, 136 V. V. -i. ssp. majus 180 200 SELANDER: Floristic Phytogeography of South-Western Lule Lappmark Vaccinium V.-i. ssp. minus 90, 171, 180 Viola epipsila 14, 56, 89, 95 Valeriana sambucifolia 11, 81, 102, 121, 141, 145 V. ep. ssp. repens 89 Veratrum album 164, 173 V. mirabilis 13, 96, 103, 121, 176 Veronica alpina 10, 48, 54, 55, 67, 72, 139, 168 V. montana 13, 68, 95, 120, 141, 169 V. Anagallis-aquatica 7 5 V. palustris 14, 96 V. fruticans 12, 66, 87, 120, 138, 141, 143, 151, 152, 153, V. Riviniana 5 156 V. rupestris 16, 92, 102, 120, 121, 136, 141, 142, 145, 147 V. officinalis 14, 96, 104, 105, 121 Viscaria alpina 10, 48, 56, 62, 66, 104, 112, 156, 170, 173 V. pumila 14, 67, 72, 164, 168, 171 V. alp. v. oelandica 62 V. scutellata 14, 97, 102, 140 Woodsia alpina 12, 67, 120, 141, 143, 151, 161, 169, 170, V. serpyllifolia 14, 34, 47, 49, 97 172 V. tenella 14, 48, 93, 136, 141, 161, 166 W. glabella 12, 22, 48, 56, 85, 119, 120, 138, 141, 153, 169, V. W ormskioldi 7 2 170, 172 Vicia Cracca 69 W. ilvensis 13, 85, 91, 120, 135, 144, 171 Viola biflora 10, 53, 57, 68, 69, 139, 166 STEN SELANDER: Floristic Phytogeography of South-Western Lule Lappmark (Swedish Laplancl) I. ERRATA P. 28, left column, line 37: p. 155, should be p. 150 P. 42, right 9: fig. 12, should be fig. 13 " P. 47, left 11: wheras, should be whereas " P. 74, right 40: Brandegeei , should be Brandegei " P. lOO, 41: mountin, should be mountain " P. 101, left 16: born, should be borne P. 103, right 1: changes, should be change P. 114, left 6: p. 84, should be p. 85 " P. 118, 38: l945a p. 131, 1945, should be l942a p. 131, l945a " P. 160, 17: occuring, should be occurring " P. 180, right 26: Hult.), should be Hult. SVENSKA VAXTSOCIOLOGISKA SALLSKAPETS HANDLI NGAR I. H. OsvALD, Die Vegetation des Hochmoores Komosse. 6. Tx. C. E. FRms, Die RoUe des Gesteinsgrundes bei xxn+436 s. 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