(PDF) Vascular flora of Inari Lapland. 9. Geraniaceae

REPORTS FROM THE KEVO

SUBARCTIC RESEARCH STATION

(Rep. Kevo Subarctic Res. Stat.)

Vol. 25 2019

CONTENTS

KEVO SUBARCTIC RESEARCH INSTITUTE, UNIVERSITY OF TURKU

YRJÖ MÄKINEN, MIKKO PIIRAINEN, UNTO LAINE†, JAAKKO NURMI, SAINI

HEINO and LASSE ISO-IIVARI: Vascular flora of Inari Lapland. 9.

Geraniaceae – Primulaceae..............................................................................................3-164

Reports from the Kevo Subarctic Research Station

(Rep. Kevo Subarctic Res. Stat.)

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ISBN XXX-XXX-XX-XXXX-X

Turku 2019

*** Painopaikka *** [Uniprint Suomen Yliopistopaino Oy] ***

ISSN 0453-7831

Unto Laine (1930 – 2018) in memoriam

This part of the flora is dedicated to our friend and colleague Mr. Unto Laine, Lic. Phil.,

who died on the 27

th

of January, 2018, at the age of 87. He was a keen nature lover and

enthusiast and an accurate observer and researcher of nature. He was widely interested as

well in vascular plants, mosses and lichens as in birds and butterflies.

Unto participated already in 1954 in the first botanical expedition in Inari Lapland

organized by Turun Eläin- ja Kasvitieteellinen Seura (The Zoological and Botanical

Society of Turku). His special research subject was the flora of the Kevojoki canyon.

Together with his wife Kaija he made many excursions both in Inari and Utsjoki during

more than 50 years. His studies about the vascular and moss flora in the area have

constituted an important basis for the flora of Inari Lapland.

In the picture Kaija and Unto are looking for Botrychium’s in Utsjoki village in 2007.

Vascular flora of Inari Lapland. 9.

Geraniaceae – Primulaceae

YRJÖ MÄKINEN, MIKKO PIIRAINEN, UNTO LAINE†, JAAKKO

NURMI, SAINI HEINO and LASSE ISO-IIVARI

MÄKINEN, YRJÖ¹, PIIRAINEN, MIKKO³, LAINE, UNTO¹†, NURMI, JAAKKO³,

HEINO, SAINI² and ISO-IIVARI, LASSE²: Vascular flora of Inari Lapland. 9.

Geraniaceae – Primulaceae. Rep. Kevo Subarctic Res. Stat. 25: 3-164. 2019. – Distribution

and ecology of 72 species, subspecies and established hybrids of Geraniaceae (3), Linaceae

(1), Balsaminaceae (1), Malvaceae (2), Clusiaceae (1), Violaceae (8), Tamaricaceae (1),

Elatinaceae (1), Onagraceae (11), Haloragaceae (2), Hippuridaceae (1), Cornaceae (1),

Apiaceae (11), Diapensiaceae (1), Pyrolaceae (6), Ericaceae (15), Empetraceae (2), and

Primulaceae (4) in Inari Lapland, northernmost Finland are described, with notes on their

morphology, variation, taxonomy, hybridization and dependence on culture. Kalmia

polifolia Wangenh. is presented as new to Finland, and Cicuta virosa L. var. virosa as new

to Inari Lapland. Four species, Androsace septentrionalis L., Heracleum sphondylium L. s.

str., Peucedanum palustre (L.) Moench and Viola rupestris F. W. Schmidt, which

sometimes have been reported from Inari Lapland, are here not accepted to the flora of the

province.

KEY WORDS: Aegopodium – Andromeda – Androsace – Angelica – Anthriscus –

Arctostaphylos – Calluna – Carum – Cassiope – Chaerophyllum – Cicuta – Circaea –

Cornus – Diapensia – distribution maps – Elatine – Empetrum – Epilobium – Erodium –

Finnish Lapland – floristics – Geranium – Heracleum – Hippuris – Hypericum – Impatiens

– Inari – Kalmia – Ledum – Linum – Loiseleuria – Lysimachia – Malva – Moneses –

Myricaria – Myriophyllum – Orthilia – Pastinaca – Petroselinum – Peucedanum –

Phyllodoce – Pimpinella – Primula – Pyrola – Rhododendron – Trientalis – Utsjoki –

Vaccinium – Viola.

¹ Botanical Museum, Biodiversity Unit, University of Turku, FI-20014 TURUN

YLIOPISTO, FINLAND

² Kevo Subarctic Research Institute, Biodiversity Unit, University of Turku, FI-20014

TURUN YLIOPISTO, FINLAND

³ Botany Unit, Finnish Museum of Natural History, University of Helsinki, P.O.Box 7, FI-

00014 HELSINGIN YLIOPISTO, FINLAND

Introduction

This paper is the ninth in the series

describing the vascular flora of Inari

Lapland. The study area is situated in N

Finland, at ca. 69° N and 27° E. The first

paper (Kallio et al. 1969) described the

area, investigation methods, various terms

and symbols in detail. Seven papers have

been published between 1971 and 2011

(Kallio et al. 1971, Kallio & Mäkinen 1975,

1978a, Mäkinen et al. 1982, 1998, 2005,

2011). A tentative list of all the vascular

plants is given in Mäkinen & Kallio (1979).

During 2009-2017 minor excursions

were made in the study area, adding 351

floristically studied 1×1 km² squares.

Except for the authors, contributions to the

4 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

flora have been made by several persons

(see Acknowledgements). Observations in

the study area have been made from 6486

squares altogether, comprising 28.0 % of

the total 23138 squares of Inari Lapland

(boundary squares counted as whole

squares). Of the studied squares, 4284 (66.0

%) are in Inari and 2202 (34.0 %) in

Utsjoki; 761 are (at least mainly) in the

alpine belt (177 in Inari, 584 in Utsjoki),

2265 in the birch belt (796 in Inari, 1469 in

Utsjoki), and 3460 in the coniferous zone

(3311 in Inari, 149 in Utsjoki).

Some of the squares are very incompletely

studied. There are over 600 squares with only 1-3

taxa and 1089 squares with less than 20 taxa. Of

these, 801 squares (73.6 %) are in Inari and 288 (26.4

%) in Utsjoki. Especially the common species have

often been omitted. As a result, their relative

frequencies appear to be clearly lower than they

should be. In the present paper, this especially

concerns the commonest taxa like Empetrum nigrum

subsp. hermaphroditum, Vaccinium myrtillus, V.

uliginosum and V. vitis-idaea.

When calculating the relative frequencies,

boundary squares have been counted as half squares.

On the basis of the relative frequencies, seven

frequency classes are denoted with Latin numerals: I

(0.000-0.015) very rare; II (0.016-0.062) rare; III

(0.063-0.140) rather rare; IV (0.141-0.249)

scattered; V (0.250-0.390) fairly frequent; VI (0.391-

0.562) frequent; VII (0.563-1.000) very frequent.

Significant differences have been marked with

asterisks (* – ***), non-significant differences have

been omitted.

The number of all 1×1 km² squares in which the

species has been found is given in parentheses after

the frequency group, followed by the relative

frequency. Thus, under Primula stricta, Rare (131;

0.020) means a total number of 131 squares where

the species has been found, the relative frequency

being 0.020.

In the paragraph InL ref., Kevo XX % refers to

Heikkinen & Kalliola (1990) and gives the

percentage of those 1×1 km² squares where the taxon

has been found in the Kevo Strict Nature Reserve.

InL XX % refers to Mäkinen & Kallio (1979) and

gives the percentage of those 10×10 km² squares

where the taxon has been found in Inari Lapland. In

the latter reference, 0 % indicates a value between 0

% and 0.5 %. The number XX sq. gives the number

of those 10×10 km² squares in which the taxon

occurs in Inari Lapland according to Lahti et al.

(1995). For taxa missing in the references no value is

given.

The value for FMF (Floristic Mapping of

Finland) gives the relative frequency of those 10×10

km² FMF-squares, where the species has been

reported in the total of 229 squares of Inari Lapland

(a combined total of 191 whole squares and 76 partial

borderline squares, see Kallio et al. 1971: 74).

In the paragraph Vertical distribution, the letters

a, b and c refer to the alpine belt, the birch belt and

the coniferous zone, respectively.

As in the previous papers of the flora, the

division of the families and their order follows Flora

Europaea (2, 3), and the nomenclature is according to

Hämet-Ahti et al. (1998, 2005a and 2005b). The

names of mosses, lichens and fungi are mostly

according to the references cited. The names of

localities in Inari Lapland mainly follow the

catalogue of Iso-Iivari (1977). The biogeographical

provinces in Fennoscandia are according to Flora

Nordica 1 (2000). The references to herbaria follow

Holmgren et al. (1990) and Thiers (2015). The

specimens of the former TURA have been annexed to

TUR in 2000, and are referred to as specimens of

TUR. YME refers to the private herbarium of Yrjö

Mäkinen. The documents in the Botanical Database

’Kastikka’ of the Finnish Museum of Natural History

are referred to as “Kastikka 2018” or “Kastikka doc.

xxxxx” (single records).

The geographical coordinates are according to

the Finnish National Uniform Coordinate System

(YKJ, Heikinheimo & Raatikainen 1971, 1981). The

zone number ”3” has been added to the easting grid-

coordinate to distinguish the coordinates from the

EUREF-FIN coordinate system (see Saarenmaa et al.

2008).

The author is indicated in the end of the text of

each taxon as follows: SH (Saini Heino), UL (Unto

Laine), YM (Yrjö Mäkinen), JN (Jaakko Nurmi), MP

(Mikko Piirainen). Lasse Iso-Iivari is responsible for

the distribution maps and calculating the frequencies.

GERANIACEAE

Erodium cicutarium (L.) L’Hér.

Introduced, very rare

Map 1

Distribution. A very variable species complex,

originating from the Mediterranean region (Fl. Eur.

2: 202, SKK III: 34, Hultén & Fries 1986: 1089).

Widely distributed in Europe, W Asia and North

America, but almost missing in E Asia and NW

North America (Hultén 1971b: 220, map 212).

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 5

Spread as a weed in other continents (Hultén & Fries

l.c., map 1275). Fairly common in Denmark and S

Sweden, more or less casual and scattered

northwards, almost missing in the northern parts

(Hultén 1971a: map 1190, Mossberg & Stenberg

2003: 374). In Finland a fairly rare archaeophyte in

the S part, a rare casual northwards (Hämet-Ahti et

al. 1998: 309, Lampinen & Lahti 2018).

Very rare in Troms and Finnmark, partly as a

German polemochore (Benum 1950, Hultén 1971a:

map 1190, Engelskjøn & Skifte 1995: 134, Lid & Lid

2005: 522). No records from Pechenga, very rare in

the Kola Peninsula (Fl. Murm. IV: 172) and Kittilä

Lapland (Hämet-Ahti et al. 1998: 309, Lampinen &

Lahti 2018, 1 specimen in TUR); no records from

Sompio or Enontekiö Lapland.

InL ref. Recorded as a casual alien in

InL (Hämet-Ahti et al. 1998: 309). Not

mentioned in Mäkinen & Kallio (1979).

H 1, TUR 1 spec.

Very rare (2; 0.000). Inari: I (1;

0.000), Utsjoki: I (1; 0.000). Inari: (1) Inari

– Ivalo road by Lake Ukonjärvi, 3 km S of

Kirakkaköngäs rapids, 1 specimen on a

road verge sown in the previous year

(7631:3518, 1973 J. Suominen 3555, H

476765). Utsjoki: (2) Kevo Subarctic

Research Station, E side of the main

building, 2 specimens in a newly sown

lawn (7742:3500, 2000 M. Alanen, TUR

571610). – Also collected on the Russian

side in a war-time German camp site near

the Electric Power Plant of Jäniskoski in

the former Inari Lapland area (1957 C. E.

Sonck, TUR 259175). Southern

hemerochore.

FMF 0.009.

Vertical distribution. c: I (2; 0.001).

Range ca. 90 m (Kevo) – 120 m

(Ukonjärvi). Silvine.

Ecology. The species is a casual weed

probably arrived among lawn seed.

Dependence on culture. All the

occurrences have been results of human

introduction. The occurrences in Inari

Lapland have been very short-lived. In

Jäniskoski Erodium was a polemochore,

and as the collection dates from 1957, the

species obviously persisted, at least as

seeds, for more than ten years (cf. Parnela

1985). On the Finnish side an

ephemerophytic anthropochore.

JN

Geranium pusillum L.

Introduced, very rare

Map 2

Distribution. Originating from Europe and SW Asia,

spread by man e.g. to North and South America, but

missing from large parts of Asia (Hultén 1971b: 192,

Hultén & Fries 1986: 1088 and map 1270). Most of

Europe except for the extreme north (Fl. Eur. 2: 198).

A scattered to fairly common, established

archaeophyte in the southernmost Fennoscandia, rare

and casual northwards (Hultén 1971a: map 1185,

SKK III: 28, Hämet-Ahti et al. 1998: 306, Mossberg

& Stenberg 2003: 372).

Three localities in Troms, one in Finnmark

(Sør-Varanger, Benum 1958: 282, Lid & Lid 2005:

521). Not found in Pechenga, very rare in the Kola

Peninsula (Fl. Murm. IV: 170). Missing in Sompio,

Kittilä and Enontekiö Lapland, a rare casual in

Koillismaa (Söyrinki & Saari 1980: 112).

InL ref. By the river in the garden of

the Tourist Hotel among other southern

species (Vainio 1947).

InL 0 %, 2 sq. H 2, TUR 1 spec.

Very rare (2; 0.000). Inari: I (2;

0.000). (1) Kiilopää Training Center, in the

yard among newly sown grass (7584:3519,

1968 P. Kallio, field list); (2) Ivalo, in the

abandoned garden of the Tourist Hotel

(7618:3522, 1945 K. Vainio, H 394281,

691294), and on waste ground in an unbuilt

site (7619:3522, 1966 P. Kallio, TUR

278595). The Ivalo locality is incorrectly in

the square 762:352 in Lampinen & Lahti

(2011) and earlier versions of the atlas, but

corrected in Lampinen et al. (2012)

onwards. Southern hemerochore.

FMF 0.007.

Vertical distribution. c: I (2; 0.001).

Range ca. 120 m (Ivalo) – 330 m

(Kiilopää). Silvine.

Ecology. In Inari Lapland the species

is an ephemeral casual weed. The

6 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

specimens collected by K. Vainio from

Ivalo are fruiting and with well-developed

seeds. Thus the species may have persisted

for a year or two in the locality. The

collection of P. Kallio consists of only one

non-flowering leaf-rosette.

Dependence on culture. In Kiilopää

the species had obviously arrived among

lawn seed. In Ivalo the older occurrence

was reported as polemochorous (Vainio

1947, cf. also Suominen 1979: 58, Parnela

1985). Ephemerophytic anthropochore,

partly polemochore.

JN

Geranium sylvaticum L.

Indigenous, frequent

Map 3

Distribution. European – W Asian, with four closely

related species (earlier regarded as subspecies) in

Europe (Fl. Eur. 2: 196, Aedo 2009, 2012) and close

relatives in central and E Asia and NW North

America (Hultén & Fries 1986: map 1263). Common

to very common in most of Fennoscandia except for

the coastal parts of the southernmost Norway and

Sweden (Hultén 1971a: map 1189, Roweck 1981:

297, Hämet-Ahti et al. 1998: 306, Mossberg &

Stenberg 2003: 370, Lid & Lid 2005: 518).

Common to very common in Troms and

Finnmark (Norman I(1): 272, Dahl 1934: 362,

Benum 1958: 283, Engelskjøn & Skifte 1995: 134,

Lid & Lid l.c.), 33 localities in the Rastigaissa area

(Ryvarden 1969: 33). Scattered to common in

Pechenga and the Kola Peninsula (Fellman 1831:

321, Kontuniemi 1932: 25, Söyrinki 1939a: 280, Fl.

Murm. IV: map 64, Alm et al. 1997: 40, Mäkinen

2002: 21). Common to very common in Sompio,

Kittilä and Enontekiö Lapland (Fellman 1835: 276,

Hjelt & Hult 1885: 125, Wainio 1891: 55, Hult 1898:

166, Roivainen 1923: 291, Hustich 1940c: 57,

Montell 1948, 1962: 123, Pertola 1961: 35, Piirainen

& Piirainen 1991b, Hämet-Ahti et al. 1998: 306,

Lampinen & Lahti 2018), but less frequent in the

northwesternmost alpine parts of Enontekiö (Lindén

1943: 73, Laine 1958: 83, Väre et al. 2008: 76,

Lampinen & Lahti l.c.).

InL ref. Common to very common

(Kihlman 1884: 99, Wainio 1891: 55,

Mikkola 1941: 30), particularly in the S

parts of Inari (Kujala 1964: 70). Common

in the Lemmenjoki area (Klockars &

Luther 1938, Rahkonen 1968: 18) and

along the Ivalojoki (Kujala 1962: 176) and

Vaskojoki (Rautava 1969: 30), very

common in the NE parts of Inari (Såltin

1958). Fairly common to common in W

Utsjoki (Laine et al. 1955: 129, Kallio &

Mäkinen 1957: 26, Laine 1964: 113).

Scattered but locally abundant along the

Kevojoki canyon (Laine 1970(II): 123).

Kevo 53.7 %, InL 84 %, 240 sq. H 12,

JYV 3, KUO 1, OULU 4, TUR 35, YME 8

spec.

Frequent (2688; 0.411). Inari: V

(1681; 0.388), Utsjoki: VI (1007; 0.457).

Difference***. Whole area, but fairly rare

in the basin of Lake Inari.

FMF 0.927.

Vertical distribution. a: IV (128;

0.177), b: VI (1264; 0.555), c: V (1296;

0.369). Differences***. Range 15 m

(Pulmankijärvi, 7762:3539) – ca. 500 m

(Peäldoajvi, 7675:3484, already mentioned

by Kihlman 1884: 99). The species has a

wide altitudinal amplitude, e.g. in Pite

lappmark, Sweden, from the coniferous

zone to the low alpine belt (Wistrand 1962:

123). Tr 1100 m (Engelskjøn & Skifte

1995: 134), Fnm 669 m (Norman I(1):

272), EnL 950 m (Laine 1958: 83). Silvike

in Mäkinen & Kallio (1979: 16). Clearly

commonest in the birch belt and rarest in

the alpine belt. Silvike.

Ecology. Geranium sylvaticum is one

of the most important components in many

herb-rich forest types and tall-herb meadow

communities in N Finland and adjacent

regions (cf. Söyrinki & Saari 1980: 112,

Fremstad & Normann 1982: 4). Such

communities have been described e.g. from

Swedish Lapland (Fries 1913: 98-102),

from the coniferous and birch belts of

Pechenga and Inari Lapland (Kujala 1929:

52-59, cf. Kontuniemi 1932: 9-10), from

the alpine belt of Pechenga (Kalliola 1932:

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 7

42-47, 63-64, 1939: 105-114), from the

subarctic and arctic-alpine areas of the

Rybachy Peninsula (Kalela 1939: 63-64,

222-232), from the birch belt of N

Fennoscandia (Hämet-Ahti 1963a: 89-99),

and from the Oulanka National Park,

Koillismaa (Söyrinki et al. 1977: 26-41).

Geranium-Dryopteris-Myrtillus type

(GDrMT) described by Kujala (1929: 80) is

represented by one study site in the spruce

forest of the lower W slope of Sarmitunturi

in SE Inari, and Geranium-Uliginosa type

(GUT) by three sites in S Inari, Laanila

(Kujala 1929: 86-88).

In Inari Lapland typical habitats of G.

sylvaticum are herb-rich meadows and

willow thickets along brooks and rivers and

by lake shores, and herb-rich forests both in

the coniferous zone and in the birch belt

(cf. Heikkinen & Kalliola 1989: 22). G.

sylvaticum also grows in stony talus slopes

under steep precipices as well as in rich

mires with shallow peat layer (cf. Cajander

1903: 16). Together with Filipendula

ulmaria, the species is an indicator of

wooded mires in N Finland (Kotilainen

1951: 132). In the alpine belt it usually

grows by brooks and in springy places.

G. sylvaticum is an important

constituent in several herb-rich associations

in the Kevojoki canyon (Laine 1970(I):

108). E.g. in the N part in the mouth of the

Tsarsejohka associates include

Anthoxanthum alpinum, Astragalus alpinus,

Deschampsia cespitosa, Rubus arcticus and

Viola biflora. SW of Lake Njaggaljavrrik

Geranium grows together with Cirsium

helenioides, Cornus suecica, Filipendula

ulmaria, Melica nutans, Rubus saxatilis and

Trollius europaeus. In the harsh conditions

near the southern end of the canyon

associates include e.g. Alchemilla

glomerulans, Arabis alpina, Bistorta

vivipara, Saussurea alpina and Viola

biflora.

G. sylvaticum also spreads as an

apophyte into oligohemerobic habitats, e.g.

in the margins of roads and semicultural

meadows around Lapp dwellings (Vinnamo

1963: 10, Helander 1965: 64, Vanhatalo

1965: 121, cf. Ahti & Hämet-Ahti 1971:

65, Dorogostaiskaya 1972: 123, Alm et al.

1997: 40). The main companions e.g. in the

seminatural field at Thule (Inari,

Kaamanen) were Achillea millefolium,

Carex brunnescens, Deschampsia

cespitosa, Festuca ovina, Ranunculus acris,

Solidago virgaurea and Stellaria graminea

(Helander 1965: appendix 4a).

Around the Kevo Subarctic Research

Station, flowering usually starts in the latter

half of June or beginning of July (Alanen

2007), the earliest date being June 14 in

2002 and the latest July 11 in 2000. In

unfavorable years flowering may be

delayed to the end of July. Castrén (1803)

reported that flowering in Utsjoki started as

late as on July 28 in 1795. The flowering

usually lasts for ca. one month, and first

seeds ripen usually in the end of July

(Alanen l.c.). In Pechenga flowering had

already started on July 1 in 1930 and was

profuse; fruits were partly ripe on August 5

(Kontuniemi 1932: 25). Reproduction from

seeds was fairly plentiful. Flowering is

usually common also in the alpine belt

(Söyrinki 1939a: 280, Hustich 1940c: 57).

In the slopes of the fjelds Saana and Jehkats

in Enontekiö Lapland, first flowers were

detected as early as on June 11-13 in 1986

(Uotila 1987).

Regarded as amphicline (Benum 1958:

283, Laine 1970(II): 123, Mäkinen &

Kallio 1979: 16), Ca-indifferent (Roweck

1981: 297), or as weakly or somewhat

calciphilous (Arwidsson 1943: 224, Laine

1958: 83, Wistrand 1962: 123).

Considering the preference for eutrophic

habitats, rather slightly basocline than

amphicline.

8

REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Fig. 1. Different color forms of Geranium sylvaticum L. in Utsjoki, Puksala. Photo 9.7.2003 M. Alanen.

Parasites. The rusts Puccinia leveillei

Mont. and P. morthieri Körn. have been

found a few times both in Inari and Utsjoki

(Mäkinen 1964b: 169).

Morphology and taxonomy. In S

Finland flowers of G. sylvaticum have

predominantly intensively reddish-purple or

violet-purple petals, but in N Fennoscandia

flowers with light purple to lilac or white

petals (usually with purple veins) are also

common and often prevailing (Montell

1910, 1962: 123, Auer 1938, Lundman

1948, Benum 1958: 283, Vaarama &

Jääskeläinen 1967: 27, Ahti & Hämet-Ahti

1971: 65, SKK III: 23, Engelskjøn & Skifte

1995: 134, Alm 2000, Nilsson 2000: 143;

Fig. 1). In Inari Lapland this was already

noticed by Kihlman (1884: 99): “Varietates

lilacina et parviflora multis locis, ex. gr. ad

Tenojoki, vix minus freqventer qvam forma

genuina proveniunt”. Remarks on light-

colored or white flowers have also been

made by Wainio (1891: 55) from

Törmänen, Laine et al. (1955: 129) from W

Utsjoki, Kujala (1962: 176) from the lower

Ivalojoki, Koivistoinen (1964: 49) from

Peäldoajvi, and Laine (1970(II): 31) from

the Kevojoki canyon.

In a population by the upper Repojoki

in Inari in a moist brook side grove, 184

white-flowered and 53 purple-flowered

plants were counted in 1969. On the basis

of this, the collectors suggest that the white

color may depend on one dominant allele

only (Y. Mäkinen 69-196 & L. Nurmi,

TUR 173130). On the other hand, in the

vicinity of the Kevo Subarctic Research

Station in Utsjoki, the populations have

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 9

usually approximately equal amounts of

light-colored and dark purple flowers, and

in some populations, e.g. in a herb-rich

forest by the Tsieskuljohka, light-colored

flowers are almost or totally lacking

(Alanen 2003). The genetic background of

the variation can obviously not be

explained only as a result of one dominant

allele affecting the flower color. In any

case, this variation has no greater

significance from the systematic point of

view (cf. Roweck 1981: 297). – The names

f. albiflorum Blytt and f. sublilacinum C. G.

Westerl. have sometimes been used for

plants with white or lilac flowers,

respectively.

The populations of G. sylvaticum are

usually gynodioecious: some plants have

rudimentary stamens or none at all, and are

thus functionally female. The flowers are

also considerably smaller than the

hermaphrodite ones (Vaarama &

Jääskeläinen 1967: 12, Fl. Eur. 2: 196, Fig.

2). Such pistillate, small-flowered plants

have sometimes been called var.

parviflorum Blytt. Intermediate plants with

1-9 functional anthers in a flower may also

occur (Vaarama & Jääskeläinen l.c.,

Volkova et al. 2007).

The amount of pistillate plants in

different populations varies from near zero

to ca. one quarter of the individuals

(Vaarama & Jääskeläinen 1967: 29,

Asikainen & Mutikainen 2003). The results

concerning the proportions of

hermaphrodite, intermediate and female

plants in different areas are so far

contradictory. In Finland Vaarama &

Jääskeläinen (l.c.) found no clinal

differences in the proportions from south to

north, while according to Asikainen &

Mutikainen (2003) female plants are more

frequent in northern populations. On the

contrary, in European Russia the proportion

of hermaphrodites was higher in the arctic

than in the temperate region, where

intermediates play a significant role

(Volkova et al. 2007). The proportion of

different gender morphs seems to depend at

least partly on edaphic factors (Asikainen

& Mutikainen 2003, Volkova et al. 2007).

The amount of female plants varies

also within Inari Lapland. In the study of

Vaarama & Jääskeläinen (l.c.) two

populations (158 individuals) from Inari

had 14.3 % female plants, while in three

populations (707 individuals) from Utsjoki

their amount was only 1.8 %. In the

herbarium material of H, TUR and YME,

there are 8 female plants in a total of 58

Fig. 2. Flowers of Geranium sylvaticum L. Left: a hermaphrodite flower with (at least mostly) functional

stamens. Utsjoki, Kevo, 11.7.2003. Right: a pistillate flower with shorter petals and short, rudimentary

staminodes. Utsjoki, Puksala, 26.6.2003. Photos M. Alanen.

10

REP. KEVO SUBARCTIC RES. STAT. 25. 2019

individuals from Inari Lapland.

Possible causes for the maintenance of

gynodioecy in the populations of G.

sylvaticum have been studied by Asikainen

& Mutikainen (2003, 2005a, 2005b),

Ramula & Mutikainen (2003), Ramula et

al. (2007), Varga et al. (2009) and Varga &

Kytöviita (2010a, 2010b).

A very strange form of G. sylvaticum

was collected in Utsjoki near the mouth of

the Tsarsejohka in a copse of Prunus padus

(77419:34996, 1956 U. Laine, TUR

104274, Fig. 3). It had very deeply divided

leaves with narrow, entire lobes, and

whitish, very narrow (ca. 1 mm broad)

petals. Stamens were normally developed.

Two similar plants were detected in the

locality.

Dependence on culture. The species

occurs as an apophyte around dwellings, in

pastures and seminatural meadows and

along trails and roads (see above).

Hemeradiaphore in Mäkinen & Kallio

(1979). Slightly hemerophilous.

JN

LINACEAE

Linum usitatissimum L.

Introduced, very rare

Map 4

Distribution. Cultivated oil seed and fiber plant, not

known wild; widely as a casual throughout Europe

(Fl. Eur. 2: 209).

Casual in most of Fennoscandia, except for the

northernmost parts and mountain areas (Mossberg &

Stenberg 2003: 375). In Norway scattered up to

Troms (three localities in Tromsø town); found also

in Sør-Varanger, Finnmark (Alm et al. 2004: 79, Lid

& Lid 2005: 523). Two localities in the Kirovsk area

in the Murmansk region (Fl. Murm. IV: map 63).

One record in Sompio Lapland, not known in

Kittilä or Enontekiö Lapland (Lampinen & Lahti

2018).

InL ref. Presented as new to Inari

Lapland in Lampinen & Lahti (2007).

H 1 spec.

Very rare (1; 0.000). Inari: I (1;

0.000). Ivalo, ca. 7 km NW of the village in

the communal garbage dump in Vittakuru

(7625:3517, 1999 M. Piirainen 4173, H

730375). Southern hemerochore.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Alt. ca. 190 m. Silvine.

Ecology. Collected only once as a

casual introduction. Part of the plants (12

exx. on the herbarium sheet) were

flowering at the time of collecting August

22, some had already rather well-developed

capsules but with flat undeveloped seeds.

The species is sometimes used in bird seed

mixtures, which is one possibility as the

Fig. 3. A deviating form of Geranium

sylvaticum L. with deeply divided, narrow lea

f

lobes and petals (Utsjoki, Tsarsejohka, 1956 U.

Laine, TUR 104274

)

.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 11

origin. It may also be cultivated as an

annual ornamental.

Dependence on culture. Cultivation

experiments in a small scale in Inari were

reported by Parvela (1932: 75).

Ephemerophytic anthropochore.

MP

BALSAMINACEAE

Impatiens glandulifera Royle

Introduced, very rare

Map 5

Distribution. A widely cultivated garden plant,

originating from Himalaya; rapidly and widely

naturalized in Europe (Fl. Eur. 2: 240).

A frequent and largely naturalized escape in the

southern parts of Fennoscandia, rare in the north

(Mossberg & Stenberg 2003: 385). In Sweden

throughout the country except for the northernmost

inland (Larsson & Martinsson 1998); first recorded in

Norrbotten in 1993, given already from 151 squares

of 5×5 km² by Stenberg (2010: 546). In Norway up to

Troms (Lid & Lid 2005: 533, Artsdatabanken 2015);

19 records from Tromsø, spreading (Alm et al. 2004:

79). No records from the Murmansk region.

One record in Sompio Lapland, not known from

Kittilä Lapland or Enontekiö Lapland; in Finland

locally abundant as far north as Kemijärvi, Outer

Ostrobothnia, ca. 15 km N of the Arctic Circle

(Lampinen & Lahti 2018). Hämet-Ahti et al. (1998:

310) do not give the species north of Outer

Ostrobothnia, but it is spreading rapidly, as shown by

e.g. the numerous records from Koillismaa in

Lampinen & Lahti (l.c.).

InL ref. Presented as new to Inari

Lapland in Lampinen & Lahti (2011).

Very rare (1; 0.000). Inari: I (1;

0.000). Ivalo, one plant in forest margin on

wasteland with garden refuse (7619:3522,

2006 M. Piirainen, H-Arch., Kastikka doc.

606881). Southern hemerochore.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Alt. ca. 125 m. Silvine.

Ecology. Cultivated as an annual

ornamental, recorded only once as a garden

escape. In S Finland and neighboring areas

Impatiens glandulifera is an invasive alien

in nutrient rich moist habitats but hardly

able to invade natural habitats in Inari

Lapland.

Dependence on culture. Ephemero-

phytic anthropochore.

MP

MALVACEAE

Malva pusilla Sm.

M. rotundifolia L. p. p.

Introduced, very rare

Map 6

Distribution. Originally restricted mainly to E

Europe and W Asia, introduced as a weed in central

and N Europe and North America as well as SW

Africa and New Zealand (Hultén & Fries 1986: 1091,

map 1305, Fl. Eur. 2: 251, SKK III: 77). An

established but fairly rare archaeophyte in the

southernmost Fennoscandia, nowadays less frequent

than earlier, northwards rare and casual (Hultén

1971a: map 1226, SKK III l.c., Nurmi 1987, Hämet-

Ahti et al. 1998: 222, Mossberg & Stenberg 2003:

392). Three localities in the Far North of Russia

(Dorogostaiskaya 1972: 125).

Not found in Troms, one locality in Finnmark

(Deatnu, Lid & Lid 2005: 539). No localities in

Pechenga, three in the Kola Peninsula (Fl. Murm. IV:

map 67). Missing in Sompio, Kittilä and Enontekiö

Lapland, a rare casual polemochore in Koillismaa

(Ahti & Hämet-Ahti 1971: 66) and Keret Karelia

(Herlin 1944b, Söyrinki 1956: 29, Sokolov & Filin

1996: 113).

InL ref. Ivalo in 1945 (Vainio 1947,

Mäkinen & Kallio 1979: 16). In Lahti et al.

(1995) and Lampinen & Lahti (2011) the

locality of this reference is incorrectly

placed in the square 762:352 instead of

761:352.

InL 0 %, 2 sq. H 1 spec.

Very rare (1; 0.000). Inari: I (1;

0.000). Ivalo, in the abandoned garden of

the Tourist Hotel together with other

southern species like Geranium pusillum,

Lotus corniculatus and Trifolium campestre

(7618:3522, 1945 K. Vainio, H 398437,

Vainio 1947). Southern hemerochore.

12 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Alt. ca. 120 m. Silvine.

Ecology. The species is an ephemeral

nitrophilous casual weed in Inari Lapland.

Although collected in the end of August,

the specimen is only in an early stage of

flowering, and was probably not able to

produce viable seeds.

Dependence on culture. The species

is regarded as a German polemochore

(Vainio 1947, cf. also Suominen 1979: 60).

Ephemerophytic polemochore.

JN

Malva sylvestris L.

Introduced, very rare

Not mapped

Distribution. Originally from Europe, W Asia and N

Africa, widely cultivated and introduced in large

parts of the world, also in the S hemisphere (Hultén

& Fries 1986: 1091, map 1302, Fl. Eur. 2: 250). As

an escape or casual in the southernmost

Fennoscandia, mostly along the coasts, rare inland

(Hultén 1971a: map 1227, Hämet-Ahti et al. 1998:

222, Mossberg & Stenberg 2003: 392).

One locality in Troms, none in Finnmark (Lid

& Lid 2005: 539). No records in Pechenga or the

Kola Peninsula (Fl. Murm. IV: 170), neither in

Sompio, Kittilä nor Enontekiö Lapland (Hämet-Ahti

et al. l.c., Lampinen & Lahti 2018).

InL ref. The record in Nurmi (1987)

and Hämet-Ahti et al. (1998: 222) is based

on the specimen in H (see below). Not

mentioned in Parvela (1932: 79), Mäkinen

& Kallio (1979) or Lampinen & Lahti

(2018).

H 1 spec.

Very rare (1; 0.000). Inari: I (1;

0.000). “Inari, in an oat field” (1906 T.

Itkonen, H 549663). The exact locality

missing in the specimen label (possibly

Inari village, 764:350). Southern

hemerochore.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Concluding from the habitat, probably from

the coniferous zone. Silvine.

Ecology. In Inari Lapland the species

was either an ephemeral casual weed or an

escape.

Dependence on culture. The species

mostly occurs as an escape in Finland, only

rarely as a grain immigrant (Suominen

1979: 59). However, considering the

habitat, it is possible that the species

arrived in Inari with oat seed.

Ephemerophytic anthropochore.

JN

CLUSIACEAE

Hypericum maculatum Crantz

Introduced, very rare

Map 7

Distribution. Originally Eurasiatic; mainly restricted

to Europe (Hultén 1962: map 113, Hultén & Fries

1986: map 1316). Common in S Fennoscandia,

mostly casual N of ca. 64° N (Hultén 1971a: map

1230, Roweck 1981: 305, Hämet-Ahti et al. 1998:

155, Mossberg & Stenberg 2003: 396).

Two localities in Troms (Benum 1958: 285);

very rare in Finnmark, e.g. in Sør-Varanger and

Neiden (Dahl 1934: 364, Vorren 1968 (with e.g.

Vicia sepium), Alm 1992, Piirainen 1997d, Alm &

Piirainen 2000a, Alm et al. 2000c). Rare in N

Finland, only two localities in Kittilä Lapland,

missing in Sompio Lapland and Enontekiö Lapland

(Hjelt 1911: 65, Ahti & Hämet-Ahti 1971: 66,

Hämet-Ahti et al. 1998: 155, Lampinen & Lahti

2018; H, OULU, TUR). A few localities in Pechenga

and the Kola Peninsula (Ramenskaya & Andreeva

1982: 297, H, KPABG), one locality S of

Kandalaksha (Fl. Murm. IV: 192); very rare in the

Kovda area (Sokolov & Filin 1996: 113).

InL ref. In Inari Lapland as a casual

alien found only before 1951 (Hämet-Ahti

et al. 1998: 155). However, the date is

probably erroneous, and the information

concerns the observation from near Ivalo in

1971 (see below).

InL 0 %. TUR 2, YME 1 spec.

Very rare (1; 0.000). Inari: I (1;

0.000). Ivalo, gravelly roadside between

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 13

Törmänen and Kerttuoja, a few sterile

specimens (7614:3521, 1971 Y. Mäkinen

71-724, TUR 118985, 185771, YME 5753).

Southern hemerochore.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Alt. ca. 130 m. Silvine.

Ecology. Reported as a polemochore

(Ahti & Hämet-Ahti 1971: 66, Tammilehto

1991, Ulvinen 1996, Piirainen 1997e) and

as a rye seed casual (Suominen 1979: 40).

Arrived to Ivalo probably in the connection

of road improvement works, with e.g.

Arabidopsis suecica and Fumaria

officinalis.

Dependence on culture. Ephemero-

phytic anthropochore.

YM

VIOLACEAE

Viola arvensis Murray

Introduced, very rare

Map 8

Distribution. Origin in SE Europe and adjacent parts

of Asia, spread with cultivation all over Europe and

to other continents (Hultén & Fries 1986: 1094, map

1338). Archaeophytic and common in S

Fennoscandia, in Finland to the Arctic Circle (Hultén

1971a: map 1249, Lampinen & Lahti 2018);

neophyte in the north (Fl. Nord. 6: 21).

Rare in Swedish Lapland, mainly casual and

only in man-made habitats (Roweck 1981: 315, Fl.

Nord. 6: 21). In N Norway scattered and probably

casual north to Troms (Lid & Lid 2005: 548, Fl.

Nord. 6: 21), where the species is very rare and partly

a German polemochore (Benum 1958: 287,

Engelskjøn & Skifte 1995: 137); no records from

Finnmark. Very rare in the Murmansk region (two

localities in Fl. Murm. IV: map 71, Söyrinki 1956:

23, Ramenskaya & Andreeva 1982: 299); one record

from Pechenga (Yläluostari), 1927 A. Cajander (H

400007).

Very rare in N Finland, missing from Enontekiö

Lapland, over 10 localities from both Sompio

Lapland and Kittilä Lapland (Lampinen & Lahti

2018, Kastikka 2018, TUR). One record from Kolari

in Hjelt & Hult (1885: 123); very rare and casual in

Muonio (Montell 1945a: 87, 1962: 123). Scattered in

the Kuusamo district, e.g. in fallow fields, kitchen

gardens and flower beds (Ahti & Hämet-Ahti 1971:

66).

InL ref. Sparse at Ivalo by the guest

house 1925 (Linkola 1929: 209).

InL 1 %, 2 sq. H 2, YME 1 spec.

Very rare (6; 0.001). Inari: I (5;

0.001), Utsjoki: I (1; 0.000). Inari: (1) S of

Ivalo, roadside between Törmänen and

Kerttuoja, sparsely on gravelly waste

ground (7614:3521, 1971 Y. Mäkinen,

YME 8656); (2) Ivalo, by the guest house

(7618:3522, 1925 K. Linkola, Linkola

1929); (3) Inari – Ivalo road near the S end

of Lake Ukonjärvi, one plant on a newly

sown road side lawn (7628:3518, 1973 J.

Suominen 3550, H 477717); (4) Inari

village, one small flowering plant on a

compost heap (7646:3501, 1962 E.

Tourunen, Helander 1965); (5) Kaamanen,

Toivoniemi, one poor plant in a newly built

lawn (7665:3504, 2000 M. Riikonen, H-

Arch., Kastikka doc. 352565). Utsjoki: (6)

by the main road near Sirma between

Vetsikko and Nuorgam, on a soil heap

(7771:3517, 2005 H. Väre & H.

Kaipiainen, H 807508). Southern

hemerochore.

FMF 0.020.

Vertical distribution. b: I (1; 0.000),

c: I (5; 0.001). Range ca. 50 m (Sirma) –

160 m (Toivoniemi). Silvine.

Ecology. Only as a casual introduction

on road verges and yard areas. At

Toivoniemi probably introduced with lawn

seed.

Dependence on culture. Ephemero-

phytic anthropochore.

MP

Viola biflora L.

Indigenous, scattered

Map 9

Distribution. Eurasiatic and NW North American,

arctic to subarctic-montane (Hultén 1958: 49, Hultén

14 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

& Fries 1986: 1094, map 1336). In Fennoscandia

along the whole Scandes; in central and N Norway

also in the coastal areas, in central and N Sweden

also in the W parts of the coastal provinces. In

Finland only in the four northernmost provinces

(Hultén 1971a: map 1250, Nilsson 2000: 144, Lid &

Lid 2005: 547, Fl. Nord. 6: 17).

Very common – common in Troms (Benum

1958: 287, Engelskjøn & Skifte 1995: 137).

Throughout most of Finnmark, missing from the

border areas near Kautokeino and scattered in Sør-

Varanger (Dahl 1934: 366), common in the

Rastigaissa area (Ryvarden 1969: 33). In the Kola

Peninsula common along the N and NE coastal areas

and in the Khibiny Mountains, otherwise rare (Fl.

Murm. IV: map 71, Hultén 1971a: map 1250,

Ramenskaya & Andreeva 1982: 299, Mäkinen 2002:

15). Common in Pechenga: Kammikivi area (Kalliola

1932: 106), numerous records from the Rybachy

Peninsula (Kalela 1939) and other coastal areas of

Pechenga (Kastikka 2018).

Very rare in Sompio Lapland (Pertola 1961: 35,

Rintanen 1968: 282, Ulvinen & Varkki 1998: 104)

and Kittilä Lapland (Montell 1910: 156, 1945a: 87,

1962: 123); a few dots in both provinces in Lampinen

& Lahti (2018). In Enontekiö common in the NW

part, rare or largely missing in the SE (Montell 1910,

Hustich 1940c: 57, Lindén 1943: 73, Piirainen &

Piirainen 1991b, Lampinen & Lahti l.c.).

InL ref. In Utsjoki along riversides on

sandy, slightly elevated soil (Castrén 1803).

Common in the mountains and along rivers

in Utsjoki (Fellman 1835: 281). Rather

common along the rivers Vaskojoki,

Inarijoki, Teno and Utsjoki; very common

in the subalpine belt, collected also in the

alpine belt (Kihlman 1884: 97). Rather

common to common in the valleys of the

large rivers, in the Muotkatunturit fjelds

and northwards also along smaller

tributaries, in the northern high fjelds up to

the alpine belt (Mikkola 1941: 31). Rather

rare along the upper course of the Ivalojoki

downstream at least to the Törmänen area,

missing from the lower course (Kujala

1962: 176); all the localities by the

Ivalojoki are in the coniferous zone.

Klockars & Luther (1938) give four

localities from the upper reaches and

tributaries of the Lemmenjoki, Hustich

(1942a: 225) gives two localities from the

Muotkatunturit fjelds in Inari and six along

the Utsjoki. Scattered (Laine et al. 1955:

130) or rather frequent (Kallio & Mäkinen

1957: 26) in the W parts of Utsjoki, rather

frequent in the Kevo Strict Nature Reserve

(Laine 1970(II): 124). Widely missing from

the Lake Inari basin.

Kevo 39.7 %, InL 52 %, 146 sq. H 35,

JYV 3, KUO 7, OULU 13, TUR 73, VOA

1, YME 7 spec.

Scattered (1102; 0.165). Inari: III

(310; 0.070), Utsjoki: V (792; 0.353).

Difference***. Locally rather common in

the W parts of Inari, rare in the south and

almost missing from the Lake Inari basin,

absent in the areas E of the lake. Fairly

common in Utsjoki. Northern.

FMF 0.579.

Vertical distribution. a: IV (144;

0.198), b: V (707; 0.307), c: III (251;

0.067). Differences***. Most frequent in

the subalpine region, missing from the

extensive coniferous lowlands. However,

within its area the species grows also in the

coniferous zone along rivers. Range 15 m

(Lake Pulmankijärvi, 7762:3539) – ca. 540

m (Kuovdaoaivi, 7729:3476). Tr 1269 m

(Norman I(1): 167, Engelskjøn & Skifte

1995: 137), Fnm: Rastigaissa 800 m

(Ryvarden 1969: 33), EnL 1050 m (Laine

1958: 83, Väre & Partanen 2009: 67).

Silvike.

Ecology. In the alpine belt in Finnish

Lapland Viola biflora grows in several

associations, mostly in tall-herb scrub, rich

meadow-like snow-beds and eutrophic

dwarf shrub heaths but also in

mesoeutrophic peat meadows (Kalliola

1939: 258). The species is common in

river- and brook sides, where it often

descends to the subalpine and coniferous

belts, also to lakeshores. Usually it grows in

rich meadows, herb-rich forests or scrub in

the flood zone, on sandy or stony banks, or

on humus accumulated in crevices of

riverside rocks. The species is common also

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 15

in seminatural meadows, where it often

grows together with V. epipsila, forming

sometimes even continuous colonies in the

marginal parts of the meadows towards the

birch forest.

Though V. biflora is locally rather

common, its habitats are mostly

characterized by better edaphic conditions

than in the average. The species is most

common in mesic and moist meadows on

slightly basic soil in the subalpine and

alpine belt. In the alpine belt it grows

typically in rich snow-beds, but also in

other kinds of moist meadows, with e.g.

Bistorta vivipara, Geranium sylvaticum,

Thalictrum alpinum and Trollius

europaeus. It thrives in a wide range of

habitats; Gjærevoll (1956) used it as one of

the nominal species for several snow-bed

associations, calciphilous, non-calciphilous,

hygrophilous and non-hygrophilous.

V. biflora favors the most maritime

parts of the area. It demands a humid

climate, but maximum summer

temperatures are not a decisive factor

(Rintanen 1968: 282). The species favors

also a humid microclimate and thrives well

in half-shaded places.

Flowering starts in Troms very early

after the snow has melted (Benum 1958:

287), in the alpine belt in Pechenga in the

beginning of July; by that time it is already

over in the subalpine belt (Söyrinki 1939a:

289). According to Valle (1933b) flowering

was at its best on June 21, 1929 and almost

ending on June 29, 1930. In 1880 on July

25 the flowering was already over along the

Vaskojoki in Inari, but still going on in

shady places in the subalpine belt on

Harremahtsohkka fjeld in Utsjoki on

August 6 (Kihlman 1884: 97). In the

Utsjoki valley flowering starts around mid-

June. In Pechenga, fertility and seed

production are good and it takes less than

35 days from flowering to seed ripening

(Söyrinki 1939a: 289). The significance of

the cleistogamic flowers has not been

studied in the area. Vegetative propagation

with the help of the rhizomes is

insignificant (Söyrinki 1939a: 291).

Amphicline (Laine 1970(II): 124) or

basocline (Mäkinen & Kallio 1979: 16). In

N Sweden V. biflora is slightly calciphilous

(Arwidsson 1943: 225), indifferent

(Roweck 1981: 312, Nilsson 2000: 144), or

indifferent to weakly calciphilous

(Wistrand 1962: 124); amphicline in Troms

(Benum 1958: 287). Basocline.

Parasites. The rust Puccinia alpina

Fuck. has been found several times on

Viola biflora in Utsjoki (Mäkinen 1964b:

165).

Dependence on culture. Clearly

hemerophilous and part of seminatural

meadow vegetation (Vanhatalo 1965: 121).

Hemeradiaphore.

MP

Viola canina L.

V. canina subsp. ruppii (All.) Schübl. &

Martens

V. montana auct., V. canina subsp. montana

auct.

?V. nemoralis Kütz.

Indigenous, rare

Map 10

Distribution. Large parts of Europe and Asia, also in

Greenland (perhaps an early introduction), mainly

boreal and boreonemoral (Hultén 1958: 102, Hultén

& Fries 1986: map 1328). Common in most of

Fennoscandia, scattered in the southernmost parts

and in the north and northeast (Fl. Eur. 2: 274, Hultén

1971a: map 1257, SKK III: 116–118, Hämet-Ahti at

al. 1998: 160, Nilsson 2000: 146, Lid & Lid 2005:

551, Fl. Nord. 6: 38, Krok & Almquist 2013: 316).

Rather common in the lowland areas in Troms

and rare in the inland (Benum 1958: 288–289),

probably common in Finnmark but only in the

lowlands (Norman I(1): 159, II(1): 111, Dahl 1934:

365; as either V. canina coll. or V. montana coll.).

More or less common throughout the Kola Peninsula

(Fl. Murm. IV: map 69) and the Paatsjoki area

(Wainio 1891: 56), rare or very rare in Pechenga

16 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

fjelds (Söyrinki 1939a: 288), very rare in the Lutto

area (Roivainen 1923: 291).

Rather rare to rather common in Sompio and

Kittilä Lapland, rare in Enontekiö Lapland where

almost exclusively in the NW part (Hjelt & Hult

1885: 122, Wainio 1891: 56, Hult 1898: 167, Montell

1962: 123, Laitinen & Ohenoja 1990: 24, Lampinen

& Lahti 2018).

InL ref. Common by rapids along

rivers (Fellman 1835: 281). Scattered in

Inari Lapland (Kihlman 1884: 97, Mikkola

1941: 31). Scattered to frequent in S parts

of the province, e.g. along the Ivalojoki

(Wainio 1891: 56, Kujala 1962: 177). Four

localities on sandy ground along the

Näätämöjoki (Såltin 1958). Rare in W

Utsjoki (Kallio & Mäkinen 1957: 26).

InL 12 %, 47 sq. H 12, JYV 1, OULU

2, TUR 34, YME 9 spec.

Rare (176; 0.026). Inari: II (151;

0.035), Utsjoki: I (25; 0.008).

Difference***. The distribution is clearly

concentrated in the valleys of the Ivalojoki

and the Teno and the relatively low areas

NE of Lake Inari. Lowland.

FMF 0.170.

Vertical distribution. a: I (1; 0.001),

b: II (47; 0.018), c: II (128; 0.036).

Differences***. Most common in the

coniferous zone, but ca. 25 % of the records

have been made in the subalpine birch belt.

Range 45 m (Teno valley at Pajukoste,

7771:3519) to ca. 480 m (Raututunturit

fjelds, Hult 1898: 167). Tr 600 m (V.

canina coll., Engelskjøn & Skifte 1995:

137), Fnm 120 m (Ryvarden 1969: 33),

EnL 600 m (Väre & Partanen 2009: 68).

Silvine.

Ecology. In Inari Lapland, Viola

canina grows mainly on different kinds of

shores. Typical habitats are dryish or mesic

river banks or meadows, preferably on

sandy and gravelly soils. It thrives also on

stony and gravelly places by rapids.

Sometimes it is met in luxuriant riverside

forests or shrubberies. It grows also on

lakeshore banks or dryish shore meadows.

It may be found also in brook side

meadows, but only exceptionally in natural

meadows at a distance from watercourses.

Hult (1898: 167) reported it from Dryas

heath in the low alpine belt in the

Raututunturit fjeld area at 480 m.

Flowering plants have been found in

Utsjoki in the beginning of July (Castrén

1803, Alanen 2007). Seeds ripen in

Pechenga in mid-August, but seedlings are

rather rare. No vegetative spreading stated

(Söyrinki 1939a: 288).

Viola canina favors a warm

microclimate, as is seen in its habitat choice

in sheltered river valleys. Most habitats are

rather open. The species is not very

demanding as to the soil properties but

favors unpaludified mineral soils.

Amphicline (Benum 1958: 288, Mäkinen &

Kallio 1979: 16), possibly calciphilous in

Pite lappmark (Arwidsson 1943: 225,

Wistrand 1962: 124), partly so in

Koillismaa (Söyrinki & Saari 1980: 113).

Amphicline.

Morphology and taxonomy. Viola

canina is represented in Inari Lapland by

the northern race of the species, often

accepted at the subspecific level under the

name subsp. montana (L.) Hartm. The

northern race of V. canina has a distinct

geographical area where it differs

morphologically well from the southern

populations by its taller and more erect

habit, size and shape of stipules, shape of

leaf blades and larger flowers. Where the

races meet, they seem to form a transition

zone where it is not always possible to keep

them reliably apart. Other opinions have

been expressed e.g. by Marcussen (2007,

and in Fl. Nord. 6: 38).

Already e.g. Lindberg (1958: 106)

showed that the name V. montana had been

misused. Finally, Nikitin’s (1988) formal

lectotypification made the epithet montana

unavailable in its traditional meaning, and it

seems that now there may be no name

available at the subspecific level for

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 17

northern V. canina. Because of the

nomenclatural confusion, the name V.

montana L. has now been rejected; for the

reasons, see Danihelka et al. (2010).

Nikitin (1995) lectotypified the name

Viola canina with a specimen from

Linnaeus’ Lapland herbarium. He

suggested the name V. nemoralis Kütz. to

be used at the specific level instead of the

misused name V. montana, or to accept

only one polymorphic species V. canina s.

lat. Unaware of Nikitin’s lectotypification,

Jonsell & Jarvis (2002) suggested a

specimen from the Clifford herbarium as

the lectotype for V. canina, and stated that

the Lapland herbarium specimen belongs to

subsp. montana. According to the

nomenclatural code, Nikitin’s

lectotypification must be followed, but at

present it seems difficult to decide the exact

meaning of the name (see also Jarvis 2007:

923).

Dependence on culture. A few

records from sandy roadsides. Hemera-

diaphore.

MP

Viola epipsila Ledeb.

Indigenous, frequent

Map 11

Distribution. Almost circumpolar with at least two

subspecies: subsp. epipsila from N Europe to W

Siberia, and subsp. repens (Turcz. ex Trautv. & C. A.

Mey.) W. Becker in NE Eurasia and NW North

America (Hultén & Fries 1986: 1093, map 1334).

Common in the N parts of Fennoscandia (Hultén

1971a: map 1254, Fl. Nord. 6: 23), but e.g. in S and

central Finland scattered to very rare (Nilsson 2000:

144, Fl. Nord. 6: 24).

Common throughout N Norway (Benum 1958:

289, map 392, Lid & Lid 2005: 547, Fl. Nord. 6: 23),

in Finnmark more common in the inland (Dahl 1934:

365, Elven et al. 2013: 448), missing in the NE parts

of the Varanger Peninsula (Hultén 1971a: map 1254).

Common throughout most of the Kola Peninsula (Fl.

Murm. IV: map 70, Mäkinen 2002: 15) but less so in

the northernmost parts (Hultén l.c.); relatively rare

e.g. in meadow vegetation in the Rybachy Peninsula

(Kalela 1939).

Very common in Sompio and Kittilä Lapland

and most of Enontekiö Lapland, fairly common in the

NW parts of Enontekiö (Lindén 1943: 73, Virtanen &

Väre 1990, Piirainen & Piirainen 1991b, Lampinen &

Lahti 2018, Väre et al. 2010: 111).

InL ref. Common in Inari Lapland

(Kihlman 1884: 97, Mikkola 1941: 30), in

S parts of the province (Wainio 1891: 56),

along the Ivalojoki (Kujala 1962: 176) and

in the Viipustunturit – Maarestatunturit

fjeld area in the Lemmenjoki National Park

(Klockars & Luther 1938), common in the

Peäldoajvi fjeld area (Koivistoinen 1964:

50); in W Utsjoki common (Laine et al.

1955: 130) to rather common (Kallio &

Mäkinen 1957: 26), scattered in the Kevo

Strict Nature Reserve (Laine 1970(II):

124).

Kevo 66.7 %, InL 90 %, 249 sq. H 26,

JYV 1, KUO 4, OULU 6, TUR 45, YME 2

spec.

Frequent (3193; 0.490). Inari: VI

(1985; 0.460), Utsjoki: VI (1208; 0.551).

Difference***. The distribution is rather

even, and most negative records seem to

concentrate to the Lake Inari area and other

less fertile coniferous forests in the S and

SE parts of the province. Whole area.

FMF 0.958.

Vertical distribution. a: V (243;

0.331), b: VII (1357; 0.601), c: VI (1593;

0.454). Differences***. Most common in

the birch belt and least common in the

alpine belt. Range 15 m (Pulmankijärvi,

7762:3539) – ca. 500 m (Peäldoajvi,

7675:3484). Tr 828 m (Engelskjøn & Skifte

1995: 137), Fnm 580 m (Ryvarden 1969:

33), EnL 1000 m (Laine 1958: 83). Silvike.

Ecology. Viola epipsila grows in mesic

to wet sites especially on river-, brook- and

lakesides and flood or fen meadows; in

birch and willow thickets (often between

sedge hummocks) and luxuriant birch

forests close to shores; along springs and

spring brooks; in swamp forests; and in the

18 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

margins and on tussocks of rich fens. Along

the Kevojoki it grows in gravelly river

shores, slopes below steep precipices and

willow thickets with, e.g., Bistorta

vivipara, Pinguicula vulgaris and

Selaginella selaginoides (Laine 1970(II):

124). It is common also in mown or grazed

seminatural meadows.

The species is common up to the

uppermost part of the birch belt, ascending

up to 100 m above it (Kihlman 1884: 97).

In Troms it is seldom found above the

wood limit (Benum 1958: 289), and in

Pechenga and Kittilä Lapland it is rather

sparse in the low alpine belt (Söyrinki

1939a: 285, Hustich 1940c: 57). It is less

common in wider alpine areas, perhaps

because of lack of suitable habitats rather

than the altitude itself.

Flowering starts around mid-June in

the Kevojoki valley. In Pechenga the first

flowers open in mid-July in the alpine belt;

by that time the flowering is already almost

over in the subalpine belt. Fertility and seed

production are good in Pechenga; seeds

ripen in more favorable habitats in late

July, in snow-bed communities in late

August (Söyrinki 1939a: 285). Vegetative

propagation is possible with the help of the

rhizomes (Söyrinki 1939a: 286, Heikkinen

& Kalliola 1990: 34).

The species is not very demanding but

benefits from soils with a raised mineral

nutrient content. Amphicline (Laine

1970(II): 124, Mäkinen & Kallio 1979: 16)

or basophilous (more notably in the more

southern parts of its distribution area; Fl.

Nord. 6: 24), amphicline in Troms (Benum

1958: 289) and Pite lappmark (Arwidsson

1943: 225). Amphicline.

Parasites. The rust Puccinia

fergussonii Berk. & Br. is fairly common

on Viola epipsila in Inari Lapland

(Mäkinen 1964b: 168).

Morphology and taxonomy. Elven et

al. (2013: 450) pointed out that there are

morphological differences in the southern

and northern populations of Viola epipsila

in Norway (cf. Fl. Nord. 6: 24). Elven

(2016) even draw the conclusion that the

northern populations represent subsp.

repens, usually regarded as a NW North

American – Siberian taxon, and the

nominal subspecies would be missing in N

Norway. In lack of available published

research result V. epipsila is here treated as

one entity in Finland.

Viola epipsila and V. palustris are

often very similar and they also hybridize

(see under V. × ruprechtiana). The best

diagnostic characters for V. epipsila in the

area are somewhat triangular leaf blades

which have an acute or acuminate apex (cf.

Fl. Nord. 6: 24) and a more or less hairy

lower surface, bracteoles which are usually

located high up in the upper half of the

pedicels and relatively large flowers (the

spurred petal at least 13 mm in well-

developed plants, the spur ca. twice the

length of the sepal appendages or longer).

Often the uppermost part of the pedicels

and, more rarely, sepal appendages may be

hairy. This character does not seem to

correlate significantly with leaf hairiness,

and the indumentum of both leaves and

pedicels may also vary within populations.

Plants with V. epipsila type leaf shape,

bracteoles and flowers and with good seeds

but almost totally glabrous leaves have

been collected from Inari, Menesjärvi SW,

along the rivulet Hahpatanoja (7627:3473,

2007 H. Väre, H 812872). Similar plants

have also been recorded from Inari, S of

Nellim, SW end of Lake Kontosjärvi

(7627:3563, 2010 J. Nurmi 10-36, TUR

603669) and N end of Lake Talasjärvi

(7629:3554, 2010 J. Nurmi, Kastikka doc.

608709). Plants with white flowers have

been collected from Inari, by the

Suomujoki (1904 A. Torckell, H 410353).

Dependence on culture. Common in

home meadows and other seminatural

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 19

meadows. Hemeradiaphore.

MP

Viola palustris L.

Indigenous, rare

Map 12

Distribution. Amphi-Atlantic, concentrated in the

area from W central Europe and NW Europe to the

Ural Mountains, also in Iceland, S Greenland and NE

America; plants from Spain, France and part of the

British Isles belong to subsp. juressi (Wein) Coutinho

(Hultén 1958: 122, Hultén & Fries 1986: map 1333,

Elven 2016).

Common in the S parts of Scandinavia and

Finland, scattered in the north. In Norway common in

the coastal areas to Troms, fairly rare in Finnmark

(Benum 1958: 290, Hultén 1971a: map 1259,

Engelskjøn & Skifte 1995: 137, Nilsson 2000: 144,

Lid & Lid 2005: 547, Fl. Nord. 6: 25); in Finnmark

mainly in the coastal lowlands, especially in the east,

where recorded only up to ca. 30 m a.s.l. (Dahl 1934:

364). Scattered along the coastal areas of the Kola

Peninsula, more common along the White Sea coast;

rare in the inland (Fl. Murm. IV: map 68, Hultén

1971a: map 1259). In Pechenga along the Lutto but

not abundantly (Roivainen 1923: 291), rather

frequent to frequent along the lower course of the

Paatsjoki (Wainio 1891: 56), rare in the Pechenga

fjelds (Söyrinki 1939a: 286); 14 records in Kastikka

(2018) from Pechenga.

Mainly scattered in the N parts of Sompio and

Kittilä Lapland, more common in the coniferous zone

than in the subalpine belt; rather rare in Enontekiö

Lapland (Hjelt & Hult 1885: 122, Hult 1898: 167,

Hustich 1940c: 57, Lindén 1943: 73, Montell 1962:

123, Kujala 1964: 74, Piirainen & Piirainen 1991b,

Lampinen & Lahti 2018).

InL ref. Scattered in Inari Lapland up

to the lower limit of the alpine belt, main

distribution in the valleys of the larger

rivers (Kihlman 1884: 96, Mikkola 1941:

30). Scattered to rather frequent along the

Ivalojoki (Wainio 1891: 56, Kujala 1962:

176), rather rare in the Viipustunturit –

Maarestatunturit fjeld area in the

Lemmenjoki National Park (Klockars &

Luther 1938). Three localities in the NE

parts of Inari (Såltin 1958). Rather rare

(Kallio & Mäkinen 1957: 26) or rare (Laine

et al. 1955: 130) in W Utsjoki, very rare

(Laine 1970(II): 124) or rare (Heikkinen &

Kalliola 1990: 34) in the Kevo Strict Nature

Reserve.

Kevo 2.8 %, InL 16 %, 61 sq. H 6,

OULU 1, TUR 11, YME 3 spec.

Rare (246; 0.035). Inari: II (160;

0.037), Utsjoki: II (86; 0.031). Clearly

concentrated in the larger river valleys,

avoiding the fjeld areas between them; a

large gap also in most of the Lake Inari

basin and the lowland areas north of it –

this might, however, be due to less

intensive field mapping. Lowland.

FMF 0.359.

Vertical distribution. a: I (6; 0.008),

b: II (70; 0.025), c: II (170; 0.047).

Difference a-b**, a-c***, b-c***. Most

common in the coniferous zone, largely

missing in the alpine belt. Range 15 m

(Pulmankijärvi, 7762:3539) – ca. 500 m

(Viibusoaivi, 7617:3457). Tr 925 m

(Norman I(1): 154), Fnm 805 m (Norman

II(1): 106), EnL 1050 m (Väre & Partanen

2009: 69). Silvike.

Ecology. In Inari Lapland, Viola

palustris grows mainly along river and

brook shores in the flood zone, rarely also

on lake shores. It prefers sandy and

gravelly soils but may grow also on wet

and rather tight moss mats close to the

waterline. The vegetation is usually rather

scarce, but may be a low-grown meadow or

thin forest. Typical associates are e.g.

Galium uliginosum, Juncus trifidus and

Viola epipsila, and the hepatics Blasia

pusilla, Nardia geoscyphus and Scapania

irrigua. In Inari Lapland V. palustris is

almost totally missing from several habitats

given from neighboring areas, as wet

forests and moss grasslands (Ahti &

Hämet-Ahti 1971: 66), peatlands (Dahl

1934: 364), bogs and snow-beds (Norman

II(1): 106, Benum 1958: 290).

Flowering starts approximately in late

June in the Kevojoki valley (Alanen 2007).

20 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

In Pechenga, flowering is over by late July,

seeds are formed in early August and ripen

before the end of the growing period,

seedlings are common in suitable habitats,

and also vegetative propagation by the

rhizomes is good (Söyrinki 1939a: 287).

The species is not very demanding. In

Inari Lapland it is confined almost solely to

moist soil in the littoral zone of rivers and

brooks. Amphicline (Laine 1970(II): 124,

Mäkinen & Kallio 1979: 16). According to

Hultén (1958: 122) acidophilous, lacking

on calcareous soil; amphicline in Troms

(Benum 1958: 290) and Pite lappmark

(Wistrand 1962: 124). Amphicline.

Morphology and taxonomy.

According to Ahti & Hämet-Ahti (1971:

66), Viola palustris is very homogenous in

the Kuusamo district and less variable than

in the more southern parts of Finland; it

also remains small-grown in late season

differing in this respect from V. epipsila.

This seems to be largely true also in Inari

Lapland. Morphological variation is rather

small, probably partly due to the limited

habitat range. Plants are usually small-

grown, totally glabrous and have small

rounded leaves and small flowers.

Relatively tall-grown specimens with hairy

pedicels just below the flowers – which is

not uncommon in V. epipsila – have been

collected at Raja-Jooseppi (7598:3559,

1979 J. Nurmi, TUR 267799); the capsules

are dehisced and partly broken but no seeds

are left in the specimen. The plants seem to

represent normal V. palustris, with 3 or 4

rounded leaves and bracteoles well below

the middle of the pedicels. – For further

discussion, see under V. × ruprechtiana.

Dependence on culture. Hemera-

diaphore.

MP

Viola rupestris F. W. Schmidt subsp. rupestris

Not accepted for Inari Lapland

Not mapped

Distribution. Eurasiatic, mainly central and E

European to W Asian (Hultén & Fries 1986: map

1323). Scattered to rather common in S Scandinavia

as well as S and SE Finland, the northernmost

localities in Koillismaa (scattered) and Sompio

Lapland (one locality in Pelkosenniemi, 1878 E.

Wainio, TUR 71984) (Hultén 1971a: map 1263,

Ulvinen 1985, Hämet-Ahti et al. 1998: 158, Nilsson

2000: 145, Lid & Lid 2005: 549, Fl. Nord. 6: 28,

Lampinen & Lahti 2018). Very rare in the S and E

part of the Kola Peninsula, mainly along the coasts

(Fl. Murm. IV: map 68, Hultén l.c.).

InL ref. “Ivalo (A. Torckell in herb. A. L.

Backman; A. L. Backman comm.)” (Hjelt 1911: 24);

“rr., the only record from the Ivaloj[oki], Kultala area

(Torckell)” (Mikkola 1941: 31; translated from

Finnish). Ivalo, as a dot on distribution maps in Jalas

(1950: map 18) and Hultén (1971a: map 1263), with

no other information. Recorded from Inari Lapland in

Hiitonen (1933: 413), but not in Hämet-Ahti et al.

(1998: 158), Fl. Nord. 6 or Lampinen & Lahti (2018);

the record in Kastikka (doc. 728327) is based on the

specimen in H (see below). The record in Mäkinen &

Kallio (1979: 16) is based on Mikkola (l.c.).

H 1 spec.

Ecology. No information of the ecology in Inari

Lapland is available. The locality given on the

specimen label is in the coniferous zone (alt. ca. 130

m). The species is usually confined to fairly open,

dry habitats, preferably on sandy, gravely or rocky

substrates and it is at least slightly basophilous (SKK

III: 110, Fl. Nord. 6: 28).

Notes. The only existing herbarium specimen of

Viola rupestris from Inari Lapland (H 826165)

originates in the former herbarium HSI (annexed to H

in 2000). The sheet has been labeled with a printed

form titled “Herbarium A. L. Backman”, filled in

with Backman’s handwriting: “ad Ivalo, VII.1902.

leg. A. Torckell”. The base for the locality

information in Mikkola (1941: 31) is not known,

though it seems to be based on the same specimen.

The sheet is part of an exchange or gift lot from

Torckell to Backman, as there are also specimens of

some other species in HSI/H collected by Torckell in

1902 from Inari Lapland, and labeled by Backman in

the same way. There are no duplicates of the V.

rupestris collection in Finnish herbaria. On the

contrary, there are two collections of the species in H

by Torckell, labeled with his own handwriting: ”Ta

[South Häme], Lampis, Evois, VI.1902, A. Torckell”,

the other representing var. rupestris, the other var.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 21

glaberrima (Murb.) Hyl. As these seem to be the

only specimens of the species collected and labeled

by him in 1902, it could be possible that Backman

has made a mistake in copying the label data. The

specimen from Backman’s herbarium belongs to var.

rupestris and looks very much the same as Torckell’s

specimen from Lammi (Lampis). As the Ivalo area is

floristically rather well known and no other records

of V. rupestris are known there, the taxon is here not

accepted to the flora of the province, until further

evidence is received.

MP

Viola × ruprechtiana Borbás

V. × fennica F. Nyl.

V. epipsila × palustris

Indigenous, probably very rare

Map 13

Distribution. Probably rather widely in the common

area of the parental species (the boreal circumpolar

V. epipsila and amphi-Atlantic V. palustris), but

distribution insufficiently known. Common in

Scandinavia (Mossberg & Stenberg 2003: 399; not

mapped). Intermediates between V. epipsila and V.

palustris recorded from several provinces of S

Norway and from N Norway, but only plants from

the south have been verified as hybrids by

intermediate chromosome numbers (Lid & Lid 2005:

574, Fl. Nord. 6: 51). In Sweden in most mainland

provinces, in Finland possibly common in all

provinces (Fl. Nord. l.c., Hämet-Ahti et al. 1998:

161). No unambiguous references from the Kola

Peninsula, but may be given as V. × hyperborea (V.

epipsila × epipsiloides) in Fl. Russia (9: 305) from

the area (cf. Väre 2007: 478).

Very rare in Sompio Lapland and scattered in

Kittilä and Enontekiö Lapland (Lampinen & Lahti

2018), but most records based on field notes (hybrid

chromosome numbers recorded in Finland only from

the south, with one putative exception from Sompio

Lapland in Sorsa 1965: 6).

InL ref. An easily arising and

establishing hybrid aggregate, locally rather

common in the Kevojoki valley (Laine

1970(II): 49); but the few specimens later

redetermined as V. palustris.

1 sq. (772:349; not accepted here). H 1

spec.

Very rare (1; 0.000). Inari: I (1;

0.000). Only one record accepted here,

based on a specimen consisting of two

sheets from Ivalo, Kyrö (ca. 7620:3523,

1902 A. Torckell, H 400219, H 400220).

Recorded in the field lists from the

Kevojoki (7722:3491) in 1967 and

Toivoniemi (7663:3501) in 2000, but not

included in this treatment in the lack of

specimens. The hybrid has clearly been

neglected during the field work for the Inari

Lapland flora. 41 records in H-Arch. from

field notes made during the 2000’s for the

atlas of the vascular plants of Finland

(Lampinen & Lahti 2018), with only one

voucher specimen collected as the hybrid

(Lemmenjoki National Park, Njurgalahti,

Aivvetjávri E, thin-peated mire, 7634:3471,

2007 H. Väre 17749, H 812854), but later

redetermined as V. epipsila (det. M.

Piirainen 2011). These records are mainly

from the large river valleys (Lemmenjoki,

Vaskojoki, Inarijoki, Kevojoki). In the lack

of voucher specimens, they are not included

in this treatment. Lowland.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

The only specimen has been collected from

the coniferous zone. Alt. ca. 120 m. EnL

820 m (H 812428, determination not

checked). Silvine.

Ecology. No reliable information of

the ecology in Inari Lapland. Probably

amphicline.

Morphology and taxonomy. V. ×

ruprechtiana is usually characterized as

rather variable and largely intermediate

between the parental species in size, leaf

form and leaf indumentum, position of the

bracts, and flower size and color. It is

mainly sterile, failing to set capsules and

seeds (e.g. Sorsa 1965: 15, Brandrud &

Borgen 1986, Hämet-Ahti et al. 1998: 161,

Kuta 1991: 16, Fl. Nord. 6: 51), but it has

been suggested that long-lasting vegetative

propagation may give the plants enough

time to produce also good seeds, though

rarely (SKK III: 126). Plants on the two

22 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

herbarium sheets from Ivalo show variation

as to leaf indumentum, flower size and spur

length, and part of the plants have slightly

hairy pedicels just below the flowers

(which is rather common in V. epipsila in

Inari Lapland). Thus, the population is

better understood as a variable hybrid

swarm than a mixed collection of the two

parental species. As the hybrid has largely

been neglected in the field work, collecting

of herbarium specimens of suggested

hybrid populations would be badly needed.

As most herbarium specimens of V.

epipsila, V. palustris and V. × ruprechtiana

have been collected during flowering time,

special attention should be paid to

document possible fertility of the plants.

The correct binary name for the taxon

is probably V. × fennica (Väre 2007: 478).

Dependence on culture. Probably

hemeradiaphore.

MP

Viola tricolor L. var. tricolor

Introduced, very rare

Map 14

Distribution. Origin in Europe and adjacent areas in

N Africa and Asia, introduced in other continents

(Hultén & Fries 1986: 1094, map 1337). Native and

common in S Fennoscandia up to central Norway and

Sweden, in Finland only in the south (Hultén 1971:

1266, Fl. Nord. 6: 19).

In Swedish Lapland only in man-made habitats

and often casual (Roweck 1981: 314). In Norway

common along the coastal areas north to Troms

(Benum 1958: 291, Engelskjøn & Skifte 1995: 138,

Nilsson 2000: 144, Alm et al. 2004: 80, Lid & Lid

2005: 548, Fl. Nord. 6: 19); scattered in Finnmark,

mainly in settlements (Dahl 1934: 366); in W

Finnmark partly native and locally common

especially in Alta and Loppa (Alm et al. 1993: 535,

1994: 224). The first record from E Finnmark in 1925

(Kirkenes; Linkola 1929), 9 localities in Zizka (1985:

57), still rare today and almost solely on waste land

(Alm et al. 1993: 536, 1994: 224). Infrequent in the

Murmansk region, mostly as a weed (Ramenskaya &

Andreeva 1982: 299, Ramenskaya 1983: 118); 6

localities in Pechenga (Kontuniemi 1930, Fl. Murm.

IV: map 71, specimens at H, LE, KPAPG), first

record in 1912 by Polilov (LE).

Very rare in the neighboring provinces of N

Finland (Lampinen & Lahti 2018). One record from

Sompio Lapland (Kelujärvi, 1996 T. Ulvinen, the

floristic archives at OULU). After World War I

found in several villages and farm sites in Kittilä

Lapland (Montell 1945a, 1962: 123) and Enontekiö

Lapland (Montell 1948), but disappeared rather soon;

after World War II only seven records from Kittilä

Lapland and four records from Enontekiö Lapland

(Kastikka 2018). Very rare in the Kuusamo district,

as a weed in arable land (Kastikka 2018), but may be

an old alien (Ahti & Hämet-Ahti 1971: 66).

InL ref. Two dots in Lampinen et al.

2014 and newer versions of the atlas, based

on two specimens in TUR.

InL 2 %, 3 sq. H 1, TUR 2, YME 2

spec.

Very rare (4; 0.001). Inari: I (4;

0.001). (1) 13.5 km NE from Ivalo, ca. 20

specimens by the road, in an area planted

with Pinus sylvestris (7625:3533, 1965 Y.

Mäkinen, TUR 131104, YME 8567); (2)

SW of Inari village, at a German camp site

close to Kivioja (7644:3497, 1949 G.

Marklund, H 404132); specimen collected

from “S of the road to Solojärvi, at a World

War II German camp site” (1962 A.

Vuoristo, TUR 72198) is possibly from the

same locality; (3) Muddusjärvi, at Käck’s

house, several plants (7653:3493, 1965 P.

Siltanen, field list); (4) Sevettijärvi, E end

of Lake Luolajärvi, one plant as a weed in

Lappalainen’s yard area (7732:3577, 1962

Y. Mäkinen, YME 8566). Vuoristo’s record

from Solojärvi road cannot be located

precisely, as there were several German

camps in the area, and no exact locality is

given. The record in Lahti et al. (1995)

from Utsjoki is erroneous: there was a

coding mistake in the database, and thus the

species is known only from Inari. Southern

hemerochore.

FMF 0.015.

Vertical distribution. b: I (1; 0.000),

c: I (3; 0.001). Range 90 m (Luolajärvi) –

150 m (Muddusjärvi). Silvine.

Ecology. In the area only in man-made

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 23

habitats on disturbed soil. The species has

been able to persist for at least 20 years in

the Inari – Solojärvi area, and at least for

three years at Muddusjärvi.

Dependence on culture. Ephemero-

phytic anthropochore; partly epoikophytic

polemochore.

MP

Viola × wittrockiana Gams ex Nauenb.

& Buttler

Introduced, very rare

Map 15

Distribution. A widely cultivated garden hybrid,

parents including at least V. altaica Ker Gawl., V.

lutea Huds. and V. tricolor L. (Nauenburg & Buttler

2007).

A frequent escape in the southern parts of

Fennoscandia, rare in the north. In Sweden up to

Norrbotten in the north (recorded from 68 25 km²

squares in Stenberg 2010: 408), in Norway up to

Troms (Fl. Nord. 6: 22). Four records from Tromsø

in Alm et al. (2004: 83). In Lid & Lid (2005: 548)

also given from W Finnmark: Hasvik and E

Finnmark: Sør-Varanger; recorded by Alm et al.

(1993: 535) as V. tricolor f. maxima from Sør-

Varanger: Elvenes. No records from the Murmansk

region in Fl. Murm. IV or Ramenskaya & Andreeva

(1982).

The northernmost record in Finland outside InL

is from Kemijärvi (Lampinen & Lahti 2018).

Recorded by J. Jalas (H-Arch.) in 1980 from Kittilä

Lapland: Kittilä, Pallastunturi hotel area (no

herbarium specimen; considered to be cultivated

(Kastikka doc. 437416).

InL ref. Four notes in H-Arch., see

below.

OULU 1 spec.

Very rare (4; 0.001). Inari: I (4;

0.001). Inari: (1) Ivalo, Lintumaa, refuse

recycling station (7614:3521, 2010 J.

Särkkä, H-Arch., Kastikka doc. 609367);

(2) Ivalo, one plant on gravelly waste land

in a depot area (7619:3521, 2006 M.

Piirainen, H-Arch., Kastikka doc. 606879);

(3) Ivalo, numerous plants on garden

throw-out in a forest margin (7619:3522,

2006 M. Piirainen, H-Arch., Kastikka doc.

606881) and two plants on sandy ground in

the shrub section of the plant nursery

‘Ivalon Taimitupa’ (7619:3522, 2010 J.

Särkkä 164B/10, OULU 10003883); (4)

Angeli, in a potato field (ca. 7648:3446,

1985 J. Jalas, H-Arch.; considered to be

cultivated in Kastikka (doc. 438807).

Southern hemerochore.

FMF 0.007.

Vertical distribution. c: I (4; 0.001).

Range ca. 120 m (Ivalo) – 200 m (Angeli).

Silvine.

Ecology. Cultivated as an annual

ornamental, very rarely recorded as a casual

escape. Hardly able to survive for more

than one or two years.

Dependence on culture. Ephemero-

phytic anthropochore.

MP

TAMARICACEAE

Myricaria germanica (L.) Desv.

Indigenous, very rare

Map 16

Distribution. Boreal Eurasiatic, mainly in central

Europe and Fennoscandia, extending to the Pyrenees,

E Spain, central Italy and S Ukraine, with an

outlaying site on the lower Volga; the population in

central Asia is distinguished as M. bracteata Royle

(Hultén & Fries 1986: 1094, map 1343, Fl. Eur. 2:

294, Fl. Nord. 6: 61).

In Fennoscandia three distinct areas (Norman

I(1): 451, Hiitonen 1933: 408, Hultén 1971a: map

1239, SKK III: 91, Roweck 1981: 317, Hämet-Ahti

et al. 1998: 193, Mossberg & Stenberg 2003: 405).

The northernmost area extends from Troms to

Finnmark (Dahl 1934: 364, Rønning 1954, Benum

1958: 286, Roweck l.c., Alm 1993b, Nilsson 2000:

146, Elven et al. 2013: 261). The nearest occurrence

to Utsjoki is in Tana, at a distance of ca. 50 km (SKK

III: 91, Alm l.c., Lid & Lid 2005: 553).

Anthropochorous in S Sweden (Hultén & Fries 1986:

1094). – The reports from Enontekiö Lapland

(Suomalainen 1913, Laine 1950, Nuotio 1950, the

last two reporting the find by N. Arlin in 1949, TUR

70804) are erroneous or have remained unverified.

24 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

InL ref. The only localities of

Myricaria in Finland are in Utsjoki, Lake

Pulmankijärvi area, where it was first

discovered by O. Heikinheimo and O.

Wirkkula on the W shore of the lake in the

mouth of the Kalddasjohka in 1920 (H

399180, Anonymous 1921a, b, Koivisto

1936, Hagman 1953).

InL 1 %, 4 sq. H 8, KUO 6, OULU 7,

TUR 28, YME 2 spec.

Very rare (12; 0.002). Utsjoki: I (12;

0.006). Two separate but close areas in the

valley of Lake Pulmankijärvi: (A) the

lowermost course of the Kalddasjohka in

the middle W shore of Lake Pulmankijärvi,

S of Vappula farmstead; (B) the shores of

the Pulmankijoki S of Lake Pulmankijärvi

and two roadside occurrences close to the S

end of Lake Pulmankijärvi. A detailed list

of occurrences below, mainly according to

Rautiainen (1991, 1996, Ryttäri & Kettunen

1997: 195).

(1) 7755:3539: E shore of the

Pulmankijoki 850 m N of Moresveijohka

mouth, a few shrubs on a stony river shore;

E shore of the Pulmankijoki 750 m SSE of

the Tavrajohka mouth, ca. 200 shrubs on a

sandy riverside; at the mouth of the

Tavrajohka, hundreds of shrubs on a sandy

river shore; (2) 7756:3539: E shore of the

Pulmankijoki, thousands of shrubs, forming

a continuous stand towards the mouth of

the Luossajohka; (3) 7757:3539:

Luossajohka between the Skiihpajohka and

the Pulmankijoki, and 600 m SSW of the

Luossajohka mouth, 23 large shrubs on the

riverside sand; (4) 7758:3539: sandbanks of

the Pulmankijoki in the mouth of the

Luossajohka, and the shore of the

Pulmankijoki 800 m N of the Luossajohka,

over 100 large and several hundreds of

smaller shrubs; (5) 7758:3540: N shore of

the Luossajohka near the mouth, several

large shrubs; (6) 7759:3539: a few shrubs

1–2 km N of the Luossajohka mouth, in a

dry and moss-growing old brook canyon;

along the Pulmankijoki 30 large and several

tens of smaller shrubs; (7) 7760:3539:

shore of the Pulmankijoki 1.4 km SSE from

the S end of Lake Pulmankijärvi, one large

shrub and 30 saplings on sand-mixed

gravel; (8) 7761:3539: SW side of the

Pulmankijärvi road, by the electric power

line on a newly exposed roadside, ca. 40

bushes in two groups (inventory of

threatened plants, 2003 T. Länsman & S.

Keränen); (9) 7762:3538: ca. 1 km from the

S end of the lake, W side of the

Pulmankijärvi road, ca. 10 bushes in the

road margin (inventory of threatened

plants, 2003 T. & T. Länsman); (10)

7762:3539: ca. 600 m SE of the

Pulmankijoki mouth, E shore, two small

shrubs on a sandbank; (11) 7767:3536:

Kalddasjohka, several hundreds of shrubs

600 m SW of the road bridge; (12)

7767:3537: mouth of the Kalddasjohka,

several hundreds of shrubs in three separate

areas. Northern.

FMF 0.009.

Vertical distribution. b: I (12; 0.006).

Range 15 m (7762:3539) – 70 m

(7762:3538). Tr 230 m (Benum 1958: 286),

Fnm 53 m (Norman I(1): 451). In S

Norway, Dovre, Myricaria goes up to 800-

1000 m (Nilsson 2000: 146, Lid & Lid

2005: 553). All Finnish localities are in the

birch belt. Silvine.

Ecology. Myricaria is a pioneer

species along the river shores both in

central Europe, Norway and in Finland. It

favors low and open, periodically flooded

riversides. It is often a pioneer plant which

forms thick but often short-lived

shrubberies on the river shores. In Utsjoki it

grows on glacifluvial deposits by rivers,

mostly on gravel or sand. The meandering

of the rivers continuously creates new

habitats while the old sites are decreasing.

At the mouth of the Kalddasjohka

Myricaria also grows on hemerobic

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 25

habitats like gravel pits, roadsides and

erosion fields (Fig. 4).

The seeds of Myricaria are very light

and equipped with hair balls and are thus

easily distributed by wind (cf. Lempiäinen

1981, Rautiainen 1991). Due to the

effectively spreading and germinating

seeds, Myricaria often forms rows of

shrubs (or at least seedlings) on riversides,

but it may disappear in a few years due to

an exceptionally long and high flood. Often

it has to give way to other shrubs like Salix

lapponum or S. phylicifolia.

Mostly Myricaria grows without any

associates, but in old and dried side valleys

of small brooks it has Astragalus alpinus,

Empetrum nigrum subsp. hermaphroditum,

Festuca ovina, F. rubra, Vaccinium vitis-

idaea as companions. On open riversides

Myricaria requires gravel shores or perhaps

gravel mixed with fine sand; sand alone is

too unstable for large shrubs.

The maximum height of Myricaria

shrubs around Lake Pulmankijärvi is ca.

160 cm, and the width may exceed one

meter. The general features of the

Pulmankijoki – Lake Pulmankijärvi valley

are described by Mansikkaniemi (1964).

Amphicline/Ca-indifferent (Mäkinen &

Kallio 1979: 16, Nilsson 2000: 146).

Amphicline.

Dependence on culture. At the mouth

of the Kalddasjohka, Myricaria benefits of

sandy and gravelly roadsides, and it thrives

well in gravel pits. Similarly it behaves also

in S and central Sweden. Hemeradiaphore.

YM

Fig. 4. Myricaria germanica (L.) Desv. flowering on the gravelly shore of the Kalddasjohka, Lake Pulmanki-

j

ärvi, Utsjoki. Photo 23.6.1991 S. Heino.

26 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

ELATINACEAE

Elatine hydropiper L. s. str.

E. hydropiper subsp. gyrosperma Fr., E.

gyrosperma (Fr.) Meinsh.

E. spathulata Gorski

Indigenous, very rare

Map 17

Distribution. Mainly boreal Eurasiatic, but the total

distribution incompletely known especially in the

east (Fl. USSR XV: 265, Fl. Eur. 2: 296, SKK III: 87,

Hultén & Fries 1986: map 1345, Fl. Nord. 6: 67). In

Fennoscandia unevenly distributed, in general

avoiding calcareous districts. Very rare – rare in

Denmark and in the whole of Norway, rather

common – scattered N to Dalarna in S Sweden as

well as the coast of the Gulf of Bothnia and the

Finnish mainland N to the Arctic Circle. Elsewhere

in Fennoscandia mainly very rare or lacking

(Samuelsson 1934: 145 (map), Hultén 1971a: map

1237, Uotila 1974, SKK III: 88 (map), Roweck 1981:

317, Engelskjøn & Skifte 1995: 136, Hämet-Ahti et

al. 1998: 156, Mossberg & Stenberg 2003: 406, Lid

& Lid 2005: 554).

Sør-Varanger in E Finnmark (Lid & Lid 2005:

554, Elven et al. 2013: 183), Lake Hirvasjärvi in

Imandra Lapland in the Kola Peninsula (1901 W. M.

Axelsson & V. Borg, H, TUR; Hjelt 1911: 76,

Hiitonen 1933: 408). A few localities in Kittilä

Lapland and Sompio Lapland (Wainio 1891: 55,

Montell 1962: 123, Lampinen & Lahti 2018). No

records from Enontekiö Lapland. A few records from

Koillismaa, one in the Oulanka National Park

(Söyrinki & Saari 1980: 114, Lampinen & Lahti l.c.).

InL ref. Ivalo, Törmänen (Kujala 1962:

176), Ivalo, Alempi Akujärvi (Siltanen

1967: 159), Tirro, Kuotsamo, Junnas by the

Vaskojoki (Rautava 1964: 87, 1969: 129).

InL 2 %, 3 sq. H 1, TUR 2 spec.

Very rare (5; 0.001). Inari: I (5;

0.001). (1) Ivalo, Törmänen, an oxbone

pond constricted from the Ivalojoki

(7614:3520, Kujala 1962: 176); (2) Ivalo,

Lake Alempi Akujärvi (7621:3527, 1964 P.

Siltanen, H 504141, TUR 216646, Siltanen

1967: 158, Rintanen 1976: 144); (3)

Junnas, the lower course of the Vaskojoki

(7646:3486, 1961 E. Rautava, fig. 60 in

Rautava 1969: 129); (4) Lake Inari,

Kaikunuora, Tyllylahti bay (7647:3536,

1970 M. Varjonen & Y. Mäkinen, field

list); (5) Muddusjärvi, Toivoniemi,

Mukkavuopaja, at the mouth of the

Kaamasjoki (7665:3504, 1977 C. E. Sonck,

TUR 258119, 2000 M. Riikonen, H-Arch.,

Kastikka doc. 352567). Herbarium

specimens confirmed by P. Uotila.

Southern.

FMF 0.022.

Vertical distribution. c: I (5; 0.001).

Range ca. 120 m (Alempi Akujärvi) – 145

m (Vaskojoki by Junnas). Silvine.

Ecology. Elatine hydropiper grows in

Lake Alempi Akujärvi mainly in shallow

water at boat harbors at a depth of 10-50

cm on silty – muddy bottom together with

Alopecurus aequalis, Eleocharis acicularis,

Potamogeton berchtoldii, Ranunculus

reptans and Subularia aquatica (Siltanen

1967: 159). In the lowermost course of the

Vaskojoki by Junnas the associates include

Callitriche palustris, Equisetum fluviatile,

Ranunculus reptans and Subularia aquatica

(Rautava 1964: 87, 1969: 130). E.

hydropiper is a weak competitor which

suffers from the eutrophication of waters

and overgrowth. The most typical habitats

in Inari Lapland are in rather eutrophic

shallow bays of lakes and quiet parts of

rivers. Mesotrophic – fairly eutrophic

(Linkola 1932: 95, Samuelsson 1934: 140,

Maristo 1941: 117), basocline (Mäkinen &

Kallio 1979: 16). Slightly basocline.

Morphology and taxonomy. All the

herbarium specimens seen have horseshoe-

shaped seeds and represent thus E.

hydropiper s. str. (excluding E.

orthosperma Düben; Fl. Nord. 6: 67).

Dependence on culture. Probably

hemeradiaphore.

UL

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 27

ONAGRACEAE

Circaea alpina L.

Indigenous, very rare

Map 18

Distribution. Boreal circumpolar (Fl. USSR XV:

474, Hultén 1971b: 344 and map 86, SKK III: 184,

Hultén & Fries 1986: map 1353, Fl. Nord. 6: 88).

Rather common in the coastal area of Norway N to

Troms, widely distributed in S Sweden and locally in

S Finland. Elsewhere in Fennoscandia scattered to

very rare and absent e.g. in Enontekiö Lapland and in

the Kola Peninsula, a few localities in Keret Karelia,

Kandalaksha area and Russian Koillismaa, Alakurtti

(Fl. Murm. IV: 225, Hultén 1971a: map 1293,

Roweck 1981: 319, Ramenskaya & Andreeva 1982:

307, Nilsson 2000: 150, Mossberg & Stenberg 2003:

410, Lid & Lid 2005: 567, Fl. Nord. 6: 89).

Fairly common in Troms (Norman I(1): 443,

Benum 1958: 297, Engelskjøn & Skifte 1995: 141).

Rare – very rare in Finnmark (Dahl 1934: 369,

Ryvarden 1967, Artsdatabanken 2015). Very rare in

Sompio Lapland, but several localities in Kittilä

Lapland and in Koillismaa (Hjelt & Hult 1885: 128,

Auer 1938, Montell 1962: 124, Hiltunen 1992,

Hämet-Ahti et al. 1998: 294, Lampinen & Lahti

2018; H, OULU, TUR).

InL ref. Inari, Paatari, Pyhäjärvi,

Uutela farm-yard (Laine 1964: 114, Laine

& Nurmi 1971).

InL 0 %, 1 sq. TUR 3 spec.

Very rare (2; 0.000). Inari: I (1;

0.000), Utsjoki: I (1; 0.000). Inari: (1)

Vaskojoki area, on the E side of the

abandoned Uutela farmhouse (7646:3459,

1960 K. Karinkanta, TUR 73212, 73213).

Utsjoki: (2) N side of Sarja farmstead on

the E side of Lake Pulmankijärvi

(7767:3539, 1964 P. Siltanen, TUR

223013). Also found on the Norwegian side

in Finnmark, Deatnu, Polmak, on the NE

shore of Lake Polmakvatn (Pulmankijärvi)

at the foot of W-facing cliffs less than one

kilometer from the Finnish border (1965 P.

Kallio, TUR 131099). The species was

collected in the same or a nearby locality

already by O. Dahl in 1901 (O,

Artsdatabanken 2015, see also Dahl 1934:

369). Lowland.

FMF 0.007.

Vertical distribution. b: I (1; 0.000),

c: I (1; 0.000). Alt. ca. 50 m (Sarja) – 180

m (Uutela). Tr 490 m (Engelskjøn & Skifte

1995: 141). Silvine.

Ecology. In Uutela farm Circaea

alpina was found as a small stand in a

shallow, shady depression near the fence.

Near Sarja farmstead by Lake

Pulmankijärvi the plant was growing in a

rather luxuriant grove, and on the

Norwegian side in a shady, stony grove

under the cliffs. Basocline (Benum 1958:

297, Mäkinen & Kallio 1979: 17, Nilsson

2000: 150). Basocline.

Dependence on culture. Ahemerobe.

UL

Epilobium adenocaulon Hausskn.

Introduced, very rare

Map 19

Distribution. Originally North American; now

widely distributed by human activity and naturalized

as a weed and an anthropochore particularly in

Europe, but more detailed data lacking due to

taxonomic and nomenclatural difficulties (Fl. Eur. 2:

311, SKK III: 156, Hultén & Fries 1986: 1097, map

1371, Fl. Nord. 6: 120, Elven 2016). Common to

fairly common and locally frequent in SW Norway

and particularly in S Sweden and S Finland,

elsewhere in Fennoscandia scattered to very rare or

absent, but continuously spreading towards the north

(Piispala 1964, Suominen 1969b, Hultén 1971a:

1273, Tolmatchev 1980: 50, SKK III l.c., Hämet-Ahti

et al. 1998: 298, Mossberg & Stenberg 2003: 419).

Very rare in Troms, almost exclusively in

Tromsø town and its immediate vicinity (Reiersen &

Sortland 1991, Engelskjøn & Skifte 1995: 141 (as E.

watsonii Barbey), Sortland 1997); recorded once in

Finnmark (Kirkenes, Alm & Piirainen 2000b).

Apatity and Kirovsk in the Kola Peninsula (Fl.

Murm. IV: map 73). In Lampinen & Lahti (2018)

two records from Sompio Lapland (Pelkosenniemi

2002 A. Varkki, OULU and 2014 R. Lampinen, H-

Arch.) and one from Kittilä Lapland (Kittilä 2009 J.

Särkkä, H-Arch.); earlier specimens from Muonio

(1924 J. Montell, TURA, 1927 J. Montell, VOA) are

regarded as cultivated plants (Montell 1962: 124). No

records from Enontekiö Lapland.

28 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

InL ref. Presented as new to Inari

Lapland in Lampinen et al. (2014).

TUR 1 spec.

Very rare (1; 0.000). Utsjoki: I (1;

0.000). About ten flowering and fruiting

specimens in the center of Utsjoki village

near the bridge over the Teno (7759:3501,

2000 S. Heino & U. Laine, TUR 361486,

361487). Southern hemerochore.

FMF 0.002.

Vertical distribution. b: I (1; 0.000).

Alt. ca. 70 m. Silvine.

Ecology. As a weed with

Cardaminopsis arenosa chiefly in plantings

of Dianthus superbus, Salix lanata and

Spiraea spp. The seeds very likely arrived

with the soil or garden plants from the

south. No later records.

Morphology and taxonomy. The

taxonomy of Epilobium adenocaulon –

ciliatum -complex is by no means clear.

Elven (Lid & Lid 2005: 561, Elven 2016)

includes E. adenocaulon in E. ciliatum Raf.

subsp. ciliatum. He states (Elven 2016) that

E. ciliatum is a highly variable species,

which predominantly self-pollinates, often

establishing locally distinctive races, which

have been interpreted as different species.

According to him, these are only randomly

selected and completely interfertile

components of a complex intergrading

series of forms in North America.

However, the interpretation of Flora

Nordica 6 (: 120-122) is here followed,

separating E. adenocaulon from E. ciliatum

e.g. on the grounds of the mode of

branching, presence of turions and flower

color.

In the center of Utsjoki village one

individual (TUR 361487) with white

flowers was detected among the majority of

purplish-flowered plants of E.

adenocaulon. This specimen was originally

determined as E. ciliatum and the

determination accepted by T. Karlsson in

2009. However, concerning other

morphological characteristics, the white-

flowered plant is identical with the

purplish-flowered plants in the locality, e.g.

all plants are unbranched with reddish

brown leaves and equal flower size. Hence

the conclusion is that the specimen

represents the rare white-flowered form of

E. adenocaulon, earlier recorded e.g. in

Oulu (Fl. Nord. 6: 122).

Dependence on culture. Ephemero-

phytic anthropochore.

UL

Epilobium alsinifolium Vill.

E. origanifolium Lam.

E. alpinum auct. p.p.

Indigenous, very rare

Map 20

Distribution. European arctic-montane with one

locality in W Greenland (Hultén 1950: 46, Fl. Eur. 2:

311, Hultén & Fries 1986: map 1370). Rather

common – scattered in most of Norway, scattered in

central and N Sweden as well as north of Lake

Oulujärvi in Finland, rare – very rare in NNE

Fennoscandia (Hultén 1971a: map 1275, Kytövuori

1972: 197, SKK III: 169, Roweck 1981: 328,

Mossberg & Stenberg 2003: 418, Lid & Lid 2005:

564, Fl. Nord. 6: 113).

In Troms fairly common – scattered on the

coast, rarer inland and in Finnmark (Norman I(1):

434, Dahl 1934: 368, Benum 1958: 293, Ryvarden

1969: 33, Engelskjøn & Skifte 1995: 139). Here and

there in Pechenga and the Kola Peninsula (Fellman

1831: 310, Valle 1933a, Kalela 1939: 371, Fl. Murm.

IV: map 74). Scattered in Sompio and Kittilä Lapland

(Hjelt & Hult 1885: 128, Hult 1898: 165, Montell

1910, Hustich 1940c: 57, Kujala 1961, Montell 1962:

124). Rare in most of Enontekiö Lapland (Jalas 1949,

Harviainen et al. 1968, Piirainen 1996b, Lampinen &

Lahti 2018). Numerous localities in Koillismaa and

the adjacent Russian Karelia (Söyrinki 1956: 27,

Söyrinki & Saari 1980: 116).

InL ref. Locally frequent in cold

springs by open or fairly shaded banks of

cold-watered brooks and rivers (“Ad omnes

fere fontes frigidas copiose”, Fellman 1835:

259, as E. alpinum var. montanum).

Frequent in reg. subsylvatica by springs and

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 29

shady, marshy places close to rivers at

many places, e.g. by Muddusjärvi,

Paksusammali (= Ollila), and reaching the

lower reg. alpina in the fjeld

Karegasnjarga-Ailigas (Kihlman 1884: 102,

as E. origanifolium). Kultala and Törmänen

(Wainio 1891: 51). Rather rare in Laanila

district and by the Sotajoki, Moberginoja

(Mikkola 1941: 31). Very rare by the

Ivalojoki (Kujala 1962: 177), in the

Vaskojoki area (Laine 1964: 114) and in

the W part of Utsjoki (Laine et al. 1955:

130, Laine 1970(II): 126, Heikkinen &

Kalliola 1990: 34).

Kevo 2.8 %, InL 6 %, 24 sq. H 10,

JYV 1, KUO 2, OULU 4, TUR 22, VOA 2,

YME 8 spec.

Very rare (63; 0.009). Inari: I (41;

0.009), Utsjoki: I (22; 0.010). The

occurrences are concentrated mostly to the

W and S parts of Inari Lapland, where

suitable localities are mainly situated, but

missing in the Lake Inari basin. Lowland.

FMF 0.152.

Vertical distribution. a: I (2; 0.003),

b: I (31; 0.013), c: I (29; 0.008).

Differences a-b*, b-c*. Range ca. 120 m

(Törmänen, 7614:3520) – 380 m

(Petronellankukkulat, Jomppastenoja,

7619:3449). Tr 889 m (Norman I(1): 435,

Engelskjøn & Skifte 1995: 139), Fnm 623

m (Norman I(1): 436), EnL 740 m (H.

Väre, pers. comm.). In Fennoscandia rarely

ascending to the tree line. Silvine.

Ecology. The occurrences are almost

totally concentrated around cold springs

and along spring-fed brooklets (Kytövuori

1972: 196, Fl. Nord. 6: 113). E.

alsinifolium belongs to their typical plants

along with other vascular plants and

hygrophilous mosses. It often forms tight

stands by means of hypogeal and aquatic

stolons. On steeper slopes the spring-water

flows rapidly along subterranean channels

breaking out at the foot of fjelds, often

through a rather thick and continuous moss

cover. At the base of the NW-facing slope

of the Kuoihkkarikjorbmi rivulet

(7732:3498) in the Kevojoki valley, E.

alsinifolium occurs abundantly together

with, e.g., Arabis alpina, Chrysosplenium

tetrandrum, Cystopteris montana,

Epilobium davuricum, E. hornemannii and

Saxifraga stellaris, as well as with the

mosses Brachythecium rivulare,

Campylium stellatum, Cinclidium stygium,

Cratoneuron decipiens, Dicranella

palustris, Philonotis seriata and

Warnstorfia sarmentosa (Laine 1970(II):

126). At Pystykurkkio rapids in the

Skiehttsamjohka (7627:3424), the

associates include such exacting vascular

plants as Chrysosplenium tetrandrum,

Luzula parviflora, Milium effusum and

Veronica serpyllifolia subsp. humifusa. On

the slopes of the Luovvosvarri fjeld in N

Utsjoki (7753:3530) the companions are

e.g. Athyrium filix-femina, Geum rivale and

Pinguicula alpina. In 1965, the flowering

was there still only beginning on July 19, at

the same time when E. hornemannii was

already in the fruiting stage.

E. alsinifolium is generally considered

indifferent (Benum 1958: 293, Wistrand

1962: 126, Nilsson 2000: 148), basocline –

slightly basophilous (Kytövuori 1972: 197,

Mäkinen & Kallio 1979: 17, Roweck 1981:

328) or even basocole (Laine 1970(II):

126). Slightly basocline.

Morphology and taxonomy. The

following hybrids have been collected:

E. alsinifolium × hornemannii: Inari:

(1) Tolosjoki, E of Tolospää (7601:3510,

1968 Y. Mäkinen, YME 6161); (2)

Kietsimäjoki, Stuorraboggi (7637:3427,

1965 U. Laine, TUR 133030); (3) Inari

village, Iso-Luosmajärvi (7654:3502, 1982

C. E. Sonck, TUR 269903); Utsjoki: (4)

Kaamasmukka, N of the road to

Karigasniemi, Pajuniemi (7697:3480, 1984

C. E. Sonck, TUR 279248); (5)

Karegasnjarga-Ailigas (770:346, 1961 J.

30 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Tammisola, TUR 152138). – Laine et al.

(1955: 130) mentioned a locality in the NW

side of Meäddemvarri, by a spring-water

brook of the Pullinoja rivulet (7705:3456),

but the specimen (1954 Y. Mäkinen, YME

6160) has later been determined as E.

alsinifolium.

E. alsinifolium × palustre: Inari: (1)

Raja-Jooseppi, Nohkimajärvi (7608:3556,

1972 Y. Mäkinen, YME 6158); (2) Angeli,

Kaska-Saddehvarri (7656:3465, 1960 U.

Laine, TUR 73360, 133007).

Dependence on culture. Not found in

places affected by human activity.

Ahemerobe.

UL

Epilobium anagallidifolium Lam.

E. alpinum auct. p.p.

Indigenous, rare

Map 21

Distribution. Arctic-alpine circumpolar (Hultén

1958: 240, Meusel et al. 1978: 299, Hultén & Fries

1986: map 1367, Fl. Nord. 6: 113). Mostly common

– fairly common in the Scandes and the adjacent

mountain areas up to the coast of the Arctic Ocean,

less common in NE Fennoscandia (Gjærevoll 1950:

426, Hultén 1971a: map 1276, Kytövuori 1972: 196,

SKK III: 176, Roweck 1981: 325, Hämet-Ahti et al.

1998: 301, Mossberg & Stenberg 2003: 418, Lid &

Lid 2005: 563, Fl. Nord. l.c.).

Fairly common in Troms and locally in W

Finnmark (Norman I(1): 428, Dahl 1934: 367,

Benum 1958: 294); only six localities in the

Rastigaissa area (Ryvarden 1969: 33). In Pechenga

numerous localities chiefly in the alpine belt

(Kalliola 1932: 106, 1939: 142-159, Söyrinki 1939a:

294, Alm et al. 1998: 135; see also Kalela 1939). In

the Kola Peninsula scattered in the coastal region and

the Khibiny Mountains (Fellman 1831: 310 as E.

alpinum, Hultén 1971a: map 1276, Fl. Murm. IV:

map 73).

Very rare in the Saariselkä area in Sompio

Lapland (Rintanen 1968: 283) and in the

Pallastunturi – Ounastunturi National Park in Kittilä

Lapland (Hjelt & Hult 1885: 127, Hustich 1937a:

102, 1940c: 57, Montell 1962: 124). In Enontekiö

Lapland mainly in the NW alpine region (Montell

1910, Jalas 1949, Kallio 1949, Laine 1958: 84,

Virtanen & Väre 1990, Piirainen & Piirainen 1991b,

Kämäräinen 1998, Väre et al. 2015, Lampinen &

Lahti 2018). The southernmost localities are at

Yllästunturi in Kolari (KiL), in Riekkohauta by the

Kairijoki in Savukoski (SoL) and by the Auermajoki

in the Tuntsa area in Salla (Ks) (Ulvinen 1962,

Rintanen 1968: 283, SKK III: 176, Lampinen &

Lahti l.c.). The locality in S Kittilä by the Ounasjoki

(747:341, Lampinen & Lahti 2017) is a mistake.

InL ref. In the older literature

concerning Inari Lapland E.

anagallidifolium is often included in the

Linnaean collective species E. alpinum,

together with E. lactiflorum and obviously

also E. hornemannii (e.g. Fellman 1835:

259, Kihlman 1884: 102, Wainio 1891: 51;

cf. Mela & Cajander 1906: 418-419; see

below). The oldest herbarium specimen

from Inari Lapland is possibly a specimen

without a locality collected by Jacob

Fellman in 1820-1830 during his vicarage

in Utsjoki and Inari (H 819340); the next

oldest specimen is from Utsjoki, Outakoski,

from the shore of Lake Akkojavrre

(Akujärvi, 7722:3469, 1906 H. Ranckén, H

405969, 405971, TUR 381610). The first

reliable literature records of E.

anagallidifolium are obviously two

occurrences from the Lemmenjoki National

Park: Morgam-Viibus and

Soabbekeäldimoaivi (Klockars & Luther

1938). Rather rare in Inari Lapland

(Mikkola 1941: 31). Rare – very rare in W

and NW parts of Utsjoki parish (Hustich

1942a: 225, Laine et al. 1955: 130, Kallio

& Mäkinen 1957: 26, Laine 1970(II): 125,

Kuitunen 1984, Heikkinen & Kalliola

1990: 34). Very rare in the Peäldoajvi fjeld

area (Koivistoinen 1964: 51).

Kevo 15.7 %, InL 20 %, 70 sq. H 8,

JYV 1, KUO 3, OULU 2, TUR 25, YME

11 spec.

Rare (295; 0.046). Inari: I (49; 0.012),

Utsjoki: III (246; 0.115). Difference***.

The frequency is highest in W Utsjoki due

to its vast alpine areas and the occurrences

in Finnmark (Dahl 1934: 367, Ryvarden

1969: 33). Northern with a clearly oceanic

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 31

tendency.

FMF 0.302.

Vertical distribution. a: IV (125;

0.171), b: III (146; 0.067), c: I (24; 0.007).

Differences***. Most of the localities

between the upper part of the birch belt and

the lower part of the alpine belt. Range ca.

60 m (E side of Luosnjársuolo, 7763:3509)

– ca. 460 m (Soabbekeäldimoaivi,

7633:3451). In Scandinavia mostly in the

alpine belt (Björkman 1939: 61, Arwidsson

1943: 226, Gjærevoll 1956: 168, 179,

Benum 1958: 294). Tr 1200 m (Engelskjøn

& Skifte 1995: 139), Fnm 640 m (Ryvarden

1969: 33), EnL 1050 m (Laine 1958: 84,

Väre & Partanen 2009: 130). Alpike.

Ecology. In the Fennoscandian

mountains E. anagallidifolium belongs to

the typical components of the snow-bed

vegetation as a characteristic chionophilous

hygrophyte (e.g. Gjærevoll 1950: 393,

Kytövuori 1972: 196). It thrives especially

well by cold springs and melt-water brooks

and rills flowing from snow-patches lying

above. However, it often descends along

brooks and small rivers to the forest region

(“regio alpina descensa”), e.g. in the

uppermost part of the Kevojoki valley

(Laine et al. 1955: 130, Laine 1970(II):

125, Heikkinen & Kalliola 1990: 46). In

general, the species occurs in sites where

the peat layer is thin or totally lacking, and

there are open places free of competition.

Sometimes the plant may occur as almost

continuous stands along brooks on gently

sloping hillsides together with hygrophilous

bryophytes and hepatics.

E.g. in the Kevojoki valley the typical

”descensa” habitats are enriched with

electrolytes rich in calcium and

magnesium. However, the properties of the

bedrock do not seem to be important to E.

anagallidifolium in its distribution (cf.

Benum 1958: 294, Wistrand 1962: 125,

Rintanen 1968: 283). Characteristic

associates in the Kevo Strict Nature

Reserve are Cerastium cerastoides,

Epilobium hornemannii, Gnaphalium

supinum and Juncus biglumis, sometimes

also Phleum alpinum, Selaginella

selaginoides and Veronica alpina. On the

slopes of the Galldoaivi and Guorboaivi

fjelds in E Utsjoki the following

chionophilous and cryophilous species may

belong to the associates: Agrostis mertensii,

Cassiope hypnoides, Luzula wahlenbergii,

Salix herbacea and Veronica alpina, as

well as the bryophytes and hepatics

Anthelia juratzkana, Bryum

pseudotriquetrum, Conostomum

tetragonum, Diplophyllum taxifolium,

Kiaeria starkei, Pleuroclada albescens and

Pohlia wahlenbergii (cf. Kalliola 1939:

259, Gjærevoll 1956: 179, Wistrand 1962:

125). Sometimes E. anagallidifolium has

been found on the shores of oligotrophic

lakes and pools in the subalpine birch belt,

e.g. by Vuongelijärvi (= Iso-Luosmajärvi,

7658:3508) and Kuortahjärvi (7662:3517).

Considered amphicline/Ca-indifferent

(Laine 1970(II): 125, Mäkinen & Kallio

1979: 17, Nilsson 2000: 148). Amphicline.

Parasites. The species was found

infected by the rust Puccinia scandica

Johans. on the Geidnogaissa mountain in E

Finnmark, Norway, near the Finnish border

(1963 Y. Mäkinen, TUR 180841). The rust

Puccinia epilobii DC. has been reported on

E. anagallidifolium from Utsjoki; the report

from Inari probably refers to E.

hornemannii as the host (Rainio 1926: 248,

Mäkinen 1964b: 167).

Morphology and taxonomy. The

Linnaean name Epilobium alpinum has

earlier been used for one or more of four

different northern species of Epilobium,

namely E. alsinifolium, E. anagallidifolium,

E. hornemannii and E. lactiflorum. The

name is often used as a synonym of E.

anagallidifolium (cf. Hultén 1958: 240, Fl.

Murm. IV: 214, Hämet-Ahti et al. 1998:

301). However, the lectotype belongs to the

32 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

plant usually known as E. lactiflorum, and

to avoid confusion, the epithet alpinum is

therefore rejected (see Hylander 1945: 243,

Hoch et al. 1995, Fl. Nord. 6: 116).

E. anagallidifolium varies considerably

in appearance depending on the moisture

and the nutrient content of the habitat.

Furthermore, the young and tiny specimens

of the first year may cause

misidentifications especially with E.

hornemannii. – Two specimens from the

Teno valley in Utsjoki, from the mouth of

the Suohpajohka (7750:3473, 1955 Y.

Mäkinen, YME 6147) and SE of Välimaa

(7770:3517, 1969 U. Laine, TUR 175662)

obviously represent the hybrid E.

anagallidifolium × hornemannii, and the

specimens from the NW side of

Pursatsohkka (7732:3464, 1970 Y.

Mäkinen, YME 6151) and from the

Suohpajohka (7749:3474, 1990 Y. Mäkinen

90-673, YME 19069) the hybrid E.

anagallidifolium × palustre.

Dependence on culture. Ahemerobe.

UL

Epilobium angustifolium L.

Chamerion angustifolium (L.) Holub

Chamaenerion angustifolium (L.) Scop.

Indigenous, frequent

Map 22

Distribution. Boreal circumpolar; “one of the most

completely circumpolar of all plants” (Hultén 1971b:

88, Hultén & Fries 1986: map 1355). In

Fennoscandia common almost everywhere (Hultén

1971a: map 1277, Kujala 1964: 76, Roweck 1981:

320, Mossberg & Stenberg 2003: 414).

Common – very common in Troms and

Finnmark (Norman I(1): 423, Dahl 1934: 366,

Benum 1958: 293, Ryvarden 1969: 33, Engelskjøn &

Skifte 1995: 139) and everywhere in N Finland,

Lapland and the Kola Peninsula (Fellman 1831: 310,

Hjelt & Hult 1885: 127, Hult 1898: 165, Lindén

1943: 75, Kallio 1949, Fl. Murm. IV: map 77,

Piirainen & Piirainen 1991b, Piirainen 1996a, b, Alm

et al. 1997: 41, 1998: 135, Mäkinen 2002: 20,

Lampinen & Lahti 2018. Cf. also Roivainen 1923:

291, Valle 1930, 1933a, b, Montell 1945a, 1962: 124,

Söyrinki 1956: 28, Rintanen 1962, Ahti & Hämet-

Ahti 1971: 67, Söyrinki & Saari 1980: 114, Hämet-

Ahti et al. 1998: 296). Common in the Kovda area

(Sokolov & Filin 1996: 115).

InL ref. Common – very common

(Castrén 1803, Fellman 1835: 259, Kihlman

1884: 102, Wainio 1891: 50, Mikkola

1941: 31). Ivalojoki common (Kujala

1962: 177), Vaskojoki fairly common

(Laine 1964: 114), Viipustunturit –

Maarestatunturit fairly common (Klockars

& Luther 1938). W Utsjoki scattered (Laine

et al. 1955: 130) – common (Kallio &

Mäkinen 1957: 26). Kevojoki scattered

(Kalliola 1937a: 29, Laine 1970(II): 126,

Heikkinen & Kalliola 1990: 34). On 18.6 %

of the roadsides in Inari (Helander 1965:

64).

Kevo 37.2 %, InL 90 %, 251 sq. H 4,

JYV 2, OULU 2, TUR 6, YME 9 spec.

Frequent (2868; 0.437). Inari: VI

(2005; 0.465), Utsjoki: V (863; 0.381).

Difference***. A common vascular plant

and rather evenly distributed over the area.

Only locally absent in the highest fjeld

areas (Paistunturit, Jeskaddam and

Kaldoaivi fjelds in Utsjoki), and in the

open, extensive swamp areas. Whole area.

FMF 0.965.

Vertical distribution. a: IV (148;

0.204), b: VI (987; 0.428), c: VI (1733;

0.492). Differences***. Range 15 m (Lake

Pulmankijärvi, 7762:3539, shore of the

Teno at Nuorgam, 7779:3535) – ca. 600 m

(Karegasnjarga-Ailigas, Lanka,

7705:3460). The decrease in the frequency

in the alpine areas is rather due to the lack

of suitable habitats, not to the altitude itself.

Tr 1099 m (Norman I(1): 418), Fnm 636 m

(Norman I(1): 421), EnL 1000 m (Laine

1958: 84, Väre & Partanen 2009: 127).

Vertical ubiquitous.

Ecology. As to the habitat, Epilobium

angustifolium is not exacting. On the moist

gravelly or sandy shores of brooks and

rivers it grows almost everywhere, with e.g.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 33

Parnassia palustris, Potentilla palustris,

Rubus saxatilis, Solidago virgaurea and

Trientalis europaea. It avoids shade and is

never dominating or forming large uniform

stands in its natural habitats. In all vertical

belts the species clearly favors warm

southern slopes, and due to the long

rhizomes it is also able to compete

successfully on dry and stony habitats in

the alpine belt. Several authors describe

occurrences on warm cliffs and stony rock

slopes often under the nests of birds of

prey, especially on the coast and in the river

valleys, and also on unstable scree localities

(Norman I(1): 423, Roivainen 1923: 291,

Dahl 1934: 366, Kalliola 1939: 259, 286,

Hustich 1940c: 57, Lindén 1943: 75,

Benum 1958: 293, Rintanen 1962,

Wistrand 1962: 126, Laine 1964: 114,

Mäkinen 1964b: 168, Lundqvist 1968: 123,

124, Karlsson 1973: 178, Heikkinen &

Kalliola 1990: 34).

The species especially favors burnt

areas (Roivainen 1923: 291, Kujala 1964:

76), and in Utsjoki also birch forest areas

defoliated or destroyed by Epirrita

autumnata (Kallio & Lehtonen 1973).

The species is an old apophyte being a

constant species in old Lapp and Skolt

villages and fire places (Hustich 1936b,

Mikkola 1941: 31, Suominen 1975, Alm et

al. 1997: 41, Alm & Piirainen 1997a). It

forms large and continuous stands on sandy

roadsides and in the villages around houses.

It also occurs in Inari as an old

polemochore (cf. Vorren 1968).

Cleve (1898: 54), Sylvén (1906: 150),

Kujala (1926: 126) and Kontuniemi (1932:

23, 47) describe the germination of the

seeds, which usually takes place in the

spring (see also SKK III: 182). The species

generally flowers abundantly from the end

of June until the middle of August, but

there is large yearly and site-depending

variation in the flowering time (cf. Valle

1930, Hustich 1942a, Söyrinki 1956: 28). –

Amphicline/Ca-indifferent (Laine 1970(II):

126, Mäkinen & Kallio 1979: 17, Nilsson

2000: 147). Amphicline.

Parasites. The species is infected by

two rust species. Pucciniastrum epilobii

Otth (extremely common in S Finland) has

been found only in Inari, Laanila (Rainio

1926: 254, Mäkinen 1964b: 172). The rare

Puccinia gigantea Karst. (a few collections

in Utsjoki and one in Polmak, Finnmark,

Mäkinen 1964b: 168) always occurs at the

foot of steep south bluffs. It is an

interesting example of how the distribution

of the parasite is influenced by local

macroecological factors.

Morphology and taxonomy. The N

European plants belong to subsp.

angustifolium. Mosquin (1966: 169) studied

the morphological variation of the species

in N America and described a new

subspecies, subsp. circumvagum Mosquin.

It has broader leaves and abaxially

pubescent leaf midveins. Subsp.

angustifolium is tetraploid (functionally

diploid) with 2n = 36 chromosomes. Subsp.

circumvagum is octoploid (functionally

tetraploid) with 2n = 72 chromosomes; it

occurs at lower latitudes in Asia and North

America (Elven 2016). Mitotic disturbances

occur, due to accessory chromosomes

(Laane 1965: 176, 1967). Fernald (1918)

described the variation of E. angustifolium

in Greenland, where most plants belong to

var. intermedium Wormsk. (cf. also

Jørgensen et al. 1958: 88). According to

Wistrand (1962: 126) and Roweck (1981:

320), var. spectabi1e Simm. with large

flowers occurs in Swedish Lapland. Most

probably occasional large-flowered plants

do not deserve a varietal status.

There are numerous forms of flower

color. Some observations point to an

environmental (climatic) action in the

manifestation of the genes in question; e.g.

Hustich (1936a) states that there is a white-

flowered form in Kittilä, but it is not

34 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

appearing every year.

f. albiflorum Hausskn. Inari:

Menesjärvi-Pokka road (7601:3462, YME

6132), Ivalo center (7619:3522, YME

6129; 7617:3521 Kujala 1962: 177).

Utsjoki: Karigasniemi, N part (7703:3454),

Keneskoski parking place, roadside

(7737:3502).

f. roseum Lindb. fil. Inari: Menesjärvi-

Pokka road (7600:3462, TUR 187931),

Lake Inari, Ukonkivi (7650:3511).

Narrow-leaved forms, which are

considered native, have been collected in

Inari, brook side at Ahmaoja between

Nellim and Raja-Jooseppi (7630:3568,

YME 6130) and Nanguvuono,

Laitavaaranselkä (7639:3532, YME 6131),

and in Utsjoki, Mielkejohka N of Kevojärvi

(7747:3500, YME 25248). No doubt they

are more common.

Dependence on culture. The leaves

are known to produce excellent tea-like

drink (“Ivan tšai”), and the young shoots

and flowers are edible either raw or cooked

(cf. Piippo 2004: 223, Moisio et al. 2006:

55). The author does not know of any

nutrimental use of the plant in Inari

Lapland. Strongly hemerophilous.

YM

Epilobium ciliatum Raf.

E. americanum Hausskn.

E. rubescens auct. non Rydb., E.

adenocaulon Hausskn. subsp. rubescens

(Rydb.) Hiitonen, nomen illeg.

E. saximontanum auct. non Hausskn.

E. pseudorubescens A. K. Skvortsov

Introduced, very rare

Map 23

Distribution. Originally a weakly known North

American species, locally well established and partly

naturalized in N Europe (Fl. Eur. 2: 311, Hultén &

Fries 1986: map 1372, Fl. Nord. 6: 122), probably

also in Great Britain (Blamey & Grey-Wilson 1989:

264). However, the total distribution is incompletely

known even in North America due to the taxonomic

and nomenclatural confusion with some other N

American Epilobium species (Fl. Eur. l.c., Rickett

1969: 228, SKK III: 158, Voss 1985: 622, Fl. Nord.

l.c.). Locally fairly common and partly established at

least in SW Finland as well as in S Sweden and most

of Denmark. Elsewhere in Fennoscandia rare – very

rare or lacking but spreading to new areas (Piispala

1964, Hultén 1971a: map 1290, Hämet-Ahti et al.

1998: 298, Mossberg & Stenberg 2003: 419, Alm et

al. 2004: 84, Fl. Nord. l.c.).

Very rare in Troms and Finnmark (Elven &

Solstad 2000, Lid & Lid 2005: 561). Not reported

from Pechenga Lapland or the Kola Peninsula. One

record in Sompio Lapland (2013 J. Särkkä, Kastikka

doc. 730312, Lampinen et al. 2014, 2018) and Kittilä

Lapland (as a garden weed and an escape in Muonio,

Montell 1945a, 6 specimens in TUR); not found in

Enontekiö Lapland (Lampinen & Lahti 2018).

InL ref. Utsjoki (Mäkinen & Kallio

1979: 17), Inari (Lampinen et al. 2015).

InL 0 %, TUR 1, YME 1 spec.

Very rare (3; 0.000). Inari: I (1;

0.000), Utsjoki: I (2; 0.001). Inari: (1)

Ivalo, garden center “Ivalon TaimiTupa”,

as a weed on sand (7619:3522, 2010 J.

Särkkä, H-Arch., Kastikka doc. 767836).

Utsjoki: (2) At least six plants just N of

Tsieskula farmstead on the E side of the

Utsjoki (7740:3501, 1961 Y. Mäkinen,

YME 6023); (3) in the greenhouses of the

Kevo Subarctic Research Station

(7741:3500, 1976-1977 S. Heino, TUR

261701, 261703). – The dot in 775:350 in

Lampinen et al. (2014, 2015) and

Lampinen & Lahti (2016, 2017) refers to a

white-flowered E. adenocaulon (see the

section “Morphology and taxonomy” under

that species). Southern hemerochore.

FMF 0.009.

Vertical distribution. c: I (3; 0.001).

Range ca. 75 m (Tsieskula) – 120 m

(Ivalo). Silvine.

Ecology. A casual alien on disturbed

ground in a road margin in the Tsieskula

locality or a weed in greenhouses and a

garden center. The seeds probably arrived

with soil or garden plants.

Morphology and taxonomy. See the

discussion under E. adenocaulon.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 35

Dependence on culture. Ephemero-

phytic anthropochore.

UL

Epilobium collinum C. C. Gmel.

Indigenous, very rare

Map 24

Distribution. Boreal montane European endemic

(Hultén & Fries 1986: 1096, map 1360, Fl. Nord. 6:

99). Common – fairly common in most of W

Fennoscandia, decreasing to the north. In Finland

rather common in the south, scattered – rare

northwards to Lake Oulujärvi, very rare – absent in

the northernmost provinces (Hultén 1971a: map

1278; Roweck 1981: 322, Hämet-Ahti et al. 1998:

297, Mossberg & Stenberg 2003: 415, Lid & Lid

2005: 559, Fl. Nord. l.c.).

Rather common in Troms (Norman I(1): 427,

Benum 1958: 294), scattered in W Finnmark, very

rare in E Finnmark (Norman l.c.: 428, Dahl 1934:

367, Lid & Lid l.c. 559). As a garden weed in

Muonio, Kittilä Lapland (Montell 1945a, TUR

387060). No reports from Pechenga, the Kola

Peninsula, Sompio Lapland, Enontekiö Lapland or

Keret Karelia. Erroneously reported from Koillismaa

(Ahti & Hämet-Ahti 1971: 67), the correct

determination being Epilobium montanum (cf.

Hämet-Ahti et al. 1998: 297, Fl. Nord. 6: 99,

Lampinen & Lahti 2018).

InL ref. Linkkapahta (Linkepakti) by

the Kevojoki and Kenespahta (Kanespakti)

by the Utsjoki (Kallio 1961 (see below),

Laine 1965 and 1970(II): 124, Mäkinen &

Kallio 1979: 17). The oldest observation

made by Paavo Jokela & Eino Pallari in

1952.

Kevo 0.3 %, InL 1 %, 2 sq. H 1, OULU

3, TUR 4, YME 1 spec., all from

Linkkapahta cliffs.

Very rare (1; 0.000). Utsjoki: I (1;

0.000). Linkkapahta cliffs in the N part of

the Kevojoki canyon (7734:3499, possibly

also in the adjoining square 7734:3498;

1952 P. Jokela & E. Pallari, H 406274,

OULU 45662-45664; 1955 U. Laine, TUR

73496, 73498; 1958 R.-L. Piitulainen, TUR

73497; 1958 C. E. Sonck, TUR 261640). –

The presumed record in Kenespahta

precipice on the E shore of Lake Kenesjärvi

(7735:3503, Kallio 1961) is dubious. Kallio

(l.c.) includes the species in the list of

plants growing in the precipices of the

Kevojoki and Utsjoki, but does not specify

the locality unambiguously. The possible

occurrence of Kenespahta has been

searched later without result, and no

specimen exists. Southern disjunct.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Alt. ca. 150 m. Tr 350 m (Engelskjøn &

Skifte 1995: 140). Silvine.

Ecology. In the Linkkapahta localities,

Epilobium collinum grows on the W-facing

hillside here and there in small separate

stands on a few lowermost rock ledges and

on the uppermost part of the scree, where

trickling water rich in nutrients temporarily

moistens the humus-rich soil of weathered

gravel. The yearly amounts of individuals

vary considerably depending on the drought

periods of the summer. For instance, in

1958 U. Laine observed almost a hundred

flowering and fruiting plants, but in 1965

only 32 partly non-flowering specimens. At

the foot of the rock-face in a coppice

dominated by Epilobium angustifolium, E.

palustre, Poa nemoralis, Potentilla crantzii

and Solidago virgaurea, there are numerous

exacting, nitrophilous and thermophilous

plants such as Elymus mutabilis, Erysimum

strictum, Lappula deflexa, Luzula

pallescens, Melica nutans, Paris

quadrifolia, Potentilla norvegica, Stellaria

longifolia and Urtica dioica subsp.

sondenii. In crevices and rock-ledges also

such cryptogams as Encalypta

streptocarpa, Lobaria pulmonaria,

Peltigera venosa, Physconia muscigena,

Ptilium crista-castrensis, Rhodobryum

roseum, Rhytidium rugosum and Syntrichia

ruralis belong to the typical hillside species

(cf. Wistrand 1962: 125, Lundqvist 1968:

72, Karlsson 1973: 86). – Basocline (Laine

1970(II): 125, Mäkinen & Kallio 1979: 17),

36 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

indifferent or weakly calcifile (Nilsson

2000: 147). Thermophilous, basocline.

Morphology and taxonomy. The

hybrid Epilobium collinum × palustre has

been found at Linkkapahta cliffs

(7734:3499, 1958 C. E. Sonck, TUR

261642, conf. S. Snogerup 2005, Fl. Nord.

6: 127).

Dependence on culture. Ahemerobe.

UL

Epilobium davuricum Fisch. ex Hornem.

incl. E. arcticum Sam.

Indigenous, rare

Map 25

Distribution. Arctic-montane circumpolar with a

continental tendency (Hultén 1971b: map 33, Hultén

& Fries 1986: map 1366). Fairly common in the

Scandes, in the adjacent Norwegian coastal area and

in Swedish Lapland. Scattered to rather rare in

central Sweden as well as N of Lake Oulujärvi in

Finland and in the northernmost part of Fennoscandia

(Kytövuori 1969: 35, Hultén 1971a: map 1279,

Roweck 1981: 324, Hämet-Ahti et al. 1998: 299,

Mossberg & Stenberg 2003: 417, Lid & Lid 2005:

563, Fl. Nord. 6: 104).

Fairly common in Troms and most of Finnmark

(Norman I(1): 440, II(1): 273, Dahl 1934: 368,

Benum 1958: 295, Gjærevoll 1990: 62, Engelskjøn &

Skifte 1995: 140); however, only one locality in the

Rastigaissa fjeld area (Ryvarden 1969: 33). Rare –

very rare in the low-alpine and subalpine belts of

Pechenga including the Rybachy Peninsula (Wainio

1891: 51, Kalela 1939: 426, 429, 449, Söyrinki

1939a: 293, Alm et al. 1998: 135); scattered – rare in

the Kola Peninsula and the Kandalaksha area (Fl.

Murm. IV: map 76, Hultén 1971a: map 1279).

Several localities in E Saariselkä and Sompio

Lapland (Wainio 1891: 51, Hult 1898: 165,

Roivainen 1923: 292, Ulvinen 1962). Locally

frequent in Kittilä and Enontekiö Lapland (Hjelt &

Hult 1885: 127, Hustich 1940c: 57, Lindén 1943: 75,

Montell 1910, 1962: 124). Fairly common in the

Kuusamo district (Auer 1938, Söyrinki & Saari 1980:

116). The southernmost occurrences in Finland are in

Central Ostrobothnia and in North Karelia (Hämet-

Ahti et al. 1998: 299, Lampinen & Lahti 2018).

InL ref. Verkkojärvi and Harremah-

tsohkka (Kihlman 1884: 102, as E. lineare).

Koarvikodds, Louhioja rapids in the

Ivalojoki and Laanila (Mikkola 1941: 31).

Very rare in the Peäldoajvi fjeld area

(Koivistoinen 1964: 50). Seven localities in

the Lemmenjoki – Vaskojoki area

(Klockars & Luther 1938, Laine 1964:

114). Four sites in W Utsjoki (Laine et al.

1955: 130). Ten occurrences in the Kevo

Strict Nature Reserve (Laine 1970(II): 125,

Heikkinen & Kalliola 1990: 34).

Kevo 2.8 %, InL 20 %, 73 sq. H 26,

JYV 2, KUO 1, OULU 5, TUR 50, YME

26 spec.

Rare (188; 0.029). Inari: II (115;

0.027), Utsjoki: II (73; 0.033). E.

davuricum is almost absent on the shores

and islands of Lake Inari and by the

watercourses of the Ivalojoki (cf. Kujala

1962) and Kietsimäjoki – Inarijoki, as well

as in the Vätsäri and Kalddoaivi wilderness

areas. Whole area.

FMF 0.359.

Vertical distribution. a: II (15;

0.021), b: II (87; 0.039), c: II (86; 0.024).

Differences a-b*, b-c**. Range 15 m (E

shore of Lake Pulmankijärvi, 7767:3538) –

ca. 420 m (Tsuomasvarri, 7755:3548). Only

seldom ascending to the tree line. Tr 880 m

(Engelskjøn & Skifte 1995: 140), Fnm 441

m (Norman (I)1: 441), EnL 720 m (Väre &

Partanen 2009: 128). Silvike.

Ecology. E. davuricum is a week

competitor requiring fairly free and open

habitats created by inundation and freezing

and melting of the ground. Furthermore, it

favors muddy, silty and sandy mineral soil

by springs and spring-fed brooklets. In the

region of Saariselkä, the species occurs in

flat, moist frost ground sites and in stony

sites with flowing spring water (Rintanen

1970a: table 1 and 4). It often grows as

solitary individuals or small groups in the

edges of quagmires and meso-eutrophic

fens and mires together with the mosses

Campylium stellatum, Dicranella palustris,

Philonotis spp., Pseudobryum cinclidioides,

Sphagnum warnstorfii and Tomentypnum

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 37

nitens. Typical vascular companions in the

coniferous zone are Agrostis mertensii,

Bartsia alpina, Carex capillaris, C. dioica,

Epilobium hornemannii, E. palustre,

Equisetum variegatum, Luzula sudetica,

Pinguicula vulgaris, Selaginella

selaginoides and Viola epipsila. In the

lowermost alpine belt of Tsuomasvarri fjeld

the exacting calciphilous plants Carex

parallela, Pinguicula alpina and Saxifraga

aizoides grow in the immediate vicinity of

E. davuricum.

E. davuricum is often considered

exacting – rather exacting (Pesola 1928:

159, Björkman 1939: 62, Arwidsson 1943:

226, Wistrand 1962: 125, Kytövuori 1969:

52, Gjærevoll 1990: 63, Nilsson 2000:

148). However, in Inari Lapland it grows

both in the calcium-poor granulite area and

on calcareous schists and hence appears to

be less demanding, being indifferent or at

most slightly basocline (Laine 1970(II):

125, Mäkinen & Kallio 1979: 17). Slightly

basocline.

Morphology and taxonomy. E.

davuricum has usually a fairly well-defined

habit, and it does not show any

morphologically or taxonomically

significant variation. However, tiny white-

flowered specimens with opposite leaves

resemble small plants of E. palustre var.

lapponicum.

Dependence on culture. The species

thrives also in man-made habitats, e.g. in

roadside ditches, in gravel pits and swampy

clearings. Hemeradiaphore.

UL

Epilobium hornemannii Rchb.

E. alpinum auct. p.p.

Indigenous, scattered

Map 26

Distribution. Arctic-montane, incompletely

circumpolar with a large gap in N Asia (amphi-

Atlantic and amphi-Pacific), but like many other

Epilobium species, critical and often misidentified

(Benum 1958: 296, Hultén 1958: 196, Hultén & Fries

1986: map 1368, Lid & Lid 2005: 563, Fl. Nord. 6:

116). Arctic and subarctic Europe (Fl. Eur. 2: 311).

Most of Fennoscandia except for S Sweden and S

Finland, in Finland fairly common S to Kainuu

(Hultén 1971a: map 1282, Hämet-Ahti et al. 1998:

300, Mossberg & Stenberg 2003: 418, Fl. Nord. l.c.,

Lampinen & Lahti 2018).

Common in Troms (Benum 1958: 296,

Engelskjøn & Skifte 1995: 140), fairly common in

Finnmark, but mainly in coastal regions (Norman

I(1): 434, Dahl 1934: 367), only one locality in the

Rastigaissa area (Ryvarden 1969: 33). More or less

common throughout Pechenga and the Kola

Peninsula (Fellman 1831: 310, Wainio 1891: 51,

Valle 1933a, Fl. Murm. IV: map 75, Mäkinen 2002:

20), but rarer in the alpine belt (Kalliola 1932: 106,

Söyrinki 1939a: 299, Alm et al. 1998: 135). Rare in

the Lutto area and E Saariselkä (Roivainen 1923:

291). One locality mentioned in Keret Karelia by

Söyrinki (1956: 28).

Rather rare to fairly common in Sompio and

Kittilä Lapland (Hjelt & Hult 1885: 128, Wainio

1891: 51, Hult 1898: 165, Hustich 1940c: 57, Kujala

1961, Montell 1962: 124, Lampinen & Lahti 2018).

In Enontekiö Lapland fairly common in the birch

belt, scattered in the alpine belt (Lindén 1943: 75,

Laine 1958: 84, Piirainen & Piirainen 1991b,

Lampinen & Lahti l.c.).

InL ref. Common (“in regionibus

lapinis [sic! = alpinis?] frequens”, Fellman

1835: 259, as E. alpinum). Several

localities up to the upper limit of the birch

belt (Kihlman 1884: 102). Inari, Törmänen

(Wainio 1891: 51). Fairly common in the

whole area and in all belts (Mikkola 1941:

31). Rather rare to scattered along the

Ivalojoki and mainly in the upper and

middle parts of the river (Kujala 1962:

177). Rather rare in the NE parts of Inari

(Såltin 1958), mostly found along brooks

and rivers. Scattered to fairly common in

the Lemmenjoki area (Klockars & Luther

1938, Rahkonen 1968: 18), but only five

localities reported in the northernmost pine

forest area in the Vaskojoki basin in W

Inari (Laine 1964: 114). Rather rare in the

Peäldoajvi fjeld area (Koivistoinen 1964:

51). In W Utsjoki rather rare to fairly

common (Laine et al. 1955: 130, Kallio &

38 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Mäkinen 1957: 26), scattered in the Kevo

Strict Nature Reserve (Laine 1970(II):

126).

Kevo 33.3 %, InL 55 %, 158 sq. H 75,

JYV 1, KUO 7, OULU 4, TUR 125, VOA

1, YME 14 spec.

Scattered (951; 0.146). Inari: III (449;

0.104), Utsjoki: IV (502; 0.229).

Difference***. The most common of the

northern Epilobium species in Inari

Lapland. In the whole province, but most

frequent in the fjeld areas and almost

missing in the Lake Inari basin. Whole

area.

FMF 0.674.

Vertical distribution. a: IV (115;

0.156), b: IV (526; 0.233), c: III (310;

0.089). Differences***. Range 15 m (S end

of Lake Pulmankijärvi, 7762:3539) – ca.

500 m (N side of Viibusoaivi, 7617:3457).

The main distribution is clearly in the birch

belt. EnL 1270 m (Lindén 1943: 75

”Haltitschohko W”, possibly on Norwegian

side?), 900 m (Laine 1958: 84), Tr 1000 m

(Engelskjøn & Skifte 1995: 140), Fnm 420

m (Dahl 1934: 368). Vertical ubiquitous

(cf. Kytövuori 1972: 196).

Ecology. The habitat requirements of

E. hornemannii are much like those of E.

alsinifolium, but with a clearly wider

amplitude. E. hornemannii is typical around

moss covered edges of springs and ponds,

often dominated by the moss Cratoneuron

commutatum. It often grows together with

Epilobium alsinifolium, but usually in

somewhat drier parts around the springs

than the latter. E. hornemannii also grows

in moist depressions and willow thickets

especially in the low-alpine belt, along

brook banks, near snowbeds, in wet, mossy

slopes and at the base of steep cliffs where

the soil remains moist. It is found even in

swampy bogs and mire margins. It is a

weak competitor, and prefers open

communities with a sparse field layer.

In Inari Lapland the associates of E.

hornemannii include both hygrophytes and

mesophytes. Typical companions in

mesotrophic localities are Angelica

archangelica, Bistorta vivipara, Carex

vaginata, Equisetum variegatum, Juncus

biglumis, Luzula sudetica, Parnassia

palustris and Stellaria borealis. In more

eutrophic habitats the associates may

include Carex capillaris, Chrysosplenium

tetrandrum, Saxifraga aizoides, S. stellaris,

Selaginella selaginoides, Thalictrum

alpinum, Tofieldia pusilla and Viola

biflora. In the Kevojoki valley E.

hornemannii grows together with, e.g.,

Arabis alpina, Epilobium anagallidifolium

and Eriophorum scheuchzeri, as well as the

mosses Bryum weigelii, Drepanocladus

badius and Philonotis tomentella (Laine

1970(II): 126). In the Peäldoajvi fjeld area

E. hornemannii is almost always

accompanied by Stellaria borealis and the

mosses Dicranella squarrosa and

Drepanocladus exannulatus (Koivistoinen

1964: 51). At upper elevations E.

hornemannii occurs along small gravelly

brooks or in moist rock-crevices by brooks,

often with Epilobium anagallidifolium,

Salix herbacea and Veronica alpina, in the

Kistuskaidi area also with Rhodiola rosea.

In the snow-bed localities it may be

accompanied by Carex lachenalii, Cassiope

hypnoides, Saxifraga stellaris, Sibbaldia

procumbens and Viola biflora.

E. hornemannii flowers earlier than E.

alsinifolium, and therefore their hybrids are

not particularly common despite the fact

that both species are often growing

together. Near the Kevo Subarctic Research

Station in Utsjoki in 2004 flowering started

on July 1 (Alanen 2007).

E. hornemannii is sometimes

considered basocline (Pesola 1928: 159,

Mäkinen & Kallio 1979: 17, Lid & Lid

2005: 563), but mostly amphicline/Ca-

indifferent (Björkman 1939: 62, Arwidsson

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 39

1943: 226, Benum 1958: 296, Wistrand

1962: 126, Laine 1970(II): 126, Roweck

1981: 326, Nilsson 2000: 147, Fl. Nord. 6:

116). Amphicline.

Parasites. The species is sometimes

infected by the rust Puccinia epilobii DC.

(Mäkinen 1964b: 167).

Morphology and taxonomy. Young

and tiny specimens of the first year may be

misidentified as E. anagallidifolium. – The

chromosome number 2n = 36 has been

determined from material collected in

Utsjoki (a road verge by the Raessijoki, N

of Kevo Subarctic Research Station, TUR

73648; Laine et al. 1974).

E. hornemannii hybridizes fairly easily

with several other species of Epilobium.

The following hybrids have been recorded

in Inari Lapland:

E. alsinifolium × hornemannii, see

under E. alsinifolium.

E. anagallidifolium × hornemannii, see

under E. anagallidifolium.

E. hornemannii × palustre is a fairly

common hybrid in the coniferous region of

Lapland (Montell 1962: 124, SKK III:

174). In Inari Lapland it has been found

along the Ivalojoki at least in a few

localities (“st r”, Kujala 1962: 177) usually

in the company of the parent species. Other

localities: Inari: (1) Moberginoja brook,

eutrophic spring fen (759:349, 1925 E.

Mikkola, TUR 73731); (2) S of Lake

Menesjärvi, shore of the

Kaddsatsuohppamjohka (7619:3475, 1983

Y. Mäkinen, YME 14294); (3) Majavaoja

rivulet N of Lake Hammasjärvi

(7630:3492, 1976 Y. & L. Mäkinen, YME

6080); (4) Tuulispää, Tammukkalampi

pond, by a spring brook (7640:3504, 1972

C. E. Sonck, TUR 263708); (5) Lake Iso

Pielpajärvi, NE corner (7652:3504, 1972 C.

E. Sonck, TUR 261913); Utsjoki: (6) S of

Lake Allamarasjavri (7700:3463, Laine et

al. 1955: 130); (7) S side of Karegasnjarga-

Ailigas (7704:3461, Laine et al. l.c.); (8)

NW side of Meäddemvarri, by a spring-

water brook of the Pullinoja rivulet

(7705:3456, 1954 Y. Mäkinen, YME 6072,

Laine et al. l.c.); (9) W side of

Njavgoairoavvi (7713:3475, Laine et al.

l.c.); (10) W of Kevo, upper course of the

Kaskamus Madjoksuorgi (7734:3489, 1993

Y. Mäkinen, YME 21919); (11) Kutuniemi

– Patoniva, shore of the Rassejohka

(7743:3500, 1964 U. Laine, TUR 125280);

(12) N of Vaisjeäggi, tributary of the

Hannojohka (7753:3509, 1961 Y. Mäkinen,

field list); (13) Teno valley, ca. 1 km S of

Kaava, brook side (7762:3494, 1991 K. &

R. Alho, K. & U. Laine, TUR 321564);

(14) Teno valley, brook side N of Kaava

(7764:3494, 1991 K. & R. Alho, K. & U.

Laine, TUR 319150).

Dependence on culture. E.

hornemannii spreads easily in wet, man-

made habitats like ditches in home fields,

wet tracks, roadsides and edges of routes

used in winter (cf. Kytövuori 1972: 198).

The species is also found in localities rich

in humus and nutrients, as around cow

sheds and on garbage and dung heaps.

Hemeradiaphore (Mäkinen & Kallio 1979:

17) Hemeradiaphore or slightly hemero-

philous.

UL, JN

Epilobium lactiflorum Hausskn.

E. alpinum L., nom. rej.

Indigenous, very rare

Map 27

Distribution. Amphi-Atlantic, arctic-montane

(Hultén 1958: 194, Hultén & Fries 1986: map 1369).

Common – fairly common in the Scandes and in

most of Norway, scattered – rare elsewhere in N

Fennoscandia (Hultén 1971a: map 1283, Roweck

1981: 327, Mossberg & Stenberg 2003: 418, Lid &

Lid 2005: 564, Fl. Nord. 6: 116).

Fairly common in Troms, scattered – rather rare

in Finnmark (Norman I(1): 430, II(1): 267, Dahl

1934: 367, Benum 1958: 296, Engelskjøn & Skifte

1995: 140), but only one locality in the Rastigaissa

40 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

fjeld area (Ryvarden 1969: 33). Rare – very rare in

the subalpine and alpine belts of the Lutto area (1921

H. Roivainen, TUR 73755, Roivainen 1923: 292), in

Pechenga and in the Kola Peninsula (Kalliola 1932:

106, Kalela 1939: 161, 266, Söyrinki 1939a: 259,

297, Fl. Murm. IV: map 74, Alm et al. 1998: 135).

Three localities in the Salla district on the Russian

side (Rintanen 1968: 283, 1970b: 360, SKK III: 173).

No records in Sompio Lapland, very rare in the

Pallastunturi – Ounastunturi National Park in Kittilä

Lapland (Rantaniemi 1921c, Hustich 1936c, 1940c:

57, Montell 1962: 124, Lampinen & Lahti 2018), not

found elsewhere in the province. In Enontekiö

Lapland mainly in the NW fjeld area (Lindén 1943:

75, Jalas 1949, Kallio 1949, Lammes 1991, Piirainen

& Piirainen 1991b, Lampinen & Lahti l.c.).

InL ref. One locality in the S part of the

Kevo Strict Nature Reserve (Laine

1970(II): 125, Heikkinen & Kalliola 1990:

34).

Kevo 0.3 %, InL 2 %, 3 sq. TUR 7.

Very rare (7; 0.001). Utsjoki: I (7;

0.003). (1) A springy brook-bed on the SE

slope of Suohppasoaivi fjeld in the upper

course of the Kevo (7711:3476, 1966 U.

Laine, TUR 163241); (2) a springy

subalpine valley between Tuoljivaldem-

roavvi and Tsuodjaroavvi fjelds

(7727:3521, 1965 P. Siltanen, TUR

172533); (3) a brook valley in the S slope

of Kaldoaivi fjeld (774:353, 1959 R.

Nikoskelainen, TUR 595696); (4) a stony

and springy valley between Varddoaivi and

Guovloaivi fjelds (7765:3522, 1959 P.

Siltanen, TUR 197778); (5) on the S shore

of a springy depression at the base of

Guovloaivi fjeld (7766:3522, 1959 P.

Siltanen, TUR 119114); (6) at the base of

the E slope of Jovnnaleägeoaivi fjeld

(7766:3531, 1975 K. & U. Laine, J. Nurmi,

field record); (7) E slope of Avvatšohkka

fjeld (7766-7767:3532, 1975 K. & U.

Laine, J. Nurmi, TUR 293509, 293510). A

local concentration of occurrences in

Guovloaivi – Varddoaivi – Avvatšohkka

wilderness area in NNE Utsjoki. – The

determination of specimens confirmed by

T. Karlsson. Northern.

FMF 0.020.

Vertical distribution. a: I (5; 0.007),

b: I (1; 0.000). Difference**. Range 225 m

(Guovloaivi) – 410 m (Suohppasoaivi).

Most localities in the lower part of the

alpine belt. Tr 900 m (Engelskjøn & Skifte

1995: 140), Fnm 650 m (Dahl 1934: 367),

EnL 900 m (Laine 1958: 84, Väre &

Partanen 2009: 129). Alpine.

Ecology. All the occurrences are rather

scanty and situated in springy places,

preferably on mineral soil. The following

associates were recorded on the sloping

meadow of Suohppasoaivi fjeld: Bistorta

vivipara, Carex lachenalii, Epilobium

hornemannii, Pinguicula vulgaris,

Ranunculus nivalis, Saxifraga stellaris and

Veronica alpina. In NNE Utsjoki typical

companions are e.g. Bartsia alpina, Carex

lachenalii, Equisetum pratense, Parnassia

palustris, Pinguicula alpina and Stellaria

borealis, sometimes also Carex capillaris,

Milium effusum, Thalictrum alpinum,

Vahlodea atropurpurea and Viola biflora.

E. lactiflorum is usually considered

amphicline, indifferent or slightly

basophilous (Dahl 1934: 367, Arwidsson

1943: 226, Wistrand 1962: 126, Laine

1970(II): 126, Mäkinen & Kallio 1979: 17;

see also Fl. Nord. 6: 117). In Inari Lapland

slightly basocline.

Dependence on culture. Ahemerobe.

UL

Epilobium palustre L.

Indigenous, scattered

Map 28

Distribution. Circumpolar in several races (Hultén

1971b: 124, Hultén & Fries 1986: map 1365).

Common to fairly common in the whole

Fennoscandia but less common in alpine areas and

the N part of the Kola Peninsula (Hultén 1971a: map

1287, Mossberg & Stenberg 2003: 417, Lid & Lid

2005: 563, Fl. Nord. 6: 101). In Finland very

common to common throughout the country (Kujala

1964: 75, Hämet-Ahti et al. 1998: 299, Lampinen &

Lahti 2018).

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 41

In Troms and Finnmark common throughout

the area, especially in the lowlands (Norman I(1):

437, II(1): 271, Dahl 1934: 368, Benum 1958: 297,

Engelskjøn & Skifte 1995: 141), five localities in the

Rastigaissa area (Ryvarden 1969: 33). Common in

the Kola Peninsula (Fl. Murm. IV: map 76).

Scattered to rather common in the inner parts of

Pechenga, the Lutto area (Roivainen 1923: 292),

throughout the coniferous zone (Valle 1933a: 270,

Alm et al. 1997: 41, Mäkinen 2002: 20) north to the

Arctic Ocean (Wainio 1891: 51); rare in the alpine

belt in the Kammikivi area (Kalliola 1932: 106) and

the Pechenga fjelds (Söyrinki 1939a: 291).

Common in Finnish Lapland (Fellman 1835:

259). Rather common in Sompio and Kittilä Lapland

(Hjelt & Hult 1885: 127, Hult 1898: 165, Hustich

1940c: 57, Montell 1962: 124, Laitinen & Ohenoja

1990: 22, Hämet-Ahti et al. 1998: 299, Lampinen &

Lahti 2018). Common in the E parts of Enontekiö

Lapland, less common towards the W, and scattered

in the Lake Kilpisjärvi area (Lindén 1943: 75,

Piirainen & Piirainen 1991b, Lampinen & Lahti l.c.).

InL ref. Rather common in the

coniferous zone and subalpine birch belt

(Kihlman 1884: 102, Mikkola 1941: 31).

Scattered in the whole Ivalojoki valley

(Kujala 1962: 177). Scattered to rather rare

in the Viipustunturit – Maarestatunturit

fjeld area (Klockars & Luther 1938). Rather

rare in the Peäldoajvi fjeld area

(Koivistoinen 1964: 50), rare in the NE

parts of Inari (Såltin 1958). Rather rare in

W Utsjoki (Kallio & Mäkinen 1957: 26),

scattered in the Paistunturit area (Laine et

al. 1955: 130), rather rare in the Kevo Strict

Nature Reserve (Laine 1970(II): 125).

Kevo 49.3 %, InL 75 %, 208 sq. H 25,

KUO 2, OULU 5, TUR 49, YME 12 spec.

Scattered (1449; 0.222). Inari: IV

(747; 0.174), Utsjoki: V (702; 0.317).

Difference***. Rather evenly distributed

throughout most of the area but locally less

common in the E and NE parts of the Lake

Inari basin. Whole area.

FMF 0.850.

Vertical distribution. a: IV (119;

0.165), b: V (699; 0.308), c: IV (631;

0.180). Difference a-b***, b-c***. Range

15 m (Lake Pulmankijärvi, 7762:3539) –

525 m (Kalgoaivi, 7670:3473). Tr 925 m

(Engelskjøn & Skifte 1995: 141), Fnm 490

m (Norman I(1): 440), EnL 850 m (Väre &

Partanen 2009: 128). Vertical ubiquitous.

Ecology. Epilobium palustre grows in

many kinds of wet and moist habitats on

peat or mineral soil. It is most common

along the shores of rivers and brooks, often

on moist sand or gravel, but is often met

also on lakeshores. It grows also e.g. on

cliff terraces with dribbling water. In the

Kevojoki valley it grows often together

with e.g. Agrostis mertensii, Bartsia alpina,

Galium uliginosum, mosses Blindia acuta,

Sanionia uncinata and lichens Baeomyces

rufus and Cetrariella delisei (Laine

1970(II): 125). It thrives also in swamps,

meso- and eutrophic mires, paludified

meadows and wet Salix thickets. It is often

found along spring brooks and on spring-

effected bare mineral soil. Rarely it grows

in small hollows or belts of bare frost

ground at bog edges (Rintanen 1970a:

Table 1).

The flowering takes place in July;

flowering specimens have been collected

from the end of June to mid-August. In the

surroundings of the Kevo Subarctic

Research Station first flowers are seen in

mid-July (Hustich 1942a). In the Pechenga

fjelds Epilobium palustre flowers in late

July – early August, and seeds are

dispersed ca. 6 weeks later (Söyrinki

1939a: 292). Overwintering takes place by

turions formed at the tips of long and

slender stolons. Turions serve also as a

means of vegetative spreading, transported

e.g. by moving water (Kytövuori 1969: 51,

SKK III: 165, Fl. Nord. 6: 101). The

species is usually said to be

amphicline/indifferent (Arwidsson 1943:

226, Benum 1958: 297, Wistrand 1962:

125, Laine 1970(II): 125, Mäkinen &

Kallio 1979: 17, Nilsson 2000: 149), but in

Kuusamo it seems to be weakly

calciphilous (Kytövuori 1969: 54).

Amphicline.

42 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Morphology and taxonomy. Both the

genetic variation and habitat modification

of the species are wide. Several

infraspecific forms and varieties are named

in the literature, but most of them seem to

have no greater taxonomic value. Only one

entity, var. lapponicum Wahlenb., is

accepted here in addition to the nominal

variety (cf. Hämet-Ahti et al. 2005a: 65;

however, the nomenclature needs

verifying). The plants are small (10–20

cm), slender and unbranched, with several

short internodes near the base; leaves are

upright, almost appressed to the stem, short,

linear with a rounded tip and with stomata

also on the upper surface (missing from the

nominal variety); flowers are usually

solitary, white or pale pinkish and larger

than those of the nominal variety; seeds are

large and almost smooth. Habitats of var.

lapponicum are wet, mainly eutrophic

mires. Only records from herbarium

specimens are accepted (the localities are

not mapped separately): Inari: (1) Sallijoki

N of the Repojoki, rich fen (7598:3456,

1983 J. Savola, TUR 274186), (2) ca. 2 km

N of the Repojoki, E of the Lemmekäsjoki,

sloping rich fen (7600:3447, 1983 J.

Savola, TUR 274187), (3) Kultala

(7602:3487, 1878 E. Wainio, TUR 74049),

(4) shore of the Ivalojoki (ca. 761:352,

1956 A. Valta, TUR 569045), (5) Kyrö

(=Ivalo) (762:52, 1878 E. Wainio, TUR

74048), (6) shore of Lake Ahvenjärvi by

the Vaskojoki (7644:3473, 1937 I. Mikola,

H 407478), Utsjoki: (7) Muotkatunturit, NE

part of Tuolba Saraoaivi, alpine brook side

(7692:3474, 1988 Y. Mäkinen 88-320 & L.

Mäkinen, YME 16932), (8) Muotkatunturit,

upper Sarajohka, moist mossy brook side

(7692:3475, 1988 Y. Mäkinen 88-325 & L.

Mäkinen, YME 16934), (9) Karigasniemi,

ca. 500 m S from W end of Pasijärvi,

peatland on tarn shore (7701:3462, 1954 N.

Tarén, TUR 170925; 1954 Y. Mäkinen,

YME 6063), (10) peatland N of the small

tarn N of Kâskamuš Tsieskuljavri

(7739:3505, 1960 P. Siltanen, TUR 73527).

Four hybrids of Epilobium palustre are

known from the study area, namely E.

alsinifolium × palustre, E. anagallidifolium

× palustre, E. collinum × palustre, and E.

hornemannii × palustre; see under E.

alsinifolium, E. anagallidifolium, E.

collinum and E. hornemannii, respectively.

Dependence on culture. Epilobium

palustre favors open habitats and the

nominal variety var. palustre is sometimes

met in ditches, wet semicultural meadows

and other moist sites influenced by man.

Hemeradiaphore.

MP

HALORAGACEAE

Myriophyllum alterniflorum DC.

Indigenous, rather rare

Map 29

Distribution. Boreal, amphi-Atlantic (Hultén 1958:

252). Widely spread in most of Europe with a clearly

oceanic and northern tendency (Fl. Eur. 2: 312;

Meusel et al. 1978: 302, Blamey & Grey-Wilson

1989: 266). Less common in N America, chiefly in

the eastern parts (Gleason 1952: 600, Hultén 1958:

map 234, Hultén & Fries 1986: map 1375).

Rather common and often locally abundant in

most of Fennoscandia. Declining – rare in Denmark

and the southernmost part of Sweden, especially in

the islands and the coastal provinces of the Baltic

Sea, but avoiding brackish water and calcareous soil.

Scattered localities in the Scandes, N Sweden and

arctic Fennoscandia (Samuelsson 1934: 41,

Arwidsson 1943: 226, Hultén 1971a: map 1296, SKK

III: 189, Roweck 1981: 331, Hultén & Fries 1986:

map 1375, Nilsson 2000: 150, Mossberg & Stenberg

2003: 420, Fl. Nord. 6: 152, Haeggström &

Haeggström 2010: 267). One of the commonest and

most widely distributed aquatic plants in

Fennoscandia (Samuelsson l.c.). No confirmed

records east of Fennoscandia (Hultén 1958: map 234,

Hultén & Fries l.c.).

Infrequent in Troms (Norman I(1): 444, Benum

1958: 298, Engelskjøn & Skifte 1995: 142). Rather

rare – rare in Finnmark, even if locally abundant in

some larger rivers in Karasjok and Kautokeino

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 43

(Norman I(1): 445, Dahl 1934: 370, Lid & Lid 2005:

569), only three occurrences in the Rastigaissa fjeld

area (Ryvarden 1969: 33). In Pechenga scattered

localities in lakes and rivers in the Lutto area and the

Paatsjoki river system (Wainio 1891: 50, Alm et al.

1997: 41). A few localities in the Kola Peninsula and

Kandalaksha area (Fl. Murm. IV: map 78,

Tolmatchev 1980: 57, Mäkinen 2002: 20). Fairly

common in Sompio Lapland, Kittilä Lapland and

Enontekiö Lapland as well as Koillismaa (Hjelt &

Hult 1885: 128, Hult 1898: 165, Roivainen 1923:

292, Lindén 1943: 76, Pesola 1952, Salonen 1956,

Montell 1962: 124, Söyrinki & Saari 1980: 117,

Rintanen 1976 and 1982a: 253, Laitinen & Ohenoja

1990: 23, Piirainen & Piirainen 1991b, Hämet-Ahti et

al. 1998: 302, Lampinen & Lahti 2018). Also in

Keret Karelia (Söyrinki 1956: 28, Sokolov & Filin

1996: 115).

InL ref. Utsjoki, Mandojärvi (1880 A.

Arrhenius & A. O. Kihlman, H 409070). In

the coniferous zone and the birch belt fairly

common but mostly sterile (Kihlman 1884:

103). Näätämö (Hjelt 1911: 374),

Morgamjärvi, Ravadasjärvi and Vaskojoki

in the Lemmenjoki National Park (Klockars

& Luther 1938). Fairly abundant in several

larger rivers (Mikkola 1941: 32). Lake

Inari, Ukonselkä, Vastusjärvi, Talvitupa-

järvi (Maristo 1941: 79, 80, table).

Muddusjärvi (Tuomikoski 1950). Five

localities in SW Utsjoki (Laine et al. 1955:

130). Ivalojoki, fairly common and

abundant especially in the lower course

(Kujala 1962: 177). Obviously rare in the

Peäldoajvi fjeld area (Koivistoinen 1964:

52). Only four observations in the Angeli –

Vaskojoki anorthosite area (Laine 1964:

114), but evidently partly overlooked.

Frequent and abundant in numerous well-

studied lakes and quiet waters in Inari and

Utsjoki (Kallio 1959a: 169, Nyman 1964,

Rautava 1964: 84, 1969: 115, Siltanen

1964: 51, 1967: 162). Rare in the Kevo

Strict Nature Reserve (Laine 1970(II): 127,

Heikkinen & Kalliola 1990: 34). In 33 of

the 38 lakes studied by Rintanen (1982a:

253).

Kevo 3.3 %, InL 65 %, 182 sq. H 20,

KUO 1, OULU 3, TUR 28, YME 5 spec.

Rather rare (852; 0.132). Inari: IV

(712; 0.167), Utsjoki: II (140; 0.061).

Difference***. Clearly commoner in Inari.

Strongly concentrated in Lake Inari basin

and the river systems of the Ivalojoki and

Vaskojoki. In the Utsjoki river system,

most continuous between the lakes

Kenesjärvi – Puksaljärvi – Kevojärvi –

Mantojärvi (e.g. Nyman 1964, Siltanen

1964: 51). The rarity at higher altitudes,

e.g. in the fjeld areas of Muotkatunturit,

Paistunturit and Peäldoajvi, is due to the

lack of suitable water basins. The species is

most likely partly overlooked in the

Inarijoki – Teno watercourse area. Further-

more, M. alterniflorum avoids extensive

swamp areas. – Part of the observations is

based on drifting shoots. Lowland.

FMF 0.789.

Vertical distribution. a: I (11; 0.015),

b: III (157; 0.070), c: IV (684; 0.195).

Differences*** Range ca. 15 m (S side of

Lake Pulmankijärvi, 7762:3539) – 425 m

(Stuor Avdshigas, 7684:3463). Tr 694 m

(Engelskjøn & Skifte 1995: 142), Fnm 380-

400 m (Dahl 1934: 370), EnL 700 m (H.

Väre, pers. comm.). M. alterniflorum is

prevalently a species of the coniferous

zone, often ascending to the subalpine birch

belt but very seldom to the low-alpine one.

Silvike.

Ecology. Typical habitats of M.

alterniflorum are sheltered and shallow,

base-poor – oligotrophic, clear and oxygen-

rich lakes, pools and tarns. Furthermore, the

species forms extensive, nearly pure stands

in slowly flowing extensions and bays

(river-lakes) of greater rivers on silty, sandy

or even coarse gravelly bottom, sometimes

also in dystrophic tarns. The growing depth

varies between 20 and 350 cm (Nyman

1964, Rautava 1964: 87, 1969: 115,

Siltanen 1964: 51, 1967: 164). The species

is often predominant in its habitats. In

chilly and rainy summers the stands remain

almost sterile, whereas in favorable warm

44 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

years they may flower profusely. The

species flourishes also in the rapids of

strongly flowing rivers. In general, the

flowering starts not until July or the

beginning of August.

In the shallow littoral zone M.

alterniflorum forms tight, homogenous

submerged stands along shores

(“Gürtelbildung”) or grows in the gaps of

the isoetid zone together with Isoëtes spp.,

Eleocharis acicularis, Ranunculus reptans,

Sparganium hyperboreum and Subularia

aquatica. Typical associates in deeper

waters are e.g. Potamogeton alpinus, P.

gramineus, P. perfoliatus, Sparganium

angustifolium and Utricularia vulgaris, in

lake habitats also Isoëtes lacustris (Maristo

1941: 79). In small pools on rocky bottom

by the Kevojoki Hippuris vulgaris,

Ranunculus confervoides and R. peltatus

are its companions (Laine 1970(II): 127).

Also the bryophytes Fontinalis dalecarlica,

Scorpidium scorpioides, Warnstorfia

fluitans and W. trichophylla as well as the

algae Chara fragilis and Nitella opaca may

belong to the associates.

Thunmark (1931: 85) considers the

species very oligotrophic, Samuelsson

(1934: 62) oligotrophic, Wistrand (1962:

127) indifferent. In any case, M.

alterniflorum favors infertile and nutrient-

poor waters, but thrives also in slightly

eutrophic ones (Benum 1958: 298,

Engelskjøn & Skifte 1995: 142). pH

amplitude 6.2-7.6 (Maristo 1941: 241,

Table 9). Amphicline (Laine 1970(II): 127,

Mäkinen & Kallio 1979: 17). Amphicline –

slightly acidocline.

Morphology. Exceptionally long-

leaved specimens have been collected from

Lake Njammijärvi in E Inari (7675:3568,

1986 J. Savola, TUR 312974).

Dependence on culture. M. alterni-

florum avoids dirty, polluted water.

Hemeradiaphore – slightly hemerophobe.

UL

Myriophyllum sibiricum Kom.

M. spicatum L. subsp. squamosum Laest.

ex Hartm.

M. exalbescens Fernald, M. spicatum L.

subsp. exalbescens (Fernald) E. Murray

Indigenous, very rare

Map 30

Distribution. Probably circumboreal; the total

distribution area incompletely known. N American

and Russian authors have considered M. sibiricum a

separate species for quite a long time, European and

particularly Fennoscandian taxonomists have not

recognized it at all or included it in M. spicatum L. as

a variety or subspecies (Fernald 1919, Löve 1961,

Faegri 1982a, 1982b, Waern & Pekkari 1980, Ceska

& Ceska 1986, Fl. Nord. 6: 155). Now M. sibiricum

is unanimously accepted as a separate species.

In the coastal provinces in most of Finland and

Sweden fairly common, rarer inland. Absent – very

rare in Denmark, SW Sweden and most of central

and S Norway (Wistrand 1962: 126, SKK III: 187,

Roweck 1981: 331, Hultén & Fries 1986: map 1374,

Hämet-Ahti et al. 1998: 301, Nilsson 2000: 150,

Mossberg & Stenberg 2003: 420, Lid & Lid 2005:

569, Fl. Nord. 6: 154).

In Troms numerous collections from S and

central parts, in Finnmark scattered and mainly in S

inland (Engelskjøn & Skifte 1995: 142, Lid & Lid

2005: 569, Elven et al. 2013: 264). In the earlier

papers from N Norway M. sibiricum is included in

M. spicatum (Norman I(1): 445, II(1): 277, Dahl

1934: 370, Benum 1958: 299). M. spicatum s.lat. is

rare to very rare in Pechenga (Wainio 1891: 50,

Söyrinki 1939a: 302, Mikkola 1941: 32 (Paatsjoki);

TUR) and in the Kola Peninsula (Hultén 1971a: map

1297, Fl. Murm. IV: map 78, Tolmatchev 1980: 55).

Because M. spicatum s. str. is lacking in Finnish

Lapland (Hämet-Ahti et al. 1998: 301, Lampinen &

Lahti 2018), it is most likely that the references in

literature concern M. sibiricum. It grows in numerous

eutrophic Stratiotes lakes in Sompio Lapland and

especially in Kittilä Lapland (Salonen 1954, 1956,

Montell 1962: 124, Rintanen 1976, 1982a: 253,

Lampinen & Lahti l.c.). Several finds in Koillismaa

and Keret Karelia (Pesola 1952b, Söyrinki 1956: 28,

Söyrinki & Saari 1980: 116, Sokolov & Filin 1996:

115, Lampinen & Lahti l.c.).

InL ref. Old information concerning M.

spicatum from Inari Lapland refers to M.

sibiricum (single misidentified observations

of M. alterniflorum may be included).

“Enare Lapland” (Fellman 1835: 286).

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 45

Lake Inari, Ukonselkä (Maristo 1941:

table). Angeli, Inarijoki (Laine 1964: 114).

Keptujoki and Vaskojoki (7 and 19

localities, Rautava 1964: 84, 1969: 112,

table, 1971: table, Rintanen 1982b). Lake

Pieni Iivananjärvi and Lake Suuri

Iivananjärvi (Siltanen 1967: 160, 1977

Rintanen in H-Arch., Kastikka 2018,

Lampinen & Lahti 2018, see also Mäkinen

& Kallio 1979: 17).

InL 3 % (as M. spicatum), 11 sq. H 3,

OULU 1, TUR 5, YME 6 spec.

Very rare (19; 0.003). Inari: I (19;

0.004). (1) Ivalojoki, Ivalolommol

(7589:3427, 1970 Y. Mäkinen, field list,

YME); (2) S end of Lake Kietsimäjärvi

(7615:3423, 1982 Y. Mäkinen, YME); (3)

Nellim, Vuopaja, Rantala (7641:3551, 1970

Y. Mäkinen, field list, YME); (4-8) Paatari,

in the mouth and the lowermost course of

the Vaskojoki (7645:3485, 7646:3485,

7646:3486, 7647:3486, 7648:3486, 1961 U.

Laine, TUR 74459, 119476, Rautava 1964:

84; the localities along the Kettujoki are

referable only to the square 764:348); (9-

10) Angeli, Vuopionsuu, Inarijoki

(7648:3446, 1960 U. Laine, field list, TUR

74460; 7649:3446, 1965 Y. Mäkinen, field

list, YME); (11) Lake Inari, at the mouth of

Siskelvuono ca. 4 km SE from the church

(ca. 7646:3506, 1939 L. Alariesto, H

168040), and (possibly from the same

locality) Ukonselkä, ca. 5 km SE from the

church (764:350, Maristo 1941: table); (12-

15) Tirro, along the Vaskojoki (7650:3487,

7651:3480, 7651:3487, 7652:3486, 1961 U.

Laine, TUR 119475, Rautava l.c.); (16) N

of Angeli, Lake Pieni Iivananjärvi

(7653:3449, 1977 T. Rintanen, H 497034,

Rintanen 1982b); (17-18) N of Angeli,

Lake Suuri Iivananjärvi (7654:3449,

7655:3449, 1965 P. Siltanen, field list,

1965 Y. Mäkinen, field list, 1967 P.

Siltanen, TUR 272651, YME, 1977 T.

Rintanen, H 497059); (19) Kielajoki,

Alempi Honkavuoma (7687:3487, 1986 Y.

Mäkinen, field list, YME). – The specimen

in OULU has not been available.

The amounts and the list of localities

above include only observations based on

specimens or otherwise considered

trustworthy. In addition, there are a few

records in the field notes e.g. from the

Näätämöjoki and the Teno, but without

specimens they are not taken into account

here. The most copious occurrences are

concentrated in the SW part of Inari in the

lower course of the Vaskojoki and in the

lakes Pieni and Suuri Iivananjärvi.

Lowland.

FMF 0.029.

Vertical distribution. b: I (1; 0.000),

c: I (18; 0.005). Difference**. Almost

totally restricted to the coniferous zone.

Range 120 m (Lake Inari) – ca. 315 m

(Lake Kietsimäjärvi). Tr 694 m

(Engelskjøn & Skifte 1995: 142). Silvine.

Ecology. In the rivers Vaskojoki and

Kettujoki M. sibiricum grows on soft

muddy or fine sandy bottom mixed with

organic sediments, chiefly in a depth of 20-

140 cm (Rautava 1964: 84, 1969: 113). The

majority of the occurrences are in shallow

and sheltered parts of rivers with almost

standing water. The vegetative turions

allow the plant to grow in streams that

freeze in the winter. Typical associates in

the lower course of the Vaskojoki are

Equisetum fluviatile, Hippuris vulgaris,

Potamogeton gramineus, Sparganium

angustifolium and Utricularia vulgaris;

sometimes also Butomus umbellatus,

Callitriche hermaphroditica, Potamogeton

berchtoldii and Ranunculus confervoides.

Exacting companions in the Inarijoki by

Vuopionsuu at Angeli are Callitriche

hermaphroditica and Potamogeton

filiformis.

M. sibiricum avoids dystrophic and

oligotrophic waters. In the SW part of Inari

including Angeli district and the catchment

area of the Vaskojoki the bedrock is largely

46 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

composed of mafic volcanic rocks and

anorthosite. This obviously affects the

water chemistry in the area resulting in

more suitable conditions for demanding

aquatic plants. According to Maristo (1941:

241, Table 9), the pH amplitude of M.

sibiricum is 6.8-8.3, which is clearly higher

than that of M. alterniflorum. Basocline

(Mäkinen & Kallio 1979: 17), somewhat

calciphilous (Nilsson 2000: 150).

Basocline.

Morphology. Easily detachable, cone

like, scaly turions are present in almost all

herbarium specimens.

Dependence on culture. In Lake Inari

found close to landing places.

Hemeradiaphore.

UL

HIPPURIDACEAE

Hippuris vulgaris L.

Indigenous, rather rare

Map 31

Distribution. Almost circumpolar (Hultén 1971b:

122, Hultén & Fries 1986: map 1376). In

Fennoscandia fairly common but scattered – rare in

the extreme north (Hultén 1971a: map 1300,

Björkman 1939: 79, Arwidsson 1943: 227, Wistrand

1962: 127, Roweck 1981: 332, Mossberg & Stenberg

2003: 421).

Scattered in Troms and Finnmark (Norman I(1):

447, Dahl 1934: 369, Ryvarden 1969: 33), e.g.

Karasjok, Sør-Varanger, Vardø. Rare to fairly rare in

Pechenga and in the Kola Peninsula (Kalliola 1932:

96, 106, Fl. Murm. IV: map 79, Alm et al. 1997: 41).

Common – scattered in Finnish Lapland (Hjelt &

Hult 1885: 128, Hult 1898: 165, Hjelt 1911: 378,

Roivainen 1923: 292, Lindén 1943: 76, Laine 1958:

85, Kujala 1961, Montell 1962: 124, Hämet-Ahti et

al. 1998: 399, Lampinen & Lahti 2018), also found in

the alpine belt (Linkola 1932: 90, Söyrinki 1939a:

303, 1939b: 36, Piirainen & Piirainen 1991b).

Fairly common in the Oulankajoki and

Aventojoki in Kuusamo (Söyrinki & Saari 1980:

117). Common – fairly common in in the Kovda area

and in Keret Karelia (Söyrinki 1956: 28, Sokolov &

Filin 1996: 116).

InL ref. “in aquis stagnantibus fq”

(Fellman 1835: 245). Vaskojoki, Kettu-

Matti, Mantojärvi (Kihlman 1884: 103);

Ivalojoki (Wainio 1891: 50); Ravadasjärvi,

Vaskojoki (Klockars & Luther 1938); in all

three studied Carex-type lakes (Inari,

Vastusjärvi, Talvitupajärvi: Maristo 1941:

table). Ivalojoki 8 localities (Kujala 1962:

177). Vaskojoki fairly common (Laine

1964: 114, Rautava 1964: 77). In 8 lakes of

38 lakes studied by Rintanen (1982a: 251).

Leämmashjoki area (Kuitunen 1984). W

Utsjoki scattered (Laine et al. 1955: 130) –

very rare (Kallio & Mäkinen 1957: 26),

Kevojärvi fairly rare (Siltanen 1964: 47),

Mantojärvi scattered (Nyman 1964).

Kevo 4.4 %, InL 55 %, 166 sq. H 21,

JYV 2, KUO 1, OULU 1, TUR 30, YME 2

spec.

Rather rare (740; 0.111). Inari: III

(589; 0.136), Utsjoki: II (151; 0.061).

Difference***. Especially in the valleys of

larger rivers (the lower course of the Teno,

Utsjoki, Vetsijoki, Kaamasjoki, Ivalojoki,

Repojoki) and on the shores of Lake Inari.

Whole area.

FMF 0.714.

Vertical distribution. a: I (11; 0.015),

b: II (146; 0.061), c: IV (583; 0.163).

Differences***. Range 15 m

(Pulmankijärvi, 7762:3539) – 460 m

(Paistunturit, the uppermost Tsarsejohka in

the alpine belt, 7731:3479). Avoids large

alpine areas. Tr 810 m (Engelskjøn &

Skifte 1995: 142), Fnm 560 m (Ryvarden

1969: 33), EnL 800 m (Laine 1958: 85,

Väre & Partanen 2009: 156). 800 m in

Peljekaise in Swedish Lapland (Wistrand

1962: 127). Silvike.

Ecology. Hippuris vulgaris grows in

small lakes and ponds and in bays of larger

lakes, usually in small stands but may

occasionally cover areas of several square

meters. It is usually submerged at the depth

of 10-100 cm, with e.g. Callitriche

palustris, Isoëtes echinospora and

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 47

Ranunculus reptans (Siltanen 1964: 47) or

with Equisetum fluviatile, Potamogeton

gramineus, Ranunculus peltatus,

Calliergon giganteum, Warnstorfia

exannulata and W. fluitans (Laine 1970(II):

127; cf. Nyman 1964). Although the

species mostly grows on muddy bottom, it

favors spring-fed waters (Rintanen 1976,

1982a: 251).

According to Laine (1970(II): 127) the

species forms tight stands in rivers

especially in narrow places. In the

Vaskojoki it forms locally small groups in

which the stems are very densely and

exceed 1 m in length (Rautava 1964: 77).

Similar “Hippuris vulgaris Wiesen” have

been described by Eurola (1967: 41) in

Sodankylä especially in the bends of the

rivers.

Amphicline and Ca-indifferent (Benum

1958: 300, Wistrand 1962: 127, Laine

1970(II): 127, Mäkinen & Kallio 1979: 17,

Nilsson 2000: 150). Rarely considered

weekly calciphilous (Pesola 1928: 159), but

in Inari Lapland it is amphicline.

Morphology. A narrow-leaved and

lax-stemmed modification grows in all

streaming habitats, and seems to favor more

alpine localities.

Dependence on culture. Hemera-

diaphore.

YM

CORNACEAE

Cornus suecica L.

Chamaepericlymenum suecicum (L.) Asch.

& Graebn.

Indigenous, very frequent

Map 32

Distribution. Amphi-Atlantic (Hultén & Fries 1986:

map 1380); N Europe, also in N America and E Asia

(Hultén 1958: 256). Common in N Fennoscandia and

on the Atlantic shore (in Finland N of Lake

Oulujärvi); scattered – rare in S and central Finland

(Hultén 1971a: map 1302, Wistrand 1962: 127,

Kujala 1964: 76, Hämet-Ahti et al. 1998: 312,

Mossberg & Stenberg 2003: 422).

Common in Troms and Finnmark (Norman I(1):

533, Dahl 1934: 373, Benum 1958: 300, Vorren

1968, Ryvarden 1969: 33, Alm 1993c). Common –

scattered everywhere in N Finland, Pechenga and the

Kola Peninsula (Hjelt & Hult 1885: 130, Hult 1898:

164, Hjelt 1911: 247, Kalela 1939, Kalliola 1939:

259, Söyrinki 1939a: 306, Hustich 1940c: 58, Lindén

1943: 74, Laine 1958: 85, Pertola 1961: 35, Montell

1962: 124, Hämet-Ahti 1963a: 70, Ahti & Hämet-

Ahti 1971: 67, Söyrinki & Saari 1980: 117, Piirainen

& Piirainen 1991b, Piirainen 1996b, Alm et al. 1997:

41, 2000c, Roivainen 1923: 292, Fl. Murm. IV: map

90, Mäkinen 2002: 21). Common also in the Kovda

area (Sokolov & Filin 1996: 120).

InL ref. “Ubique vulgaris” (Fellman

1835: 251). Very common – scattered

(Castrén 1803, Kihlman 1884: 105, Wainio

1891: 53, Mikkola 1941: 32, Hämet-Ahti

1963a: e.g. 110, Haapasaari 1988: Tables

21, 23). Viipustunturit – Maarestatunturit

fairly common (Klockars & Luther 1938),

Ivalojoki common and abundant (Kujala

1962: 177), likewise Vaskojoki (Laine

1964: 114); Lake Hietajärvi area (Kvist

1978: 53). W Utsjoki fairly common (Laine

et al. 1955: 130) – common (Kallio &

Mäkinen 1957: 26, Laine 1970(II): 127,

Heikkinen & Kalliola 1990: 9).

Kevo 67.5 %, InL 94 %, 258 sq. H 23,

JYV 2, KUO 4, OULU 6, TUR 42, VOA 1,

YME 2 spec.

Very frequent (3940; 0.605). Inari: VI

(2411; 0.562), Utsjoki: VII (1529; 0.691).

Difference***. One of the commonest

vascular plants in Inari Lapland (cf.

Mäkinen & Kallio 1979: 6), avoiding only

the highest barren fjeld tops and some

extensive and wet swamp areas (cf.

Ruuhijärvi 1960: 154, 355). Whole area.

FMF 0.980.

Vertical distribution. a: VI (342;

0.470), b: VII (1688; 0.740), c: VI (1910;

0.547). Differences***. Very clearly

commonest in the birch belt, decreasing in

frequency both downwards and upwards;

especially the upward decreasing is very

48 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

rapid. The dominance in the birch belt is a

common phenomenon; according to Lindén

(1943: 74), the species is very common in

the birch belt and fairly common in the

lower alpine belt, and according to

Björkman (1939: 51, 1965: 36) it is rarer in

regio sylvatica than in the subalpine belt.

Range ca. 15 m (Lake Pulmankijärvi,

7762:3539) – 600 m (Karegasnjarga-

Ailigas, 7705:3460). Tr 700 m (Engelskjøn

& Skifte 1995: 142), Fnm 595 m (Norman

I(1): 541), EnL 810 m (Väre & Partanen

2009: 131). Pallas-Ounastunturit ca. 610 m

(Hustich 1937a: 62). Vertical ubiquitous.

Ecology. The main distribution in Inari

Lapland is clearly in the subalpine birch

forests. The species is most abundant in the

flooded birch woods along rivers and

brooks, but it also grows on moist moss-

rich slopes and below riverside cliffs, with,

e.g., Calamagrostis lapponica, Equisetum

pratense, Gymnocarpium dryopteris,

Lycopodium annotinum, Rubus arcticus, R.

saxatilis, and Vaccinium myrtillus (cf.

Kujala 1962: 177, Laine 1970(II): 127).

Along the river shores it often forms almost

pure stands. Also in Inari Lapland, Cornus

is an important member in the submaritime

birch heath forests (Hämet-Ahti 1963a: 76).

In the alpine belt of Pechenga it grows in

e.g. Empetrum-Myrtillus-Stereocaulon

association (Kalliola 1939: 213) and in

moss-rich Empetrum-Myrtillus association

(Kalliola 1939: 220), but is also a member

of the Trollius-Geranium association

(Kalliola 1939: 111). In the alpine belt the

common associates include Anthoxanthum

alpinum, Phyllodoce caerulea and

Vaccinium myrtillus.

Kontuniemi (1932: 24) describes the

flowering and seed production (cf. also

Sylvén 1906: 136, Valle 1930, Söyrinki

1939a: 308, Hustich 1940c: 58). In Inari

Lapland, Cornus normally begins flowering

during the first days of July (cf. Valle l.c.

and Hustich l.c.). The earliest date for

flowering near the Kevo Subarctic

Research Station was on June 7 in 2002

(Alanen 2007).

The ecology of Cornus suecica should

be studied more in detail. It is quite

possible that there are various ecological

races. According to Söyrinki & Saari

(1980: 117) it mainly occurs on N facing

rocky slopes, while Dahl (1934: 373)

mentions that it prefers sunny sides.

Furthermore, in Finland it generally prefers

acid substrate and avoids eutrophic areas

(Söyrinki & Saari l.c.). According to

Karlsson (1973: 86) and Nilsson (2000:

150) it is clearly or slightly acidocline,

whereas Arwidsson (1943: 227) and

Wistrand (1962: 127) considered it possibly

somewhat calciphilous. Amphicline in Inari

Lapland (Laine 1970(II): 127, Mäkinen &

Kallio 1979: 17). Amphicline.

Morphology. A rare red-“flowered”

form occurs, in which the uppermost leaves

have a reddish tinge (Inari: Virtaniemi,

Kantojärvi, 7658:3567, YME 4653).

Dependence on culture. Cornus

suecica is a common apophyte (e.g.

Vanhatalo 1965: 122, Ahti & Hämet-Ahti

1971: 67). In Inari Lapland it is most

abundant in the grove-like zone between

birch forests and old hemerobic

semicultural meadows, avoiding drier

habitats. Hemeradiaphore (cf. Kujala 1964:

77).

YM

APIACEAE

Aegopodium podagraria L.

Introduced, very rare

Map 33

Distribution. Europe and adjacent Asia, introduced

in North America (Hultén & Fries 1986: 1099, map

1395; cf. Roweck 1981: 339). Common in S Sweden

and S Finland, rare – scattered elsewhere, in the

extreme north only occasional (Kujala 1964: 77,

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 49

Hultén 1971a: map 1328, Hämet-Ahti et al. 1998:

319, Mossberg & Stenberg 2003: 433).

Rare in Troms (Benum 1950, 1958: 302,

Engelskjøn & Skifte 1995: 143), very rare in

Finnmark (e.g. Gamvik and Sør-Varanger: Lid & Lid

2005: 583). Very rare also in Pechenga and the Kola

Peninsula (H, KPABG, Mäkinen 2002: 21) and in the

Kovda area (Sokolov & Filin 1996: 119). Very rare

in Sompio Lapland, Kittilä Lapland and Enontekiö

Lapland as well as in Koillismaa (Ahti & Hämet-Ahti

1971: 68, Hämet-Ahti et al. 1998: 319, Lampinen &

Lahti 2018).

InL ref. InL 0 %. H 1, TUR 1 spec.

Very rare (4; 0.001). Inari: I (4;

0.001). (1) Ivalo centrum, garden waste

dumping place in a wood (7619:3521, 2006

M. Piirainen, H-Arch., Kastikka doc.

606880); (2) Virtaniemi, brook side at the

Russian border, deciduous grove

(7647:3558, 2001 M. Piirainen 4656, H

733441 and 2001 S. Keränen, H-Arch.,

Kastikka doc. 388725); (3) Kaamanen, as a

weed in an old garden at Thule (7668:3507,

2004 S. Heino & U. Laine, TUR 580417);

(4) Sevettijärvi cemetery (7715:3562, 1977

J. Jalas, H-Arch., Kastikka doc. 436624).

Southern hemerochore.

FMF 0.015.

Vertical distribution. c: I (4; 0.001).

Range ca. 100 m (Sevettijärvi) – 150 m

(Thule). Silvine.

Ecology. Only found as sterile in Inari

Lapland; similarly also in the Kovda area

(Sokolov & Filin 1996: 119). Probably

arrived as a garden weed and persisting

with the long horizontal rhizomes.

Dependence on culture. Epoikophytic

anthropochore.

YM

Angelica archangelica L. subsp.

archangelica

Archangelica norvegica Rupr.

Archangelica officinalis Hoffm.

Indigenous, rather rare

Map 34

Distribution. Angelica archangelica L. is Eurasiatic,

growing in N and E Europe, N Asia and the

Himalaya (Fl. Eur. 2: 357, Fl. Nord. 6: 183). It is

divided into two subspecies, subsp. archangelica and

subsp. littoralis (Wahlenb.) Thell. In Scandinavia the

nominal subspecies is earlier understood to occur

mainly in the Scandes, extending to S Norway

(Hultén 1971a: map 1344, Roweck 1981: 344, Hultén

& Fries 1986: map 1422, Mossberg & Stenberg

2003: 444), but according to Fl. Nord. (6: 184) the

distribution is wider than earlier considered, reaching

the southernmost Sweden and Denmark. In the

northernmost provinces of Fennoscandia only subsp.

archangelica is known (Fl. Nord. l.c.); in N Finland

it is common – rare down to the Arctic Circle

(Hämet-Ahti et al. 1998: 322, Lampinen & Lahti

2018).

In N Norway fairly common in Troms (Benum

1958: 303, Engelskjøn & Skifte 1995: 144), in

Finnmark fairly common – scattered (Dahl 1934:

372); 5 localities in the Rastigaissa area (Ryvarden

1969: 33). Scattered – fairly rare in the Kola

Peninsula (Fl. Murm. IV: map 88, Pobedimova et al.

1959: 589, Alm et al. 1998: 135, Mäkinen 2002: 21).

Also in Pechenga (Valle 1933a, Kvist 1978: 53);

common in the Kovda area (Sokolov & Filin 1996:

120).

In N Finland rare – locally common in Sompio

and Kittilä Lapland (Wainio 1891: 53, Hustich

1940c: 58, Auer 1944, Kotilainen 1951: 82, Kujala

1961, Montell 1962: 124, Ulvinen 1962, Virtanen

1990, Lampinen & Lahti 2018), common in most of

Enontekiö Lapland (Lindén 1943: 74, Laine 1958:

86, Piirainen & Piirainen 1991b, Piirainen 1996b,

Hämet-Ahti et al. 1998: 322, Lampinen & Lahti l.c.).

According to Rintanen (1967: 203), A. archangelica

has spread to N Finland from two directions, from

NW and NE. In Inari Lapland, the migration from

NW reached only the Ivalojoki valley, the rest is due

to migration from the Kola Peninsula.

InL ref. ”Frequentissime” (Fellman

1835: 255). Fairly frequent close to the

rivers, but often scantily (Kihlman 1884:

105). Ivalojoki fairly common (Wainio

1891: 53), Ivalojoki and Tolosjoki, Sotajoki

(Mikkola 1941: 32). NE Utsjoki fairly

50 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

common – scattered (Laine et al. 1955: 130,

Kallio & Mäkinen 1957: 26). Vaskojoki

(Klockars & Luther 1938, Laine 1964:

114), Sotajoki and Laanila (Rintanen 1967:

203), and the uppermost Kietsimäjoki

(7613:3422, Kulmala 1999).

Kevo 23.7 %, InL 38 %, 117 sq. H 19,

JYV 1, OULU 3, TUR 22, YME 4 spec.

Rather rare (728; 0.108). Inari: II

(253; 0.057), Utsjoki: IV (475; 0.209).

Difference***. Very clearly distributed

mainly along the large rivers:

Pulmankijoki, Teno, Vetsijoki, Utsjoki,

Kevojoki in Utsjoki commune, and

Näätämöjoki, Inarijoki-Kietsimäjoki and

Ivalojoki in Inari commune. Almost totally

missing in the E and SE parts of Inari; the

only exception is the mouth of the brook

Möyryoja in SE Lake Inari area (2 km E of

Kultalahti, 7651:3549), where the species

grew at the remains of a war-time building

and was probably not originally native.

The species is most frequent and

abundant in the Kevo Strict Nature

Reserve, which offers plenty of suitable

brook sides. The species avoids high alpine

areas, which are often too dry for the

species. On the other hand, it is also absent

in large open swamp areas. Northern.

FMF 0.449.

Vertical distribution. a: II (41;

0.057), b: IV (448; 0.194), c: III (239;

0.065). Differences: a-b***, b-c***. Range

15 m (Lake Pulmankijärvi, 7763:3539) –

475 m (S Paistunturit SW of Kuivi, the

uppermost Kamajohka by

Kamajohkeädsaoaivi, 7725:3476). Tr 1070

m ( Norman I(1): 521, Engelskjøn & Skifte

1995: 144), Fnm 595 m (Norman I(1):

522), EnL 900 m (Laine 1958: 86, Väre &

Partanen 2009: 132). Mainly silvine, and

most common in the birch belt.

Ecology. The most typical habitats

especially in the subalpine birch belt are the

inundation zones along rivers and brooks,

as well as the margins of springs. Along the

rivers the species often grows in the shelter

of big stones right at the water margin, but

it often also proceeds a few tens of meters

into the inundated grove-like, luxurious

birch forest. The habitats normally offer a

rich supply of electrolytes, which is

necessary for the plant. The most common

associates in the Kevojoki area are Bartsia

alpina, Calamagrostis phragmitoides and

Trollius europaeus, with the mosses

Plagiomnium ellipticum, Rhytidiadelphus

triquetrus and Sanionia uncinata (Laine

1970(II): 127). Also willows (Salix glauca,

S. phylicifolia, S. lapponum) may provide

the mechanical shelter which A.

archangelica may need.

In the spruce area of the southernmost

part of Inari Lapland, the species prefers

“hollows and valleys dominated by the

spruce” (Rintanen 1967). Occasionally the

species proceeds into the alpine belt, often

growing right at the snow-beds and small

brooks (cf. Söyrinki 1939a: 304 in

Pechenga).

Fairly seldom A. archangelica may

proceed on the brook-like margins along

highways and roads, and very seldom it is

found on gravelly waste fields (e.g. in Inari

village, 7647:3501, YME 1370); on such

habitats it is very low (at most 20-30 cm)

and mostly remains sterile.

Söyrinki (1939a: 304) has described

the flowering, the ripening and the

distribution of the seeds in the alpine belt of

Pechenga. He states that the circumstances

for the generative reproduction are

favorable; this also holds true for the

occurrence in the subalpine belt in Inari

Lapland. Ojala (e.g. 1984, 1986a, b) has

studied in detail the variation, reproduction

and the life history strategies of the species.

Ca-indifferent or weekly calciphilous

(Arwidson 1943: 228, Benum 1958: 303,

Wistrand 1962: 129, Nilsson 2000: 152), in

Inari Lapland amphicline (Laine 1970(II):

127) or basocline (Mäkinen & Kallio 1979:

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 51

17). As a rule the pH value of the habitats is

over 5. Slightly basocline.

Morphology. The chromosome

number 2n=22 has been counted from two

populations in Utsjoki (Uhtsa Skallovarri

7747:3506 and Padosjohka 7754:3477,

Ojala 1986b).

Dependence on culture. The species

is an old vegetable and medicine plant,

which was collected in the nature by the

Lapps and also cultivated in house gardens

(Rosberg 1891: 38, Mehus 1969, Nickul

1970: 96). In Norway, the oldest

regulations concerning its cultivation date

back to the year 1154 (Kallio et al. 1978).

With the increase of the cultivation of

“modern” vegetables its use has declined,

but it is still known in every Lapp house as

“Olbmoporramrassi” (“an edible plant for

man”, cf. Häyrén 1925, Parvela 1931: 74,

77, 78, 1932: 12, Rintanen 1967, Kallio et

al. 1978). Young leaves and inflorescences

have been collected and used (raw) as a

salad, or mixed with warm reindeer milk.

Leaves and roots (Radix angelicae)

include proteins, vegetable oils and

minerals, especially potassium, in

abundance (Isotalo 1971: 33, Table 7,

Taskinen & Nykänen 1975). Piippo (2005a:

187) lists several tens of diseases which

have been cured with Angelica extract, and

likewise presents a great number of useful

chemical compounds which have been

identified in Angelica juice. The species

was cultivated in the Pakatti Experimental

Garden, Kittilä in 1975-1977 (Kallio et al.

1978); in central Europe it is commonly

cultivated (Piippo 2005a: 186).

Hemeradiaphore, occasionally archaeo-

phytic.

YM

Angelica sylvestris L.

Introduced, very rare

Map 35

Distribution. Eurasiatic, rare – occasional in the

north of Fennoscandia (Hultén 1971a: map 1347,

Roweck 1981: 342, Hultén & Fries 1986: map 1421,

Hämet-Ahti et al. 1998: 322, Mossberg & Stenberg

2003: 444).

Fairly common in Troms and W Finnmark,

absent e.g. in Sør-Varanger (Dahl 1934: 371, Benum

1958: 303, Engelskjøn & Skifte 1995: 144, Alm et al.

2000c). Rare – scattered in Pechenga and in the Kola

Peninsula (Mikkola 1941: 32, Fl. Murm. IV: map 87,

Pobedimova et al. 1959: 589, Alm et al. 1998: 135;

Mäkinen 2002: 21 e.g. E of Kandalaksha, scattered

along the Kolvitsa). Scattered in Keret Karelia

(Söyrinki 1956: 28), rare in the Kovda area (Sokolov

& Filin 1996: 120).

Rare but locally common in Sompio and Kittilä

Lapland, very rare in Enontekiö Lapland (Hjelt &

Hult 1885: 130, Wainio 1891: 53, Auer 1944,

Montell 1945a, Kotilainen 1951: 134, Kujala 1964:

77, Hämet-Ahti et al. 1998: 322, Lampinen & Lahti

2018; H, OULU, TUR). Scattered – common in

Koillismaa, e.g. common in the Oulanka National

Park (Paatela 1953: 65, Ahti & Hämet-Ahti 1971: 68,

Söyrinki & Saari 1980: 118).

InL ref. InL 2 %, 3 sq. TUR 1, YME 1

spec.

Very rare (4; 0.001). Inari: I (3;

0.001), Utsjoki: I (1; 0.000). Inari: (1)

Ivalo, bus station (7619:3522, 2006 H.

Väre, H-Arch., Kastikka doc. 529039); (2)

Virtaniemi, along the main road 1 km SW

of the Frontier Guard Station, an old war-

time encampment site, by small ponds, one

sterile specimen (7645:3557, 1965 Y.

Mäkinen, field list, YME 1377); (3)

Kaamanen, Kaamas-Aitta camping area,

shore of the Kaamasjoki, one flowering

specimen with Silene dioica (7669:3507,

1965 U. Laine, TUR 127933-4). Utsjoki:

(4) Mieraslompolo, one specimen in the

house yard (7723:3508, 1973 P. Kallio,

KEVO). Southern hemerochore.

FMF 0.015.

Vertical distribution. c: I (4; 0.001).

Range 120 m (Ivalo) – 150 m (Kaamanen).

Tr 789 m (Norman I(1): 517, Engelskjøn &

52 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Skifte 1995: 144), Fnm 346 m (Norman

I(1): 518). Above the forest line in N

Sweden (Roweck 1981: 342). Silvine.

Ecology. The species has been found

both on recent and war-time garbage dumps

with, e.g., Anthriscus sylvestris, Glechoma

hederacea, Heracleum sibiricum,

Pimpinella saxifraga (7645:3557), and with

Cirsium palustre (7723:3508). The native

habitats, which are missing in Inari

Lapland, are often considered eutrophic and

the species somewhat calciphilous (Pesola

1928: 94, Arwidsson 1943: 227, Kotilainen

1951: 134, Ahti & Hämet-Ahti 1971: 68),

whereas Nilsson (2000: 151) considers the

species indifferent or weakly calciphilous,

and Wistrand (1962: 128) Ca-indifferent.

Dependence on culture. Epoikophytic

anthropochore, partly polemochore.

YM

Anthriscus sylvestris (L.) Hoffm.

Chaerophyllum silvestre L.

Indigenous and introduced, rare

Map 36

Distribution. Europe, N Asia, N Africa

(Dorogostaiskaya 1972: 126, SKK III: 203, Hultén &

Fries 1986: map 1388). Very common over most of

Fennoscandia, in Finland common up to the Arctic

Circle (Hultén 1971a: map 1312, Roweck 1981:

335).

In Troms a very common weed around

inhabited places (Norman I(1): 529, Benum 1958:

300, Engelskjøn & Skifte 1995: 143). In Finnmark

mostly on the coast (Dahl 1934: 372); Neidenfjord

(Kihlman 1884: 61), Laevvajokka mouth (Ryvarden

1969: 33); ca. 30 localities in Sør-Varanger and the

Varanger Peninsula (Zizka 1985: 38, 87). Native or

an established anthropochore in Pechenga (Parvela

1930: 74), Valle 1931, Kontuniemi 1932: 22, Kvist

1978: 53, Alm et al. 1997: 29, 41). In the Kola

Peninsula scattered in the population centers (Fl.

Murm. IV: map 82, Mäkinen 2002: 21). In Kovda

area fairly common (Sokolov & Filin 1996: 118), in

Keret Karelia scattered (Söyrinki 1956: 28).

In N Finland rather rare – scattered – fairly

common in Sompio and Kittilä Lapland, rare in

Enontekiö Lapland (Hjelt & Hu1t 1885: 130, Wainio

1891: 53, Linkola 1929, Parvela 1932: 12, 78,

Hustich 1936b, 1940c: 58, Auer 1944, Paatela 1953:

64, 81, Laine 1958: 85, Montell 1962: 124, Ahti &

Hämet-Ahti 1971: 67, Piirainen & Piirainen 1991b,

Hämet-Ahti et al. 1998: 316, Lampinen & Lahti

2018). Common in Koillismaa, also as a native plant

in the valley of the Oulankajoki (Söyrinki & Saari

1980: 117).

InL ref. Ivalojoki, e.g. Kuttura,

Härkäsaari, Tolosenniitty, Törmänen, Ivalo,

mouth of the river (Wainio 1891: 53,

Mikkola 1941: 32, Kujala 1962: 177). A

few localities in W Utsjoki (Laine et al.

1955: 130, Kallio & Mäkinen 1957: 26);

numerous localities both in Inari (Linkola

1929, Helander 1965: 65) and in Utsjoki

(Vanhatalo 1965: 122). According to

Helander so common in the Inari village

that it is impossible to list the localities

separately.

InL 16 %, 12 sq. H 5, JYV 1, OULU 1,

TUR 11, YME 5 spec.

Rare (141; 0.021). Inari: II (116;

0.027), Utsjoki: I (25; 0.009).

Difference***. Clearly commoner in the

south, where it often occurs abundantly in

every village on courtyards, field margins

and garbage heaps, also on grass lawns.

Since 1960, the abundance has been

increasing in most of the localities. In

Utsjoki still missing in some smaller

villages. Indigenous and southern

hemerochore; in a few Utsjoki localities of

northern origin.

FMF 0.249.

Vertical distribution. b: I (20; 0.007),

c: II (121; 0.034). Difference***. Range 20

m (Lake Pulmankijärvi, Niemelä,

7770:3537) – 360 m (Lake Kulpakkojärvi,

yard meadow by the ruins of an old hut,

7615:3489). Tr 830 m (Norman I(1): 529,

Engelskjøn & Skifte 1995: 143), Fnm 521

m (Norman I(1): 531), EnL 850 m (Laine

1958: 86). In Pite lappmark to 630 m

(Arwidsson 1943: 227). Silvine.

Ecology. The species has most1y

arrived to yards and hay fields from the

south with hay seed; it favors sites with rich

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 53

nitrogen and calcium supply, e.g. compost

heaps and walls of old cattle sheds (cf.

Pesola 1928: 95, Hustich 1936b, Wistrand

1962: 127). In the yard of Thule Anthriscus

covered large areas in 1965 (Helander

1965: 65).

In Pechenga the seeds as a rule

germinate well and produce seedlings

(Kontuniemi 1932: 22). The seeds are also

spread by the birds (Dahl 1934: 372).

The species may be indigenous in a

few riverside localities in Inari; there is one

apparently native locality along the upper

Lismajoki (E end of Lismajärvi,

7592:3428): a very luxurious riverside

meadow without any observable human

activity; the species grows there with e.g.

Elymus caninus, Milium effusum, Lactuca

sibirica and Polemonium acutiflorum.

According to Zizka (1985: 38) the species

favors oceanic climate. Basocline (Mäkinen

& Kallio 1979: 17), calciphilous in natural

localities (Nilsson 2000: 150). Basocline.

Parasites. The rust Puccinia

chaerophylli Purton has been found in

Norway close to the Finnish border on the

shore of the Tana between the villages

Polmak and Horma (Mäkinen 1964b: 166).

Morphology and taxonomy. It has

been suggested that in Fennoscandia there

are two, or even several races, which are,

however, morphologically inseparable (cf.

Björkman 1939: 20, Hustich 1940c: 58,

Wistrand 1962: 127, Nilsson 2000: 151).

Their responses to human activities may be

different. The specimens found by T. Laine

in Enontekiö at 850 m were flowering and

over 1 m in height (Laine 1958: 86).

Dependence on culture. Anthriscus

has been cultivated in flower beds in

Muonio and Pechenga (Parvela 1931: 74,

1932: 12, 78). In the Kuusamo district it is

also a polemochore (Ahti & Hämet-Ahti

1971: 67). Probably most of the

occurrences around Nellimö area are of

polemochorous origin; several of them have

been found on abandoned military camps. It

is also probable that raising hay has

strongly increased the frequency of

Anthriscus (Helander 1965: 65). In Laiti,

Pajuranta (7761:3505) the species is

reported to grow since the beginning of

World War II (Vanhatalo 1965: 122).

Established anthropochore, partly

polemochore; rarely native.

YM

Carum carvi L.

Introduced, very rare

Map 37

Distribution. Originally Eurasiatic, now circumpolar

(introduced in N America; Hultén & Fries 1986: map

1414, Hultén 1971b: 218, Dorogostaiskaya 1972:

128, SKK III: 214, Roweck 1981: 341).

Fennoscandian lowland area up to the Arctic Circle,

scattered – rare in the extreme north (Benum 1958:

301, Hultén 1971a: map 1324, Engelskjøn & Skifte

1995: 143), decreasing in frequency in Finnmark

towards east (e.g. Elvenes, Vardø, Sør-Varanger;

Norman I(1): 510-511, Dahl 1934: 370, Zizka 1985:

39, map 21).

Rare in Pechenga (H; Linkola 1929, Valle

1933a: f. atrorubens), the Rybachy Peninsula

(Pobedimova et al. 1959: 589) and the Kola

Peninsula (Fl. Murm. IV: map 83, Alm et al. 1997:

41, Mäkinen 2002: 21 locally even very abundant). In

Kovda area fairly common (Sokolov & Filin 1996:

119; cf. Fellman 1831: 306).

Mostly rare but established in Sompio and

Kittilä Lapland, very rare and casual in Enontekiö

Lapland (Hjelt & Hult 1885: 130, Wainio 1891: 54,

Hult 1898: 164, Montell 1910, 1945a, 1962: 125,

Linkola 1929, Parvela 1932: 36, Ahti & Hämet-Ahti

1971: 68, Hämet-Ahti et al. 1998: 321, Lampinen &

Lahti 2018).

InL ref. Ivalojoki, Tolosenniitty

(Kujala 1962: 177). 4 localities in Ivalo and

Inari villages (Helander 1965: 66), 6 in

Utsjoki (Vanhatalo 1965: 122).

InL 7 %, 16 sq. H 1, OULU 1, TUR 6,

YME 3 spec.

Very rare (29; 0.004). Inari: I (24;

0.006), Utsjoki: I (5; 0.002). Difference*.

Mainly in old villages around Lake Inari.

54 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Well established in most of its places, but

does not seem to proceed to new localities,

possibly due to the weak production or poor

germination of seeds. Also Wistrand (1962:

128) states that it seldom occurs as a

neophyte. Southern (in Utsjoki possibly

also a northern) hemerochore.

FMF 0.079.

Vertical distribution. b: I (3; 0.001, c:

I (26; 0.007). Difference***. Range 25 m

(Nuorgam, 7778:3533) – 310 m (Laanila

Exp. Station, 7590:3516). Tr 500 m

(Engelskjøn & Skifte 1995: 143), Fnm 124

m (Norman I(1): 510). Silvine.

Ecology. In most of the localities in

Utsjoki, Carum is a relic of old distribution,

and in any case spread before the World

War II. In Nuorgam (7778:3533), it grows

at the chapel with e.g. Alchemilla

murbeckiana, Capsella bursa-pastoris and

Gentiana nivalis, and at the Sarja farmstead

(7766:3539; on the shore of Lake

Pulmankijärvi) with Achillea millefolium

and Capsella; in both places,

Tripleurospermum maritimum subsp.

phaeocephalum belongs to the associates.

In Inari, the species occurs in the yards of

several Lapp houses, also on the ruins of

houses burned during 1944-1945. Near

Ivalo (7624:3533), Carum was found along

the Veskoniemi road, as a weed in a

timothy cultivation, with Silene dioica and

Trifolium pratense.

Only in a few cases, the species has

spread further on the sides of recent roads.

Carum is cultivated in a few places and is

able to remain for tens of years. In Muonio,

the species is probably everywhere of

cultivated origin (Montell 1962: 125; cf.

Roweck 1981: 342).

Dependence on culture. Partly spread

during the World War II. Epoikophytic

anthropochore (cultivated and escaped),

partly polemochore.

YM

Chaerophyllum prescottii DC.

C. bulbosum L. subsp. prescottii (DC.)

Nyman

Introduced, very rare

Map 38

Distribution. Central and E Europe, especially

Russia (Hultén 1971a: map 1309, Hultén & Fries

1986: map 1386). Nowadays absent in Swedish

Lapland (Roweck 1981: 335) as well as in Troms and

Finnmark (Mossberg & Stenberg 2003: 428). A few

localities in the Kola Peninsula (Fl. Murm. IV: map

82); scattered in Keret Karelia (Söyrinki 1956), and

one locality along the Lutto (Roivainen 1923: 292).

Whole Finland (Hämet-Ahti et al. 1998: 316),

but very rare and casual in the S parts, established

only in the north. In Koillismaa and Sompio Lapland

a common weed in villages (Wainio 1891: 53,

Linkola 1929, Hämet-Ahti 1967b, Ahti & Hämet-

Ahti 1971: 67, Hämet-Ahti et al. 1998: 316,

Lampinen & Lahti 2018; H, OULU, TUR). Scattered

in Kittilä Lapland, very rare in Enontekiö Lapland

(Hjelt & Hult 1885: 130, Hult 1898: 164, Montell

1945a, 1948, 1962: 124, Kujala 1961, Lampinen &

Lahti l.c.; H, TUR).

InL ref. Ivalo, and Laanila Tourist

House (Linkola 1929, Mikkola 1941: 32).

Ivalo, two localities (Helander 1965: 64).

InL 2 %, 5 sq. H 3, JYV 1, TUR 6,

YME 4 spec.

Very rare (8; 0.001). Inari: I (8;

0.002). (1) Laanila Tourist House

(7589:3516, 1925 E. Mikkola, TUR

75304); (2) Laanila Experimental Station

(7590:3516, 1964 T. Ahti, H 409722); (3)

Ivalo, Koppelontie, ditch in a potato field at

Ilokyrö (7620:3522, 1961 E. Tourunen,

TUR 73305) and near the “Casket House”

(7620:3522, Helander 1965: 64); (4)

Nellimö, Koskela house by the river, small

stands in several locations (7641:3553,

1971 Y. Mäkinen 71-553, TUR 185775,

YME 1427) and ca. 50 specimens in a tight

stand on a waste riverside slope (2000 Y.

Mäkinen 00-491, YME 26115); (5)

Muddusniemi, abundantly on a grassy

meadow at the ruins of an old farmstead,

with numerous other anthropochores

(7664:3500, 1968 P. Kallio & Y. Mäkinen

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 55

68-1246, TUR 162788, YME 1428); (6)

Valpurinniemi, a few plants by the wall of a

building (7664:3502, 2000 U. Laine & S.

Heino, TUR 361430, 361440); (7)

Toivoniemi, ca. 30 specimens, waste place

near the shore (7665:3504, 1968 Y.

Mäkinen 68-1234, YME 1429); (8)

Kaamanen, Thule, large stand near the road

(7668:3507, 1981 P. Kallio). – One

specimen from Inari without exact locality

(1951 A. Ojala, JYV 22243). Southern

hemerochore.

FMF 0.018.

Vertical distribution. c: I (8; 0.002).

Range 120 m (Ivalo area) – 310 m

(Laanila). Silvine.

Ecology. All the localities are

characterized by intensive and old meadow-

type cultivation. The associates include,

e.g., Carduus crispus, Dianthus superbus

and Silene vulgaris. Also in Keret Karelia

the distribution is in connection with old

and intensive agriculture. Amphicline

(Mäkinen & Kallio 1979: 17). Amphicline.

Dependence on culture. The species

is clearly an eastern newcomer, which has

also been cultivated since the end of the

19th century (Nordling 1884a: 307, 1884b:

314, Elfving 1897: 96). One of the early

cultivation sites is in Toivoniemi on the

shore of Lake Muddusjärvi (Parvela 1932:

39), where the species has survived at least

to the year 2000. However, it does not

show any tendencies to spread, e.g., there

are no localities along new roads. – Only

one locality is clearly due to Russian

military camps during the World War II (cf.

Heikkinen 1948, 1959). Epoikophytic

anthropochore, partly polemochore.

YM

Cicuta virosa L.

Indigenous, very rare

Map 39

Distribution. Eurasiatic (Hultén 1971b: 152, Hultén

& Fries 1986: map 1412; in Europe mainly in N and

E parts. Fairly common – scattered in S and central

Finland and Sweden, rare N of the Arctic Circle

(Hultén 1971a: map 1322, Mossberg & Stenberg

2003: 442, Roweck 1981: 340, Hämet-Ahti et al.

1998: 321).

Missing in Troms, two localities in Finnmark,

Kautokeino (Engelskjøn & Skifte 1995, Lid & Lid

2005: 582, Elven et al. 2013: 153). Rare in the Kola

Peninsula: a few localities in the Kirovsk area and on

the S coast (Fl. Murm. IV: 246: map 83). Scattered –

fairly common in the S parts of Sompio and Kittilä

Lapland, rare in the N and NE parts (Hjelt & Hult

1885: 130, Wainio 1891: 54, Hjelt 1911: 224,

Ruuhijärvi 1960: 352, Montell 1962: 124, Hämet-

Ahti et al. 1998: 321, Lampinen & Lahti 2018). In

Enontekiö Lapland only in the S parts (Hustich

1936a: 160, Lampinen & Lahti l.c.). Rather rare in

Koillismaa (Ahti & Hämet-Ahti 1971: 68, Hämet-

Ahti et al. l.c., Lampinen & Lahti l.c.), six localities

in Keret Karelia in Söyrinki (1956: 28), five more in

Kastikka (2018).

InL ref. Very rare along the Ivalojoki

near the mouth, not flowering (as var.

angustifolia, Kujala 1962: 177). Vaskojoki

area (Kallio 1961: 99, Laine 1964: 114).

Lower course of the Ivalojoki, Lake Alempi

Akujärvi (Siltanen 1967: 168).

InL 2 %, 5 sq. H 3, KUO 1, TUR 5,

YME 3 spec.

Very rare (14; 0.002). Inari: I (13;

0.003), Utsjoki: I (1; 0.000). Difference*.

Inari: (1) Ivalojoki, Umpimukka [?]

(possibly 761:352, 1932 K. Enwald, KUO

59536); (2) NE shore of Lake Ylempi

Akujärvi (7620:3529, 1963 Y. Mäkinen,

TUR 75408, YME 1433, 1973 C. E. Sonck,

TUR 259474; T. Rintanen, H-Arch.,

Kastikka doc. 412045, as var. angustifolia);

(3) mouth of the Ivalojoki S of Jänkälä

(7621:3524, 1962 P. Siltanen, field list); (4)

S shore of Lake Alempi Akujärvi

(7621:3527, 1962 Y. Mäkinen, YME

1434); (5) SE shore of Lake Alempi

Akujärvi (7621:3528, 1982 C. E. Sonck, H

56

REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Fig. 5. Cicuta virosa L. var. angustifolia (Kit.)

Wimm. & Grab. (Inari, Lake Ylempi Akujärvi, 1963

Y. Mäkinen, TUR 75408).

581973, determined as var. angustifolia,

det. P. Uotila 2009, TUR 280698); (6) N

shore of Lake Ylempi Akujärvi

(7621:3529, Rintanen 1976); (7) mouth of

the Ivalojoki, oxbone lake S of Vaarala

(7623:3524, 1962 P. Siltanen, field list,

1962 Y. Mäkinen, YME 26464); (8)

Mielikköjänkkä (7627:3528, 2001 Y.

Mäkinen & A. Rantio-Lehtimäki, field list;

2005 L. Laasonen, H 838061, determined

as var. angustifolia, det. M. Piirainen

2013); (9) Veskoniemi road, copious in a

wet meadow N of the N end of Lake Iso

Mielikköjärvi (7628:3528, 2001 Y.

Mäkinen, field list); (10) Veskoniemi,

fishing harbor (7633:3526, 2008 H. Väre,

H-Arch., Kastikka doc. 578238, as var.

virosa); (11) Vaskojoki, 4 km N of Lake

Paadarjärvi, twin ponds Peräkkäisjärvet SE

of Pahtavaara, five flowering specimens on

the W shore of the S pond (7647:3482,

1960 U. Laine, TUR 75409-75410,

determined as var. virosa, det. U. Laine

2010; 2008 H. Kaipiainen & H. Väre, H-

Arch., Kastikka doc. 578218, as var.

virosa); (12) Toivoniemi, Umpisuuvuopaja

– Mukkavuopaja, W side of Lake

Vastusjärvi, several flowering exx. in a

large wet meadow (7665:3503, 2000 S.

Heino, K. & U. Laine, TUR 361411-

361414, as var. virosa); (13) Lake

Aksujärvi (768:349, 1958 R. Ruotsalo, H

261127, determined as var. angustifolia,

det. P. Uotila 2009). Utsjoki: (14) Teno by

Kaivojoki, N of Rovisuvanto, springy bog

(7711:3455, 1987 J. & J. Lampolahti, field

list, no specimen). – The determination of

the varieties in the field lists without a

specimen must be considered uncertain.

Southern lowland.

FMF 0.029.

Vertical distribution. c: I (12; 0.003).

All localities in the coniferous zone. Range

120 m (Lake Akujärvi) – 235 m

(Peräkkäisjärvet). Silvine.

Ecology. All localities are in wet

meadows on shores of ponds, small lakes or

slowly flowing quiet waters of river loops;

they are permanently wet, and always

inundated in the spring. The most common

associates are Carex globularis, C.

rostrata, Eleocharis palustris, Equisetum

fluviatile, Galium palustre, Lysimachia

thyrsiflora and Potentilla palustris.

Cicuta virosa does not flower every

year, and easily remains undetected.

Roweck (1981: 340) points out that it may

also spread from rhizome pieces, loosened

by ice in the spring.

The species is usually regarded as

amphicline, Ca-indifferent or mesotrophic

(Pesola 1928: 110, Linkola 1933, Wistrand

1962: 128, Mäkinen & Kallio 1979: 17,

Roweck 1981: 340), rarely as favoring Ca

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 57

(Söyrinki & Saari 1980: 117). Slightly

basocline.

Morphology and taxonomy. Cicuta

virosa includes two varieties in

Fennoscandia, var. virosa and var.

angustifolia (Kit.) Wimm. & Grab.

According to Fl. Nord. (6: 208), var.

angustifolia is smaller in every respect and

has fewer umbellules and flowers than var.

virosa, and its stem and flowers are often

partly purplish.

In Fennoscandia var. virosa has a more

southerly distribution, while var.

angustifolia is prevailing in the N parts.

The latter is usually the only variety

reported in literature from Inari Lapland

(Kujala 1962: 177, Hämet-Ahti et al. 1998:

321, Mossberg & Stenberg 2003: 442, Fl.

Nord. 6: 208). However, according to

Kastikka (2018), Lampinen & Lahti (2018)

and specimens in H and TUR, both

varieties have been found there.

Most specimens from Inari Lapland

have fairly narrow leaf segments and

somewhat fewer flowers than plants from

S Finland, but the variation is considerable

(see Table 1). The specimen from Lake

Ylempi Akujärvi (Y. Mäkinen, TUR

75408, Fig. 5, specimen no. 1 in Table 1) is

a good representative of var. angustifolia,

while the plant from Lake Vastusjärvi (S.

Heino, K. & U. Laine, TUR 361411-

361414, Fig. 6, specimen no. 7 in Table 1)

is a typical var. virosa. Also the sterile

specimen in KUO from the Ivalojoki (1932

K. Enwald, KUO 59536) has fairly broad

leaf segments and may belong to var.

virosa. Other specimens in Table 1 are

Fig. 6. Cicuta virosa L. var. virosa, the top and the base of the specimen (Inari, Toivoniemi, 2000 S. Heino, K.

& U. Laine, TUR 361411, 361414).

58 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

more or less intermediate between the two

varieties. The variation in Inari Lapland is

obviously clinal and probably partly

environmental, but the material is too

scanty to draw further conclusions.

Cultivation experiments would be needed

to clarify the nature of the variation in the

species (see also Fl. Nord. 6: 207, Elven et

al. 2013: 154-155).

Dependence on culture. Hemera-

diaphore.

YM, JN

Table 1. Variation of Cicuta virosa var. angustifolia and var. virosa according to Fl. Nord. (6: 208) and in

seven specimens from Inari Lapland in H and TUR: 1. Lake Ylempi Akujärvi (TUR 75408), 2. Lake Alempi

Akujärvi (H 581973), 3. Lake Alempi Akujärvi (TUR 280698), 4. Mielikköjänkkä (H 838061), 5.

Peräkkäisjärvet ponds (TUR 75409), 6. Peräkkäisjärvet ponds (TUR 75410), 7. Lake Vastusjärvi (TUR

361411-361414).

var.

angusti‐

folia

var.virosa 1.2.3.4.5.6.7.

a)Stemlength,

cm

40–60 50–110 >42 >49 >36 >37 >53 >55 110

b)Leafblade,

length,cm

9.5–12 14–25 9.5 13 13 14.5 10 10 15

c)Leafblade,

breadth,cm

4–6 5–22 6 9 10 8 8 7.5 8.5

d)Apicallobe,

length,mm

17–44 30–88 30 34 32 37 28 40 70

e)Apicallobe,

breadth,mm

2–5 4.5–14 3.5 3 5 5 3 4 10

f)Apicallobe,

length/breadth

6.9–13 3.1–8.1 8.6 11.3 6.4 7.4 9.3 10.0 7.0

g)Umbellules

perumbel

8–21 13–32 13 19 18 19 23 24 28

h)Flowersper

umbellule

20–35 34–64 35 39 32 37 42 31 42

i)Colorofstem ±purple green p (p) g (p) (p) (p) g

j)Colorof

petals/anthers

±purple white/pale p p (p) (p) w w w

Totalrating

(seebelow)

0– –30 5.0 9.1 13.6 12.5 12.5 12.6 25.0

Rating for each characteristic has been calculated as X=3*(A-Bang)/(Bvir-Bang), where A =

value of a single characteristic a-h in each specimen (stem length estimated for specimens

1-6), Bang = value of an extreme var. angustifolia, and Bvir = value of an extreme var.

virosa. The values of Bang and Bvir used for each characteristic are: a) stem length: 40 cm

(Bang) – 110 cm (Bvir), b) length of leaf blade: 10 cm – 20 cm, c) breadth of leaf blade: 4 cm

– 15 cm, d) length of apical lobe: 20 mm – 70 mm, e) breadth of apical lobe: 2 mm – 10

mm, f) length/breadth ratio of apical lobe: 12 – 4, g) number of umbellules per umbel: 9 –

25, h) number of flowers per umbellule: 22 – 55, i) color of stem: purple (0) – green (3), j)

color of petals and anthers: purple (0) – white or pale (3). The sum of the values received is

presented in the lowermost row. An extreme var. angustifolia specimen will receive a

value close to 0, and an extreme var. virosa a value close to 30.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 59

Heracleum sibiricum L. var. sibiricum

H. sphondylium L. subsp. sibiricum (L.)

Simonk.

Introduced, very rare

Map 40

Distribution. Europe, W Siberia (Benum 1958: 305,

Roweck 1981: 347, Hultén & Fries 1986: map 1429).

Common – fairly common in S Fennoscandia (esp. in

the east; Hultén 1971a: map 1354, Mossberg &

Stenberg 2003: 446, Fl. Nord. 6: 228). Rare –

scattered in N Finland (Montell 1945a, 1962: 125,

Ahti & Hämet-Ahti 1971: 68, Suominen 1979: 63,

Söyrinki & Saari 1980: 118, Parnela 1985, Hämet-

Ahti et al. 1998: 324).

Rare in Troms (Benum 1958: 305, Engelskjøn

& Skifte 1995: 145), in Finnmark e.g. in Alta and

Sør-Varanger (Dahl 1934: 372). Scattered in

Pechenga and the Kola Peninsula in towns and

villages, small stands often by the walls of buildings

(Fl. Murm. IV: map 89, Alm et al. 1997: 41,

Piirainen 1997d, Piirainen et al. 1997, Mäkinen 2002:

21). Rather rare in Sompio and Kittilä Lapland and

mainly in the S and W parts, a rare casual in

Enontekiö (Linkola 1929, Hämet-Ahti et al. 1998:

324, Fl. Nord. 6: 229, Lampinen & Lahti 2018; H,

TUR). Rare along the Lutto (Roivainen 1923: 292),

common in the Kovda area (Sokolov & Filin 1996:

120).

InL ref. Törmänen, two localities 1961

(Kujala 1962: 177). The oldest specimen

collected in Kaamanen, Thule (7668:3507,

1937 E. Häyrén, H 412787). “Typical in the

yards and waste heaps of almost every

house in Inari commune” (Helander 1965:

66).

InL 4 %, 13 sq. H 3, OULU 2, TUR

10, YME 8 spec.

Very rare (33; 0.005). Inari: I (32;

0.008), Utsjoki: I (1; 0.000). Difference***.

The observation from Utsjoki

(Karigasniemi, riverside in the village

center, 7702:3454, 1987 J. & J. Lampolahti,

field list) is unfortunately without a

specimen. Southern hemerochore.

FMF 0.059.

Vertical distribution. c: I (33; 0.009).

Range 120 m (Inari village, 7647:3501) –

320 m (Saariselkä, by the S road to the

resort village, 7592:3516). Tr 80 m

(Engelskjøn & Skifte 1995: 145). Silvine.

Ecology. The largest stand is probably

in Ivalo, W end of the Tourist Hotel, partly

on a sandy yard, partly on a grassy

meadow. In 2007 the stand covered an area

of ca. 12 m². All the localities are on

cultural ground, but Kujala (1962: 177)

found the species also among shore shrubs

along the Ivalojoki in Törmänen.

Morphology and taxonomy.

Heracleum sibiricum is currently included

in H. sphondylium as a subspecies H.

sphondylium L. subsp. sibiricum (L.)

Simonk. (Fl. Nord. 6: 227, Elven 2016, see

also Fl. Eur. 2: 365). Specimens with setose

fruits (referred to as var. chaetocarpum H.

Neumayer & Thell., Hämet-Ahti et al.

1998: 324) have not been found in Inari

Lapland; all specimens collected belong to

var. sibiricum.

Dependence on culture. According to

many local inhabitants, the species arrived

in the area during the World War II (cf.

Helander 1965: 66). It is a typical

polemochore, which, however, is still

actively spreading mainly along roadsides

(further south it spreads extensively along

railroads, Suominen 1979: 63). Along the

Nellimö road by the Mustola saw-mill

(7633:3546) it occurred in 1962 with

Anthriscus sylvestris, Carum carvi,

Galeopsis speciosa, Silene dioica,

Trifolium pratense and T. repens. Also in

Muonio village, Kittilä Lapland, the species

has appeared during the World War II.

Roweck (1981: 348) describes the nature of

the hemerochorous occurrences in Swedish

Lapland. Epoikophytic anthropochore,

mainly polemochore.

YM

60 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Heracleum sphondylium L. s. str.

H. sphondylium L. subsp. sphondylium

Not accepted for Inari Lapland

Not mapped

Distribution. W and C Europe – N Africa (Fl. Eur.

2: 366, Fl. Nord. 6: 228). Fairly rare to scattered in

the S parts of Fennoscandia (Fl. Nord. l.c.), not

mentioned from the Kola Peninsula in Fl. Murm. IV

or Ramenskaja & Andreeva (1982). In Finland a rare,

established anthropochore in the south, and as a

German polemochore in the N part of the country in

Tornio, Hyrynsalmi and Kuusamo (Fl. Nord. l.c.).

Part of the records in Lampinen & Lahti (2018) from

N Finland are dubious due to the fact that H.

sphondylium and H. sibiricum have often been

regarded as conspecific and the exact identity of

other than specimen data is very uncertain.

InL ref. Heracleum sphondylium s. str. is

mentioned as a rare casual in Inari Lapland in Hämet-

Ahti et al. (1998: 324) and mapped with two dots

(760:351, 765:349) in Lampinen et al. (2015 and

earlier versions of the atlas), dots removed in

Lampinen & Lahti 2016. The information was based

on two specimens collected in Inari in 1963. The

other specimen from “Törmänen, 100 m E from

Niemelä, field margin” (correctly 7614:3519, 1963 T.

Niemelä, OULU 47991) was determined as H.

sibiricum by L. Fröberg in 2009. We have not seen

the other specimen ("Inari, in ruderatis", 1963 P.

Jokela, OULU, information according to Kastikka

doc. 98505856) and we are not aware of its possible

new determinations. Without the specimen, the

species is here not regarded as belonging to the flora

of Inari Lapland (cf. also Fl. Nord. 6: 228).

MP

Pastinaca sativa L. subsp. sativa

Introduced, very rare

Map 41

Distribution. Originally European (Hultén & Fries

1986: map 1428, Hämet-Ahti et al. 1998: 323,

Mossberg & Stenberg 2003: 446). Rare in S Finland

as an escape, but established and locally common in

the southernmost provinces (Hultén 1971a: map

1352).

Rare in Troms and Finnmark (Benum 1950,

1958: 304, Roweck 1981: 347); Sør-Varanger (Lid &

Lid 2005: 592). The northernmost occurrences in

Finland in Outer Ostrobothnia according to Hämet-

Ahti et al. (1998: 323) and Lampinen & Lahti (2018),

but recorded also in Inari Lapland (Mäkinen & Kallio

1976: 17).

InL ref. Recorded as an introduced

casual in Inari (Mäkinen & Kallio 1976: 17,

37).

InL 0 %.

Very rare (1; 0.000). Inari: I (1;

0.000). Gravelly roadside between

Törmänen and Kerttuoja, a few rosettes

(7614:3521, 1971 Y. Mäkinen, pers.

comm.). Southern hemerochore.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Alt. 135 m. Silvine.

Dependence on culture. Cultivated in

Inari (Elfving 1927: 152); according to

Nordling (1884a: 307, 1884b: 315) the

species grows well in Inari and produces a

fair yield. In Pechenga it thrives poorly

(Parvela 1931: 60). To the locality

mentioned above it has spread in

connection with the road improvement

works, with e.g. Carum carvi and

Hypericum maculatum as companions.

Further south, it may occur both as a

German and Russian polemochore

(Suominen 1979: 63). Ephemerophytic

polemochore.

YM

Petroselinum crispum (Mill.) Fuss

Introduced, very rare

Map 42

Distribution. SE Europe and W Asia (Hämet-Ahti et

al. 1998: 320). Cultivated almost throughout Finland,

rarely found as an escape from cultivation or casual

in dumps, mostly in the southernmost Finland

(Hämet-Ahti et al. 1998: 320, Lampinen & Lahti

2018).

InL ref. As an introduced casual in the

southernmost Inari (Mäkinen & Kallio

1976: 17, 37).

InL 0 %. YME 1 spec.

Very rare (1; 0.000). Inari: I (1;

0.000). Laanila, Laaninhovi, as a weed in a

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 61

flower bed (7590:3516, 1971 Y. Mäkinen

71-701, YME 1494). Not cultivated in this

locality. Southern hemerochore.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Alt. ca. 310 m. Silvine.

Dependence on culture. Parsley

thrives well in cultivation both in Inari and

Utsjoki (Grotenfelt 1897: 256, Ahola 1929,

Parvela 1932: 92). In the 2000’s the author

has seen parsley beds e.g. at the Utsjoki

vicarage, Kevo Station, Tsieskula garden,

in several localities in Kaamanen

(Jokitörmä camping area, Toivoniemi and

Thule; cf. also Parvela l.c.), and in several

house gardens in the centers of Inari and

Ivalo (incl. the School for Domestic

Sciences). The specimen collected in

Laanila was almost flowering, which

indicates that parsley was able to

overwinter and complete its 2-year life-

cycle there. According to Heikkinen

(1959), it has been found as a polemochore

in Hyrynsalmi, Kainuu. Ephemerophytic

anthropochore.

YM

Peucedanum palustre (L.) Moench

Not accepted for Inari Lapland

Not mapped

Distribution. Eurasiatic (Hultén & Fries 1986: map

1426). Common to scattered in S and central parts of

Sweden and Finland, in Norway in the southernmost

parts only (Hultén 1971a: map 1351, Mossberg &

Stenberg 2003: 445, Lid & Lid 2005: 591). Very rare

in the Kola Peninsula (Fl. Murm. IV: 267, map 89,

Hultén l.c.). The northernmost Finnish localities in

Kittilä and Sodankylä and one square in Inari

(Lampinen & Lahti 2018, see below).

InL ref. Included in the preliminary list of the

vascular plants of Inari Lapland as an introduced

casual, based on a single record (Mäkinen & Kallio

1979: 17, 38). However, no specimen exists. The

record is most probably either due to

misidentification or an erroneous note made in the

field. Unfortunately the record has been included in

the maps in Hämet-Ahti et al. (1984: 268, 1986: 285,

1998: 323).

There are also field notes of Peucedanum in H-

Arch. from six nearby localities in SW Inari, S part of

the Lemmenjoki National Park (7600-7608:3451-

3453, 1988-1990 S. Valtiala, Kastikka doc. 210000-

210001, 210540-210543). They are included in

Lampinen & Lahti (2018) and earlier versions of the

atlas. In the lack of specimens, the field notes are

here regarded as dubious. They probably represent

erroneous marks in the field lists (possibly displacing

Potentilla palustris, which is missing in all the lists in

question).

No specimens from Inari Lapland in H, JYV,

KUO, OULU, TUR, VOA or YME.

The species is not accepted in the flora of the

province, until further evidence is received.

YM

Pimpinella saxifraga L.

Introduced, very rare

Map 43

Distribution. Europe and W Asia (Benum 1958:

302, Hultén & Fries 1986: map 1394); introduced in

N America. In Fennoscandia mainly in the southern

and central parts (Hultén 1971a: map 1327, Roweck

1981: 338, Mossberg & Stenberg 2003: 433). In

Finland common up to ca. 65° N (Kujala 1964: 77,

Hämet-Ahti et al. 1998: 319).

Rare in Troms, very rare in Finnmark, e.g.

Neiden and Sør-Varanger (Norman I(1): 512, Benum

1958: 301, Vorren 1968, Engelskjøn & Skifte 1995:

143, Lid & Lid 2005: 583). Very rare in Pechenga

and the Kola Peninsula (Fl. Murm. IV: map 83,

Hultén 1971a: map 1327). Very rare in Sompio and

Kittilä Lapland, no records from Enontekiö Lapland

(Montell 1945a, 1962: 125, Hämet-Ahti et al. 1998:

319, Lampinen & Lahti 2018; H, OULU, TUR). Very

rare also in Keret Karelia (Söyrinki 1956: 28) and in

the Kuusamo district (Ahti & Hämet-Ahti 1971: 68,

Lampinen & Lahti l.c.).

InL ref. As an introduced casual in SE

Inari (Mäkinen & Kallio 1976: 17, 38,

Hämet-Ahti et al. 1998: 319).

InL 0 %, 1 sq. YME 1 spec.

Very rare (2; 0.000). Inari: I (2;

0.000). (1) Virtaniemi, ca. 1 km SW of the

Frontier Guard, old German military camp

area, ca. 10 sterile leaf rosettes by small

ponds (7645:3557, 1965 Y. Mäkinen, YME

62 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

1508); (2) Toivoniemi, meadow (766:350,

1897 A. W. Granit & B. R. Poppius, H

410290). Southern hemerochore.

FMF 0.007.

Vertical distribution. c: I (2; 0.001).

Range ca. 130 – 150 m. Tr 101 m (Norman

I(1): 512). Silvine.

Dependence on culture. Ephemero-

phytic polemochore.

YM

DIAPENSIACEAE

Diapensia lapponica L.

Indigenous, rather rare

Map 44

Distribution. Amphi-Atlantic, mainly arctic-

montane. NE North America and N Europe to N and

central Urals and Obi Bay, with an outpost in

Scotland (Roger 1952, Benum 1958: 315, Elven

2016). In Fennoscandia from the mountains of S

Norway and Härjedalen in Sweden northwards to N

Norway, N Finland and the Kola Peninsula (Hultén

1971a: map 1387, SKK III: 318, Gjærevoll 1990: 55,

Nilsson 2000: 152, Mossberg & Sternberg 2003:

450). The closely related, amphi-Pacific D. obovata

(F. Schmidt) Nakai in NE Asia and NW North

America is often regarded as a subspecies subsp.

obovata (F. Schmidt) Hultén (Hultén 1958: map 204,

1968: 736, Hultén & Fries 1986: 1102, map 1435).

Fairly common in Troms and Finnmark mostly

in mountain areas, but especially in Finnmark also in

the lowland along the coast (Norman I(2): 878, Dahl

1934: 381, Benum 1958: 315, map 435, Alm 1993d).

Very common in the Rastigaissa area (Ryvarden

1969: 34). In Pechenga fairly common, several sites

near Köngäs (Wainio 1891: 44, Söyrinki 1939a:

339), in the Lutto area rare – rather rare (Roivainen

1923: 293). In the Kola Peninsula fairly evenly

distributed in the northern parts along the coast, in

the inner parts especially in the area of the Khibiny

Mountains (Fellman 1831: 305, Fl. Murm. IV: map

110, Hultén 1971a: map 1387). Southwards in the

surroundings of Kandalaksha N of the Arctic Circle

and on the top of the fjeld Sallatunturi in Kuolajärvi

near the Finnish border (Wainio l.c., Pesola 1918:

245, Fl. Murm. l.c.). The southernmost locality in

Russian Karelia at 64°57’ N (Piirainen et al. 2005).

In Sompio Lapland only in the NE corner in the

fjelds of E Saariselkä (Hult 1898: 163, Rintanen

1968: 287, Lampinen & Lahti 2018). In NW Kittilä

Lapland in the fjelds Pallastunturit and Olostunturi,

the southernmost locality in Finland (Hustich 1940c:

59, Montell 1962: 125, Lampinen & Lahti l.c.). In

Enontekiö Lapland rare in the fjeld area of Pallas-

Ounastunturit in the south, but common – fairly

common in NW Enontekiö (Hustich l.c., Lindén

1943: 78, Laine 1958: 90, Virtanen & Väre 1990,

Kämäräinen 1998, Lampinen & Lahti l.c.).

InL ref. “In regionibus subalpinis et

collibus elevatis ad Utsjoki, Enare et

Enontekis frequens” (Fellman 1835: 252).

Common in the alpine belt, rarer in the

subalpine belt (Kihlman 1884: 111). Fairly

rare (five localities listed) in the

Lemmenjoki area (Klockars & Luther

1938). Three localities along the Teno

(Hustich 1942a). Scattered – fairly frequent

in W and SW Utsjoki (Laine et al. 1955:

130, Kallio & Mäkinen 1957: 26).

Scattered in the Peäldoajvi fjeld area in the

alpine belt, two localities in the birch belt

(Koivistoinen 1964: 59). Ten localities in

SE Inari, mainly in the fjelds of Saariselkä

(Rintanen 1968: 287). In the Kevo Strict

Nature Reserve concentrated in the S part

(Laine 1970(II): 133, Heikkinen & Kalliola

1990: 47).

Kevo 13.5 %, InL 35 %, 103 sq. H 40,

JYV 3, KUO 8, OULU 14, TUR 44, VOA

3, YME 4 spec.

Rather rare (641; 0.099). Inari: II

(174; 0.041), Utsjoki: IV (467; 0.214).

Difference ***. The distribution in Inari

Lapland concentrates to the N and W fjeld

areas including the Muotkatunturit fjelds,

and to the NE fjeld areas. In the southern

parts of Inari Lapland Diapensia occurs

more scarcely, and the sites are mainly

confined to the fjeld areas of Saariselkä and

Maaresta- and Viipustunturit. The species

is almost lacking in the basin of Lake Inari

and the boggy lowlands N of it.

In the vicinity of the Kevo Subarctic

Research Station, Diapensia is common in

the fjeld Jesnalvarri and N of it, whereas E

of the Station, e.g. in the fjeld area of

Juovva-Skallovarri and its surroundings,

Diapensia is very rare, although there are

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 63

suitable, wide fjeld heaths. It is almost

missing in the fjelds around the Vetsikko

bog area including even the Kuorboaivi

fjeld in the east. Northwards in the

Kalddoaivi fjeld area and N of it the species

is fairly frequent. Northern.

FMF 0.432.

Vertical distribution. a: VI (374;

0.495), b: III (235; 0.104), c: I (31; 0.009).

Differences ***. Frequent in the alpine

belt, but rather rare in the birch belt and

very rare in the coniferous zone. Range ca.

80 m (Anaraspakti, 7776:3529) – 600 m

(Karegasnjarga-Ailigas, Lanka,

7705:3460). Tr 1480 m (Engelskjøn &

Skifte 1995: 151), Fnm 781 m (Norman

I(2): 882), EnL 1100 m (Laine 1958: 90),

possibly 1340 m (Lindén 1943: 78

Haltitschohko on the border of Finland and

Norway). Alpike.

Ecology. Diapensia lapponica is a

plant of the alpine belt. It is growing in

windswept, dry, open, gravelly sites on the

top of heaths, ridges and fjeld plateau (Fig.

7). Without the shelter of snow it is

exposed to hard winds and large changes in

daily temperatures. Against these extreme

conditions the thick main root fastens the

short-grown, only 2–6 cm high cushions to

the ground. The plants do not form large

stands, they mostly grow solitary or in

small groups.

Diapensia occurs as a characteristic

species of Arctostaphylo-Cetrarion nivalis

alliance, a community which is widely

distributed on oceanic as well as

continental mountains (Gjærevoll 1990:

56). Kalliola (1939: 174–181) considered

Diapensia to be a constant species in the

lichen-rich Diapensia-Loiseleuria-

Empetrum association of the extremely

xerophilous Loiseleurieto-Arctostaphylion

group, together with Arctostaphylos alpina,

Betula nana, Empetrum nigrum, Juncus

trifidus, Loiseleuria procumbens,

Vaccinium vitis-idaea and the cryptogams

Fig. 7. Cushions of Diapensia lapponica L. growing on the dry, windswept alpine heath of Kistuskaidi fjelds,

N

W Utsjoki. Photo 7.6.2002 S. Heino.

64 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Alectoria nigricans, Cetraria nivalis and

Polytrichum piliferum. Other usual

companions are Carex bigelowii, Luzula

arcuata, Salix herbacea, Racomitrium

microcarpon, Alectoria ochroleuca,

Cetraria cucullata and Stereocaulon

paschale (Laine 1970(II): 133).

Diapensia also occurs in the birch belt

and (although very rarely) in the coniferous

zone, growing between stones in crevices

and terraces of rocky precipices (Roivainen

1923: 293, Rintanen 1968: 287, Kallio &

Mäkinen 1957: 26), e.g. at the N end of

Puksalskaidi near the Kevo Subarctic

Research Station. In the Peäldoajvi fjeld

area the few localities in the birch belt are

in snow-bed areas with thick snow cover

(Koivistoinen 1964: 59).

Diapensia flowers from June to August

(Hämet-Ahti et al. 1998: 215). According

to the phenological observations from 1977

onwards (Kevo archives), on Puksalskaidi

near the Kevo Subarctic Research Station

the flowering usually starts during the

second week of June. The earliest date

recorded was May 30 in 1984 and the latest

June 25 in 1982. Flowering lasts about two

weeks, and the tiny, light seeds ripen in the

end of August and September (M. Alanen,

pers. comm.). At Saana in Enontekiö

Diapensia was flowering at the height of

600–800 m on June 11–13, 1986 (Uotila

1987).

Diapensia is common both on acid and

neutral soil. It mostly occurs on soils poor

in lime, but it may also be an important

constituent of Dryas-heaths (Gjærevoll

1990: 55). Considered indifferent or

amphicline (Arwidsson 1943: 232, Benum

1958: 315, Wistrand 1962: 132, Nilsson

2000: 152), sometimes acidocline (Laine

1970(II): 133, Mäkinen & Kallio 1979: 17).

Amphicline.

Parasites. In Diapensia cushions the

uppermost leaves live one or two years, but

inside the cushions there are withered

leaves and dead, grey parts of old shoots. In

these dead parts a small crustaceous lichen

Lecidea diapensiae Th. Fr. may appear as

small black spots. This is the only known

substrate for this lichen (SKK III: 319).

Morphology and taxonomy. The

treatment of Diapensia lapponica and D.

obovata as separate species is based on

their almost full allopatry and several

differential characteristics: growth form

(pulvinate in D. lapponica, prostrate and

mat-forming in D. obovata) and differences

in shape, color and surface structure of

leaves, bracteoles and sepals (Hultén 1958:

222, Nesom 2009: 338, Elven 2016). The

difference between the species is small, but

fairly clear.

Dependence on culture. Ahemerobe.

SH

PYROLACEAE

Moneses uniflora (L.) A. Gray

Pyrola uniflora L.

Indigenous, rare

Map 45

Distribution. Widespread boreal circumpolar;

mostly morphologically homogenous, but in the

Pacific N America represented by var. reticulata

(Nutt.) Blake with strongly reticulated leaves (Hultén

1971b: 120, Hultén & Fries 1986: map 1442). In W

Europe rare and absent from many islands and the

extreme south (Fl. Eur. 3: 4, SKK III: 251). Common

in most of Fennoscandia, rarer in the northernmost

and coastal parts (Hultén 1971a: map 1365, Hämet-

Ahti et al. 1998: 214, Mossberg & Stenberg 2003:

452).

In Troms and Finnmark mainly confined to the

pine forests in the inland valleys and climatically

favorable fjord districts including the valley of the

Pasvikelva in the easternmost Finnmark (Norman

II(1): 442, Dahl 1934: 375, Benum 1958: 305, Hultén

1971a: map 1365, Engelskjøn & Skifte 1995: 146,

Nilsson 2000: 155, Lid & Lid 2005: 599, map 421,

Elven et al. 2013: 256). In Pechenga rare – fairly rare

in the Lutto area, in the Kola Peninsula common in

the Kandalaksha area, rare or missing elsewhere

(Fellman 1831: 311, Roivainen 1923: 292, Fl. Murm.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 65

IV: map 91, Hultén 1971a: map 1365, Alm et al.

1998: 135, Mäkinen 2002: 17).

In Finnish Lapland scattered in moist localities

[“locis succosioribus passim”] (Fellman 1835: 263).

In the forest areas of Sompio and Kittilä Lapland rare

to scattered (Hjelt & Hult 1885: 137, Wainio 1891:

45, Hult 1898: 163). Rare in Muonio (Montell 1948,

1962: 125). Scattered to frequent in the Finnish

Lapland up to S and SE parts of Inari (Kujala 1961,

1964: 78, Lampinen & Lahti 2018). Very rare in

Enontekiö (Piirainen 1996b, Lampinen & Lahti l.c.).

InL ref. Scattered in the upper

coniferous zone (“in reg. subsylvatica p”)

and along shores (Kihlman 1884: 110).

Inari, Törmänen, Veskoniemi and

Ruoptuinvaara (Wainio 1891: 45). Inari,

Palopää (Kujala 1929: 114). In the

Lemmenjoki area rare to scattered

(Klockars & Luther 1938, Rahkonen 1968:

18). Two localities (Kuttura, Törmänen)

along the Ivalojoki and four in the

Vaskojoki area (Kujala 1962: 177, Laine

1964: 114). N of the continuous pine forest

line found only in the pine forest area along

the Kevojoki and Utsjoki (Kallio 1959b:

20, 1961: 103, Laine 1970(I): 70), e.g. by

the Keneskoski rapids (Kallio & Mäkinen

1957: 26). Seven localities along the

Kevojoki and five along the Utsjoki (Laine

1970(II): 128, 213).

Kevo 0.6 %, InL 23 %, 84 sq. H 10,

OULU 3, TUR 32, YME 5 spec.

Rare (335; 0.053). Inari: III (310;

0.073), Utsjoki: I (25; 0.012).

Difference***. Practically all localities

south of the continuous pine forest line or

inside the local pine region of the Utsjoki-

Kevojoki valley (cf. Kallio et al. 1971: 84).

Southern.

FMF 0.443.

Vertical distribution. a: I (1; 0.001),

b: I (21; 0.010), c: III (313; 0.090).

Difference a-b*, a-c***, b-c***. Almost

exclusively in the coniferous zone. Range

ca. 90 m (Lake Puksaljavri, 7738:3501) –

410 m (Joenkielinen, 7625:3464, ”fjällhed

vid trädgränsen” 1937 B. Pettersson, H

413302). Tr 373 m (Norman I(1): 760),

Khibiny Mountains 360 m (Alm et al.

1998: 135), EnL 500 m (Väre & Partanen

2009: 101). Silvine.

Ecology. The occurrences of Moneses

uniflora are concentrated in edaphically

favorable localities. It prefers fresh,

nutrient-rich habitats often with a rich moss

cover, and avoids dry and unfertile habitats

(cf. Kujala 1964: 78). It even tolerates

slightly paludified conditions (Kotilainen

1951: 135, Kujala l.c., SKK III: 251). By

the Ivalojoki it grows in herb-rich forests

and coppices, as well as herb-rich spruce

mires (Kujala 1962: 177). Along the

Kevojoki it is found in older pine forests on

fresh heaths slightly moistened by spring

waters, in the company of, e.g., Empetrum

nigrum subsp. hermaphroditum, Phyllodoce

caerulea and Tofieldia pusilla (Laine

1970(II): 128). Moneses is often found near

lake shores, brooks, rivers and by rapids

(cf. Benum 1958: 305). By the Lemmenjoki

it grows in shady pine forests in the mouths

of brooks (Rahkonen 1968: 18). In Inari,

Lintumaa in 2013 it was growing in a fresh

brook-side coppice with, e.g., Carex

cespitosa, C. loliacea, Corallorhiza trifida,

Elymus caninus, Geranium sylvaticum,

Phegopteris connectilis and Prunus padus.

According to the specimens collected,

the flowering in Inari Lapland usually takes

place in July. The earliest flowering

specimens have been collected in the end of

June around the Midsummer, the latest in

the beginning of August. Capsules are

commonly developed.

Moneses is considered amphicline or

Ca-indifferent (Wistrand 1962: 129, Laine

1970(II): 128, Mäkinen & Kallio 1979: 17,

Roweck 1981: 357, Nilsson 2000: 155) or

slightly basocline (Pesola 1928: 89, 159).

The localities of Moneses are usually rich

in nutrients (Kujala 1964: 78). In the

Oulanka National Park it is common in the

area of basic rocks (Söyrinki & Saari 1980:

120). Like some of its companions, the

66 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

species seems to be somewhat demanding

also in Inari Lapland. Probably slightly

basocline.

Parasites. The rust fungus

Pucciniastrum pyrolae Diet. has been

found once on Moneses in Inari (Mäkinen

1964b: 172).

Dependence on culture. Ahemerobe.

JN

Orthilia secunda (L.) House

Pyrola secunda L., Ramischia secunda (L.)

Garcke

Indigenous, scattered

Map 46

Distribution. Boreal circumpolar with two

subspecies, subsp. secunda in Eurasia and North

America, and subsp. obtusata (Turcz.) Böcher in

Siberia and N North America (Hultén 1971b: 138,

Hultén & Fries 1986: map 1441). The latter is

sometimes regarded as a separate species O. obtusata

(Turcz.) H. Hara (Elven 2016). On the other hand,

the whole complex may be seen as one variable

species without subordinate taxa (Fl. N. Am. 8: 388).

Common in most of Fennoscandia, but with slightly

continental tendency with fewer occurrences in the

maritime parts in N and W (Dahl 1934: 374, Hultén

1971a: map 1364, Roweck 1981: 355, Hämet-Ahti et

al. 1998: 213, Mossberg & Stenberg 2003: 452, Lid

& Lid 2005: 598, Lampinen & Lahti 2018).

Widespread throughout Troms (Benum 1958:

307, Engelskjøn & Skifte 1995: 147) and Finnmark

(Norman II(1): 440, Dahl 1934: 374), 7 localities

recorded in the Rastegaissa area (Ryvarden 1969:

34). In Pechenga fairly common (Kontuniemi 1932:

30), common in the E parts of the Luttojoki area but

rare in the west (Roivainen 1923: 292). Rare to

scattered in the alpine belt of Pechenga (Kalliola

1932: 106, Söyrinki 1939a: 316). Common in the

Kola Peninsula (Fellman 1831: 311), but mainly

concentrated in the S and central parts (Fl. Murm. IV:

map 92, Hultén 1971a: map 1364, Mäkinen 2002:

17).

Common in Finnish Lapland (Fellman 1835:

263). In the coniferous forests of Sompio and Kittilä

Lapland common (Hjelt & Hult 1885: 136) to

scattered (Hult 1898: 163). Fairly rare in Muonio

(Montell 1962: 125). Scattered to common in the

Finnish Lapland S of Enontekiö and Inari (Kujala

1964: 80, Lampinen & Lahti 2018), rare in Enontekiö

Lapland except for the southernmost part (Lindén

1943: 78, Piirainen 1996b, Väre et al. 2010: 115,

Lampinen & Lahti l.c.).

InL ref. Fairly frequent in the

coniferous zone and birch belt, only hardly

extending to the alpine belt (Kihlman 1884:

110). In the W parts of Inari and Utsjoki

fairly rare or rare, but possibly overlooked

(Laine et al. 1955: 130, Laine 1964: 115),

fairly frequent in Kallio & Mäkinen (1957:

26). Fairly rare in NE Inari (Såltin 1958).

Fairly rare to rare in the Lemmenjoki area,

the upper course of the Ivalojoki and along

the Kevojoki (Klockars & Luther 1938,

Kujala 1962: 177, Rahkonen 1968: 18,

Laine 1970(II): 129, Heikkinen & Kalliola

1990: 34), scattered to fairly frequent in the

middle and lower course of the Ivalojoki

(Wainio 1891: 46, Kujala l.c.). Two

localities in the Peäldoajvi fjeld area and by

the Nukkumajoki (Koivistoinen 1964: 53,

Suominen 1975).

Kevo 9.7 %, InL 59 %, 192 sq. H 8,

JYV 1, KUO 2, OULU 3, TUR 23, YME 2

spec.

Scattered (1030; 0.156). Inari: IV

(726; 0.169), Utsjoki: III (304; 0.129).

Difference***. Rather evenly distributed

throughout most of the area but largely

missing in the alpine parts of Utsjoki.

Whole area.

FMF 0.782.

Vertical distribution. a: II (38;

0.052), b: III (302; 0.128), c: IV (690;

0.195). Differences***. Range 15 m (Lake

Pulmankijärvi, 7762:3539) – ca. 550 m

(Ladnjoaivi, 7631:3446, Klockars & Luther

1938). Rarely extending to the alpine belt

(cf. Kihlman 1884: 110, Hustich 1940c: 58,

Benum 1958: 307). Tr 590 m (Engelskjøn

& Skifte 1995: 147), Fnm 633 m (Norman

I(1): 759), EnL 750 m (Väre & Partanen

2009: 101). Silvike.

Ecology. Orthilia secunda prefers

humid or moist, humus-rich soil. It grows

in different types of coniferous and birch

forests but prefers fresh and herb-rich ones.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 67

It is mostly found in coppices along brooks

and rivers, on humus-rich or sandy-gravelly

banks, even on stony ground. It is

sometimes found in the not too wet margins

of mires, but is mostly missing in vast mire

areas. Along the Ivalojoki and Lemmenjoki

its habitats include fresh heath forests,

shrubberies and herb-rich coppices along

the shores (Kujala 1962: 177, Rahkonen

1968: 18), by the Kevojoki it grows on

stony shores, heaths and sandy banks with,

e.g., Bistorta vivipara, Juncus trifidus and

Luzula spicata (Laine 1970(II): 129). In the

subalpine forests of Pechenga it is found in

fresh heath forests and herb-rich forests,

e.g. in Dryopteris-Myrtillus, Geranium-

Myrtillus and Geranium types (Kujala

1929: 51-55, Kontuniemi 1932: 30). In the

alpine belt of Pechenga the species is rare

and the habitats include fjeld heaths, heath

meadows and shady places under steep

rocky precipices (Kalliola 1932: 106,

Söyrinki 1939a: 316).

In Utsjoki flowering takes place in

July, the earliest date in Kevonniemi in

1999–2006 being July 1, the latest July 28;

ripe seeds were observed around the middle

of September (Alanen 2007). In the

subalpine belt in Pechenga plants flowered

from early July to early August in 1930

(Kontuniemi 1932: 30). The few alpine

occurrences often do not flower at all

(Söyrinki 1939a: 316, SKK III: 261).

Seedlings are rarely seen, but vegetative

propagation from subterranean stolons is

effective (Kontuniemi l.c., Söyrinki 1939a:

317).

Orthilia is sometimes regarded as

slightly basophilous (Arwidsson 1943: 228,

Lid & Lid 2005: 598). Although it avoids

the most meager substrates (cf. Kujala

1964: 80), it is usually considered

amphicline or indifferent (Pesola 1928:

160, Benum 1958: 307, Wistrand 1962:

129, Roweck 1981: 356, Nilsson 2000:

155). In Inari Lapland it is regarded as

amphicline (Laine 1970(II): 129, Mäkinen

& Kallio 1979: 17). Amphicline.

Parasites. In Inari Lapland Orthilia is

often infected by the rust Pucciniastrum

pyrolae Diet.; the rust Chrysomyxa pirolata

Wint. has been found once in Inari (Rainio

1926: 253, Mäkinen 1964b: 157, 173).

Dependence on culture. Orthilia may

sometimes grow on road verges and in

seminatural Lapp meadows. Hemera-

diaphore.

JN

Pyrola chlorantha Sw.

Indigenous, very rare

Map 47

Distribution. Widespread boreal-montane, amphi-

Atlantic with a tendency towards circumpolar

distribution, but with a wide gap in E Siberia (Hultén

1958: 268, Hultén & Fries 1986: map 1438).

Common in the southernmost parts of Sweden

and Finland, scattered towards the north and rare

north of the Arctic Circle. Scattered through most of

Norway, rare in Troms and only a few localities in

Finnmark (Dahl 1934: 374, Lid 1950, Benum 1958:

306, Hultén 1971a: map 1359, Alm et al. 1995,

Engelskjøn & Skifte 1995: 146, Lid & Lid 2005: 598,

Elven et al. 2013: 325). One locality in the Lutto area

in Pechenga (Roivainen 1923: 292). In the Kola

Peninsula only known from a few localities in the

Kandalaksha area (Fl. Murm. IV: map 93, Hultén

1971a: map 1359).

Very rare in Sompio and Kittilä Lapland

(Wainio 1891: 46, Hämet-Ahti et al. 1998: 212,

Lampinen & Lahti 2018), missing in Enontekiö

Lapland; the dots in NW Enontekiö in Lampinen et

al. (2012 and the earlier versions of the atlas) are

erroneous.

InL ref. In pine forest by Lake

Pyhäjärvi (Kihlman 1884: 110, Mikkola

1941: 33, Laine 1964: 115).

InL 1 %, 2 sq. H 2, TUR 4 spec.

Very rare (6; 0.001). Inari: I (4;

0.001), Utsjoki: I (2; 0.001). Inari: four

localities in three squares in a limited area

in the Vaskojoki area: (1) pine heath ca. 2

km SW of Pyhäjärvi (7644:3455, 1974 J.

Savola, H 320437, TUR 231218); (2)

68 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

sloping pine forest WSW of Pyhäjärvi

(7645:3456, 1960 U. Laine, TUR 76856)

and heath forest by the W end of Pyhäjärvi

(7645:3456, 1965 P. Siltanen, TUR

164833); (3) in pine forest by Pyhäjärvi

(7645:3457, 1880 A. Arrhenius & A.O.

Kihlman, H 413516) and ca. 100 m from

the former main building of Pyhäjärvi

farmstead towards the Vaskojoki

(7645:3457, 1960 E. Rautava, TUR 76857);

(4) one locality ca. 6 km NW of the former

ones in the subalpine birch belt (7650:3452,

1968 U. Laine & J. Nurmi, field list).

Utsjoki: two localities in the birch belt: (5)

SE of Luobmosjavrrik 1 km E of

Leämmasvarri top, one specimen

(7703:3471, 1991 Y. Mäkinen & M.

Airakorpi, field list), and (6) 2 km S of

Skallovarri reindeer fencing along the

upper Kardejohka rivulet, a small stand

with one flowering plant on ca. 0.25 m2 in

Empetrum heath (7746:3507, 1999 Y.

Mäkinen & M. Yli-Pere, field list).

Recorded also between Marastsobma and

Kallovarri in W Utsjoki (7741:3475, 1997

Y. Mäkinen & H. Joutsenlahti, field list);

however, this locality is not accepted here,

as it is the only record in the alpine belt and

no specimen exists. Lowland.

FMF 0.015.

Vertical distribution. b: I (3; 0.001),

c: I (3; 0.001). Both in the coniferous zone

and in the birch belt. Range 180 m

(Pyhäjärvi, 7645:3456) – ca. 375 m

(Leämmasvarri , 7703:3471). Tr 150 m

(Engelskjøn & Skifte 1995: 146), Fnm ca.

250 m (Elven et al. 2013: 325). Silvine.

Ecology. In Inari Lapland, Pyrola

chlorantha is restricted to dry heath

vegetation, and is found both in pine and

birch forests. Its distribution in N Finland is

mainly determined by climatic conditions,

and it shows a continental tendency. It

grows mainly in warm sites on till and

sand, and is missing from paludified or

moist soils (Kujala 1964: 79). It can form

small patches with its runners. The plants

are probably susceptible to cold weather

conditions during autumn and winter, as

flower buds are formed already during the

previous growing season. Flowers are not

nectariferous, and pollinating insects have

not been seen visiting them in Finland

(SKK III: 258).

Amphicline in Pite lappmark (Wistrand

1962: 129), most localities in Koillismaa

with some lime effect (Söyrinki & Saari

1980: 120), basocline in Inari Lapland

(Mäkinen & Kallio 1979: 17). In Inari on

anorthosite bedrock together with some

other calciphilous plants (Laine & Nurmi

1971). Basocline.

Dependence on culture. Ahemerobe.

MP

Pyrola media Sw.

Indigenous, very rare

Map 48

Distribution. Mainly boreal European, from N

Ireland, Scotland, Scandinavia and Central Europe to

the Urals, scattered in W parts of Siberia but

distribution insufficiently known there (Hultén &

Fries 1986: map 1437). More or less common in S

Fennoscandia to ca. 65°N, getting rare towards the

north (Hultén 1971a: map 1361, Mossberg &

Stenberg 2003: 450, Lid & Lid 2005: 597), in

Finland most common in the SE parts of the country

(Lampinen & Lahti 2018).

In Troms rather rare and mainly in the coastal

and fjord areas in the lowland (Benum 1958: 306,

Engelskjøn & Skifte 1995: 146). Rather rare in the W

parts of Finnmark, only a few widely scattered

localities in the east; mainly coastal (Norman II(1):

438, Dahl 1934: 373). Missing in Pechenga (Fl.

Murm. IV: map 95, Hultén 1971a: map 1361), only

in a limited area in the W and SW of the Kola

Peninsula.

Rare in Sompio and Kittilä Lapland, in

Enontekiö Lapland only a few localities in the NW

(Hämet-Ahti et al. 1998: 212, Lampinen & Lahti

2018).

InL ref. Very rare in Inari Lapland: one

locality in Inari, Palopää (7597:3523,

Kujala 1929: 114); only three known

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 69

localities in Laine (1964: 114, 1970(II):

128): Palloaivi in the Vaskojoki area,

Sirratsohkka in the Kevo Strict Nature

Reserve and Juovva-Skallovarri shortly E

of it.

InL 2 %, 11 sq. H 10, TUR 14, YME 9

spec.

Very rare (28; 0.004). Inari: I (19;

0.004), Utsjoki: I (9; 0.004). Most localities

in the W part of the province, centered in

the Lemmenjoki, Vaskojoki and Kevojoki

areas. Whole area.

FMF 0.079.

Vertical distribution. a: I (4; 0.006),

b: I (18; 0.008), c: I (6; 0.001). Difference

a-c*, b-c***. Range 120 m (Raja-Jooseppi

SW, 7596:3553) – ca. 500 m (Morgam-

Viibus, 7618:3455). Tr 580 m (Engelskjøn

& Skifte 1995: 146), Fnm 280 m (Norman

II(1): 438), EnL 540 m according to

specimens in H. Vertical ubiquitous.

Ecology. Most localities of Pyrola

media in Inari Lapland are in subalpine

birch forests, but it grows also in the lower

part of the alpine belt and in the coniferous

zone. Typical habitats are dwarf-shrub

heaths with a well-developed moss layer.

The distribution in the north is probably

determined by climatic factors. The habitat

requirements of P. media are much the

same as those of P. chlorantha, but it seems

to favor more nutrient-rich and mesic, and

in some cases even calciferous localities

(Kujala 1964: 79, SKK III: 253, Söyrinki &

Saari 1980: 118). In Laanila, the lower E

slope of Palopää (7597:3523), the species

was recorded in a relatively mesic pine

forest characterized by a rich dwarf shrub

layer of Vaccinium myrtillus, V.

uliginosum, V. vitis-idaea, Empetrum

nigrum (s. lat.) and Calluna vulgaris, the

abundance of Deschampsia flexuosa and

mosses and hepatics (e.g. Pleurozium

schreberi, Dicranum fuscescens, Lophozia

spp.) and lichens (mainly Cladonia spp.)

(Calluna-Uliginosa site type; Kujala 1929:

114). In a locality in E Utsjoki, ca. 2 km E

of Giikkentsopma (7727:3522), P. media

was growing in a poor dwarf shrub heath in

the subalpine belt, dominated by Betula

nana, Empetrum nigrum subsp.

hermaphroditum and Vaccinium myrtillus;

other vascular plants were e.g. Carex

bigelowii, Vaccinium uliginosum and V.

vitis-idaea.

Norman (II(1): 439), Benum (1958:

306) and Nilsson (2000: 153) regarded P.

media as amphicline or Ca-indifferent,

Wistrand (1962: 129) as possibly

calciphilous. In Inari Lapland it is regarded

as amphicline (Mäkinen & Kallio 1979:

17). Amphicline.

Dependence on culture. All the

known localities are untouched by man.

Ahemerobe.

MP

Pyrola minor L.

Indigenous, frequent

Map 49

Distribution. Boreal-montane circumpolar (Hultén

1971b: 100, Hultén & Fries 1986: map 1436).

Common throughout Fennoscandia (Hultén 1971a:

map 1362, Roweck 1981: 350, Mossberg & Stenberg

2003: 450, Lid & Lid 2005: 597), in Finland most

common in the E and N parts of the country

(Lampinen & Lahti 2018).

In Troms common in suitable habitats in the

whole area (Benum 1958: 307, Engelskjøn & Skifte

1995: 146). More or less evenly distributed

throughout Finnmark (Norman II(1): 439, Dahl 1934:

373); 31 localities in the Rastegaissa area (Ryvarden

1969: 33). In Pechenga scattered to rather common in

the inner parts, Luttojoki area (Roivainen 1923: 292),

common in the alpine belt in the Kammikivi area

(Kalliola 1932: 106) and Pechenga fjelds (Söyrinki

1939a: 314); rather common north to the Arctic

Ocean (Wainio 1891: 46). Scattered (Fellman 1831:

311) to common in the Kola Peninsula, but less

frequent in the northernmost parts (Fl. Murm. IV:

map 96, Hultén 1971a: map 1362).

Common in Finnish Lapland (Fellman 1835:

263). Rather common (Hjelt & Hult 1885: 136, Hult

1898: 163, Montell 1962: 125) to very common

70 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

(Lampinen & Lahti 2018) in Sompio and Kittilä

Lapland. Common or rather common in Enontekiö

Lapland (Laine 1958: 86, Virtanen 1990, Piirainen &

Piirainen 1991b, Hämet-Ahti et al. 1998: 212, Väre et

al. 2008: 76, Lampinen & Lahti l.c.), fairly common

in the birch belt in the province, rather rare in the

alpine belt (Lindén 1943: 78).

InL ref. Common in the whole

coniferous zone and birch belt, also in the

lower part of the alpine belt along rivulets

(Kihlman 1884: 110). Scattered in the

Ivalojoki valley, most common along the

upper course (Kujala 1962: 177). Scattered

in the Viipustunturit – Maarestatunturit and

Lemmenjoki area (Klockars & Luther

1938, Rahkonen 1968: 18). Common in the

Peäldoajvi fjeld area (Koivistoinen 1964:

53), W Utsjoki (Kallio & Mäkinen 1957:

26) and in the Paistunturit fjeld area (Laine

et al. 1955: 130), scattered in the Kevo

Strict Nature Reserve (Laine 1970(II):

128).

Kevo 51.5 %, InL 88 %, 244 sq. H 10,

JYV 2, KUO 1, OULU 7, TUR 54, YME 6

spec.

Frequent (2566; 0.392). Inari: V

(1463; 0.341), Utsjoki VI (1103; 0.495).

Difference***. Rather evenly distributed

throughout most of the area but clearly

most common in W Utsjoki. Whole area.

FMF 0.947.

Vertical distribution. a: V (262;

0.357), b: VI (1135; 0.497), c: V (1169;

0.334). Difference a-b***, b-c***. Range

15 m (Lake Pulmankijärvi, 7762:3539) –

ca. 550 m (Peäldoajvi, 7675:3484). Tr 950

m (Engelskjøn & Skifte 1995: 146), Fnm

570 m (Norman II(1): 439), EnL 1000 m

(Laine 1958: 86). Vertical ubiquitous.

Ecology. Pyrola minor grows in many

kinds of habitats but prefers humid to moist

and humous soils. It is usually scattered or

rather sparse in its localities. In the birch

belt and coniferous zone it is common

especially in relatively nutrient-rich woods,

on rocky slopes and along brooks and

rivers. On river and brook shores it grows

as well in grassy and meadow-like habitats,

often also in paludified meadows, as on

occasionally flooded sand bars and on

gravel close to the shoreline. Sometimes it

is found sparsely at the edges of

minerotrophic mires and in oligotrophic

and mesotrophic fens.

Pyrola minor has no clear preferences

as to topography or the type of the bedrock.

In the subalpine belt in Pechenga it is most

common in forests rich in herbs, and

belongs to the main species in the

Geranium-Myrtillus and Geranium types

(Kujala 1929: 53, 55, Kontuniemi 1932:

30). It grows commonly in the alpine belt,

and the limiting factor in the upper parts of

the relatively low fjelds e.g. in Utsjoki is

usually the dryness of the soil rather than

elevation. In the alpine belt it often grows

in late-snow sites along brooks and rivulets.

In Scandinavian snow-beds P. minor is

mainly found in habitats poor in calciphiles

and hygrophiles (heath series, alliance

Deschampsio-Anthoxanthion; Gjærevoll

1956: 29–86). In the alpine belt in the

Kammikivi area in Pechenga the species

prefers snow-beds (Kalliola 1932: 106). In

the meadow vegetation in the Rybachy

Peninsula it is most common in moist true

meadows (Kalela 1939: Table 25) and

snow meadows (l.c.: Tables 45 and 46).

In the surroundings of the Kevo

Subarctic Research Station flowering starts

in late June (Alanen 2007). In the subalpine

belt in Pechenga plants flower from early

July to early August (Kontuniemi 1932:

30), in the alpine belt flowering usually

starts in late July, and seed ripens regularly

only in favorable sites; in harsher

conditions seed ripens only during the most

favorable growing periods (Söyrinki 1939a:

315). The species is probably mainly self-

pollinating; flowers have no nectar and they

are usually not visited by pollinating insects

(SKK III: 253). Seedlings are rare at least

in the alpine and subalpine belts in

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 71

Pechenga, and propagation is mostly

vegetative (Kontuniemi l.c., Söyrinki l.c.).

In Scandinavia the species is slightly

basophilous (Lid & Lid 2005: 597) or

amphicline (Norman II(1): 439, Wistrand

1962: 129, Arwidsson 1943: 228, Benum

1958: 307, Nilsson 2000: 153). In Inari

Lapland it is regarded as amphicline (Laine

1970(II): 128, Mäkinen & Kallio 1979: 17).

Amphicline.

Parasites. The species is sometimes

infected by the rust species Chrysomyxa

pirolata Wint. in Inari Lapland, and more

often by the rust Pucciniastrum pyrolae

Diet. (Rainio 1926: 253, Mäkinen 1964b:

157).

Morphology and taxonomy. Hybrids

with P. rotundifolia are rare. They resemble

P. media, but have partly reduced anthers

and are seed-sterile. Specimens have been

collected from Inari: (1) SE of the

Lemmenjoki ca. 2 km NE of Nihasanskaidi,

mesic birch grove (7614:3454, 1970 S.

Heinonen & J. Nurmi, TUR 172566); (2)

Morgam-Viibus, brook side meadow in

regio alpina (7617:3456, 1959 R. Kalliola,

H 414035; det. G. Knaben & T. Engelskjøn

1965); (3) Rahajärvi, Aunionlahti, mouth of

Ronkajoki (7629:3507, 1972 Y. Mäkinen,

YME 6853); (4) Uhtsa-Keätkepassi, upper

part of the birch belt (7685:3471, 1976 U.

Laine & J. Nurmi, TUR 271157) and from

Utsjoki: (5) N end of Lake Luomushjärvi,

birch heath (7710:3471, 1973 U. Laine & J.

Nurmi, TUR 223175); and (6) N slope of

Tsierromvarri, in eutrophic, slightly

meadow-like heath (ca. 7752:3477, 1974 L.

Oksanen, H 491152; det. L. Hämet-Ahti

1979).

Dependence on culture. Pyrola minor

is often seen in moderately disturbed road-

cuttings and roadsides and in seminatural

meadows by dwellings. Hemeradiaphore.

MP

Pyrola rotundifolia L.

Incl. P. norvegica Knaben, P. rotundifolia

var. norvegica (Knaben) Hyl., P.

rotundifolia subsp. norvegica (Knaben)

Hämet-Ahti, P. grandiflora subsp.

norvegica (Knaben) Á. Löve & D. Löve

Incl. P. rotundifolia f. chloranthoides Norrl.

P. grandiflora auct.

Indigenous, rare

Map 50

Distribution. Pyrola rotundifolia belongs to a

circumpolar aggregate of several closely related taxa

(Hultén 1958: 142, Hultén & Fries 1986: map 1440,

Elven 2010, 2016). Common in most of

Fennoscandia (weakly bicentric along the Scandes)

(Hultén 1971a: map 1360, Mossberg & Stenberg

2003: 451, Lid & Lid 2005: 598), scattered in N

Finnish Lapland (Hämet-Ahti et al. 1998: 213,

Lampinen & Lahti 2018).

In Troms common in the lowland and up to the

low-alpine belt (Benum 1958: 307, Engelskjøn &

Skifte 1995: 147). Rather common in Finnmark

especially in the lowlands (Norman I(1): 749, II(1):

436, Dahl 1934: 374, Lid & Lid 2005: 598). One

locality in the Rastigaissa area (Ryvarden 1969: 34).

In Pechenga scattered to rather common in the

Kammikivi fjeld area (Kalliola 1932: 106), rare in the

Pechenga fjelds (Söyrinki 1939a: 313), rather

common in the Rybachy Peninsula (Fl. Murm. IV:

map 94, Kalela 1939 in several tables). In the Kola

Peninsula mainly along the northern coastal areas,

scattered in the Khibiny Mountains, in the vicinity of

Kandalaksha and along the S coast (Fellman 1831:

311, Wainio 1891: 46, Fl. Murm. IV: map 94, Hultén

1971a: map 1360, Alm et al. 1998: 135, Mäkinen

2002: 17).

In Sompio and especially in Kittilä Lapland

rather common, rarer in Enontekiö Lapland (Hjelt &

Hult 1885: 136, Hult 1898: 163, Hustich 1940c: 58,

Lindén 1943: 78, Jalas 1949: 95, Laine 1958: 86,

Kujala 1961, Montell 1962: 125, Laitinen & Ohenoja

1990: 23, Lammes 1991, Piirainen 1996b, Hämet-

Ahti et al. 1998: 213, Lampinen & Lahti 2018).

InL ref. Given as common (“locis

umbrosis frequens”) by Fellman (1835:

262), but this seems to be an

overestimation. Scattered in the SW parts

of the province, otherwise rare to very rare

(Mikkola 1941: 33). Scattered along the

Ivalojoki, most common in the upper

72 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

course (Kujala 1962: 177). Rather rare to

scattered in the Viipustunturit –

Maarestatunturit area (Klockars & Luther

1938), rare in the brook side vegetation in

the Lemmenjoki National Park (Rahkonen

1968: 18). Rare along the lower course of

the Vaskojoki (Laine 1964: 115). Rather

rare in the southern part of W Utsjoki

(Laine et al. 1955: 130), rare in Paistunturit

fjeld area (Kallio & Mäkinen 1957: 26) and

in the Kevojoki valley (Laine 1970(II):

128).

Kevo 4.7 %, InL 26 %, 81 sq. H 17,

KUO 1, OULU 3, TUR 48, YME 15 spec.

Rare (240; 0.035). Inari: II (156;

0.034), Utsjoki: II (84; 0.036). Most

localities are in the SW and NW parts of

the province, largely missing in the central

and E parts. Lowland.

FMF 0.344.

Vertical distribution. a: I (10; 0.014),

b: II (120; 0.052), c: II (110; 0.029).

Difference a-b***, a-c*, b-c***. Range ca.

20 m (E side of Lake Pulmankijärvi,

7768:3538) – 440 m (Muotkatunturit,

Bartačohkka, 7674:3464). Tr 930 m

(Engelskjøn & Skifte 1995: 147), Fnm ca.

300 m (Ryvarden 1969: 34), EnL 900 m

(Laine 1958: 86). Silvike.

Ecology. Pyrola rotundifolia grows

most typically in mesic habitats close to

watercourses: luxuriant shore meadows and

birch forests along rivers, brooks and on

lakeshores. It is also met in other luxuriant

forests and mesic heath vegetation,

especially in areas with basic bedrock.

Typical associates among vascular plants

are e.g. Astragalus frigidus, Cornus

suecica, Geranium sylvaticum, Phyllodoce

caerulea and Rubus arcticus, mosses

Barbilophozia hatcheri, B. lycopodioides

and Brachythecium salebrosum, and lichens

Cladonia ecmocyna, Peltigera aphthosa

and P. scabrosa (Laine 1970(II): 128). In

the alpine belt it grows in Dryas heaths and

in moist sloping meadows. It grows also in

eutrophic fens and in fen meadows. In the

alpine belt of the Pechenga fjelds it is most

common in tall-herb meadows and tall-herb

Salix shrub (Geranieto-Cirsium

heterophylli; Kalliola 1939: 105), Dryas

heaths and herb-rich meadows (Söyrinki

1939a: 313), and in the coniferous zone it

belongs to the characteristic taxa of the

moist herb-rich Pyrola-Saussurea forest

site type of Kujala (1929: 60). Flowering

takes place in July. Vegetative propagation

is common by underground stolons.

Pyrola rotundifolia is usually regarded

as basophilous/calciphilous (Arwidsson

1943: 228, Knaben 1943: 5, Selander

1950b: 14, Benum 1958: 307, Knaben &

Engelskjøn 1968: 31, SKK III: 255, Nilsson

2000: 154, Lid & Lid 2005: 598). In Inari

Lapland it is regarded as (slightly)

basocline (Laine 1970(II): 129, Mäkinen &

Kallio 1979: 17). Basocline.

Morphology and taxonomy. The

plants in the study area belong to subsp.

norvegica, which is the predominant race in

N Fennoscandia and probably endemic to N

Europe (Hultén 1958: 142, Hultén & Fries

1986: map 1440, Elven 2010, Elven 2016

as P. grandiflora subsp. norvegica).

Elsewhere in N Finland it is rather common

in the central parts of Lapland, very rare

south to Oulu Ostrobothnia and Kainuu.

The nominal subspecies is southern (the

only race in S Finland); it is rare in Kittilä

and Sompio Lapland and possibly missing

in Enontekiö Lapland (Hämet-Ahti et al.

1998: 213, Lampinen & Lahti 2018). The

subspecies have not been separated in the

printed field lists used during the field

work, and records of subsp. rotundifolia

without a voucher specimen cannot be

accepted here. In Inari Lapland, three

herbarium specimens have been determined

as subsp. rotundifolia: Inari: (1) center of

Ivalo main village, luxuriant riverside

forest (7619:3522, 1957 A. Kaisla, TUR

77167; det. T. Engelskjøn & G. Knaben

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 73

1966), (2) ca. 2 km E of the mouth of the

Vuobmaveäijohka, luxuriant shrub

(7664:3452, 1965 U. Laine, TUR 148198)

and (3) Utsjoki: Karigasniemi, lower slope

S of Meäddemvarri (ca. 7704:3455, 1954

N. Tarén, TUR 294672; det. U. Laine).

However, these specimens represent rather

typical subsp. norvegica with broad and

blunt bracts and sepals and few-veined

petals; also their habit is that of subsp.

norvegica. In addition to the locality (1)

above, there are two dots representing three

records in the distribution map of subsp.

rotundifolia in Lampinen & Lahti (2018),

based on field notes and with no voucher

specimens: Inari: (4) Morgamjärvi SW

(7619:3453, 2007 H. Väre & H.

Kaipiainen, H-Arch., Kastikka doc.

564724), (5) Kultahamina (7619:3454,

2007 H. Väre & H. Kaipiainen, H-Arch.,

Kastikka doc. 564725) and Utsjoki: (6)

mouth of the Fiellokeädggejohka

(7716:3485, 2008 H. Väre & K. Syrjänen,

H-Arch., Kastikka doc. 578205).

About hybrids with Pyrola minor, see

under that species.

Dependence on culture. Practically all

records are from more or less natural

habitats. Ahemerobe.

MP

ERICACEAE

Andromeda polifolia L.

Indigenous, very frequent

Map 51

Distribution. Polymorphic subarctic to boreal

circumpolar (Hultén 1971b: 172, Tolmatchev 1980:

133, Hultén & Fries 1986: map 1456, Fl. North

America 8: 504, Lid & Lid 2005: 603, Elven 2016).

Common throughout Scandinavia, less frequent only

in the southernmost parts of Sweden and in the

Varanger Peninsula in the north (Hultén 1971a: map

1373, Lid & Lid 2005: 603). Very common in most

of Finland, especially in the north (Hämet-Ahti et al.

1998: 210, Lampinen & Lahti 2018).

Common throughout Troms and Finnmark up to

the lower alpine belt (Norman I(1): 720, Dahl 1934:

378, Benum 1958: 310, Alm 1993a: 10, Engelskjøn

& Skifte 1995: 149), 44 localities in the Rastigaissa

area (Ryvarden 1969: 34). Common in Pechenga

(Roivainen 1923: 293, Kalliola 1932: 107, Kalela

1939, Söyrinki 1939a: 326) and in the Kola Peninsula

(Fellman 1831: 311, Fl. Murm. IV: map 102,

Mäkinen 2002: 16).

Common everywhere in Finnish Lapland

(Fellman 1835: 262). Common in Sompio and Kittilä

Lapland (Hjelt & Hult 1885: 136, Hult 1898: 163,

Hustich 1940c: 58, Montell 1962: 125, Hämet-Ahti et

al. 1998: 210, Lampinen & Lahti 2018) and in

Enontekiö Lapland, but scattered in the higher fjeld

areas (Lindén 1943: 77, Laine 1958: 89, Piirainen &

Piirainen 1991b, Väre et al. 2015, Lampinen & Lahti

l.c.).

InL ref. Common in the coniferous

zone and subalpine belts, rather common in

the alpine belt (Kihlman 1884: 109). Very

common and often rather abundant to

abundant in the whole area (Mikkola 1941:

34). Common in the S parts (Wainio 1891:

45). Common in most of the Ivalojoki

(Kujala 1962: 177) and Vaskojoki valleys

(Laine 1964: 115). Rather common in the

Viipustunturit – Maarestatunturit area

(Klockars & Luther 1938). Common in the

Peäldoajvi fjeld area (Koivistoinen 1964:

55) and in W Utsjoki (Kallio & Mäkinen

1957: 26), rather common in the

Paistunturit fjeld area (Laine et al. 1955:

130); common in the Kevo Strict Nature

Reserve but less so in the alpine belt (Laine

1970(II): 130, Heikkinen & Kalliola 1990:

34).

Kevo 74.4 %, InL 94 %, 257 sq. H 19,

JYV 1, KUO 1, OULU 2, TUR 31, YME 2

spec.

Very frequent (4184; 0.646). Inari: VII

(2719; 0.634), Utsjoki: VII (1465; 0.671).

Difference**. Whole area.

FMF 0.978.

Vertical distribution. a: VII (439;

0.591), b: VII (1586; 0.708), c: VII (2159;

0.619). Difference a-b***, b-c***. Range

15 m (Lake Pulmankijärvi 7762:3539) –

600 m (Karegasnjarga-Ailigas, Lanka,

74 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

7705:3460). Tr 900 m (Benum 1958: 311),

Fnm 730 m (Ryvarden 1969: 34), EnL 950

m (Laine 1958: 89). Silvike.

Ecology. The habitat variation of

Andromeda polifolia is very wide. It

belongs to the most oligotrophic mire

plants (Ruuhijärvi 1960: 33), but it is

common in several kinds of meso- and

eutrophic fens as well. It is not confined to

any special mire types, though it is most

common in low-sedge bogs and fens,

Fuscum bogs and Eriophorum vaginatum

pine bogs (SKK III: 279). Typical

associates in bogs are e.g. Betula nana,

Empetrum nigrum subsp. hermaphroditum,

Ledum palustre, Rubus chamaemorus,

Vaccinium vitis-idaea, Dicranum

elongatum, Sphagnum fuscum, Cladonia

arbuscula, C. stellaris and Cetraria nivalis,

in wet fens e.g. Carex chordorrhiza, C.

rostrata, C. rotundata, Eriophorum

angustifolium, Sphagnum compactum and

S. lindbergii, and in more eutrophic mires

also e.g. Carex dioica, Menyanthes

trifoliata, Trichophorum alpinum,

Calliergon stramineum, Paludella

squarrosa, Scorpidium revolvens and

Sphagnum teres (Ruuhijärvi 1960).

Andromeda polifolia grows also in lake

and river shores, moist rocks and moist low

alpine heaths. It is also common e.g. in

arctic dwarf shrub meadows in N Pechenga

(Kalela 1939: 296).

Flower buds are formed already in the

autumn. Flowering starts in the lowlands in

late May to early June, in the alpine belt in

early July (Valle 1930, Söyrinki 1939a:

326, SKK III: 279, Alanen 2007). Self-

pollination is probably a rule (SKK III:

279). In Pechenga, seeds start to ripen in

late July, but seed production is usually

very poor, and vegetative propagation by

the long rhizomatous basal parts is

probably more important (Söyrinki 1939a:

326). Usually regarded as indifferent

(Arwidsson 1943: 230, Wistrand 1962: 131,

Nilsson 2000: 157), in the Oulanka

National Park the pH in its habitats varies

between 3.6–7.1 with the average at 5.4

(Söyrinki & Saari 1980: 121). In Inari

Lapland amphicline (Laine 1970(II): 130)

or acidocline (Mäkinen & Kallio 1979: 17).

Acidocline.

Morphology and taxonomy. Typical

lowland plants are 10–20 cm tall and their

leaves are ca. 20–30 mm × 2–4 mm. Plants

in alpine habitats are often very small, less

than 10 cm, and they have short and

narrow, almost needle-like leaves ca. 10–20

mm × 1–2 mm. Good examples are three

specimens from Utsjoki: (1) alpine heath

NE of Tsoagis Jottijärvi (Goškkes

Johttejávri, 7720:3521, 1958 A. Korhonen,

TUR 77542); (2) Kenesvaara (773:350,

1958 C. E. Sonck, TUR 263690); and (3)

Kevo, Mielgitsohkka (Mielketsohkka), on

moist mosses on a rocky S slope

(7746:3495, 1970 S. Fagerlund, TUR

186537). This kind of plants have

sometimes been called var. acerosa C.

Hartm. Tolmatchev (1980: 134, map 56)

included also the N Fennoscandian arctic-

alpine variants in subsp. pumila V. M.

Vinogr., which is usually understood only

as a Siberian taxon, or Siberian – North

American as in Elven (2016). However, the

N Finnish alpine plants probably deserve

no taxonomical recognition, as the variation

seems to be totally clinal. – White-flowered

plants have been recorded by L. Iso-Iivari

in 2002 in W Utsjoki, the Akujoki valley

(7719:3463).

Dependence on culture. Hemera-

diaphore.

MP

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 75

Arctostaphylos alpina (L.) Spreng.

Arbutus alpina L., Arctous alpina (L.) Nied.

Indigenous, frequent

Map 52

Distribution. Circumpolar arctic-montane, widely

distributed in Eurasia and N America (Hultén 1971b:

50, Hultén & Fries 1986: map 1455). Common

throughout most of Norway, the Scandes and the

northernmost parts of Fennoscandia, but less frequent

in Inari Lapland and adjacent eastern parts of

Finnmarksvidda in Norway (Hultén 1971a: map

1375, Mossberg & Stenberg 2003: 458, Lid & Lid

2005: 603).

Common to very common in Troms (Benum

1958: 311, Engelskjøn & Skifte 1995: 149). Common

in Finnmark (Dahl 1934: 377, Alm 1993a: 12), 53

localities in the Rastigaissa area (Ryvarden 1969:

34). Common in the alpine and subalpine areas of

Pechenga, rather rare to scattered in the coniferous

zone (Roivainen 1923: 293, Kalliola 1932: 107, Valle

1933a: 271, Söyrinki 1939a: 327). Common

throughout the Kola Peninsula (Fl. Murm. IV: map

104, Hultén 1971a: map 1375).

Common to very common in Sompio and

Kittilä Lapland in the alpine and subalpine belts, rare

to scattered in the upper parts of the forest belt (Hjelt

& Hult 1885: 135, Wainio 1891: 45, Hult 1898: 163,

Hustich 1937a: 102, 1940c: 58, Laine 1958: 89,

Montell 1962: 125, Rintanen 1968: 286, Lampinen &

Lahti 2018). Common to very common in Enontekiö

Lapland (Hustich 1937a: 102, 1940c: 58, Lindén

1943: 77, Piirainen & Piirainen 1991b, Lampinen &

Lahti l.c.). The common distribution extends to ca.

Muonio–Kuusamo line in the south, scattered further

south to N Outer Ostrobothnia and S Koillismaa,

three close localities in NE Oulu Ostrobothnia

(Kujala 1964: 81, Lampinen & Lahti l.c., Kastikka

2018).

InL ref. Common in Inari Lapland

(Fellman 1835: 262, Mikkola 1941: 34),

very common in the subalpine and alpine

belts (Kihlman 1884: 109). Rather common

in the Ivalojoki and Veskoniemi areas, also

at the Paatsjoki (Wainio 1891: 45).

Common in the SE parts of Inari Lapland

(Rintanen 1968: 286) and in the

Viipustunturit – Maarestatunturit area, also

occurring in the forest belt (Klockars &

Luther 1938). Rather common in the

Peäldoajvi fjeld area (Koivistoinen 1964:

56). In W Utsjoki rather common (Laine et

al. 1955: 130) to common (Kallio &

Mäkinen 1957: 26), rather rare outside the

alpine belt in the Kevo Strict Nature

Reserve (Laine 1970(II): 130).

Kevo 79.9 %, InL 86 %, 240 sq. H 38,

JYV 6, KUO 5, OULU 8, TUR 43, VOA 3,

YME 2 spec.

Frequent (3236; 0.502). Inari: VI

(1774; 0.414), Utsjoki: VII (1462; 0.677).

Difference***. Common nearly throughout

the area but less frequent in the coniferous

zone in the Lake Inari basin. Whole area.

FMF 0.936.

Vertical distribution. a: VII (593;

0.788), b: VII (1508; 0.681), c: V (1135;

0.326). Differences***. Most common in

the alpine and birch belts, significantly

more infrequent in the coniferous zone.

Range 15 m (Lake Pulmankijärvi,

7762:3539) – ca. 600 m (Karegasnjarga-

Ailigas, Lanka, 7705:3460). Tr 1075 m

(Norman I(1): 716), Fnm 884 m (Norman

I(1): 718), EnL 1050 m (Laine 1958: 89,

Väre & Partanen 2009: 97). Vertical

ubiquitous.

Ecology. Arctostaphylos alpina is

typical in dry lichen heaths. It is most

abundant in the continental types of the

alpine and subalpine vegetation, which is

seen in its scarcity in the coastal areas of

the oceanic subzone of the north

Fennoscandian mountain birch forest

vegetation (Hämet-Ahti 1963a: 65) and

lacking from the submaritime birch forests

(l.c.: 77). However, the species may be

locally common also in the oceanic

subalpine Empetrum (sET) association,

which is characterized by a dense cover of

Empetrum nigrum subsp. hermaphroditum

(l.c.: 70). Typical habitats in the birch belt

are the continental subalpine Empetrum-

Lichenes (sELiT) and Empetrum-Lichenes-

Pleurozium (sELiPlT) associations. In these

patchy vegetation types A. alpina is mostly

confined to dry open patches between birch

groups, together with e.g. Arctostaphylos

76 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

uva-ursi, Calamagrostis lapponica, Carex

bigelowii, Diphasiastrum complanatum,

Empetrum nigrum subsp. hermaphroditum,

Juncus trifidus, Loiseleuria procumbens,

Luzula spicata, Pohlia nutans, Polytrichum

juniperinum, P. strictum, Cetraria

islandica, Cladonia bellidiflora, C.

stellaris, C. uncialis and Stereocaulon

paschale (l.c.: 65). Other typical associates

among vascular plants are e.g.

Deschampsia flexuosa, Festuca ovina,

Vaccinium uliginosum and V. vitis-idaea.

Arctostaphylos alpina is common in

several oligotrophic associations of the

alpine heath vegetation, even in deflation

sites (Kalliola 1939, Haapasaari 1988).

Typical associates in addition to those

mentioned above are e.g. Dicranum

fuscescens, Polytrichum piliferum,

Alectoria nigricans, A. ochroleuca,

Cetraria nivalis and Cladonia mitis. The

species is common also in dryish Dryas

heaths, as recorded also from Pechenga

(Kalliola 1932: 23, 1939: 118–131,

Söyrinki 1939a: 327) and in dryish dwarf

shrub meadows rich in calcium (Kalela

1939: 288–294). It is less common in more

mesic sites characterized by Vaccinium

myrtillus, e.g. the Empetrum-Myrtillus

association described by Kalliola (1939:

220), and avoids localities with late snow.

The species is rather common also on

raised hummocks in bogs and in the

marginal parts of aapa mires. Koivistoinen

(1964: 56) mentions the species in this kind

of habitats in the subalpine belt in the

Peltotunturi area but not in the coniferous

zone. In peatland vegetation it belongs to

the ombrotrophic and oligotrophic group of

hummock-level bog species (Eurola et al.

1995: 67). Lumiala (1939) explained its

peatland occurrences in Koillismaa, Salla,

by the local microclimate, which resembles

alpine conditions in day-time warming and

night-time cooling.

Flowering starts in the surroundings of

the Kevo Subarctic Research Station in late

May and lasts till mid-June, the earliest

recorded date being May 22, the latest June

30. Seed ripening starts usually in early

August, in early summers already in mid-

July, and lasts till mid-September (Alanen

2007). In Pechenga flowering is usually

over by the end of June, and all the seeds

are ripe by the end of August. Seed

production is good and seedlings are

common on open mineral soil, and the

species may also propagate vegetatively by

the rooting branches (Söyrinki 1939a: 327–

328). The species is one of the most showy

participants of the autumn coloring with its

bright red or orange (more yellow in the

shade), and its abundance is best detectable

at this time of the year. The fruits are juicy

and edible as well as important

nourishment for fjeld birds (willow grouse

and ptarmigan).

Amphicline/indifferent in relation to

nutrients and calcium (Norman II(1): 421,

Arwidsson 1943: 230, Benum 1958: 311,

Wistrand 1962: 131, Laine 1970(II): 131,

Mäkinen & Kallio 1979: 17, Roweck 1981:

369, Nilsson 2000: 157), both on acid and

base-rich soil (Lid & Lid 2005: 603).

Rintanen (1968: 286) gives the pH range

4.1–4.6, but basic soils are more or less rare

in his study area. Amphicline.

Parasites. Often attacked by the

“swelling” fungus Exobasidium

arctostaphylii. Also the rust Pucciniastrum

sparsum (Wint.) Jørst. has been found

several times (Kari 1936: 15, Mäkinen

1964b: 173).

Morphology. A morph with light

green summer leaves, orange autumn color

(though in an open site) and red berries also

when ripe is known from the top of

Jesnalvárri fjeld close to the Kevo

Subarctic Research Station (7743:3498,

2000 M. Alanen, TUR 571611). Some

specimens have exceptionally small (e.g.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 77

ca. 11–15 × 6–7 mm; Lutto, Suomuköngäs

in Sompio Lapland, 7595:3553, 1906 A.

Renvall, H 415914) or narrow leaf blades

(e.g. ca. 20 × 5 mm; Vaistsohkka,

7745:3511, 1956 N. Tarén, TUR 165699).

Dependence on culture. Occasionally

on heavily trampled trails. Hemera-

diaphore.

MP

Arctostaphylos uva-ursi (L.) Spreng.

Arbutus uva-ursi L.

Indigenous, scattered

Map 53

Distribution. Circumpolar, with a gap in the

Beringian area. Rather variable in N America, but the

nominal race is distributed throughout the area

(Hultén 1971b: 158, Hultén & Fries 1986: map

1454). Common throughout Fennoscandia except for

the mountainous areas of the Scandes and the far

north (Hultén 1971a: map 1376, Hämet-Ahti et al.

1998: 210, Mossberg & Stenberg 2003: 458, Lid &

Lid 2005: 603).

In Troms scattered in the lowland and up to the

tree line, more common in the E and SE parts; rare in

the alpine regions (Benum 1958: 311, Engelskjøn &

Skifte 1995: 150). In Finnmark scattered to rather

common in the inland especially along the larger

rivers, more or less rare in the coastal areas (Dahl

1934: 377, Alm 1993a: 15), eight localities in the

Rastigaissa area (Ryvarden 1969: 34). Only two dots

in Pechenga in Fl. Murm. (IV: map 103), but

scattered to rather frequent especially in the W parts

of the Lutto area in S part of the province (Roivainen

1923: 293); two records from Pechenga in Mäkinen

(2002: 16), and nine localities in Kastikka (2018).

Common in the S parts of the Kola Peninsula, rare in

the north (Fl. Murm. l.c., Hultén 1971a: map 1376).

Common in Keret Karelia (Sokolov & Filin 1996:

123).

Mostly common in Sompio and Kittilä Lapland

in the forest belts (Hjelt & Hult 1885: 135, Wainio

1891: 44, Hult 1898: 163), especially in the N parts

(Kujala 1964: 82, Lampinen & Lahti 2018), scattered

in the lower parts of the alpine belt (Hustich 1940c:

58), less common in Muonio (Montell 1962: 125).

Common in the E part of Enontekiö Lapland, rather

rare in the NW parts (Lindén 1943: 77, Piirainen &

Piirainen 1991b, Lampinen & Lahti l.c.).

InL ref. Common in Inari Lapland

(Fellman 1835: 262), common and

abundant in the forest belts, rare in the

alpine belt (Kihlman 1884: 109, Mikkola

1941: 34). Rather common (Wainio 1891:

44) or scattered (Kujala 1962: 177) along

the Ivalojoki, also in the Paatsjoki area

(Wainio l.c.), common in the Viipustunturit

– Maarestatunturit area in the Lemmenjoki

National Park (Klockars & Luther 1938).

Rather rare in the Peäldoajvi fjeld area

(Koivistoinen 1964: 55). In W Utsjoki

rather rare (Laine et al. 1955: 130) to

scattered but common on sandy heaths at

the W shore along the lower course of the

Utsjoki, rare along the Teno (Kallio &

Mäkinen 1957: 26), rather rare in the Kevo

Strict Nature Reserve (Laine 1970(II):

130).

Kevo 10.7 %, InL 64 %, 188 sq. H 15,

JYV 2, TUR 13, YME 4 spec.

Scattered (1434; 0.217). Inari: IV

(1041; 0.241), Utsjoki: IV (393; 0.169).

Difference***. Scattered almost throughout

the area except for the E parts of the Lake

Inari basin and the alpine parts of Utsjoki,

where the species is largely missing. Whole

area.

FMF 0.764.

Vertical distribution. a: III (66;

0.089), b: IV (492; 0.217), c: IV (876;

0.244). Difference a-b***, a-c***, b-c*.

Commonest in the coniferous zone and the

subalpine birch forest belt, significantly

more infrequent in the alpine belt, missing

from the larger alpine areas in Utsjoki.

Range 15 m (E shore of Lake

Pulmankijärvi, 7767:3538) – ca. 580 m

(Ladnjoaivi fjeld, 7631:3446). Tr 1000 m

(Engelskjøn & Skifte 1995: 150), Fnm 544

m (Norman I(1): 719), EnL 950 m (Väre &

Partanen 2009: 96). Silvike.

Ecology. Arctostaphylos uva-ursi

grows mainly in open forests on sand or

gravel and on rock surfaces with a thin

mineral soil, but also on till. It prefers

78 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

forest openings and sunny slopes on eskers

and sandy – gravelly river banks etc.; cf.

Roweck (1981: 367). The species is at its

commonest in Finland in a broad belt

reaching from SW Inari Lapland to NE

Sompio Lapland (Kujala 1964: 82, map

139; Lampinen & Lahti 2018). It favors

heath forests rich in lichens and dwarf

shrubs, and is a constant but rather scarce

member of Vaccinium-Empetrum-Cladonia

pine heath association described by Kujala

(1929: 72) from Pechenga. Often in the

company of A. alpina (Mikkola 1941: 34).

The species is clearly continental in

Finnmark (Dahl 1934: 377). Continentality

increases towards the SW part of Inari

Lapland (Tuhkanen 1980: 78), which is

also partly visible in the higher frequency

of the species towards SW.

Arctostaphylos uva-ursi has no

underground vegetative means of

propagation and is therefore destroyed by

forest fires. However, it regenerates easily

from the seed bank, and seed germination is

also enhanced by fire (Kujala 1926: 24,

Söyrinki & Saari 1980: 121). The species

has a good ability to activate dormant

meristems in the above-ground shoots after

mechanical damage (Salemaa et al. 1999),

which probably is advantageous e.g. against

trampling by reindeer.

Flowering starts in the surroundings of

the Kevo Subarctic Research Station during

the first half of July and lasts till the end of

the month, the earliest recorded date being

May 22 and the latest July 9. Seed ripens

mostly during the first half of September

(August 21 – September 25) (Alanen 2007).

Though the species is significantly less

common in the alpine belt, fruiting has

been recorded at least to ca. 500 m a.s.l. in

the Peäldoajvi fjeld area (7675:3484,

Kihlman 1884: 109).

Basocline in Troms (Benum 1958:

311), often on calcareous soils in Finnmark

(Norman II(1): 423, Dahl 1934: 377), the

Oulanka National Park in Koillismaa

(Söyrinki & Saari 1980: 121) and in the

Kilpisjärvi area in Enontekiö Lapland

(SKK III: 285), possibly slightly

calciphilous (Arwidsson 1943: 230) or

indifferent (Wistrand 1962: 131) in Pite

lappmark. Calciphilous according to

Nilsson (2000: 157), indifferent according

to Roweck (1981: 368). Acidocline in Inari

Lapland (Mäkinen & Kallio 1979: 17); in

the Kevojoki area, Laine (1970(II): 130)

considers the species amphicline.

Thermophilous, amphicline.

Morphology. In the Kevojoki canyon,

f. angustifolia Moe has been found as very

rare (Laine 1970(II): 31; SE of Lake Ylä-

Njaggaljärvi, 7726:3495, 1965 U. Laine,

TUR 130975), with leaf blades ca. 15 × 3–4

mm.

Dependence on culture. In more

southern parts of Finland, Arctostaphylos

uva-ursi is often apophytic e.g. on road and

railroad banks and in forest cutting areas

(Kujala 1964: 82, Ahti & Hämet-Ahti 1971:

69, SKK III: 284). Also in Inari Lapland it

spreads in forest clearings and road side

sowings. In Enontekiö Lapland it has been

recorded on disturbed soil in a dumping

place (Piirainen 1996a). Hemeradiaphore.

MP

Calluna vulgaris (L.) Hull

Erica vulgaris L.

Indigenous, very frequent

Map 54

Distribution. Europe and adjacent parts of Asia,

introduced in N America and New Zealand (Hultén

& Fries 1986: 1447). In Fennoscandia very common

except for the mountainous areas of the Scandes

especially north of the Arctic Circle and N parts of

Finnmark and the Kola Peninsula (Hultén 1971a:

map 1386, Lid & Lid 2005: 605). In Finland very

common almost throughout the country except for

NW Lapland (Hämet-Ahti et al. 1998: 209,

Lampinen & Lahti 2018).

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 79

In Troms very common from the outermost

islands to the fjords, less frequent to the heads of the

fjords, seldom in the valleys and partly absent from

the interior mountain plains (Benum 1958: 312,

Engelskjøn & Skifte 1995: 150). In W Finnmark

mainly in the coastal areas, rare in the inland, in E

Finnmark increasing from the coast towards the

inland; in Sør-Varanger mainly in the inland (Dahl

1934: 381, Alm 1993a: 16). Very common in the

inner parts of Pechenga, Luttojoki area (Roivainen

1923: 293), missing from the alpine belt in the

Kammikivi area but present in the near-by subalpine

belt (Kalliola 1932: 107), a few localities in W

Pechenga (Valle 1933a: 271); common north to the

Arctic Ocean (Wainio 1891: 45), but only three dots

in Pechenga in Fl. Murm. (IV: map 105) and ca. 20

records from not more than ca. one dozen separate

subareas in Kastikka (2018). Common in the S parts

of the Kola Peninsula, rare or almost missing in the

northernmost and easternmost parts (Fl. Murm. l.c.).

Common everywhere in Finnish Lapland

(Fellman 1835: 260). Very common in Sompio and

Kittilä Lapland (Hjelt & Hult 1885: 135, Hult 1898:

164, Montell 1962: 125, Kujala 1964: 83, Hämet-

Ahti at al. 1998: 209, Lampinen & Lahti 2018). Very

common in the E parts of Enontekiö Lapland, in the

west only a few localities in the Könkämäeno river

valley and Lake Kilpisjärvi area (Lampinen & Lahti

l.c.).

InL ref. Common in the whole

coniferous zone, scattered to rather

common in the subalpine belt, often also

above the birch limit (Kihlman 1884: 109).

Very common in the SE parts of the

province, scattered in the N parts (Kujala

1964: 83). Common and abundant in the

whole Ivalojoki valley but probably less so

closer to the river mouth (Kujala 1962:

178). Rather common in the Viipustunturit

– Maarestatunturit area (Klockars & Luther

1938). Rather common in the Peäldoajvi

fjeld area (Koivistoinen 1964: 57), W

Utsjoki (Kallio & Mäkinen 1957: 26),

scattered in the Paistunturit fjeld area

(Laine et al. 1955: 130), rather rare in the

canyon area of the Kevo Strict Nature

Reserve (Laine 1970(II): 132).

Kevo 69.1 %, InL 92 %, 246 sq. H 21,

JYV 1, OULU 1, TUR 18, YME 1 spec.

Very frequent (3795; 0.582). Inari: VII

(2833; 0.659), Utsjoki: VI (962; 0.430).

Difference***. Rather evenly distributed

throughout most of the area but locally

almost missing in the fjeld areas in N

Utsjoki. Whole area.

FMF 0.960.

Vertical distribution. a: V (279;

0.381), b: VI (1257; 0.555), c: VII (2259;

0.643). Differences***. Range 15 m (Lake

Pulmankijärvi, 7762:3539) – ca. 540 m

(Peäldoajvi, 7675:3484). Tr 664 m

(Norman I(1): 734), Fnm 428 m (Norman

I(1): 734, but see Alm 1993a: 17), EnL 650

m (Väre & Partanen 2009: 90). Silvike.

Ecology. Calluna vulgaris grows

mainly in poor heath forests dominated by

Pinus sylvestris heath and/or Betula

pubescens subsp. czerepanovii. The species

prefers dryish sandy or gravelly soils.

Typical associates are Deschampsia

flexuosa, Empetrum nigrum subsp.

hermaphroditum, Vaccinium myrtillus, V.

vitis-idaea, Barbilophozia spp., Dicranum

fuscescens, D. majus, Polytrichum

piliferum, Cetraria delisei, Cladonia

coccifera, C. stellaris, Nephroma arcticum

and Stereocaulon paschale. It is often met

also on till in more mesic forests and river

and lake shores, accompanied by also e.g.

Carex vaginata, Festuca ovina, Geranium

sylvaticum, Molinia caerulea, Pyrola

minor, Solidago virgaurea, Trientalis

europaea and Vaccinium uliginosum.

Kujala (1929: 83–88) described three

associations along this gradient

characterized by the abundance of Calluna

vulgaris from the Ivalo – Laanila area.

Other typical habitats are paludified forests

and pine bogs where Calluna grows mainly

on mats or hummocks of Sphagnum

fuscum, together with e.g., Carex

paupercula, Eriophorum vaginatum and

Trichophorum cespitosum. It may grow

also in meadow vegetation, especially

along shores and in depressions with

spring-time melt water flow. It grows

mainly in the coniferous zone and

80 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

subalpine belt but it is not uncommon in the

lower parts of the alpine belt. The species

favors a humid climate, is light-demanding

and needs a good winter-time snow

protection. In the upper subalpine and

lower alpine belt it grows sometimes in

sites with late snow and a scarce

vegetation, and often with some almost

bare patches of frost soil.

The coverage of the species in Finnish

forests has declined since the 1950’s, the

main reasons being closing-up of forest

vegetation, and forest cutting combined

with soil preparing especially in the north

(Salemaa 2000). This is probably visible

also in the SE parts of Inari Lapland.

The flowering period in the

surroundings of the Kevo Subarctic

Research Station is from late July to mid-

September (Alanen 2007). The species is

said to be clearly calcifugous (SKK III:

307), possibly acidocline (Laine 1970(II):

132) or amphicline/indifferent (Wistrand

1962: 132; Mäkinen & Kallio 1979: 17,

Nilsson 2000: 155). It is abundant in the

Angeli anorthosite area in W Inari, but this

seems to be rather due to the high ground

water level than soil chemistry (Laine &

Nurmi 1971). Acidocline.

Morphology. Plants with white

flowers (f. alba (Weston) Braun-Blanq.) are

relatively common in the area (e.g. 12 out

of 18 specimens in TUR and 7 out of 21

specimens in H represent this form);

mentioned also e.g. from the Kevo Strict

Nature Reserve (Laine 1970(II): 132).

Dependence on culture. Seedlings are

common in trampled places, where open

soil is exposed. Hemeradiaphore.

MP

Cassiope hypnoides (L.) D. Don

Andromeda hypnoides L., Harrimanella

hypnoides (L.) Coville

Indigenous, rare

Map 55

Distribution. Arctic, amphi-Atlantic with an oceanic

tendency (Hultén 1958: 34, Hultén & Fries 1986:

map 1449). Common throughout the Scandes, with a

rather sharp limit east of the mountain range (Hultén

1971a: map 1371, Mossberg & Stenberg 2003: 454,

Lid & Lid 2005: 602).

Common in Troms (Benum 1958: 310,

Engelskjøn & Skifte 1995: 149) and Finnmark

chiefly in the alpine areas, in Finnmark scattered to

rather rare in the outermost coastal areas, less

common in the N part of Finnmarksvidda and very

rare in the south (Norman II(1): 427, Dahl 1934: 379,

Alm 1993a: 18); 75 localities in the Rastigaissa area

(Ryvarden 1969: 34). Rare to scattered in the alpine

and subalpine areas in Pechenga (Roivainen 1923:

293, Kalliola 1932: 107, Valle 1933a: 271, Söyrinki

1939a: 324). In the Kola Peninsula common mainly

in the high inland fjeld areas, only a few localities

along the N and E coasts (Fl. Murm. IV: map 101,

Hultén 1971a: map 1371).

Common in the fjeld areas of the NE parts of

Sompio Lapland (Hult 1898: 164, Vuori & Pertola

1959a, Rintanen 1968: 286, Lampinen & Lahti

2018). In Kittilä Lapland only in the Pallastunturit

area (Hustich 1940c: 58, Montell 1962: 125,

Lampinen & Lahti l.c.). Very common in NW

Enontekiö Lapland, fairly common in the central

parts, rare in the east (Hustich 1940c: 58, Lindén

1943: 78, Laine 1958: 89, Piirainen & Piirainen

1991b, Lampinen & Lahti l.c.).

InL ref. Not rare in the fjeld areas

(Fellman 1835: 262), common in the alpine

belt, descending to the limit of the birch

belt (Kihlman 1884: 109), and along brook

sides also in the subalpine belt (Mikkola

1941: 34). Common in the SE parts of Inari

Lapland (Rintanen 1968: 286), rather rare

in the Viipustunturit–Maarestatunturit area

in snow-beds in the alpine belt (Klockars &

Luther 1938). Scattered in the Peäldoajvi

fjeld area in the alpine and birch belts

(Koivistoinen 1964: 55). In W Utsjoki

rather rare to scattered (Laine et al. 1955:

130, Kallio & Mäkinen 1957: 26), mostly

in the southern part of the Kevo Strict

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 81

Nature Reserve (Laine 1970(II): 130,

Heikkinen & Kalliola 1990: 47); only one

locality in Hustich (1942a: 225).

Kevo 22.6 %, InL 24 %, 67 sq. H 30,

TUR 43, YME 4 spec.

Rare (356; 0.056). Inari: I (59; 0.014),

Utsjoki: III (297; 0.139). Difference***.

Scattered in the high fjeld areas

(Raututunturit, Viipustunturit–Maaresta-

tunturit, Muotkatunturit, Paistunturit).

Northern.

FMF 0.275.

Vertical distribution. a: V (257;

0.345), b: II (92; 0.042), c: I (7; 0.002).

Differences***. Fairly common in the

alpine belt, rare in the subalpine belt, only a

few records in the coniferous zone e.g. in

the valley of the Kevojoki (Laine 1970(I):

62). Range ca. 100 m (Paddajohka,

7763:3488) – ca. 600 m (Karegasnjarga-

Ailigas, Lanka, 7705:3460). Tr 1540 m

(Engelskjøn & Skifte 1995: 149), Fnm

1010 m (Norman II(1): 428), EnL 1325 m

(Väre & Partanen 2009: 93). Alpike.

Ecology. In Inari Lapland, most

habitats are in the alpine belt and in a lesser

extent also in the subalpine belt, usually in

sites with fairly good moisture throughout

most of the vegetation period, typically in

snow-beds and other moist depressions, on

rocks by rapids and along brooks (Kallio

1959b: 29, Laine 1970(II): 130). On the

other hand, in Gjærevoll’s (1956: 29)

classification the species belongs to the

heath series, i.e. being only season-

hygrophilous. It is common in several kinds

of snow-bed communities, but at least in

Pechenga and in the Scandes especially

characteristic to relatively late-melting sites

characterized by Salix herbacea and Carex

bigelowii (Kalliola 1939: 162, Gjærevoll

1956: 398 and Table 9). Other typical

associates among vascular plants are e.g.

Bistorta vivipara, Gnaphalium supinum,

Sibbaldia procumbens and Tofieldia

pusilla, among mosses e.g. Bryum

pseudotriquetrum, Dicranoweisia crispula

and Pohlia wahlenbergii, small hepatics

and lichens like Baeomyces placophyllus,

Cetraria delisei, Cladonia macrophylla, C.

stricta, Solorina crocea and crustose

lichens. Cassiope may descend to the birch

belt along watercourses, and is met close to

the waterline on relatively open sites on

moist gravel.

The species is not demanding as to the

nutrient content of the soil (Rintanen 1968:

286). It prefers humid and cool micro-

climate, and according to Dahl (1951: 36),

it cannot tolerate maximum temperatures

exceeding +26 °C – a likely reason for the

sharp limit of the distribution at the E

margin of the Scandes. However, if the soil

is humid enough, evaporation will cool

down the surface temperatures to a more

tolerable level (Rintanen 1968: 286).

In Utsjoki, flowering starts usually in

mid-June (Alanen 2007). In the alpine belt

in Pechenga the flowering is at its best in

early July, in late-melting sites not until the

end of the month, and seed ripens in ca.

eight weeks; seed production and

germination are mainly good, but seedlings

in natural habitats are not very abundant;

vegetative regeneration by rooting branches

is common (Söyrinki 1939a: 324–326).

Amphicline/indifferent (Arwidsson

1943: 230, Benum 1958: 310, Wistrand

1962: 131, Laine 1970(II): 130, Mäkinen &

Kallio 1979: 17, Roweck 1981: 361,

Nilsson 2000: 155, Lid & Lid 2005: 602) or

somewhat indifferent (“preferential

species” in the series poor in calciphiles;

Gjærevoll 1956: 28). pH range 4.2–5.1

(Kalliola 1939: 164, Gjærevoll 1956: 126–

127, 136, Rintanen 1968: 286), but no

records from the study area. Amphicline.

Dependence on culture. Ahemerobe.

MP

82

REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Kalmia polifolia Wangenh.

Introduced, very rare

Map 56

Distribution. Native to eastern North America: NE

U.S.A. and most of Canada except for British

Columbia, Yukon and Nunavut (Fl. North America 8:

485), cultivated for ornament in Europe and

naturalized in SE England (Fl. Eur. 3: 10, Blamey &

Grey-Wilson 1989: 292, Valdés 2009). No earlier

records from Finland, other Nordic countries or NW

Russia outside cultivation.

InL ref. New to Inari Lapland.

TUR 1 spec.

Very rare (1; 0.000). Utsjoki: I (1;

0.000). Utsjoki: Kevo, one small stand ca.

400 m SW from the Kevo Subarctic

Research Station (7741:3500, 2005 M.

Alanen, TUR 149020). Hemerochore.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Alt. 105 m. Silvine.

Ecology. The habitat of Kalmia

polifolia in Kevo is an experimental field

for testing birch species and provenances,

cleared of a dwarf shrub bog. The species

was probably introduced either as seed or

seedling(s) with birches brought from

Schefferville, Quebec, Canada in 1978.

Kalmia was not found until 2005, probably

due to a long juvenile period in the harsh

conditions. There may be only one clone of

the species, but the underground parts of

the plant have not been checked. In 2005,

there were a total of 10 aerial shoots in two

small groups at a distance of ca. 30 cm

from each other. Due to the presence of dry

old capsules and counted from the annual

growth, it seems that the plant had flowered

for the first time in 2002. In 2005,

flowering had already begun on June 27,

and was ending on July 8. In 2006, it

started on June 14 and continued at least for

two weeks (Fig. 8). Part of the capsules did

not develop in 2005, but there were two

Fig. 8. Kalmia polifolia as a partly established anthropochore in an experimental field close to the Kevo

Subarctic Research Station. NB: Remnants of the capsules of the previous year on the right branch. Photo

28.6.2006 S. Heino.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 83

capsules with ripe seeds by the end of

August. Flower buds for the next season

were visible in September. However, it

seems that the plants are not thriving very

well. In late July 2011, the plants were still

extant and with unripe new capsules, but

the condition of the stand was not checked

more thoroughly.

Dependence on culture. Probably

accidentally introduced in 1978. Because of

the life form of the plant, it is difficult to

judge the state of establishment. Probably

ephemerophytic anthropochore.

MP

Ledum palustre L.

Rhododendron palustre (L.) Kron & Judd,

non Turcz.

Ledum tomentosum Stokes, Rhododendron

tomentosum Harmaja

Indigenous, very frequent

Map 57

Distribution. Continental circumpolar (Fl. USSR

XVIII: 30, Dutilly et al. 1953: 82, Jørgensen et al.

1958: 90, Wiggins & Thomas 1962: 285, Porsild

1964: 130, Ohwi 1965: 695, Calder & Taylor 1968:

462, Hultén 1968: 717, 1971b: 80, map 71, Savile

1969, Hustich 1970, Andrulaїtis et al. 1976: 80,

Malyšev 1976: 80, Mäkinen & Kallio 1978, Roweck

1981: 364, Hultén & Fries 1986: map 1451, Aiken et

al. 2007). A gap in E Greenland, Iceland, most of the

British Isles and W Norway (Hultén & Fries 1986:

map 1451). In Fennoscandia mainly in Sweden and

Finland; in Norway almost totally confined to the

north (Hultén 1971a: map 1367, Norman I(1): 748,

Dahl 1934: 376, Rønning 1954, Benum 1958: 308,

Ryvarden 1969: 34, Mossberg & Stenberg 2003:

455). Two subspecies: subsp. palustre (N Europe – N

Asia) and subsp. decumbens (Aiton) Hultén (NE Asia

– N America – W Greenland); see under Morphology

and taxonomy.

Very rare in the inner parts of Troms, scattered

– fairly frequent in the inner parts of Finnmark (Lid

& Lid 2005: 602, Elven et al. 2013: 337). Rather rare

– fairly common in Pechenga (Valle 1930, 1933a,

Kalliola 1932: 106, Söyrinki 1939a: 317, Piirainen

1996b, Alm et al. 1997: 41). Common in the Kola

Peninsula (Fl. Murm. IV: 288, map 97, Mäkinen

2002: 17), Keret Karelia (Söyrinki 1956: 31) and

Kovda area (Sokolov & Filin 1996: 123).

Mostly common to very common in Kittilä,

Sompio and SE parts of Enontekiö Lapland, scattered

to fairly rare in the NW parts of Enontekiö (Hjelt &

Hult 1885: 136, Wainio 1891: 45, Hult 1898: 164,

Hjelt 1919: 395, Roivainen 1923: 293, Hustich

1937a: 62, 1940c: 58, Lindén 1943: 78, Ruuhijärvi

1960: e.g. 154-157, Pertola 1961: 35, Montell 1962:

125, Hämet-Ahti 1963a: 110, 113, Kujala 1964: 81,

Piirainen & Piirainen 1991b, Lampinen & Lahti

2018).

InL ref. Common (Kihlman 1884:

110), fairly common (Mikkola 1941: 33).

In the upper Lemmenjoki fairly common

(Klockars & Luther 1938). In W Utsjoki

fairly rare (Laine et al. 1955: 130) –

common (Kallio & Mäkinen 1957: 26).

Along the Ivalojoki rare but fairly common

at the mouth (Kujala 1962: 177), in the

Vaskojoki area very common (Laine 1964:

115). In the Kevojoki area scattered – fairly

abundant, but rare in the most alpine parts

(Laine 1970(II): 129, Heikkinen & Kalliola

1990: 48).

Kevo 72.2 %, InL 95 %, 258 sq. H 13,

KUO 1, OULU 2, TUR 21, YME 22 spec.

Very frequent (4510; 0.695). Inari: VII

(3034; 0.707), Utsjoki: VII (1476; 0.670).

Difference**. Although the species is fairly

evenly distributed over the area, it is

somewhat more frequent in Inari. Whole

area.

FMF 0.982.

Vertical distribution. a: VI (342;

0.467), b: VII (1698; 0.749), c: VII (2470;

0.709). Differences***. Range 15 m (S end

of Lake Pulmankijärvi, 7762:3539) – ca.

600 m (Karegasnjarga-Ailigas, Lanka,

7705:3460); “Hammasuro” 473 m

(Kihlman 1884: 110). Tr 900 m

(Engelskjøn & Skifte 1995: 148), Fnm 391

m (Norman I(1): 748), EnL 825 m (Väre &

Partanen 2009: 91). In Pallas-Ounastunturit

often above the forest line (Hustich 1940c:

58), 690 m (Hustich 1937a: 62). Vertical

ubiquitous.

Ecology. In N Finland Ledum palustre

84 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

is a typical and often dominant species in

numerous bog types, forming often tight

and uniform stands. Söyrinki et al. (1977:

21, 22, 24) describe a Ledum-Uliginosum

association, which belongs to fresh heath

forests, and (l.c. 80, 81) a Ledum dominant

pine bog especially in the area of spruce

forests. Even as a bog species, Ledum

usually forms stands around bog margins

near the mineral soil. Ruuhijärvi (1960: e.g.

128, 130, 131) describes “Ledum-reiche

normale Reisermoore”, in which Ledum

belongs to the most important species. It

also occurs in the alpine belt in many dwarf

shrub communities, but is not dominating

(cf. Kalliola 1939: 260). In N Utsjoki, it

may also grow on dry birch and pine

slopes. Often it grows also on bluff shelves

and rock crevices, together with e.g.

Calamagrostis lapponica, Epilobium

angustifolium and Solidago virgaurea.

In the alpine belt L. palustre flowers

clearly less than in the birch forest areas

(cf. Söyrinki 1939a: 317); it flowers in the

alpine belt but does not produce seeds

(Norman II(1): 434). For flowering times in

Pechenga, see Valle 1930. – Ledum is

usually considered slightly acidocline

(Mäkinen & Kallio 1979: 17, Nilsson 2000:

156), in the Kevojoki valley amphicline

(Laine 1970(II): 129). Slightly acidocline.

Parasites. Ledum palustre is attacked

by the rust fungus Chrysomyxa woroninii

Tranz. (II), observed in seven localities in

the Lake Inari basin (Mäkinen 1964b: 157).

The rust occurs here independent of host-

alternation and overwinters in the

uredostage (the aeciostage on Picea abies

has also been collected in Inari).

Morphology and taxonomy. Recent

phylogenetic studies and cladistical

analyses have shown that Ledum must be

regarded as a subsection of Rhododendron

(Kron & Judd 1990, Kurashige et al. 2001).

The correct name for Ledum palustre L. is

thus Rhododendron tomentosum (Stokes)

Harmaja (Harmaja 1990, 1991). In modern

floras this name is unanimously accepted

(Georgson et al. 1997: 479, Rydberg &

Wanntorp 2001: 454, Bertilsson et al. 2002:

445, Mossberg & Stenberg 2003: 455, Lid

& Lid 2005: 601, Fröberg 2006: 501,

Edqvist & Karlsson 2007: 539, Tyler et al.

2007: 421, Hæggström & Hæggström

2010: 192, Jonsell 2010: 468, Lidberg &

Lindström 2010: 342, Stenberg 2010: 568,

Blomgren et al. 2011: 419, Elven et al.

2013: 336, Löfgren 2013: 532). Here,

however, the nomenclature of Hämet-Ahti

et al. (1998) is followed.

The NE Asian – N American subsp.

decumbens has been suggested to occur

also in N Asia and N Europe, including N

Fennoscandia (Hultén 1971b: 80, map 71,

Fl. Arct. URSS VIII: 112, map 45, Hultén

& Fries 1986: 1104, map 1451). However,

it is omitted or not accepted for Europe by

most authors, even Hultén himself (Benum

1958: 308, Fl. Murm. IV: 287-288, Hultén

1968: 717, 1971a: map 1367, Fl. Eur. 3: 9,

Roweck 1981: 369, Hämet-Ahti et al. 1998:

209, Mossberg & Stenberg 2003: 455, Lid

& Lid 2005: 602, Elven et al. 2013: 338,

Elven & Murray in Elven 2016).

Descriptions of the subspecies are given

e.g. by Wiggins & Thomas (1962: 285),

Porsild (1964: 130), Böcher et al. (1968:

148) and Aiken et al. (2007). Keys for the

subsection Ledum are provided in Harmaja

(1991) and Voss (2011). In subsp.

decumbens the young twigs are brown and

puberulent, glabrescent in age. The leaves

are 5-20 mm long, 2-3 mm wide, oblong to

oblong-linear, somewhat acute, shiny and

glabrescent above, woolly beneath, margins

strongly revolute. Petals are smaller than in

subsp. palustre; flowers are nodding and

calyx-lobes with hook-like reflexed apices;

and fruiting pedicels are abruptly hooked at

the apex. Plants in some collections from N

Finland resemble this description by their

habit. However, they have larger

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 85

inflorescences, longer pedicels and larger

flowers and capsules than N American and

NE Asian subsp. decumbens, and their

pedicels are more or less straight or only

curved. There also seems to be a gradual

transition from tall and relatively broad-

leaved plants in S Finland to smaller and

more narrow-leaved plants in the north. At

the same time, the plants occupy dryer

habitats in heath forests and subalpine

heaths.

In Canada, subsp. decumbens grows on

heaths and rocky places (Hultén 1968:

717); in British Columbia it is restricted to

dry alpine habitats (Calder & Taylor 1968:

462). In Alaska it occurs mainly on slopes,

river banks and dry tundra (Wiggins &

Thomas 1962: 285). In the Canadian Arctic

Archipelago (Porsild 1964: 130), subsp.

decumbens usually grows “in not too wet

dwarf shrub- or moss-lichen heath or on

sunny cliffs and ledges”. In Siberia, the

usual sites are in high mountains on stony

slopes, and on gravelly tundra (Andrulaїtis

et al. 1976: 80). In Japan, Ohwi (1965: 695)

mentions high moors in alpine regions as

typical sites for subsp. decumbens. Small

and narrow-leaved plants from Inari

Lapland have been collected from, e.g. pine

bog; moist, partly paludified site in

subalpine belt; and transitional zone

between Pinus woodland and shore mire

with big hummocks. These habitats do not

differ from those of taller and more broad-

leaved plants.

Elven et al. (2013: 338)

comprehensively discuss the existence of

subsp. decumbens in Fennoscandia and

come to the conclusion that it must be

excluded from Fennoscandia. Elven &

Murray in Elven (2016) suspect some

confusion between subsp. decumbens and

dry heath ecotypes or modifications of

subsp. palustre in N European Russia and

Fennoscandia, and according to them, also

in N European Russia the occurrence of

subsp. decumbens remains to be

convincingly documented (Elven 2016).

In lack of any well-defined

morphological or ecological differences we

accept only subsp. palustre in Inari

Lapland.

A deviating form, f. dilatatum

Wahlenb., has been found in N Finland

(also in Inari Lapland), in Pechenga and the

Kola Peninsula (Hiitonen 1933: 570, Valle

1933a, Lindén 1943: 78). It has flat, fairly

broad leaves which have only sparsely

brown hairs; the leaves may resemble those

of Vaccinium uliginosum.

Dependence on culture. The species

makes use of free habitats along the Utsjoki

highway, and also flowers abundantly in

these sites (Vanhatalo 1965: 123).

Hemeradiaphore.

YM, MP

Loiseleuria procumbens (L.) Desv.

Azalea procumbens L., Kalmia procumbens

(L.) Gift, Kron & P.F. Stevens ex Galasso,

Banfi & F. Conti

Indigenous, fairly frequent

Map 58

Distribution. Circumpolar arctic-alpine, with a wide

gap in Siberia (Hultén & Fries 1986: map 1452); may

also be regarded as amphi-Atlantic (Hultén 1958:

200). Common throughout the Scandes in Norway

and Sweden, and the northernmost parts of Finland,

scattered in the more isolated mountain areas E and S

of the main area (Hultén 1971a: map 1369).

Common in the Swedish part of the Scandes

and most of Norway but in the far south mainly in the

mountain areas, from Trøndelag northwards also in

lowlands (Mossberg & Stenberg 2003: 453, Lid &

Lid 2005: 603). Common in Troms (Benum 1958:

308, Engelskjøn & Skifte 1995: 148) and Finnmark

(Norman II(1): 432, Dahl 1934: 375, Alm 1993a: 30);

86 localities in the Rastigaissa area (Ryvarden 1969:

34). Common to very common in the alpine and

subalpine areas in Pechenga, rather rare to scattered

in the coniferous zone (Roivainen 1923: 292, Kalliola

1932: 106, Valle 1933a: 270, Söyrinki 1939a: 319).

Rather common in the northernmost parts of the Kola

Peninsula, in the mountain areas in the W south to

86 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

the surroundings of Kandalaksha, and along the E

coast (Fl. Murm. IV: map 98, Hultén 1971a: map

1369).

Common in the fjeld areas of the northern parts

of Sompio and Kittilä Lapland in the alpine and

subalpine belts, descends also to the coniferous zone;

rare in the solitary southern fjelds, e.g. Luosto (SoL),

Kätkätunturi and Yllästunturi (KiL) (Hjelt & Hult

1885: 136, Wainio 1891: 45, Hustich 1937a: 103,

1940c: 58, Vuori & Pertola 1959a, Montell 1962:

125, Rintanen 1968: 283, Lampinen & Lahti 2018).

The southernmost localities in Finland are in central

Kittilä Lapland, southern Sompio Lapland and SE

Koillismaa, one locality in NE Outer Ostrobothnia

(Noitatunturi fjeld; Lampinen & Lahti l.c., Kastikka

2018). Common to very common in Enontekiö

Lapland, except for the southernmost parts where

only scattered (Hustich 1937a: 103, 1940c: 58,

Lindén 1943: 78, Laine 1958: 87, Piirainen &

Piirainen 1991b, Lampinen & Lahti l.c.).

InL ref. “So common in Utsjoki that

mountains turn red in the early summer”

(Fellman 1835: 253), very common and

abundant in the alpine belt, common to

rather common in the subalpine belt

(Kihlman 1884: 110, Mikkola 1941: 33).

Mentioned from the Hammastunturit fjelds

and Veskoniemi area by Wainio (1891: 45).

Common in the SE parts of Inari Lapland

(Rintanen 1968: 283) and in the

Viipustunturit–Maarestatunturit area, also

occurring in the forest region (Klockars &

Luther 1938). Several localities near

Kultala in the valley of the Ivalojoki

(Kihlman 1884: 110), very rare along the

central course of the river (Kujala 1962:

177); abundant localities in heath forests in

the area of the Ivalojoki (Rintanen 1968:

283). Rather common in the Peäldoajvi

fjeld area (Koivistoinen 1964: 53). In W

Utsjoki rather common (Laine et al. 1955:

130, Kallio & Mäkinen 1957: 26) to

common (Kuitunen 1984), in the Kevo

Strict Nature Reserve very common

especially in the alpine areas (Heikkinen &

Kalliola 1990: 35), but rather rare in the

canyon area itself (Laine 1970(II): 129);

“rarer than expected” in the coniferous

forests by the Utsjoki (Hustich 1942a: 225).

Kevo 78.8 %, InL 61 %, 179 sq. H 59,

JYV 6, OULU 14, TUR 78, VOA 1, YME

8 spec.

Fairly frequent (2161; 0.338). Inari:

IV (771; 0.181), Utsjoki: VII (1390; 0.650).

Difference***. Very common throughout

the N parts of Inari Lapland. In Inari parish

almost exclusively confined to the fjeld

areas, almost missing from the Lake Inari

basin except for the low outer islands of

Lake Inari (Mikkola 1941: 33). Northern.

FMF 0.705.

Vertical distribution. a: VII (640;

0.841), b: VII (1233; 0.570), c: III (290;

0.083). Differences***. Very common in

the alpine and birch belts, rather rare in the

coniferous zone. Range 15 m (Lake

Pulmankijärvi, 7762:3539) – ca. 600 m

(Karegasnjarga-Ailigas, Lanka,

7705:3460). Tr 1150 m (Benum 1958:

308), Fnm 827 m (Norman I(1): 744), EnL

1050 m (Laine 1958: 87, Väre & Partanen

2009: 94). Alpike.

Ecology. Loiseleuria procumbens is a

typical plant of dry windswept fjeld heaths,

as well on till as on gravel and sand. It is

most common in the alpine belt at the top

of hills and ridges, and on open dry gravel

fields. In Haapasaari’s (1988: App. Table

1.) material, typical habitats are deflation

sites (and other windswept sites), where it

forms tight prostrate patches and is often

accompanied by Arctostaphylos alpina,

Empetrum nigrum subsp. hermaphroditum,

Juncus trifidus, Vaccinium vitis-idaea,

Polytrichum piliferum, Alectoria nigricans,

A. ochroleuca, Cetraria nivalis, Cladonia

mitis and Sphaerophorus globosus.

Oksanen & Virtanen (1995: 18) named this

kind of continental deflation sites as their

Empetrum-Loiseleuria type (ELT), which is

typical to hemiarctic and low arctic ridges

(l.c.: 37, figs. 25 and 26), and considered

them identical with Kalliola’s (1939: 175)

lichen-rich Diapensia-Loiseleuria-

Empetrum association. The species is

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 87

common also in other chionophobous

communities described e.g. by Haapasaari

(1988), most notably in the subcontinental

arctic Empetrum-Cetraria nivalis type and

its hemiarctic counterpart Empetrum-

Lichenes type. Other associates in these two

vegetation types are e.g. Betula nana,

Calamagrostis lapponica, Dicranum

elongatum, D. fuscescens, D. scoparium,

Polytrichum hyperboreum, Cetraria

cucullata, C. ericetorum, Cladonia

uncialis, Ochrolechia frigida and O.

geminipara; Cladonia stellaris may also be

locally common. Kalliola (1939: 174)

grouped this kind of extremely xerophilous

vegetation types as the alliance

Loiseleurieto-Arctostaphylion. Loiseleuria

may also grow in moister habitats, even in

moderate snow beds, where it, however, is

scarce and usually sterile (Söyrinki 1939a:

319). Sometimes it is also met in

oligotrophic bogs.

Loiseleuria also grows on rocky slopes

and on the gravelly shores of brooks and

lakes, it is characteristic of the low outer

islands of Lake Inari (Mikkola 1941: 33).

Towards the north it becomes more

common also in open patches in birch and

pine forests. In the coniferous zone it is not

concentrated in the lower parts of river and

brook shores but grows at or above the

upper flood margin in normal heath

vegetation (Vuori & Pertola 1959a,

Rintanen 1968: 283). In bogs it may be

found on dry hummocks.

Flowering usually starts during the first

half of June in the surroundings of the

Kevo Subarctic Research Station, in early

summers already in late May, and lasts for

3–4 weeks. Seeds ripen from late August

till late September (Alanen 2007). In the

alpine belt in Pechenga the flowering takes

place ca. two weeks later, and seeds ripen

in ca. two months; seed production and

germination are good (Söyrinki 1939a:

319–321).

Amphicline/indifferent (Norman II(1):

432, Arwidsson 1943: 229, Benum 1958:

308, Wistrand 1962: 130, Roweck 1981:

365, Laine 1970(II): 129, Nilsson 2000:

156), both on acid and base-rich soil (Lid &

Lid 2005: 602). pH range 3.9–5.2 (Rintanen

1968: 283); with a wide pH range, but the

weight is at the acidic side (SKK III: 272);

acidocline/calcifuge (Fl. Eur. 3: 10,

Mäkinen & Kallio 1979: 17). Acidocline.

Morphology. White-flowered plants

are rather rare but probably unrecorded;

only one herbarium specimen: Utsjoki,

Kaimmioaivi (7735:3477, 1989 M.

Piirainen 564 & P. Piirainen, H 645245).

Field notes made by L. Iso-Iivari 2002

(7756:3548) and 2004 (7714:3482);

recorded from Utsjoki also by Kallio &

Mäkinen (1957: 26).

Dependence on culture. Hemera-

diaphore.

MP

Phyllodoce caerulea (L.) Bab.

Andromeda caerulea L., Bryanthus

caeruleus (L.) Dippel, Menziesia caerulea

(L.) Sw.

Indigenous, frequent

Map 59

Distribution. Arctic-montane, amphi-Atlantic or

circumpolar with some gaps in the distribution

(Hultén 1958: 228, Hultén & Fries 1986: map 1453).

Common along the Scandes and in the northernmost

parts of Fennoscandia, in the north also in lowlands

(Hultén 1971a: map 1370, Roweck 1981: 365–366,

Nilsson 2000: 156, Mossberg & Stenberg 2003: 453,

Lid & Lid 2005: 602). Otherwise in Europe only in

isolated stations in Scotland and the Pyrenees (Fl.

Eur. 3: 10, Blamey & Grey-Wilson 1989: 292).

Common and mostly abundant in Troms but

less common in the SW lowland (Benum 1958: 309,

Engelskjøn & Skifte 1995: 149). Common over most

of Finnmark but missing in parts of the coastal areas

(Dahl 1934: 375, Alm 1993a: 33); 65 localities in the

Rastigaissa area (Ryvarden 1969: 34). Common in

the alpine and subalpine areas in Pechenga, but rather

rare to scattered in the coniferous zone (Roivainen

88 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

1923: 292, Kalliola 1932: 106, Valle 1933a: 270,

Söyrinki 1939a: 322). Common in the N part of the

Kola Peninsula and in the mountain areas in the W

and NW (Fellman 1831: 309, Söyrinki 1956: 31, Fl.

Murm. IV: map 99, Hultén 1971a: map 1370,

Tolmatchev 1980: 123).

Common everywhere in Finnish Lapland

(Fellman 1835: 262). Common to very common in

the N parts of Sompio and Kittilä Lapland in the

alpine and subalpine belts, rare to scattered in the

upper parts of the coniferous zone (Wainio 1891: 45,

Hult 1898: 164, Hustich 1937a: 103, 1940c: 58,

Kujala 1961, Montell 1962: 125, Rintanen 1968: 284,

Lampinen & Lahti 2018). Common to Pallastunturit

and Saariselkä fjeld areas in the south, scattered

further south to southern Sompio Lapland and

Koillismaa (Hjelt & Hult 1885: 136, Auer 1944,

Kujala 1961, Lampinen & Lahti l.c.). Common to

very common in Enontekiö Lapland (Lindén 1943:

78, Laine 1958: 87, Virtanen & Väre 1990, Piirainen

& Piirainen 1991b, Lampinen & Lahti l.c.). Three

localities in N and NE Outer Ostrobothnia (Kastikka

2018, Lampinen & Lahti l.c.).

InL ref. Common in Inari Lapland

(Mikkola 1941: 33), common to very

common in the subalpine and alpine belts,

scattered in the coniferous zone (Kihlman

1884: 110). Rather rare along the Ivalojoki

and in Veskoniemi (Wainio 1891: 45,

Kujala 1962: 177), mentioned from the

Sotajoki area by Rintanen (1968: 284).

Common to very common in the

Viipustunturit – Maarestatunturit area

(Klockars & Luther 1938), rather common

in the Vaskojoki area (Laine 1964: 115).

Common in the Peäldoajvi fjeld area

(Koivistoinen 1964: 54). Rather common

between Suolisvuono and Pakanajoki

(Kihlman l.c.). In SW Utsjoki very

common (Laine et al. 1955: 130) to

common (Kallio & Mäkinen 1957: 26,

Kuitunen 1984), scattered in the Kevojoki

canyon (Laine 1970(II): 129).

Kevo 95.3 %, InL 76 %, 208 sq. H 55,

JYV 8, KUO 13, OULU 15, TUR 77, VOA

3, YME 10 spec.

Frequent (3153; 0.490). Inari: V

(1479; 0.346), Utsjoki: VII (1674; 0.775).

Difference***. Common nearly throughout

the area but largely missing in the Lake

Inari basin. Whole area.

FMF 0.830.

Vertical distribution. a: VII (652;

0.871), b: VII (1690; 0.760), c: IV (811;

0.235). Differences***. Most common in

the alpine and subalpine belts, significantly

less frequent in the coniferous zone. Range

15 m (Lake Pulmankijärvi, 7762:3539) –

ca. 600 m (Karegasnjarga-Ailigas, Lanka,

7705:3460). Tr 1390 m (Engelskjøn &

Skifte 1995: 149), Fnm 884 m (Norman

I(1): 739), EnL 1150 m (Laine 1958: 87).

Vertical ubiquitous.

Ecology. Phyllodoce caerulea is

typical to dryish and mesic alpine heaths

and shows slight oceanic or hygric

tendencies (Kalliola 1932: 106, Hämet-Ahti

1963a: 65, SKK III: 273, Haapasaari 1988:

70), however Kärenlampi & Kauhanen

(1972: 89) suggested that it may be

concentrated to the subxeric parts of the

moisture gradient. Haapasaari (l.c.) named

an arctic treeless heath type dominated by

the species as Phyllodoce type,

characterized by a mosaic-like structure

composed of Phyllodoce patches and

intermediate ground-layer spaces

dominated by either Stereocaulon paschale

and several Cladonia species or by

bryophytes. In exposed sites the ground

may be relatively bare. Betula nana,

Empetrum nigrum subsp. hermaphroditum,

Vaccinium myrtillus and V. vitis-idaea are

usually present. The only constant

herbaceous vascular plant is Juncus

trifidus, in Haapasaari’s relevées from Inari

Lapland also e.g. Deschampsia flexuosa

and Festuca ovina are present. In these

relevées the most common bryophytes are

Dicranum fuscescens, Polytrichum

juniperinum, P. piliferum and P. strictum,

and lichens Cetraria ericetorum, C. nivalis,

Cladonia bellidiflora, C. coccifera, C. mitis

and Stereocaulon paschale. In Kalliola’s

material from Pechenga Phyllodoce is most

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 89

common in moss-rich alpine heaths, where

it is met e.g. in several associations of his

Phyllodoceto-Vaccinion myrtilli (Kalliola

1939, e.g.: 214, 218, 220, 227). Oksanen &

Virtanen (1995: 12) named also a

Phyllodoce-Myrtillus group of treeless

heaths where the field layer is dominated

by evergreen or semi-evergreen dwarf

shrubs.

Typical habitats in the birch belt are

the continental subalpine Empetrum-

Lichenes (sELiT), Empetrum-Lichenes-

Pleurozium (sELiPlT) and Empetrum-

Myrtillus (sEMT) associations, with e.g.

Deschampsia flexuosa, Empetrum nigrum

subsp. hermaphroditum, Festuca ovina,

Vaccinium uliginosum and V. vitis-idaea,

and cryptogams Dicranum scoparium,

Polytrichum juniperinum, P. piliferum,

Barbilophozia hatcheri, B. lycopodioides,

Cladonia stellaris, Peltigera aphthosa and

Stereocaulon paschale. Phyllodoce is

relatively often met also in oceanic Cornus-

Empetrum-Myrtillus birch forests

(CoEMT), but it is lacking in the

submaritime birch forests (Hämet-Ahti

1963a: 40–44, 50, 72, 77).

Phyllodoce grows also in Dryas heaths,

rock crevices, dwarf shrub bogs, meadow

heaths, low-herb meadows along brooks

and rivers, but rarely in snow-beds, and in

heath forests and gravelly riversides in the

coniferous zone. Oksanen & Virtanen

(1995: 50) pointed out that soil

temperatures have an important role in

habitat preferences for the species: it favors

sites with a snow cover thick enough to

prevent soil freezing, which gives it

advantage in early-spring growth start.

Flowering starts in the surroundings of

the Kevo Subarctic Research Station

usually in mid-June and lasts till early July,

the earliest recorded date being May 29, the

latest July 13 (Alanen 2007); the best time

for flowering is between June 28 and – July

7 (Hustich 1942a: 220). E.g. in 1797

flowering was recorded on June 26, and in

1795 on July 10 in Utsjoki (Castrén 1803).

Flowers are possibly largely autogamous

(Hagerup 1954). Seeds ripen usually in late

August to early September (Alanen 2007).

In Pechenga flowering is usually at its best

in late June – early July (Valle 1930,

1933b, Söyrinki 1939a: 322), and seeds

ripen in early September. Seedlings are

common though not abundant; vegetative

propagation may probably happen only in a

small scale because of the weak or missing

adventive root system (Söyrinki l.c.: 322-

323). On the other hand, the weak

adventive root system favors the species in

competition against e.g. Vaccinium vitis-

idaea on instabile soils. Kujala (1964: 81)

was of the opinion that seedling

establishment demands bare soil, but

Rintanen (1968: 284) pointed that the

occurrences in paludified forests may

suggest this is not the case.

Amphicline/indifferent (Arwidsson

1943: 229, Benum 1958: 309, Wistrand

1962: 130, Laine 1970(II): 130, Mäkinen &

Kallio 1979: 17, Nilsson 2000: 156), both

on acid and base-rich soil (Roweck 1981:

366, Lid & Lid 2005: 602). Amphicline.

Morphology. White-flowered plants

have been recorded from Inari, Peäldoajvi

(767:348, 1950 K. Laurila, H 556489), and

from Utsjoki, three localities in the W parts

(Kallio 1954, Laine et al. 1955: 130), NE of

Njavgoaivi (7713:3475, 2002 L. Iso-Iivari,

not.), Stuorra Piesvarri (7714:3464, 1954

N. Tarén, TUR 165712), and plants with

nearly white flowers from SE of

Skirratsohkka (7743:3495, 1971 K. & U.

Laine, TUR 264139) and NE Kistuskaidi

(7764:3486, 2002 L. Iso-Iivari, not.).

Dependence on culture. Spreads

readily to bare soils excavated by e.g. gold-

diggers (Rintanen 1968: 284). Probably

benefits from heavy reindeer-grazing (SKK

III: 273). Hemeradiaphore.

MP

90 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Rhododendron lapponicum (L.)

Wahlenb.

Indigenous, very rare

Map 60

Distribution. Amphi-Atlantic (Hultén 1958: 200, Lid

& Lid 2005: 601); might also be regarded as a

circumpolar arctic-montane plant with a wide gap in

NE Europe and W Siberia (Hultén & Fries 1986:

1103, map 1450). The plants from NE Siberia and

NW North America probably represent a different

taxon, R. lapponicum subsp. alpinum (Glehn) A.P.

Khokhr. (R. parvifolium Adams, see Elven 2016). In

Fennoscandia very isolated and bicentric (Fries 1913:

319, Nordhagen 1936b: 96; Hultén 1971a: map 1368,

Rune 1963: 239, Berg 1963: 161, Roweck 1981: 362,

Nilsson 2000: 156, Mossberg & Stenberg 2003: 454);

the southern area is small and restricted.

Fairly common in Troms (Benum 1958: 308),

in Finnmark only in the western parts (Dahl 1934:

376, Gjærevoll 1990: 97). Neither found in Pechenga

nor in the Kola Peninsula. Rare in NW Enontekiö

Lapland (Fellman 1835: 262, Laine 1958: 87, Hultén

1971, map 1368, Väre et al. 2008: 71, 77, 2010: 110,

116, Lampinen & Lahti 2018; H, TUR). Locally

common in Swedish Lapland, e.g. in Lule and Pite

lappmark (Björkman 1939: 122, 1965: 78, Arwidsson

1943: 229, Wistrand 1962: 130, Karlsson 1973: 88).

InL ref. First recorded in Inari Lapland

by Helenius (1948), who found the species

on the fjeld Kistuskaidi in NW Utsjoki (NE

end of Kistuskaidi 453 m, N slope at ca.

400 m a.s.l., 1948 O. Helenius, H 414826).

The record is the easternmost one in

Europe. In the other collections from that

time (1948 P. Rantasuo, OULU 48799,

1949 L. Itkonen, OULU 48800, 1949 R.

Parviainen, OULU 48797, 1949 P.

Rantasuo, OULU 48798, and 1949 A.

Salonen, H 615940) only “Kistuskaidi” is

mentioned in the label and they can only be

located to the 10×10 km² square 776:348.

Since 1956, the species has been found in a

few more areas, all belonging to the same

fjeld group. In most sites the species has

been sparse, with only a few or at most a

few tens of individuals. The localities of

1956 are not precise due to the inaccurate

maps available at that time.

The report from Tsuomasvarri, NE

Utsjoki (e.g. Lahti et al. 1995, Ryttäri &

Kettunen 1997: 227, Lampinen & Lahti

2010 and earlier versions of the atlas) is

based on an incorrect interpretation of the

text of Kalliola (1961: 117). Also the dot in

Inari (Hultén 1971a: map 1368, Gjærevoll

1990: 97) is erroneous. Of the three 10×10

km² squares in Lahti et al. (1995) only one

(776:348) is correct, others (776:351 and

776:354) are erroneous.

InL 0 %, 3 sq. H 2, OULU 4, TUR 3,

YME 1 spec.

Very rare (4; 0.001). Utsjoki: I (4;

0.002). Rhododendron lapponicum grows

in three areas on the N slope of Kistuskaidi

fjelds: (1) the westernmost top, on Dryas

heath above the steep N slope, rather

sparsely, with Carex capillaris, C.

glacialis, C. rupestris (7762:3482, 1956 Y.

Mäkinen, TUR 78417); (2) 1 km ENE of

the middle top, bare ground on dry alpine

heath, ca. 25 individuals, with Carex

capillaris, C. rupestris, Dryas octopetala

(7762:3484, 1956 Y. Mäkinen, TUR

78418); (3) N side of the E top, dry, N

facing slope in the alpine belt, alt. 260 m

(7763:3486, 2001 M. Alanen, TUR

565117); (4) N of the E top, sparsely on dry

Dryas heath, with Carex bigelowii, C.

glacialis, C. rupestris, Pinguicula alpina

(7764:3486, 1956 Y. Mäkinen, YME 4984,

and 2001 M. Alanen, field list). The

easternmost area is the largest one,

covering ca. 17 hectares and including four

sub-areas: the largest one is at an altitude of

ca. 250 m, with one smaller area above and

two smaller ones below the main area

(Alanen 2010). The first record from Inari

Lapland (1948 O. Helenius, see above) is

probably from this or the previous square. –

Without flowers, the plant is very difficult

to see in the field; the leaves of Loiseleuria

procumbens, Phyllodoce caerulea,

Vaccinium vitis-idaea, and especially

Arctostaphylos alpina may have the same

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 91

coloring as those of R. lapponicum (Alanen

2010). Northern.

FMF 0.002.

Vertical distribution. a: I (4; 0.005).

Range 190 m – ca. 350 m. The altitude

mentioned in Helenius’ specimen (400 m

a.s.l.) is probably based on the erroneous

maps of the time; the highest top in the NE

part of Kistuskaidi is less than 350 m. – Tr

1170 m (Engelskjøn & Skifte 1995: 148),

Fnm 436 m (Norman I(1): 747), EnL 950 m

(Urtasvarri, Laine 1958: 87, Väre &

Partanen 2009: 91). In N Norway and in N

Sweden R. lapponicum typically occurs in

the lower alpine and in the subalpine belt,

between ca. 250 m and 500 m (Dahl 1934:

376, Björkman 1939: 122, 194, 1965: 78),

rising up to 1320 m in Lule lappmark

(Selander 1950b: 112). Alpine.

Ecology. Typical sites on Kistuskaidi

are gravelly half-open heaths, with several

basocole or at least basocline vascular

plants, like Carex capillaris, C. parallela,

C. rupestris, Dryas octopetala and

Saxifraga oppositifolia. Also Diapensia

lapponica, Loiseleuria procumbens, and (in

moister places) Pinguicula alpina belong to

the companions (Helenius 1948). The total

plant cover is often only 50-70 %.

There is often only a thin snow cover

in the habitats of R. lapponicum during the

winter. As a result, the branches seldom

grow higher than 5-7 cm. The species

spreads effectively with subterraneous,

horizontal runners, which may be more

than 1 meter long. The flowering starts

already in the beginning of June and ends

in the end of June. The first flowers open

when the +5°C thermal sum at Kevo has

reached the value 90 (Alanen 2010; Fig. 9).

The total number of inflorescences in the

easternmost area in 2002 was 875, each

with 2.9 flowers in the average according to

the count in 2001.

Fig. 9. A flowering Rhododendron lapponicum L. with opened capsules of the previous year on Kistuskaidi

fjelds, Utsjoki. Photo 7.6.2002 S. Heino.

92 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

The species is generally described as

strongly basocole (Fries 1925: 8, Björkman

1939: 122, Wistrand 1962: 130, Mäkinen &

Kallio 1979: 17, SKK III: 269, Nilsson

2000: 156). Several authors stress that it

clearly requires calcium in the substrate

(e.g. Björkman l.c.). The acidity of the soil

has been found to be pH (4.8 –) 7.2 – 8.1

(Karlsson 1973: 38), pH (4.8 –) 5.2 – 7.3 (–

7.7) (Lunde 1962: 65), pH 6.1 – 6.8

(Coombe & White 1951: 40), pH 5.8 – 6.0

(Laine 1958: 87). Gjærevoll (1961: 23)

points out that R. lapponicum grows on Mt.

Javreoaivi, the richest fjeld in N

Scandinavia. However, the species may

also occur in oligotrophic sites together

with more acidophilous species (Karlsson

1973: 88). In S Norway R. lapponicum has

several localities in pine forests (Gjærevoll

1990: 98). Basocole.

Dependence on culture. Because of

the rarity of the species and the small size

of the populations, the plant is protected by

law in Finland and classified as near

threatened (NT) (Rassi et al. 2010: 200, see

also Ryttäri & Kettunen 1997: 226 and

Ryttäri et al. 2012: 359). Threatened by

picking and erosion of the habitats caused

by trampling. Hemerophobe.

YM

Vaccinium microcarpum (Turcz. ex

Rupr.) Schmalh.

Oxycoccus microcarpus Turcz. ex Rupr.,

Vaccinium oxycoccos L. subsp.

microcarpum (Turcz. ex Rupr.) A. Blytt &

O. C. Dahl

Indigenous, frequent

Map 61

Distribution. Boreal circumpolar (Hultén 1971b:

140, Hultén & Fries 1986: map 1459). Common in N

Fennoscandia, scattered in the S (Hultén 1971a: map

1381, Roweck 1981: 373, Mossberg & Stenberg

2003: 457).

Common – scattered in Troms and Finnmark

(Norman I(1): 710, Dahl 1934: 380, Benum 1958:

314, Ryvarden 1969: 34, Lid & Lid 2005: 607).

Common in Pechenga and the Kola Peninsula

(Kalliola 1932: 107, Valle 1933a, Fl. Murm. IV: map

112, Alm et al. 1997: 41, Mäkinen 2002: 17).

Scattered in Keret Karelia (Söyrinki 1956: 30),

common in the Kovda area (Sokolov & Filin 1996:

124).

Common all over Finnish Lapland except for

the fjeld areas (Kihlman 1884: 109, Hjelt & Hult

1885: 135, Wainio 1891: 46, Hult 1898: 163,

Roivainen 1923: 293, Hustich 1940c: 59, Lindén

1943: 77, Söyrinki 1939a: 330, Ruuhijärvi 1960,

Montell 1962: 125, Kujala 1964: 83, Söyrinki &

Saari 1980: 121, Piirainen & Piirainen 1991b,

Piirainen 1996b, Hämet-Ahti et al. 1998: 211,

Lampinen & Lahti 2018).

InL ref. “Copiosissime usque ad

terminum betulae” (Kihlman 1884: 109); in

W Utsjoki fairly rare (Laine et al. 1955:

130, Kallio & Mäkinen 1957: 26);

numerous swamps (Ruuhijärvi 1960).

Ivalojoki very common (Kujala 1962: 177),

Vaskojoki fairly common – scattered

(Laine 1964: 115), Kevojoki rather rare

(Laine 1970(II): 131).

Kevo 36.4 %, InL 89 %, 250 sq. H 11,

JYV 1, KUO 2, TUR 21, YME 1 spec.

Frequent (3100; 0.479). Inari: VI

(2288; 0.532), Utsjoki: V (812: 0.375).

Difference***. Although the species seems

to occur fairly evenly in whole area, it is

however very significantly commoner in

Inari. This mainly depends on the fact that

it avoids alpine fjeld areas, which are

commoner in Utsjoki than in Inari. Whole

area.

FMF 0.974.

Vertical distribution. a: IV (117;

0.162), b: VI (1089; 0.488), c: VI (1894;

0.540). Differences***, and thus the main

distribution is clearly in the coniferous zone

and the birch belt (cf. Heikkinen & Kalliola

1990: 35). The smaller abundance in the

alpine belt is at least partly connected with

the lack of suitable habitats. Range ca. 20

m (E side of Lake Pulmankijärvi,

7768:3538) – ca. 600 m (Karegasnjarga-

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 93

Ailigas, Lanka, 7705:3460). Tr 600 m

(Engelskjøn & Skifte 1995: 150), Fnm 528

m (Norman I(1): 713), EnL 780 m (Väre &

Partanen 2009: 98). In Pite lappmark very

rarely above the forest line (720 m, 780 m;

Arwidsson 1943: 231, Wistrand 1962: 132).

Silvike.

Ecology. Vaccinium microcarpum

thrives on markedly drier substrate than V.

oxycoccos (cf. Söyrinki 1939a: 330, Kujala

1964: 83, Roweck 1981: 373, Nilsson

2000: 158); this is in contrast with the

observation of Montell (1962: 125) made in

Muonio that the species often grow

together. They may occur in the same

10×10 m² area, but the actual habitat is

always clearly drier: V. microcarpum grows

usually on Sphagnum fuscum hummocks,

often in similar habitats as Drosera

rotundifolia and in the company of it (see

Mäkinen et al. 2005: 38); also Carex

pauciflora, Pinguicula villosa, Rubus

chamaemorus, Trichophorum cespitosum,

and in the S parts of Inari Lapland,

Empetrum nigrum subsp. hermaphroditum

may grow as associates. In the

classification of Kalliola (1939: 235) V.

microcarpum belongs to the ”Verband

Oxycocceto-Rubion chamaemori”, of which

he says: ”Dieser Verband ist der

oligotrophste und azidiphilste von allen”. V.

microcarpum may however also grow in

moss-rich mesotrophic fens with e.g.

Andromeda polifolia. In the alpine belt the

habitats are more often not on hummocky

but on even ground. Acidocole (Mäkinen &

Kallio 1979: 17), acidocline (Laine

1970(II): 132, Nilsson 2000: 158).

Acidocline or even acidocole.

Morphology and taxonomy. There

are three specimens in TUR collected in NE

Inari and identified as V. microcarpum ×

oxycoccos: (1) by the Harrijoki E of Lake

Nitsijärvi (7686:3547, 1960 J. Suominen,

TUR 78515), (2) 1 km S of Kuosnaoajvi

(7698:3553, 1960 E. Antikainen, TUR

78680, det. J. Nurmi 2010) and (3) E of

Karhuvaara (7710:3552, 1960 M-L.

Wallenius, TUR 78514).

Dependence on culture. Hemera-

diaphore.

YM

Vaccinium myrtillus L.

Indigenous, very frequent

Map 62

Distribution. Boreal Eurasiatic (Hultén & Fries

1986: map 1462). Common in N Fennoscandia

(Hultén 1971a: map 1377, Roweck 1981: 377,

Hämet-Ahti et al. 1998: 211, Mossberg & Stenberg

2003: 459).

Common and abundant almost everywhere in

Troms, Finnmark, Pechenga and the Kola Peninsula

(Norman I(1): 703, Hult 1898: l63, Benum 1958:

312, Dahl 1934: 380, Lid & Lid 2005: 607; Fl.

Murm. IV: map 106, Kontuniemi 1932: 40,

Roivainen 1923: 293, Alm et al. 1998: 135, Mäkinen

2002: 17), and in the Kovda area (Sokolov & Filin

1996: 124). Common in Sompio, Kittilä and

Enontekiö Lapland (Hjelt & Hult 1885: 135, Lindén

1943: 77, Montell 1962: 125, Pertola 1961: 125,

Laine 1958: 89, Hämet-Ahti 1963a, Virtanen & Väre

1990, Piirainen & Piirainen 1991b, Lahti et al. 1995,

Piirainen 1996b, Hämet-Ahti et al. 1998: 211,

Lampinen & Lahti 2018).

InL ref. All authors describe V.

myrtillus as common to very common in

Inari Lapland (Fellman 1835: 259, Kihlman

1884: 108, Wainio 1891: 46, Klockars &

Luther 1938, Laine et al. 1955: 130, Kallio

& Mäkinen 1957: 26, Kujala 1962: 177,

Laine 1964: 115, Laine 1970(II): 131,

Heikkinen & Kalliola 1990: 35, Kulmala

1999). The species belongs to the main

components of the Finnish forest vegetation

(Kujala 1964: 82).

Kevo 98.3 %, InL 94 %, 260 sq. H 11,

OULU 1, TUR 9, YME 3 spec.

Very frequent (5145; 0.793). Inari: VII

(3294; 0.766), Utsjoki: VII (1851; 0.846).

Difference***. Like many common plants,

V. myrtillus seems to be more frequent in

Utsjoki. This, however, is probably an error

94 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

due to the greater amount of incompletely

studied squares in Inari (see Introduction).

Whole area.

FMF 0.985.

Vertical distribution. a: VII (652;

0.868), b: VII (1930; 0.856), c: VII (2563;

0.736). Difference a-c***, b-c***. Range

15 m (Lake Pulmankijärvi, 7762:3539) –

ca. 600 m (Karegasnjarga-Ailigas, Lanka,

7705:3460). – Tr 1100 m (Engelskjøn &

Skifte 1995: 150), Fnm 860 m (Ryvarden

1969: 34), EnL 1000 m (Laine 1958: 89,

Väre & Partanen 2009: 99). Pite lappmark

1385 m (Karlsson 1973: 160). Vertical

ubiquitous, but commonest in the alpine

and birch belts.

Ecology. The most typical sites for V.

myrtillus are oligotrophic dwarf-shrub

heaths. It usually grows in small

depressions which are slightly moister than

the surrounding heath. In localities like this

V. myrtillus often forms small uniform

stands. It has given name for numerous

associations; typical associates belong to

the subalpine Empetrum-Myrtillus type

(Hämet-Ahti 1963a: 52; cf. Laine & Nurmi

1971), oceanic Cornus-Empetrum-Myrtillus

type (Hämet-Ahti 1963a: 70, 74), Cornus-

Myrtillus type (Hämet-Ahti 1963a: 74, 81),

Geranium-Myrtillus type (Kujala 1929: 52),

meadow-heath forests (Hämet-Ahti 1963a:

70, 74), and Verband Phyllodoceto-

Vaccinion myrtilli (Kalliola 1939: 209).

This small selection of various types (cf.

further Kalela 1939: 94-398, Roweck 1981:

377, Haapasaari 1988: 30-129) indicates

that V. myrtillus is common in numerous

subalpine and boreal associations, and is

also an important and characteristic species

in all of them. Other common associates

include Calamagrostis lapponica,

Deschampsia flexuosa and Linnaea

borealis (Laine 1970(II): 131).

Vaccinium myrtillus is also common in

the alpine belt but avoids the highest and

sterile alpine areas where it may, however,

grow at the margins of melting snow-beds,

often in the company of V. uliginosum. It

also avoids open and wet swamps and

flooded margins of brooks and rivers.

Karlsson (1973: 60) points out its

preference to southern expositions.

In the 19th century the flowering of

Vaccinium myrtillus in Utsjoki started

mostly in the second half of June (Moberg

1885: 145). During 1995-2006 around the

Kevo Subarctic Research Station the

earliest date recorded was May 28, the

latest June 21 (Alanen 2007). In the alpine

belt flowering may start as late as in the

middle of July. The berries begin to ripen in

the S part of Inari Lapland already in the

middle of July, in the Kevo area usually in

the end of July or in the beginning of

August (Alanen l.c.). Kontuniemi (1932:

40) describes the flowering and seed

germination in Pechenga.

In N Sweden ”neither amphicline nor

strongly acidophilous” (Karlsson 1973: 60).

Indifferent/amphicline (Laine 1970(II):

131, Mäkinen & Kallio 1979: 17, Nilsson

2000: 158). Amphicline.

Parasites. In Inari Lapland the species

is generally attacked by the powdery

mildew Podosphaera myrtillina (Schub.)

Kunze, which is probably the commonest

powdery mildew parasite in Finland.

However, it is very inconspicuous, because

the tiny cleistothecia (diameter 60-70 µm)

are always produced on the lower surface

of the leaves, and mildew-like conidial

mycelium is very rarely present. Especially

in the Lake Inari area this parasite is

common; there are 5 specimens in TUR

collected in Utsjoki and 34 specimens

collected in Inari. See Mäkinen 1969.

The rust Pucciniastrum vaccinii

(Wint.) Jørst. is also a common parasite,

although it is markedly rarer in Lapland

than in S Finland (Rauhala 1959: 148). Of

this rust, there are 5 collections from

Utsjoki and 6 from Inari in TUR, and in

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 95

addition 6 from Finnmark, Tana (see

Mäkinen 1964b: 173, 1964c). Kari (1936:

15) mentions it also from Pechenga (3

specimens in TUR).

The parasitic genus Exobasidium is

represented on V. myrtillus by three species

(Nannfeldt 1981: 39), of which two occur

in Inari Lapland. E. myrtilli Siegm. infects

leaves, which are enlarged and colored

white-cream to bright red. There are three

collections in TUR: two from Inari

(7629:3471 and 7729:3581) and one from

Utsjoki (7741:3475). E. aequale Sacc. is a

high-alpine species, which has only once

been found in Utsjoki (Kistuskaidi,

7762:3484). It affects annual shoots, which

turn white – red.

Morphology and taxonomy. A form

without the waxy surface in the berries (f.

epruinosum Asch. & Magn.) occurs

scattered in Inari Lapland. A thick-leaved

and stout ecotype (?) has been observed

twice in the Lake Inari area (e.g. Lake Inari,

Maurasaaret, Korkia Maura, 7639:3526,

1969 Y. Mäkinen 69-384 & L. Nurmi, TUR

172896). It resembles the hybrid V.

myrtillus × vitis-idaea, but deviates e.g. in

the serrature of the leaf margins.

Dependence on culture. The berries

of V. myrtillus are a widely known and

valued nature product, which are collected

by local people all over the distribution

area. The total yield in 1986 in Inari

Lapland was estimated as 11.64 million

kilograms (Kujala et al. 1987: 20), of which

only about 0.23 % was collected (Kujala et

al. 1987: 35). The amount of the yield

varies greatly from year to year, depending

e.g. on the weather conditions during the

flowering period. Thus, in 1987 the

commercial yield was zero, and in 1988 the

total amount of blueberries collected was

only 350 kg (Kujala et al. 1989). Puikko

(1992) has studied the usage of the natural

berries in Inari Lapland, and stresses e.g.

that the berries do not accumulate heavy

metals. The berries and the leaves contain

numerous flavoproteins, anthocyanins and

organic acids (Piippo 2005a: 204) and have

been used in innumerable ways also in

Lapland as herbal medicines. Hemera-

diaphore.

YM

Vaccinium oxycoccos L.

Oxycoccus palustris Pers.

O. quadripetalus Braun-Blanq.

Indigenous, rare

Map 63

Distribution. Nearly circumpolar (Benum 1958: 314,

Hultén 1971b: 140, Hultén & Fries 1986: map 1458).

Common in S and central Fennoscandia, rare in the

north (Hultén 1971a: map 1382, Roweck 1981: 372,

Nilsson 2000: 158, Mossberg & Stenberg 2003: 457),

missing in Iceland (Löve 1983: 300, Hultén & Fries

l.c.).

Very rare in Troms and Finnmark, e.g. Nesseby

and Sør-Varanger (Norman I(1): 710, Dahl 1934:

380, Benum 1958: 314, Alm et al. 1997: 41, Lid &

Lid 2005: 607). Fairly common in S Pechenga

(Mikkola 1941: 34). Numerous localities in the Kola

Peninsula, the northernmost ones close to the

Rybachy Peninsula (Fl. Murm. IV: map 111,

Mäkinen 2002: 17). Scattered in Keret Karelia

(Söyrinki 1956: 30); “often” in the Kovda area

(Sokolov & Filin 1996: 124). Scattered – fairly

common in the Lutto area (Roivainen 1923: 293).

In N Finland scattered – common up to the

coniferous forest boundary (Hjelt & Hult 1885: 135,

Wainio 1891: 46, Hult 1898: 163, Hjelt 1919: 297,

Ruuhijärvi 1960, Kujala 1961, 1964: map 143,

Montell 1962: 125, Söyrinki & Saari 1980: 121, SKK

III: 303, Hämet-Ahti et al. 1998: 211, Lampinen &

Lahti 2018). In Enontekiö Lapland rare (e.g.

Ruuhijärvi 1960, Hämet-Ahti et al. l.c., Lampinen &

Lahti l.c.).

InL ref. “ad lac. Jevjejärvi inarensem et

in reg. subalp. alpium Tuorpumoaivi et

Harimatschokka” (Kihlman 1884: 109).

Törmänen, Kyrö, Veskoniemi (Wainio

1891: 46), seven localities in SE Inari

(Kujala 1964: map 143). Ivalojoki at

Lismajoki mouth (Kujala 1962: 177), four

localities in the Vaskojoki area (Laine

1964: 115).

96 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

In the Lemmenjoki area scattered –

fairly rare (Klockars & Luther 1938), but V.

microcarpum is not mentioned at all. In the

upper Kevojoki two uncertain localities in

Heikkinen & Kalliola (1990: 48:

7710:3480, 7711:3481).

Kevo 0.6 %, InL 28 %, 84 sq. H 6,

OULU 1, TUR 31, YME 9 spec.

Rare (333; 0.052). Inari: III (321;

0.076), Utsjoki: I (12; 0.005).

Difference***. Clearly concentrated in and

around the basin of Lake Inari. The

northern limit of V. oxycoccos

approximately coincides with the northern

pine forest line (cf. Kallio 1961, Kallio et

al. 1971: 84). Southern.

FMF 0.396.

Vertical distribution. b: I (26; 0.012),

c: III (307; 0.088). Difference b-c***.

Range 120 m (S fork of Nanguvuono ca. 2

km E of Mielikköjärvi, 7627:3530) – 320 m

(E of Litmuorvaara, 7598:3468). Silvine.

Ecology. V. oxycoccos favors clearly

moister substrates than V. microcarpum;

the habitats are often between Sphagnum

fuscum hummocks. Likewise, as Drosera

rotundifolia is a companion of V.

microcarpum (cf. Mäkinen et al. 2005: 36),

D. longifolia is a companion of V.

oxycoccos. Often the habitats of the latter

are covered by a water layer 1-5 cm thick.

Other typical associates include Carex

canescens, C. limosa, Eriophorum

angustifolium, Juncus filiformis, Potentilla

palustris and Trichophorum alpinum.

Especially in the S parts of the area, V.

oxycoccos may also grow on mesotrophic

swamps (“Scorpidium-Rimpibraunmoore”,

Ruuhijärvi 1960: 118-119), characterized

by Carex chordorrhiza, Juncus stygius,

Loeskypnum badius, Scorpidium revolvens

and S. scorpioides.

V. oxycoccos generally flowers

abundantly, and also regularly produces

berries at least in habitats around the Lake

Inari basin. – Acidocline (Mäkinen &

Kallio 1979: 18). Acidocline.

Dependence on culture. V. oxycoccos

is collected as a valuable berry, at least in

the southern parts of the Lake Inari basin

(cf. Eklund 1926, Ervi 1956, Ruuhijärvi

1974). Two persons in Nellim (Mr. Lasse

Männistö, Mr. Esa Saijets) told that they

preferred to collect the berries (with hand

pickers) in the spring when the long stems

with well-preserved berries are floating in

water. It is also known that bears eat the

berries in the spring (cf. Ruuhijärvi 1974).

Hemeradiaphore.

YM

Vaccinium uliginosum L.

Indigenous, very frequent

Map 64

Distribution. V. uliginosum is a very complicated

polymorphic complex and includes numerous

subordinate taxa (Hultén 1971a: map 1378, 1378a;

1971b: 332). Subsp. uliginosum is arctic and boreal

circumpolar (Hultén 1948: 1261). In Fennoscandia

common and mainly a lowland plant (Roweck 1981:

375, Hultén & Fries 1986: map 1461).

Common – very common all over N

Fennoscandia (Fellman 1835: 259, Hjelt & Hult

1885: 135, Norman I(1): 709, Hult 1898: 163, Hjelt

1919: 292, Roivainen 1923: 293, Dahl 1934: 380,

Kalela 1939: 115-491, Kalliola 1932: 107, Lindén

1943: 77, Söyrinki 1956: 30, Benum 1958: 313,

Laine 1958: 89, Montell 1962: 125, Wistrand 1962:

131, Hämet-Ahti 1963a: 115, Kujala 1964: 82,

Söyrinki & Saari 1980: 122, Roweck 1981: 375,

Virtanen & Väre 1990, Piirainen & Piirainen 1991b,

Piirainen 1996b, Hämet-Ahti et al. 1998: 211,

Mossberg & Stenberg 2003: 459, Lampinen & Lahti

2018), as well as in the Kola Peninsula (Fl. Murm.

IV: map 107, Mäkinen 2002: 17).

InL ref. Common – very common in

Inari Lapland according to all published

floristic studies: Kihlman 1884: 108,

Wainio 1891: 46, Klockars & Luther 1938

(uppermost Lemmenjoki), Laine et al.

1955: 130 and Kallio & Mäkinen 1957: 26

(W and NW Utsjoki), Kujala 1962: 177

(Ivalojoki), Laine 1964: 115 (Vaskojoki),

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 97

Laine 1970(II): 131 (Kevojoki), Kuitunen

1984 (W Utsjoki), Heikkinen & Kalliola

1990: 35 (Kevo Strict Nature Reserve).

Kevo 98.1 %, InL 95 %, 260 sq. H 16,

KUO 1, OULU 2, TUR 24, YME 5 spec.

Very frequent (5269; 0.811). Inari: VII

(3381; 0.787), Utsjoki: VII (1888; 0.859).

Difference***. One of the commonest

species and fairly evenly distributed in the

map. There are over 1200 squares of 1×1

km² in the area in which V. uliginosum has

not been recorded, most probably mainly

due to incompletely studied squares (see

Introduction). Whole area, although very

significantly commoner in Utsjoki than in

Inari.

FMF 0.985.

Vertical distribution. a: VII (655;

0.865), b: VII (1958; 0.870), c: VII (2656;

0.761). Difference a-c***, b-c***.

Although V. uliginosum is common in all

altitudinal belts, it seems to favor the alpine

and birch belts. Range 15 m (S end of Lake

Pulmankijärvi, 7762:3539) – ca. 600 m

(Karegasnjarga-Ailigas, Lanka,

7705:3460). Tr 1330 m (Engelskjøn &

Skifte 1995: 150), Fnm 781 m (Norman

I(1): 709), EnL 1100 m (Laine 1958: 89).

Lule lappmark, Sweden 1410 m (Karlsson

1973: 159). Ascending generally to greater

heights than either V. myrtillus or V. vitis-

idaea. Vertical ubiquitous.

Ecology. In the pine forest areas, V.

uliginosum occurs almost without

exception in boggy or at least in fresh

associations; however, it avoids large open

fen-like areas. Often it is one of the most

important species in the inundation zone

along rivers and lakes. Especially along

rivers it forms shrub-like stands and

reaches regularly over 50 cm, even 70 cm

in height (cf. Kujala 1962: 177, Laine

1970(II): 131, Haapasaari 1988: 30-129).

According to the measurements by the

author, the diameter at the base of the

branching stem may reach 11 mm.

Common associates are e.g. Betula

nana, Ledum palustre, Andromeda polifolia

and Eriophorum angustifolium (Laine

1970(II): 131). The growth sites in the pine

area and birch belt are decisively moister

than those of V. myrtillus or V. vitis-idaea.

V. uliginosum is dominating in the Ledum-

Uliginosum association, and common also

in the Hylocomium-Myrtillus and

Empetrum-Myrtillus associations in the

Oulanka National Park (Söyrinki & Saari

1980: 122).

In the alpine belt, however, the

moisture requirements turn around. The

species usually prefers dry habitats and

avoids e.g. the inundation zones. This has

been noted by several botanists, e.g.

Kontuniemi 1932: 41 (“often on rather dry

sites”), Mikkola 1941: 34 (“in alpine belt

regularly on dryish gravelly grounds”),

Dahl 1934: 380 (“dels på fuktige, dels på

torre steder”), Karlsson 1973: 159 (“prefers

dry habitats and S-facing sites”). There may

be two different ecotypes in question: one

preferring riversides in the forested belts,

the other preferring dry gravelly ground in

the alpine belt.

Around the Kevo Subarctic Research

Station V. uliginosum starts flowering

around the middle of June, usually ca. one

week later than V. myrtillus, but the first

berries are ripe almost at the same time in

the end of July or in the beginning of

August (Alanen 2007). Söyrinki (1939a:

334) describes the flowering and ripening

of the seeds in the alpine zone of Pechenga;

there are only weak possibilities for

generative increase. – V. uliginosum is

weakly calciphilous according to Pesola

(1928: 160), and may also grow on

ultrabasic ground (Roweck 1981: 376).

Indifferent or somewhat calciphilous

(Nilsson 2000: 158). In Inari Lapland it is

considered amphicline (Laine 1970(II):

131, Mäkinen & Kallio 1979: 18).

Amphicline.

98 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Parasites. V. uliginosum is generally

infected by members of three parasitic

genera. Pucciniastrum vaccinii (Wint.)

Jørst. has been collected in numerous

localities (Mäkinen 1964b: 173, 1964c);

there are 13 specimens in TUR from Inari

Lapland. The powdery mildew

Podosphaera major (Juel) Blumer is

probably common over the area, but it is

inconspicuous and there are only seven

collections from Inari Lapland in TUR

(Mäkinen 1969).

Of the genus Exobasidium (see

Nannfeldt 1981) there are three species

which attack V. uliginosum and which

occur in Inari Lapland. E. vaccinii-uliginosi

Boud., which is “strictly bound to high

latitudes and high altitudes” (Nannfeldt

1981: 64), is the commonest; there are nine

specimens in TUR, of which seven have

been collected in the Lake Inari area and

two in NE Inari near Lake Iisakkijärvi. E.

pachysporum Nannf. (“all over the Nordic

countries”) has been collected at Lake

Kevojärvi, Kutuniemi (7742:3500, 1995 M.

Alanen, TUR 116152). E. expansum Nannf.

has been found at the Tshieskuljohka near

Kevojärvi (7740:3502, 1995 R. Kapanen &

R. Kosonen, TUR 114410).

Morphology and taxonomy. A

narrow-leaved form (f. angustifolium

Hiitonen) has been collected three times:

(1) Inari, Ivalojoki, Kuttura, Kalaoja

(7592:3480, 1985 Y. Mäkinen, YME

21513); (2) Inari, W of Pasasjärvi,

Rautujärvi, forming a stand of ca. 1 sq.

meter (7618:3503, 1996 Y. Mäkinen, YME

23155); (3) Utsjoki, Puksalskaidi 4 km S of

the Kevo Subarctic Research Station

(7738:3500, 1971 U. Laine, TUR 214063).

E.g. Hiitonen (1933: 567) reported this

form from SW Finland, and Montell (1962:

125) has found it in Muonio, N Finland.

V. uliginosum subsp. uliginosum has

2n = 48 chromosomes (now regarded as an

octoploid); this distinguishes it from the

tetraploid V. uliginosum subsp.

microphyllum (Lange) Tolm. (=V.

gaultherioides Bigelow) with 2n = 24

chromosomes (Löve & Löve 1966: 46,

Engelskjøn & Knaben 1971: 21). The latter

has been found in the northernmost coastal

Norway, Kirovsk – Kandalaksha area and

in the Rybachy Peninsula and Kola

Peninsula (Benum 1958: 313, Fl. Murm.

IV: map 107, Elven 2016). Specimens

morphologically resembling subsp.

microphyllum have also been collected

from NW Finland, Enontekiö Lapland, SW

slope of the Saana fjeld (767:325, 1958 O.

Hiidensalo, TUR 578307) and S slope of

the Háldi fjeld (ca. 7700:3274, 1977 S.

Vuokko, H 469210), and from Inari

Lapland, E part of Utsjoki, S slope of

Luovvosvarri fjeld, brook side (7754:3520,

1958 M. Kovanen, TUR 78769). In

addition, Kuitunen (1984) reported subsp.

microphyllum from Karegasnjarga-Ailigas,

from barren ground near the top, but there

is no specimen. However, plants

morphologically resembling microphyllum-

type in the Fennoscandian mountains have

been counted to have 2n = 48 chromosomes

(Borgen & Elven 1983), so the existence of

subsp. microphyllum in Finland remains

still to be demonstrated (cf. Elven 2016).

Dependence on culture. According to

Piippo (2005a: 133), the berries are used

especially to cure the troubles in the

digestive system; the berries have also been

eaten in Lapland. According to the

preliminary enquiries made in Inari in

1990, among ten households, eight

considered the berries poisonous and in two

houses they were collected in small

amounts and eaten. Hemeradiaphore.

YM

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 99

Vaccinium vitis-idaea L.

Rhodococcum vitis-idaea (L.) Avr.

Indigenous, very frequent

Map 65

Distribution. The collective species is circumpolar

(Hultén & Fries 1986: 1104). Subsp. vitis-idaea (the

lowland “race”) is Eurasiatic, and subsp. minus

(Lodd.) Hultén (the arctic-montane “race”) mainly N

American and arctic Siberian.

Subsp. vitis-idaea is common all over the

Fennoscandian lowland and also on the mountains

(Hultén 1971a: map 1379, 1971b: 78, 332, Roweck

1981: 374). Common – very common in all floristic

studies concerning N Fennoscandia, e.g.: Fellman

1835: 260, Hjelt & Hult 1885: 135, Norman I(1):

691, Hult 1898: l63, Dahl 1934: 380, Björkman

1939: 158, Kalliola 1939: 107, Söyrinki 1939a: 331,

Arwidsson 1943: 230, Benum 1958: 313, Montell

1962: 125, Wistrand 1962: 131, Hämet-Ahti 1963a:

e.g. 112, Laine 1958: 89, Kujala 1964: 83, Ryvarden

1969: 34, Söyrinki & Saari 1980: 122, Virtanen &

Väre 1990, Piirainen & Piirainen 1991b, Piirainen

1996b, Hämet-Ahti et al. 1998: 211, Mossberg &

Stenberg 2003: 458, Lid & Lid 2005: 605, Lampinen

& Lahti 2018. The same applies to the Kola

Peninsula (Fl. Murm. IV: map 108, Alm et al. 1997:

41, 1998: 135, Mäkinen 2002: 17), and the Kovda

area (Sokolov & Filin 1996: 124).

InL ref. Common to very common in

Inari Lapland: Kihlman 1884: 108, Wainio

1891: 46, Klockars & Luther 1938 (upper

Lemmenjoki), Laine et al. 1955: 130 and

Kallio & Mäkinen 1957: 26 (W and NW

Utsjoki). Kujala 1962: 177 (Ivalojoki),

Laine 1964: 115 (Vaskojoki), Vanhatalo

1965: 124 (Utsjoki), Laine 1970(II): 131

(Kevojoki), Kuitunen 1984 (W Utsjoki),

Heikkinen & Kalliola 1990: 35 (Kevo Strict

Nature Reserve).

Kevo 99.2 %, InL 95 %, 260 sq. H 22,

KUO 2, OULU 3, TUR 19, YME 11 spec.

Very frequent (5257; 0.809). Inari: VII

(3366; 0.783), Utsjoki: VII (1891; 0.860).

Difference***. Seemingly commoner in

Utsjoki than in Inari; this is probably

mainly due to the larger amount of

incompletely studied squares in the latter.

Whole area.

FMF 0.985.

Vertical distribution. a: VII (661;

0.873), b: VII (1952; 0.865), c: VII (2644;

0.759). Difference a-c***, b-c***. Range

15 m (S end of Pulmankijärvi, 7762:3539)

– ca. 600 m (Karegasnjarga-Ailigas, Lanka,

7705:3460). V. vitis-idaea is clearly

commoner in the alpine and birch belts than

in the coniferous zone. Tr 1380 m

(Engelskjøn & Skifte 1995: 150), Fnm 910

m (Rastigaissa, Ryvarden 1969: 34), EnL

1270 m (Väre & Partanen 2009: 98). In

Finnmark higher up than V. uliginosum or

V. myrtillus (Dahl 1934: 380). Sarek

National Park in N Sweden 1510 m

(Karlsson 1973: 58). V. vitis-idaea often

rises up to the summits of the fjelds

(Söyrinki 1956: 30). Vertical ubiquitous.

Ecology. V. vitis-idaea is the main

dwarf-shrub on all gravel ridges (Mikkola

1941: 34), and also its amplitude in Finland

is greater than with any other forest or bog

plant (Kujala 1964: 83). It is one of the

most dominant species in the Vaskojoki

area, often growing together with

Empetrum nigrum subsp. hermaphroditum,

Deschampsia flexuosa and Festuca ovina

(Laine 1964: 115). It occurs almost

everywhere, and is missing only in moist

riverside groves. It is not, however, a strong

competitor, and this may be one of the

reasons why it is actually commoner in the

subalpine belt than in the pine forest area.

Only near the mouth of the Ivalojoki it

grows ”on a meadow” (Kujala 1962: 177).

The cowberry has high frequencies in

several associations in the subalpine birch

forests, e.g. in the subalpine Empetrum-

Lichenes type, subalpine Empetrum-

Lichenes-Pleurozium type and subalpine

Empetrum-Myrtillus type (Hämet-Ahti

1963a: 40, 44, 50, 54, 107, 115). It is so

common that no association is called ”the

cowberry type”.

In the surroundings of the Kevo

Subarctic Research Station, V. vitis-idaea

starts flowering on an average in the second

100 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

half of June, ca. two weeks later than V.

myrtillus (Alanen 2007). Several authors

state that the flowering is often very scanty

and the generative reproduction is weak

(e.g. Kontuniemi 1932: 40 concerning the

Pechenga subalpine area; cf. also Söyrinki

& Saari 1980: 122). Söyrinki (1939a: 331)

has studied the generative reproduction in

the alpine belt in Pechenga and stresses that

seedlings are found very seldom, and the

germination ability of the seeds is very

weak. One of the reasons might be that the

ripening of the berries takes a long time,

almost two weeks longer than with V.

myrtillus (Alanen l.c.), and the berries are

often unripe when they are already covered

by the first snow layer and the temperature

has permanently dropped below zero. The

berries often remain edible up to the next

summer (cf. Kihlman 1884: 108, Hjelt

1919: 294).

According to Pesola (1928: 160), V.

vitis-idaea is clearly but weakly calciphobe.

Classified as acidophilous by Dahl (1934:

380) and indifferent by Nilsson (2000:

158). According to Karlsson (1973: 158) it

is amphicline and prefers dry substrate and

south-facing slopes. In Inari Lapland it is

considered amphicline (Laine 1970(II):

131) or acidocline (Mäkinen & Kallio

1979: 18). Acidocline.

Parasites. The species is commonly

attacked by several species of parasitic

fungi. The rust Pucciniastrum vaccinii

(Wint.) Jørst. (see Rainio 1926: 254,

Mäkinen 1964b: 173, 1964c) is common

over Inari Lapland; there are 32 specimens

in TUR (Inari 13, Utsjoki 19). The powdery

mildew Podosphaera major (Juel) Blumer

(see Mäkinen 1969) is very rare on this host

(the main host is V. uliginosum); it is

reported by Blumer (1967: 159) in

Germany. Apparently V. vitis-idaea has no

”own” species of Podosphaera; it is rarely

infected also by P. myrtilli (Ahti 1968).

Exobasidium vaccinii (Fuck.) Woron.

is fairly common: 38 specimens over the

area in TUR (see also Nannfeldt 1981: 63).

Mycosphaerella rubefaciens B. Erikss. is

probably common; its description (Eriksson

1974: 217) was based on a specimen

collected at the Kevo Subarctic Research

Station (7741:3500, 1965 P. Tapaninen,

TUR 53926, UME, UPS). It causes the

upper parts of the stems to turn red and

slightly swollen.

Many parasitic fungi decrease the yield

produced by the cowberry, such as

Exobasidium vaccinii, Helminthosporium

vaccinii, Gibbera vaccinii (Paal & Paal

1989: 77). Also parasitic insects, especially

Rhopobota naevana (Lepidoptera), may

decrease the yield through weakening of

the plant (Paal & Paal 1989: 81-82). The

book of the Paals’ contains a lot of useful

information on the ecology of the

cowberry.

Morphology and taxonomy. The

mainly N American subsp. minus has a few

sites in Europe and Asia (Hultén 1971a:

map 1379). Subsp. minus has one

confirmed locality in the Kola Peninsula

(Ponoi Lapland, Orlow; 1889 A. O.

Kihlman, TUR 78857). Fl. Murm. IV: map

109 gives over 20 localities, e.g. in the

Rybachy Peninsula (Pechenga). Subsp.

minus occurs also in Finnmark (cf. Lid &

Lid 2005: 605) and might occur on high

fjelds in N Utsjoki; according to Elven

(2016) subsp. minus is scattered in N

Fennoscandia. In any case, the distribution

boundaries between the two subspecies are

not everywhere clear.

A deviating form, f. sterile H. V.

Rosend. (f. longicaule auct.), has been

found in a few localities in the southern

part of Inari Lapland. Specimens in TUR:

(1) Inari, Alempi Paksupetäjäjärvi,

“dominant race in some places on fairly dry

pine heaths” (7638:3509, 1976 Y. Mäkinen,

TUR 287369); (2) Inari, NE of Virtaniemi,

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 101

NW of Harrijärvi, close to the Soviet

border, “growing in an area of ca. 200 m²,

on dry pine-growing slope” (7659:3570,

1982 Y. Mäkinen, TUR 269934); (3) Inari,

6 km S of Kaamanen village, a few stands

on pine heath (7663:3505, Y. Mäkinen,

TUR 125876). The same form was

apparently described by Hämet-Ahti

(1963a: 112): “A striking feature is that V.

vitis-idaea is tall and slender ... commonly

attaining 30 cm (sometimes 50 cm) in

height”. According to observations made, f.

sterile has few flowers (if any!) and may

produce few berries; it often remains totally

sterile. It often forms stands of 1 m² in area,

but may be dominant even on an area of

several hectares, e.g. in Inari, Romopää,

Tolonen and Martinkotavaara (7604:3522,

7607:3513 and 7628:3516, 2012 J. Nurmi,

field lists).

Sometimes most of the leaves are

exceptionally large. Such plants have been

collected in Inari, E of Menesjärvi, NW

slope of Vaatimenseisomapää, luxurious

slope (7633:3482, 1976 Y. Mäkinen 76-16,

TUR 287412). The broadness of the leaves

may be due to a fungal infection

(Exobasidium? Mycosphaerella?), or to an

occasional mutation in the leaf growth

point.

Dependence on culture. Kujala et al.

(1987, 1989) report an estimated yearly

total yield of cowberry in Inari Lapland as

11.8 million kilograms, of which only a

small amount was collected: the

commercial yields in 1987 were 82396 kg,

in 1988 50470 kg, in 1989 15930 kg.

Local Sámi people also used cowberry

jam as a preservative in other jams: a layer

of 1 cm is enough to prevent the growth of

molds and other infections. The chemical

composition of the berries of subsp. vitis-

idaea and subsp. minus is different (Hultén

1971b: 78); the berries include e.g. several

organic acids (glucosides of benzoic acid),

which act as mold-preventing compounds

(Piippo 2005b: 43). The berries do not

accumulate heavy metals, either (Puikko

1992). Paal & Paal (1989: 52) present

various methods for the estimation of the

cowberry yield in the coenopopulations of

the plant.

Cultivation experiments with cowberry

have been conducted also at the Kevo

Subarctic Research Station (Kärenlampi

1973, Lehmushovi 1976, 1977, Nousiainen

et al. 1978).

Hemeradiaphore in Inari Lapland

according to Mäkinen & Kallio (1979: 18),

but here rather regarded as hemerophobe

(cf. Kujala 1964: 83).

YM

EMPETRACEAE

Empetrum nigrum L.

Indigenous, very frequent

A complicated species complex with a boreal to

arctic circumpolar distribution. There are different

views of the number of taxa included. Usually E.

nigrum L. is understood as a separate taxon from E.

rubrum Vahl ex Willd. in the Southern Hemisphere

and mostly also from E. eamesii Fernald & Wiegand

and E. atropurpureum Fernald & Wiegand in North

America (Elven 2016). E. nigrum is usually divided

into two subspecies (or species), the diploid (2n =

26), dioecious subsp. nigrum (E. nigrum L. s. str.)

with a more southerly distribution, and the tetraploid

(2n = 52), monoecious subsp. hermaphroditum

(Lange ex Hagerup) Böcher (E. hermaphroditum

Lange ex Hagerup) prevailing in the northern

latitudes. However, the connection between the

ploidy level and the occurrence of different sex types

is not as straightforward as sometimes presumed, and

also the distinction between monoecious and

dioecious plants is not always clear (Hultén 1971b:

86; see below). Tetraploids seem to have arisen

several times from different combinations of

diploids, and thus the tetraploid level is probably

highly polyphyletic (Elven 2016). In recent

treatments, the complex is therefore sometimes dealt

collectively without named subordinate taxa (e.g.

Elven l.c.). Here, however, the taxonomic view and

nomenclature of Hämet-Ahti et al. (1998) is

followed.

JN

102 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Empetrum nigrum L. subsp.

hermaphroditum (Hagerup) Böcher

E. hermaphroditum Hagerup

Indigenous, very frequent

Map 66

Distribution. Boreal circumpolar, mountains of

central Europe (Hultén 1971b: 86, Hultén & Fries

1986: map 1464, Roweck 1981: 379). A tetraploid

taxon (2n = 52), sometimes separated at species level

(E. hermaphroditum) from the diploid plants (E.

nigrum s. str., 2n = 26). Subsp. hermaphroditum is a

typical example of the phenomenon that polyploidy

increases towards arctic and boreal conditions. – The

two taxa have not been separated in older literature,

therefore the list of references below begins only in

ca. 1940.

In Fennoscandia common in the north, but rare

or absent S of ca. lat. 60° N (“Limes Norrlandicus” in

Sweden) and in the SW lowland of Norway (Hultén

1971a: map 1385a, Mossberg & Stenberg 2003: 459,

Lid & Lid 2005: 609). Common in N Fennoscandia

according to all published studies concerning the area

(e.g. Björkman 1939: 61, Hustich 1940c: 58,

Ruuhijärvi 1960: 354-359, Montell 1962: 125,

Wistrand 1962: 132, Söyrinki & Saari 1980: 123,

Mossberg & Stenberg 2003: 459; a comprehensive

list of references in Hultén 1971b: 339).

Very common in Troms and Finnmark

(Ryvarden 1969: 34, Engelskjøn & Skifte 1995: 151,

Lid & Lid 2005: 609); a common dwarf shrub also on

palsa bogs (Vorren 1967: 25). Common in Pechenga

and the Kola Peninsula (Fl. Murm. IV: map 66,

Mäkinen 2002: 17) as well as in the Kovda area

(Sokolov & Filin 1996: 112). Common to very

common in Sompio, Kittilä and Enontekiö Lapland

(Laine 1958: 90, Pertola 1961: 61, Virtanen & Väre

1990, Piirainen & Piirainen 1991b, Hämet-Ahti et al.

1998: 215, Lampinen & Lahti 2018).

InL ref. Very common in the

Lemmenjoki and Vaskojoki areas

(Klockars & Luther 1938, Laine 1964:

115), as well as in the Peäldoajvi fjeld area

(Koivistoinen 1964: 58). Common and

abundant along the Ivalojoki (Kujala 1962:

178). In NW Utsjoki very common (Laine

et al. 1955: 130, Kallio & Mäkinen 1957:

26) as well as in the Kevojoki area (Laine

1970(II): 132, Heikkinen & Kalliola 1990:

35, 48).

Kevo 98.1 %, InL 96 %, 261 sq. (as E.

nigrum). H 16, JYV 2, OULU 2, TUR 53,

YME 17 spec.

Very frequent (5233; 0.806). Inari: VII

(3342; 0.779), Utsjoki: VII (1891; 0.861).

Difference***. There are a few squares in

which the species is not listed (cf. map 67),

but this is mainly due to incomplete

mapping (see Introduction); the relative

frequency at least in Utsjoki is close to

1.000. Probably the only areas where the

plant is truly missing are the large open

waters of Lake Inari! The numbers above

include 49 squares in which Empetrum has

been determined only to the species level.

Whole area.

FMF 0.985.

Vertical distribution. a: VII (664;

0.876), b: VII (1953; 0.868), c: VII (2616;

0.752). Difference a-c***, b-c***. Range

15 m (Pulmankijärvi, 7762:3539) – ca. 600

m (Karegasnjarga-Ailigas, Lanka,

7705:3460). Tr 1190 m (Engelskjøn &

Skifte 1995: 151), Fnm 948 m (Norman

I(2): 935, as E. nigrum), 920 m (Ryvarden

1969: 34, as E. hermaphroditum), EnL

1150 m (Laine 1958: 90). Vertical

ubiquitous.

Ecology. Empetrum nigrum subsp.

hermaphroditum is very common all over

Inari Lapland, and also in all vertical belts.

Hämet-Ahti (1963: 37-78) has described a

subalpine Empetrum type (sET) and a

submaritime Empetrum type (ET); in

addition, subsp. hermaphroditum is an

important factor in four further mountain

forest types. Kalliola (1939: 185) described

(in alpine belt) an Empetrum-Cetraria

nivalis association, which, according to

him, is the most widely distributed plant

association in the Finnish fjeld areas; this

holds true also for both Inari and Utsjoki

(in fact, Kalliola was using the name E.

nigrum, but most probably meaning subsp.

hermaphroditum).

According to Ruuhijärvi (1960: 265),

in Finnish Lapland subsp. hermaphroditum

is continental, subsp. nigrum suboceanic;

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 103

this is in contrast with Ahti et al. (1968:

201), who state that "a continuous

Empetrum hermaphroditum heath is

prevalent on the very coast". In the

Torneträsk area, N Sweden, subsp.

hermaphroditum forms, together with

Rubus chamaemorus, an important

vegetation type on the higher parts of the

mires (Sonesson 1970: 38).

In the Kevojoki valley (Laine 1970(II):

132) subsp. hermaphroditum is very

common, and also vertically ubiquitous. It

grows on various dwarf-shrub heaths, often

with Deschampsia flexuosa, Festuca ovina

and Vaccinium vitis-idaea, with the mosses

Dicranum fuscescens, D. scoparium,

Polytrichum piliferum, and with the lichens

Cetraria nivalis, Cladonia deformis and C.

elongata.

Almost regularly subsp. herma-

phroditum favors the mesic patches under

birches, cf. Hämet-Ahti 1963a: 46. The

subspecies also occurs on the open or half-

open gravelly patches on fjeld tops, with,

e.g., Carex glacialis and Luzula arcuata,

but avoids the sandy patches.

E. nigrum subsp. hermaphroditum

generally flowers and produces berries in

abundance (cf. Laine 1970(II): 132).

Söyrinki (1939a: 308-313) describes the

autecology (under E. nigrum!) in Pechenga

in detail, including the generative and

vegetative reproduction. About the ecology,

see further Söyrinki & Saari 1980: 122 and

Roweck 1981: 379. – Amphicline or Ca-

indifferent (Laine 1970(II): 132, Nilsson

2000: 159), acidocline (Mäkinen & Kallio

1979: 18). Acidocline.

Parasites. In Inari Lapland subsp.

hermaphroditum is attacked by the rust

fungus Chrysomyxa empetri Schroter (only

II is known; Rainio 1926: 253, Mäkinen

1964b: 157; TUR). C. empetri is also

collected in all surrounding provinces

(Sompio, Kittilä and Enontekiö Lapland in

Finland, Finnmark in Norway and

Pechenga in Russia; TUR).

Morphology and taxonomy.

Arwidsson (1935) was the first to realize

the significance of the two Empetrum taxa.

Marklund (1940) was one of the first

botanists to describe in detail the

differences between these two taxa.

According to Arwidsson (1943: 231)

and Knaben (1966), the pollen tetrads of

subsp. hermaphroditum are markedly

greater than those of subsp. nigrum (22-34

µm in subsp. nigrum, 34-47 µm in subsp.

hermaphroditum). It is generally believed

that subsp. hermaphroditum is always

bisexual. According to Knaben (1966) this

is not necessarily always true: there are

strains in which all the flowers are

unisexual, or in which some flowers are

bisexual, others unisexual. In S Norway

(close to Oslo), the proportion of such

deviating flowers was 4-8 %. Unisexual

flowers in subsp. hermaphroditum have

also been found in Inari Lapland. Whether

they are able to "hybridize" with bisexual

hermaphroditum, is not known.

Dependence on culture. The berries

of subsp. hermaphroditum (crowberry)

have been a highly valued nature product in

Lapland (see Puikko 1992). In 1990’s,

crowberry-cowberry jam was priced as 12-

14 €/kg, and the crowberry juice as 5-8

€/liter. The berries have also been used as a

valued medicine among northern nations

(SKK III: 314, Piippo 2005b: 61), e.g. they

include more ascorbic acid and trace

elements than the berries of Vaccinium

myrtillus or V. vitis-idaea. There are,

however, differences in the chemical

contents between the various races

(Arwidsson 1943: 231). Hemeradiaphore.

YM

104 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Empetrum nigrum L. subsp. nigrum

Indigenous, scattered

Map 67

Distribution. Boreal circumpolar, in central Europe

extending south almost to the Alps (Hultén 1971b:

86, 339, Hultén & Fries 1986: map 1463). A diploid

unisexual taxon (2n = 26). Earlier the tetraploid E.

nigrum subsp. hermaphroditum was not separated

from subsp. nigrum, see above.

In Fennoscandia distributed on the Atlantic

coast; in S Sweden up to ca. 60° N; and over the

whole of Finland, rare or missing in central and N

Sweden and in the Scandes (Arwidsson 1935, 1943:

128, Hultén 1971a: map 1385, Roweck 1981: 378,

Hämet-Ahti et al. 1998: 215, Mossberg & Stenberg

2003: 459, Lid & Lid 2005: 609).

A few localities in Troms and Finnmark

(Hämet-Ahti 1963a: 40, Mäkinen 1964a, Engelskjøn

& Skifte 1995: 151). The northernmost locality in

Norway was reported at Tanabru, Polmak (70° 14' N,

Hämet-Ahti 1963a: 35, Mäkinen 1964a). Later the

author found subsp. nigrum in Berlevåg (sandy spots

in a rich fjeld meadow by the main road, 70° 52' N,

2005 Y. Mäkinen 05-211, TUR 583568), which is

probably the northernmost record reported so far. –

Two localities on the southern coast of the Kola

Peninsula (Fl. Murm. IV: map 66), six localities in

Mäkinen (2002: 17). Very rare in the Kovda area

(Sokolov & Filin 1996: 112). Scattered – fairly rare

in Sompio Lapland, Kittilä Lapland and Enontekiö

Lapland (Montell 1962: 125, Piirainen 1996b,

Hämet-Ahti et al. 1998: 215, Lampinen & Lahti

2018). Several tens of specimens in H, TUR, YME

from all surrounding provinces, including Finnmark

and Pechenga.

InL ref. Laine (1964: 115) has found

the subspecies in four localities along the

Vaskojoki, and (1970(II): 132) one locality

in the Kevojoki area, in the mouth of the

Kamajohka, where it grew in abundance

with Diphasiastrum complanatum, Juncus

trifidus and Vaccinium vitis-idaea.

InL 44 %. H 19, TUR 63, YME 91

spec.

Scattered (934; 0.141). Inari: IV (852;

0.196), Utsjoki: II (82; 0.034).

Difference***. E. nigrum subsp. nigrum is

a southern plant both on the basis of its

horizontal as well as vertical distribution: it

is clearly more common in Inari especially

along the larger rivers like the Inarijoki,

Ivalojoki, Kaamasjoki and Vaskojoki. In

Utsjoki it occurs almost exclusively in the

valleys of large rivers, mainly along the

Inarijoki–Teno watercourse and the

Utsjoki. Southern.

FMF 0.632.

Vertical distribution. a: I (4; 0.006),

b: II (139; 0.061), c: IV (791; 0.220).

Differences***. Mainly a plant of the

coniferous zone, very rare in the alpine belt

and rare in the birch belt. Range ca. 20 m

(E side of Lake Pulmankijärvi, 7768:3538)

– ca. 600 m (Karegasnjarga-Ailigas, Lanka,

7705:3460). EnL 410 m (Väre & Partanen

2009: 102). Silvine.

Ecology. E. nigrum subsp. nigrum very

seldom grows in closed communities, but

prefers open or half-open, warm and sandy

slopes. In several cases it occurs on

recently burned areas. Often it favors

sandy-clayey, somewhat disturbed riverside

grounds, remaining, however, outside the

inundation zone. In a few cases it grows on

the wind-swept and open tops of dry

hummocks of palsa bogs (e.g. on Nuvvos-

Ailigas, W Utsjoki), while subsp.

hermaphroditum dominates their lower and

moister margins. It appears that subsp.

nigrum is a weak competitor in Inari

Lapland, and it is not a constituent member

of any plant association.

In the alpine localities of the Nuvvos-

Ailigas fjeld complex, E. nigrum subsp.

nigrum grows fairly sparsely on dry

gravelly alpine heath, with Carex glacialis,

Empetrum nigrum subsp. hermaphroditum

and Luzula arcuata, e.g. in the summit area

close to the antenna (7748:3472, 1990 Y.

Mäkinen, YME).

E. nigrum subsp. nigrum favors sandy

heaths, and often, in the southern parts of

Inari Lapland, it grows on sandy riversides

together with pine. However, it is by no

means dependent on pine forests but thrives

equally well on sandy birch heaths, and

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 105

may also be the dominant dwarf-shrub on

half-open sandy heaths. Acidocline

(Mäkinen & Kallio 1979: 18). Amphicline.

Parasites. In TUR there are three

collections of the rust Chrysomyxa empetri

Schröter (II) on subsp. nigrum: Utsjoki,

Outakoski (G. Marklund); Inari, Ivalo (G.

Marklund); Inari, Angeli (U. Laine).

Morphology and taxonomy.

Marklund (1940) gives a good comparison

concerning the vegetative differences

between subsp. nigrum and subsp.

hermaphroditum. The branches of subsp.

nigrum are normally clearly longer than

those of subsp. hermaphroditum, and of

brownish-reddish or yellowish color. The

distance of the leaves in young branches is

greater than in subsp. hermaphroditum; if

the habitat is not profitable, the leaves may

be crowded as in subsp. hermaphroditum.

The white "midrib" should be more

prominent in subsp. hermaphroditum, but

in field conditions this is very difficult to

observe because the opening and closing of

leaves depends on the atmospheric moisture

and on the air temperature.

Dependence on culture. The berries

of Empetrum have been used for a long

time. They contain trace elements more

than the berries of Vaccinium myrtillus or

V. vitis-idaea (Piippo 2005b: 61); see under

subsp. hermaphroditum. Mainly the

tetraploid subsp. hermaphroditum is used:

there is a clear difference both in the flavor

and the content of trace elements in the

favor of subsp. hermaphroditum.

The subspecies has effectively spread

on sandy highway banks and takes often

advantage of temporary tractor tracks. At

Darjokka (Finnmark, Norway) it proceeds

almost to the alpine belt along the margins

of a temporary tractor road. Hemero-

philous.

YM

PRIMULACEAE

Androsace septentrionalis L.

Not accepted for Inari Lapland

Not mapped

Distribution. A circumpolar, continental species

complex. Widely distributed in Eurasia and North

America, extending to the Arctic in North America

and N Asia (Hultén & Fries 1986: 1106, map 1472).

Scattered in the S and central parts of Fennoscandia,

one locality in the Russian side of Koillismaa

(Paanajärvi district, Ruskeakallio, Kravchenko 2007:

133, Kastikka 2018) and in the Kola Peninsula N of

the Arctic Circle (Hiitonen 1933: 558, Hultén 1971a:

map 1396, H). In Finland a rare, mostly casual

anthropochore, seldom established or considered to

be native (Kemppainen et al. 1991, Ryttäri &

Kettunen 1997: 57, Hämet-Ahti et al. 1998: 217,

Ryttäri et al. 2012: 39). The northernmost Finnish

locality in Oulu, where it probably arrived with

ballast and is now extinct (Väre et al. 2005: 446).

InL ref. There is a dot in S Inari (approximately

in Ivalo) in Hultén (1971a: map 1396) and Hultén &

Fries (1986: map 1472), although in Hultén (1971b:

map 118) it is missing. The record was included in

Hämet-Ahti et al. (1986: 193). However, its origin is

unclear (cf. Kemppainen et al. 1991: 114), and hence

the dot has been removed in Hämet-Ahti et al. (1998:

217). The species is here not regarded as belonging to

the flora of Inari Lapland.

JN

Lysimachia thyrsiflora L.

Naumburgia thyrsiflora (L.) Rchb.

Indigenous, very rare

Map 68

Distribution. Boreal circumpolar, rare in NE Asia

and NW North America, in Europe missing or rare in

the S and W parts (Hultén 1971b: 112, Fl. Eur. 3: 27,

Hultén & Fries 1986: map 1479). In Fennoscandia

common in the southernmost parts of Norway and

most of Sweden and Finland up to the Arctic Circle,

scattered – rare northwards and almost lacking in the

mountain areas (Hultén 1971a: map 1401, Roweck

1981: 383, Hämet-Ahti et al. 1998: 219, Mossberg &

Stenberg 2003: 464, Lid & Lid 2005: 614).

Rare in Troms and Finnmark (Norman I(2):

871, Dahl 1934: 383, Benum 1958: 317, Edvardsen

& Elvebakk 1981, Lid & Lid 2005: 614). Rare in

Pechenga (Hiitonen 1933: 559, Alm et al. 1997: 42,

106 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Fl. Murm. V: map 2), in the Kola Peninsula scattered

occurrences mostly in the S parts (Fl. Murm. l.c.). In

Sompio and Kittilä Lapland scattered to fairly

common in the southern parts, fairly rare to rare

northwards (Fellman 1835: 253, Hjelt & Hult 1885:

139, Wainio 1891: 44, Hult 1898: 163, Hustich

1936a, Salonen 1956, Kujala 1961, Montell 1962:

125, Rintanen 1976: 142, 1982a: 251, Laitinen &

Ohenoja 1990: 23, Lampinen & Lahti 2018). In

Enontekiö Lapland rare and only in the S parts

(Hustich 1936a, Montell 1948, Rintanen 1982a: 251,

Hämet-Ahti et al. 1998: 219, Lampinen & Lahti l.c.).

InL ref. Vastusjärvi (Kihlman 1884:

115), “in littore lacus ad Kyrö” (Wainio

1891: 44), Sotajoki 2 km from the mouth

(Mikkola 1941: 35). Two localities in the

upper course of the Ivalojoki, scattered in

the lowermost course (Kujala 1962: 178).

Three localities in the Vaskojoki basin

(Laine 1964: 116, Rautava 1964: 90). In

only one lake (Valkkojärvi) of the 38 lakes

studied by Rintanen (1982a: 251) in InL.

InL 8 %, 29 sq. H 3, KUO 1, OULU 2,

TUR 10, YME 14 spec.

Very rare (60; 0.010). Inari: I (60;

0.014). The vast majority of the localities is

concentrated around Lake Inari, especially

in the S parts, and along the Ivalojoki,

mostly along its lower course. The rest of

the finds are mainly along the Vaskojoki

and Kaamasjoki, and their tributaries. No

localities known from Utsjoki, although

sometimes misunderstood so on the

grounds of the list of southern species in

Kallio (1961: 98). Southern.

FMF 0.141.

Vertical distribution. b: I (3; 0.001),

c: II (57; 0.017). Difference***. Two

localities in the birch belt, both in the basin

of the upper Vaskojoki. Most localities

below 150 m. Range 120 m (several

localities around Lake Inari) – 295 m

(Stuorrabogejávri, 7635:3430). Tr “not

above 30 m” (Engelskjøn & Skifte 1995:

152), Fnm ca. 370 m (Artsdatabanken

2015), EnL ca. 390 m (Hustich 1936a).

Silvine.

Ecology. Lysimachia thyrsiflora grows

in the grassy margins of lakes and rivers, in

damp, periodically flooded meadows along

rivers and brooks, and marshy meadows by

shallow lakes. It often grows in shallow

water, and is partly inundated at least

during high water level. The growth depth

varies between 0 and 40 cm, in regulated

waters even down to 70 cm (Rintanen

1976: 142). The habitats are similar all over

N Fennoscandia (cf. Benum 1958: 317,

Wistrand 1962: 133, Söyrinki & Saari

1980: 124). Typical associates in the

riverside localities, like those by the

Sallijoki (7597:3456) and Syysjoki

(7685:3509), include Caltha palustris,

Carex aquatilis, Equisetum fluviatile,

Filipendula ulmaria, Galium uliginosum,

Juncus filiformis, Menyanthes trifoliata,

Parnassia palustris and Potentilla

palustris. In marshy and swampy lake

shores, as in its northernmost locality Lake

Alempi Kivivuopajajärvi near the

Kaamasjoki (7687:3501) and on a

Sphagnum fringe by a small lake S of Lake

Valkkojärvi (7663:3512), Lysimachia often

grows with other southern species like

Galium trifidum, Salix myrtilloides and

Vaccinium oxycoccos. In these sites also

Carex laxa, C. tenuiflora and Molinia

caerulea may belong to the companions.

In Inari Lapland Lysimachia thyrsiflora

is often sterile, but under favorable

conditions flowering is not unusual. Five

specimens in TUR and YME are fertile. In

most localities along the Ivalojoki recorded

by Kujala (1962: 178) the plants were

sterile. Only one occurrence in the lower

course of the river on a polluted, lush shore

near dwellings was fertile (1961 R.

Ruotsalo, H 418067, cf. Kujala l.c.).

Flowering does not usually start earlier than

in the latter half of July.

The two records in the birch belt are

both on the E side of the amphibolite fjelds

of Kietsimävaarat in an edaphically

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 107

favorable area in the upper Vaskojoki basin

(cf. Rautava 1964: 71, 1971). On the other

hand, Lysimachia thyrsiflora is clearly

more abundant along the less eutrophic

Ivalojoki than along the Vaskojoki. In

Kittilä it is fairly common in the eutrophic

Stratiotes-lakes (Salonen 1956), in the

Oulanka National Park it grows in localities

as well rich as poor in nutrients (Söyrinki &

Saari 1980: 124). Considered

amphicline/indifferent (Wistrand 1962:

133, Mäkinen & Kallio 1979: 18).

Amphicline or slightly basocline.

Dependence on culture. In the lower

course of the Ivalojoki Lysimachia

thyrsiflora may benefit from nutrients of

slight pollution caused by man (see above).

On the other hand, around Ivalo some

occurrences may have been destroyed due

to the development and growth of the

village. In any case, most of the localities in

Inari Lapland are not affected by man.

Hemerophilous in Mäkinen & Kallio (1979:

18). Hemeradiaphore.

JN

Primula stricta Hornem.

Indigenous, rare

Map 69

Distribution. Arctic, amphi-Atlantic, sometimes

regarded as incompletely circumpolar (Benum 1958:

317; Hultén 1958: 190, SKK III: 326, Hultén & Fries

1986: 1105). In N Europe from Iceland eastwards to

Novaja Zemlja and the Pechora river basin, widely in

N North America, but missing in Asia except for one

locality in the Yamal Peninsula (Fl. USSR XVIII:

170, Hultén 1958: map 171, Fl. Eur. 3: 17, Hultén &

Fries 1986: map 1469). Because of taxonomic

difficulties in the group (see the paragraph

“Morphology and taxonomy” below), the distribution

area is somewhat unclear (cf. also Hultén 1958: 190).

In Fennoscandia a weakly bicentric distribution

pattern, S part in the fjeld area of S Norway and

central Sweden (Oppland – Jämtland), N part north-

and eastwards from Nordland and the N fjelds of

Sweden to N Finland and the Kola Peninsula;

avoiding coastal areas in W and NW (Hultén 1971a:

map 1392, Roweck 1981: 381, Hultgård 1993,

Nilsson 2000: 160, Mossberg & Stenberg 2003: 461,

Lid & Lid 2005: 613).

In Troms frequent only in the N parts, and

mainly in inland, in Finnmark mostly in the lowland

and river valleys in the mainland (Norman II(2): 470,

Dahl 1934: 382, Benum 1958: 316, Hultén 1971a:

map 1392, Engelskjøn & Skifte 1995: 152, Elven et

al. 2013: 315). In Pechenga and the Kola Peninsula

along the coasts and in the fjeld area around Kirovsk

and Monchegorsk (Fl. Murm. V: map 1). A few

localities in Koillismaa in Finland and adjacent parts

of Russia (Keret Karelia, Lake Pjaozero), in Sompio

Lapland along the Kitinen, missing in Kittilä and

Enontekiö Lapland (Hult 1898: 163, Söyrinki 1956:

29, Rintanen 1968: 288, Hultén l.c., Söyrinki & Saari

1980: 122, Laitinen & Ohenoja 1990: 23, Hämet-

Ahti et al. 1998: 216, Lampinen & Lahti 2018; the

dot in Enontekiö in Lampinen & Lahti 2010 and

earlier versions of the atlas is an error).

InL ref. Along the Ivalojoki and

Vaskojoki especially in the middle course

(Kihlman 1884: 115, Wainio 1891: 44

(Kultala, Törmänen), Mikkola 1941: 35,

Kujala 1962: 178, Laine 1964: 115 (a pupil

specimen from the mouth of the Palojoki),

Rintanen 1968: 288), Lake Hietajärvi (1899

C. Fontell, H 417543, Kvist 1978: 54).

Paksusammali by the Teno (Kihlman l.c.).

Linkkapahta (Linkepakti) by the Kevojoki

(Kalliola 1937: 29, Laine et al. 1955: 130).

Four localities along the Teno in W

Utsjoki, four on the precipices by the N

Kevojoki, Tsuoggajoki and Lake

Kenesjärvi (Kallio & Mäkinen 1957: 26,

Laine 1970(II): 213).

Kevo 0.6 %, InL 10 %, 42 sq. H 32,

JYV 1, KUO 7, OULU 16, TUR 42, YME

10 spec.

Rare (130; 0.020). Inari: II (103;

0.025), Utsjoki: I (27; 0.011).

Difference***. Most localities along the

rivers Ivalojoki, Juutuanjoki, Näätämöjoki,

Vetsijoki, and the Teno. A few localities on

lake shores, mainly on the shores of Lake

Inari. Five localities on steep cliffs in

Utsjoki: two (Linkkapahta 7734:3499 and

Könkäänpahta 7738:3497) by the N part of

the Kevojoki, two by Lake Kenesjärvi

(Kenespahta 7734:3503 and 7735:3503)

108 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

and one by the Tsuoggajoki (7725:3503).

Considering the overall distribution in

Fennoscandia, the distribution pattern could

be described as montane (cf. Mäkinen &

Kallio 1979: 18), but in Inari Lapland P.

stricta is better regarded as a lowland

species.

FMF 0.152.

Vertical distribution. b: I (32; 0.013),

c: II (98; 0.028). Difference***. Mainly in

the coniferous zone. Range 55 m (mouth of

the Vetsijoki, 7765:3511) – 260 m (middle

course of the Repojoki, 7597:3449). Tr 960

m (Engelskjøn & Skifte 1995: 152), Fnm

374 m (Norman I(2): 875). Silvine.

Ecology. Most of the occurrences of

Primula stricta in Inari Lapland are on river

shores (cf. Kallio 1958), occasionally also

on lake shores. It grows on open shore

meadows, sandy or stony shore banks, and

crevices of cliffs e.g. by rapids. The banks

are regularly inundated and worn by

breaking ice in the spring, which keeps

them partly open and suitable for such a

weak competitor as P. stricta, which is

never copious in its localities. Usual

companions in these sites are Astragalus

alpinus, Bartsia alpina, Carex capillaris,

Equisetum variegatum, Parnassia palustris,

Pinguicula vulgaris, Selaginella

selaginoides and Tofieldia pusilla.

Another type of localities are the

shelves and ledges of W facing rock

precipices of steep cliffs (“pahta”)

preferably on basic bedrock. They are all

found in the gorges and valleys of the

Utsjoki and its tributaries. The weathering

rock surface in these localities and the

seeping water rich in nutrients make the

habitats favorable for P. stricta (Laine

1970(II): 213). Typical associates e.g. at

Linkkapahta are Campanula rotundifolia,

Carex capillaris, Carex norvegica subsp.

inferalpina, Draba daurica, Pinguicula

vulgaris, Poa glauca and Saxifraga nivalis.

In Finland similar localities are found in

Kuusamo (SKK III: 326), and they are

characteristic also in Scandinavia (Dahl

1934: 382, Benum 1958: 316, Wistrand

1962: 133, Roweck 1981: 381).

The seed production of this self-

compatible plant is fairly good in the study

area, and seedlings have been found

copiously e.g. in the localities along the

Vetsijoki. The species is also capable of

vegetative reproduction by short stolons

from overwintering buds.

Primula stricta favors substrate fairly

rich in nutrients and containing at least

some calcium. The species is therefore

usually considered basocline (Arwidsson

1943: 232, Benum 1958: 316, Wistrand

1962: 133, Laine 1970(II): 133, Mäkinen &

Kallio 1979: 18, Roweck 1981: 381,

Nilsson 2000: 160, Lid & Lid 2005: 613).

Basocline.

Morphology and taxonomy. Primula

stricta is a highly polyploid member of

Primula sect. Aleuritia Duby subsect.

Aleuritia. The subsection includes 21

arctic-alpine species with a basic

chromosome number x = 9, and comprises

at least five different ploidy levels from the

diploid 2n = 18 (e.g. P. farinosa L.) to P.

stricta, which is usually reported as 14-

ploid with 2n = 126 chromosomes (Löve &

Löve 1956: 125, Jørgensen et al. 1958: 92,

Lid & Lid 2005: 611, Guggisberg et al.

2006, Kelso 2009: 294). However,

deviating numbers between 2n = ca. 87 and

2n = 136 have also been reported

(Sokolovskaya & Strelkova 1960, Laane

1967: 52, Hultgård 1990: 123, Guggisberg

et al. l.c., Kelso l.c.). A count made from

material collected near the mouth of the

Vetsijoki (1973 U. Laine, TUR 243028)

gave the usual number 2n = 126 (Fig. 10).

Most Aleuritia polyploids are probably

of hybrid origin (Hultgård 1993). P. stricta

is most likely an allopolyploid, but its

origin in Europe and in N America may be

different (Guggisberg et al. 2006). In

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 109

Europe the progenitors are probably P.

scandinavica Bruun and P. scotica Hook.

Primula stricta consists of two slightly

different races in Europe, one mainly in W

Scandinavia, the other with a more NE

distribution (Hultgård 1993). The former

includes the type of the species and is

named var. stricta. It is slender, with

relatively narrow, efarinose leaves (farinose

in most species of Aleuritia), with fairly

few (1-6) flowers 4-6 mm in diameter and

long pedicels especially in the fruiting

stage. The latter is stouter, with broader,

somewhat farinose leaves, with more (5-10)

flowers 5-9 mm in diameter and shorter

pedicels (Hiitonen 1933: 557, Hultgård

1993, Nilsson 2000: 160, Lid & Lid 2005:

610). The name P. stricta var. obesior

Norman is usually applied to this taxon. It

is probably very highly polyploid (2n =

>180, Lid & Lid 2005: 613, but see also

Laane 1967 and Elven et al. 2013: 316). Its

status is still disputable; it is sometimes

regarded as a subspecies and may deserve

even a specific rank (Lid & Lid l.c.). It is

sometimes said to be linked by intermediate

forms with var. stricta (Dahl 1934: 382),

but according to Elven (2016),

intermediates are not yet documented.

Var. obesior has a few localities in

Troms, more in Finnmark especially

towards the east, and it is possibly the only

race in the northernmost European Russia

(Hiitonen 1933: 557, Lid & Lid 2005: 613,

Alm et al. 1995, Alm & Piirainen 1997b:

51, 60, Elven et al. 2013: 316, Elven 2016).

Although Inari Lapland is close to the

presumed area of the variety, it has not

been recorded there or elsewhere in

Finland. There is one specimen in OULU

under the name P. stricta collected from

“Ivalo, shore meadow” (1924 V. Isola,

OULU 49924) which resembles var.

obesior: it has a stout stem, faintly farinose

younger leaves and ten rather large flowers

on the main stem. However, the other

specimen on the same sheet is a typical var.

stricta. On the grounds of a single,

uncertain specimen it is too daring to

include var. obesior in the Finnish flora.

Dependence on culture. Primula

stricta is rarely met in places influenced by

man. There is one locality at Luhkkarpaihki

by the Teno (7738:3464) on seminatural

ground (Kallio & Mäkinen 1957: 26), but

the competition in such localities is usually

too heavy for P. stricta.

In Sompio Lapland P. stricta has

almost disappeared on the shores of the

Kitinen as the result of power plant

construction and water regulation

(Rautiainen et al. 2002: 135, Ryttäri et al.

2012: 274). Due to the decline and

fragmentation of the area of occupancy, P.

stricta is now classified as endangered

(EN) in Finland (Rassi et al. 2010: 199). At

present the majority of the Finnish

localities are situated in Inari Lapland,

where the rivers are still mostly flowing

free. There may be some threat because of

gold mining along the Ivalojoki as well as

the building of new houses and dwellings

on the shores, but most of the localities in

Fig. 10. Root-tip mitosis in Primula stricta with 2n =

126 chromosomes (an arrow pointing two

chromosomes overlapping each other). Voucher July

28, 1973 U. Laine (“gravelly shore of the Vetsijoki

ca. 1 km S of the highway”, TUR 243028). Del. A.

Lehmushovi.

110 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Inari Lapland are not threatened by man. –

Hemeradiaphore in Mäkinen & Kallio

(1979: 18). Hemerophobe.

JN

Primula veris L.

Introduced, very rare

Map 70

Distribution. European – W Asian with several

subspecies, subsp. veris endemic to Europe. In most

of Europe except for the extreme north and most of

the Mediterranean region (Fl. Eur. 3: 16, Hultén &

Fries 1986: map 1467). In Fennoscandia fairly

common in E Denmark, S part of Sweden, SE part of

Norway and SW Finland, scattered or rare, mainly as

an escape or casual, northwards up to the Arctic

Circle and in most of Norway (Hultén 1971a: map

1393, Hämet-Ahti et al. 1998: 216, Mossberg &

Stenberg 2003: 460, Lid & Lid 2005: 611).

Two specimens obviously as garden escapes

from Troms, Tromsø (TROM, T. Alm in an e-mail

February 14, 2014), five other observations from

Troms (Artsdatabanken 2015), two finds in Finnmark

(Sør-Varanger and Vadsø, Lid 1950, Nordhagen

1964). None of these accepted in Lid & Lid (2005:

611). Several specimens from Tromsø were earlier

regarded as P. veris (Benum 1950, 1958: 317, Hultén

1971a: map 1393), but belong to P. elatior (L.) Hill

(Engelskjøn & Skifte 1995: 152, Lid & Lid 2005:

611). No records from Pechenga or the Kola

Peninsula, one locality in the Kutsajoki area in Keret

Karelia (Fl. Murm. V: 31, map 1). No records from

Sompio or Kittilä Lapland (Lampinen & Lahti 2018;

the observation of P. veris in Kittilä in Lampinen &

Lahti 2013 and earlier versions of the atlas belongs to

P. elatior), one locality in Enontekiö, Iitto in a

meadow margin (H-Arch., H, Kastikka doc.

98329270, specimen not seen).

InL ref. Recorded as an old casual alien

in Inari Lapland (Hämet-Ahti et al. 1998:

216).

InL 1 %, 1 sq. H 1 spec. – The

frequency in Mäkinen & Kallio 1979: 18 is

incorrect and should be 0 %.

Very rare (1; 0.000). Inari: I (1;

0.000). Inari: in a former German camp site

near Kivioja rivulet (7644:3497, 1949 G.

Marklund, H 417719, Kastikka doc.

128561). – Many occurrences of subgenus

Primula (P. elatior, P. veris, P. vulgaris) in

N Scandinavia belong to P. elatior (e.g.

Parnela 1985, Engelskjøn & Skifte 1995:

152, Kastikka 2018, see also above).

Unfortunately it has not been possible to

find Marklund’s specimen in the collections

of H to check its identity, but his

determination is most probably correct. It is

accepted also in Hämet-Ahti et al. (1998:

216). Southern hemerochore.

FMF 0.004.

Vertical distribution. c: I (1; 0.000).

Alt. ca. 140 m. Silvine.

Ecology. The species was a casual

alien in a German camp site.

Dependence on culture. In N

Scandinavia Primula veris is usually a

casual introduced with grass seed or an

escape from cultivation (Benum 1958: 317,

SKK III: 320, Roweck 1981: 380). In Inari

it was a short-lived polemochore possibly

arrived with hay imported for fodder. Other

polemochorous occurrences of P. veris

introduced in N Finland by German troops

are known from Kn Hyrynsalmi, PeP

Tornio and Ks Kuusamo (Ahti & Hämet-

Ahti 1971: 69, Parnela 1985, Kastikka

2018). Ephemerophytic polemochore.

JN

Trientalis europaea L.

Lysimachia europaea (L.) U. Manns &

Anderb.

Indigenous, very frequent

Map 71

Distribution. Boreal Eurasia and NW North

America, with two subspecies, subsp. europaea in

Europe, N Asia and a restricted area in the interior of

NW North America, and subsp. arctica (Fisch.)

Hultén in the easternmost Asia and NW North

America (Hultén 1968: 751, Hiirsalmi 1969: 122,

Hultén & Fries 1986: 1106, map 1480). Common to

very common all over Fennoscandia (Hultén 1971a:

map 1404, Roweck 1981: 384, Hämet-Ahti et al.

1998: 220, Mossberg & Stenberg 2003: 465, Lid &

Lid 2005: 615).

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 111

Common to very common in Troms and

Finnmark (Norman I(2): 866, Dahl 1934: 383,

Benum 1958: 317, Engelskjøn & Skifte 1995: 153,

Lid & Lid l.c.), 39 localities in the Rastigaissa area

(Ryvarden 1969: 34). Common to very common in

Pechenga and the Kola Peninsula (Fellman 1831:

309, Kalliola 1932: 107, Kontuniemi 1932: 33,

Söyrinki 1939a: 342, Fl. Murm. V: map 3, Alm et al.

1997: 42, Mäkinen 2002: 17). Common to very

common in Sompio, Kittilä and Enontekiö Lapland

(Fellman 1835: 259, Hjelt & Hult 1885: 139, Wainio

1891: 44, Hult 1898: 163, Roivainen 1923: 293,

Hustich 1940c: 59, Lindén 1943: 78, Pertola 1961:

36, Montell 1962: 125, Piirainen & Piirainen 1991b,

Hämet-Ahti et al. 1998: 220, Lampinen & Lahti

2018), fairly common in the alpine belt of Enontekiö

Lapland (Lindén l.c., Laine 1958: 90).

InL ref. Common to very common

(Kihlman 1884: 115, Wainio 1891: 44,

Mikkola 1941: 35). Fairly common to

common in the Lemmenjoki area (Klockars

& Luther 1938, Rahkonen 1968: 20),

common along the Ivalojoki (Kujala 1962:

178) and Vaskojoki (Rautava 1969: 30).

Very common in the NE parts of Inari

(Såltin 1958). Fairly common to very

common in W Utsjoki (Laine et al. 1955:

130, Kallio & Mäkinen 1957: 27), scattered

but “possibly overlooked” according to

Laine (1964: 116). Fairly common and

evenly distributed along the Kevojoki

canyon (Laine 1970(II): 133).

Kevo 78.2 %, InL 92 %, 259 sq. H 9,

JYV 1, OULU 5, TUR 19, YME 2 spec.

Very frequent (4231; 0.649). Inari: VII

(2605; 0.605), Utsjoki: VII (1626; 0.737).

Difference***. Somewhat less frequent in

the area around Lake Inari and W and S of

it. Whole area.

FMF 0.982.

Vertical distribution. a: VII (452;

0.613), b: VII (1746; 0.768), c: VII (2033;

0.581). Difference a-b***, b-c***. Range

15 m (Pulmankijärvi, 7762:3539) – ca. 500

m (Peäldoajvi, 7675:3484). Tr 1010 m

(Engelskjøn & Skifte 1995: 153), Fnm 562

m (Norman I(2): 871), EnL 950 m (Laine

1958: 90). In Pite lappmark, Sweden,

common up to the low alpine belt, but

rapidly decreasing in the middle alpine belt

(Arwidsson 1943: 232, Wistrand 1962:

133). In the Khibiny Mountains in the Kola

Peninsula, sparse in the low alpine belt and

missing in the middle alpine belt (Alm et al.

1998: 135). Very frequent in all altitudinal

belts, but clearly commonest in the birch

belt (cf. Klockars & Luther 1938, Laine

1970(II): 133). Vertical ubiquitous.

Ecology. Trientalis europaea is one of

the commonest woodland plants in Finland,

growing in many kinds of coniferous and

deciduous forests, spruce mires, pine bogs

and even in treeless fens and alpine

meadows (Roivainen 1923: 293, Kalliola

1939: 116, Hämet-Ahti 1963a, Kujala

1964: 85, Hiirsalmi 1969: 128, Ahti &

Hämet-Ahti 1971: 69, Söyrinki et al. 1977).

In Inari Lapland it is common in almost all

coniferous and subalpine forest types

(Kujala 1929: 104-118, Hämet-Ahti

1963a). It occurs as well in the dry and

unfertile subalpine Empetrum-Lichenes

type (sELiT, Hämet-Ahti 1963a: 37-49) as

in the luxuriant meadow forest by

Keneskoski in Utsjoki (Hämet-Ahti 1963a:

62, 91-96). In the driest forest types in the

subalpine belt Trientalis is almost totally

confined to the mesic patches under the

birches with e.g. Linnaea borealis,

Pedicularis lapponica, Pleurozium

schreberi and Peltigera aphthosa (Hämet-

Ahti 1963a: 46). Also, according to

Hiirsalmi (1969: 129-132), in Inari Lapland

Trientalis is less frequent in dry heath

forests than in mesic ones and its

occurrences are less contiguous.

In the Kevojoki canyon Trientalis has

profited from greater light intensity and

increased amount of nutrients after the

destruction of the birch canopy caused by

the mass occurrence of the Autumnal Moth

Epirrita autumnata (Borkhausen) in the

middle of 1960’s (Laine 1970(II): 133, see

Kallio & Lehtonen 1973). Also in the

Peäldoajvi fjeld area Trientalis was

112 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

exceptionally abundant in those sample

plots where birch canopy was destroyed by

the moth (Koivistoinen 1964: 59).

In swampy habitats Trientalis prefers

drier parts with a thin peat layer, and

usually grows on the hummocks (Cajander

1903: 16, Hiirsalmi 1969: 130). E.g. in the

Lemmenjoki National Park it is found in

somewhat paludified habitats (Rahkonen

1968: 20). In the alpine belt it is mostly

concentrated along turfy places near brooks

(Kihlman 1884: 115). Trientalis also grows

in slopes and screes and in flooded stony or

pebbly sites by rivers (Hiirsalmi 1969: 130,

Laine 1970(II): 133). In higher altitudes

sufficient snow cover is important (cf.

Roweck 1981: 384).

Flowering began in Utsjoki on July 4

in 1795, and on June 27 in 1797 because of

a warm period in the end of June (Castrén

1803). In 1996-2006 in Utsjoki, Kevo,

dates were remarkably earlier, probably

partly reflecting the ongoing climate

change: the earliest date recorded was June

3 and the latest June 24 (Alanen 2007).

Although flowering is often profuse, seed

production is scarce and seedlings are met

only rarely (Kontuniemi 1932: 33, Söyrinki

1939a: 343-344, cf. Hiirsalmi 1969: 154-

157). On the other hand, vegetative

reproduction and regeneration from long,

thread-like rhizomes is very effective,

especially in open habitats like road banks

and forest fire areas (Kujala 1926: 28,

Söyrinki l.c., Vanhatalo 1965: 124,

Hiirsalmi 1969: 157-159).

Amphicline/Ca-indifferent (Arwidsson

1943: 232, Benum 1958: 317, Wistrand

1962: 133, Laine 1970(II): 133, Mäkinen &

Kallio 1979: 18, Nilsson 2000: 161, Lid &

Lid 2005: 615). Amphicline.

Morphology and taxonomy. In

Lapland Trientalis seems to be more

variable than in S Finland (Hiirsalmi 1969:

163-164, Laine 1970(II): 31). The variation

in the vegetative parts is largely

environmental, but e.g. the length/breadth

ratio of the whorl leaves is partly

genetically determined. In Inari Lapland

they are often relatively shorter and the leaf

tip is more obtuse than in S Finland

(Hiirsalmi 1969: 161). In this respect the

plants show some similarity with the E

Asian and N American subsp. arctica.

There is considerable, genetically

determined variation also in the floral

organs. E.g. petals are often somewhat

shorter and narrower in the north, although

the variation is vast (Hiirsalmi 1969: 162-

164). Especially in higher altitudes and in

the alpine belt petals are often pink or

reddish (“f. rosea Neum.”) and even leaves

may be reddish or brownish (Montell 1910,

1962: 125, Valle 1933a: 271, Arwidsson

1943: 232, Laine 1958: 90, Kujala 1962:

178, Hiirsalmi l.c., Roweck 1981: 384).

The coloring is caused by anthocyanin

pigments. Their amount seems to depend

largely on external factors, especially on

light and temperature conditions (Hiirsalmi

1969: 165). Also the low-growing plant,

mf. minor (Brenn.) Mela & Cajander, often

met in the alpine belt is an environmental

modification (Hiirsalmi 1969: 166).

Dependence on culture. Trientalis

grows as an apophyte in meadows, forest

margins and clearings, seminatural

pastures, road sides and yards (Ahti &

Hämet-Ahti 1971: 69, Alm et al. 1997: 42,

Vinnamo 1963: 10). In these habitats it

benefits from increased light and

diminished competition. In Nukkumajoki,

Inari, it was recorded in all Lappish

medieval winter villages, although it was

missing in the surrounding pine heath

(Suominen 1975). The species may rapidly

colonize open road margins by vegetative

reproduction, flowering profusely in a few

years (Vanhatalo 1965: 124). Also in the

Kuusamo district it is sometimes clearly

hemerophilous in meadows and forest

margins (Ahti & Hämet-Ahti 1971: 69). On

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 113

the other hand, it may suffer from human

activities and forest fires (Kujala 1964: 85).

Hemeradiaphore in Mäkinen & Kallio

(1979: 18). Hemeradiaphore.

JN

Acknowledgements. We are grateful to Torbjørn

Alm for information concerning Primula veris in

TROM, Annikki Kestilä and Annu Ruotsalainen for

arranging the loan from OULU, Raino Lampinen for

information in Kastikka, Aaro Lehmushovi for the

drawing of the chromosomes of Primula stricta,

Nelly Llerena for scanning the herbarium sheets, and

Outi Vainio for information on the specimens in

KUO.

We express our best thanks to the following

persons, who have made contributions to the floristic

study of Inari Lapland during 2002-2017: Krõõt

Aasamaa, Esko Aikio, Marjaana Alanen, Kaija Helle,

Timo Hietanen, Riikka Juutinen, Heidi Kaipiainen-

Väre, Sinikka Kallio, Markku Kantola, Jouko

Karjalainen, Antti Karlin, Tuula-Riitta Karlin, Paula

Karlström, Esa Karpoff, Leena Kaskisto, Vesa

Kaskisto, Matti Kouvo, Tuomo Kuitunen, Hannu

Kulmala, Kyösti Laaksonen, Erkki Laasonen, Leena

Laasonen, Kaija Laine, Ulla Liimatainen, Liisa

Mäkinen, Anu Munne, Aimo Myllynen, Esteri

Ohenoja, Olli Osmonen, Riitta Parviainen, Matti

Piiparinen, Petteri Polojärvi, Maire Puikko, Pekka

Rahko, Ropi Rahko, Pekka Rautiainen, Arto

Saikkonen, Jari Särkkä, Kimmo Syrjänen, Eini

Tallgren, Erkki Toivonen, Saara Tynys, Henry Väre,

Jukka Vauras, Aune Veersalu, Mia Vuomajoki, and

Jukka Ylikörkkö.

References

AEDO, C. 2009: Geranium. – In: Aedo, C.,

Estébanez, B. & Navarro, C. (ed.); with

contributions from Raab-Straube, E. von &

Parolly, G.: Geraniaceae. Euro+Med Plantbase –

the information resource for Euro-Mediterranean

plant diversity. – Published on the Internet

http://ww2.bgbm.org/EuroPlusMed/. – Accessed

on 7.3.2012.

AEDO, C. 2012: Geranium Taxonomic Information

System. – Published on the Internet

http://www.geranium.es/index.php. – Accessed

on 7.3.2012.

AHOLA, J. 1929: Kasvitarha tuottaa sadon

napapiirin takanakin. – Kotiliesi 1929: 202-204,

217.

AHTI, T. 1968: Micromycetes (Peronosporaceae,

Erysiphales and Uredinales) new to the province

of Kuusamo, N.E. Finland. – Karstenia 8: 5-8.

AHTI, T. & HÄMET-AHTI, L. 1971:

Hemerophilous flora of the Kuusamo district,

northeast Finland, and the adjacent part of

Karelia, and its origin. – Annales Botanici

Fennici 8(1): 1-91.

AHTI, T., HÄMET-AHTI, L. & JALAS, J. 1968:

Vegetation zones and their sections in

northwestern Europe. – Annales Botanici Fennici

5(3): 169-211.

AIKEN, S. G., DALLWITZ, M. J., CONSAUL, L.

L., MCJANNET, C. L., BOLES, R. L., ARGUS,

G. W., GILLETT, J. M., SCOTT, P. J., ELVEN,

R., LEBLANC, M. C., GILLESPIE, L. J.,

BRYSTING, A. K., SOLSTAD, H. & HARRIS,

J. G. 2007: Flora of the Canadian Arctic

Archipelago: Descriptions, Illustrations,

Identification, and Information Retrieval. – NRC

Research Press, National Research Council of

Canada, Ottawa. Published on the Internet

http://nature.ca/aaflora/data. – Accessed on

14.5.2012.

ALANEN, M. 2003: Geranium sylvaticum:

Eriväristen ja -kokoisten kukkien osuudet

Utsjoella 2003. – Manuscript in the Archives of

the Kevo Research Station. 8 pp.

ALANEN, M. 2007: Flowering and seed ripening of

the most common plant species around the Kevo

biological and weather station in Utsjoki

commune 1995-2006. –

http://www.kevo.utu.fi/tutkimus/fenologia_kksie

m.html. – Accessed on 5.5.2007.

ALANEN, M. 2010: Jotakin Rhodosta. – Manuscript

in the Archives of the Kevo Research Station. 3

pp.

ALM, T. 1992: Floraen i Finnmark. 5.

Perikumfamilien (Hypericaceae). – Polarflokken

16 (1): 149-152.

ALM, T. 1993a: Floraen i Finnmark. 6. Lyngfamilien

(Ericaceae). – Polarflokken 17(1): 7-54.

ALM, T. 1993b: Floraen i Finnmark. 8.

Tamariskfamilien (Tamaricaceae). – Polarflokken

17(1): 115-118.

ALM, T. 1993c: Floraen i Finnmark. 9.

Kornellfamilien (Cornaceae). – Polarflokken

17(1): 119-122.

ALM, T. 1993d: Floraen i Finnmark. 10.

Fjellprydfamilien (Diapensiaceae). –

Polarflokken 17(1): 123-126.

ALM, T. 2000: Nye funn av karplanter med

avvikende farge på blomster og baer i Nord-

Norge 1998-2000. – Polarflokken 24: 225-230.

ALM, T. & PIIRAINEN, M. 1997a: Vaarlamasaari –

changes in the cultural landscape of former

Finnish farm sites on a Paatsjoki/Pasvik River

114 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

island (Pechenga, Russia) during the last 60

years. – Memoranda Societatis pro Fauna et Flora

Fennica 73(2): 37-44.

ALM, T. & PIIRAINEN, M. 1997b: Den finske

botanikeren A. A. Arrhenius' opptegnelser fra

Varanger 1880. – Polarflokken 21(1): 45-67.

ALM, T. & PIIRAINEN, M. 2000a: Krigsspredte

arter i Sør-Varanger, Finnmark: knollerteknapp

Lathyrus linifolius. – Blyttia 58: 174-177.

ALM, T. & PIIRAINEN, M. 2000b: Amerikamjølke

(Epilobium adenocaulon) på Kirkenäs – ny for

Finnmark. – Polarflokken 24(1): 3-9.

ALM, T., ALSOS, I., BRÅTHEN, K. A. &

SOMMERSEL, G-A. & ØIESVOLD, S. 1993:

Bidrag til floraen i Finnmark. – Polarflokken

17(3): 519-537.

ALM, T., ALSOS, I. G., BRÅTHEN, K. A.,

NORDTUG, B., OFTEN, A. & SOMMERSEL,

G-A. 1994: Bidrag til floraen i Finnmark. Nyfunn

i 1994, med særlig vekt på kulturspredte arter. –

Polarflokken 18(2): 189-226.

ALM, T., ALSOS, I. G. & OFTEN, A. 1995: Rike

skoglier på Altenes i Alta, Finnmark. –

Polarflokken 19: 119-130.

ALM, T., ALSOS, I. G., KOSTINA, V. A., OFTEN,

A. & PIIRAINEN, M. 1997: Cultural landscapes

of some former Finnish farm sites in the

Paaz/Pasvik/Paatsjoki area of Pechenga, Russia.

– Tromura, Naturvitenskap 82: 1-42.

ALM, T., ALSOS, I. G., OFTEN, A. & PIIRAINEN,

M. 1998: Botaniska intryck från Kolahalvön

(Botanical impressions from the Kola peninsula).

– Svensk Botanisk Tidskrift 92: 121-137.

ALM, T., OFTEN, A. & PIIRAINEN, M. 2000c:

Krigsspredte arter i Sør-Varanger, Finnmark: stor

gjeldkarve Pimpinella major. – Blyttia 58: 120-

124.

ALM, T., GAMST, S. B., GAMST, U. B. &

SORTLAND, A. B. 2004: Kulturspredte arter i

Tromsø (Troms) ved starten av et nytt årtusen. 1.

Innledning og artsomtaler: Hampefamilien

(Cannabaceae) til skjermplantefamilien

(Apiaceae). – Polarflokken 28(1): 3-98.

ANDRULAЇTIS, C. Y., VODOPJANOVA, N. C.,

IVANOVA, M. M., KISELEVA, A.

A.,MALYSEV, L. I. & PETROTSENKO, Y. N.

1976: Sostav flory Putorana. – In: Izdatelstvo

"Nauka". – Novosibirsk. Pp. 40-162.

ANONYMOUS 1921a: Suomalaisen Eläin- ja

Kasvitieteellisen Seuran Vanamon kokous

20.XI.1920. – Luonnon Ystävä 25(1): 19-21.

ANONYMOUS 1921b: Mötet den 4 december 1920.

– Meddelanden af Societas pro Fauna et Flora

Fennica 47: 30-32.

ARTSDATABANKEN 2015: Species Map Service

1.6, provided by the Norwegian Biodiverstiy

2015. – Published on the Internet

artskart.artsdatabanken.no/default.aspx. –

Accessed on 8.12.2015.

ARWIDSSON, T. 1935: Empetrum hermaphroditum

(Lange) Hagerup och E. nigrum L. s.str. i

Norden. – Bot. Not. 1935: 181-183.

ARWIDSSON, T. 1943: Studien über die

Gefässpflanzen in den Hochgebirgen der Pite

Lappmark. – Acta Phytogeographica Suecica 17:

1-274.

ASIKAINEN, E. & MUTIKAINEN, P. 2003: Female

frequency and relative fitness of females and

hermaphrodites in gynodioecious Geranium

sylvaticum (Geraniaceae). – American Journal of

Botany 90(2): 226-234.

ASIKAINEN, E. & MUTIKAINEN, P. 2005a: Pollen

and resource limitation in a gynodioecious

species. – American Journal of Botany 92: 487-

494.

ASIKAINEN, E. & MUTIKAINEN, P. 2005b:

Preferences of pollinators and herbivores in

gynodioecious Geranium sylvaticum. – Annals of

Botany 95(5): 879-886.

AUER, A. V. 1938: Kasvistollisia havaintoja

Pohjois-Suomesta II. – Memoranda Societatis pro

Fauna et Flora Fennica 14: 110-123.

AUER, A. V. 1944: Kasvistollisia havaintoja

Pohjois-Suomesta III. – Memoranda Societatis

pro Fauna et Flora Fennica 19: 44-57.

BENUM, P. 1950: Nyare plantefunn i Nord-Noreg. –

Blyttia 8: 1-11.

BENUM, P. 1958: The flora of Troms Fylke. A

floristic and phytogeographical survey of the

vascular flora of Troms fylke in Northern

Norway. – Tromsø Museums Skrifter 6: 1-402 +

546 maps.

BERG, R. Y. 1963: Disjunksjoner i Norges fjellflora

og de teorier som er framsatt til forklaring av

dem. – Blyttia 21: 133-177.

BERTILSSON, A., ARONSSON, L.-E., BOHLIN,

A., BÖRJESON, G., GEIJER, M., IVARSSON,

R., JANSON, O. & SAHLIN, E. 2002:

Västergötlands flora. – SBF-förlaget, Lund. 743

pp.

BJÖRKMAN, G. 1939: Kärlväxtfloran inom Stora

Sjöfallets Nationalpark jämte angränsande delar

av norra Lule Lappmark. – Kungliga Svenska

Vetenskapsakademiens Avhandlingar i

Naturskyddsärenden 2: 1-224.

BJÖRKMAN, G. 1965: Tillägg till kärlväxtfloran

inom Stora Sjöfallets Nationalpark jämte

angränsande delar av norra Lule Lappmark. –

Kungliga Svenska Vetenskapsakademiens

Avhandlingar Naturskyddsärenden 21: 1-128.

BLAMEY, M. & GREY-WILSON, C. 1989: The

illustrated flora of Britain and Northern Europe. –

London. 544 pp.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 115

BLOMGREN, E., FALK, E. & HERLOFF, B. (eds.)

2011: Bohusläns Flora. – Föreningen Bohusläns

Flora. Lund. 731 pp.

BLUMER, S. 1967: Echte Mehltaupilze

(Erysiphaceae). Ein Bestimmungsbuch fur die in

Europa vorkommenden Arten. – Gustav Fischer

Verlag. Jena. 436 pp.

BÖCHER, T. W., HOLMEN, K. & JAKOBSEN, K.

1968: The flora of Greenland. – Copenhagen. 312

pp.

BORGEN, L. & ELVEN, R. 1983: Chromosome

numbers of flowering plants from northern

Norway and Svalbard. – Nordic Journal of

Botany 3: 301–306.

BRANDRUD, K. H. & BORGEN, L. 1986: Viola

epipsila, V. palustris and their hybrid in SE

Norway. – Acta Universitatis Upsaliensis,

Symbolae Botanicae Upsalienses 27(2): 19-24.

CAJANDER, A. K. 1903: Beiträge zur Kenntniss der

Vegetation der Hochgebirge zwischen Kittilä und

Muonio. – Fennia 20(9): 1-37.

CALDER, J. A. & TAYLOR, R. L. 1968: Flora of

the Queen Charlotte islands. 1. Systematics of the

vascular plants. – Canada Department of

Agriculture, Monograph no. 4. 659 pp.

CASTRÉN, S. 1803: Observationer gjorde i Utsjoki

Lappmark åren 1795 och 1797. – Kongliga

Finska Hushållnings-Sällskapets Handlingar 1:

344-355.

CESKA, A. & CESKA, O. 1986: Notes on

Myriophyllum (Haloragaceae) in the Far East:

The identity of Myriophyllum sibiricum

Komarov. – Taxon 35(1): 95-100.

CLEVE, A. 1898: Studier öfver några svenska

växters groningstid och förstärkningsstadium. –

Dissertation. Uppsala. 98 pp.

COOMBE, D. E. & WHITE, F. 1951: Notes on

calcicolous communities and peat formation in

Norwegian Lappland. – Journal of Ecology 39:

33-65.

DAHL, O. 1934: Floraen i Finnmark fylke. – Nyt

Magazin for Naturvidenskaberne 69: 1-430 +

XVII Pl.

DAHL, E. 1951: On the relation between summer

temperature and the distribution of alpine

vascular plants in the low-lands of Fennoscandia.

– Oikos 3(1): 22-52.

DANIHELKA, J., HOF, K. VAN DEN,

MARCUSSEN, T. & JONSELL, B. 2010: Viola

montana and V. persicifolia (Violaceae). – Taxon

59(6): 1900-1902.

DOROGOSTAISKAYA, E. V. 1972: Sornye

rasteniya Krajnego Severa SSSR. – Rastitel'nost'

Krajnego Severa SSSR i ee osvoenie 13: 1-182.

DUTILLY, A., LEPAGE, E. & DUMAN, M. 1953:

Contribution à la flore du basin de la Baie

d’Ungava. – Catholic University of America,

Contribution of the Arctic Institute 4F. 104 pp.

EDQVIST, M. & KARLSSON, T. (eds.) 2007:

Smålands flora. – SBF-förlaget. Uppsala. 880 pp.

EDVARDSEN, H. & ELVEBAKK, A. 1981:

Vurdering av botaniske verneverdier langs

Alta/Kautokeino-vassdraget. – Polarflokken 5:

90-103.

EKLUND, O. 1926: Zur Systematik und Verbreitung

der Gattung Oxycoccus Hill in Fennoscandia

orientalis. – Acta Soc. Fauna Flora Fennica

55(4): 1-16.

ELFVING, F. 1897: Anteckningar om kulturväxterna

i Finland. – Acta Societatis pro Fauna et Flora

Fennica 14(2): 1-116.

ELFVING, F. 1927: Tärkeimmät viljelyskasvit. 2.

painos. – Osakeyhtiö Valistus. Helsinki. 284 pp.

ELVEN, R. 2010: Norsk vintergrønn og

legevintergrønn – Pyrola norvegica og P.

rotundifolia – en kommentar. – Blyttia 68: 173-

178.

ELVEN, R. (ed.) 2016: Panarctic Flora. Annotated

Checklist of the Panarctic Flora (PAF). Vascular

Plants. – Published on the Internet

http://nhm2.uio.no/paf/. – Accessed on

23.2.2016.

ELVEN, R. & SOLSTAD, H. 2000: Blygmjølke

(Epilobium ciliatum) i Finnmark, og litt

kommentarer om aggressive immigranter. –

Polarflokken 24: 89-96.

ELVEN, R., FREMSTAD, E. & PEDERSEN, O.

2013: Distribution maps of Norwegian vascular

plants. IV. The eastern and northeastern

elements. – Academika Publishing. Trondheim.

489 pp.

ENGELSKJØN, T. & KNABEN, G. 1971:

Chromosome numbers of Scandinavian arctic-

alpine plant species. III. – Acta Borealia, A

Scientia 28: 1-30.

ENGELSKJØN, T. & SKIFTE, O. 1995: The

vascular plants of Troms, North Norway. Revised

distribution maps and altitude limits after Benum:

The flora of Troms fylke. – Tromura

Naturvitenskap 80: 1-227.

ERIKSSON, B. 1974: On ascomycetes on

Diapensiales and Ericales in Fennoscandia. 2.

Pyrenomycetes. – Svensk Botanisk Tidskrift.

68(2): 192-234.

ERVI, L. O. 1956: Karpalolajien morfologiasta ja

viljelymahdollisuuksista Suomessa. – Acta

Agralia Fennica 92: 1-148.

EUROLA, S. 1967: Über die Vegetation der

Alluvialwiesen im Gebiet der geplanten Stauseen

von Lokka und Porttipahta im Finnischen

Lappland. – Aquilo, Ser. Botanica 5: 1-119.

116 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

EUROLA, S., HUTTUNEN, S. & KUKKO-OJA, K.

1995: Suokasvillisuusopas. 2. painos. – Oulanka

Reports 14: 1-85.

FAEGRI, A. 1982a: Et bortglemt fennoscandisk

tusenblad (Myriophyllum) taxon. (A

Fennoscandian taxon that has fallen into

oblivion). – Blyttia 40: 149-153.

FAEGRI, A. 1982b: The Myriophyllum spicatum

group in north Europe. – Taxon 3: 467-471.

FELLMAN, J. 1831: Index plantarum

phanerogamarum in territorio Kolaënsi lectarum.

– Bulletin de la Société Impériale des

Naturalistes de Moscou 3: 299-328.

FELLMAN, J. 1835: Index plantarum in Lapponia

Fennica lectarum. A. Plantae Phanerogamae. –

Bulletin de la Société Impériale des Naturalistes

de Moscou 8: 245-289.

FERNALD, M. L. 1918: American variations of

Epilobium, section Chamaenerion. – Rhodora 20:

1-10.

FERNALD, M. L. 1919: Two new Myriophyllum

and a species new to the United States. –

Rhodora 21: 120-124.

Flora Arctica URSS. VIII Geraniaceae –

Scrophulariaceae. 1980. Eds. Tolmatchev, A. I. &

Jurtzev, B. A. – Leningrad. 336 pp.

Flora Europaea 2. 1968. Eds. Tutin, T. G., Heywood,

V. H., Burges, N. A., Valentine, D. H., Walters,

S. M. & Webb, D. A. – Cambridge. 455 pp.

Flora Europaea 3. 1972. Eds. Tutin, T. G., Heywood,

V. H., Burges, N. A., Valentine, D. H., Walters,

S. M. & Webb, D. A. – Cambridge. 370 pp.

Flora Murmanskoy Oblasti IV. 1959. Ed. Poyarkova,

A. I. – Moskva & Leningrad. 394 pp.

Flora Murmanskoy Oblasti V. 1966. Eds. Gorodkov,

B. N. & Poyarkova, A. I. – Moskva & Leningrad.

548 pp.

Flora Nordica 1. 2000. Ed. Jonsell, B. – The Bergius

Foundation. Stockholm. 344 pp.

Flora Nordica 6. 2010. Eds. Jonsell, B. & Karlsson,

T. – The Swedish Museum of Natural History.

Stockholm. 298 pp.

Flora of North America north of Mexico 8. 2009. Ed.

Flora of North America Editorial Committee. –

New York. 585 pp.

Flora of Russia. The European part and bordering

regions. Vol. 9. 2006. Ed. N. N. Tzvelev. –

London. 722 pp.

Flora USSR XV. 1949. Eds. Shishkin, B. K. &

Bobrov, E. G. – Moskva – Leningrad. 742 pp. +

maps.

Flora USSR XVIII. 1952. Eds. Shishkin, B. K. &

Bobrov, E. G. – Moskva-Leningrad. 802 pp. +

maps.

FREMSTAD, E. & NORMANN, Ö. 1982:

Inventering av rik lövskog i Troms. (Summary:

Registrations of localities with rich deciduous

forests in Troms, Northern Norway). – Tromura,

Naturvitenskap 34: 1-97.

FRIES, T. 1913: Botanische Untersuchungen im

nördlichsten Schweden. Ein Beitrag zur Kenntnis

der alpinen und subalpinen Vegetation in Torne

Lappmark. – Vetenskapliga och Praktiska

Undersökningar i Lappland anordnade af

Luossavaara-Kiirunavaara Aktiebolag 1: 1-361.

FRIES, T. 1925: Die Rolle des Gesteingrundes bei

der Verbreitung der Gebirgspflanzen in

Skandinavien. – Svenska Växtsociologiska

Sällskapets Handlingar 6: 1-17.

FRÖBERG, L. 2006: Blekinges flora. – SBF-

förlaget, Uppsala. 856 pp.

GEORGSON, K., JOHANSSON, B., JOHANSSON,

Y., KUYLENSTIERNA, J., LENFORS, I. &

NILSSON, N.-G. 1997: Hallands flora. – Lund.

798 pp.

GJÆREVOLL, O. 1950: The snow-bed vegetation in

the surroundings of Lake Torneträsk, Swedish

Lappland. – Svensk Botanisk Tidskrift 44(2):

387-440.

GJÆREVOLL, O. 1956: The plant communities of

the Scandinavian alpine snow-beds. – Kongel.

Norske Vidensk. Selsk. Skr. 1956(1): 1-405.

GJÆREVOLL, O. 1961: XIII International

Phytogeographical Excursion to Finnmark and

Troms 26.7.-5.8.1961. – Trondheim. 26 pp.

GJÆREVOLL, O. 1990: Alpine plants. – In: Berg, R.

Y., Fægri, K. & Gjærevoll, O. (eds.): Maps of

distribution of Norwegian vascular plants. –

Trondheim. 123 pp. + 37 pl.

GLEASON, H. A. 1952: Illustrated Flora of

Northeastern United States and adjacent Canada

2. – Lancaster, PA. 655 pp.

GROTENFELT, G. 1897: Suomen maanviljelys.

Yleiskatsaus kuvilla ja kartoilla varustettu. –

Söderström. 328 pp.

GUGGISBERG, A., MANSION, G., KELSO, S. &

CONTI, E. 2006: Evolution of biogeographic

patterns, ploidy levels, and breeding systems in a

diploid-polyploid species complex of Primula. –

New Phytologist 171(3): 617-632.

HAAPASAARI, M. 1988: The oligotrophic heath

vegetation of northern Fennoscandia and its

zonation. – Acta Botanica Fennica 135: 1-219 +

Tables 1-23.

HÆGGSTRÖM, C-A. & HÆGGSTRÖM, E. 2010:

Ålands flora. 2nd ed. – Ekenäs. 528 pp.

HAGERUP, O. 1954: Autogamy in some drooping

Bicornes flowers. – Botanisk Tidsskrift 51: 103-

116.

HAGMAN, M. 1953: Några iaktagelser över

Myricaria germanica (L.) Desv. i Utsjoki, Li. –

Memoranda Societatis pro Fauna et Flora

Fennica 28: 4-7.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 117

HÄMET-AHTI, L. 1963a: Zonation of the mountain

birch forests in northernmost Fennoscandia. –

Annales Botanici Societatis Zoologicae

Botanicae Fennicae Vanamo 34(4): 1-127.

HÄMET-AHTI, L. 1967b: Chaerophyllum bulbosum

L. ssp. bulbosum and ssp. prescottii (DC.)

Nyman in Finland. – Annales Botanici Fennici

4(4): 417-421.

HÄMET-AHTI, L., SUOMINEN, J., ULVINEN, T.,

UOTILA, P. & VUOKKO, S. (eds.) 1984:

Retkeilykasvio (Field Flora of Finland). –

Suomen Luonnonsuojelun Tuki Oy. Helsinki.

544 pp.

HÄMET-AHTI, L., SUOMINEN, J., ULVINEN, T.,

UOTILA, P. & VUOKKO, S. (eds.) 1986:

Retkeilykasvio (Field Flora of Finland). Ed. 3.

Suomen Luonnonsuojelun Tuki Oy. Helsinki.

598 pp.

HÄMET-AHTI, L., SUOMINEN, J., ULVINEN, T.

& UOTILA, P. (eds.) 1998: Retkeilykasvio (Field

Flora of Finland). Ed. 4. – Finnish Museum of

Natural History, Botanical Museum. Helsinki.

656 pp.

HÄMET-AHTI, L., KURTTO, A., LAMPINEN, R.,

PIIRAINEN, M., SUOMINEN, J., ULVINEN,

T., UOTILA, P. & VÄRE, H. 2005a: Lisäyksiä ja

korjauksia Retkeilykasvion neljänteen

painokseen. – Lutukka 21(2): 41-85.

HÄMET-AHTI, L., KURTTO, A., LAMPINEN, R.,

PIIRAINEN, M., SUOMINEN, J., ULVINEN,

T., UOTILA, P. & VÄRE, H. 2005b: Lisäyksiä ja

korjauksia Retkeilykasvion neljänteen

painokseen. Jälkimmäinen osa: auktoreita ja

synomyymejä. – Lutukka 21(3): 109-116.

HARMAJA, H. 1990: New names and nomenclatural

combinations in Rhododendron (Ericaceae). –

Annales Botanici Fennici 27: 203-204.

HARMAJA, H. 1991: Taxonomic notes on

Rhododendron subsection Ledum (Ledum,

Ericaceae), with a key to its species. – Annales

Botanici Fennici 28: 171-173.

HARVIAINEN, S., EKKO, P., BORG, M. &

FAGERSTÉN, R. 1968: Kasvillisuustietoja

Termislehdosta (EnL). – Luonnon Tutkija 72: 51-

52.

HÄYRÉN, E. 1925: Spridda anteckningar om

kulturväxterna i Lapland. – Trädgårdsodlaren

1925: 188-190.

HEIKINHEIMO, O. & RAATIKAINEN, M. 1971:

Paikan ilmoittaminen Suomesta talletetuissa

biologisissa aineistoissa. – Annales Entomologici

Fennici 37(1a): 1-30 + 1 map.

HEIKINHEIMO, O. & RAATIKAINEN, M. 1981:

Ruutukoordinaattien ja paikannimien käyttö

Suomessa. (Grid reference and names of

localities in the recording of biological finds in

Finland). – Notulae Entomologicae 61: 133-154.

HEIKKINEN, L. 1948: Saksalaiset sotajoukot

kasvien levittäjinä. – Luonnon Tutkija 52(1): 25-

26.

HEIKKINEN, L. 1959: Sota-ajan tulokaskasvistosta

Hyrynsalmella. – Memoranda Societatis pro

Fauna et Flora Fennica 34: 57-71.

HEIKKINEN, R. & KALLIOLA, R. 1989:

Vegetation types and map of the Kevo nature

reserve, northernmost Finland. – Kevo Notes 8:

1-39.

HEIKKINEN, R. & KALLIOLA, R. 1990: The

vascular plants of the Kevo Nature Reserve

(Finland); an ecological-environmental approach.

– Kevo Notes 9: 1-56.

HELANDER, E. 1965: Piirteitä Inarin

kyläkasvistosta. – M. Sc. Thesis, Department of

Biology, University of Turku. 124 pp. + 22 maps.

HELENIUS, O. 1948: Lapin alppiruusu

(Rhododendron lapponicum) Inarin Lapista. –

Luonnon Tutkija 52(4): 127.

HERLIN, N. 1944b: Beitrag zur Kenntniss der Flora

in der Provinz Karelia Keretina in Ost-Karelien. –

Memoranda Societatis pro Fauna et Flora

Fennica 19: 91-103.

HIIRSALMI, H. 1969: Trientalis europaea L. A

studyof the reproductive biology, ecology and

variation in Finland. – Annales Botanici Fennici

6: 119-173.

HIITONEN, I. 1933: Suomen kasvio. – Helsinki. 771

pp + 1 map.

HILTUNEN, R. 1992: Kuulumisia Luostolta. –

Lutukka 8(3): 88-90.

HJELT, H. 1911: Conspectus Florae Fennicae. IV. –

Acta Societatis pro Fauna et Flora Fennica 35(1):

1-411.

HJELT, H. 1919: Conspectus Florae Fennicae. V. –

Acta Societatis pro Fauna et Flora Fennica 41(1):

1-502.

HJELT, H. & HULT, R. 1885: Vegetationen och

floran i en del av Kemi Lappmark och norra

Österbotten. – Meddelanden af Societas pro

Fauna et Flora Fennica 12: 1-159.

HOCH, P. C., LIEVENS, A. W., RAVEN, P. H. &

JARVIS, C. E. 1995: Proposal to reject the name

Epilobium alpinum L. (Onagraceae). – Taxon 44:

237-239.

HOLMGREN, P. K., HOLMGREN, N. H. &

BARNETT, L. C. 1990: Index Herbariorum. Part

1: The herbaria of the World. Ed. 8. – Bronx,

New York. 693 pp.

HULT, R. 1898: Växtgeografiska anteckningar från

den finska Lappmarkens skogsregioner. – Acta

Societatis pro Fauna et Flora Fennica 16(2): 1-

200.

HULTÉN, E. 1948: Flora of Alaska and Yukon 8. –

Lunds Universitets Årsskrift N.F. 2, 44(1): 1201-

1341.

118 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

HULTÉN, E. 1958: The amphi-atlantic plants and

their phytogeographical connections. – Kungliga

Svenska Vetenskapsakademiens Handlingar 4,

7(1): 1-340 + 279 maps.

HULTÉN, E. 1962: The circumpolar plants I. –

Kungliga Svenska Vetenskapsakademiens

Handlingar 4, 8(5): 1-275.

HULTÉN, E. 1968: Flora of Alaska and neighboring

territories. A manual of the vascular plants. –

California. 1008 pp.

HULTÉN, E. 1971a: Atlas of the distribution of

vascular plants in northwestern Europe. Ed. 2. –

Stockholm. 56 + 531 pp.

HULTÉN, E. 1971b: The circumpolar plants. II.

Dicotyledons. – Kungliga Svenska

Vetenskapsakademiens Handlingar 13(1): 1-463.

HULTÉN, E. & FRIES, M. 1986: Atlas of North

European vascular plants north of the Tropic of

Cancer. I, II, III. – Koeltz Scientific Books.

Königstein. 1172 pp.

HULTGÅRD, U-M. 1990: Polyploidy and

differentiation in N European populations of

Primula subgenus Aleuritia. – Sommerfeltia 11:

117-135.

HULTGÅRD, U-M. 1993: Primula scandinavica and

P. stricta – patterns of distribution, variation,

reproduction strategies and migrations. – Opera

Botanica 121: 35-43.

HUSTICH, I. 1936a: Botaniska notiser från västra

Lappland. 1. Växtlokaler från skogsregionen. –

Memoranda Societatis pro Fauna et Flora

Fennica 11: 154-161.

HUSTICH, I. 1936b: Botaniska notiser från västra

Lappland. 2. Floran kring en timmerkoja i Kittilä

Lappmark. – Memoranda Societatis pro Fauna et

Flora Fennica 11: 162-165.

HUSTICH, I. 1936c: Botaniska notiser från västra

Lappland. 3. Några för alpina regionen på Pallas-

och Ounastunturi nya kärlväxter. - Memoranda

Societatis pro Fauna et Flora Fennica 11: 166-

170.

HUSTICH, I. 1937a: Pflanzengeographische Studien

im Gebiet der niederen Fjelde im westlichen

finnischen Lappland. I. Über die Beziehung der

Flora zu Standort und Höhenlage in der alpinen

Region sowie über das Problem "Fjeldpflanzen in

der Nadelwaldregion". – Acta Botanica Fennica

19: 1-156.

HUSTICH, I. 1940c: Pflanzengeographische Studien

im Gebiet der niederen Fjelden im westlichen

finnischen Lappland. II. Über die horizontale

Verbreitung der alpinen und alpiken Arten sowie

einige Angaben über die winterlichen

Naturverhältnisse auf den Fjelden. – Acta

Botanica Fennica 27: 1-80.

HUSTICH, I. 1942a: Några växtgeografiska

anteckningar under en juliresa 1940 i Utsjoki,

norra Lappland. – Memoranda Societatis pro

Fauna et Flora Fennica 17: 215-226.

HUSTICH, I. 1970: On the phytogeography of the

eastern part of central Quebec-Labrador

Peninsula. II. – Commentationes Biologicae

Societas Scientiarum Fennica 30(7): 1-16.

HYLANDER, N. 1945: Nomenklatorische und

systematische Studien über nordische

Gefässpflanzen. – Uppsala Universitets Årsskrift

1945(7): 1-337.

ISO-IIVARI, L. 1977: Topografikartan 1:20000

nimistö Inarin Lapissa aakkosjärjestyksessä. –

Lapin tutkimuslaitos Kevo. 96 pp.

ISOTALO, A. 1971: Porojen luonnonvaraisten

rehukasvien ravintoarvoista. – Lapin

tutkimusseuran vuosikirja 12: 28-45.

JALAS, J. 1949: Floristisches aus Lapponia

enontekiensis, Lapponia tornensis und Troms

Fylke. – Archivum Societatis Zoologicae

Botanicae Fennicae Vanamo 2: 90-96.

JALAS, J. 1950: Zur Kausalanalyse der Verbreitung

einiger nordischen Os- und Sandpflanzen. –

Annales Botanici Societatis Zoologicae

Botanicae Fennicae Vanamo 24(1): 1-362.

JARVIS, C. 2007: Order out of the chaos. Linnaean

plant names and their types. – The Linnaean

Society of London & The Natural History

Museum London. London. 1016 pp.

JONSELL, L. (ed.) 2010: Upplands flora. – SBF-

förlaget. Uppsala. 895 pp.

JONSELL, B. & JARVIS, C. 2002: Lectotypification

of Linnaean names for Flora Nordica

(Brassicaceae – Apiaceae). – Nordic Journal of

Botany 22: 67-86.

JØRGENSEN, C. A., SØRENSEN, T. &

WESTERGAARD, M. 1958: The flowering

plants of Greenland. A taxonomical and

cytological survey. – Biologiske Skrifter utgivet

af Det Kongelige Danske Videnskabernes

Selskap 9(4): 1-172.

KALELA, A. 1939: Über Wiesen und wiesenartige

Pflanzengesellschaften auf der Fischerhalbinsel

in Petsamo Lappland. – Acta Forestalia Fennica

48(2): 1-523.

KALLIO, P. 1949: Eräitä kasvitietoja Kilpisjärven

seudulta ja Tornionjokivarrelta. – Archivum

Societatis Zoologicae Botanicae Fennicae

Vanamo 2: 51-55.

KALLIO, P. 1954: Turun Eläin- ja Kasvitieteellisen

Seuran Lapin retki v. 1954. – Luonnon Tutkija

58(5): 145-151.

KALLIO, P. 1958: Piirteitä Nuortijoen latva-alueen

kasvistosta. – Luonnon Tutkija 62(3): 82-89.

KALLIO, P. 1959a: Eräistä Perä-Lapin

kasvitieteellisen tutkimuksen tehtävistä.

(Summary: Some aspects on botanical

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 119

investigation of northern Lapland.) – Terra 71(4):

167-183.

KALLIO, P. 1959b: Subarktisen Suomen

kasvitieteellisiä kysymyksiä. – Lapin

tutkimusseuran vuosikirja 1: 17-33.

KALLIO, P. 1961: Zur floristisch-ökologischen

Charakteristik des östlichen Teiles von Finnish-

Fjeldlappland. – Archivum Societatis Zoologicae

Botanicae Fennicae Vanamo 16 (suppl.): 98-111.

KALLIO, P. & LEHTONEN, J. 1973: Birch forest

damage caused by Oporinia autumnata (Bkh.) in

1965-66 in Utsjoki, N Finland. – Reports from

the Kevo Subarctic Research Station 10: 55-69.

KALLIO, P. & MÄKINEN, Y. 1957:

Untersuchungen über die Flora von Utsjoki in

Nordfinnland II. – Archivum Societatis

Zoologicae Botanicae Fennicae Vanamo 12(1):

12-29.

KALLIO, P. & MÄKINEN, Y. 1975: Vascular flora

of Inari Lapland. 3. Salicaceae. – Reports from

the Kevo Subarctic Research Station 12: 66-105.

KALLIO, P. & MÄKINEN, Y. 1978a: Vascular flora

of Inari Lapland. 4. Betulaceae. – Reports from

the Kevo Subarctic Research Station 14: 38-63.

KALLIO, P., LAINE, U. & MÄKINEN, Y. 1969:

Vascular flora of Inari Lapland. 1. Introduction

and Lycopodiaceae – Polypodiaceae. – Reports

from the Kevo Subarctic Research Station 5: 1-

108.

KALLIO, P., LAINE, U. & MÄKINEN, Y. 1971:

Vascular flora of Inari Lapland. 2. Pinaceae and

Cupressaceae. – Reports from the Kevo Subarctic

Research Station 8: 73-100.

KALLIO, P., PARVIAINEN, R. & YLIAHO, H.

1978: Väinönputki – perinteinen Lapin vihannes.

– Acta Lapponica Fenniae 10: 96-100.

KALLIOLA, R. 1932: Alpiinisesta kasvillisuudesta

Kammikivialueella Petsamon Lapissa.

Kasvillisuusmonografia. (Referat: Über die

alpine Vegetation im Kammikivi-Gebiet von

Petsamo-Lappland. Eine kleine

Vegetationsmonographie. – Annales Botanici

Societatis Zoologicae-Botanicae Fennicae

Vanamo 2(2): 1-121.

KALLIOLA, R. 1937a: Dryopteris fragrans (L.)

Schott, ein für Europa neuer Farn. – Annales

Botanici Societatis Zoologicae-Botanicae

Fennicae Vanamo 9(4): 1-56.

KALLIOLA, R. 1939: Pflanzensoziologische

Untersuchungen in der alpinen Stufe Finnisch-

Lapplands. – Annales Botanici Societatis

Zoologicae-Botanicae Fennicae Vanamo 13(2):

1-328.

KALLIOLA, R. 1961: Über die Fjeldvegetation. –

Archivum Societatis Zoologicae Botanicae

Fennicae Vanamo 16(suppl.): 113-120.

KÄMÄRÄINEN, H. 1998: Kilpisjärven mahtava

Saana – kasviparatiisi. – Lutukka 14(3): 80-87.

KÄRENLAMPI, L. 1973: Biomass and estimated

yearly net production of the ground vegetation at

Kevo. In: Bliss, L. C. & Wielgolaski, F. E. (eds.):

Proceedings of the Conference on Primary

Production and Production Processes, Tundra

Biome, Dublin, Ireland, April 1973. – Tundra

Biome Steering Committee. Edmonton. Pp. 111-

114.

KÄRENLAMPI, L. & KAUHANEN, H. 1972: A

direct gradient analysis of the vegetation of the

surroundings of the Kevo Subarctic Station. –

Reports from the Kevo Subarctic Research

Station 9: 82-98.

KARI, L. E. 1936: Mikromyceten aus Finnisch-

Lappland. – Annales Botanici Societatis

Zoologicae-Botanicae Fennicae Vanamo 8(3): 1-

25.

KARLSSON, L. 1973: Autecology of cliff and scree

plants in Sarek National Park, northern Sweden.

– Växtekologiska studier 4: 1-203.

KELSO, S. 2009: Primula. In: Flora of North

America Editorial Committee (eds.): Flora of

North America North of Mexico. 8. Oxford

University Press. New York and Oxford. 585 pp.

– Published on the Internet

http://www.efloras.org/florataxon.aspx?flora_id=

1&taxon_id=250092232. – Accessed on

23.11.2014.

KEMPPAINEN, E., KETTUNEN, T., KURTTO, A.,

LAHTI, T. & UOTILA, P. 1991: Ketonukki

(Androsace septentrionalis) Suomessa – historiaa,

biologiaa ja suojelunäkökohtia. – Lutukka 7(4):

103-120.

KIHLMAN, A. O. 1884: Anteckningar om Floran i

Inari Lappmark. – Meddelanden af Societas pro

Fauna et Flora Fennica 11: 45-135.

KLOCKARS, B. & LUTHER, H. 1938: Floristiska

iakttagelser i Li, Viibus-Maarestatunturiområdet.

– Memoranda Societatis pro Fauna et Flora

Fennica 14: 45-54.

KNABEN, G. 1943: Studier over norske Pyrola-arter.

– Bergens Museums årbok 1943,

Naturvitenskapelig rekke 6: 1-18 + 1 pl.

KNABEN, G. 1966: Om kromosomvariasjon og

rasedannelse i den norske flora. – Blyttia 24: 65-

79.

KNABEN, G. & ENGELSKJØN, T. 1968: Studies in

Pyrolaceae, especially in the Pyrola rotundifolia

complex. – Årbok for Universitetet i Bergen.

Matematisk-naturvitenskapelig Serie 4/1967. 71

pp + 5 pl.

KOIVISTO, M. 1936: Saksan pensaskanerva

(Myricaria germanica) Utsjoella. – Luonnon

Ystävä 40(4): 131-132.

120 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

KOIVISTOINEN, E. 1964: Inarin Peldoaivin ja sen

lähitunturien alueen kasvistosta ja

kasvillisuudesta. – M. Sc. Thesis, Department of

Biology, University of Turku. 134 pp.

KONTUNIEMI, T. 1930: Kulttuuritulokkaita

Petsamon maantiellä. – Luonnon Ystävä 34(3):

107-108.

KONTUNIEMI, T. 1932: Metsäkasvien

siemenellisestä lisääntymisestä Petsamon

subalpiinisessa vyöhykkeessä. (Referat: Über die

fruktifikative Vermehrung der Waldpflanzen in

der subalpinen Zone von Petsamo). – Annales

Botanici Societatis Zoologicae-Botanicae

Fennicae Vanamo 2(4): 1-58.

KOTILAINEN, M. J. 1951: Über die Verbreitung der

meso-eutrophen Moorpflanzen in Nordfinnland.

– Annales Academiae Scientiarum Fennicae

Series A IV, 17: 1-162.

KRAVCHENKO, A. V. 2007: Konspekt floryi

Karelii. – Petrozavodsk. 404 pp.

KROK, T. O. B. N. & ALMQUIST, S. 2013: Svensk

flora. Fanerogamer och kärlkryptogamer. 29th ed.

Eds. Jonsell, L. & Jonsell, B. – Liber, Stockholm.

586 pp.

KRON, K. A. & JUDD, W. S. 1990: Phylogenetic

relationships within the Rhodoreae (Ericaceae)

with specific comments on the placement of

Ledum. – Systematic Botany 15(1): 57-68.

KUITUNEN, T. 1984: Läntisen Utsjoen

kasvillisuudesta kesäretkellä 1983. – Talvikki

8(2): 41-47.

KUJALA, M., KYNSILEHTO, K., OHENOJA, E.,

SAASTAMOINEN, O. & SEPPONEN, P. 1987:

Lapin läänin luonnonmarja- ja sienivarat, niiden

satoarvot, hyödyntäminen ja sivuansiollinen

merkitys. – Polar-Marsi 86, Final report. –

Pellervo-Seuran Markkinatutkimuslaitos. 60 pp.

KUJALA, M., MALIN, A., SAASTAMOINEN, O.,

LOHINIVA, S. & NIVA, A. 1989: Pohjois-Lapin

kuntien metsämarja- ja sienitutkimus vuosina

1987-1988 (Pohla-Project). – Pellervo-Seuran

Markkinatutkimuslaitos. 101 pp.

KUJALA, V. 1926: Untersuchungen über den

Einfluss von Waldbränden auf die

Waldvegetation in Nord-Finnland. –

Communicationes ex Instituto Quaestionum

Forestalium Finlandiae 10(5): 1-41 + Tafel I-II.

KUJALA, V. 1929: Untersuchungen über Waldtypen

in Petsamo und in angrenzenden Teilen von

Inari-Lappland. – Communicationes ex Instituto

Quaestionum Forestalium Finlandiae 13(9): 1-

125 + 11 Tables + 1 Map.

KUJALA, V. 1961: Havaintoja Tepastojoen (KemL)

varsien kasvistosta. – Luonnon Tutkija 65(5):

149-152.

KUJALA, V. 1962: Ivalojoen ja sen varsien

kasvistosta. (Zusammenfassung: Über die Flora

des Flusses Ivalojoki und seiner Ufer in Finnisch-

Lappland). – Archivum Societatis Zoologicae

Botanicae Fennicae Vanamo 16(2): 163-193.

KUJALA, V. 1964: Metsä- ja suokasvilajien

levinneisyys- ja yleisyyssuhteista Suomessa.

Vuosina 1951-1953 suoritetun valtakunnan

metsien III linja-arvioinnin tuloksia. –

Communicationes Instituti Forestalis Fenniae

59(1): 1-137 + 196 maps.

KULMALA, H. 1999: Kasvihavaintoja Kietsimä- ja

Repojokien latvoilta Inarista. – Lutukka 15(1):

27-29.

KURASHIGE, Y., ETOH, J-I., HANDA, T.,

TAKAYANAGI, K. & YUKAWA, T. 2001:

Sectional relationships in the genus

Rhododendron (Ericaceae): evidence from matK

and trnK intron sequences. – Plant Systematics

and Evolution 228(1-2): 1-14.

KUTA, E. 1991: Biosystematic studies on Viola sect.

Plagiostigma: III. Biometrical analysis of the

Polish populations of V. epipsila, V. palustris and

their spontaneous hybrids. – Fragmenta Floristica

et Geobotanica 35: 5-34.

KVIST, G. 1978: C. W. Fontells resa till finska och

ryska Lappmarken 1899. – Memoranda Societatis

pro Fauna et Flora Fennica 54: 41-57.

KYTÖVUORI, I. 1969: Epilobium davuricum Fisch.

(Onagraceae) in Eastern Fennoscandia compared

with E. palustre L. A morphological, ecological

and distributional study. – Annales Botanici

Fennici 6(1): 35-58.

KYTÖVUORI, I. 1972: The alpine group of the

genus Epilobium in northernmost Fennoscandia.

A morphological, taxonomical and ecological

study. – Annales Botanici Fennici 9(x): 163-203.

LAANE, M. 1965: Kromosomundersøkelser hos

noen norske plantearter. – Blyttia 23(4): 169-189.

LAANE, M. 1967: Kromosomundersøkelser i Øst-

Finnmarks flora II. – Blyttia 25: 45-54.

LAHTI, T., LAMPINEN, R. & KURTTO, A. 1995:

Suomen putkilokasvien levinneisyyskartasto.

Versio 2.0. (Atlas of Finnish vascular plants.

Version 2.0). – Finnish Museum of Natural

History, Botanical Museum. Helsinki. 23 pp. +

1604 maps as a database.

LAINE, A. 1950: Turun Eläin- ja Kasvitieteellinen

Seura. Vuosikokous 24.2.1950. - Luonnon

Tutkija 54(2): 68.

LAINE, T. 1958: Floristis-ekologisia tutkimuksia

Urtasvaarrin ja sen ympäristötunturien alueella

Enontekiön luoteisosissa. – M. Sc. Thesis,

Department of Biology, University of Turku. 145

pp.

LAINE, U. 1964: Über die floristischen Züge der

nördlichen Waldgrenze der Kiefer im Westteil

von Inari-Lappland. – Reports from the Kevo

Subarctic Research Station 1: 94-123.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 121

LAINE, U. 1965: Alustavia tuloksia Kevojoen

pahtojen kasvistollisesta kartoitustyöstä. – Lapin

Tutkimusseuran vuosikirja 6: 45-58.

LAINE, U. 1970: Kevojoen laakson floora I-II. – Lic.

Phil. Thesis, Department of Biology, University

of Turku. 201 + 258 pp.

LAINE, U. & NURMI, J. 1971: Factors affecting

vegetation and flora of anorthosite and granulite

areas in western Inari, Finnish Lapland. – Reports

from the Kevo Subarctic Research Station 8: 104-

115.

LAINE, U., LEHMUSHOVI, A. & NURMI, J. 1974:

Chromosome numbers of phanerogams in Inari

Lapland and adjacent regions. – Rep. Kevo

Subarctic Res. Stat. 11: 79-89.

LAINE, U., LINDGREN, L. & MÄKINEN, Y. 1955:

Havaintoja Utsjoen pitäjän länsiosan kasvistosta.

– Archivum Societatis Zoologicae Botanicae

Fennicae Vanamo 9(2): 120-135.

LAITINEN, J. & OHENOJA, E. 1990: Kitisen ranta-

ja vesikasvillisuudesta Pelkosenniemen

Suvannossa. – Memoranda Societatis pro Fauna

et Flora Fennica 66: 1-24.

LAMMES, T. 1991: Luoteis-Enontekiön

ylhiötunturialueen kasvistosta – valikoituja

poimintoja. – Lutukka 7(3): 67-80.

LAMPINEN, R. & LAHTI, T. 2007: Kasviatlas

2006. – University of Helsinki, Finnish Museum

of Natural History, Botanical Museum, Helsinki.

Electronic distribution maps available at

http://www.luomus.fi/kasviatlas.

LAMPINEN, R. & LAHTI, T. 2010: Kasviatlas

2009. – University of Helsinki, Finnish Museum

of Natural History, Botanical Museum, Helsinki.

Electronic distribution maps available at

http://www.luomus.fi/kasviatlas.

LAMPINEN, R. & LAHTI, T. 2011: Kasviatlas

2010. – University of Helsinki, Finnish Museum

of Natural History, Botanical Museum, Helsinki.

Electronic distribution maps available at

http://www.luomus.fi/kasviatlas.

LAMPINEN, R. & LAHTI, T. 2013: Kasviatlas

2012. – University of Helsinki, Finnish Museum

of Natural History, Helsinki. Electronic

distribution maps available at

http://www.luomus.fi/kasviatlas.

LAMPINEN, R. & LAHTI, T. 2016: Kasviatlas

2015. – University of Helsinki, Finnish Museum

of Natural History, Helsinki. Electronic

distribution maps available at

http://www.luomus.fi/kasviatlas.

LAMPINEN, R. & LAHTI, T. 2017: Kasviatlas

2016. – University of Helsinki, Finnish Museum

of Natural History, Helsinki. Electronic

distribution maps available at

http://www.luomus.fi/kasviatlas.

LAMPINEN, R. & LAHTI, T. 2018: Kasviatlas

2017. – University of Helsinki, Finnish Museum

of Natural History, Helsinki. Electronic

distribution maps available at

http://www.luomus.fi/kasviatlas.

LAMPINEN, R., LAHTI, T. & HEIKKINEN, M.

2012: Kasviatlas 2011. – University of Helsinki,

Finnish Museum of Natural History, Helsinki.

Electronic distribution maps available at

http://www.luomus.fi/kasviatlas.

LAMPINEN, R., LAHTI, T. & HEIKKINEN, M.

2014: Kasviatlas 2013. – University of Helsinki,

Finnish Museum of Natural History, Helsinki.

Electronic distribution maps available at

http://www.luomus.fi/kasviatlas.

LAMPINEN, R., LAHTI, T. & HEIKKINEN, M.

2015: Kasviatlas 2014. – University of Helsinki,

Finnish Museum of Natural History, Helsinki.

Electronic distribution maps available at

http://www.luomus.fi/kasviatlas.

LARSSON, C. & MARTINSSON, K. 1998:

Jättebalsamin Impatiens glandulifera i Sverige –

invasionsart eller harmlös trädgårdsflykting? –

Svensk Botanisk Tidskrift 92: 329-345.

LEHMUSHOVI, A. 1976: Puolukka –

uutuusmarjakasvi pellolla. – Puutarha-Uutiset

28(51/52): 1070-1071.

LEHMUSHOVI, A. 1977: Puolukan marjontaan

vaikuttavista tekijöistä. – Puutarha 80(2): 122-

123.

LEMPIÄINEN, T. 1981: Pensaskanerva. – In Kallio,

P. & al. (eds.): Kasvien maailma 4: 1521.

Keuruu.

LID, J. 1950: Nye plantefunn 1945-1949. – Blyttia 8:

41-53.

LID, J. & LID, D. T. 2005: Norsk flora. Ed. 7. Ed. R.

Elven. – Oslo. 1230 pp.

LIDBERG, R. & LINDSTRÖM, H. 2010: Medelpads

flora. – SBF-förlaget. Uppsala. 736 pp.

LINDBERG, H. 1958: Växter kända från Norden i

Linnés herbarium. Plantae e serptentrione

cognitae in herbario Linnaei. – Acta Botanica

Fennica 60: 1-133.

LINDÉN, J. 1943: Bidrag till kännedomen om

vegetation och flora inom Enontekis Lappmarks

björk- och fjällregioner. – Acta Societatis pro

Fauna et Flora Fennica 63(1): 1-82.

LINKOLA, K. 1929: Lapin tulokaskasvistosta

kesällä 1925. – Luonnon Ystävä 33(6): 199-210.

LINKOLA, K. 1932: Alueellista lajitilastoa

vesiemme putkilokasveista. – Luonnon Ystävä

36(2): 86-101.

LINKOLA, K. 1933: Regionale Artenstatistik der

Süsswasserflora Finnlands. – Annales Botanici

Societatis Zoologicae-Botanicae Fennicae

Vanamo 3(5): 3-13.

122 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

LÖFGREN, L. 2013: Närkes flora. – SBF-förlaget.

Uppsala. 744 pp.

LÖVE, Á. 1961: Some notes on Myriophyllum

spicatum. – Rhodora 63(749): 139-145.

LÖVE, Á. 1983: Flora of Iceland. – Reykjavik. 403

pp.

LÖVE, Á. & LÖVE, D. 1956: Cytotaxonomical

conspectus of the Icelandic Flora. – Acta Horti

Gotoburgensis 20(4): 65-291.

LÖVE, Á. & LÖVE, D. 1966: Cytotaxonomy of the

alpine vascular plants of Mount Washington. –

University of Colorado Studies, Series in Biology

24: 1-74.

LUMIALA, O. V. 1939: Etwas über das Vorkommen

der Arctostaphylos alpina (L.) Spr. in der Regio

silvatica im Tuntsagebiet (Ks, Salla). – Annales

Botanici Societatis Zoologicae-Botanicae

Fennicae Vanamo 11(3): 1-4.

LUNDE, T. 1962: An investigation into the pH-

amplitude of some mountain plants in the county

of Troms. – Acta Borealia. A. Sci. 20: 1-105.

LUNDMAN, B. 1948: Något om den regionala

variationen av blomfärgen hos

midsommarblomster (Geranium silvaticum L.). –

Svensk Botanisk Tidskrift 42: 153-157.

LUNDQVIST, J. 1968: Plant cover and environment

of steep hillsides in Pite Lappmark. – Acta

Phytogeographica Suecica 53: 1-153 + 31 Tables.

MÄKINEN, Y. 1964a: Floristic observations in

Finnmark (northern Norway). – Reports from the

Kevo Subarctic Research Station 1: 124-128.

MÄKINEN, Y. 1964b: On Finnish micromycetes 3.

Uredinales of Inari Lapland. – Reports from the

Kevo Subarctic Research Station 1: 155-177.

MÄKINEN, Y. 1964c: On Finnish micromycetes 4.

On the distribution of rusts in Finland. – Annales

Botanici Fennici 1: 214-219.

MÄKINEN, Y. 1969: On Finnish micromycetes 8.

Erysiphales of Inari Lapland. – Reports from the

Kevo Subarctic Research Station 5: 109-116.

MÄKINEN, Y. 2002: Floristic observations in

western Kola Peninsula, NW Russia. – Kevo

Notes 12: 1-33.

MÄKINEN, Y. & KALLIO, P. 1978: Floristic

mapping of the Schefferville area. – In: Müller-

Wille, L. (ed.): Environmental and socio-

economic comparisons of subarctic regions

(Fenno-Scandia and Ungava-Labrador). Pp. 22-

25.

MÄKINEN, Y. & KALLIO, P. 1979: The vascular

plants of Inari Lapland, Finland. – Kevo Notes 4:

1-45.

MÄKINEN, Y., KALLIO, P., LAINE, U. & NURMI,

J. 1982: Vascular flora of Inari Lapland. 5.

Urticaceae – Caryophyllaceae. – Reports from

the Kevo Subarctic Research Station 18: 10-94.

MÄKINEN, Y., KALLIO, P., LAINE, U. & NURMI,

J. 1998: Vascular flora of Inari Lapland. 6.

Nymphaeaceae – Papaveraceae. – Reports from

the Kevo Subarctic Research Station 22: 25-86.

MÄKINEN, Y., LAINE, U., HEINO, S. & NURMI,

J. 2005: Vascular flora of Inari Lapland. 7.

Brassicaceae – Grossulariaceae. – Reports from

the Kevo Subarctic Research Station 23: 1-95.

MÄKINEN, Y., LAINE, U., HEINO, S., ISO-

IIVARI, L. & NURMI, J. 2011: Vascular flora of

Inari Lapland. 8. Rosaceae and Fabaceae. –

Reports from the Kevo Subarctic Research

Station 24: 3-126.

MALYŠEV, L. I. 1976: Materialy k poznaniju

osobennostei sostava i genezisa gornyh

subarktitseskih flor Sibiri. – In: Malyšev, L.I.

(ed.): Flora Putorana. Nauka, Novosibirsk. 243

pp.

MANSIKKANIEMI, H. 1964: Main features of the

glacial and postglacial development of Pulmanki

valley in northernmost Finland. – Reports from

the Kevo Subarctic Research Station 1: 322-337.

MARCUSSEN, T. 2007: Lifiol er død – leve engfiol!

En kritisk morfologisk gjennomgang av

komplekset i Norge. – Blyttia 65: 195-207.

MARKLUND, G. 1940: Beobachtungen über

Empetrum hermaphroditum (Lange) Hagerup und

E. nigrum L. s. str. – Memoranda Societatis pro

Fauna et Flora Fennica 16: 74-77.

MARISTO, L. 1941: Die Seetypen Finnlands auf

floristischer und Vegetationsphysiognomischer

Grundlage. – Annales Botanici Societatis

Zoologicae-Botanicae Fennicae Vanamo 15(5):

1-314 + Table.

MEHUS, H. 1969: Kvann – en gammel nytteplante. –

Ottar 59: 3-8.

MELA, A. J. & CAJANDER, A. K. 1906: Suomen

kasvio. 5. painos. – Suomalaisen kirjallisuuden

seuran toimituksia 53: 1-763.

MEUSEL, H., JÄGER, E., RAUSCHERT, S. &

WEINERT, E. 1978: Vergleichende Chorologie

der Zentraleuropäischen Flora. 2. Karten. –

Fischer. Jena. Pp. 259-421.

MIKKOLA, E. 1941: Manuscript concerning the

flora of Inari Lapland and Saariselkä. – Archives

of the Kevo Subarctic Research Institute. 41 pp.

MOBERG, A. 1885: Fenologiska anteckningar.

Klimatologiska iakttagelser i Finland. Andra

delen. År 1856-1875. – Bidrag till kännedom af

Finlands Natur och Folk 41: 1-318.

MOISIO, S., MÄKINEN, Y., TUOMINEN, M. &

VAURAS, J. 2006: Luonnonyrttiopas. –

Opetushallitus. 68 pp.

MONTELL, J. 1910: Några anmärkningsvärda

kärlväxter från Muonio och Enontekis. –

Meddelanden af Societas pro Fauna et Flora

Fennica 36: 152-157.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 123

MONTELL, J. 1945a: Bidrag till kännedomen om

adventiv- och ogräsfloran i Muonio socken,

Lapponia kemensis. – Memoranda Societatis pro

Fauna et Flora Fennica 20: 82-91.

MONTELL, J. 1948: Några anmärkningsvärda

växtfynd, de flesta från södra Enontekis. –

Memoranda Societatis pro Fauna et Flora

Fennica 24: 178-183.

MONTELL, J. 1962: Vegetationen och floran i

Muonio socken. (Zusammenfassung:Über die

Vegetation und Flora im Kirchspiele Muonio in

Nordfinnland). – Memoranda Societatis pro

Fauna et Flora Fennica 37: 70-130.

MOSQUIN, T. 1966: A new taxonomy for

Epilobium angustifolium L. (Onagraceae). –

Brittonia 18(2): 167-188.

MOSSBERG, B. & STENBERG, L. 2003: Den nya

nordiska floran. – Stockholm. 928 pp.

NANNFELDT, J. A. 1981: Exobasidium, a

taxonomic reassessment applied to the European

species. – Symbolae Botanicae Upsalienses

23(2): 1-72.

NAUENBURG, J. D. & BUTTLER, K. P. 2007:

Validierung des Namens Viola wittrockiana. –

Kochia 2: 37-41.

NESOM, G. L. 2009: Diapensiaceae (Diapensia,

Galax, Pyxidanthera, Shortia). – In: Flora of

North America Editorial Committee (eds.): Flora

of North America North of Mexico. Vol. 8.

Oxford University Press, New York and Oxford.

585 pp. – Published on the Internet

http://www.efloras.org/florataxon.aspx?flora_id=

1&taxon_id=250092268. – Accessed on

17.4.2015.

NICKUL, K. 1970: Saamelaiset kansana ja

kansalaisina. – Suomalaisen Kirjallisuuden

Seura. Helsinki. 304 pp.

NIKITIN, V. V. 1988: On the typification of Viola

montana (Violaceae). – Botanicheskii Zhurnal

SSSR 73(11): 1536-1542.

NIKITIN, V. V. 1995: Critical notes on taxonomy

and nomenclature of some European species of

the section Trigocarpea of the genus Viola

(Violaceae). – Botanicheskii Zhurnal SSSR

80(7): 84-96.

NILSSON, Ö. 2000: Nordisk fjällflora. 5th ed. –

Stockholm. 272 pp.

NORDHAGEN, R. 1936b: Skandinavias fjellflora og

dens relasjoner til den siste istid. – Nord. (19.

Skand.) Naturforskarmöte i Helsingfors 1936:

93-124.

NORDHAGEN, R. 1964: Om Anemone nemorosa L.

ved Vadsø og andre norske plantefunn gjort av

avdøde sorenskriver Jakob Bredrup. – Blyttia

22(2): 73-77.

NORDLING, X. W. 1884a: Berättelse om

odlingsförsök verkställda i Inari Lappmark år

1874 – 1883. – Kungliga Finska Hushålls

Sällskaps Handlingar 1878-1883 (2), bil. KK:

303-309.

NORDLING, X. W. 1884b: Berättelse om jordbruket

å Toivoniemi modellfarm i Inari Lappmark år

1883. – Kungliga Finska Hushålls Sällskaps

Handlingar 1878-1883 (2), bil. LL: 310-317.

NORMAN, J. M. 1894-1901: Norges arktiske flora.

I(1-2), II(1-2). – Kristiania. 1487 + 623 pp.

NOUSIAINEN, H., TERÄS, I. & VIRAMO, J. 1978.

Mustikka ja puolukka – hyönteispölytteiset

metsämarjamme. – Suomen Luonto 37(2): 91-94.

NUOTIO, H. 1950: Pensaskanerva (Myricaria

germanica) Enontekiön Lapista. – Luonnon

Tutkija 54(3): 97.

NURMI, J. 1987: Suomen malvakasvit. – Lutukka

3(3): 67-75.

NYMAN, A. 1964: Aquatic vegetation of Lake

Mantojärvi in Inari Lapland, Finland. – Reports

from the Kevo Subarctic Research Station 1: 60-

68.

OHWI, J. 1965: Flora of Japan. – Washington, D.C.

1067 pp.

OJALA, A. 1984: Variation of Angelica archangelica

subsp. archangelica (Apiaceae) in northern

Fennoscandia. 1. Variation in fruit morphology. –

Annales Botanici Fennici 21: 103-115.

OJALA, A. 1986a: Variation of Angelica

archangelica subsp. archangelica (Apiaceae) in

northern Fennoscandia. 3. Interpopulational

variation in reproductive and life-history

characters. – Annales Botanici Fennici 23: 11-21.

OJALA, A. 1986b: Variation of Angelica

archangelica subsp. archangelica (Apiaceae) in

northern Fennoscandia. 4. Pattern of geographic

variation. – Annales Botanici Fennici 23: 23-31.

OKSANEN, L. & VIRTANEN, R. 1995:

Topographic, altitudinal and regional patterns in

continental and suboceanic heath vegetation of

northern Fennoscandia. – Acta Botanica Fennica

153: 1-80.

PAAL, T. & PAAL, J. 1989: Struktura

tsenopopuljatsii brusniki Vaccinium vitis-idaea l:

– Akademija nauk Estonskoi SSR. Tallinn

'Valgus'. 212 pp.

PAATELA, J. 1953: Maamme heinänurmien

botaanisesta koostumuksesta. (Summary: On the

botanical composition of the tame-hayfields in

Finland.) – Acta Agralia Fennica 79(3): 1-128.

PARNELA, A. 1985: Henkiinheränneitä saksalaisia

sotatulokkaita Hyrynsalmen Kangasjärvellä. (The

revival of some German polemochores at

Kangasjärvi, Hyrynsalmi, E Finland). – Lutukka

1(2): 56-59.

PARVELA, A. A. 1930: Oulun läänin viljelyskasvit.

Niiden historia ja nykyinen levinneisyys. I.

124 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Yleinen osa. – Annales Societatis Zoologicae-

Botanicae Fennicae Vanamo 13(1): 1-354.

PARVELA, A. A. 1931: Petsamon alueen

viljelyskasvien historia ja levinneisyys. – Acta

Agralia Fennica 23(2): 33-98.

PARVELA, A. A. 1932: Oulun läänin viljelyskasvit.

Niiden historia ja nykyinen levinneisyys. II.

Erikoisosa. (Referat: Über die Kulturpflanzen im

Län Oulu (Uleåborg), ihre Geschichte und

Verbreitung. II. Spezieller Teil.) – Annales

Botanici Societatis Zoologicae-Botanicae

Fennicae Vanamo 2(5): 1-144.

PERTOLA, E. 1961: Züge aus der Flora und

Vegetation im Ostteil des Saariselkä-Massives im

finnischen Ostlappland. – Annales Universtitatis

Turkuensis Ser. A II, 27: 1-38.

PESOLA, V. 1918: Huomattavia kasvilöytöjä N-

Kuusamosta ja Kuolajärveltä. – Meddelanden af

Societas pro Fauna et Flora Fennica 44: 229-246.

PESOLA, V. 1928: Kalsiumkarbonaatti

kasvimaantieteellisenä tekijänä Suomessa.

(Summary: Calcium carbonate as a factor in the

distribution of plants in Finland.) – Annales

Societatis Zoologicae-Botanicae Fennicae

Vanamo 9(1): 1-246 + Appendix 1-14.

PESOLA, V. 1952b: Havaintoja Kuusamon ja Sallan

pitäjien (Ks) vesikasvillisuudesta. (Summary:

Notes on the aquatic vegetation in the communes

of Kuusamo and Salla, NE Finland.) – Archivum

Societatis Zoologicae Botanicae Fennicae

Vanamo 6(2): 102-104.

PIIPPO, S. 2004: Luonnon lääkeyrtit 2. – Tammi.

Helsinki. 310 pp.

PIIPPO, S. 2005a: Luonnon lääkeyrtit 3. – Tammi.

Helsinki. 277 pp.

PIIPPO, S. 2005b: Luonnon lääkeyrtit 4. – Tammi.

Helsinki. 328 pp.

PIIRAINEN, M. 1996a: Itä-Enontekiön

kaatopaikkakasveista. – Lutukka 12(2): 62-63.

PIIRAINEN, M. 1996b: Termislehto – keidas

keskellä karua Enontekiön Maanselkää. –

Lutukka 12(3): 67-72.

PIIRAINEN, M. 1997d: Paatsjoen laakson

kulttuurikasvistoa tutkimassa. –

Luonnontieteellinen keskusmuseo, vuosikirja

1997: 33-40.

PIIRAINEN, M. 1997e: Centaurea scabiosa och

några andra växtfynd vid Grense Jakobselv, Sør-

Varanger. – Polarflokken 21: 279-282.

PIIRAINEN, M. & PIIRAINEN, P. 1991b:

Enontekiön Ropin kasveista ja kasvillisuudesta. –

Lutukka 7(3): 87-96.

PIIRAINEN, M., OFTEN, A. & ALM, T. 1997:

Sammakonleinikki ja muita jätelietekuoppien

kasveja Petsamossa 1995–1996. – Lutukka 13(2):

51-54.

PIIRAINEN, M., KRAVCHENKO, A. V. &

UOTILA, P. 2005: O nahodke Diapensia

lapponica (Diapensiaceae) v Respublike Kareliâ.

– Botaničeskij Žurnal 90: 63-66.

PIISPALA, E. 1964: Epilobium adenocaulon

Hausskn. and E. rubescens Rydb. in

Ostfennoskandien. – Annales Botanici Fennici 1:

36-46.

POBEDIMOVA, E. G., STANITSHEVA, O. N. &

DROZDOVA, I. N. 1959: O rasteniyah,

sobrannyh v 1956 na poberezhyah Barentseva i

Belogo Morei. – Bot. Mat. Gerb. Bot. Inst.

Komarova Akad. Nauk SSSR 19: 572-594.

PORSILD, A. E. 1964: Illustrated flora of the

Canadian Arctic Archipelago. 2nd ed. – National

Museum of Canada, Bulletin 146: 1-218.

PUIKKO, M. 1992: Inarin luonnonmarjojen ja -

sienten jalostaminen ja koemarkkinointi. –

Maatilahallitus. 35 pp.

RAHKONEN, R. 1968: Lemmenjoen

kansallispuiston puronvarsikasvistosta. – B. Sc.

Thesis, Department of Biology, University of

Turku. 33 pp.

RAINIO, A. J. 1926: Uredinae Lapponicae. –

Annales Societatis Zoologicae-Botanicae

Fennicae Vanamo 3: 239-267.

RAMENSKAYA, M. L. 1983: Analyz flory

Murmanskoj oblasti i Karelii. – Nauka.

Leningrad. 216 pp.

RAMENSKAYA, M. L. & ANDREEVA, V. N.

1982: Opredelitel' vysshikh rastenij Murmanskoj

Oblasti i Karelii. – Nauka. Leningrad. 435 pp.

RAMULA, S. & MUTIKAINEN, P. 2003: Sex

allocation of females and hermaphrodites in the

gynodioecious Geranium sylvaticum. – Annals of

Botany 92: 207-213.

RAMULA, S., TOIVONEN, E. & MUTIKAINEN,

P. 2007: Demographic consequences of pollen

limitation and inbreeding depression in a

gynodioecious herb. – International Journal of

Plant Sciences 168: 443-453.

RASSI, P., HYVÄRINEN, E., JUSLÉN, A. &

MANNERKOSKI, I. (eds.) 2010: The 2010 Red

List of Finnish Species. – Ympäristöministeriö &

Suomen ympäristökeskus, Helsinki. 685 pp.

RAUHALA, A. 1959: Enumeratio Uredinearum

Fennicarum et distributio hucusquae cognita

earum in provinciis phytogeographicis

Fennoscandiae orientalis. – Kuopion Luonnon

Ystäväin Yhdistys Julkaisuja Sarja B 3(3): 1-181.

RAUTAVA, E. 1964: Über die Wasservegetation des

Flusses Vaskojoki im nördlichsten Finnland. –

Reports from the Kevo Subarctic Research

Station 1: 69-93.

RAUTAVA, E. 1969: Tutkimuksia Vaskojoen

vesikasvillisuudesta I. – Phil. Lic. Thesis,

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 125

Department of Biology, University of Turku. 198

+ 2 pp.

RAUTAVA, E. 1971: Tutkimuksia Vaskojoen

vesikasvillisuudesta II. – Phil. Lic. Thesis,

Department of Biology, University of Turku. 122

+ 30 pp.

RAUTIAINEN, V-P. 1991: Pensaskanervan

(Myricaria germanica) esiintyminen Utsjoella

vuonna 1991. – Manuscript, Botanical Museum,

University of Turku. 22 pp.

RAUTIAINEN, V-P. 1996: Pensaskanerva – Koillis-

Utsjoen jokivarsien erikoisuus. –

Toimintakertomus 1996. Botanical Museum,

University of Turku. Pp. 17-20.

RAUTIAINEN, V-P., RYTTÄRI, T., KURTTO, A.

& VÄRE, H. (eds.) 2002: Putkilokasvien

uhanalaisuuden arviointi – lajikohtaiset

perustelut. – Suomen ympäristö 593: 1-194.

REIERSEN, J. & SORTLAND, A. 1991:

Amerikamjølke (Epilobium adenocaulon) funnet

i Nord-Norge. – Polarflokken 15(2): 147-148.

RICKETT, H. W. 1969: Wild flowers of the United

States III(1). – New York. 274 pp.

RINTANEN, T. 1962: Havaintoja Tuntsajoen latva-

alueen kasvistosta. – Luonnon Tutkija 66(2): 48-

51.

RINTANEN, T. 1967: On the distribution of some

boreal plants in Eastern Lapland. – Aquilo, Ser.

Botanica 6: 197-208.

RINTANEN, T. 1968: The distribution of fjeld plants

in eastern Lapland. – Annales Botanici Fennici

5(4): 225-305.

RINTANEN, T. 1970a: On the vegetation and

ecology of frost ground sites in eastern Finnish

Lapland. – Annales Botanici Fennici 7(1): 1-24.

RINTANEN, T. 1970b: Floristic areas distinguished

in eastern Finnish Lapland on the basis of fjeld

plant distribution. – Annales Botanici Fennici

7(4): 353-374.

RINTANEN, T. 1976: Lake studies in eastern

Finnish Lapland. I. Aquatic flora: Phanerogams

and Charales. – Annales Botanici Fennici 13(3):

137-148.

RINTANEN, T. 1982a: Botanical lake types in

Finnish Lapland. – Annales Botanici Fennici

19(4): 247-274.

RINTANEN, T. 1982b: Järvitutkimusaineisto Lapin

läänistä. – Manuscript, Floristic Archives,

Botanical Museum, University of Helsinki.

ROGER, J. G. 1952: Diapensia lapponica L. in

Scotland. – Transactions of the Botanical Society

of Edinburgh 36 (1): 34-36.

ROIVAINEN, H. 1923: Tietoja kasvillisuudesta sekä

putkilo- ja lehtisammalkasvistosta keskisen

Luttojoen seuduilla. (Referat: Beobachtungen

über die Vegetation und die Gefässpflanzen- und

Laubmoosflora in der Gegend des Flusses

Luttojoki in Finnisch-Lappland.) – Annales

Societatis Zoologicae-Botanicae Fennicae

Vanamo 1(8): 229-304.

RØNNING, O. I. 1954: Some new plant finds from

Arctic Norway. – Acta Borealia, A. Scientia 7: 4-

10.

ROSBERG, J. 1891: Nordöstra Sodankylä. –

Geografiska Förenings Tidskrift 3: 1-59.

ROWECK, H. 1981: Die Gefässpflanzen von

Schwedisch-Lappland. – Flora et Vegetatio

Mundi, VIII. Vaduz. 804 pp.

RUNE, O. 1963: Lapplands flora. – In: Curry-

Lindahl, K. (ed.): Natur i Lappland. Uppsala. Pp.

211-250.

RUUHIJÄRVI, R. 1960: Über die regionale

Einteilung der nordfinnischen Moore. – Annales

Botanici Societatis Zoologicae Botanicae

Fennicae Vanamo 31(1): 1-360.

RUUHIJÄRVI, R. 1974: Soiden karpalosadoista. –

Suo 25: 25-30.

RYDBERG, H. & WANNTORP, H.-E. 2001:

Sörmlands flora. – Botaniska Sällskapet i

Stockholm. Stockholm. 776 pp.

RYTTÄRI, T. & KETTUNEN, T. (eds.) 1997:

Uhanalaiset kasvimme. – Suomen

Ympäristökeskus. Helsinki. 335 pp.

RYTTÄRI, T., KALLIOVIRTA, M. & LAMPINEN,

R. (eds.) 2012: Suomen uhanalaiset kasvit. –

Tammi. Helsinki. 384 pp.

RYVARDEN, L. 1967: Bidrag til Finnmarks flora

III. – Blyttia 25: 55-60.

RYVARDEN, L. 1969: The vascular plants of the

Rastigaissa area (Finnmark, Northern Norway). –

Acta Borealia A, Scientia 26: 1-56.

SAARENMAA, H., KAHANPÄÄ, J., LAMPINEN,

R., LAHTI, T., HEIKKINEN, M., KOVANEN,

J., HÄKLI, P. & PUUPPONEN, J. 2008:

Luonnontieteellisten havaintojen sijainnin

ilmoittaminen EUREF-FIN-koordinaatistossa.

Luonnontieteellisen keskusmuseon suositus. –

Luonnon Tutkija 112: 144-150.

SALEMAA, M. 2000: Calluna vulgaris. Kanerva. –

In: Reinikainen, A., Mäkipää, R., Vanha-

Majamaa, I. & Hotakainen, J-P.: Kasvit

muuttuvassa metsäluonnossa. Tammi, Helsinki.

Pp. 109-111.

SALEMAA, M., VANHA-MAJAMAA, I. &

GARDNER, P. J. 1999: Compensatory growth of

two clonal dwarf shrubs, Arctostaphylos uva-ursi

and Vaccinium uliginosum in a heavy metal

polluted environment. – Plant Ecology 141: 79-

91.

SALONEN, J. 1954: Vesikasvihavaintoja

Kuusamosta (Ks) ja Kittilästä (KmL). – Luonnon

Tutkija 58(5): 156-157.

SALONEN, J. 1956: Über das Vorkommen der

Hydrophyten in den Stratiotes-Seen in Kittilä,

126 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Finnisch-Lappland. – Archivum Societatis

Zoologicae Botanicae Fennicae Vanamo 10(2):

146-152.

SÅLTIN, H. 1958: Havaintoja Inarin pitäjän

koillisosan kasvistosta. – Manuscript in the

archives of the Kevo Subarctic Research

Institute, University of Turku. 15 pp.

SAMUELSSON, G. 1934: Die Verbreitung der

höheren Wasserpflanzen in Nordeuropa

(Fennoskandien und Dänemark). – Acta

Phytogeographica Suecica 6: 1-211.

SAVILE, D. B. O. 1969: Interrelationships of Ledum

species and their rust parasites in Western

Canada and Alaska. – Canadian Journal of

Botany 47: 1085-1100.

SELANDER, S. 1950b: Floristic phytogeography of

South-Western Lule Lappmark (Swedish

Lapland). II. Kärlväxtfloran i sydvästra Lule

Lappmark. – Acta Phytogeographica Suecica 28:

1-152 + 488 maps.

SILTANEN, P. 1964: The aquatic flora and

vegetation of Lake Kevojärvi. – Reports from the

Kevo Subarctic Research Station 1: 41-59.

SILTANEN, P. 1967: Inarin Lapin vesikasvien

ekologiasta. – M. Sc. Thesis, Department of

Biology, University of Turku. 229 pp. +

Appendix.

SKK III = Suuri Kasvikirja III. 1980 (ed. J. Jalas). –

Otava. Helsinki. 944 pp.

SOKOLOV, D. D. & FILIN, V. R. 1996: Opredelitel'

sosudistyh rastenij okrestnostej Belomorskoj

biologitsheskoj stantsii Moskovskogo

universiteta. – Fizitsheskij fakul'tet Moskovskogo

universiteta. Moskva. 170 pp.

SOKOLOVSKAYA, A. P. & STRELKOVA, O. S.

1960: Geograficheskoe rasprostranenie

poliploidnykh vidov rastenii v Evraziatskoi

Arktike. – Botanicheskii Zhurnal SSSR 45: 369-

381.

SONESSON, M. 1970: Studies on mire vegetation in

the Torneträsk area, Northern Sweden. III.

Communities of the poor mires. – Opera

Botanica 26: 1-120.

SORSA, M. 1965: Hybridization of Palustres violets

in Finland. – Annales Academiae Scientiarum

Fennicae A IV, 86: 1-18.

SORTLAND, A. 1997: Veikantfloraen på Tromsøya,

august 1997. – Polarflokken 21(2): 145-158.

SÖYRINKI, N. 1939a: Studien über die generative

und vegetative Vermehrung der Samenpflanzen

in der alpinen Vegetation Petsamo-Lapplands. II.

Spezieller Teil. – Annales Botanici Societatis

Zoologicae-Botanicae Fennicae Vanamo 14(1):

1-406 + 1 Karte.

SÖYRINKI, N. 1939b: Beobachtungen über die

Gefässkryptogamenflora der Petsamofjelde

(Lapponia petsamoensis). – Annales Botanici

Societatis Zoologicae-Botanicae Fennicae

Vanamo 11(3): 26-35.

SÖYRINKI, N. 1956: Kasvistosta Oulankajoen –

Pääjärven alueella Kieretin Karjalassa. (Referat:

Über die Flora im Gebiet von Oulankajoki-

Pääjärvi, Karelia Keretina, Ostkarelia.) – Annales

Botanici Societatis Zoologicae Botanicae

Fennicae Vanamo 27(2): 1-118.

SÖYRINKI, N. & SAARI, V. 1980: Die Flora im

Nationalpark Oulanka, Nord-Finnland. – Acta

Botanica Fennica 114: 1-149.

SÖYRINKI, N., SALMELA, R. & SUVANTO, J.

1977: Oulangan kansallispuiston metsä- ja

suokasvillisuus. (Summary: The forest and mire

vegetation of the Oulanka National Park,

northern Finland.) – Acta Forestalia Fennica 154:

1-150 + vegetation map.

STENBERG, L. 2010: Norrbottens flora II. – SBF-

förlaget. Uppsala. 792 pp.

SUOMALAINEN, E. W. 1913: Chamaeorchis alpina

(L.) Rich. i Kilpisjärvi-trakten, Le. –

Meddelanden af Societas pro Fauna et Flora

Fennica 39: 38.

SUOMINEN, J. 1969b: The vegetation of railway

yards and adjacent storage areas in Finland. –

Annales Botanici Fennici 6(4): 353-367.

SUOMINEN, J. 1975: Kasvipeitteestä saamelaisten

muinaisilla talvikylän paikoilla. – Luonnon

Tutkija 79: 92-94.

SUOMINEN, J. 1979: The grain immigrant flora of

Finland. – Acta Botanica Fennica 111: 1-108.

SYLVÉN, N. 1906: Om de svenska dikotyledonernas

första förstärkningsstadium. I. – Kungliga

Svenska Vetenskapsakademiens Handlingar B

40(2): 1-348.

TAMMILEHTO, V. 1991: Tornion

Hirsikankaanmäen saksalaiskasvien vaiheita. –

Lutukka 7(2): 35-39.

TASKINEN, J. & NYKÄNEN, L. 1975: Chemical

composition of Angelica root oil. – Acta chemica

Scandinavica B 29(7): 757-764.

THIERS, B. 2015 [continuously updated]: Index

Herbariorum: A global directory of public

herbaria and associated staff. New York

Botanical Garden's Virtual Herbarium. –

Published on the Internet

http://sweetgum.nybg.org/ih/. – Accessed on

5.3.2015.

THUNMARK, S. 1931: Der See Fiolen und seine

Vegetation. – Acta Phytogeographica Suecica 2:

1-198.

TOLMATCHEV, A. I. 1980: Arkticheskaya flora

SSSR. 8. Geraniaceae-Scrophulariaceae. –

Leningrad. 336 pp.

TUHKANEN, S. 1980: Climatic parameters and

indices in plant geography. – Acta

Phytogeographica Suecica 67: 1-110.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 127

TUOMIKOSKI, R. 1950: Huomioita putkilokasveista

Inarin Muddusjärveltä 28.-31.VII.1950. –

Manuscript in the archives of the Kevo Subarctic

Research Institute, University of Turku. 4 pp.

TYLER, T., OLSSON, K-A., JOHANSSON, H. &

SONESSON, M. (eds.) 2007: Floran i Skåne.

Arterna och deras utbredning. – Lunds Botaniska

Förening. Lund. 778 pp.

ULVINEN, T. 1962: Savukosken selkosilta. –

Luonnon Tutkija 66(4): 107-117.

ULVINEN, T. 1985: Hietaorvokki (Viola rupestris)

Pelkosenniemen Kalkkivaaralla – ikivanhan

virheen oikaisu. – Lutukka 1(2): 40-42.

ULVINEN, T. 1996: Tornion Kalkkimaan

saksalaiskasveista ja vähän muistakin. – Lutukka

12(4): 99-109.

ULVINEN, T. & VARKKI, A. 1998: Uusia tietoja

Pohjois-Suomen kasvistosta 1.

Alkuperäiskasveja. – Lutukka 14(4): 99-114.

UOTILA, P. 1974: Elatine hydropiper L. aggr. in

northern Europe. – Memoranda Societatis pro

Fauna et Flora Fennica 50: 113-123.

UOTILA, P. 1987: Alkukesän 1986 kukkijoita

Kilpisjärvellä. – In: Kurtto, A. (ed.):

Kasvihavaintoja. – Lutukka 3(2): 62-63.

VAARAMA, A. & JÄÄSKELÄINEN, O. 1967:

Studies on gynodioecism in the Finnish

populations of Geranium silvaticum L. – Annales

Academiae Scientiarum Fennicae A. IV, 108: 1-

39.

VAINIO, K. 1947: "Saksalaiskasveista" Pohjois-

Suomessa. – Luonnon Tutkija 51(5): 172-173.

VALDÉS, B., with contributions from Raab-Straube,

E. von & Parolly, G. 2009: Ericaceae. – In:

Euro+Med Plantbase – the information resource

for Euro-Mediterranean plant diversity. –

Published on the Internet

http://ww2.bgbm.org/EuroPlusMed/PTaxonDetai

l.asp?NameCache=Kalmia%20polifolia&PTRefF

k=7100000. – Accessed on 2.2.2012.

VALLE, K. J. 1930: Kasvifenologisia havaintoja

Petsamosta. – Luonnon Ystävä 34(2): 56-61.

VALLE, K. J. 1931: Petsamon kolttakylien

rikkaruohokasvistosta. – Luonnon Ystävä 35(5):

171.

VALLE, K. J. 1933a: Kasvitietoja Petsamosta. –

Memoranda Societatis pro Fauna et Flora

Fennica 8: 259-274.

VALLE, K. J. 1933b: Kasvifenologisia havaintoja

Lapin matkalta kesällä 1930. – Luonnon Ystävä

37(1): 20-21.

VANHATALO, P. 1965: Havaintoja Utsjoen

kyläkasvistosta. – M. Sc. Thesis, Department of

Biology, University of Turku. 145 pp. +

Appendix 1-13.

VÄRE, H. 2007: Typification of names published by

the Finnish botanist Fredrik Nylander. – Annales

Botanici Fennici 44: 465-480.

VÄRE, H. & PARTANEN, R. 2009: Suomen

tunturikasvio. – Metsäkustannus Oy. Helsinki.

255 pp.

VÄRE, H., KAIPIAINEN-VÄRE, H. &

SYRJÄNEN, K. 2015: Kuonjarvarrin ja

lähituntureiden kalkkiylänköjen kasvit. – Lutukka

31(4): 99-112.

VÄRE, H., ULVINEN, T., VILPA, E. &

KALLEINEN, L. 2005: Oulun kasvit –

Piimäperältä Pilpasuolle. – Norrlinia 11: 1-512.

VÄRE, H., KAIPIAINEN, H. & SYRJÄNEN, K.

2008: Toskalharji – Enontekiön suurtuntureiden

aatelia. – Lutukka 24(3): 67-83.

VÄRE, H., SYRJÄNEN, K. & KAIPIAINEN-

VÄRE, H. 2010: Porojärven tunturialueen kasvit.

– Lutukka 26(4): 103-121.

VARGA, S. & KYTÖVIITA, M-M. 2010a: Gender

dimorphism and mycorrhizal symbiosis affect

floral visitors and reproductive output in

Geranium sylvaticum. – Functional Ecology

24(4): 750-758.

VARGA, S. & KYTÖVIITA, M-M. 2010b:

Mycorrhizal benefit differs among the sexes in a

gynodioecious species. – Ecology 91(9): 2583-

2593.

VARGA, S., KYTÖVIITA, M-M. & SIIKAMÄKI,

P. 2009: Sexual differences in response to

simulated herbivory in the gynodioecious herb

Geranium sylvaticum. – Plant Ecology 202: 325-

336.

VINNAMO, S. 1963: Kevonsuun kentän kasvistosta

ja kasvillisuudesta. – M. Sc. Thesis, Department

of Botany, University of Turku. 34 pp. + 2 maps.

VIRTANEN, R. 1990: Kasvistohavaintoja Pallas-

Ounastunturin kansallispuiston pohjoisosassa. –

Lutukka 6(3): 81-86.

VIRTANEN, R. & VÄRE, H. 1990: Haltin kasvisto.

– Lutukka 6(2): 35-41.

VOLKOVA, P., RUDAKOVA, V. & SHIPUNOV,

A. 2007: Sex ratios in populations of Geranium

sylvaticum in European Russia. – Plant Species

Biology 22(2): 125-128.

VORREN, K-D. 1967: Evig tele i Norge. – Ottar 51:

1-26.

VORREN, K-D. 1968: Polemochorer i Neiden. –

Blyttia 26: 11-14.

VOSS, D. H. 2011: A key for Ledum in

Rhododendron. – Journal of the American

Rhododendron Society 65(2): 99-101.

VOSS, E. G. 1985: Michigan flora 2. Dicots

(Saururaceae – Cornaceae). – Ann Arbor. 724 pp.

VUORI, P. & PERTOLA, E. 1959a: Kasvihavaintoja

itäiseltä Saariselältä (KemL). – Luonnon Tutkija

63(1): 24-25.

128 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

WAERN, M. & PEKKARI, S. 1980: A forgotten

Myriophyllum species in Norden. – Meddelanden

Växtbiologiska institutionen Uppsala 1980(3):

97-98.

WAHLENBERG, G. 1812: Flora Lapponica. –

Berolini. 550 pp. + Table I-XXVI.

WAINIO, E. A. 1891: Notes sur la flore de la

Laponie finlandaise. – Acta Societatis pro Fauna

et Flora Fennica 8(4): 1-90.

WIGGINS, I. L. & THOMAS, J. H. 1962: A flora of

the Alaskan Arctic Slope. – Toronto. 425 pp.

WISTRAND, G. 1962: Studier i Pite Lappmarks

kärlväxtflora med särskild hänsyn till skogslandet

och de isolerade fjällen. – Acta Phytogeographica

Suecica 45: 1-211 + 168 maps.

ZIZKA, G. 1985: Botanische Untersuchungen in

Nordnorwegen I. Anthropochore Pflanzenarten

der Varangerhalbinsel und Sør-Varanger. –

Dissertationes Botanicae 85: 3-102.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 129

Map 1 ERODIUM CICUTARIUM

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Map 2 GERANIUM PUSILLUM

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130 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 3 GERANIUM SYLVATICUM

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Map 4 LINUM USITATISSIMUM

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REP. KEVO SUBARCTIC RES. STAT. 25. 2019 131

Map 5 IMPATIENS GLANDULIFERA

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Map 6 MALVA PUSILLA

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132 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 7 HYPERICUM MACULATUM

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Map 8 VIOLA ARVENSIS

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REP. KEVO SUBARCTIC RES. STAT. 25. 2019 133

Map 9 VIOLA BIFLORA

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Map 10 VIOLA CANINA

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134 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 11 VIOLA EPIPSILA

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Map 12 VIOLA PALUSTRIS

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REP. KEVO SUBARCTIC RES. STAT. 25. 2019 135

Map 13 VIOLA X RUPRECHTIANA

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Map 14 VIOLA TRICOLOR VAR. TRICOLOR

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136 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 15 VIOLA X WITTROCKIANA

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Map 16 MYRICARIA GERMANICA

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REP. KEVO SUBARCTIC RES. STAT. 25. 2019 137

Map 17 ELATINE HYDROPIPER S. STR.

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Map 18 CIRCAEA ALPINA

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138 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 19 EPILOBIUM ADENOCAULON

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Map 20 EPILOBIUM ALSINIFOLIUM

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REP. KEVO SUBARCTIC RES. STAT. 25. 2019 139

Map 21 EPILOBIUM ANAGALLIDIFOLIUM

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Map 22 EPILOBIUM ANGUSTIFOLIUM

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140 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 23 EPILOBIUM CILIATUM

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Map 24 EPILOBIUM COLLINUM

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REP. KEVO SUBARCTIC RES. STAT. 25. 2019 141

Map 25 EPILOBIUM DAVURICUM

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Map 26 EPILOBIUM HORNEMANNII

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142 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 27 EPILOBIUM LACTIFLORUM

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Map 28 EPILOBIUM PALUSTRE

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REP. KEVO SUBARCTIC RES. STAT. 25. 2019 143

Map 29 MYRIOPHYLLUM ALTERNIFLORUM

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Map 30 MYRIOPHYLLUM SIBIRICUM

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144 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 31 HIPPURIS VULGARIS

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Map 32 CORNUS SUECICA

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777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 145

Map 33 AEGOPODIUM PODAGRARIA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 34

ANGELICA ARCHANGELICA SUBSP. ARCHANGELICA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

146 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 35 ANGELICA SYLVESTRIS

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 36 ANTHRISCUS SYLVESTRIS

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 147

Map 37 CARUM CARVI

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 38 CHAEROPHYLLUM PRESCOTTII

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

148 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 39 CICUTA VIROSA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 40 HERACLEUM SIBIRICUM VAR. SIBIRICUM

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 149

Map 41 PASTINACA SATIVA SUBSP. SATIVA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 42 PETROSELINUM CRISPUM

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

150 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 43 PIMPINELLA SAXIFRAGA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 44 DIAPENSIA LAPPONICA SUBSP. LAPPONICA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 151

Map 45 MONESES UNIFLORA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 46 ORTHILIA SECUNDA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

152 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 47 PYROLA CHLORANTHA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 48 PYROLA MEDIA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 153

Map 49 PYROLA MINOR

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 50 PYROLA ROTUNDIFOLIA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

154 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 51 ANDROMEDA POLIFOLIA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 52 ARCTOSTAPHYLOS ALPINA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 155

Map 53 ARCTOSTAPHYLOS UVA-URSI

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 54 CALLUNA VULGARIS

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

156 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 55 CASSIOPE HYPNOIDES

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 56 KALMIA POLIFOLIA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 157

Map 57 LEDUM PALUSTRE

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 58 LOISELEURIA PROCUMBENS

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

158 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 59 PHYLLODOCE CAERULEA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 60 RHODODENDRON LAPPONICUM

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 159

Map 61 VACCINIUM MICROCARPUM

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 62 VACCINIUM MYRTILLUS

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

160 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 63 VACCINIUM OXYCOCCOS

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 64 VACCINIUM ULIGINOSUM

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 161

Map 65 VACCINIUM VITIS-IDAEA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 66

EMPETRUM NIGRUM SUBSP. HERMAPHRODITUM

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

162 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 67 EMPETRUM NIGRUM SUBSP. NIGRUM

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 68 LYSIMACHIA THYRSIFLORA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 163

Map 69 PRIMULA STRICTA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

Map 70 PRIMULA VERIS

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

778778

777777

776776

775775

774774

773773

772772

771771

770770

769769

768768

767767

766766

765765

764764

763763

762762

761761

760760

759759

758758

757757

164 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

Map 71 TRIENTALIS EUROPAEA

EUR

50

770

FMF

40 50

760

770

41

42

43

44

45

46

47

48

49

50

51

52

53

54

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REP. KEVO SUBARCTIC RES. STAT. 25. 2019 165

Reports from the Kevo Subarctic Research Station

(Rep. Kevo Subarctic Res. Stat.)

ISSN 0453-7831

Vol. 1 (1964)

KALLIO, P.: The Kevo Subarctic Research Station of the University of Turku, 9-40.

SILTANEN, P.: The aquatic flora and vegetation of Lake Kevojärvi, 41-59.

NYMAN, A.: Aquatic vegetation of Lake Mantojärvi in Inari Lapland, Finland, 60-68.

RAUTAVA, E.: Über die Wasservegetation des Flusses Vaskojoki im nördlichsten Finnland, 69-93.

LAINE, U.: Über die floristischen Züge der nördlichen Waldgrenze der Kiefer im Westteil von Inari-

Lappland, 94-123.

MÄKINEN, Y.: Floristic observations in Finnmark (Northern Norway), 124-128.

MÄKINEN, L.: On the morphology of Primula sibirica Jacq. ssp. finmarchica (Jacq.) Hult., 129-131.

HAKULINEN, R.: Beobachtungen über die Flechtenflora und Flechtenvegetation von Utsjoki, Nordfinnland,

132-139.

KUKKONEN, I.: Facts and speculations about the factors affecting the distribution of Anthracoidea scirpi as

a parasite of Trichophorum caespitosum, 140-148.

ERIKSSON, M.: Larger fungi on dunes in Finland, 149-154.

MÄKINEN, Y.: On Finnish micromycetes 3. Uredinales of Inari Lapland, 155-177.

KALLIO, P. & KANKAINEN, E.: Notes on the macromycetes of Finnish Lapland and adjacent Finnmark,

178-235.

KUPIAS, R.: On filamentous blue-green algae of brooks of Lapland, 236-249.

SILVOLA, T.: On the land molluscs of the Kevojoki River valley in Finnish Lapland, 250-268.

JUSSILA, R.: Occurrence of macrolepidoptera in the biotopes of the Kevojoki area in Inari Lapland (Finland),

269-278.

LEHTINEN, P. T.: The Phalangids and Pseudoscorpionids of Finnish Lapland, 279-287.

LINDQVIST, O. V.: The spider fauna of the cliffs in eastern Finnish Lapland, 288-291.

BAGGE, P.: A freshwater amphipod Gammarus lacustris Sars in Utsjoki, Finnish Lapland, 292-294.

LAINE, H.: Notes on some southern bird species found in the vicinity of Kevo in Utsjoki, Finnish Lapland,

295-300.

SCHANTZ, M. von & IVARS, L.: Über die Zusammensetzung des ätherischen Öles von Thymus serpyllum

ssp. tanaënsis (Hyl.) Jalas, 301-307.

UNGERSON, J. & SCHERDIN, G.: Über die plötzlichen Änderungen in dem Tagesverlauf der

Photosynthese und der Atmung unter natürlichen Bedingungen, 308-321.

MANSIKKANIEMI, H.: Main features of the glacial and postglacial development of Pulmanki valley in

northernmost Finland, 322-337.

SYRILÄ, S.: Retreat of the continental ice and fluvioglacial erosion features on the Tuolbanjaugoaivi fjeld in

northernmost Finland, 338-345.

PETÄJÄ, A.: Depth charts of some lakes in Utsjoki, Finnish Lapland, 346-349.

VESANEN, E.: An outline of the development of the seismograph station network and of the study of the

seismicity of the Baltic Shield, 350-356.

PETÄJÄ, A.: Simplified apparatus for recording waterlevel fluctuations, 357-358.

Vol. 2 (1965)

JUSSILA, R.: The Ichneumonidae of the Kevojoki area in Inari Lapland (Finland), 3-186.

Vol. 3 (1966)

HUSTICH, I.: On the forest-tundra and the northern tree-lines, 7-47.

HEIKKILÄ, H. & KALLIO, P.: On the problem of subarctic basidiolichenes I, 48-74.

MÄKINEN, Y.: On the macroecology of some rust fungi, 75-84.

KALLIO, P. & KÄRENLAMPI, L.: Observations on the lichens of Labrador and Ungava, 85-100.

166 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

HAKULINEN, R. & ULVINEN, T.: Asahinea chrysantha (Tuck.) Culb. et Culb. in Fennoscandien, 101-105.

PIHAKASKI, S.: Studies on northern Luzula multiflora (Retz.) Lej. races, 106-131.

RAUDASKOSKI, M.: Studies on the karyology of the smut Anthracoidea limosa (Ustilaginales), 132-142.

PIHAKASKI, K.: Ecological and morphological studies on Luzula arcuata (Wg.) Sw., 143-176.

KALLIO, P. & KANKAINEN, E.: Additions to the mycoflora of northernmost Finnish Lapland, 177-210.

HAAPASAARI, M.: The genus Gymnomitrion Corda in Finland, 211-235.

LAINE, H.: On some influences of the Atlantic and the Arctic Ocean upon the bird fauna of Utsjoki, Finnish

Lapland, 236-258.

NUORTEVA, P.: Observations on the diel periodicity of flight by Lonchaea laxa Collin (Dipt., Lonchaeidae)

in subarctic conditions, 259-260.

SILVOLA, T.: Quantitative observations on the avifauna of the Kevojoki River valley, 261-273.

SEPPÄLÄ, M.: Recent ice-wedge polygons in eastern Enontekiö, northernmost Finland, 274-287.

Vol. 4 (1967 - 1969)

MANSIKKANIEMI, H. (1967): Geomorphological analysis of Pulmanki-Tana valley in Lapland, 7-31.

BAGGE, P. (1968): Ecological studies on the fauna of subarctic waters in Finnish Lapland, 32-83.

JUSSILA, R. (1968): Distribution of Ichneumonidae (Hymenoptera) at different altitude belts in Finnish

Lapland, 84-89.

HEIKKILÄ, H. & KALLIO, P. (1969): On the problem of subarctic basidiolichenes II, 90-97.

VUORISALO, A. (1969): Conjugatophyceae of Utsjoki, Finnish Lapland, 98-111.

ERIKSSON, J. & STRID, Å. (1969): Studies in the Aphyllophorales (Basidiomycetes) of Northern Finland,

112-158.

HUSTICH, I. (1969): Notes on the growth of pine in northern Finland and Norway, 159-170.

Vol. 5 (1969)

KALLIO, P., LAINE, U. & MÄKINEN, Y.: Vascular flora of Inari Lapland. 1. Introduction and

Lycopodiaceae - Polypodiaceae, 1-108.

MÄKINEN, Y.: On Finnish micromycetes. 8. Erysiphales of Inari Lapland, 109-116.

Vol. 6 (1970)

MANSIKKANIEMI, H.: Deposits of sorted material in the Inarijoki-Tana river valley in Lapland, 1-63.

Vol. 7 (1970 - 1971)

KÄRENLAMPI, L. (1970): Distribution of chlorophyll in the lichen Cladonia alpestris, 1-8.

KÄRENLAMPI, L. (1970): Morphological analysis of the growth and productivity of the lichen Cladonia

alpestris, 9-15.

MANSIKKANIEMI, H. (1970): The sinuosity of rivers in northern Finland, 16-32.

KÄRENLAMPI, L. (1971): Studies on the relative growth rate of some fruticose lichens, 33-39.

KÄRENLAMPI, L. (1971): On methods for measuring and calculating the energy flow through lichens, 40-

46.

KÄRENLAMPI, L. & PELKONEN, M. (1971): Studies on the morphological variation of the lichen

Cladonia uncialis, 47-56.

KOPONEN, S. (1971): On the abundance relations of mesofaunal groups in the ground layer of three subarctic

habitats, 57-59.

HAUKIOJA, E. (1971): Summer schedule of some subarctic passerine birds with reference to postnuptial

moult, 60-69.

MÄKINEN, Y. (1971): On Finnish micromycetes. 9. Sphaerotheca drabae Juel on Saxifraga nivalis in

Finnish Lapland, 70-73.

EUROLA, S. (1971): The driftwoods of the Arctic Ocean, 74-80.

Vol. 8 (1971)

ANDREEV, V. N.: Methods of defining overground phytomass on vast territories of the Subarctic, 3-11.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 167

BLÜTHGEN, J.: Die Dokumentation der Herbstfärbung und ihre floristisch-systematische Differenzierung in

Lappland, 12-21.

GROVES, J. W. & ELLIOTT, M. E.: Notes on fungi from Northern Canada VI. Additional records of

Discomycetes, 22-30.

HARE, F. K.: Snow-cover problems near the Arctic tree-line of North America, 31-40.

HAVAS, P.: The water economy of the bilberry (Vaccinium myrtillus) under winter conditions, 41-52.

HOLTMEIER, F-K.: Waldgrenzstudien im nördlichen Finnisch-Lappland und angrenzenden Nordnorwegen,

53-62.

KALLIO, P. & HEINONEN, S.: Influence of short-term low temperature on net photosynthesis in some

subarctic lichens, 63-72.

KALLIO, P., LAINE, U. & MÄKINEN, Y.: Vascular flora of Inari Lapland. 2. Pinaceae and Cupressaceae,

73-100.

KÄRENLAMPI, L.: Weight loss of leaf litter on forest soil surface in relation to weather at Kevo Station,

Finnish Lapland, 101-103.

LAINE, U. & NURMI, J.: Factors affecting vegetation and flora of anorthosite and granulite areas in western

Inari, Finnish Lapland, 104-115.

MIKOLA, P.: Reflexion of climatic fluctuations in the forestry practices of Northern Finland, 116-121.

OHENOJA, E.: The larger fungi of Svalbard and their ecology, 122-147.

RYVARDEN, L.: Studies in the Aphyllophorales of Finnmark, Northern Norway, 148-154.

SIRÉN, G. & HARI, P.: Coinciding periodicity in recent tree rings and glacial clay sediments, 155-157.

TRANQUILLINI, W. & MACHL-EBNER, I.: Über den Einfluss von Wärme auf das

Photosynthesevermögen der Zirbe (Pinus cembra L.) und der Alpenrose (Rhododendron ferrugineum L.) im

Winter, 158-166.

Vol. 9 (1972)

SEPPÄLÄ, M.: Peat at the top of Ruohttir fell, Finnish Lapland, 1-6.

KALLIO, P., SUHONEN, S. & KALLIO, H.: The ecology of nitrogen fixation in Nephroma arcticum and

Solorina crocea, 7-14.

MANSIKKANIEMI, H.: Flood deposits, transport distances and roundness of loose material in the Tana river

valley, Lapland, 15-23.

WIELGOLASKI, F. E. & KJELVIK, S.: The methodology of net primary production investigations in

Norwegian IBP tundra studies, 24-27.

LEMMETYINEN, R.: Nest defense behaviour in the arctic tern Sterna paradisaea towards stuffed nest

predators on Spitsbergen, 28-31.

KOPONEN, S.: On the spiders of the ground layer of a pine forest in Finnish Lapland, with notes on their

diurnal activity, 32-34.

STRID, Å.: Aspects on the Daedaleopsis Schroet. complex (Polyporaceae) in Fennoscandia and Denmark, 35-

43.

VOIPIO, P.: Problems of cold adaptation in the Red squirrel Sciurus vulgaris, 44-49.

KÄRENLAMPI, L.: Comparisons between the microclimates of the Kevo ecosystem study sites and the Kevo

Meteorological Station, 50-65.

KÄRENLAMPI, L.: Factor analytic studies on the vegetation of the surroundings of the Kevo Subarctic

Station, 66-72.

HAUKIOJA, E. & NYGRÉN, K.: Short-distance movements in the grey-sided vole (Clethrionomys

rufocanus), 73-77.

KÄRENLAMPI, L.: On the relationships of the Scots pine annual ring width and some climatic variables at

the Kevo Subarctic Station, 78-81.

KÄRENLAMPI, L. & KAUHANEN, H.: A direct gradient analysis of the vegetation of the surroundings of

the Kevo Subarctic Station, 82-98.

RAUTAVA, E.: Amphiphytic and aquatic moss vegetation in the rivers Vaskojoki and Kettujoki in Finnish

Lapland, 99-107.

Vol. 10 (1973)

BUNNELL, F. L., KÄRENLAMPI, L. & RUSSELL, D. E.: A simulation model of lichen-Rangifer

interactions in Northern Finland, 1-8.

168 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

HAUKIOJA, E.: Weight development, consumption and egestion of Dineura virididorsata (Hym.,

Tenthredinidae) larvae, 9-13.

KOPONEN, S.: On the mining insects of the mountain birch in northernmost Fennoscandia, 14-19.

KOPONEN, S.: Herbivorous invertebrates of the mountain birch at Kevo, Finnish Lapland, 20-28.

HAUKIOJA, E., KOPONEN, S. & OJALA, H.: Local differences in birch consumption by invertebrates in

northern Norway and Finland, 29-33.

KALLIO, S.: The ecology of nitrogen fixation in Stereocaulon paschale, 34-42.

KALLIO, P. & HEINONEN, S.: Ecology of Rhacomitrium lanuginosum (Hedw.) Brid., 43-54.

KALLIO, P. & LEHTONEN, J.: Birch forest damage caused by Oporinia autumnata (Bkh.) in 1965-66 in

Utsjoki, N Finland, 55-69.

VAARAMA, A. & VALANNE, T.: On the taxonomy, biology and origin of Betula tortuosa Ledeb., 70-84.

Vol. 11 (1974)

ALEXANDER, V., BILLINGTON, M. & SCHELL, D.: The influence of abiotic factors on nitrogen fixation

rates in the Barrow, Alaska, arctic tundra, 3-11.

BARR, M. E.: The Cucurbitaria sorbi Karsten complex, 12-15.

HAUKIOJA, E.: Measuring consumption in Eriocrania (Eriocraniidae, Lep.) miners with reference to

interaction between the leaf and the miner, 16-21.

HAUKIOJA, E. & HEINO, J.: Birch consumption by reindeer (Rangifer tarandus) in Finnish Lapland, 22-

25.

HINNERI, S.: Podzolic processes and bioelement pools in subarctic forest soils at the Kevo Station, Finnish

Lapland, 26-34.

HUSTICH, I.: Common species in the northern part of the Boreal Region of Canada. An essay, 35-41.

KALLIO, S. & VARHEENMAA, T.: On the effect of air pollution on nitrogen fixation in lichens, 42-46.

KJELVIK, S. & WIELGOLASKI, F. E.: Biomass, nutrient content and energy of some dwarf shrubs in a

Norwegian subalpine birch forest, 47-51.

KOPONEN, S.: On the occurrence and ecology of Eriocrania spp. (Eriocraniidae) and other mining insects of

the birch in northernmost Fennoscandia in 1973, 52-64.

KOPONEN, S. & OJALA, H.: On the mesofauna of the field layer of three subarctic habitats, 65-71.

LÄHDE, E.: Rate of decomposition of cellulose in forest soils in various parts of the Nordic countries, 72-78.

LAINE, U., LEHMUSHOVI, A. & NURMI, J.: Chromosome numbers of phanerogams in Inari Lapland and

adjacent regions, 79-89.

MÄKINEN, Y. & OIKARINEN, H.: Cultivation of cloudberry in Fennoscandia, 90-102.

MANSIKKANIEMI, H.: Some methods of measuring fluvial transportation load, alluvial fan of Jomppala in

Utsjoki, Finland, 103-111.

VAARAMA, A. & LAINE, U.: The southern element in the moss flora of Utsjoki, Inari Lapland, North

Finland, 112-125.

Vol. 12 (1975)

HAUKIOJA, E. & HAKALA, T.: Herbivore cycles and periodic outbreaks. Formulation of a general

hypothesis, 1-9.

HINNERI, S.: On the water chemistry of the Utsjoki River System, and its significance for the evaluation of

edaphic conditions in the drainage area, 10-24.

KALLIO, P.: Leccinum scabrum (Fries) S. F. Gray subsp. tundrae Kallio, a new subspecies from Lapland, 25-

27.

KALLIO, P. & KALLIO, S.: Nitrogen fixation by free-living micro-organisms in the Kevo district, 28-33.

KARCZEWSKI, A.: Morphology and textural-structural features of ground and hummocky moraine in the

Paistunturit area, Finnish Lapland, 34-44.

KOPONEN, S. & OJALA, M-L.: Quantitative study of invertebrate groups in the soil and ground layer of the

IBP sites at Kevo, northern Finland, 45-52.

DOROGOSTAISKAYA, E. V.: Weeds of the Far North of the U.S.S.R., 53.

HIPPA, H. & KOPONEN, S.: On the damage caused by the species of Galerucella (Col., Chrysomelidae) on

cloudberry (Rubus chamaemorus L.) in Finland and northern Norway. 54-59.

KALLIO, H.: Chemical constituents of the volatile aroma compounds in Rubus arcticus L. subsp. stellatus

(Sm.) Boivin, with reference to Rubus arcticus L. subsp. arcticus, 60-65.

REP. KEVO SUBARCTIC RES. STAT. 25. 2019 169

KALLIO, P. & MÄKINEN, Y.: Vascular flora of Inari Lapland. 3. Salicaceae, 66-105.

Vol. 13 (1976)

SEPPÄLÄ, M.: Periglacial character of the climate of the Kevo region (Finnish Lapland) on the basis of

meteorological observations 1962-71, 1-11.

ALEXANDER, V. & KALLIO, S.: Nitrogenase activity in Peltigera aphthosa and Stereocaulon paschale in

early spring, 12-15.

KALLIO, S., KALLIO, P. & RASKU, M-L.: Ecology of nitrogen fixation in Peltigera aphthosa (L.) Willd.

in North Finland, 16-22.

FONG, D. W. & BAL, A. K.: Histological differences between chilled and nonchilled seeds in Rubus

chamaemorus L., 23-25.

HAUKIOJA, E. & ISO-IIVARI, L.: Local and annual variation in secondary production by Dineura

virididorsata (Hym., Tenthredinidae), 26-32.

ISO-IIVARI, L. & KOPONEN, S.: Insect catches by light trap compared with geomagnetic and weather

factors in subarctic Lapland, 33-35.

HIPPA, H., KOPONEN, S. & NEUVONEN, S.: Population dynamics of the form of Galerucella

nymphaeae-complex (Col., Chrysomelidae) living on cloudberry in northern Finland, 36-39.

HIPPA, H. & KOPONEN, S.: Distribution of the species of Galerucella (Col., Chrysomelidae) on cloudberry

in Fennoscandia, 40-43.

HAUKIOJA, E. & NIEMELÄ, P.: Does birch defend itself actively against herbivores?, 44-47.

KOPONEN, S.: Spider fauna (Araneae) of Kevo area, northernmost Finland, 48-62.

KARUNEN, P. & KALLIO, P.: Seasonal variation in the total lipid content of subarctic Dicranum

elongatum, 63-70.

Vol. 14 (1978)

HAUKIOJA, E. & SALOVAARA, R.: Summer weight of reindeer (Rangifer tarandus) calves and its

importance for their future survival, 1-4.

HAUKIOJA, E., NIEMELÄ, P., ISO-IIVARI, L., OJALA, H. & ARO, E-M.: Birch leaves as a resource

for herbivores. I. Variation in the suitability of leaves, 5-12.

KOPONEN, S.: Notes on herbivorous insects of the birch in southern Greenland, 13-17.

HIPPA, H., KOPONEN, S. & LAINE, T.: On the feeding biology of Coccinella hieroglyphica L. (Col.,

Coccinellidae), 18-20.

HAUKIOJA, E., NIEMELÄ, P. & ISO-IIVARI, L.: Birch leaves as a resource for herbivores. II. Diurnal

variation in the usability of leaves for Oporinia autumnata and Dineura virididorsata, 21-24.

KARUNEN, P., HEINONEN, S. & LYLY, O.: Persistence of mecoprop in northern and southwestern

Finland, 25-30.

HIPPA, H., KOPONEN, S. & OSMONEN, O.: Role of bees (Hym., Apidae) in pollination of the cloudberry

(Rubus chamaemorus L.) in northern Fennoscandia, 31-37.

KALLIO, P. & MÄKINEN, Y.: Vascular flora of Inari Lapland. 4. Betulaceae, 38-63.

OKSANEN, L.: Lichen grounds of Finnmarksvidda, northern Norway, in relation to summer and winter

grazing by reindeer, 64-71.

VALANNE, T.: Soft-wood rooting experiments with Betula, 72-75.

Vol. 15 (1979)

FORSÉN, K.: Aroma constituents of Angelica archangelica. Variations in the composition of the essential

root oil of strains of var. norvegica and var. sativa, 1-7.

HIPPA, H. & KOPONEN, S.: Experiments on biological control of leaf beetles (Col., Chrysomelidae) on the

cloudberry (Rubus chamaemorus L.), 8-10.

INKI, M. & VALANNE, T.: The F1 hybrid between Betula glandulosa, subsection Nanae and Betula

pendula, subsection Albae, 11-18.

KOPONEN, S. & LINNALUOTO, E. T.: Flight periods and abundance of some moths caught by light traps

in subarctic Finnish Lapland, 1972-78, 19-26.

LEHTONEN, J. & YLI-REKOLA, M.: Field and ground layer vegetation in birch forests after Oporinia

damage, 27-32.

170 REP. KEVO SUBARCTIC RES. STAT. 25. 2019

NIEMELÄ, P.: Topographical delimitation of Oporinia-damages: Experimental evidence of the effect of

winter temperature, 33-36.

NIEMELÄ, P., ARO, E-M. & HAUKIOJA, E.: Birch leaves as a resource for herbivores. Damage-induced

increase in leaf phenols with trypsin-inhibiting effects, 37-40.

REID, D. A.: Some fungi from Spitsbergen, 41-47.

WORKMAN, C.: Life cycles, growth rates and reproductive effort in lycosid and other spiders, 48-55.

Vol. 16 (1980)

HELLE, T.: Abundance of warble fly (Oedemagena tarandi) larvae in semi-domestic reindeer (Rangifer

tarandus) in Finland, 1-6.

KOPONEN, S.: Herbivorous insects of the birch in Iceland, 7-12.

KOPONEN, S.: Spider fauna in the Adventfjorden area, Spitsbergen, 13-16.

KALLIO, H., LAINE, M. & HUOPALAHTI, R.: Aroma of the berries of the hybrid Rubus stellatus Sm. x

R. arcticus L., 17-22.

NIEMINEN, M., TIMISJÄRVI, J. & LAITINEN, M.: The effects of antiparasitic treatment on the condition

of semi-domestic reindeer (Rangifer tarandus), 23-26.

NIEMELÄ, P.: Dependence of Oporinia autumnata (Lep., Geometridae) outbreaks on summer temperature,

27-30.

SULKINOJA, M. & VALANNE, T.: Polyembryony and abnormal germination in Betula pubescens subsp.

tortuosa, 31-37.

INKI, M. & VÄISÄNEN, L.: Essential oils in Betula tortuosa Ledeb. and in some other Betula species and

hybrids, 38-44.

NIEMELÄ, P., ISO-IIVARI, L., LAINE, C. & LAINE, K. J.: Food plant preference and larval growth of

larval colour forms of Entephria caesiata (Schiff.) (Lep., Geometridae), 45-48.

NIEMELÄ, P., TUOMI, J. & HAUKIOJA, E.: Age-specific resistance in trees: Defoliation of tamaracks

(Larix laricina) by larch bud moth (Zeiraphera improbana) (Lep., Tortricidae), 49-57.

Vol. 17 (1981)

AURELA, A. & PUNKKINEN, R.: Atmospheric nitrogen dioxide and northern plants, 1- 6.

OKSANEN, L. & OKSANEN, T.: Lemmings (Lemmus lemmus) and grey-sided voles (Clethrionomys

rufocanus) in interaction with their resources and predators on Finnmarksvidda, northern Norway, 7-31.

NOORDELOOS, M. E.: Notes on Entoloma (Basidiomycetes, Agaricales) in Inari Lapland, northernmost

Finland, 32-40.

SOLHØY, T. & KOPONEN, S.: Oribatei fauna (Acari) on alpine heath at Kevo, Finland, 41-43.

HIPPA, H., KOPONEN, S. & OSMONEN, O.: Flower visitors to the cloudberry (Rubus chamaemorus L.) in

northern Fennoscandia, 44-54.

HIPPA, H., KOPONEN, S. & OSMONEN, O.: Diurnal activity of flower visitors to the cloudberry (Rubus

chamaemorus L.), 55-57.

HIPPA, H., KOPONEN, S. & OSMONEN, O.: Pollen transport and pollinating efficiency of flower visitors

to the cloudberry (Rubus chamaemorus L.) in northern Fennoscandia, 58-66.

PIHAKASKI, K.: Seasonal changes in structure of mesophyll cells in subarctic Diapensia lapponica L., 67-

80.

SOLDÁN, T.: The mayflies (Ephemeroptera) of Utsjoki, northernmost Finland, 81-85.

Vol. 18 (1982)

HIPPA, H., KOPONEN, S. & ROINE, R.: Feeding preference of Coccinella hieroglyphica (Col.,

Coccinellidae) for eggs of three chrysomelid beetles, 1-4.

PRUDHOMME, T. I.: The effect of defoliation history on photosynthetic rates in mountain birch, 5-9.

MÄKINEN, Y., KALLIO, P., LAINE, U. & NURMI, J.: Vascular flora of Inari Lapland. 5. Urticaceae -

Caryophyllaceae, 10-94.

Vol. 19 (1984)

VALANNE, N. & VALANNE, T.: The development of the photosynthetic apparatus during bud burst and

leaf opening in two subspecies of Betula pubescens Ehrh., 1-10.

KULLMAN, L.: Germinability of mountain birch (Betula pubescens ssp. tortuosa) along two altitudinal

transects downslope from the tree-limit, in Sweden, 11-18.

KOPONEN, S.: Abundance of herbivorous insects on dwarf birch near the treeline in Alaska, 19-24.

HOOGESTEGER, M.: The effect of trampling on vegetation at four cottages in Torne Lapland, northern

Sweden, 25-34.

HELLE, T.: Foraging behaviour of the semi-domestic reindeer (Rangifer tarandus L.) in relation to snow in

Finnish Lapland, 35-47.

HELLE, T. & TARVAINEN, L.: Determination of the winter digging period of semi-domestic reindeer in

relation to snow conditions and food resources, 49-56.

HELLE, T. &ASPI, J.: Do sandy patches help reindeer against insects?, 57-62.

HIPPA, H. & KOPONEN, S.: Parasitism of larvae of Galerucini (Col., Chrysomelidae) by larvae of

Asecodes mento (Hym., Eulophidae), 63-65.

HIPPA, H., KOPONEN, S. & ROINE, R.: Larval growth of Coccinella hieroglyphica (Col., Coccinellidae)

fed on aphids and preimaginal stages of Galerucella sagittariae (Col., Chrysomelidae), 67-70.

KÄRPPÄ, J., KALLIO, H., PELTONEN, I. & LINKO, R.: Anthocyanins in the northern and southern

crowberry, Empetrum nigrum coll., 71-73.

KALLIO, H., NIKKOLA, P. & REUNANEN, M.: Sugar composition of the juice of crowberry, Empetrum

hermaphroditum, 74-76.

SALEMAA, H.: Chromosomes of the freshwater amphipod Gammarus lacustris G. O. Sars, 77-80.

Vol. 20 (1987)

SVEINBJÖRNSSON, B.: Biomass proportioning as related to plant size in juvenile mountain birch near

Abisko, Swedish Lapland, 1-8.

ENGELSKJØN, T. & SKIFTE, O.: Distribution and ecology of Trichophorum pumilum (Vahl) Sch. & Th.

(Cyperaceae) in Norway, 9-19.

HÖMMÖ, L. & VALANNE, T.: Cytological and morphological analyses of grafted triploid aspens (Populus

tremula L.) from the Nonabeljävri area in Finnish Lapland, 21-25.

SULKINOJA, M. & VALANNE, T.: Leafing and bud size in Betula provenances of different latitudes and

altitudes, 27-33.

LEHTONEN, J.: Recovery and development of birch forests damaged by Epirrita autumnata in Utsjoki

area, North Finland, 35-39.

OJALA, A., HINNERI, S. & YLIAHO, H.: Mineral element content of Angelica archangelica subsp.

archangelica, 41-45.

Vol. 21 (1990)

NENONEN, S-P. & MANSIKKANIEMI, H.: Observations on the spread of vegetation to road cuttings in

the far north of Finland, 1-9.

MANSIKKANIEMI, H. & LAITINEN, T.: Pattern of local wind changes in a fell region, northern Finland,

11-20.

MATSUKI, M. & MACLEAN, S. F. Jr.: The effect of temperature on the molt of Dineura virididorsata

(Hymenoptera, Tenthredinidae), 21-25.

MOLSKI, B. & DMUCHOWSKI,W.: The comparison of environmental pollution in northern Finland near

Kevo and in Poland with the use of Pinus sylvestris L. as bioindicator, 27-30.

KORTELAINEN, I.: Gammarus lacustris - herbivore or predator?, 31-34.

Vol. 22 (1998)

LIIRA, T. & HIETARANTA, J.: On the postglacial development of the bedrock precipices and talus

formations in the Keävvu river valley, northern Finnish Lapland, 1-10.

BOGACHEVA, I.: Ecological and ontogenetic heterogeneity of leaves and its role in insect-plant

relationships, 11-17.

LUNDVALL, P., NEUVONEN, S. & HALONEN, M.: Interspecific differences in the susceptibility of adult

leaf beetles (Col., Chrysomelidae) to predation byWillow warblers (Phylloscopus trochilus), 19-24.

MÄKINEN, Y., KALLIO, P., LAINE, U. & NURMI, J.: Vascular flora of Inari Lapland. 6. Nymphaeaceae

- Papaveraceae, 25-86.

Vol. 23 (2005)

MÄKINEN, Y., LAINE, U., HEINO, S. & NURMI, J.: Vascular flora of Inari Lapland. 7. Brassicaceae -

Grossulariaceae, 1-95.

HOLTMEIER, F-K.: Change in the timberline ecotone in northern Finnish Lapland during the last thirty

years, 97-113.

Vol. 24 (2011)

MÄKINEN, Y., LAINE, U., HEINO, S., ISO-IIVARI, L. & NURMI, J.: Vascular flora of Inari

Lapland 8. Rosaceae and Fabaceae, 3-126.

SAMULI HELAMA : Climate and Scots pine tree-rings in Utsjoki-Kevo district (North-East Finnish

Lapland) during the 20th century, with special emphasis on mid-summer connexions, 129-138.

ISBN XXX-XXX-XX-XXXX-X

ISSN 0453-7831

(nid.)

(pdf)

Vol. 25 (2019)

MÄKINEN, Y., PIIRAINEN, M., LAINE, U.†, NURMI, J., HEINO, S. & ISO-IIVARI, L.: Vascular

flora of Inari Lapland. 9. Geraniaceae – Primulaceae, 3-164.

Kuuluuko tähän yhteyteen risti Unto Laineen Nimen yhteyteen?

Vascular flora of Inari Lapland. 9. Geraniaceae - Primulaceae.

Abstract

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