Allium oleraceum (Alliaceae) in Finland: Distribution, habitats and accompanying vascular plant species

Abstract

The distribution of the field garlic, Allium oleraceum L., in Finland is presented according to herbaria voucher specimens, literature and field observations of the authors. The habitats of A. oleraceum, rock and dry meadows, mesic meadows, gardens and a few deciduous stands, are shortly described. The habitats are mostly influenced by man and domestic animals. The species is often found on iron age mounds, in ancient hilltop fortresses and in areas with medieval settlement, e.g. in connection with medieval castles and churches. A. oleraceum is moderately calciphilic as it is much more common in areas with calcareous bedrock or soil than in acid areas. The accompanying vascular plant flora is briefly presented.

Full text

Allium oleraceum (Alliaceae) in Finland: distribution,
habitats and accompanying vascular plant species
Carl-Adam Hæggström & Helena Åström
Hæggström, C.-A. & Åström, H., Department of Biological and Environmental Sciences,
P. O. Box 65, FI-00014 University of Helsinki, Finland
The distribution of the field garlic, Allium oleraceum L., in Finland is presented according
to herbaria voucher specimens, literature and field observations of the authors. The habi-
tats of A. oleraceum, rock and dry meadows, mesic meadows, gardens and a few decidu-
ous stands, are shortly described. The habitats are mostly influenced by man and domestic
animals. The species is often found on iron age mounds, in ancient hilltop fortresses and in
areas with medieval settlement, e.g. in connection with medieval castles and churches.
A. oleraceum is moderately calciphilic as it is much more common in areas with calcare-
ous bedrock or soil than in acid areas. The accompanying vascular plant flora is briefly
presented.
Key words: accompanying species, Allium oleraceum, distribution, Finland, habitats
Introduction
The field garlic, Allium oleraceum L., is an Euro-
pean plant growing in meadows and deciduous
woods. Its main distibution area comprises most of
Central Europe, England, southern Scandinavia
and southwestern Finland (Meusel et al. 1965,
Hultén 1971, Hultén & Fries 1986). It is quite
common in many areas, e.g. NW and SE Poland
(Kotliñska 1992) and the Czech Republik (Ducho-
slav 2001a). It has been introduced in the north-
eastern USA (Hultén & Fries 1986).
Allium oleraceum has a rather southern distri-
bution in Finland (Cedercreutz 1927, Hultén 1971,
Hämet-Ahti et al. 1998). It is quite common in the
ÅlandIslandsandfairlycommonintherestofthe
southwestern archipelago. Its northernmost sta-
tion in Finland is in the island of Replot at the west
coast at about 63º 07’ N. It is distributed about as
far north along the Swedish east coast, but it
reaches 70º N in Norway (Hultén 1971).
Allium oleraceum is regarded as an archaeo-
phyte in Finland (Suominen & Hämet-Ahti 1993,
Hämet-Ahti et al. 1998). Its connection with an-
cient and medieval settlement is obvious in some
areas, e.g. in the mainland part of the biogeo-
graphic province of Ab and in the province of Ta
(Seppänen & Rantanen 1986, Seppänen 1987,
1988, Silkkilä & Koskinen 1990). According to
Pettersson (1943) and Seppänen (1988), A. olera-
ceum has been used as a food, spice, medicinal and
cult plant during the first millennium A.D. and
during the medieval time. According to Nyman
(1868), tender verdure of A. oleraceum has been
used together with green cabbage. The young
leaves have been used as a spice in wintertime
(Hultén et al. 1958). Høeg (1976) mentions that it
has been cultivated next to the houses in Kvikne in
Norway, and thatit has been used as a spice in fish
soup in the same way as Allium schoenoprasum.
The aim of this study is to get a better knowl-
edge of the distribution of A. oleraceum in Fin-
land, its habitats and accompanying vascular plant
species.
Memoranda Soc. Fauna Flora Fennica 81:1–18. 2005

Material and methods
All voucher specimens of Allium oleraceum in the
Finnish herbaria (H, HFR, JYV, KUO, OULU,
TMP, TUR, TUR-A, VOA) were studied. During
our studies on the populations of Allium ole-
raceum, we searched for it throughout most of its
range in Finland. Further, we used written infor-
mation to complete the distribution map.
Many localities were visited according to the
information given on the labels of voucher speci-
mens. In a few cases, we were not able to find the
species, either because of imperfect information or
because the habitats were destroyed, usually by
overgrowth or construction of roads, etc. (cf. Hin-
tikka 2000). Other localities were found by
chance, often simply by visiting suitable looking
habitats. During these field studies, accompanying
vascular plant species (nomenclature according to
Hämet-Ahti et al. 1998) were annotated. Some in-
formation could also be gathered from the labels of
voucher specimens and from the literature.
Results
Distribution in Finland
The distribution of Allium oleraceum, according
to voucher specimens in Finnish herbaria (H,
HFR, JYV, KUO, OULU, TMP, TUR, TUR-A,
VOA), in 10 × 10 km National uniform grid
squares in Finland, is presented in Fig. 1. The dis-
tribution map was completed for those 10 × 10 km
squares from which only information in the litera-
ture was available (see Bonsdorff 1867, Norrlin
1871, Cajander 1902, Palmgren 1915–17, Idman
1927, Eklund 1929, 1958, Luotola 1931, Åberg
1933, Vieras 1935, Jalas 1951, Jalas & Vuorisalo
1956, Vaahtoranta 1959, 1964, Laine 1960, 1964,
2Hæggström & Åström • Memoranda Soc. Fauna Flora Fennica 81, 2005
Fig. 1. The distribution of the field garlic (Allium oleraceum) in Finland in 10 km × 10 km squares according to
voucher specimens in herbaria (H, HFR, JYV, KUO, OULU, TMP, TUR, TUR-A, VOA). n= voucher specimens
seen from the square; p= voucher specimens with inexact locality information; ¢= localities according to liter-
ature or the author CAH’s field notes; := introduced specimens in gardens; ?= uncertain report.

Parnela 1962, Kause 1972, Hakila & Kalinainen
1984, Haikonen 1986, Seppänen & Rantanen
1986, Jutila et al. 1996, Raikas 1998, Hering 1999,
Saarinen 2000).
Allium oleraceum is quite densely distributed
in the Åland Islands, in the archipelago of Ab and
the along the south coast. Many of the archipelago
and coastal stations are located along old sailing
routes (e.g. Pettersson 1943, Lemberg 1947, Saari-
salo-Taubert 1963, Kurtto & Helynranta 1998).
Another area with many finds is South Häme (Ta).
Quite many of the finds here are located along old
trade and travelling routes (Seppänen 1987).
Whereas 48 stations, of which 43 were ob-
served during the 1990s, are known in Helsinki
(Kurtto & Helynranta 1998), only two are known
in the neighbouring town of Vantaa north of Hel-
sinki (Ranta & Siitonen 1996). Both stations are so
close to two stations in Helsinki that they may be-
long to the same populations. Another anomaly is
that only one known station, in a cottage garden, is
known in the neighbouring town of Espoo west of
Helsinki, besides introduced specimens in the gar-
dens of both authors.
The oldest written information of Allium ole-
raceum growing in Finland is given by Til-Landz
(1683); his no. 369 “Porrum pratense” has been in-
terpreted by Wein (1930) as representing A. olera-
ceum. The oldest voucher specimens in the her-
baria were collected in N, Borgnäs, Laha, at the be-
ginning of July, 1851, by H. Nervander (H; cf.
Nylander 1852) and in N, Ingå, Fagervik, on Au-
gust 14, 1851, by E. Hisinger (H; cf. Hisinger
1855). There are a few voucher specimens col-
lected during the 19th century without an exact
year mentioned on the labels. Therefore, it is not
possible to judge whether some of these were col-
lected earlier than 1851. The exsiccate no. 1863,
collected by Harald Lindberg, has the date August
28, 1842 (cf. Lindberg 1944); this is a misprint as
the year ought to be 1942.
The number of collections of Allium olera-
ceum in the different biogeographical provinces
are presented in Table 1. The most intensive col-
lecting period was from 1980 onwards. The period
1960–1979 was, together with the two earliest pe-
riods (before 1900 and 1900–1919), the least in-
tensive. Although the species is most abundant
and frequent in the Åland Islands, the greatest
number of collections were made in the province
of Ab.
Habitats
The habitats of Allium oleraceum are mostly mod-
erately or strongly influenced by man and domes-
tic animals. The species is much more common in
areas with calcareous bedrock or soil than in acid
areas; in the latter the species is mostly lacking.
The habitats of Allium oleraceum in Finland
range from rock meadows and dry meadows via
Memoranda Soc. Fauna Flora Fennica 81, 2005 • Hæggström & Åström 3
Table 1. Collections of Allium oleraceum in the biogeographical provinces of Al, Ab, N, Ka, St, Ta, Sa, Oa and
Sb during different periods. The collections are according to the vouchers in Finnish herbaria (H, HFR, JYV,
KUO, OULU, TMP, TUR, TUR-A, VOA). One collection may comprise several voucher specimens, preserved
in one or a few herbaria, collected by the same person in the same place on the same day.
Period –1899 1900–1919 1920–1939 1940–1959 1960–1979 1980– Total
Biogeogr. province
Al 23 47 47 28 13 129 287
Ab 10 24 41 55 49 153 332
N23 24 20 51 23 27 168
Ka 00880723
St 32738326
Ta 8 5 13 16 10 38 90
Sa 0000033
Oa 00145515
Sb 0000101
Total per period 67 102 137 165 109 311 891

mesic meadows to deciduous stands with a closed
canopy, including very dense Corylus stands. The
species seems to be quite persistent in many of its
localities, as it still grows in places where voucher
specimens were collected even more than one hun-
dred years ago.
Rock meadows and dry meadows. These are the
main habitats in the southwestern archipelago
(Fig. 2) and south Finland. In the main islands of
Åland, Allium oleraceum is readily found in al-
most every village on rock meadows and dry
meadows next to farm houses and roads and in
pastures. In the eastern archipelago of Åland and
the Turku archipelago, the species grows chiefly in
the larger inhabited islands (cf. Eklund 1958, Skult
1960). It grows still in deserted islands, such as
Väderskär, the island of ’Stormskärs-Maja’, in Al,
Vårdö Simskäla.
Seashore meadows. The habitats for Allium ole-
raceum mentioned on the labels of voucher speci-
mens include seashore meadows. It has also been
collected on a sandy seashore, on a stony seashore
andonaFucus wrack.
Iron Age mounds, hilltop fortresses, medieval
castles and churches. Both in the Åland Islands
and on the Finnish mainland, Allium oleraceum is
strongly confined to Iron Age habitation. Espe-
cially on the mainland part of the province of Ab,
dry meadows on Iron Age mounds are very typical
habitats of Allium oleraceum. Ancient hilltop for-
tresses, e.g. Vanhalinna in Ab, Lieto (Fig. 3),
Sibbesborg in N, Sibbo and Hakoisten linna
(Hagaborg) in Ta, Janakkala have been known for
long for the occurrence of this species. It grows in
the vicinity of the ruins of the medieval castle of
Kastelholm in Åland and near the ruins of the
Bishop’s castle of Kuusisto (Kustö) in Ab, Kaa-
rina.
The 18th century Swedish sea fortress of Svea-
borg (Suomenlinna) off N, Helsingfors (Helsinki)
and the 19th century Russian fortress of Bomar-
sund on Åland harbour good populations of A. ole-
raceum on dry and rock meadows among the ram-
parts and ruins. It grows on rock and dry meadows
next to the medieval stone churches of Eckerö,
Föglö, Hammarland, Jomala, Kökar, Lumparland,
Saltvik and Sund in Åland. However, both on
Åland and the mainland of Finland, the church-
yards themselves are so well mown and weeded
that A. oleraceum is searched for in vain. The spe-
cies has not been found but near a few churches
outside the Åland Islands, for example next to the
churches of Ab, Halikko, Nådendal (Naantali) and
Tarvasjoki (locality destroyed in 2004); N, on the
ruins of the churchyard wall of the first church of
Helsinki; near the place of the old church of Oa,
Vähäkyrö. Allium oleraceum does not grow in the
churchyards of these churches, but on rock or dry
meadows in the vicinity. It has also been found
4Hæggström & Åström • Memoranda Soc. Fauna Flora Fennica 81, 2005
Fig. 2. A dense stand of Allium
oleraceum on a rock meadow.
Al, Saltvik: Kvarnbo. July 16,
2003. Photo: CAH.

next to 19th century churches, e.g. N, Hangö
(Hanko) and N, Sveaborg (Suomenlinna) off Hel-
sinki.
Medieval manors and farm houses harbour the
species; examples are Ab, Villnäs, Lemsjöholm
manor, Masku Ohensaari manor (first mentioned
in 1232), Pargas, Qvidja 15th century manor,
Sauvo-Karuna, Paddais manor (first mentioned in
1346) and Karuna manor (first mentioned in
1347), and N, Sjundeå, Svidja manor (first men-
tioned in 1420).
Roadsides and headlands. In its whole area
Allium oleraceum is frequently found in roadside
meadows. Especially in the Åland Islands, the spe-
cies is quite common on roadsides. Headland
meadows next to cultivated fields are also fairly
common habitats.
Gardens, park lawns and ruderal soils.Allium
oleraceum is often found in gardens of different
kind. The labels of voucher specimens mention
among others the garden of a vicarage, a few aban-
doned gardens, flower beds and the rock garden of
the gardener’s school of Lepaa in Ta, Tyrväntö
where it grew together with Allium carinatum L.
and A. scorodoprasum (H). The voucher speci-
mens of Ab, Lojo, Solhem were cultivated, ac-
cordingtothelabel(H).Allium oleraceum has also
been found on park lawns at manors and in towns.
At its only known station in Tb, Keuruu, Haapa-
mäki, it was collected in a shrubby place near the
railway station; the label mentions that the species
obviously has been originally planted (H). One
voucher was collected on a rock meadow in St,
Hämeenkyrö, Kyröskoski; the collector thought
that it probably was introduced because Sedum al-
bum grew in the same place (H). A voucher col-
lected in a garden in Oa, Kristinestad mentions that
it was moved from a natural stand at a roadside (H)
and another collected in Oa, Vasa, Brändö was
found on a new planted roadside meadow, the soil
of which was brought from Old Vasa (VOA).
There are also a few more or less ruderal finds,
e.g. on filling soil at carparks at the church in Al,
Sund (H) and in the ferry harbour of Al, Sottunga
(H). Allium oleraceum has obviously been intro-
duced by sea transports. It was find on ballast i N,
Pernå, Lassdal (H) and in the harbour of Brändö in
Oa, Vasa (H, VOA)
Memoranda Soc. Fauna Flora Fennica 81, 2005 • Hæggström & Åström 5
Fig. 3. The ancient hilltop fortress of Vanhalinna in Ab, Lieto. Allium oleraceum grew abundantly on almost all
dry and rock meadows around the crest of the hill. Aug 21, 2000. Photo: CAH.

Broad-leaved deciduous stands. During studies
of the wooded meadows of Nåtö island in the
1960s and 1970s (Hæggström 1983), Allium ole-
raceum was found by the author C.-A. H. with low
constancy (15%) and very low cover (0.1%) in
more or less open meadow (meadow quadrats) and
with rather high constancy (60%) and very low
cover (0.1%) in dense hazel thickets (Corylus
quadrats). Much denser stands of A. oleraceum oc-
cur, however, here and there in the luxuriant decid-
uous stands of Fraxinus excelsior, Acer plata-
noides, Ulmus glabra, Corylus avellana, etc., both
on Nåtö and on Espholm at the southern outskirts
of the town of Mariehamn (Figs. 4–5). Densities of
up to approximately 200 to 400 shoots per square
metre were annotated in late spring in both locali-
ties (Table 2). Later in the summer, most of the
shoots wilted due to the deep shade of trees and
shrubs. During the last few years, some of the
stands of Nåtö have been eaten by some animal,
presumably the roe deer (Fig. 6). In Espholm, the
stands have been heavily eaten back by sheep.
A few other stations with Allium oleraceum
have been found in dense woods in the Åland Is-
lands. Some tens of A. oleraceum specimens were
found in Föglö on the island of Bänö in 2001. They
6Hæggström & Åström • Memoranda Soc. Fauna Flora Fennica 81, 2005
Fig. 5. A dense stand com-
prising tens of square
metres of Allium oleraceum
grows in Espholm Nature
Reserve. As the first green
leaves of A. oleraceum de-
velop already during the
previous autumn, this spe-
cies is green early in the
spring, before other herbs
and grasses have devel-
oped their foliage. Dense
thickets of shrubs and some
trees were cut during the
autumn and winter of 1999–
2000 in Espholm. Most of
the ashes were then pol-
larded or repollarded. Al,
Mariehamn: Espholm, April
21, 2000. Photo: CAH.
Fig. 4. Allium oleraceum
forms a dense carpet on a
spot of 2–3 square metres
next to a hazel shrub.
Anemone nemorosa and a
few Primula veris (in front)
and Anemone ranunculoi-
des (in the background) are
in blossom. Al, Lemland:
Nåtö, 31 May, 1987. Photo:
CAH.

Memoranda Soc. Fauna Flora Fennica 81, 2005 • Hæggström & Åström 7
Table 2. Allium oleraceum and accompanying species in 1 m² quadrats in four sites in the Åland Islands, one in
Nyland and one in South Häme. Constancy and mean cover of all analysed quadrats of one site (= index sign,in
the Alandian sites only) are given as per cent. 0.5 per cent is marked with .5; a very low mean cover is marked
with .1.
A. Al. Lemland, Nåtö, dry meadows at Nåtö Biological Station, 16 quadrats, 4 analysed on 31.VII.1999,
12 analysed on 9.VI..2000 and 2 analysed on 15.VII.2003.
B. Al, Lemland, Nåtö, Själskatsudden, former closed canopy Fraxinus excelsior and Corylus avellana
stand, cleared in 1998, 8 quadrats analysed on 18.VII.1999.
C. Al, Lemland, Nåtö, Skarpbacka, dense Allium oleraceum stands under a closed deciduous tree canopy
(rather low mean cover, because of the date of analysis), 10 quadrats analysed on 4.VI.2000.
D. Al, Mariehamn, Espholm, former closed canopy deciduous wood, cleared in 1999, 12 quadrats
analysed on 9.VI.2000.
E. N, Hangö, Tvärminne, Storängsberget, rock meadows, 4 quadrats analysed in 25.VII.2000.
F. Ta, Janakkala, Hakoinen, hilltop fortress, rock meadows, 4 quadrats analysed in 9.VIII.2000.
Site A B C D E F
Trees and shrubs
Fraxinus excelsior 38820217.5
Corylus avellana 7536 10040
Ribes alpinum 13.5 17.5
Acer platanoides 407
Ulmus glabra 207
Prunus padus 174
Rosa dumalis ssp. coriifolia 8.5
Rosa majalis 25
Field layer
Allium oleraceum 1002100110034 1006100 100
Fraxinus excelsior, seedlings 17.1 100170.5 50.5
Geranium sylvaticum 1721005602676
Heracleum sibiricum 50210012 20.5 1008
Platanthera sp. 6.1 13.1 10.1 8.1
Ranunculus acris 17.5 50.5 10.1 33.5
Ranunculus auricomus coll. 61.5 63.5 60267.5
Polygonatum odoratum 88320.5 17.1 25
Poa trivialis 11.1 63.5 10.1
Taraxacum spp. 22.5 75110.1
Veronica chamaedrys 50238.1 601
Alchemilla glaucescens 17.5 50.5 8.1
Angelica sylvestris 6.1 13.5 8.1
Cerastium fontanum ssp. vulgare 33.5 50.1 8.1
Plantago lanceolata 83313.1 8.1
Primula veris 6.1 888503
Ranunculus ficaria ssp. bulbilifer 56540.5 751
Festuca rubra 44.1 8.1 25
Filipendula vulgaris 6.1 17.1 100
Achillea millefolium 83625 25
Agrostis capillaris 22275 100
Galium verum 61650 75
Potentilla argentea 22.5 25 100
Sedum acre 6.1 25 25
Anemone nemorosa 88210020 10020
Geum rivale 13.1 10.1 421
Hepatica nobilis 88310081004
Melica nutans 63.1 10.1 671
Listera ovata 13.1 10.1 504
Paris quadrifolia 382403581

8Hæggström & Åström • Memoranda Soc. Fauna Flora Fennica 81, 2005
Satureja vulgaris 13.1 10.1 251
Viola riviniana 75.5 30.5 42.5
Poa nemoralis 38.1 10.1 50
Campanula rotundifolia 11.5 13.1
Glechoma hederacea 17.1 13.1
Poa pratensis 56.5 13.1
Trifolium repens 28.1 25.1
Urtica dioica 6.1 13.1
Veronica serpyllifolia 6.1 25.1
Corydalis solida 11.1 1001
Alchemilla monticola 11.1 8.1
Anthoxanthum odoratum 61342.5
Avenula pubescens 7228.1
Dactylis glomerata 611584
Festuca pratensis 17.5 8.1
Filipendula ulmaria 11117.5
Hypericum maculatum 11.1 33.1
Poa angustifolia 39133.5
Rumex acetosa 7218.1
Allium schoenoprasum 17175
Allium scorodoprasum 33175
Arabidopsis thaliana 17.1 25
Sedum telephium ssp. maximum 33175
Vicia tetrasperma 11.1 25
Elymus repens 39.1 25
Trifolium arvense 281100
Stellaria graminea 11.5 25
Scleranthus annuus 6.1 100
Phleum pratense 6.1 50
Acer platanoides, seedlings 75160.5
Dentaria bulbifera 13.1 803
Lathyrus vernus 13.1 201
Moehringia trinervia 13.1 20.5
Carex pallescens 25.1 50.5
Convallaria majalis 5036712
Fragaria vesca 25.1 8.1
Luzula pilosa 13.1 8.1
Platanthera chlorantha 13.1 8.1
Polygonatum multiflorum 13.1 171
Sanicula europaea 13.1 8.5
Sorbus aucuparia, seedlings 38.1 25.5
Veronica officinalis 38.1 50
Deschampsia cespitosa 20.1 8.1
Gagea lutea 50.5 33.1
Geranium sanguineum 25250
Agrostis cf. gigantea 8.5 25
Satureja acinos 50 50
Viola tricolor 50 100
Allium vineale 6.1
Alopecurus pratensis 17.1
Anthriscus sylvestris 33.5
Arenaria serpyllifolia 28.5
Briza media 6.1
Bromus hordeaceus 17.5
Capsella bursa-pastoris 6.1
Cardamine hirsuta 6.1
Centaurea jacea 11.1
Cerastium glutinosum 6.1
Cerastium semidecandrum 6.1
Corylus avellana, seedling 6.1

Memoranda Soc. Fauna Flora Fennica 81, 2005 • Hæggström & Åström 9
Cynosurus cristatus 6.1
Dactylorhiza sambucina 6.5
Draba muralis 22.1
Erophila verna 6.1
Equisetum arvense 6.1
Fallopia convolvulus 6.1
Fragaria viridis 224
Galium album 17.5
Galium boreale 392
Geranium molle 11.1
Geranium pusillum 11.1
Lathyrus pratensis 281
Lolium perenne 6.1
Lotus corniculatus 6.1
Luzula campestris 11.1
Myosotis arvensis 6.1
Myosotis ramosissima 28.5
Pimpinella saxifraga 17.1
Plantago media 6.1
Platanthera bifolia 6.1
Ranunculus bulbosus 442
Saxifraga granulata 11.5
Sedum album 28.5
Trifolium medium 11 1
Trifolium pratense 281
Veronica arvensis 39.1
Vicia cracca 17.5
Vicia hirsuta 11.1
Adoxa moschatellina 13.1
Arabis hirsuta 13.1
Betula pubescens, seedlings 50.5
Carex digitata 38.1
Geum urbanum 632
Glechoma hederacea
Laserpitium latifolium 251
Maianthemum bifolium 381
Melampyrum sylvaticum 13.5
Orchis mascula 50.1
Picea abies, seedlings 13.1
Plantago major ssp. intermedia 25.1
Poa annua 381
Polygala vulgaris 13.1
Potentilla anserina 13.1
Prunella vulgaris 13.1
Ranunculus repens 13.1
Rosa sp., seedlings 13.1
Rubus saxatilis 252
Sagina procumbens 13.1
Viola mirabilis 631
Anemone ranunculoides 404
Campanula trachelium 10.1
Galium aparine 10.1
Ranunculus cassubicus 20.1
Agrimonia eupatoria 8.1
Carex flacca 25.5
Melampyrum nemorosum 50.5
Luzula multiflora 17.1
Melica picta 8.1
Potentilla erecta 8.1
Prunus padus, seedlings 8.1

grew in a dense stand of Alnus glutinosa, Fraxinus
excelsior and Malus sylvestris together with intro-
duced Allium ursinum about 350 m ENE of Erkas.
InKumlingeadenseCorylus stand with Betula
pendula pollards and Pinus sylvestris occur on a
fairly steep E slope at Lenviken. A lax stand of
A. oleraceum grows here together with quite many
shade tolerant field layer species, such as Anemone
nemorosa, Campanula trachelium, Dentaria bul-
bifera, Galium odoratum, Hepatica nobilis, La-
thyrus vernus, Listera ovata, Milium effusum, Poa
nemoralis and Viola riviniana.
The only other locality in Finland with dense
Allium oleraceum in a deciduous wood that has
come to our knowledge is the island of Rams-
holmen in N, Ekenäs. Here a patch of A. oleraceum
grow beneath Acer platanoides, Corylus avellana
and Ulmus glabra. The species has been collected
on Ramsholmen several times: 1909, 1943, 1956,
1957 and 1981 (H, TUR-A). At the road Ystads-
gatan in the town of Ekenäs, a stand of A. olerace-
um growns beneath shading Acer platanoides
trees.
In Ab, Pargas, Lofsdal, Lenholm, Allium ole-
raceum grows scattered in a deciduous woodland
with Quercus robur, Picea abies, Prunus padus,
etc. The Allium stands, are, however, not of the
same dense character as in, e.g. Nåtö and Espholm.
Some of the voucher labels report deciduous
woods or groves as habitats. However, only a few
of them mentions anything about the lignoses, e.g.
the edge of a black alder stand, a hazel stand and a
birch copse. The character of the stands of Allium
oleraceum is not reported.
10 Hæggström & Åström • Memoranda Soc. Fauna Flora Fennica 81, 2005
Ranunculus polyanthemos 17.1
Stachys sylvatica 25.5
Crepis tectorum 25
Cystopteris fragilis 25
Dryopteris filix-mas 25
Festuca ovina 25
Geranium robertianum 50
Hypericum perforatum 25
Poa compressa 25
Rubus idaeus 25
Thymus serpyllum 25
Woodsia ilvensis 25
Rumex acetosella 100
Arabis glabra 25
Linaria vulgaris 50
Leontodon hispidus 25
Lychnis viscaria 75
Fig. 6. A dense stand of
Allium oleraceum eaten by
some animal, supposedly
the roe deer. All the bulbils
and flowers were eaten. Al.
Lemland: Nåtö, Själskats-
udden, August 11, 1999.
Photo: CAH.

Accompanying species
More than 300 vascular plant species (taxa) have
been annotated growing in, or in the immediate vi-
cinity, of stands of Allium oleraceum in the bio-
geographical provinces of Al, Ab, N and Ta (Table
3). Most of them are typical of rock meadows and
dry meadows, such as Achillea millefolium,
Agrostis capillaris, Anthoxanthum odoratum,
Arabidopsis thaliana, Arabis glabra, Calama-
grostis epigejos, Campanula rotundifolia, Des-
champsia flexuosa, Dianthus deltoides, Festuca
ovina, F. rubra, Galium verum, Hypericum per-
foratum, Plantago lanceolata, Poa compressa,
Potentilla argentea coll., Rumex acetosella, Se-
dum acre, Stellaria graminea, Viola tricolor and
Woodsia ilvensis. Mesic meadow species are also
abundant: Anthriscus sylvestris, Centaurea jacea,
Dactylis glomerata, Geranium sylvaticum, Poa
pratensis, Ranunculus acris, Rumex acetosa, Ta-
raxacum spp. Trifolium medium, T. pratense and
Vicia cracca. Afew are typical of ruderal ground,
e.g. Elymus repens and Urtica dioica. Weeds oc-
cur in many localities, especially if surrounded by
arable fields: Barbarea vulgaris, Capsella bursa-
pastoris, Chenopodium album coll., Cirsium ar-
vense, Fallopia convolvulus, Galeopsis bifida,
Plantago major ssp. major, Poa annua and Vio la
arvensis.
In the southwestern provinces of Al and Ab,
the number of calciphilic species increases. Exam-
ples of calciphilic accompanying species are
Alchemilla glaucescens, Allium scorodoprasum,
A. vineale, Avenula pratensis, Briza media, Bro-
mus hordeaceus, Dactylorhiza sambucina, Draba
muralis, Filipendula vulgaris, Geranium san-
guineum, Heracleum sibiricum, Listera ovata,
Platanthera chlorantha, Primula veris, Ra-
nunculus bulbosus, Rubus caesius, Sedum album,
Seseli libanotis and Veronica spicata.
A few vouchers collected in Stmention accom-
panying species: Kokemäki, Kauvatsa, Viola tri-
color and Veronica longifolia (TUR); Pori,
Tahkoluoto, Avenula pubescens and Pimpinella
saxifraga (H); Pori (Ahlainen), Anttoora, Avenula
pubescens (H).
The following species are typical of the inland
province of Ta, e.g. Campanula glomerata, Cen-
taurea phrygia, C. scabiosa, Knautia arvensis,
Sedum annuum, Thalictrum simplex, Trifolium au-
reum, Trollius europaeus and Verbascum nigrum.
The only information available from the prov-
ince of Sb is on the label of a voucher specimen
collected in Mäntyharju: accompanying species
were Arabis glabra, Centaurea scabiosa, Di-
anthus deltoides and Sedum acre. The label of one
voucher specimen collected in Oa, Kristinestad
mentions Alopecurus pratensis and Avenula pu-
bescens.
Discussion
The most typical habitats of Allium oleraceum in
Scandinavia are dry meadows of different kind,
road side meadows and forest edges (Genberg
1977, Andersson 1981, Mascher 1990, Påhlsson
1994, Gustafsson 1996, Kraft 1996, Georgson et
al. 1997). The species occurs also in different de-
ciduous stands in Scandinavia (Lange 1864, Bor-
nebusch 1923, Almquist 1929, 1965, Jessen 1935,
Sterner 1938, 1986, Hultén et al. 1958, Weimarck
1963, Malmgren 1982, Gustafsson 1996, Georg-
son et al. 1997, Rydberg & Wanntorp 2001).
Palmgren (1915–17), who studied wooded
meadows of the Åland Islands (cf. Hæggström
1983), knew Allium oleraceum from 17 of his 20
special areas and from about 30 additional locali-
ties. The habitats of this species was according to
him (Palmgren 1915–17: 250) open meadows and
rock meadows (“ängsbackar, back- och berg-
knallar”). He did not refer to any locality with trees
and shrubs. Although the most typical habitats for
Allium oleraceum are treeless meadows, a few de-
ciduous stands with closed canopy are known both
in the Åland Islands and SW Finland.
In Central Europe, Allium oleraceum is a typi-
cal species of the vegetation class Festuco-Bro-
metea, which occurs on calcium rich soil (Ellen-
berg et al. 1991). In the Czech Republik, Allium
oleraceum frequently occurs in deciduous woods,
especially in oak-hornbeam woodland of the
Carpinion alliance (Duchoslav 2001a, 2001b).
Passarge (1985) described the Gagea pratensis –
Allium oleraceum association from northern Ger-
many.
According to Eklund (1929), Allium olera-
ceum is a hemeradiaphorous plant, but later he
(Eklund 1958) was of the opinion that it is some-
what hemerophilous; however, its distribution in
Memoranda Soc. Fauna Flora Fennica 81, 2005 • Hæggström & Åström 11

12 Hæggström & Åström • Memoranda Soc. Fauna Flora Fennica 81, 2005
Table 3. Vascular plant species often growing together with Allium oleraceum in the biogeographical provinces
of Al, Ab, N and Ta, according to the authors field notes, labels of vouchers and literature, especially Seppänen
(1989) regarding the province of Ta. * = frequent; × less frequent; .. not annotated; o = not known in the prov-
ince.The dependence on calcium (Calciphily) is chiefly according to Eklund (1946); Ca+++ = strictly limestone
dependent (German: kalkstet), Ca++ = strongly calciphilic (stark kalkhold); Ca+ = moderately calciphilic
(kalkbegünstigt).
Taxon Al Ab N Ta Calciphily
Achillea millefolium ****
Agrostis capillaris ****
Anthriscus sylvestris ****
Dactylis glomerata ****
Elymus repens ****
Festuca ovina ****
Festuca rubra ****
Galium verum ****
Juniperus communis ****
Potentilla argentea coll. ****
Rubus idaeus ****
Sedum acre ****Ca++
Stellaria graminea ****
Taraxacum spp. ****
Vicia cracca ****
Ranunculus acris **×*
Rumex acetosella **×*
Centaurea jacea **××
Poa angustifolia **××
Campanula rotundifolia *××*
Rumex acetosa *××*
Veronica chamaedrys *××*
Arabis glabra ×*×*
Hypericum perforatum ×*×*
Viola arvensis ×*×*
Agrimonia eupatoria *×××Ca+
Anthoxanthum odoratum *×××
Arabidopsis thaliana *×××
Calamagrostis epigejos *×××
Cerastium fontanum ssp. vulgare *×××
Geranium sylvaticum *×××
Plantago lanceolata *×××Ca+
Artemisia vulgaris ×*××
Trifolium medium ×*××
Carex spicata ××*×
Viola tricolor ×××*
Cystopteris fragilis ××××Ca++
Deschampsia flexuosa ××××
Fragaria vesca ××××
Galium album ××××
Geum urbanum ××××Ca+
Poa compressa ××××Ca++
Poa pratensis ××××
Polygonatum odoratum ××××
Thymus serpyllum ××××
Trifolium pratense ××××
Trifolium repens ××××
Urtica dioica ××××
Woodsia ilvensis ××××Ca++
Sedum telephium ssp. maximum ***o
Galium boreale **..*

Memoranda Soc. Fauna Flora Fennica 81, 2005 • Hæggström & Åström 13
Pimpinella saxifraga **..*
Filipendula vulgaris * * × .. Ca+
Lathyrus pratensis **..×
Lychnis viscaria .. * × *
Allium schoenoprasum *××..Ca+
Vicia tetrasperma *××..
Allium scorodoprasum * × × o Ca++
Bromus hordeaceus * × .. × Ca++
Hypericum maculatum * × .. ×
Trifolium arvense * × .. ×
Veronica arvensis * × .. ×
Alopecurus pratensis × * .. ×
Phleum pratense × * .. ×
Arenaria serpyllifolia × × .. * Ca++
Dianthus deltoides ××..*
Ranunculus polyanthemos ××..*Ca+
Scleranthus annuus ××..*
Verbascum nigrum ..××*
Saxifraga granulata ×××..Ca+
Melica nutans ××..×
Pilosella officinarum ××..×
Silene nutans ××..×Ca+
Verbascum thapsus ××..×
Myosurus minimus × .. × × Ca++
Satureja acinos .. × × × Ca+++
Avenula pubescens **....
Vicia hirsuta *....*
Angelica sylvestris * × .. ..
Heracleum sibiricum * × .. .. Ca++
Primula veris * × .. .. Ca+
Sedum album * × .. .. Ca++
Avenula pratensis * × o .. Ca++
Ranunculus bulbosus * × .. o Ca++
Veronica spicata * × .. o Ca++
Allium vineale * .. × o Ca++
Geranium sanguineum *..×oCa+
Geranium molle * .. .. × Ca++
Myosotis arvensis .. * × ..
Campanula glomerata × .. .. * Ca++
Silene vulgaris .. × .. *
Origanum vulgare ××....Ca+
Artemisia campestris × .. × .. Ca++
Erophila verna ×....×Ca+
Gagea lutea ×....×Ca+
Glechoma hederacea ×....×
Ranunculus ficaria ssp. bulbilifer ×....×Ca
Satureja vulgaris ×....×
Seseli libanotis × × o o Ca+++
Fallopia convolvulus ..××..
Fallopia dumetorum ..××..
Tanacetum vulgare ..××..
Tripleurospermum inodorum ..××..
Alchemilla glaucescens * .. .. .. Ca++
Anemone nemorosa *......
Briza media * .. .. .. Ca++
Hepatica nobilis *......Ca+
Listera ovata * .. .. .. Ca++
Myosotis ramosissima *......Ca+
Platanthera bifolia *......
Potentilla reptans * .. .. .. Ca++

the western part of the Turku archipelago and east-
ern part of the Åland Islands was fairly unaffected
by the human culture. Jessen (1935) was of the
opinion that Allium oleraceum is not especially
culturally influenced in its distribution in Den-
mark. In Malmgren’s (1982) study area, Väst-
manland in central Sweden, the species is clearly
hemerophilous. According to Sterner (1986), it is
very common on cultural soil and ruderal habitats
on the limestone island of Öland. It is slightly fa-
voured by the culture in Halland in western Swe-
den (Georgson et al. 1997). In our opinion the spe-
cies is strongly hemerophilous in Finland; very
few localities represent anything that could be
called natural vegetation.
According to Eklund (1946), Allium olera-
ceum is moderately calciphilic (German: kalk-
begünstigt) in SW Finland. Our studies confirm
this in most of the range of this species in Finland.
In the Scandinavian countries, it occurs mainly in
areas with calcium rich habitats and is absent or
rare in areas with acid bedrock (Jessen 1935,
Weimarck & Weimarck 1985, Sterner 1986, Lid &
Lid 1994). According to Ellenberg et al. (1991),
the species occurs in Central Europe on weakly
acid to weakly basic soil, but never on strongly
acid soils.
Hegi (1909) discerned four varieties and one
forma of A. oleraceum in Central Europe:
–var.angustifolium Koch; plant rather tall,
stout, with round, not flat leaves; widespread
–var.alpicola Aschers. & Graebner; plant short,
rarely taller than 30 cm, leaves round, with
rough margins, flowers rose coloured; in
mountains
–var.complanatum Fries; plant tall, stout, leaves
flat, up to 4 mm broad, groved, margins only
weakly rough
–var.pauciflorum Aschers. & Graebner; with
only 2–6 flowers
–f.sterile Beck; without flowers.
AccordingtoClaphamet al. (1987) var. compla-
natum is confined to the north of British Isles.
Malmgren (1982) reports on a stand with very
broad (7–9 mm) leaves growing at an open road-
side. According to him, flat leaves and luxuriant
growth run together. Afew voucher specimens in
the Finnish herbaria collected in the Åland Islands
areassignedtovar.complanatum.Malmio&
Erkamo (1951) also reports of a find of A. olera-
ceum f. complanatum in Ab, Uusikaupunki. Ac-
cording to Hylander (1953), broad leaves occur in
specimens growing in shade. According to our ob-
servations, specimens growing in moist and nutri-
ent rich habitats, such as ditches and manured
flower beds, are tall, stout and broad-leaved. In our
opinion such specimens represent only modifica-
tions.
According to Almquist (1929, 1965), the
Allium oleraceum is flowerless and wilts early
when growing in shady localities in the Uppsala
area. According to Malmgren (1982), the light de-
mand amplitude is large in this species, but it does
not flower in deep shade. Rydberg & Wanntorp
(2001) report that A. oleraceum grows abundantly
in shady deciduous stands and shore woods in
14 Hæggström & Åström • Memoranda Soc. Fauna Flora Fennica 81, 2005
Platanthera chlorantha *....oCa+
Rubus caesius * .. .. o Ca+++
Draba muralis * .. o o Ca++
Arabidopsis suecica .. .. .. *
Centaurea phrygia .. .. .. *
Centaurea scabiosa .. .. .. * Ca++
Cuscuta europaea .. .. .. *
Knautia arvensis .. .. .. *
Myosotis stricta .. .. .. *
Sedum annuum .. .. .. * Ca++
Silene latifolia ssp. alba .. .. .. *
Trifolium aureum .. .. .. *
Trollius europaeus .. .. .. *
Veronica verna .. .. .. *
Thalictrum simplex .. .. o * Ca+
Gagea minima ×......Ca+
Dactylorhiza sambucina × .. o o Ca++

Södermanland, Sweden, and then without flowers.
Our studies confirm this: in deep shade below a
closed deciduous tree canopy, very few specimens
develop flowers and most of the specimens wilt
quite early in the season. Thus the taxon called f.
sterile Beck (Hegi 1909) is but a modification. The
var. pauciflorum Aschers. & Graebner, with only
2–6 flowers, is a modification, too, because the
number of flowers vary in one population due to
environmental influence, e.g. drought or shade. In
our studies on populations of A. oleraceum,the
proportion of flowerless specimens and the num-
ber of flowers in flowering specimens vary from
one year to another in the same population
(Åström & Hæggström, in prep.).
In Västmanland, central Sweden, Allium ole-
raceum is unknown in grazed areas (Malmgren
1982). However, Malmgren remarks that it
occurrs in a lawn as a weed which is impossible to
eradicate. It seems to be extremely hardy regard-
ingbothgrazingandscythingintheÅlandIslands.
In Nåtö, as an example, a part of the Nature Re-
serve have been grazed, first by sheep in 1965–
1966 and 1969–72 and by cattle since 1974
(Hæggström 1983, 1990) and the stands of Allium
oleraceum have not diminished. The meadows at
NBS have been scythed regularly since 1965. The
stands of A. oleraceum comprise hundreds of
plants in vigorous condition. Since the beginning
of the 1990s, some species rich spots at NBS have
been left to develop freely until late scything in late
August or even in September.
Allium oleraceum grows in the Botanical Gar-
den of Helsinki University. It was first annotated
there by Brenner (1906) and then collected in 1915
(TMP), 1954 (H) and 1974 (H). When we tried to
find it, we thought that there was only one possible
habitat for this species, namely the lawn around a
small rock outcrop next to the old main building.
The sward was, however, mown by a lawn-mower
every year during at least the last two decades. A
few square metres were fenced in the spring of
2001 and in the summer there were about 500 well
developed specimens growing inside the fenced
area.
Kujala & Ulvinen (1964) noticed that speci-
mens on the island of Kuorsalo in South Karelia
were larger and had more flowers but fewer bulbils
than specimens in the islands of Tammio and
Ulko-Tammio farther out in the Gulf of Finland.
This is most probably due to the fact that the plants
on Kuorsalo were pentaploid (2n = 40), while on
Tammio and Ulko-Tammio tetraploid (2n = 32; cf.
Åström & Hæggström 2004).
Only few studies of accompanying species are
known to us. Almquists (1929) studies in Uppland
west of the Åland Islands has three vegetation
analyses of dry and rock meadow vegetation and
three of deciduous stands. The accompanying spe-
cies are largely the same as in Åland.
Acknowledgements. We are indebted to the keepers of the
herbaria for loan of voucher specimens. Mrs. Eeva
Hæggström, M. Sc. and Mr. Thomas Kuusela assisteddur-
ing some of the field work.
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