Ann. Bot. Fennici 37: 197–206
Helsinki 27 September 2000
ISSN 0003-3847
© Finnish Zoological and Botanical Publishing Board 2000
Taxonomic status of Saussurea alpina subsp.
esthonica (Asteraceae): phenetical analysis
Anneli Narits, Malle Leht & Jaanus Paal
Narits, A. & Paal, J., Institute of Botany and Ecology, University of Tartu, Lai Str. 40,
Tartu 51 005, Estonia
Leht, M., Institute of Zoology and Botany, Estonian Agricultural University, Riia Str.
181, Tartu 51 014, Estonia
Received 13 April 1999, accepted 8 December 1999
Narits, A., Leht, M. & Paal, J. 2000: Taxonomic status of Saussurea alpina subsp.
esthonica (Asteraceae): phenetical analysis. — Ann. Bot. Fennici 37: 197–206.
Morphological variation of Saussurea esthonica Baer ex Rupr. and S. alpina (L.) DC.
s. str. was studied with different multivariate methods. The most important characters
supporting the clustering of the specimens into groups are the shoot hight and characters correlated to it, and hairiness of leaves, while the type of trichomes in both taxa is
the same. Hence, these characters depend very much on ecological conditions and are,
therefore, taxonomically not reliable enough. It appeared also that several specimens
were morphologically intermediate between typical representatives of S. esthonica and
S. alpina s.str. Therefore, we consider it more appropriate to treat these taxa as ecogeographical subspecies: S. alpina subsp. esthonica (Baer ex Rupr.) Kupff. and S. alpina
subsp. alpina. Chromosome numbers established for S. alpina subsp. esthonica are
2n = 52, 54.
Keywords: chromosome numbers, Estonia, Latvia, morphology, multivariate methods,
Saussurea alpina, Saussurea esthonica
INTRODUCTION
The genus Saussurea comprises over 300 species
distributed mostly in Asia (Bremer 1994) where,
in the Himalayas and high mountains of China,
its primary center of diversity lies (Lipschitz
1979). The genus is represented in Europe by three
to nine or more species, depending on the species
concept used. According to Hegi (1954), only
three species occur in Europe: S. alpina (L.) DC.,
S. discolor (Willd.) DC. and S. pygmaea (Jacq.)
Spreng. Lipschitz (1976) added to the list six species found only in the European part of Russia
and in the Ukraine. The list becomes even longer,
if some subspecies of S. alpina are recognized as
species.
Saussurea alpina s. lato is a polymorphic arctic-montane species with a disjunct distribution
area in Europe and Asia (Hultén & Fries 1986).
The only representative of the genus in Estonia
198
and Latvia is S. esthonica Baer ex Rupr., a taxon
that has been defined either as a species (Lipschitz
1962, 1979, Üksip 1966, Kuusk 1978, Czerepanov
1995) or a subspecies of S. alpina (Kupffer 1902,
Hegi 1954, Lipschitz 1976, Jalas 1980). The first
data about S. esthonica was recorded by Ruprecht
(1845), who mentioned that a new Saussurea taxon was found from central Estonia by K. E. von
Baer in 1844 and described it as a new species.
An improved description of S. esthonica was published by Meyer (1855). In 1902, Kupffer found
that the differences between S. esthonica and typical arcto-alpine S. alpina were not large enough
to justify recognition of two species, and recognized it as a subspecies, S. alpina ssp. esthonica
(Baer ex Rupr.) Kupff.
At that time the nearest known localities of
Saussurea alpina s. lato were in Russia in the Arkhangelsk and Olonets provinces, and therefore the
taxa seemed to be geographically well separated.
Later, localities in the Leningrad and Novgorod
districts were found, plants from there identified as
S. alpina s. str. (Juzepchuk 1955, Sokolovskaya
1965). Hence, Lipschitz (1962) stressed that several specimens were morphologically intermediate between S. alpina s. str. and S. esthonica, and
some were so close to S. esthonica that they could
even be placed in that taxon. And since a race of S.
alpina, S. subbendorffii Herd., which is morphologically very similar to S. esthonica, was found
growing in East Siberia and Jakutia, Lipschitz has
expressed the opinion that S. esthonica is not a local Estonian endemic. According to Laasimer
(1965), S. esthonica originated from mires of the
late glacial period and belongs in the group of plants
reaching their present localites from the south.
Ingelög et al. (1993) assume that S. esthonica is a
neoendemic of the Baltic Sea region.
Saussurea alpina s. lato shows variation especially in the length of the stem, in the shape,
size, pubescence and margin of the leaves, in inflorescence type, and in the shape and pubescence
of the involucre. The recorded chromosome numbers and ploidy levels of S. alpina s. lato are also
quite variable: 2n = 26 (Rostovtseva 1983, Krogulevitsch & Rostovtseva 1984, Lid & Lid 1994),
2n = 36 (Ishikawa 1916, Lid & Lid 1994), 2n =
48 (Sokolovskaya & Strelkova 1948), 2n = 51–
52 (Favarger 1965), 2n = 52 (Löve & Löve 1948,
Lavrenko et al. 1991, Lid & Lid 1994), 2n = 54
Narits et al. •
ANN. BOT. FENNICI 37
(Sokolovskaya & Strelkova 1960, Morton 1977,
Uotila & Pellinen 1985, Lid & Lid 1994), 2n =
72–76 (Rostovtseva 1979), 2n = 76? (Lid & Lid
1994). Chromosome numbers of S. esthonica have
not been counted.
The aims of the present study were to compare the morphological variation of Saussurea alpina s. str. and S. esthonica and to answer the
following questions:
1. How well can these two taxa be distinguished
from the statistical point of view?
2. What is the chromosome number of S. esthonica?
3. What is the taxonomic status of S. esthonica?
4. Is S. esthonica an endemic taxon?
MATERIAL AND METHODS
Morphological variation of Saussurea esthonica
was studied from 187 plant specimens collected
in 1996 from eight Estonian and two Latvian populations, and from 35 specimens (two populations)
from the Herbarium of the University of Tartu
(TU) (Table 1, Fig. 1). For studying S. alpina s.
str., 43 specimens were collected in 1996 from
two Norwegian populations, 79 were obtained
from Krakow from the Herbarium of the Polish
Academy of Sciences (KRAM), and 54 specimens
from the Herbarium of the Norwegian University
of Science and Technology (TRH). The material
from KRAM originated mostly from the Tatra
Mountains, but also included single specimens
collected from Finland, Sweden, Switzerland and
western Russia (Novgorod region); all the specimens from KRAM were treated conditionally as
representatives of one population. The material
from TRH originated mainly from the counties of
Sør-Trøndelag and Nord-Trøndelag. Altogether,
222 specimens of S. esthonica and 176 specimens
of S. alpina s. str. were studied using the 26 macromorphological characters most often considered
in the diagnoses of S. alpina (Table 2). The characters were measured on herbarium material with
a binocular microscope MBS-2 and a ruler.
Micromorphological characters were studied
with a scanning electron microscope (SEM) in the
Centre for Materials Research of Tallinn Technical University. The epidermis from both sides of
ANN. BOT. FENNICI 37
•
Taxonomic status of S. alpina subsp. esthonica
199
Fig. 1. Distribution of Saussusrea alpina subsp. esthonica in Estonia and Latvia. Studied localities:
Tammiku (1), Paraspõllu
(2), Niitvälja (3), Halinga
(4), Pärnu-Jaagupi (5), Varangu (6), Kalevi (7), Ambla (8), Hageri (9), Viirika
(10) in Estonia and Pope
(11), Tukums (12) in Latvia.
Table 1. Material studied. Site types of Saussurea esthonica named according to Paal (1997). Site types of S.
alpina s. str. according to Nordhagen (1943).
—————————————————————————————————————————————————
No.
Locality
Site type
Specimens
—————————————————————————————————————————————————
S. esthonica
01. Estonia, Harjumaa county, Tammiku
Spring fen
21
02. Estonia, Harjumaa county, Paraspõllu
Spring fen
20
03. Estonia, Harjumaa county, Niitvälja
Rich paludified grassland
20
04. Estonia, Pärnumaa county, Halinga
Rich paludified grassland
20
05. Estonia, Pärnumaa county, Pärnu-Jaagupi
Molinia site type
20
06. Estonia, Lääne-Virumaa county,
Rich paludified grassland
21
07. Estonia, Jõgevamaa county, Kalevi
Rich paludified grassland
20
08. Estonia, Järvamaa county, Ambla (TU)
Rich paludified grassland
21
09. Estonia, Harjumaa county, Hageri (TU)
Rich paludified grassland
11
10. Estonia, Raplamaa county, Viirika
Spring fen
21
11. Latvia, Pope
Rich paludified grassland
5
12. Latvia, Tuckums county, Tuckums
Rich paludified grassland
20
S. alpina s. str.
13. Norway, Sør-Trøndelag county, Oppdal commune, Kongsvoll
Rich, tall-herb birch forest,
30
(trees had been cut) 900 m
14. Norway, Oppland county, Dovre commune, Hjerkinn
Minerotrophic, baserich
13
lawn mire, 1000–1200 m
15. Poland, Tatra mountains
1000–1200 m
82
16. Norway, Sør-Trøndelag and Nord-Trøndelag county
180–750 m
54
—————————————————————————————————————————————————
200
one central cauline leaf, the epidermis of involucre, and the surface of the stem of one specimen
from each population were examined on air dried
herbarium material coated with gold.
For chromosome counts, seeds from three
plants from each of the six Estonian populations
were collected. Seeds were stratified for three
months and germinated at room temperature on
moistened filter paper in Petri dishes. Chromosome counts were made from root tips pretreated for 3–4 hours at 15 °C with 0.002% 8-hydroxyquinoline, fixed in Farmer fixative, macerated in 1N HCl at 60 °C for 13–14 minutes, stained
with 1% acetoorcein and squashed in acetic acid
and glycerin (1:1). The material collected (incl.
Narits et al. •
ANN. BOT. FENNICI 37
voucher specimens) is preserved in the Herbarium
of the Institute of Zoology and Botany (TAA).
Data processing
To avoid overweighting, characters strongly depending on ecological conditions in the (micro)habitat — shoot height, length of inflorescence
and length of the lowest peduncle — were standardized by range before data processing,
xi´= (xi – xmin)/(xmax – xmin),
(1)
where xmin and xmax are the minimum and maximum values of the character.
Table 2. Characters used.
—————————————————————————————————————————————————
No.
Symbols
Character
Type
—————————————————————————————————————————————————
01
HSH/R
Height of shoot (cm) standardised by range
metric
02
NLSH
Number of leaves on shoot
count
03
NH
Number of heads (incl. those not flowering yet)
count
04
LI
Length of involucre (cm)
metric
05
WI
Width of involucre (cm)
metric
06
PB
Spot of branching (distance between the topmost inflorescence to
ordinary
the peduncle of the lowest inflorescence): 1 (< 5 cm), 2 (5–10 cm),
3 (15–20cm), 4 (25–30cm)
07
NB
Number of branches
count
08
NHSH
Number heads on the shoot
count
09
LIF/R
Length of inflorescence (cm) standardised by range
metric
10
LLP/R
Length of the lowest peduncle (cm) standardised by range
metric
11
LFBL
Length of the fifth basal leaf (cm)
metric
12
WFL
Width of the fifth leaf (cm)
metric
13
BLL
Base of lower leaves: 1: shortly narrowed, 2: slowly narrowed,
ordinary
3: unnarrowed, 4: clasping
14
LUL
Length of the topmost leaf (cm)
metric
15
WUL
Width of the topmost leaf (cm)
metric
16
SC
Stem colour: 1: green, 2: bright red, 3: dark red
ordinary
17
IC
Involucre colour: 1: bright, 2: edges dark, 3: dark
ordinary
18
NTBL
Number of teeth of the fifth basal leaf
count
19
HUPB
Hairiness of upper part of branch: 1: glabrous, 2: few hairs,
ordinary
3: hairy, 4: very hairy
20
HLPB
Hairiness of lower part of branch: 1: glabrous, 2: few hairs,
ordinary
3: hairy, 4: very hairy
21
HI
Hairiness of involucre: 1: glabrous, 2: few hairs, 3: hairy,
ordinary
4: very hairy
22
HUSBL
Hairiness of upper side of the fifth basal leaf: 1: glabrous, 2: few hairs,
ordinary
3: hairy, 4: very hairy
23
HLSBL
Hairiness of lower side of the fifth basal leaf: 1: glabrous, 2: few hairs,
ordinary
3: hairy, 4: very hairy
24
LI/WI
ratio of length and width of involucre
ratio
25
LUL/WUL
ratio of length and width of the topmost leaf
ratio
26
LFBL/WFL
ratio of length and width of the fifth basal leaf
ratio
—————————————————————————————————————————————————
ANN. BOT. FENNICI 37
•
Cluster analysis was performed with the program package SYN-TAX 5.02 (Podani 1993). The
method of incremental sum of squares (MISSQ)
with the distance for mixed data (Podani 1994)
was used.
For the calculation of common statistics,
Spearman rank correlation coefficients between
characters were used, and for the evaluation of the
importance of characters within clusters by
classifactory discriminant analysis, the SAS program package (SAS Institute Inc. 1996) was utilised.
Ordination was carried out with principal component analysis (CANOCO package, version 3.1,
Ter Braak 1990; and CANODRAW package, version 3.0, Smilauer 1992) with default parameters
settings.
For the estimation of the adjacency of clusters, the distances of all specimens, or operational
taxonomic units (OTUs), from all centroids (except the cluster to which the OTU belongs) were
calculated according to the postulate that the jth
cluster is interpreted as being adjacent to the ith
cluster if the distance between at least one of the
OTUs of the ith cluster and the centroid of the jth
cluster is smaller than the distance to the centroids
of all the other clusters (Paal & Kolodyazhnyi
1983, Paal 1994):
min D(xi, mk) = D(xi, mj),
k, k ≠ i
(2)
where xi is the vector of OTU xi, mk and mj are
vectors of the kth and jth cluster’s centroid, D =
distance (resemblance) function.
To visualise the transitions between the two
empirical taxa-clusters S. esthonica and S. alpina
s. str., and to show the distribution of specimens
located between the centroids of the clusters considered, the split window method (Parzen 1962)
was applied, as in Paal et al. (1998). The density
of the OTUs projection probability distribution
on a straight line passing through the centroids of
both clusters was calculated as
p( x ) =
201
Taxonomic status of S. alpina subsp. esthonica
1 n 1 x – xi
∑ Φ
n i =1 h h
parameter or window breadth, n is the number of
OTUs in the cluster, and xi is the projection of the
ith OTU on the line. The density of the normal
distribution was regarded as the window function.
The smoothing parameter h was calculated according to the formula
1
h = 2 s 0.05 +
n
where s is the standard error of the projection.
In order to measure the degree of distinctness
of clusters, the α-criterion (Duda & Hart 1976)
was used.
2 I2
–
1 –
πd I1
α=
8
21 – 2 nd
π d
(5)
I1 = ∑ xi – m
(6)
where
where p(x) is the distribution density in the point
x, Φ is the window function, h is the smoothing
2
x ∈X
2
I2 = ∑ ∑ xi – mi
2
(7)
i =1 x ∈X
I1 is the sum of square distances between the
centroid of a united complex of two clusters, I2 is
the sum of square distances between the sample
plots and their cluster centroids after dividing the
complex into two suboptimal parts, xi is vector of
OUT xi, m is vector of the centroid of the united
complex, mi is vector of the cluster X centroid, d
is dimensionality of the united complex, d = min(q,
n – 1), where q and n are the number of characters
and specimens in the united complex, respectively.
To acquire a better interpretation of the estimates, it is more convenient to use correponding
probabilities called coefficients of indistincness
(CI) instead of the direct values (Paal 1987, 1994):
∞
CI =
(3)
(4)
x2
100
exp
– dx
2
2π α∫
(8)
The last two analyses were made by the original SYNCONT 3.0 program composed by S. Kolodyazhnyi, J. Paal and A. Kink.
202
Narits et al. •
ANN. BOT. FENNICI 37
Fig. 2. Dendrogram of
Saussurea alpina subsp.
esthonica and S. alpina s.
str. cluster analysis. The
hierarchical levels discussed in the text are
marked with Roman numbers and letters.
RESULTS
Chromosome counts
Chromosome numbers were counted in six Estonian spontaneous populations of Saussurea esthonica. Populations in Niitvälja, Viirika, Paraspõllu
and Kalevi yielded 2n = 52, in the Pärnu-Jaagupi
population the number was 2n = 54 and in the
Tammiku population it could not be precisely determined 2n = 30(?).
Clustering
Cluster analysis grouped the data at a quite high
level into two unequal clusters (Fig. 2). Cluster I
includes the specimens estimated conventionally
as belonging to Saussurea esthonica. The cluster
branches further into two subclusters, Ia and Ib,
consisting of representatives of different S. esthonica populations. Cluster II also branches further
into two subclusters. Subcluster IIa comprises
putative intermediate specimens identified as
S. esthonica or S. alpina s. str., while in cluster
IIb, specimens of S. alpina s. str. prevail.
Also, the principal component analysis demonstrates that the whole sample consists of two
groups corresponding to Saussurea alpina s. str.
and S. esthonica, with several specimens on intermediate positions (Fig. 3). According to the first
axis, the characters with the highest scores are the
ratio of length and width of the topmost leaf (0.96),
height of the shoot (0.64), the width of the topmost leaf (–0.59), the ratio of length and width of
the fifth basal leaf (0.55), and the number of
branches (0.53). Characters with the highest scores
by the second axis are: the number of heads (0.83),
the number of branches (0.61), the length of inflorescense standardised by range (0.55), the
length of the lowest peduncle (0.45) and the ratio
of the length and width of the topmost leaf (0,45).
Conventionally estimated species-clusters of
S. esthonica and Saussurea alpina s. str. were mutually distinct (with coefficient of indistincness
CI = 0.0) (Fig. 4), while several pairs of population-clusters were not. Further, the indistinct populations were stepwise joined together until four
distinct groups were constituted: cluster 1 includes
all populations of S. esthonica, cluster 2 contains
specimens from the Kongsvoll population (SørTrøndelag, Norway) and from TRH (Sør-Trøndelag and Nord-Trøndelag, Norway), cluster 3 is
formed by specimens from the Hjerkinn population (Oppland, Norway) and cluster 4 by specimens from the Tatra mountains (Poland).
ANN. BOT. FENNICI 37
•
Taxonomic status of S. alpina subsp. esthonica
203
Characters
SEM studies showed that hairs of specimens from
both taxa, Saussurea esthonica and S. alpina s. str.,
as well as their stomata, are of the same type. Leaves
and stems are covered with long, simple, smooth
unicellular thin hairs and multicellular thick hairs,
multicellular glandular trichomes also occured.
Hairs on involucres are smooth, thin and curved.
Differences between the two taxa appeared only in
the density of hairs: plants of S. alpina are much
more hairy. The upper surface of the leaves of S.
esthonica is either glabrous or with single hairs,
the lower surface of some S. esthonica specimens
is also nearly glabrous while the surfaces of S.
alpina are always covered with trichomes.
According to the F-criterion of discriminant
analysis, the most important characters for distinguishing the species are: height of the stem, hairiness of lower surface of the fifth leaf, spot of branchTable 3. Importance of characters according to
classifactory discriminant analysis. F-criterion significance level for all character is below 0.001, except
BLL (p = 0.012), SC (p = 0.013) and LUL (p = 0.099).
Symbols as in Table 2.
————————————————————————
Characters
F-criterion
————————————————————————
HSH/R
828.86
HLSBL
299.47
LUL/WUL
223.59
PB
214.83
LLP/R
148.94
LI/WI
141.61
HUSBL
139.25
LIF/R
127.56
HLPB
125.38
LFBL
122.11
NB
118.57
WUL
117.84
HUPB
108.01
NLSH
77.53
IC
50.47
LFBL/WFL
45.44
WI
41.09
NHSH
40.02
WFL
37.63
NLSH
32.70
LI
27.53
HI
14.48
NTBL
13.80
BLL
6.30
SC
6.21
LUL
2.74
————————————————————————
Fig. 3. Classification polygons of conventionally estimated specimens of Saussurea esthonica and S.
alpina s. str. superimposed on to PCA ordination. The
first component counts for 48.6% of the variation and
the second component for 17.1%.
ing, number of branches, length of inflorescence
and length of the lowest peduncle (Table 3).
According to Spearman’s rank correlation coefficient most of the characters are well correlated
with each other, uncorrelated are only the colour
of the stem and the shape of the leaf base. Length
of inflorescence is highly correlated with length
of the lowest peduncle (r = 0.93), height of the
shoot (r = 0.75), spot of branching (r = 0.81), number of branches (r = 0.79) and number of heads
(r = 0.66). Number of branches is also strongly
correlated with spot of branching (r = 0.64) and
number of heads (r = 0.77). These characters constitute a quite dense correlation group, others stand
more or less separately.
Mean height of Saussurea alpina s. str. plants
(26.6 cm) is smaller than the height of the smallest S. esthonica specimens (Table 4). The highest
S. esthonica plants grow in Pärnu-Jaagupi (mean
74.0 cm), and the smallest in the Hageri population (44.9 cm). The most hairy plants belong to
the Kongsvoll population (Norway) and the most
glabrous ones to the Pärnu-Jaagupi population
(Estonia).
204
Narits et al. •
ANN. BOT. FENNICI 37
Fig. 4. Specimens projection probability distribution
according to the split window method. The curve
marked with dots portrays
the empirically estimated
Saussurea esthonica specimens, the curve shown
with dash-and-dot line corresponds to the S. alpina
s. str. specimens. The continuous curve above them
represents the joint sample specimens projection
probability distribution. E
and A mark the centroids
of S. esthonica and S. alpina s. str., respectively;
the dotted lines perpendicular to the line through
the cluster centroids delimits the transition zone.
DISCUSSION
According to Favarger (1965), the basic chromosome number in the genus Saussurea is x = 13.
Thus, S. esthonica is a tetraploid (2n = 52, 54)
with a tendency towards aneuploidy.
Lipschitz (1976) states that the polymorphism
of West-European Saussurea alpina s. lato is partly the result of hybridization with S. discolor. Therefore, S. esthonica could be an allotetraploid which
has arisen from hybridization between diploids
of S. alpina and S. discolor in the overlapping part
of their distribution areas, while later, in the Post
Glacial period, as Laasimer (1965) supposed, it
spread northwards to Latvia and Estonia. However, taking into account that distances between
S. discolor and S. esthonica areas are large, and
that there exists a Siberian taxon S. stubendorffii
morphologically very close to S. esthonica (Lipschitz even considered uniting them), which for
geographic reasons cannot have arisen from the
hybridization of S. alpina and S. discolor, we prefer the idea of the endemic origin of S. esthonica
by the doubling of the chromosomal set of dip-
Table 4. Mean ± standard deviation of quantitative characters. Symbols as in Table 2.
—————————————————————————————————————————————————
No.
Character
Saussurea alpina
Saussurea esthonica
—————————————————————————————————————————————————
01
HSH
26.63 ± 9.78
64.36 ± 11.91
02
NLSH
11.67 ± 2.94
15.44 ± 4.27
03
NH
6.96 ± 2.93
10.00 ± 5.41
04
LI
1.09 ± 0.09
1.17 ± 0.17
05
WI
0.88 ± 0.14
0.77 ± 0.18
07
NB
2.84 ± 1.35
4.89 ± 1.78
08
NHSH
2.85 ± 1.04
2.09 ± 0.91
09
LIF
2.47 ± 1.87
7.69 ± 5.16
10
LLP
1.58 ± 1.10
5.22 ± 3.61
11
LFBL
10.12 ± 2.56
14.13 ± 3.65
12
WFL
1.82 ± 0.57
1.44 ± 0.62
14
LUL
4.04 ± 1.29
3.87 ± 2.02
15
WUL
0.59 ± 0.32
0.26 ± 0.23
18
NTBL
13.0 ± 2.06
14.3 ± 3.89
—————————————————————————————————————————————————
ANN. BOT. FENNICI 37
•
Taxonomic status of S. alpina subsp. esthonica
loid S. discolor. This process may result in a population able to inhabit localities new in comparison with those of its diploid predecessors (Grant
1981). As the maximum elevation in the Baltic
region is only 317 m, S. esthonica occurs here in
conditions quite different from those of the arctoalpine habitats of typical S. alpina: on swampy
meadows, wooded meadows, in shrubland and in
mires (Kuusk 1978).
Morphologically the specimens clustered
clearly into two distinct clusters corresponding to
empirically identified Saussurea esthonica and
S. alpina s. str. MISSQ clustering and PCA revealed the existence of intermediates between typical S. esthonica and S. alpina s. str. specimens
(Figs. 2 and 3).
The most important characters in the conventional delimitation of Saussurea esthonica and
S. alpina s. str. were the height of the stem (even
despite of standardising this parameter by range)
and the hairiness of lower side of the fifth basal
leaf (Table 3). Both these characters depend greatly on ecological conditions — light, humidity, etc.
Therefore, it is to be expected that S. esthonica,
which grows in moist places among high grasses
or shrubs, is taller than S. alpina s. str. growing in
mountains in well lit and less humid sites. From
the taxonomical point of view height is not considered a reliable character in delimiting taxa. In
some cases even definite correlation between the
height of the plant and its geographical origin occurred — specimens of more northern origin being considerebly lower than those from further
south (Rousi 1971). Leaves of S. alpina are more
hairy than leaves of S. esthonica, while their hairs
are of the same type. Density of hairs is also a
character that depends on habitat conditions: in
dry and open windy places plants are more hairy
than those growing in shady moist habitats (Rousi
1965). Thus, morphological data taken from specimens growing in different habitats should be used
with great care (Øvstedal 1998).
Therefore, since the most important characters supporting the clustering the material into
groups are taxonomically not reliable enough and
numerous specimens with intermediate characters
exist, we cannot recognize these taxa as independent species. Hence, as they are geographically
more or less separated and grow in different ecological conditions, we consider it more appropri-
205
ate to treat these taxa as eco-geographical subspecies: Saussurea alpina subsp. esthonica (Baer
ex Rupr.) Kupff. and S. alpina (L.) DC subsp. alpina. For more detailed studies growth experiments
would be useful — comparison of the morphology of taxa, which originally inhabit ecologically
different sites, after cultivation in ecologically
identical conditions, could help to decide about
the reliability of characters: do they depend on
the habitat conditions or not.
ACKNOWLEDGEMENTS: The authors thank Dr. Eli Fremstad
who kindly collected material from two Norwegian populations and supplied us with the material from TRH. We
also thank Dr. Viesturs Òulcs from the Institute of Biology,
Latvia, for his assistance in collecting material from Latvia.
The authors are grateful to Prof. Urve Kallavus for helping
with SEM studies, Ms. Silvia Sepp for helping with SAS
and to Dr. Vello Jaaska and Ms. Silvia Sepp for critical
reading of the manuscript. The English was revised by Mr.
Ilmar Part. The study was supported by the Estonian Science Foundation (grants 2218, 3574 and 0552).
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