Polish Botanical Journal 61(1): 65–71, 2016
DOI: 10.1515/pbj-2016-0004
SAXIFRAGA AIZOIDES (SAXIFRAGACEAE) IN UKRAINE
Yuriy Kobiv
Abstract. Saxifraga aizoides L. is endangered in the Ukrainian Carpathians. It became extinct in the Chyvchyny Mts and its
only remaining metapopulation in the Chornohora Mts is in gradual decline. The paper gives the exact location, morphological
characters, population parameters and habitat conditions of the species in the Chornohora locality, and briefly discusses the
main threat factors.
Key words: Carpathians, distribution, endangered species, extinction, peripheral population, Saxifraga aizoides
Yuriy Kobiv, Institute of Ecology of the Carpathians, National Academy of Sciences of Ukraine, Kozelnytska St. 4, 79026 Lviv,
Ukraine; e-mail: ykobiv@gmail.com
Introduction
Saxifraga aizoides L. is an arctic-alpine species with a mostly amphi-Atlantic distribution
(Hultén & Fries 1986). In Europe, in addition
to the northern part of its range, the species has
a disjunct distribution in the Pyrenees, Alps, Apennines, Carpathians, and mountains of the Balkan
Peninsula (Fig. 1). Its Central European localities are confined primarily to high elevations in
the subalpine, alpine and subnival zones (up to
3000 m a.s.l.), with some stations much lower in
the mountains or even Prealpine lowlands (Meusel
et al. 1965; Hultén & Fries 1986; Lutz et al. 2000).
Saxifraga aizoides is a calcicole and mainly hygrophytous. It is restricted mostly to cold-water
springs and riparian habitats or wet rocks. It is
considered a pioneer species, which colonizes open
sites with vegetation-free gravel (Raffl et al. 2007).
In the Carpathians the species occurs in all
three (western, eastern, southern) parts of the
mountain system in Poland, Slovakia, Ukraine
and Romania (Webb & Gornal 1989; Jalas et al.
1999). It is red-listed as endangered in Ukraine
(Kobiv 2009a). This paper is intended to clarify
the distribution of S. aizoides in the Ukrainian
Carpathians and to describe its population parameters, in order to establish level of threat to this
species in the region and to provide the baseline
for further monitoring.
Material and methods
I analyzed previous publications and herbarium data on
Saxifraga aizoides before surveying its known localities in situ.
The results are from field research carried out in 1996,
2000 and 2014 in the Chornohora Mts and in 2012 in
the Chyvchyny Mts, part of the Ukrainian Carpathians.
Biometric data were acquired from 20 plants in situ; all
measurements and descriptions refer to live plants.
Fig. 1. Distribution of Saxifraga aizoides L. in Europe, according to Meusel et al. (1965), Hultén and Fries (1986) and
Jalas et al. (1999).
66
POLISH BOTANICAL JOURNAL 61(1). 2016
Fig. 2. Location of the metapopulation of Saxifraga aizoides L. in the Chornohora Mts. I–III – the largest subpopulations.
Population density was estimated from a set of 10
square plots, demarcated by randomly placing a 0.5 ×
0.5 m2 or a 0.1 × 0.1 m2 frame ten times in different
parts of a given subpopulation (Rabotnov 1964; Harper
1977). For species abundance, Table 2 uses Braun-Blanquet’s (1965) scale. Exact site locations were determined
in the WGS-84 system with a Garmin eTrex Global
Positioning System (GPS) navigator (10 m accuracy).
Calcium and magnesium content in soil was measured
complexometrically, and pH(H2O) electrometrically
(Arinushkina 1970). Vascular plant nomenclature follows Mirek et al. (2002), and bryophyte nomenclature
follows Hill et al. (2006).
Results
distribution in ukraine
In Ukraine, Saxifraga aizoides occurs only in the
Carpathians, where its two localities are known.
One was reported by Pawłowski (1948: 21) in the
Chyvchyny Mts, referring to Mądalski’s finding of
the species in the region. It is confirmed by two
sheets collected in 1935 and 1936, stored in the
Herbarium of the W. Szafer Institute of Botany,
Polish Academy of Sciences, Kraków, in Mądalski’s
personal collection (KRAM 48673–74). The labels
indicate that it occurred at 1300–1323 m a.s.l. ‘on
a high limestone cliff located in spruce forest on
the left bank of the southern stream which flows
from Polonina Gropa on Mt. Hnytesa’.
My examination of that site in June 2012 indicated that the species does not occur there any
more. The location of the site is certain because
the mentioned limestone cliff is easily found, and
the presence of other rare species [Saussurea discolor (Willd.) DC., Saxifraga corymbosa Boiss.]
that J. Mądalski collected from the same cliff in
the 1930s confirms that it is the right locality.
Its geographic coordinates are 47°44′11.9″N,
24°54′07.1″E. My survey of the mentioned stream
and nearby ones on limestone bedrock over a considerable range of elevation did not reveal any
occurrence of S. aizoides in that area at present.
Another locality is at Polonina Gadzhyna in
the Chornohora Mts (Fig. 2). It was first reported
by Zapałowicz (1889), who stated that S. aizoides
occurs there at 1570–1620 m a.s.l. Several herbarium samples have been collected since then.
Y. KOBIV: SAXIFRAGA AIZOIDES (SAXIFRAGACEAE) IN UKRAINE
It is now the only remaining locality of the species in Ukraine; the data given below refer to the
Chornohora population.
morphological characteristics
of the chornohora plants
Saxifraga aizoides is an evergreen clonal perennial. Stems prostrate, branching, radicant, ca 1 mm
in diameter, with ascending flowering and vegetative shoots in their distal parts. Proximal (>1-yearold) creeping parts of stems leafless, black, with
sparse adventitious roots. Vegetative shoots 2.5–
8.0 cm high, flowering shoots 5.5–13.5 cm high,
densely foliate. Leaves alternate, fleshy, sessile,
persistent, 4–16 × 1.5–3.0 mm, oblong-lanceolate
or linear-lanceolate, acute, apiculate or obtusate.
Leaf surface lucid, glabrous. Margins entire, with
occasional firm spinose hairs, often on the tips.
Flowering stems with short glandular indumentum.
Inflorescence a lax cyme, with 3–13 pentamerous
flowers. Receptacle dark green, 2 mm high. Sepals
free, greenish yellow, glabrous, glandular, broadly
ovate, obtuse, 4.0–5.5 × 3.5–4.5 mm. Petals bright
yellow with numerous pink spots, obovate or elliptic, longer than calyx, 4.5–6.5 × 2–3 mm in
size. Stamina 10, as long or slightly longer than
petals. Ovary semi-inferior, glabrous; carpels 2,
broadly ovoid, greenish yellow, partly fused, flattened, with divergent 2.5–3.0 mm long styles and
globose stigmas. Seeds brownish, oblong-ellipsoid,
ca 1.0 × 0.5 mm in size.
population and habitat characteristics
Saxifraga aizoides forms one metapopulation in
the Chornohora Mts, consisting of several subpopulations of different size. They are confined
67
to the northeastern and eastern aspects of a steep
conglomerate rocky face and its bottom (Fig. 2). At
present the whole metapopulation forms a ca 200 m
long intermittent chain of subpopulations restricted
to a narrow range of elevation (1560–1575 m
a.s.l.). In 1996 and 2000 the species distribution
was significantly wider. Several outlying mats of
it occurred much higher: at 1610 m a.s.l. on the
banks of a small tarn in the lower part of a postglacial cirque and on moist sites on its moraine
slope. Zapałowicz (1889) stated that at the end of
the 19th century the species occurred as high as at
1620 m a.s.l. there.
There are three large subpopulations covering
>10 m2 each, and ca 10 smaller and more or less
isolated clusters of clones scattered in their vicinity. Easternmost subpopulation I is the largest. It
stretches along a cold stream (Fig. 3) falling from
a steep (60–70°) rock and forms a strip confined to
the right bank now. At present the strip is ca 25 m
long and its width does not exceed 6 m. In 1996
and 2000 the species occurred on both sides of
the stream along a much longer strip (ca 40 m).
Regression of the main population has been noted
since 2000 at that locality (Table 1); it shows the
most rapid decrease in the number of seedlings
and flowering shoots. In consequence, the role of
seed reproduction in population replenishment has
declined. Due to that site’s NNE aspect, strong
shading from shrub vegetation, and proximity to
the stream, microclimatic conditions there are the
coldest among all the sites.
Another large subpopulation, the central one
(II), is on a gentle slope at the base of a rock face
ca 120 m east of subpopulation I. The species
occurs next to a cold-water spring on a ca 15 ×
5 m patch of gravel scree. The habitat is almost
Table 1. Dynamics of population characteristics of Saxifraga aizoides L. in subpopulation I in the Chornohora Mts.
Parameter
Density of flowering shoots per m²
Density of vegetative shoots per m²
Ratio of flowering to vegetative shoots
Density of seedlings per m²
Year
2000
2014
332 ± 18
791 ± 52
0.42
2.42 ± 1.90
178 ± 11
635 ± 35
0.28
0.65 ± 0.05
68
Fig. 3. Saxifraga aizoides L. on a stream bank in the Chornohora Mts.
unshaded and is exposed to insolation for a large
part of the day. The density of flowering shoots of
S. aizoides is the highest there (687 ± 42 per m2).
The next large easternmost subpopulation (III)
is confined to another spring ca 70 away from the
central one (II). This subpopulation, which covers
ca 15 m2, is situated on a mossy open patch among
subalpine scrub.
In all subpopulations, individual plants of
S. aizoides form lax mats resulting from intensive
vegetative proliferation. Annual clonal growth
varies from 2.5 to 12.0 cm (mean 8.7 ± 0.4 cm).
Repent, radicated parts of old shoots persist for
several years and can reach up to 40 cm in length.
The average size of the mats is ca 15 × 10 cm,
and the largest of them cover 0.25–0.50 m2. Such
mats are formed by one clone or several interwoven clones. The density of flowering shoots
in the mats is 4.2 ± 0.3 on average but can reach
12 shoots per 100 cm2. The ratio of flowering
to vegetative shoots ranges from 0.12 to 2.10
(mean 0.35).
POLISH BOTANICAL JOURNAL 61(1). 2016
Apparently the metapopulation is maintained
mostly by vegetative reproduction. Seed-related
replenishment takes place as well; there are some
seedlings and immature plants at the locality.
The spatial contour of the Chornohora metapopulation is not constant: some small outlying mats
may appear or die off ca 50–100 m away from the
above-described large subpopulations. They establish in suitable wet microhabitats by seed dispersal
from parental subpopulations. Production of viable
seeds within a clone does not necessarily require
allogamy, because self-pollination is common in
S. aizoides (Raffl et al. 2007).
Fragments of mats can be pulled away by water
flow and established on the bank downstream;
I observed this in subpopulation I and it has also
been reported from the Alps (Jenny-Lips 1948).
In the Chornohora Mts, Saxifraga aizoides is
restricted to Cratoneuro-Saxifragetum aizoidis
hygrophytous riparian or spring communities
belonging to the Cratoneurion commutati alliance (Matuszkiewicz 2001) with high abundance
of bryophytes (Table 2). This association is not
mentioned in Malynovski and Kricsfalusy’s
(2002) prodrome of high-mountain vegetation
of the Ukrainian Carpathians, as no phytosociological relevés were ever made at the locality
earlier.
The measured soil parameters in the habitats of
the large subpopulations do not vary significantly
and are as follows: pH(H20) 5.94–6.08; calcium
content 7.91–8.32 mg-eq./100 g, and magnesium
content 3.50–3.72 mg-eq./100 g. The calcium
content of the conglomerate bedrock is not high;
such a level of calcium in substrate is atypical for
S. aizoides habitats.
Discussion
Localities of Saxifraga aizoides in the Ukrainian
Carpathians are confined to the northeastern limit
of the Central European (alpine) part of the species
range; its populations are peripheral (Fig. 1). The
nearest known localities are in the Marmarosh and
Rodna Mts in the Romanian Carpathians, not far
from the Ukrainian border, at 1475–1850 m a.s.l.
(Zapałowicz 1889; Răvăruţ 1956).
Y. KOBIV: SAXIFRAGA AIZOIDES (SAXIFRAGACEAE) IN UKRAINE
69
Table 2. Floristic composition at localities of the largest subpopulations of Saxifraga aizoides L. in the Chornohora Mts.
Number of subpopulation
I
II
III
Date
Area of relevé (m²)
Elevation (m a.s.l.)
Aspect
Inclination (º)
Cover of shrub layer (%)
Cover of herb layer (%)
48°06′49.3″N
24°34′21.9″E
10.07.14
25
1560–1575
NNE
65
15
30
48°06′46.2″N
24°34′22.2″E
10.07.14
25
1560
NE
7
5
40
48°06′45.9″N
24°34′22.5″E
10.07.14
10
1565
NE
10
10
30
Cover of moss layer (%)
30
30
40
3
4
3
2
3
1
2
1
1
3
+
.
+
2
1
.
1
2
1
+
1
1
+
+
.
.
1
2
1
2
1
+
.
+
.
+
1
+
+
2
+
1
2
.
2
+
.
2
1
1
1
Geographic coordinates:
DAss. Cratoneuro-Saxifragetum aizoidis
Saxifraga aizoides
ChCl. Montio-Cardaminetea
Brachythecium rivulare
Bryum pseudotriquetrum
Epilobium alsinifolium
Other species
Alchemilla glabra
Alnus viridis
Caltha laeta
Carex sempervirens
Deschampsia caespitosa
Heliosperma quadridentatum
Ligusticum mutellina
Pedicularis verticillata
Salix bicolor
Swertia punctata
Veratrum album
Campylium stellatum
Rhytidiadelphus triquetrus
Sporadic species: Bartsia alpina III(+), Cardamine pratensis subsp. rivularis I(+), Cirsium waldsteinii III(2), Crepis paludosa
III(1), Juncus triglumis I(+), Rumex alpinus II(+), Salix silesiaca III(1), Tussilago farfara III(+), Viola biflora I(+).
The Ukrainian locality of S. aizoides in the
Chyvchyny Mts was at an elevation (ca 1300 m
a.s.l.) atypically low for the Eastern Carpathians,
representing the lowermost limit of the species
distribution in the region. Thus, its extinction at
that locality may be due to climate change, in addition to past logging. Climate-induced extinctions
of small populations of arctic-alpine species at the
limits of their geographical and ecological ranges
have been noted in different mountain systems
(Lesica & McCune 2004; Marcysiak 2010). In
the 20th and early 21st centuries, the significant
decline or extinction of the following arctic-alpine
species have also been recorded in the Ukrainian
Carpathians: Dryas octopetala L., Hedysarum
hedysaroides (L.) Schinz. & Thell., Pedicularis
oederi Vahl, P. verticillata L., Oxyria digyna (L.)
Hill, and Saxifraga oppositifolia L. (Kobiv 2009b,
2011, 2013, 2014).
Another factor apparently contributing to the
shrinking of the Chornohora population of Saxifraga aizoides is the gradual natural replacement
of open patches of vegetation with shrubs [Alnus
viridis (Chaix) DC., Salix spp.], resulting from
a significant reduction of grazing at that locality.
Similar secondary successional changes tending
70
toward restoration of primary vegetation after massive abandonment of traditional land uses (grazing
and mowing) are taking place across vast tracts
in mountains all over Europe (MacDonald et al.
2000), including the Carpathians (Kuemmerle
et al. 2008). In the Ukrainian Carpathians this
process threatens many herbaceous, mostly lowgrowing plants occurring at the edge of their range
and confined to grasslands or other open communities, such as Erigeron atticus Vill., Erysimum
witmannii subsp. transsilvanicum (Schur) P. W.
Ball., Festuca rupicola subsp. saxatilis (Schur)
Rauschert, Minuartia verna subsp. oxypetala
(Woł.) G. Halliday, Potentilla thuringiaca Link,
Sempervivum marmoreum L., Silene zawadzkii
Herbich, Thlaspi dacicum Heuff. and Th. kovatsii
Heuff. (Kobiv 2014).
The locality of Saxifraga aizoides in the Chornohora Mts is not included in the reserve zone
of Carpathian National Nature Park. Because the
intensity of grazing at the site is very low recently,
it is not a question of limiting pastoral pressure to
prevent the further decline of its metapopulation.
On the contrary, post-anthropogenic succession
triggered by the decrease of grazing, as well as climate warming, are the main threats to the species.
Saxifraga aizoides is endangered in Ukraine.
It has become extinct in the Chyvchyny Mts. Its
only remaining metapopulation in the Chornohora
Mts is in decline. The rate of that decline is slow,
however, so its complete extinction at that locality
cannot be expected in the mid-term perspective.
Acknowledgements. I am grateful to the anonymous
reviewers for helpful remarks and suggestions on the
manuscript. This publication was prepared and published
with financial support from the National Academy of
Sciences of Ukraine (grant no. 0113U001435) and the
Ministry of Education and Science of Ukraine (grant
no. 112U001296).
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Received 2 June 2015