Boreal environment research 17: 219–236
issn 1239-6095 (print) issn 1797-2469 (online)
© 2012
helsinki 29 June 2012
vascular plants on the islands and peninsulas of maloe more
(lake Baikal): patterns of diversity and species turnover
victor v. chepinoga1), vitali e. Zverev2), elena l. Zvereva2) and
mikhail v. Kozlov2)*
1)
2)
Department of Botany and Genetics, Irkutsk State University, 1 Karl Marks Str., Irkutsk 664003,
Russia
Section of Ecology, Faculty of Biology, FI-20014 University of Turku, Finland (*corresponding author’s
e-mail: mikoz@utu.fi)
Received 19 Apr. 2011, final version received 10 Aug. 2011, accepted 1 Aug. 2011
chepinoga, v. v., Zverev, v. e., Zvereva, e. l. & Kozlov, m. v. 2012: vascular plants on the islands and
peninsulas of maloe more (lake Baikal): patterns of diversity and species turnover. Boreal Env. Res.
17: 219–236.
Unique biota of the Lake Baikal region face many threats due to increasing human activities. We documented spatial patterns in diversity of vascular plants, explored effects of
natural (bird colonies) and human-induced (tourism) disturbances on species richness
of semi-desert and steppe-desert plant communities of 12 islands and 4 peninsulas, and
estimated species turnover within a 30-year period. Floras of surveyed islands/peninsulas
contained 9 to 143 species; species–area relationship followed the power law model. Species richness did not change between 1979 and 2009, but the proportion of ruderal species
doubled during this period. Mean relative turnover rate was 1.17% of species per year. The
islands with large bird colonies had lower species richness than the islands with small or
no colonies. Imposing restrictions on tourist visitation to at least three islands (Zamogoj,
Khubyn and Khunuk) is a feasible way to conserve substantial part of regional biodiversity.
Introduction
Lake Baikal is unique in many characteristics,
including its size, location, quality of water, and
geological history (Moore et al. 2009). When it
was included in the World Heritage list, the need
for the research and monitoring activities of the
lake was specifically stressed by the International Union for Conservation of Nature (UNEP
2006). Still, we were unable to locate any recent
study exploring spatial patterns and temporal
changes in plant communities near the Baikal
shoreline, although regional diversity of vascular
plants is reasonably well documented (Zarubin
et al. 2005, Chepinoga et al. 2008).
Editor in charge of this article: Jaana Bäck
Island ecosystems are favourite objects of
ecological research due to their unique biological features (Whittaker 2007). First data
on vegetation of Olkhon Island, the largest of
Lake Baikal islands, were collected in 18th century (Galazii and Molozhnikov 1982). Ushkanji
Islands were first explored by botanists in 1914
(Sukachev and Poplavskaya 1914), and several
of Maloe More islands — only in 1978 (Petrochenko 1987). To our knowledge, flora of the
remaining islands (different sources list 6 to 47
islands in Lake Baikal; Gusev 1974) have not
been explored.
Pollution is generally seen as the largest
threat to biodiversity of Lake Baikal and its
220
Chepinoga et al.
•
Boreal env. res. vol. 17
Fig. 1. landscapes of lake Baikal islands: (a) malyi tojnak island, (b) northern part of Zamogoj island, (c) gull
colony on the eastern part of oltrek island and view on Borga-Dagan island, and (d) semi-arid plant community on
the southern part of Zamogoj island.
watershed (Sansom 2004, Moore et al. 2009).
The most acute problems are associated with the
pulp and paper mill in Baikalsk (Tretyakova and
Bazhina 2000, Voinikov et al. 2008) and with
contamination of the Selenga river flowing into
Baikal (Khazheeva et al. 2008). However, rapidly increasing tourism (from nearly 50 000 visitors in 2000 to nearly 250 000 in 2004; Rosabal
and Debonnet 2005) has been imposing substantial pressure on terrestrial ecosystems. This in
particular concerns the shores of the Maloe More
area, a shallow part of Lake Baikal between its
northwestern shore and Olkhon Island, which
hosts the largest number of natural objects determining the recreational value of the region.
Untouched and pristine nature is the principal tourist attraction of Lake Baikal. Islands
of the Maloe More area (Figs. 1 and 2) are
located close to the mainland, and are perceived
by the visitors as natural beauties. However at
present, tourism activities are implemented in a
disorganised and uncontrolled way, thus creating problems of disturbance and pollution and
often damaging important natural areas (Rosabal
and Debonnet 2005, Markova et al. 2008). In
2005–2006 more than 70 localities were used
for tourism, totalling about 700 ha impacted by
housing and beach activities (Romanova 2007).
Vegetation along several dozens of kilometres of
the shoreline is severely disturbed (Markova et
al. 2008, and pers. obs.).
The primary goals of our study were to
document spatial patterns in diversity of vascular
plants, explore effects of natural (bird colonies)
and human-induced (tourism) disturbances on
richness of semi-desert and steppe-desert plant
communities of 12 islands and 4 peninsulas,
and estimate species turnover on a subset of
five islands within a 30-year period. We use our
results to assess the conservation value of Maloe
More islands and to develop recommendations
for protection of regional biodiversity.
Boreal env. res. vol. 17 • Flora of the Baikal islands
221
Fig. 2. the map of the
study area. 1 = tojnak
island, 2 = malyi tojnak
island, 3 = Bolshoi tojnak
island, 4 = Khunuk island,
5 = sarminskaya Peninsula, 6 = Khubyn island,
7 = shara-Dagan island,
8 = oltrek island, 9 =
Borga-Dagan island, 10
= Zamogoj island, 11 =
Ujuga Peninsula, 12 =
nameless Peninsula, 13
= nameless Peninsula, 14
= Kharantsy island, 15 =
modoto island, 16 = edor
island. insert: position of
the study area within lake
Baikal.
Material and methods
Plant sampling
Study area
We visited the study area on 31 July–3 August
2009. Surveys of vascular plants were conducted
simultaneously by all of us (four persons). On
small islands and on all peninsulas sampling
was discontinued individually by each collector
when she/he decided that the chances of locating
previously unrecorded species were minor. In
practice, we attempted to stop searching when
no new species were recorded during the last 5
minutes. However, practical constrains forced
us to allocate fixed time for surveying the largest
islands (Khubyn, Oltrek, and Zamogoj); for sampling effort (person-hours of work) see Table 1.
One of us (VVC, the expert in regional flora)
recorded common species using pre-printed
forms, and collected only those specimens, identification of which required laboratory investigation. Three other collectors sampled aboveground parts of each species seen on their way;
these samples were identified by VVC on the day
of collection. Species found by each collector
were recorded separately. Materials collected in
the course of this work are deposited in the Herbarium of the Irkutsk State University (IRKU).
Plant nomenclature follows Chepinoga et al.
(2008); plant attribution to endangered or ruderal
species follows Zarubin (2001) and Chepinoga
et al. (2008), respectively.
Lake Baikal is located in southern Siberia. The
study region (Fig. 2), situated within an area
of approximately 53°02´–53°15´N and 106°45´–
107°27´E, belongs to the Irkutsk Oblast of
Russia. Maloe More is about 70 km in length
and covers about 800 km2 between the mainland
and Olkhon Island (the largest island of Lake
Baikal). This area includes 13 rocky islands and
two alluvial islands. In this paper, we use transliterations of Russian geographical names from
the most detailed map available to us (East Siberian Aerial Land-Surveying Enterprise 2007).
Climate of the Maloe More region is arid,
with annual precipitation of 230 mm or less.
Annual temperature is –1.2 °С; frost-free period
lasts 110–127 days. This climate is typical for
dry steppe regions. Additionally, small islands
are exposed to strong winds, up to 40 m s–1
in autumn (Ladeishchikov 1977). Consistently
with low level of precipitation and strong wind
impact, islands of Maloe More area are treeless
(except for Zamogoj Island; Fig. 1b); dominant
plant communities (Fig. 1d) are classified as various kinds of steppes (Petrochenko 1987). Both
1979 (the year when Yu. N. Petrochenko conducted his surveys) and 2009 (when we surveyed
the islands) were slightly warmer than average.
222
Table 1. characteristics of study areas, sampling efforts, and observed and estimated species richness of vascular plants.
characteristics of study areas
no. name
long. eb
is
is
is
is
Pns
is
is
is
is
is
Pns
Pns
Pns
is
is
is
53°02´02´´
53°04´24´´
53°04´40´´
53°05´11´´
53°05´39´´
53°05´48´´
53°09´07´´
53°09´37´´
53°09´46´´
53°10´38´´
53°09´11´´
53°09´24´´
53°08´59´´
53°14´04´´
53°14´09´´
53°14´41´´
106°46´02´´
106°49´38´´
106°50´05´´
106°51´39´´
106°52´04´´
106°56´31´´
106°58´09´´
106°59´21´´
106°59´59´´
107°06´26´´
106°57´32´´
106°56´57´´
106°56´24´´
107°24´31´´
107°26´26´´
107°26´39´´
max.
area (m2) Distance recreation Birds
elevation
(km)e
(m)c
[20]
[15]
12
1.2
0.7
33
[15]
35
[15]
77
[20]
1.5
[5]
12
5
[20]
6120
6575
65150
6110
5260
87110
4255/100d
134500
5020
490100
14455
20
2095
42800
1655
3240/2000d
0.30
0.86
1.45
1.00
0
0.34
0.67
1.35
1.49
2.63
0
0
0
0.17
0.27
0.96
0.50
0.25
0.50
0.50
0.25
0
0
0.75
0
0.25
2
0
1.75
0.75
0
0
0
0.25
0.50
0.25
0.25
0.25
1.75
0.25
2
0.25
0.25
0
0
1
0.75
2
3.0
3.0
4.0
2.0
2.0
4.0
0.2
5.7
0.5
8.0
2.0
0.1
2.0
3.0
1.0
1.0
4
4
4
4
4
4
1
4
3
4
4
1
4
4
4
4
estimated
species
richnessh
T
Sobs
Q1
Q2
Sjack Schao
165
160
198
181
157
240
9
335
29
385
226
20
173
205
71
37
64
53
80
70
55
89
9
141
13
143
98
20
70
83
28
12
19
8
24
16
13
21
9
45
3
37
33
20
17
27
9
3
9
5
18
17
7
18
0
25
4
26
24
0
23
16
5
0
85
79
61
58
104
92
84
76
69
64
109
98
–
–
189 171
15
14
181 163
131 115
–
–
84
75
111 100
38
34
16
15
types of sampled areas: is = island; Pns = peninsula. b Geographical co-ordinates refer to central parts of the sampled areas. c height data in brackets are based on
visual estimation. d total area of an island/surveyed part of an island. e the shortest distance between the island and mainland shorelines. f means of four observations
(consult the text). g T = total number of incidences (i.e., sums of species’ records across all samples), Sobs = observed species richness (all samples pooled), Q1 = the
number of species found in one sample only, Q2 = the number of species found in two samples only. h Sjack = calculated by jackknife method, Schao = calculated by chao2
method.
a
Boreal env. res. vol. 17
lat. nb
sample
characteristicsg
•
tojnak
malyi tojnak
Bolshoi tojnak
Khunuk
sarminskaya Kosa
Khubyn
shara-Dagan
oltrek
Borga-Dagan
Zamogoj
mys Ujuga
nameless
nameless
Kharantsy
modoto
edor
typea
sampling effort no. of
(person-hours) samples
Chepinoga et al.
01
02
03
04
05
06
07
08
09
10
11
12
13
14
15
16
impact scoresf
Boreal env. res. vol. 17 • Flora of the Baikal islands
Collection of additional information
Areas of islands and of the investigated parts
of peninsulas (Table 1) were determined either
from space photographs (available at GoogleEarth) or from measurements conducted during
fieldwork (small islands and peninsulas: nos. 4,
5, 11, 12 and 13 in Table 1). Impacts of tourism
and colonies of herring gull (Larus argentatus
Pontoppidan) on sampling areas were estimated
by averaging scores given individually by each of
four observers. Tourism and recreational activities: 0 = no visible traces of visitation; 1 = rubbish or other signs of visitation were occasionally
seen; 2 = trampled vegetation, paths, scrap-heaps
and bonfire places frequent across the island/
peninsula. Bird colonies: 0 = absent; 1 = present
but affecting minor part of island/peninsula; 2 =
affecting more than a half of island/peninsula.
Data analyses
Both species numbers and the sampled areas
were log-transformed prior the regression analysis. Effects of isolation (islands vs. peninsula),
recreation (low vs. high), and colonial birds
(low vs. high) on floristic diversity were tested
by ANCOVA using sampled area as a covariate
(SAS Institute 2009). Breakpoint regression was
calculated using Excel macros developed by
Lomolino and Weiser (2001).
Species lists were analysed with Integrated
Botanical Information System (Zverev 2007);
indicatory values for soil fertility and moisture
characteristics were calculated using ecological scales of south-Siberian plant species (after
Korolyuk 2006). Similarities in species composition between study areas were described using
the Jaccard index, i.e., the number of common
species divided by the total number of species recorded at both sites. A dendrogramm was
constructed using Statistica for Windows with
WPGMA algorithm (Weighted Pair Group using
Arithmetic Averages).
Changes in overall species richness and species turnover during the past 30 years were
explored for five islands by comparing our species lists with the lists based on surveys of 1979
(Klimina 1980, Petrochenko 1987). Absolute
223
(TA) and relative (TR) turnover rates were calculated using the following formulae (Panitsa et
al. 2008):
TA = (I + E)/2t,
TR = [(I + E)/t(S1979 + S2009)] ¥ 100,
where t is the period between censuses (i.e., 30
years in our case), E is the number of species
observed only in 1979 (i.e., extinct between
1979 and 2009), I is the number of species
observed only in 2009, S1979 and S2009 are the
total numbers of species recorded in 1979 and
2009, respectively. Numbers of species recorded
in 1979 and 2009 were compared using a paired
t-test (SAS Institute 2009).
Overall (i.e., expected) species richness was
calculated from the numbers of species recorded
by one of four observers only (‘singletons’,
Q1 in Table 1), and by two of four observers
(‘doubletons’, Q2). To estimate the numbers
of yet undiscovered species we employed the
Chao2 and jackknife methods, which showed the
best performance in several comparative studies (Walther and Morand 1998, King and Porter
2005).
Variables whose distributions did not fulfill
the normality assumption were analysed by nonparametric tests, including Spearman’s rank correlation (rS) and a Kruskal-Wallis test (SAS corr
and npar1way procedures, respectively; SAS
Institute 2009).
Results
Diversity of vascular plants
We recorded 269 species of vascular plants (see
Appendix). Species richness adjusted for sampling areas did not differ between islands and
peninsulas (Table 2).
Numbers of recorded species (Table 1)
ranged from 9 (Shara-Dagan Island) to 143
(Zamogoj Island). Elymus sibiricus was found in
all 16 study areas; 76 species were each recorded
a the single study area. The highest proportions
of these ‘unique’ species were found on Khunuk
Island (15.7% of the entire species list) and
Zamogoj Island (12.6%).
Chepinoga et al.
224
Fig. 3. relationship between the observed number
of species (S) and area (A) of the surveyed territory.
Filled circles = islands with high impacts of colonial
birds; empty circles = islands/peninsulas with low or no
impacts of colonial birds. regression is based on data
from islands/peninsulas with low or no impacts of colonial birds: logS = 2.52 (± 0.22) + 0.188 (± 0.023) ¥ logA;
r 2 = 0.88, p < 0.0001.
Areas with large bird colonies had significantly smaller species richness than areas with
small or no colonies (Table 2; least square means
adjusted for island areas: 28 and 65 species,
respectively). The current impact of tourism and
recreational activities did not cause detectable
changes in plant diversity (Table 2).
The breakpoint model fitted the entire data
set (Fig. 3) slightly better than the log-log linear
model (r2 = 0.654 and 0.609, respectively). However, when islands heavily affected by colonial birds were excluded from the analysis, the
•
Boreal env. res. vol. 17
species-area relationship was better described
by the log-log model (Fig. 3). Accounting for
the distance to the mainland did not improve the
model (data not shown).
The jackknife estimation method generally
predicted higher values of species richness (127%
of the observed species number) than the Chao2
method (116%). The ratio between the predicted
and observed numbers of species did not differ
between islands where we allocated a fixed time
for the surveys and where surveys were continued
until discoveries of new species became very
infrequent (ANOVA, jackknife: F1,12 = 0.03, p =
0.87; Chao2: F1,12 = 0.01, p = 0.91). This ratio also
did not depend on the sampled area (jackknife: r =
0.14, n = 14, p = 0.62; Chao2: r = –0.01, n = 14,
p = 0.97). Consistently, the slopes of the log-log
species-area regressions did not differ between the
observed species richness and the two estimates
of the expected numbers of species (ANCOVA:
F2,38 = 0.14, p = 0.87).
Similarities between sampled areas
WPGMA dendrogramm revealed an isolated
position of Shara-Dagan and Modoto Islands,
and identified (at the similarity level 0.25) three
main clusters (Fig. 4). The first cluster included
small rocky islands (Borga-Dagan and Edor) and
a nameless stony peninsula. Species-poor (9 to
28 species) floras of these rocky habitats, two
of which are heavily affected by colonial birds,
have high indicator values for soil fertility and
Table 2. effects of isolation, tourism, and bird colonies on diversity of vascular plants (ancova, log-transformed
values, type iii sums of squares). For characteristics of the sampled areas see table 1.
classificatory variable
source
df
mean square
F
isolation
isolation (islands vs. peninsulas)
area (= covariate)
isolation ¥ area
error
Birds (high vs. low impact)
area (= covariate)
Birds ¥ area
error
tourism (high vs. low impact)
area (= covariate)
tourism ¥ area
error
1
1
1
14
1
1
1
14
1
1
1
9
0.533
5.370
0.262
0.310
0.931
4.037
0.301
0.125
0.020
0.948
0.002
0.012
1.72
17.33
0.84
–
7.44
32.25
2.41
–
1.74
81.01
0.17
–
Birds colonies
tourism
p
0.21
0.0013
0.38
–
0.02
0.0001
0.15
–
0.23
< 0.0001
0.69
–
Boreal env. res. vol. 17 • Flora of the Baikal islands
225
Fig. 4. Floristic similarities
between sampled areas.
For names of islands and
peninsulas, see Fig. 2.
low indicator values for moisture (Fig. 5).
The largest cluster included six islands and
two peninsulas with high diversity of habitats
and high (64–143) numbers of recorded species
(Fig. 4). In terms of ecological requirements,
these floras are very similar to each other, demonstrating moderate indicator values for both
soil fertility and moisture (Fig. 5).
The final cluster, consisting of two small
islands (Toinak and Khunuk) and one peninsula (Sarminskaya Kosa) (Fig. 4), combines
floras with moderate numbers of species (53–70)
which have the lowest requirements for soil fertility combined with the highest indicator values
for moisture (Fig. 5).
Species turnover
A total of 284 species were recorded on a subset
of five islands that were surveyed in both 1979
and 2009 (Table 3). Among these, 228 were
found in 1979 (Petrochenko 1987), 237 in 2009
(Appendix), and 181 were common for both
surveys. We found no differences in species richness between the surveys (paired t-test: t4 = 1.81,
p = 0.15).
Mean absolute turnover rate was 1.06 species per year, and mean relative turnover rate was
1.17% of species per year, i.e., 35% mean spe-
Fig. 5. ordination of sampled areas by indicator values
for soil fertility and moisture. For names of islands and
peninsulas, see Fig. 2.
cies change between the subsequent observations.
Absolute turnover was independent of islands
area (rS = –0.10, n = 5 islands, p = 0.87), whereas
relative turnover decreased with increase in island
size (rS = –1.00, n = 5 islands, p < 0.0001). Comparisons of species’ lists from 1979 and 2009 did
not reveal any differences in indicator values for
soil fertility and moisture (p > 0.10).
Protected and ruderal species
We recorded five locally protected species: Stipa
Chepinoga et al.
226
glareosa (in 3 study areas), Oxytropis popoviana
(in 3 areas), O. tragacanthoides (in 2 areas),
Deschampsia turczaninowii (in 12 areas), and
Lilium pumilum (in 2 areas). All these species
were present on Zamogoj Island; none was found
on Tojnak, Khunuk, Shara-Dagan and Modoto
Islands; all other areas included 1–3 species.
Incidences of these species (i.e., the numbers of
areas in which they were recorded) did not differ
from incidences of all other (non-protected) species (Kruskal-Wallis test: χ21 = 0.24, p = 0.63).
The median proportion of protected species (6%)
among plants that have disappeared from island
floras between 1979 and 2009 was higher than
among immigrants (0%), but the differences
were far from the significance level (χ21 = 0.65,
p = 0.42).
We recorded 37 species classified as ruderal. Proportions of ruderal species varied from
4% (Kharantsy Island) to 31% (Sarminskaya
Peninsula); they peaked in medium-sized study
areas and showed no relationships with either
recreational loads (rS = –0.33, n = 16, p = 0.21)
or impacts imposed by bird colonies (rS = 0.23, n
= 16, p = 0.39), or distance to the nearest shore
(rS = –0.07, n = 16, p = 0.81). Incidences of
ruderal species did not differ from incidences of
all other (non-ruderal) species (Kruskal-Wallis
test: χ21 = 0.15, p = 0.70). Proportion of ruderal
species among immigrants was twice higher
than among species that disappeared from island
floras between 1979 and 2009 (13%–26% and
0%–15%, respectively; Kruskal-Wallis test: χ21 =
5.34, p = 0.02).
•
Boreal env. res. vol. 17
Discussion
Observed and estimated species
richness
It is only rarely possible to enumerate all the species present in the study area, even for vascular
plants (Connor and Simberloff 1978, Gilbert
and Lee 1980, Herwitz et al. 1996). Along with
low occurrence of some species, constrained
sampling effort unavoidably leads to the incompleteness of species’ lists. Since the number of
recorded species increases with both sampling
effort and the sampled area, uneven sampling
efforts may distort conclusions concerning species–area relationships (Preston 1979, Cam et al.
2002b).
Ecological studies commonly use three types
of methods to estimate total species richness: fitting of species-abundance distributions, extrapolation of species accumulation curves, and nonparametric estimators (Walther and Morand
1998). The use of the first method for plants is
hampered by practical impossibility to accurately quantify abundances of individual species.
The second method requires an objective measurement of sampling effort. While the number
of collected specimens is commonly used for
animals (Kozlov 1997, Willott 2001, Mauffrey
et al. 2007), this measure is hardly applicable to
field surveys of vascular plants. The use of collecting time is also questionable, because of both
collector’s personality and unavoidable uncontrolled variation in working efficiency. Therefore
the only practical choice is to use non-parametric
estimation methods.
Table 3. species turnover between 1979 and 2009.
island
species richness
turnover
no.
name
S1979
S2009
Spool
I
E
ta
tr
03
04
06
08
10
Bolshoi tojnak
Khunuk
Khubyn
oltrek
Zamogoj
89
59
63
123
136
80
70
89
141
143
120
97
102
169
168
31
38
39
46
32
40
27
13
28
25
1.183
1.083
0.867
1.233
0.950
1.400
1.679
1.141
0.934
0.681
species richness: S1979 = in 1979 (Petrochenko 1987), S2009 = in 2009 (this study), Spool = both censuses pooled.
turnover: E = number of extinct species (i.e., species observed only in 1979), I = number of immigrants (i.e., species observed only in 2009), ta = absolute turnover, tr = relative turnover (see text for the formulae).
Boreal env. res. vol. 17 • Flora of the Baikal islands
The jackknife method (in agreement with
conclusions by Walther and Morand 1998) predicted on average 10% higher values of species richness than the Chao2 method; these two
estimates can be seen as the boundaries between
which the actual value of species richness lies
(Chiarucci et al. 2003). Therefore, we conclude
that the completeness of our inventories ranged
from 77% to 89% of the potential species richness, which is very close to published estimates
for ants collected by pitfall traps (71%–90%;
King and Porter 2005) and for point counts of
birds (79%–100%; Cam et al. 2002a).
Chiarucci et al. (2003) concluded that at least
15%–30% of the total area needed to be sampled
to obtain reasonable estimates of total species
richness. During our surveys, we walked with an
average speed 2 km h–1 and recorded plants within
an approximately 4-m-wide area. With the applied
effort (Table 1) we surveyed 13% to 100% of
the areas designed for sampling. Allocation of
the fixed time to three largest islands and, consequently, relatively low coverage of these islands
by the surveying routes (13%–37%) did not
decrease completeness of our surveys. Therefore,
we conclude that when sampling is not random
but driven by ‘botanist’s internal algorithm’ (intuition) this coverage was still sufficient to avoid
underestimation which is seen as the basic problem of field surveys that cover only minor part of
the total study area (Chiarucci et al. 2003).
Impacts of colonial birds on local floras
Herring gull is the most common bird nesting on
the Maloe More islands. Island colonies of this
species totaled about 500 nests in the early 1970s
(Litvinov 1979); surveys of 1977–1984 revealed
2850–3825 gulls (Skryabin and Pyzh’yanov
1987), and since then the number of nesting gulls
has steadily increased (Pyzh’yanov 1997). However, since no recent data are available, we chose
to use a subjective rank of bird impact on our
study areas. Importantly, our estimates were generally consistent (rS = 0.59, n = 11 islands, p =
0.06) with bird numbers recorded in 1977–1984
(after Skryabin and Pyzh’yanov 1987), confirming stability of bird colonies over a long period
of time.
227
Impacts of colonial birds on vegetation are
documented for maritime islands of different
regions. Large bird colonies are usually surrounded by specific plant communities, which
consist of a few species that are able not only to
sustain heavy nitrogen and phosphorous loading,
but even benefit from it (Luther 1961, Sobey
and Kenworthy 1979, Glazkova 2009, and references therein). Vegatation of small islands hosting large bird colonies is extremely degraded
(Gillham 1953, Zelenskaya and Khoreva 2006).
Consistently with these observations, we
found more than two-fold decrease in overall species richness on islands with large gull
colonies. However, no plant species was found
exclusively on these islands, although their floras
consisted of plants with low-moisture and high
soil-fertility requirements (Fig. 5). An isolated
position of Shara-Dagan Island on the dendrogramm (Fig. 4) is explained by the absence of
information on plants growing on the upper
part of this rock, which was impossible to reach
without alpinist skills. Vegetation of the surveyed stony parts at the bottom of this rock was
less nitrofilous than on Borga-Dagan, Edor, and
Modoto Islands (Fig. 5), in spite of the presence of large gull colony on Shara-Dagan Island
(Table 1).
Species–area relationships
Although hundreds of studies fitted the relationship between the number of species and the
area of an island (or island-like fragment) with
the power-law function (reviewed by Drakare
et al. 2006), its generality has frequently been
questioned. In particular, it has been suggested
that the power law adequately describes species accumulation only in medium-sized to large
islands and fragments, while on small islands
richness may vary independently of island area
(a phenomenon called the Small Island Effect).
This hypothesis dates back to the 1960s and
was extensively debated in the past (Woodroffe
1986); the recent review by Lomolino and
Weiser (2001) renewed interest in it.
Lomolino (2000) explained the low number
of data sets that demonstrated small island effect
by low frequency of studies involving smallest
228
islands and fragments, and called for collecting
additional data. Since our study also covered
very small and isolated habitats (Table 1), we
fitted our data with both linear and breakpoint
regression models. In agreement with many earlier studies (e.g., Woodroffe 1986, Heatwole
1991, Fridley et al. 2005, Panitsa et al. 2006),
we failed to detect the small island effect in our
data set.
The slope of the power-law function fitting
our data (z = 0.188) lies within the 95% confidence limits (0.13–0.23) calculated from several
dozens of studies that used independent (i.e., not
nested) sampling scheme in non-forested terrestrial habitats (Drakare et al. 2006).
Species turnover
The dynamics of the insular floras and faunas
forms the core of the equilibrium theory of island
biogeography (MacArthur and Wilson 1967).
Many researchers attempted to estimate turnover
rates (reviewed by Panitsa et al. 2008); however,
the measured values were always approximations of real values due to unavoidable confounding effects of cryptoturnover (undetected
turnover, when both extinction and colonisation by the same species occurred between the
observations) and pseudoturnover (when species
present on the island have not been detected
during one of two censuses). Under these circumstances, accumulation of data on turnover
rates in different biomes during different periods
of time is critically needed to allow identification
of both general patterns and sources of variation
in the balance between local extinction and colonisation processes.
Yu. N. Petrochenko (pers. comm.) informed
us that his team visited each island several times,
and that surveys of larger islands lasted for several days. This clearly exceeded our own sampling effort and, therefore, we have the reasons
to believe that the completeness of the first floristic inventory (in 1979) was at least not worse
than the completeness of the second one (in
2009). This conclusion is indirectly supported by
an overall similarity in the numbers of species
recorded during these two surveys. At the same
time, species lists of 1979 and 2009 demon-
Chepinoga et al.
•
Boreal env. res. vol. 17
strated substantial differences: an average overlap was only 49% of the pooled lists (Table 3).
If we hypothesise that the larger part of differences between the surveys was due to incompleteness of the data collected in 2009, then the
highest differences should be associated with
the largest islands. However, our data yielded
the opposite pattern: the highest turnover was
found on the smallest island, flora of which was
obviously revealed more completely than floras
of the largest islands. This result agrees with
the pattern observed in Kem-Lud archipelago
(Shipunov and Abramova 2006) and fits the
predictions of the equilibrium theory, according
to which maximum turnover rates are expected
in small islands (MacArthur and Wilson 1967).
Thus, we conclude that species’ lists of 1979 and
2009 are of about the same completeness, and
that the differences between these lists are likely
to result from species turnover rather than from
methodological shortcomings.
The relative turnover rates found for our
islands (0.681% to 1.679% of species per year)
fit well the ranges reported for other islands
(Panitsa et al. 2008, and references therein).
However, in contrast to observations by Panitsa
et al. (2008), we detected an obvious trend in
species’ composition. Although changes were
random in terms of ecological requirements of
plant species, proportions of ruderal species
in local floras doubled (from 6.9% to 11.2%)
during the past 30 years. Ruderalisation can be
seen as the first indication of the increasing disturbance of island vegetation, in particular due to
creation of paths crossing steppe communities.
These paths enhance the spread of the opportunistic species and might be a threat for conserving
native flora (Godefroid and Koedam 2004). On
the other hand, climate warming in the study
region (Moore et al. 2006) may increase the risk
of establishment of ruderal species.
Conservation of local floras
Baikal region in Russia is now experiencing an
increasing environmental pressure from mass
tourism, which is an important socioeconomic
factor of regional development. Visitors to Baikal
mostly enjoy camping, fishing, beach activi-
Boreal env. res. vol. 17 • Flora of the Baikal islands
ties, walking and viewing picturesque scenery.
Eco-tourism is rare, possibly due to absence of
a charismatic flagship species, and ecologically
ignorant tourists seeking seclusion and relaxation contribute to degradation of landscapes and
floristic impoverishment (Kovtonyuk et al. 2003,
Kas’yanova 2007, Vin’kovskaya 2007). These
processes increase the importance of conservation of the Lake Baikal islands, which have so
far experienced much lower human pressure
than the mainland habitats near the shoreline.
Semi-desert and steppe-desert plant communities of the Maloe More islands include
a number of relicts (e.g., Stipa glareosa and
Oxytropis tragacanthoides), steppe species at the
northern borders of their distribution limits (e.g.,
Filifolium sibiricum and Allium burjaticum), and
other endangered and locally protected species
(Appendix). The islands remain one of few habitats which secure persistence of regional endemics, such as Deschampsia turczaninowii, Festuca
rubra subsp. baicalensis, Oxytropis popoviana,
and Artemisia ledebouriana, and of locally rare
species (e.g., Thymus pavlovii and Asplenium
altajense), generally suffering from an increase
of recreational impact on the shoreline habitats of
Baikal. Imposing restrictions on tourist visitation
to at least three islands (in the order of decreasing
importance: Zamogoj, Khubyn and Khunuk) is a
feasible way to conserve substantial part of local
biodiversity and to allow long-term monitoring
of climatic effects on structure and dynamics of
steppe plant communities at the northern limit of
the semi-arid landscapes in northern Asia.
Acknowledgements: We are grateful to Yu. N. Petrochenko
for information on the design of floristic studies conducted
during 1970s–1980s, to M. Lomolino for Excel macros, and
to anonymous reviewers for valuable suggestions. The study
was supported by the Academy of Finland (project 8126045)
and by the University of Turku strategic research grant.
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Appendix. Presence of vascular plants in sampled areas in 2009.
Plant species
Achnatherum sibiricum
Aconogonon alpinum
A. angustifolium
A. ochreatum
Agropyron cristatum
Agrostis trinii
Aizopsis aizoon
Aleuritopteris argentea
Allium burjaticum
A. ramosum
A. senescens
A. splendens
A. stellerianum
A. tenuissimum
Alyssum lenense
Amblynotus rupestris
Amethystea caerulea
Androsace incana
A. lactiflora
Anemone dichotoma
Arabis pendula
Arctopoa subfastigiata
Artemisia commutata
A. dolosa
A. dracunculus
A. frigida
A. gmelinii
A. laciniata
A. ledebouriana
A. leucophylla
A. mongolica
species
category
islands/peninsulas*
1
2
3
4
5
6
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
7
+
ruderal
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
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+
+
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+
+
9 10 11 12 13 14 15 16
+
+
+
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8
+
+
+
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+
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+
+
+
+
+
+
+
+
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+
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+
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+
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+
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+
+
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+
+
+
+
+
+
+
+
+
+
+ +
Continued
Chepinoga et al.
232
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Boreal env. res. vol. 17
Appendix. continued.
Plant species
A. monostachya
A. pubescens
A. sericea
A. vulgaris
Asplenium altajense
Aster alpinus s.l.
Astragalus chorinensis
A. inopinatus
A. lupulinus
A. suffruticosus
A. versicolor
Atragene speciosa
Axyris amaranthoides
A. hybrida
Barbarea sp.
Betula pendula
B. platyphylla
Bistorta attenuata
B. vivipara
Bromopsis inermis
B. korotkiji
B. sibirica
Bupleurum bicaule
B. scorzonerifolium
Calamagrostis epigeios
C. langsdorffii
Callitriche palustris
Campanula rotundifolia
Carduus crispus
Carex appendiculata
C. argunensis
C. duriuscula
C. korshinskyi
C. nigra
C. pediformis
C. pseudocuraica
C. rhynchophysa
C. rostrata
C. sajanensis
Chamaenerion angustifolium
Chamaerhodos altaica
C. erecta
C. grandiflora
Chenopodium album
C. aristatum
C. hybridum
C. novopokrovskianum
Chryzanthemum zawadskii
Cirsium setosum
Clausia aprica
Cleistogenes squarrosa
Comarum palustre
Corydalis impatiens
species
category
islands/peninsulas
1
2
3
4
5
+
ruderal
+
+
+
+
+
+
+
ruderal
ruderal
ruderal
6
7
8
9 10 11 12 13 14 15 16
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
ruderal
+
+
+
+
+
+
+
ruderal
ruderal
ruderal
+
+
+
+
+
+
+
+
+
+
+
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+
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+
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+
+
+
+
+
+
+
+
+
+
+
ruderal
ruderal
+
+
+
+
+
+
+
+
+
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+
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+
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+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Continued
Boreal env. res. vol. 17 • Flora of the Baikal islands
233
Appendix. continued.
Plant species
Cotoneaster melanocarpus
Critesion brevisubulatum
Cystopteris fragilis
Dasiphora fruticosa
Dasystephana decumbens
Delphinium grandiflorum
Deschampsia turczaninowii
Dianthus versicolor
Dontostemon integrifolius
D. pinnatifidus
Dracocephalum nutans
D. olchonense
D. pinnatum
Duschekia fruticosa
Elymus sibiricus
Elytrigia repens
Ephedra monosperma
Epilobium palustre
Equisetum arvense
Eremogone meyeri
Erysimum cheiranthoides
E. hieracifolium
Euphrasia pectinata
Fallopia convolvulus
Ferulopsis hystrix
Festuca lenensis
F. ovina
F. rubra ssp. rubra
F. rubra ssp. baicalensis
F. sibirica
Filifolium sibiricum
Fornicium uniflorum
Galeopsis bifida
Galium aparine
G. uliginosum
G. verum
Geranium pratense
G. sibiricum
Goniolimon speciosum
Gypsophila patrinii
Hedysarum gmelinii ssp. setigerum
Helictotrichon altaicum
H. hookeri ssp. schellianum
Heteropappus altaicus
H. biennis
Hierochloe glabra
Hylotelephium triphyllum
Hypericum gebleri
Iris humilis
Isatis oblongata
Kitagawia baicalensis
Kochia prostrata
Koeleria cristata ssp. cristata
species
category
islands/peninsulas
1
2
3
4
5
6
7
+
8
9 10 11 12 13 14 15 16
+
+
+
protected
+
+
+
+
+
+
+
+
+
+
+
+
ruderal
+
ruderal
ruderal
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
ruderal
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
ruderal
+
+
+
ruderal
ruderal
ruderal
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Continued
Chepinoga et al.
234
•
Boreal env. res. vol. 17
Appendix. continued.
Plant species
K. cristata ssp. hirsutiflora
Lamium album
Lappula redowskii
Larix ¥ czekanowskii
Lathyrus pilosus
Leontopodium leontopodioides
Leonurus deminutus
Lepidium apetalum
Leymus chinensis
Lilium pumilum
Linaria acutiloba
L. buriatica
Lupinaster pentaphyllus
Lychnis sibirica
Lycopodioides sanguinolenta
L. siberica
Minuartia stricta
M. verna
Mulgedium sibiricim
Odontites vulgaris
Orobanche caesia
Orostachys spinosa
Oxytropis coerulea
O. popoviana
O. tragacanthoides
O. turczaninovii
Papaver nudicaule
Parnassia palustris
Patrinia rupestris
P. sibirica
Pedicularis rubens
Persicaria amphibia
P. hydropiper
P. lapathifolia
Peucedanum puberulum
Phalaroides arundinacea
Phlojodicarpus sibiricus
P. villosus
Phlomis tuberosa
Pinus sylvestris ssp. kulundensis
Plantago depressa
P. media
Poa angustifolia
P. attenuata
P. palustris
P. pratensis
Polygala tenuifolia
Polygonatum odoratum
Polygonum aviculare s.l.
Populus tremula
Potentilla acaulis
P. acervata
P. anserina
species
category
islands/peninsulas
1
2
3
+
+
+
ruderal
+
protected
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
ruderal
+
+
ruderal
ruderal
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
ruderal
+
+
+
+
+
+
+
9 10 11 12 13 14 15 16
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
8
+
+
+
+
+
+
+
+
7
+
+
+
+
+
+
6
+
+
+
+
ruderal
5
+
protected
protected
+
4
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Continued
Boreal env. res. vol. 17 • Flora of the Baikal islands
235
Appendix. continued.
Plant species
P. arenosa
P. bifurca
P. conferta
P. longifolia
P. sericea
P. tanacetifolia
P. tergemina
Ptilotrichum tenuifolium
Puccinellia hauptiana
Pulsatilla patens s.l.
P. tenuiloba
P. turczaninovii
Ranunculus propinquus
Rheum rhabarbarum
Rhinanthus serotinus
Rhododendron dauricum
Ribes nigrum
Rorippa palustris
Rosa acicularis
R. majalis
Rumex acetosella
R. aquaticus
R. thyrsiflorus
Salix bebbiana
S. dasyclados
S. jenisseensis
S. rhamnifolia
S. rorida
S. taraikensis
S. viminalis
Salsola collina
Sanguisorba officinalis
Saussurea salicifolia
S. schanginiana
Saxifraga cernua
S. spinulosa
Scabiosa comosa
Schizonepeta multifida
Scorzonera austriaca
S. glabra
Scrophularia incisa
Scutellaria scordiifolia
Serratula centauroides
Silene jeniseensis
S. repens
Sisymbrium heteromallum
Sium suave
Smelowskia alba
Sonchus arvensis
Sorbus sibirica
Sphallerocarpus gracilis
Spiraea media
Stachys aspera
species
category
ruderal
islands/peninsulas
1
2
3
+
+
+
+
+
+
+
+
+
4
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
ruderal
ruderal
ruderal
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
9 10 11 12 13 14 15 16
+
+
+
+
8
+
+
+
ruderal
7
+
+
ruderal
6
+
+
+
ruderal
5
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Continued
Chepinoga et al.
236
•
Boreal env. res. vol. 17
Appendix. continued.
Plant species
Stellaria cherleriae
S. dichotoma
S. longifolia
Stipa baicalensis
S. glareosa
S. krylovii
Taraxacum ceratophorum
T. dissectum
T. mongolicum
Tephroseris integrifolia
Thalictrum appendiculatum
T. foetidum
Thermopsis lanceolata ssp. sibirica
Thlaspi arvense
Thymus baicalensis
Urtica cannabina
U. dioica
Utricularia intermedia
Valeriana officinalis
Veronica longifolia
Vicia cracca
V. nervata
Vincetoxicum sibiricum
Viola rupestris
Youngia tenuifolia
species
category
protected
islands/peninsulas
1
2
3
+
+
+
+
ruderal
+
ruderal
+
ruderal
ruderal
ruderal
4
+
+
+
+
+
+
+
+
+
+
+
6
+
+
+
+
+
+
8
9 10 11 12 13 14 15 16
+
+
+
+
+
+
+
+
+
+
+
+
+
+
7
+
+
+
+
+
+
+
+
+
+
+
+
5
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
* islands/peninsulas: 1 = tojnak island, 2 = malyi tojnak island, 3 = Bolshoi tojnak island, 4 = Khunuk island, 5 =
sarminskaya Peninsula, 6 = Khubyn island, 7 = shara-Dagan island, 8 = oltrek island, 9 = Borga-Dagan island,
10 = Zamogoj island, 11 = Ujuga Peninsula, 12 = nameless Peninsula, 13 = nameless Peninsula, 14 = Kharantsy
island, 15 = modoto island, 16 = edor island. For coordinates and other characteristics, see table 1.