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Niche analysis of orchids of serpentine and nonserpentine areas: Implications for conservation
a
To cite this article: V. Djordjevi, S. Tsiftsis, D. Lakui & V. Stevanovi (2014): Niche analysis of orchids of serpentine and
non-serpentine areas: Implications for conservation, Plant Biosystems - An International Journal Dealing with all Aspects of
Plant Biology: Official Journal of the Societa Botanica Italiana, DOI: 10.1080/11263504.2014.990534
To link to this article: http://dx.doi.org/10.1080/11263504.2014.990534
ORIGINAL ARTICLE
Faculty of Biology, Institute of Botany and Botanical Garden, University of Belgrade, Takovska 43, 11000 Belgrade, Serbia
and 2Department of Botany, School of Biology, Aristotle University of Thessaloniki, 54124 Thessaloniki, Greece
Abstract
Orchids are known for their species richness, intriguing ecology, rarity and the fact that they grow in almost all terrestrial
ecosystems. Although numerous studies about their ecology have been carried out concerning calcareous areas, little is
known about orchids that occur in serpentine habitats. The aim of this study was to investigate the ecological preferences of
orchids in serpentine and non-serpentine areas on the model of the Valjevo Mountain Range (W Serbia). Niche analysis of
orchids was performed using outlying mean index analysis. Data concerning geographical coordinates, altitude, habitat type,
inclination, bedrock type, light regime, soil moisture, acidity, nitrogen and temperature were used as explanatory variables.
Data of 33 orchid taxa from 407 localities were analysed. The most important gradients that govern orchid distribution were
geological bedrock, light regime and temperature. The results have shown that only Anacamptis morio and Gymnadenia
conopsea have statistically significantly larger populations on serpentine compared with non-serpentine bedrocks. This study
highlights the importance of serpentine habitats as orchid habitats, bearing in mind the occurrence of rare species and species
which were found exclusively in serpentine habitats.
Introduction
Understanding how environmental conditions and
species interactions determine the distribution and
abundance of species is a central problem and a
challenge in ecological and biogeographical research.
In addition, successful species conservation requires
the knowledge of their distribution patterns and
ecological preferences. On a large geographic scale,
patterns of species distributions are determined by
evolutionary and migration history together with
macroclimate. On the other hand, factors such as soil
properties, light regime, meso- and microclimate,
altitude, habitat type, disturbance, biotic factors, and
so on affect distribution and abundance of species on
a regional scale (Diez & Pulliam 2007; Blinova 2008;
Landi et al. 2009; Tsiftsis et al. 2012). The ecological
niche as defined by Hutchinson (1957) is a multidimensional hypervolume, determined by the habitat
gradients and the functional relationship among
species, within which a species maintains a viable
population. Niche analysis is a very useful tool for a
better understanding of the environmental requirements and distribution of species, as well as for
conservation planning (Tsiftsis et al. 2008; Casazza
et al. 2013).
The family Orchidaceae is one of the most speciesrich plant families with ca. 25,000 species distributed
worldwide, but it is most diverse and numerous in the
tropics and subtropics (Whigham & Willems 2003).
With the exception of the poles, extremely dry
deserts, salt marshes, and cultivated agricultural
lands, orchids grow in all terrestrial ecosystems
(Hagsater & Dumont 1996). More than 300 species
have been recorded to occur in Europe, North Africa
and Near East, with most of them distributed in
Mediterranean basin and hinterland (Hagsater &
Dumont 1996). In Serbia, 72 orchid taxa (species
and subspecies) have been recorded (Diklic 1976;
Karadzic et al. 2000; Djordjevic et al. 2010).
Although a few studies provide data on the plant
communities and geological bedrocks in which
individual orchid species occur, generally ecological
Correspondence: V. Djordjevi, Faculty of Biology, Institute of Botany and Botanical Garden, University of Belgrade, Takovska 43, 11000 Belgrade, Serbia.
Tel: 381 11 3244923. Fax: 381 11 3243603. Email: vlakiorhi@yahoo.co.uk
q 2014 Societa` Botanica Italiana
V. Djordjevic et al.
Table I. Habitat types and plant communities in which orchids were recorded.
Habitat type
Plant communities
Fagus forests
Festuco-Brometea grasslands
V. Djordjevic et al.
Results
The results of the OMI analysis are presented in
Table II. In total, 25 orchid taxa showed a significant
marginality, suggesting a considerable influence of
the selected environmental variables on species
distribution. The other seven taxa that did not show
significant marginality were those recorded in a small
number of localities. The relatively low values of the
residual tolerance indicate that the selected environmental variables strongly affect the distribution of the
orchids of the study area (Table II).
Table II. Niche parameters of orchid species of the Valjevo Mountain Range and their substrate preferences.
Taxon
Code
Occ
Inertia
OMI (%)
T1 (%)
T2 (%)
Num
Anacamptis coriophora
Anacamptis morio
Anacamptis papilionacea
Anacamptis pyramidalis
Cephalanthera damasonium
Cephalanthera longifolia
Cephalanthera rubra
Dactylorhiza incarnata
Dactylorhiza maculata subsp. maculata
Dactylorhiza maculata subsp. transsilvanica
Dactylorhiza saccifera
Dactylorhiza sambucina
Epipactis atrorubens
Epipactis helleborine
Epipactis microphylla
Epipactis palustris
Epipactis purpurata
Gymnadenia conopsea
Himantoglossum jankae
Limodorum abortivum
Neotinea tridentata
Neotinea ustulata
Neottia nidus-avis
Neottia ovata
Ophrys insectifera
Ophrys scolopax subsp. cornuta
Orchis mascula
Orchis purpurea
Orchis simia
Platanthera bifolia
Platanthera chlorantha
Spiranthes spiralis
Traunsteinera globosa
Anaccori
Anacmori
Anacpapi
Anacpyra
Cephdama
Cephlong
Cephrubr
Dactinca
Dactmama
Dactmatr
Dactsacc
Dactsamb
Epipatro
Epiphell
Epipmicr
Epippalu
Epippurp
Gymncono
Himajank
Limoabor
Neottrid
Neotustu
Neotnidu
Neotovat
Ophrinse
Ophrscco
Orchmasc
Orchpurp
Orchsimi
Platbifo
Platchlo
Spirspir
Trauglob
22
87
2
8
15
23
6
6
3
29
13
34
5
36
5
2
2
143
2
22
8
18
43
7
1
2
2
2
6
63
2
2
4
13.14
18.57
24.51
15.25
28.46
27.88
25.49
37.37
40.18
30.55
24.87
39.11
31.99
34.59
35.47
46.83
62.62
18.60
34.31
22.78
15.44
19.46
27.52
18.29
2.76
2.59
17.65
9.21
7.05
7.99
10.73
16.52
40.10
11.92
5.56
10.65
31.35
8.65
22.25
46.32
62.20
2.24
34.03
11.85
10.02
3.27
8.67
5.26
0.31
1.31
0.01
0.76
1.82
2.40
1.69
2.15
0.00
5.01
1.01
9.48
0.00
1.89
4.10
0.00
0.01
1.21
0.01
1.43
0.12
0.43
2.14
0.49
10.07
14.68
6.85
5.27
19.59
17.48
13.07
18.70
0.08
13.62
18.30
18.98
0.64
24.05
9.12
0.51
0.40
15.15
0.27
9.49
5.30
15.77
16.71
12.54
159
0
1233 (ns)
189
1
0
120
59
2
0
29
0
0
0
1
12
1
0
83
0
23
117
0
1314 (ns)
31.75
25.20
36.67
35.07
27.04
18.06
15.67
14.39
21.53
18.55
36.38
24.27
3.48
18.04
9.27
10.39
1.80
0.35
0.01
0.74
3.60
0.00
0.21
0.04
8.42
6.30
0.29
10.06
19.95
0.02
6.19
3.96
745 (ns)
1713 (ns)
57
0
0
1162 (ns)
5630 (ns)
989 (ns)
Note: Occ, occurrences; Inertia, total variability; OMI, outlying mean index (%); T1, species tolerance (%); T2, residual tolerance (%);
Num, number of random permutations; S, serpentine; L, limestone; D, diabases and spilites.
Figure 1. Canonical weights of the environmental variables. The relative importance of each variable in the analysis is showed by the length of
the arrow, whereas the direction of the arrow indicates between-variables correlations; for the abbreviations of environmental variables, see
Table III.
V. Djordjevic et al.
Table III. Pearsons correlation coefficients between the environmental variables and the first three axes of OMI analysis.
Environmental variables
Code
Latitude
Longitude
Altitude
Arrhenatheretalia elatioris grasslands
Fagus forests
Festuco-Brometea grasslands
Marshes and hygrophilous meadows
Molinion coeruleae grasslands
Nardetalia strictae grasslands
Ostrya forests
Pinus nigra forests
Pinus sylvetris forests
Querco-Carpinetum betuli forests
Quercus forests
Inclination
Diabases and spilites
Limestone
Serpentine
Light Regime
Soil moisture
Acidity
Nitrogen
Temperature
LAT
LON
ALT
Arrhenatheretalia
Fagus
Festuco-Brometea
Marshes
Molinion
Nardetalia
Ostrya
Pinus nigra
Pinus sylvetris
Querco-Carpinetum
Quercus
Inclination
Diabases
Limestone
Serpentine
Light
Moisture
Acidity
Nitrogen
Temp
OMI axis 1
OMI axis 2
OMI axis 3
0.45**
20.31**
20.49**
20.02ns
0.37**
20.30**
20.05ns
20.55**
20.23**
0.38**
0.04ns
20.15**
0.29**
0.23**
0.36**
0.22**
0.60**
20.66**
20.59**
20.40**
20.51**
0.52**
0.38**
20.32**
0.46**
20.06ns
20.24**
0.58**
20.52**
0.08ns
0.37**
0.21**
20.26**
20.07ns
0.20**
0.05ns
0.08ns
20.22**
0.39**
20.44**
0.31**
20.54**
0.54**
0.30**
0.13*
20.67**
0.53**
20.51**
0.38**
0.51**
0.28**
20.16**
0.02ns
20.03ns
0.10*
0.01ns
20.20**
20.01ns
0.07ns
20.67**
0.32**
0.14**
0.26**
20.30**
0.06ns
0.12*
0.43**
0.38**
20.33**
Discussion
Specialists and generalists
Orchid taxa tend to be relatively common close to the
centre of their geographical distribution, whereas
they become rarest near their distribution edges
(Pfeifer et al. 2010). This, in part, is the result of the
special ecological requirements of each taxon.
The results of the OMI analysis (Figure 2;
Table II) allow us to separate two major categories of
Figure 2. Weighted positions of the species along the first two axes of the OMI analysis. Species with high marginality are located far from
origin, and those with low marginality are located near the origin; for the abbreviations of taxon names, see Table II.
V. Djordjevic et al.
10
V. Djordjevic et al.
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