Ernst-Moritz-Arndt-University Greifswald The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Diploma thesis in the study programme Landscape Ecology and Nature Conservation Michael Heiß -June 2010- Supported by DAAD Michael Succow Foundation Supervised by Prof. Dr. Michael Succow Dr. Martin Flade
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I
Ernst-Moritz-Arndt-University Greifswald
The breeding bird communities of the Talish mountains (Azerbaijan)
and their response to forest degradation
Diploma thesis in the study programme Landscape Ecology and Nature Conservation
Michael Heiß
-June 2010-
Supported by DAAD Michael Succow Foundation
Supervised by Prof. Dr. Michael Succow Dr. Martin Flade
Picus canus, Phoenicurus erythrogaster or Tichodroma muraria, are up to day unclear.
Species responses to forest degradation, fragmentation and deforestation have garnered
much recent interest throughout all continents and all forest types (e.g. Edenius &
Elmberg 1996, Poulsen 2002, Sekercioglu 2002, Echeverria et al. 2007, Fuller et al.
2007b, Murakami et al. 2008). The impact of the degradation of the Caspian forest to the
fauna is unknown. No detailed studies are published yet.
Therefore, the aim of this study is to answer the following questions:
• What are the breeding bird communities of the Talish mountains and which
species do they contain?
• Which parameters are responsible for their species composition?
• How do the breeding birds respond to the degradation of the Caspian forest in
Azerbaijan?
• What are the derived nature conservation implications?
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Study site
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2 Study site
2.1 Location
The study site is located in the southernmost part of Azerbaijan at the border to Iran
(Figure 1). It includes the districts of Masally, Lenkoran, Astara and Lerik. The total area
covers 3960 km² (Statistical Committee Azerbaijan 2009).
Figure 1: Location of Azerbaijan and the study site. Note the dark green band in northern Iran and adjacent southern Azerbaijan indicating the distribution of the Caspian forest. Clearings of the nearby Caspian lowland are light green (image source: NASA 2004).
Study site The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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The altitude ranges from -24 m a.s.l. (metres above sea level) at the Caspian coast in the
East and 2492 m a.s.l. of the Kiumiurkei mountain at the Iranian border in the West
(Skvorov 1976a, Skvorov 1976b).
The study site contains several protected areas. The three largest are the Hirkan
National Park, which was established in 2004 and enlarged in April 2008 to 40358 ha
(MENR 2010), the Gizilaghaj state nature reserve at the Caspian coast, which covers
88360 ha and is an important area for wintering and breeding waterbirds (Patrikeev 2004,
Sultanov 2008, MENR 2010) and the 15000 ha sized Zuvand state nature sanctuary
(MENR 2010).
2.2 Climate
The study site can be subdivided into two differing climatic regions. The eastern slopes of
the Talish mountains down to the coastal lowland are characterised by a warm-temperate
and humid climate. The annual rainfalls exceed 1000 mm per year and peak in February
and October. The summer temperatures are warm and the winters are mild (Figure 2).
Winter means rarely drop below zero (Mammadov et al. 2007, Mühr 2007).
Figure 3: Climate diagram of Ardebil 50 km south of Zuvand upland based on data of IRIMO (2010).
Figure 2: Climate diagram of Lenkoran after Mühr (2007).
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Study site
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The Zuvand area is the second climatic region. Due to mountain ridges, which shield
humid air masses coming from the Caspian sea, the Zuvand upland is much drier and the
climatic situation is similar to the adjacent Iranian semi-desert. It has an annual mean
temperature of 4-8°C. The winters are cold and occasionally the temperature drops below
-20°C. The summers show a distinctive drought from June to October and the annual
rainfalls are between 200-400 mm per year (Mammadov et al. 2007, cf. Figure 5 in Knapp
2005). No meteorological data was available from the Zuvand region and thus the data of
Ardebil (Iran) with a similar climate was taken depicted in Figure 3.
2.3 Landscape types
The Talish mountains are subdivided into six landscape or habitat types (Figure 4). The
forest belt contains five degradation stages. Each type has its own characteristics:
Caspian lowland
From the shore of the Caspian sea to the foothills of the Talish mountains stretches the
Caspian lowland, a long but rather narrow stripe including the cities Masally, Lenkoran
and Astara (Figure 1, Figure 4). Once it was covered with subtropical broad-leaf forests
and impassable wetlands (Radde 1886a). To fight malaria and to gain land for agriculture,
forests had been cleared and a widespread system of drainage channels indicate the large
range of former wetlands (Patrikeev 2004). Nowadays, the lowland is largely covered
with fields, pastures and human settlements (Figure 5, Figure 6). The last remnants of the
former lowland forest can be found in a 91 ha sized part of the Hirkan National Park
called ‘Moscow forest’ (MENR 2010). The only further forest-like structures in the low-
land are afforestations of Quercus spec., orchards or cemeteries.
Costal habitats like lagoons, shores or large reedbeds are excluded from this study.
Study site The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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Figure 4: Schematic of the vertical and horizontal distribution of the landscape types based on Grossheim (1926), Knapp (2005) and recent Google Earth (2010) satellite images.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Study site
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Forest belt and the degradation stages
The deciduous broadleaf forest of the Talish mountains ranges from the sea level up to
1800 m a.s.l. (MENR 2004) but the upper tree line is mostly reduced to lower altitudes
due to human activities. The number of tree species is high (ca. 90) and includes many
endemic or tertiary relict species like Gleditsia caspica, Parrotia persica, Quercus casta-
2004, Hajiyev 2006). Details about forest types and tree species composition of the
Caspian forest are given in Knapp (2005).
Scharnweber et al. (2007) identified and described six degradations stages of the
Caspian forest in the Talish mountains, on which this study base:
Natural forest stage
This stage shows no signs of human activities. It is generally restricted to higher altitudes
above 500 m a.s.l., but occurs also elsewhere along steep slopes or apart from human
settlements and roads (Figure 7). The natural forest stage can be treated as a primeval
forest comparable to the Białowieża forest in Poland. These relicts can be distinguished
from other temperate deciduous or mixed forests by their large heights, multistorey profile
of stands, a diverse tree community and large amounts of dead wood (Wesołowski 2007).
I found this stage near the village Siov and the abandoned village Armudy within the
Hirkan National Park.
Figure 6: Drainage channel. Sirabil, 21.05.2008. Figure 5: The Caspian lowland is characterised by agriculture and settlements. Sirabil, 21.05.2008.
Study site The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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Slightly disturbed forest stage
The slightly disturbed forest stage is similar to the natural forest stage. The human impact
is low and often not obvious. Thus, this stage was in a few cases difficult to distinguish
from the natural forest stage. Hints for a slightly disturbed forest stage are paths, single
snags, partly missing dead wood and evidences for grazing like excrements or traces of
domestic animals or a reduced understorey. According to Scharnweber et al. (2007), are
there no differences to the natural forest stage concerning tree species composition and
stand structure. Old-growth trees are a common aspect.
Intermediate disturbed forest stage
Logging, lopping and grazing are clearly visible in this stage. This stage can be found near
villages (2-4 km) or along roads. Loose cattle or grazing flocks of sheep can regularly be
seen in the forest. The tree layer is still good developed, but with a less diverse stand
structure and some gaps in the canopy. The intermediate disturbed forest is more suffused
with light. This stage was common and widespread, especially along the main roads from
Lenkoran to Lerik and from Masally to Iardimli.
Park-like forest stage
This degradation stage is characterised by a high logging and grazing activity, which lead
to a park-like appearance (Figure 8). Trees are mostly cut down and only single old-
growth trees are left, which mostly have a chopped appearance due to lopping. Snags are
common. The foliage cover of the tree layer is low (ca. 5-10%). The herb layer is short
due to grazing. Dense forest fragments can occur. This stage can be found closer to
villages (1-2 km), on ridge tops or in plains. Large areas of this stage could be found e.g.
near Dashtatuk, Bilarsar, Tankivan or Günesli.
Shrubby woodland stage
The shrubby woodland stage is situated close to villages. It is the result of a frequently
used former forest. Large trees are almost completely removed and shrubs are dominating
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Study site
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(Figure 9). Due to lopping and pollarding, trees appear like small bushes, especially Car-
pinus betulus and Quercus castaneifolia, which rarely exceed heights of more than three
metres. Grazing intensity is very high. This stage can also be found at higher altitudes
along the upper tree line. Here, the occurrence is probably naturally triggered, as a growth
of trees is inhibited by the rougher climate (lower precipitation, lower mean annual temp-
erature, strong winds).
I neglected the described treeless bracken fern stage that is dominated by Pteridium
aquilinum. It is limited to a few small locations only and could not be adequately sampled.
The work of Rietschel (2010) confirmed the classification of Scharnweber et al. (2007).
Figure 7: Widespread natural forest stage of the Caspian forest. Siov, 26.04.2008.
Study site The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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Montane meadow belt
Montane meadows are distributed above the forest belt around the smaller towns Lerik,
Shingedulan and Iardimli. Due to deforestation, it has an open landscape character with
gentle slopes ranging from 700 to 1200 m a.s.l. (Figure 4). Typical are meadows, which
are used for hay making (Figure 10). Furthermore, agriculture and pastures are common.
Figure 8: Grazing is com-mon in the park-like forest stage. Lopped trees are visible in the background. Zunguliash, 28.05.2008.
Figure 9: Intense grazing and lopping is responsible for a shrubby appearance of the forest. Logging activity completely re-moved larger trees. Shinaband, 06.05.2008.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Study site
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Figure 10: Hay making in the montane meadow belt. Muria, 05.06.2008.
Forest-like structures are rare, but along steep slopes the shrubby woodland stage is partly
developed. Otherwise, shrubs are scattered along field paths or roads. Radde (1886a)
described the area around Lerik 130 years ago also as an open landscape with agriculture.
Zuvand
South of Lerik, the villages Mistan, Kialvaz, Allar, Gosmalian and Shonadzohla comprise
the Zuvand upland (Figure 1). Here, three landscape types can be found:
Montane semi-desert
Gentle slopes barely covered with thorny cushion-forming tragacanthic vegetation (e.g.
Tragacantum, Acantholimon) feature the montane semi-desert (Atamov et al. 2006,
Hajiyev 2006, Michael Succow Stiftung 2009) (Figure 11, Figure 12). Tree growth is
prevented due to low annual precipitations and long summer droughts (Figure 3).
Study site The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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However, a stand of Juniper excelsa in a runnel near Diwagatsh may indicate the
distribution of a former Irano-Turanian steppe-forest which can be found in the adjacent
Iran (Zohary 1973). The altitudinal distribution of the montane semi-desert ranges from
1400 to 2500 m a.s.l. (Figure 4, Figure 1). Humans use the semi-desert mainly as pastures.
At a few locations, I observed transformations into cultivated land.
Riparian forest
Embedded in the barren montane semi-desert along small rivulets, open park-like riparian
forests of willow species (Salix alba, S. purpurea, S. caprea) and Populus nigra can be
found (Michael Succow Stiftung 2009). Riparian forests range from 1200 to 1800 m a.s.l.
and are located for example around the villages Gosmalian, Mistan, Kialvaz and Govari.
They are rather narrow stripes and restricted to water containing valley bottoms (Figure
13). The inhabitants of the villages along the forest created a complex irrigation system to
water their lush and flower-rich meadows, fields and gardens (Figure 14). Large terraces
have been built and are widely used as orchards or almond plantations. The vegetation
structure of this type is similar to the open appearance of the park-like stage within the
forest belt.
Figure 11: Montane semi-desert of the Zuvand. Divagach, 08.05.2008.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Methods
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Wesche 2007), I took the square root of the dissimilarity matrix. The abundance data have
not been transformed and contains rare species.
Basing on the output of the cluster analysis, I then created a bird community table,
where the communities are arranged following an altitudinal gradient. Within the forest
belt, the order followed the intensity of degradation from natural forest stage to the
shrubby woodland stage. The socio-ecological species groups were clustered by
transposing the dissimilarity matrix to obtain a rough overview, and I finally arranged
them manually to their best fit with the help of my expert knowledge and further
observations besides the surveying time.
I also used NMDS (non-metric multidimensional scaling) to plot ordinations using the
function “metaMDS” of the package “vegan” of the statistical software package R. In
these NMDS are the transects arranged in scatter-plots regarding to their species compo-
sition and abundance (territories/km per transect) to illustrate similarities and dissimi-
larities between the transect and the gradients which influences their composition (Leyer
& Wesche 2007). NMDS base on Bray–Curtis measure (Bray & Curtis 1957) without the
square root of the dissimilarity matrix. I then used the stress-value to deter-mine the num-
ber of dimensions for the two NMDS (McCune and Grace 2002).
For identifying associations of species to communities or combinations of communi-
ties, the R package “indicspecies” (version 1.0) the function “multipatt” was applied using
1000 permutations. I rejected the indicator species analysis by Dufrene & Legendre
(1997) as it identifies species indicating only one community but not a combination of
communities.
In the following text, the code of the significance level is ‘*’ = p-value < 0.05, ‘**’ = p-
value < 0.01 and ‘***’ = p-value < 0.001.
Results The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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4 Results
I surveyed 178 km (kilometre) consisting of 94 transects (Figure 18). The length of
transects differed between 200 and 4500 metres. The average length was 1894 metres
(n=94). I excluded three transects from the cluster analysis and NMDS analysis, as their
length was below 500 m, which do not represent breeding bird communities adequately.
4.1 Species number and species richness
I found 10104 individuals of 197 bird species (Annex 1). I expect about 147 species of
them to breed in the Talish region (Table 1). (Further breeding birds of wetlands and
costal habitats are not included.)
From the faunistic aspect, the rediscovery of breeding Shikras (Accipiter badius) was a
surprising highlight, as no confirmed breeding records existed since 1933. This is the only
breeding site within the Western Palaearctic (Heiss & Gauger 2009). Further breeding
species are new to that region or rediscovered after more than a century like Prunella
ocularis, Rhodopechys sanguinea, Irania gutturalis or Bucanetes githagineus (Annex 1).
Table 1: Totals of surveyed transect length and totals of found breeding bird species per landscape types. Note that bird species can occur in more than one landscape type.
Landscape type Transect length [km] Species number
Caspian lowland 23.3 54
Natural forest stage 8.6 32
Slightly disturbed forest stage 12.1 40
Intermediate disturbed forest stage 26.1 46
Park-like stage 11.9 41
Shrubby woodland stage 20.4 59
Montane meadow belt 20.6 48
Riparian forest 15.7 44
Montane semi-desert 18.1 44
Rocky habitats 21.2 68
Total 178 147
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Results
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Results The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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However, several species listed in Agaeva (1979) or Patrikeev (2004) could not be con-
firmed in this study like Poecile hyrcana, Hippolais languida, Emberiza buchanani,
throgaster, Carpospiza brachydactyla or Tichodroma muraria.
The most species-rich landscape type are rocky habitats (Figure 19). Rather poor in
species are the less disturbed forest degradation stages (natural, slightly and intermediate
disturbed forest stages). According to this analysis, montane semi-desert and montane
meadow belt are also regarded as relatively poor in species. Interestingly, the heavily
degraded forest stages (park-like forest stage and shrubby woodland) show the highest
relative species richness.
Figure 19: Species-sampling effort relationship illustrated by the landscape types (Casp = Caspian lowland, Nat = Natural forest stage, Sli = Slightly disturbed forest stage, Int = Intermediate disturbed forest stage, Park = Park-like forest stage, Shr = Shrubby woodland stage, Mead = Montane meadow belt, Rip = Riparian forest, Mon = Montane semi-desert, Rock = Rocky habitats) and a fitted regression line (red line) indicating the same amount of relative species-richness. Landscape types above the regression line are rich in species and below are poor in species.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Results
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4.2 Breeding bird communities
Figure 20: The nine breeding bird communities of the Talish mountains represented by clusters. The cluster analysis bases on species abundance of each transect using Bray-Curtis measure (Bray & Curtis 1957) and Ward’s method (Ward 1963) as linking procedure. Cluster 6 consists of two merged clusters.
The cluster analysis identified ten breeding bird communities (Figure 20). I merged two
clusters (cluster 6) by a local reduction of the cut level, because in field both had a
shrubby appearance. Thus, I found nine breeding bird communities in the Talish moun-
tains.
The frequency table (Table 2) and the bird community table (Annex 3) show these 9
breeding bird communities and 27 socio-ecological species groups. Species of each socio-
ecological group reflect the same ecological demands and are similar distributed to the
communities. Due to the determination of clusters (communities) by breeding birds, some
transects are not within their assumed landscape type, for example transect 6 in
community 5 (Annex 3). This inaccuracy was neglected, because identifying habitats by
plants, as Schwarnweber et al. (2007) did and which I used for landscape type deter-
mination, and birds are different approaches.
Results The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
The analysis of parameters is important to understand ecological communities. For this
purpose, I carried out a NMDS analysis. This ordination is used to describe relationships
between species composition patterns and the influencing parameters onto a two
dimensional space. Figure 21 illustrates the nine breeding bird communities along two
ordination axis. The altitudinal gradient is the strongest site parameter, which is under-
lined by an approach on species-level (Figure 24). The altitudinal gradient ranges from
bottom left to top right with community 8 and 9 at the end of this vector representing the
highest altitudes. NMDS1 axis follows the vegetation structure of tree and shrub layer.
From left to right a gradient concerning naturalness of forest appearance can be derived.
Open landscape communities are on the right and forested landscapes are on the left. A
gradient following NMDS2 axis reflects the herb layer. Community 1 (Caspian lowland)
contains the best-developed herb layer, followed by community 6 (montane meadows).
Results The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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Figure 21: Ordination of breeding bird communities using a NMDS generated from abundances of all species of all transects after 20 random starts of a two-dimensional solution (final stress = 16.06, rmse = 0.002, max residual = 0.0072). Highly significant site parameters with a p-value < 0.001 are plotted as red vectors and other are black (for details, see Annex 5). Lengths of vectors indicate the significance of each parameter. Transect numbers are given in symbols. Parameter abbreviations: ALTI = altitude, DATE = date, EXPO = slope exposition (after Parker 1982), SLO = slope steepness (after Parker 1982), HH = Herb layer height, HC = Herb layer cover, SH = Shrub layer height, SC = Shrub layer cover, LTH = Lower tree layer height, LTC = Lower tree layer cover, UTH = Upper tree layer height, UTC = Upper tree layer cover. Landscape abbreviations: Casp = Caspian lowland, Nat = Natural forest, Int = Intermediate disturbed forest, Stro = Strongly disturbed forest, Shr = Shrubby woodland, Mead = Montane meadows, Rip = Riparian forest, Semi = Montane semi-desert, Rock = Rocky habitats.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Results
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Figure 22: Ordination of breeding bird communities within the forest degradation stages using a NMDS generated from abundances of all species of transects of community 2, 3, 4 and 5 after 100 random starts of a two-dimensional solution (final stress = 15.83, rmse = 0.00015, max residuals = 0.00054 after 81 tries). Highly significant site parameters with a p-value < 0.001 are plotted as red vectors and other are black (for details, see Annex 6). Lengths of vectors indicate the significance of each parameter. Transect numbers are given in symbols. Parameter abbreviations: ALTI = altitude, DATE = date, EXPO = slope exposition (after Parker 1982), SLO = slope steepness (after Parker 1982), HH = Herb layer height, HC = Herb layer cover, SH = Shrub layer height, SC = Shrub layer cover, LTH = Lower tree layer height, LTC = Lower tree layer cover, UTH = Upper tree layer height, UTC = Upper tree layer cover. Landscape abbreviations: Nat = Natural forest, Int = Intermediate disturbed forest, Stro = Strongly disturbed forest, Shr = Shrubby woodland.
Results The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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To get a deeper insight into the forest degradation stages and their explanatory vari-
ables, only transects of community 2, 3, 4 and 5 have been calculated again in a further
NMDS (Figure 22). These transects were conducted within the forest belt (Figure 4). I
excluded transect 5 and 6 as they belong to other landscape types, whereas transect 73 was
treated as belonging to community 5, as it was securely made in the shrubby woodland
stage and only due to the cluster analysis considered as belonging to community 6.
NMDS1 axis reveals a degradation gradient (Figure 22). Vectors of the tree layer are
pointing towards transects performed in natural and slightly disturbed forests and vectors
of the shrub layer into the opposite direction towards transects of the shrubby woodland
communities. The altitudinal gradient also plays an important role, but is less significant
compared to the landscape-scaled approach in Figure 21.
Each bird species and therefore each bird community has a different ecological de-
mand. The differences of each community according to their preferred vegetation struc-
tures are given in Figure 23 and the header data of Table 2. Communities 2-5 and 7 re-
present forested landscapes. The height and cover of the upper tree layer decreases conti-
nuously from community 2 to 5, whereas the cover of the shrub layer increases. This gra-
dient also appears in Figure 22. The breeding bird communities clearly show a response to
the vegetation parameters resulting in different values. Interestingly, these results are
similar to Scharnweber et al. (2007), which base on vegetation analysis. For comparison,
communities besides the forest degradation stages are also included. Community 1, 6, 8
and 9 are different. They present open landscapes with a missing or reduced tree layer.
4.4 Relative abundances of bird species
Due to the heterogeneity of several transects according to their landscape composition,
especially within the forest belt, where transects consisted often of several degradation
stages, the calculation of relative abundances could not be performed basing on bird
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Results
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Figure 23: Boxplots of height and cover of each vegetation layer for every breeding bird community.
Results The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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Figure 24: Boxplots of the altitudinal distribution of breeding bird species basing on the num
ber of territories (n≥5).
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Results
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Figure 25: Relative abundances values of the 15 most common bird species per landscape type (forest degradation stages).
Results The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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Figure 26: Relative abundances values of the 15 most common bird species per landscape type (outside the forest belt).
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Results
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communities. Thus the relative abundance values base on the landscape type deter-
mination (chapter 2.3). This was in field more precisely and therefore used for further
calculations, especially to compare the species response in different forest degradation
stages.
Figure 25 shows the relative abundances of the most common bird species within the
forest degradation stages (forest belt). The most abundant species in the lesser disturbed
forests are Periparus ater and Fringilla coelebs. However, a comparison between bird
species is theoretically not allowed, because of the different detectability among species.
This comparison is also biased towards eye-catching and loud-voiced birds.
A shift regarding species composition and relative abundance takes place in the park-
like forest stage (Figure 25). The abundance of typical forest species decline with ongoing
4.5 Response to forest degradation on species-level
I found 32 breeding bird species within the natural forest stage. The obtained species
number is the result of a total transect of 8.6 km (Table 1). 28 species are regarded as
‘forest species’. I then converted the relative abundance values of each species per forest
degradation stage into percent values to compare the impact of human activities within the
forest belt. The natural forest stage serves as a gauge that gives information about a
pristine forest without any human activities. I used the converted abundance values of the
natural forest stage as reference values constituting 100%.
Negatively affected or strongly negatively affected
Figure 27 shows a clear negative response of forest species to forest degradation. Most of
these species prefer the forest interior. A decline of about 50% according to an unaffected
population is visible in the intermediate disturbed forest stage. Some species, like
Results The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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Fringilla coelebs, are also common in the park-like forest stage, but show a rapid decline
in the shrubby woodland stage. Only 6% or less of the bird population of strongly affected
species remain in the shrubby woodland stage. 65% of the 28 forest species show a neg-
ative or strongly negative response to forest degradation.
Figure 27: Negatively and strongly negatively response of selected forest bird species to forest degradation. Values are scaled relative to the natural forest stage (reference stage) constituting 100%.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Results
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Positively affected
Generally, open landscapes and shrub formations preferring species are positively affected
by forest degradation, e.g. Luscinia megarhynchos. Only a few species, treated as forest
species or at least ‘edge species’, show a positive trend (Figure 28). These species have
low abundances in natural forests and are common in degraded forests resulting in a vast
increase of a few species, e.g. Carduelis chloris. The treatment of some of those species
as forest species might be discussible. Only 21% of the 28 forest species show a positive
response to forest degradation. Most of them show a peak in the park-like forest stage, but
decrease in the shrubby woodland stage (Figure 28).
Figure 28: Positive response of selected forest bird species to forest degradation. Values are scaled relative to the natural forest stage (reference stage) constituting 100%.
4.6 Response to forest degradation on guild-level
The guild concept is needed because it can help to identify the habitat characteristics that
determine the structure of bird communities and because it would be possible to under-
stand processes organising communities (Casenave et al. 2008).
In total, the natural and slightly disturbed forest stages reach their highest relative
abundance values within the forest belt. The shrubby woodland stage holds the lowest
values (Figure 29).
Results The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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The dominating guild within all degradation stages are cavity breeders. This guild in-
cludes typical cavity breeders, like Dendrocopos major or Cyanistes caeruleus, and semi-
cavity breeders, like Certhia familiaris. Cavity breeders reach their highest relative abun-
dances in natural and slightly disturbed forests with 25.2 and 24.8 territories per km,
respectively. In contrast, only 6.5 territories per km of cavity breeders occur in the
shrubby woodland stage. Their proportion of about 40% is in all degradation stages more
or less equal, besides the shrubby woodland stage. Here, only 17% belong to the cavity-
breeding guild owing to the lack of cavities in this stage.
Canopy breeders are common in all degradation stages with 12.0 to 16.2 territories per
km, except for the shrubby woodland stage with only 7.1 territories per km. The highest
percentages can be found in intermediate disturbed forests and park with 32% each.
Shrub breeders dominate the shrubby woodland stage. Here, the highest proportion
(37%) and relative abundance (14.2 territories per km) can be found.
Birds that breed on the ground show a decline in relative abundance and proportion
concerning a degradation gradient from high values in natural forest stage and low values
in park-like stage. On the contrary, the values of relative abundance and proportion rise
again in the shrubby woodland stage.
Figure 29: Response of nesting guilds to forest degradation. (Nat = Natural forest stage, Sli = Slightly disturbed forest stage, Int = Intermediate disturbed forest stage, Park = Park-like forest stage, Shr = Shrubby woodland stage).
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Discussion
- 44 -
5 Discussion
5.1 Breeding bird communities
The cluster analysis identified 9 breeding bird communities (Figure 20). Every community
is representing more or less one of the landscape types. Especially transects conducted
outside the forest belt show an apparent overlapping (header data of Table 2). Within the
forest belt, a differentiation of forest degradation stages by breeding bird communities is
imprecise owing to the problems of the survey method. For the cluster and NMDS
analysis each transect was treated as a single record, despite the differing transect length
and differing landscape type compositions. In a few cases, transects consisted of a mixture
of several degradation stages. For example, a 2000 m long transect contains 1500 m park-
like forest and 500 m shrubby woodland stage. The cluster algorithm may classifies this
transect to a community representing the park-like stage including 500 m of shrubby
woodland. A wrong determination of the landscape types in field could also cause blurred
results concerning the landscape composition of each community. Within the forest de-
gradation stages, it was sometimes difficult to separate the stages from each other. For
example, the natural forest stage was difficult to distinguish from the slightly disturbed
forest stage, as both are very similar. A further important point, regarding the landscape
composition of each breeding bird community, is the early date of surveying of a few
transects. The species composition of early transects (beginning of April) lacks most long-
distance migrants. These species contribute also to the classification of transects to a
community by the cluster analysis and if they are missing, the cluster analysis classifies
the transects only by the resident species. This problem could not be adequately solved by
beginning the survey when long-distance migrants completely arrived the Talish moun-
tains (end of May/beginning of June). Several residents already fed chicks and show a
reduced territorial behaviour, which leads to the opposite effect of ‘missing’ residents,
when the territorial activity of migrants peaks. The only way to solve this problem would
be a repeated surveying of each transect, but this was refused (chapter 2.6.). However, on
this landscape-scaled approach, open habitats fit to the open land breeding bird com-
munities. Within the forest belt with its degradation stages, it can only roughly be as-
Discussion The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
- 45 -
signed (header data of Table 2), but a seperation following a man-made degradation
gradient is clearly visible (Figure 21, 22).
Each community differs according to their species composition and relative abundance
values (Table 2, Annex 3). Interestingly, on this landscape-scaled approach, which se-
parated for example Caspian lowland from rocky habitats by breeding birds only, a dif-
ferentiation between forest communities (community 2, 3, 4 and 5) is evident. This is
remarkable as the clusters of the cluster analysis were cut at the same level, representing
an equal degree of similarity (Figure 20). However, the differences in bird species
composition between communities of the forest belt are low (Table 2, Annex 3). Within
the forest communities, indicator species occurred only in community 2 (natural forest):
Pyrrhula pyrrhula***, Certhia familiaris**, Ficedula semitorquata** (Annex 4). These
few indicator species are in other forest communities missing, which underlines the strong
similarity between these communities. As the cluster analysis separates clusters by species
and their abundances, hence the forest communities are mainly differentiated by the
relative abundance values. The differentiation by species inventory plays a minor role in
the forest communities. Several forest bird species, like Periparus ater, Troglodytes
troglodytes, Fringilla coelebs or Sitta europaea, reach their highest relative abundances in
community 2 representing natural forests (Table 2, Annex 3). In contrast, Tomiałojć &
Wesołowski (2004) found remarkably low abundance values in the primeval forest of
Białowieża (Poland) compared to data from man-transformed places (e.g. fragmented
forests of Western Europe). Species of community 2 are also present in communities
representing strong forest degradation, for example community 5 containing 54% of the
shrubby woodland stage. These differences of relative abundance values are responsible
for the separation of bird communities by the cluster analysis within the forest belt
(degradation stages), despite the low differences in species composition.
However, to give community 2, 3, 4 and 5 the status of a breeding bird community
might be discussable due to the similar species inventory. The separation of forest
breeding bird communities by species only is difficult or almost impossible. Here, the
main problem occurs for an practical use of this from the vegetation ecology derived
concept. Other problems are the migratory life history traits of birds. To evaluate the
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Discussion
- 46 -
quality of a landscape or a habitat by the development of a breeding bird community is
under the circumstance of a not sessile living bird inaccurate. The reason for the presence
or absence of a species might not be a loss of habitat quality on the breeding site, it can
also be hunting pressure on migration routes or quality loss on the wintering sites. Plants
are therefore more practical for the evaluation of habitat quality as they spend their whole
life on one location. Furthermore, the determination by their abundance values is not
practical in field, as it depends on a large amount of surveying time, which avoids a quick
identification of each forest community. Nevertheless, human activity created different
breeding bird communities within the forest belt, which are statistically verified in a
landscape-scaled approach.
5.2 Parameters influencing bird communities
The vegetation structure is the key parameter influencing the composition of breeding bird
communities in the Talish mountains (Figure 21-23). Heterogeneous vertical and
horizontal vegetation structures are an important factor influencing breeding bird
communities (abundance and diversity), because it offers a large number of ecological
niches for a broad variety of bird species. (Mac Arthur & Mac Arthur 1961, Mac Arthur
1968, Erdelen 1984, Tews et al. 2004, Kati & Sekercioglu 2006). Plant species diversity
of (temperate) forests is less important and has nothing to do with bird species diversity
(Mac Arthur 1961). Thus, I expect the high diversity of woody plant species in the Talish
mountains has no effect on the species diversity of bird communities, although it was not
tested in this study. For the formation of each breeding bird community is the develop-
ment of the tree layers, as the main characteristic of the vertical vegetation structure with
different tree heights and cover values, most important (header data of Table 2, Figure
23). In Figure 21 and Annex 5 are information given about the importance of each
vegetation parameter. Every vegetation parameter is highly significant in this large-scaled
approach. At a smaller-scaled approach, regarding forest communities (2, 3, 4 and 5) only,
height and cover of the upper tree layer, height of the lower tree layer and cover of the
shrub layer are among all vegetation parameters these with the highest
Discussion The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
- 47 -
significance (p-value < 0.0001) (Figure 22, Annex 6). High values of tree height and tree
cover indicate the occurrence of forests, which is obviously crucial to the bird species
composition of each community. If trees, as main contributor to the vertical vegetation
structure, are missing, completely different bird communities are developed. The resulting
open land communities consist mainly of open land species like Alauda arvensis, Hirundo
rustica or Emberiza melanocephala. This separation from open land and forest com-
munities is in agreement with other studies (e.g. Kati & Sekercioglu 2006).
A removal of forests would cause a species turnover by extirpating of most forest bird
species and colonisation of open land species (Figure 21, 30). This is also observable in
tropical rain forests after intensive logging or clear-cutting (e.g. Thiollay 1999, Patten et
al. 2010). A possible recolonisation of forest species could never bring back the whole
assemblage due to the extinction of some bird species. The British avifauna for example,
lacks of several forest bird species, for which no geographical or biological reasons exist,
probably caused by a widespread removal of forests centauries ago (Fuller et al. 2007a,
Figure 30: Breeding bird response to forest degradation. Summarised schematic of the forest degradation stages.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Discussion
- 48 -
Wesołowski 2007). Nevertheless, a complete removal of the Caspian forest in Azerbaijan
is unlikely, but the current forest degradation changes the vertical vegetation structure,
which affects most forest birds negatively (Figure 27).
The differences in the vertical vegetation structure lead to the differing appearance of
breeding bird communities, which roughly represent each degradation stage. As already
mentioned, each forest community is more differentiated by the species abundance then
the species composition. High trees and high foliage cover are typical for undisturbed
primeval forests (Wesolowski 2007). Trees provide cavities for hole nesters and their
crowns offer canopy breeders nesting opportunities. The older and larger the trees are the
more cavities are available and the higher is the abundance of cavity breeders in the
natural forest stage compared to other degradation stages (Enoksson et al. 1995, Poulsen
2002) (Figure 29). The high proportion of cavity breeders is typical for European old-
park-like forest and shrubby woodland) according to relative abundances of bird species.
From a faunistic aspect, the breeding records of Radde’s Accentor (Prunella ocularis),
Crimson-winged Finch (Rhodopechys sanguinea), White-throated Robin (Irania guttu-
ralis), Trumpeter Finch (Bucanetes githagineus) and Shikra (Accipiter badius) are of
national importance, as they are newly discovered breeding birds or rediscovered after
many decades.
The cluster analysis revealed nine breeding bird communities. They are arranged
mainly along an altitudinal gradient ranging from the Caspian lowland to montane semi-
deserts. Four of the communities are within the forest belt and a result of forest
degradation. 65% of the forest-dependent bird species are negatively affected by forest
degradation. Several species, like Eurasian Bullfinch (Pyrrhula pyrrhula), Semi-collared
Flycatcher (Ficedula semitorquata) and Eurasian Treecreeper (Certhia familiaris), are ex-
pected to become extinct with ongoing degradation. Nine endemic subspecies are
threatened.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Summary
- 56 -
The differences between natural forests and only slightly disturbed forests are low
regarding species composition and relative abundance. With further degradation, the
relative abundance and the number of forest-dependent species decreases, but the total
species number increases. The strong vertical vegetation heterogeneity of natural forests
with high and old-growth trees, multistorey profiles and a large amount of dead wood
cause a high abundance of forest-dependent species. Natural forests are horizontally
homogenous landscapes resulting in a lower total species number. In contrast, degraded
forests have a reduced vertical heterogeneity, which reduces the abundance of forest
species. The strong horizontal heterogeneity of degraded forests including shrub
formations, different-sized forest fragments and open lands (pastures) enables a broad
variety of bird species breeding opportunities resulting in high species richness.
A slightly use of the Caspian forest does not have a serious impact on the forest
avifauna. Hence, a natural and sustainable forestry, conserving a rich vertical vegetation
structure, would protect the forest avifauna and prevent a loss of global biodiversity.
7 Zusammenfassung
Primäre Laubwälder gehören global zu den am stärksten bedrohten Ökosystemen. In
Europa finden sich nur noch kleine, isolierte Reste primärer Laubwälder. Das weltweit
größte zusammenhänge Primärwaldgebiet, der Kaspische Wald, befindet sich im Norden
des Irans und im angrenzenden Aserbaidschan. In Aserbaidschan werden 40358 ha im
Hirkan National Park geschützt. Außerhalb des Nationalparks führen Viehwirtschaft und
intensive Holznutzung zu einer Degradierung und Fragmentierung des Waldes. Die
vorliegende Arbeit untersucht die Brutvogelgemeinschaften des Talisch Gebirges in
Aserbaidschan. Zudem wurde deren Verhalten unter dem Aspekt der fortschreitenden
Walddegradierung betrachtet.
Im Frühjahr 2008 führte ich die Geländeuntersuchung mittels der Transektmethode
durch. Anhand der gewonnenen Daten wurden die errechneten Brutvogelgemeinschaften
mit Umweltparametern korreliert. Dazu wurde die berechnete Abundanz pro Transekt für
Summary The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
- 57 -
die Clusteranalyse und Ordination (NMDS) benutzt. Des Weiteren verglich ich fünf
Degradierungsstadien des Waldes (Naturwald, leicht gestörter Wald, mittel gestörter
Wald, parkartiger Wald and Gebüschwald).
Unter faunistischen Gesichtspunkten sind Brutnachweise bzw. Brutzeitfeststellungen
von Steinbraunelle (Prunella ocularis), Rotflügelgimpel (Rhodopechys sanguinea),
Weißkehlsänger (Irania gutturalis), Wüstengimpel (Bucanetes githagineus) und Schikra
(Accipiter badius) von nationaler Bedeutung, da diese Arten neu entdeckt bzw. nach
vielen Jahrzehnten wiederentdeckt wurden.
Die Clusteranalyse ergab neun Brutvogelgemeinschaften. Diese sind hauptsächlich
entlang eines Höhengradienten angeordnet, welcher vom Kaspischen Tiefland bis in die
Gebirgshalbwüste verläuft. Vier der Gemeinschaften befinden sich im Waldgürtel und
sind das Ergebnis der Walddegradierung. 65% der Waldvogelarten sind von der
Walddegradierung negativ betroffen. Mehrere Arten, wie z.B. Gimpel (Pyrrhula
pyrrhula), Halbringschnäpper (Ficedula semitorquata) und Waldbaumläufer (Certhia
familiaris), werden mit fortschreitender Degradierung im Talisch Gebirge aussterben.
Neun endemische Unterarten sind bedroht.
Die Unterschiede zwischen Natur(Primär-)wäldern und nur leicht gestörten Wäldern
sind gering bezogen auf das Arteninventar und deren relativen Abundanz. Mit
zunehmender Degradierung verringert sich die Abundanz und Artenzahl von
Waldvogelarten bei einer Zunahme der Gesamtartenzahl. Die starke Heterogenität der
vertikalen Bestandsstruktur von Naturwäldern, mit hohen und alten Bäumen, vielen
Vegetationsschichten und einem hohen Anteil an Totholz, verursachen eine hohe
Abundanz von Waldvögeln. Aber ihre räumliche (horizontale) Homogenität verursacht
eine vergleichsweise geringe Gesamtartenzahl. Im Gegensatz zu Naturwäldern haben
degradierte Wälder eine verringerte Heterogenität der vertikalen Bestandsstruktur, die mit
einer Reduzierung der Abundanz von Waldvögeln einhergeht. Die starke räumliche
Heterogenität von degradierten Wäldern (z.B. durch Gebüsche, lichte und geschlossene
Waldfragmente, Viehweiden), ermöglicht einer großen Anzahl von Vogelarten
Brutmöglichkeiten, die eine hohe Gesamtartenzahl bewirken.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Summary
- 58 -
Eine geringfügige Nutzung der Kaspischen Wälder hat keinen schwerwiegenden
Einfluss auf die Waldavifauna. Daher würde eine naturnahe und nachhaltige
Forstwirtschaft, welche eine abwechslungsreiche vertikale Bestandsstruktur über weite
Flächen bewahrt, dem Schutz der Waldavifauna dienen und damit eine globale
Verarmung an Biodiversität verhindern.
8 Acknowledgements
I am grateful to Prof. Dr. Michael Succow and Dr. Martin Flade for the supervision of my
diploma thesis. The Ministry of Ecology and Natural Resources of Azerbaijan Republic
kindly granted the necessary access to the Hirkan National Park. I thank Kai Gauger for
the indispensable support during the fieldwork. I also wish to acknowledge Jan Peper for
useful critics and the brilliant introduction to multivariate statistics. Furthermore, I thank
Nigar Agaeva, Jonathan Etzold, Benjamin Herold, André Jankowski, Dr. Vladimir M.
Loskot, Prof. Dr. Michael Manthey, Dr. Hartmut Müller, Jan Peters, Tobias Scharnweber
and Sebastian Schmidt for providing literature and useful comments. The people of the
Talish mountains, especially Novrus Hüseynov and his family, Akif Aliyev and
Babakhan, also contribute with their support and overwhelming hospitality to this work.
This study could not be accomplished without the financial support of the DAAD
(German Academic Exchange Service) and the help of the Michael Succow Foundation.
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10 Appendices
Annex 1: Commented list of the observed bird species of the Talish mountains region including breeding status (b = breeding, n = non-breeding), based on own observations and interpretations, and IUCN Red List (2010) category (EN = Endangered, VU = Vulnerable, NT = Near Threatened, Least Concern not mentioned).
No. Species Comment Status IUCN1 Cygnus cygnus 1 immature bird on 20 May near Narimanabad. n 2 Cygnus olor Up to 85 individuals in wetlands near Liman. n 3 Anas querquedula 1 bird near Narimanabad on 20 May n
4 Alectoris chukar Common at rock outcrops in Zuvand upland. 1 nest with 20 eggs found near Kialakhan on 2 June.
b
5 Tetraogallus caspius On 9 July a calling bird was heard in rocks near Khozavi. b
6 Francolinus francolinus
Only 2 displaying males observed on 5 May near Tazakend (Caspian lowland).
b
7 Perdix perdix Rare. A displaying male heard on 3 April near Aliabad (montane meadow belt). 2 birds in montane semi-desert near Nalabin on 24 June.
b
8 Coturnix coturnix Singing males were common in the montane meadow belt and the Caspian lowland.
b
9 Phasianus colchicus
No observation. Only a plucking of a female or juvenile found in shrubs at steep slope near Lerik on 14 April.
b
10 Podiceps cristatus A colony of 49 breeding pairs in wetlands near Liman on 20 May. 2 chicks also present.
b
11 Phalacrocorax carbo 13 birds in wetlands near Liman on 20 May. n
12 Phalacrocorax pygmaeus
Some observations around Liman, Narimanabad and Lenkoran river.
(b) NT
13 Ardea cinerea Single birds throughout the Caspian coast and lowland. (b) 14 Ardea purpurea Found in lowland, especially in wetlands near Liman. (b) 15 Ardea alba 1 adult in costal lagoons near Narimanabad. (b) 16 Egretta garzetta Common in wetlands of the Caspian lowland. (b) 17 Ardeola ralloides Up to 30 birds on 20 May in wetlands near Liman. (b)
18 Bubulcus ibis 2 observations in the Caspian lowland in May. On 6 May 5 birds foraging in montane meadow near Aliabad.
(b)
19 Nycticorax nycticorax Regularly seen in the lowland, especially in wetland near Liman. (b)
20 Ixobrychus minutus
Occurred in the lowland. 4 migrants on 15 May in riparian forest near Gosmalijan.
b
21 Plegadis falcinellus Up to 200 birds in wetlands near Liman on 2 July. (b)
22 Ciconia nigra Occasionally seen in the Caspian lowland and the forest belt. b 23 Ciconia ciconia Breeds in villages in the lowland, but is rather rare. b 24 Pernis apivorus A few migrants seen from 7 to 14 May. (b) 25 Milvus migrans 1 bird on 10 May near Ashagy Bilnia and 1 bird on 27 May in (b)
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Moscow forest.
26 Neophron percnopterus Only a few times seen near Lerik and in Zuvand. (b) EN
27 Gyps fulvus Only observed in Zuvand. 1 near Divagach on 8 May and 16 near Mistan on 15 May.
n
28 Circaetus gallicus Regularly seen at altitudes below 1200 m a.s.l. b
29 Circus aeruginosus Breeds in the Caspian lowland. b
30 Circus macrourus Migrant. n NT
31 Aegypius monachus 1 individual on 3 April near Aliabad. n NT
32 Accipiter badius 9 territories were found in the Caspian lowland around Masally and Lenkoran (Heiss & Gauger 2008).
b
33 Accipiter nisus Nine observations from April to July . b 34 Accipiter gentilis Twice seen in April in the lower foothills. (n) 35 Buteo buteo Common in the forest and montane meadow belt. b
36 Buteo rufinus Regularly seen in Zuvand upland with territorial behavior of a pair west of Pirasora
b
37 Aquila pomarina Displaying individuals in forest near Günesli and montane meadows near Ashagy Bilnia. Regularly seen within the forest belt.
b
38 Aquila nipalensis Several migrating birds seen from 3 April to 20 Mai. n
39 Aquila chrysaetos Regularly seen in Zuvand. 1 abandoned nest found in rocks near Shonadzhola.
b
40 Aquila pennata Few observations within the forest belt. b
41 Falco naumanni A small colony with 6 breeding pairs was found on 19 April in a building in Khialakhan.
b VU
42 Falco tinnunculus Breeds in the riparian forest and in rocky habitats. Only 1 observation in the Caspian lowland on 20 May near Boladi.
b
43 Falco subbuteo Nests in the riparian forest, but was also observed in the Caspian lowland (near Tazakend, Moscow forest).
b
44 Falco biarmicus An adult seen on 6 July on rocks near Pirasora. n 45 Falco peregrinus 1 individual on 9 July west of Khozavi. (n)
46 Crex crex One singing male on 10 May in monane meadows near Ashagy Bilnia.
(b) NT
47 Gallinula chloropus
An adult with a juvenile observed at a back water of the Lenkoran river in the lowland.
b
48 Charadrius dubius
Was seen at costal lagoons near Narimanabad and Lenkoran river.
(b)
49 Himantopus himantopus 10 birds in wetlands near Liman on 10 July. b
50 Limosa limosa Migrant. 70 birds on 2 July and 30 birds on 10 July in wetlands near Liman.
n NT
51 Tringa totanus 10 birds on 2 July in wetlands near Liman. n
52 Tringa ochropus Migrant. Observed along rivulets of the riparian forest and Lenkoran river near Vel.
n
53 Actitis hypoleucos
Two displaying individuals on 22 April at Lake Xanbulan are maybe migrants. Resting birds were found in the beginning of July at several locations in the Caspian lowland (costal lagoons near Narimanabad, wetlands near Liman, Lenkoran river).
(b)
54 Arenaria 1 at a costal lagoon near Narimanabad on 20 and 24 May. n
Appendices The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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interpres
55 Philomachus pugnax 1 bird in wetlands near Liman on 10 July. n
56 Glareola pratincola
Occurred only in the lowland. 2 birds near Sirabil on 21 May and 4 birds at costal lagoons near Narimanabad on 20 May.
(b)
57 Larus cachinnans Regularly at the Caspian coast. b
58 Chroicocephalus ridibundus Regularly at costal lagoons and Lenkoran river (lowland). (n)
59 Chroicocephalus genei 30 individuals on 20 May near Liman. (n)
60 Thalasseus sandvicensis 10 birds on 20 May at the Caspian coast near Narimanabad. n
61 Sterna hirundo A few observations in wetlands near Liman and the Caspian coast.
n
62 Sternula albifrons A few observations in wetlands near Liman and the Caspian coast.
n
63 Chlidonias hybrida Common, with up to 2500 individuals in wetlands near Liman. b
64 Chlidonias leucopterus
Common in wetlands near Liman. 1 migrating flock with 150 individuals was seen on 2 May at 2300 m a.s.l. near Mistan.
(b)
65 Stercorarius parasiticus An adult seen at the Caspian coast near Narimanabad. n
66 Columba livia A wild bird was seen on 1 June near Lialiakeran (Zuvand). b 67 Columba oenas Common in the forest belt. b
68 Columba palumbus
Common in the riparian forest. Only 3 observations within the forest belt with 1 singing male near Günesli and 1 singing male near Tankivan.
b
69 Streptopelia turtur
A few singing males observerd in the montane meadow belt and Caspian lowland.
b
70 Streptopelia decaocto Occasionally seen in the Caspian lowland, e.g. Masally. b
71 Streptopelia senegalensis 1 bird at the Lenkoran river near Vel on 10 July. b
72 Cuculus canorus Common in the Talish mountains at all altitudes. b
73 Otus scops Several singing males with the montane meadow belt and the riparian forest. Only 1 singing male in the lowland near Hirkan village on 3 July.
b
74 Bubo bubo 3 fledged juveniles found on 4 and 5 July in a rocky cliff east of Mistan.
b
75 Strix aluco Almost everywhere heard at night within the forest belt. b
76 Athene noctua Seen in montane meadow belt near Bilaband and the riparian forest.
b
77 Caprimulgus europaeus
First seen on 22 April near lake Xanbulan. Singing males were found from the Caspian lowland to rocky habitats throughout all habitat types. 2 chicks were found on 11 July near Vel (lowland).
b
78 Tachymarptis melba
Regularly seen along rocky cliffs e.g. Pirasora, Nalabin, Mistan and Khozavi.
b
79 Apus apus Breeds in rocky habitats and towns throughout all altitudes. b
80 Alcedo atthis Occurred common along channels of the Caspian lowland. A few observations were made along rivers within the forest belt.
b
81 Merops persicus 1 colony with about 100 pairs found north of Kumbashi (Caspian lowland).
b
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82 Merops apiaster
During migration common throughout the Talish mountains. Breeds only at sandy cliffs of the montane semi-desert and along rivers of the lowand. Only small colonies were found with 2-3 pairs each.
b
83 Coracias garrulus Displaying individuals were found in the Caspian lowland and rarely in the montane meadow belt.
b NT
84 Upupa epops Regularly seen in villages throughout the Talish mountains, especially in riparian forest, montane meadow belt and Caspian lowland.
b
85 Jynx torquilla Several singing males within the forest belt, especially the park-like degradation stage.
b
86 Dendrocopos minor
Rare breeder of the forest belt. Occurred in the lowland only in the Moscow forest.
b
87 Dendrocopos major Common in the forest belt. b
88 Dendrocopos syriacus
Common in the riparian forest, but was also observed in the Caspian lowland near Kumbashi and Tazakend.
b
89 Dryocopus martius
1 singing male on 27 April in primeval forest near Siov at 800 m a.s.l.
b
90 Picus viridis Common in the forest belt and riparian forest. b
91 Lanius collurio Common wherever shrubs are present, especially in the montane meadow belt, Caspian lowland, riparian forest and shrubby woodland stage.
b
92 Lanius isabellinus Migrant. 1 individual on 9 May near Ambu and 1 individual on 18 May near Shinaband.
n
93 Lanius minor Common in the riparian forest, but occurred also in the montane meadow belt and Caspian lowland.
b
94 Lanius senator Only a few single birds in rocky habitats, riparian forest, montane meadow belt and Caspian lowland.
b
95 Garrulus glandarius Regularly seen in the forest belt. b
96 Pica pica Common breeder of Caspian lowland, montane meadow belt and riparian forest.
b
97 Pyrrhocorax pyrrhocorax A pair was several times observed in rocks west of Pirasora. b
98 Corvus frugilegus Common around Kialvas, where it breeds in a colony in riparian forest.
b
99 Corvus cornix Common breeder in open habitats, e.g. Caspian lowland, montane meadow belt, riparian forest.
b
100 Corvus corax Regularly seen in the forest belt and rocky habitats, where it breeds.
b
101 Melanocorypha calandra
Occurred in the montane meadow belt near Aliaband and Muria and in the Caspian lowland west of Masally.
b
102 Melanocorypha bimaculata Regularly in montane semi-desert above 1500 m a.s.l. b
103 Calandrella rufescens
Locally common in the Caspian lowland around Sirabil, Sarchuvar and west of Masally.
b
104 Galerida cristata Only 1 bird seen west of Masally. b 105 Lullula arborea Common in the montane meadow belt and Zuvand. b
106 Alauda arvensis Common in the montane meadow belt around Lerik and montane semi-deserts and subalpine meadows e.g. around Mistan, Orand and Pirasora. Did not occur below 1000 m.a.s.l.
b
Appendices The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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107 Eremophila alpestris
Breeds in the montane semi-desert. The first fledged juvenile was observed on 15 May near Mistan.
b
108 Ripparia ripparia Common migrant. No breeding colonies found. b
109 Ptyonoprogne rupestris Regularly seen at steep, rocky cliffs. 2 nests found near Nalabin. b
110 Hirundo rustica Breeds in villages and is common in the Caspian lowland. b 111 Delichon urbicum Breeds in rocky habitats and towns throughout all altitudes. b 112 Periparus ater Common breeder of the forest belt. b 113 Parus major Common throughout the Talish mountains. b
114 Cyanistes caeruleus Common within the forest belt and riparian forest. b
115 Remiz pendulinus Occurred in the Caspian lowland. b
116 Aegithalos caudatus Regularly seen in the forest belt. b
117 Sitta europaea Common in the forest belt. b 118 Sitta neumayer Common in rocky habitats. b
119 Certhia familiaris Only 4 singing males observed. 2 near Siov in natural forest and 2 near Sifiakeran in slightly disturbed forest.
b
120 Troglodytes troglodytes
Common in the forest belt. A rare breeder in riparian forest with a nest found near Mistan.
b
121 Cinclus cinclus Breeds along rivers throughout the Talish mountains except the Caspian lowland.
b
122 Cettia cetti
This species was distributed in two regions. It was common in riparian forest above 1200 m a.s.l., e.g. near Gosmalian, Shonadzhola and Mistan and occurred also in the Caspian lowland near Boladi and Liman along channels.
b
123 Locustella naevia Rare. 1 singing male on 28 April near Sim at a forest clearing was maybe a migrant. Furthermore, 1 singing male on 5 June near Bilaband in the montane meadow belt.
n
124 Locustella fluviatilis
Migrant. On 16 to 17 May a singing male in riparian forest near Gosmalian. On 19 May a singing male near Kumbashi. 2 singing males near Boladi on 20 May.
n
125 Acrocephalus schoenobaenus
Was treated as a breeding bird, but is probably only a migrant. Singing males were regularly seen along channels of the lowlands, but no observation was done after 1 June.
b
126 Acrocephalus scirpaceus
Due to the hidden behavior and the similar song with Hippolais pallida maybe overlooked. Only 3 observations. A singing male on 27 May in a park-like forest stage near Piran and a singing male in the riparian forest near Mistan on 1 June are migrants. A further singing male on 20 May near Boladi (Caspian lowland) occupied probably a territory.
b
127 Acrocephalus dumetorum
1 singing male on 19 May in bushes near Kumbashi. This species is not listed in Patrikeev (2004), but obviously a regular migrant in autumn (Gauger 2005). This is the first spring record to Azerbaijan.
n
128 Acrocephalus palustris
Was treated as a breeding bird, but is probably only a migrant. Was seen mostly in the riparian forest, but occurred rarely also in the montane meadow belt and the lowland from 11 May to 1 June.
(b)
129 Acrocephalus arundinaceus
Common along channels and reedbeds in the lowland. 1 territory was found near Aliabad with a singing male from 6 May to 5 June.
b
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Appendices
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130 Hippolais pallida Common in the Caspian lowland. First observed on 5 May. b
131 Hippolais icterina Common in lower altitudes of the forest belt (0-500 m a.s.l.). Singing males observed from 23 April to 21 June.
b
132 Phylloscopus trochilus Several singing migrants from mid-April to mid-May. n
133 Phylloscopus collybita
Maybe only a migrant. Several singing males heard at the begining of April. However, 4 singing males heard near Shinaband on 5 June.
n
134 Phylloscopus sindianus
Some singing males found in shrubs at higher altitudes above 1000 m a.s.l., but no sure signs of breeding.
(b)
135 Phylloscopus nitidus Common in the forest belt. b
136 Sylvia atricapilla Common in the forest belt. b
137 Sylvia borin Migrant. 1 bird on 16 May near Gosmalian, 1 bird on 17 May near Mistan and 1 bird on 18 May near Pirasora.
n
138 Sylvia communis Common in riparian forest. b
139 Sylvia curruca Several singing males in rocky habitats with bushes and in dense shrubs above 1000 m a.s.l.
b
140 Sylvia nisoria Probably an uncommun breeder in dense shrubs of higher altitudes. Observed near Shinaband, Mistan and Khosavi.
b
141 Sylvia mystacea Breeds rarely at a few locations in the Caspian lowland. Occurred as migrant also elsewhere, e.g. Zuvand.
b
142 Muscicapa striata Common in the forest belt and riparian forest. b
143 Ficedula hypoleuca
A male bird was observed on 4 April in riparian forest near Divagach.
n
144 Ficedula semitorquata Regularly seen in less degraded forests, especially around Siov. b NT
145 Ficedula parva Common breeder of the forest belt. Was at 3 April already present.
b
146 Erithacus rubecula Common in the forest belt. b
147 Luscinia megarhynchos
Common in montane meadow belt, riparian forest, shrubby woodland stage and Caspian lowland.
b
148 Irania gutturalis
Breeds in rocky habitats with bushes above 2000 m a.s.l. in Zuvand upland. 2 singing males and a female found on 30 May west of Orand. A total of 4 pairs found near Mistan. 2 families observed on 4 July with 3 and 2 juveniles each.
b
149 Cercotrichas galactotes
On 21 May 3 birds near Garibljar and on 24 May 1 singing male near Boladi.
b
150 Phoenicurus ochruros Common in rocky habitats. b
151 Phoenicurus phoenicurus Regularly in park-like forest stage and riparian forest. b
152 Saxicola rubetra Rare. Observed e.g. at forest clearings near Siov or near Mistan and Pirasora in Zuvand.
b
153 Saxicola torquatus Breeds in rocky habitats of Zuvand upland. b
154 Oenanthe oenanthe
Locally common in rocky habitats of Zuvand, e.g. near Pirasora. Also a rare breeder of the montane meadow belt.
b
155 Oenanthe finschii Common at rocky outcrops of Zuvand, especially near Pirasora, Kialvas and Kialakhan.
b
156 Oenanthe Rare breeder in rocky habitats of Zuvand. b
Appendices The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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hispanica melanoleuca
157 Oenanthe isabellina Common breeder of montane semi-desert. b
158 Monticola saxatilis Common in rocky habitats. b
159 Monticola solitarius
This species was only regularly seen in rocks between Shinaband and Nalabin. A pair was observed on 22 June near Kialakhan (K. Gauger pers. comm.)
b
160 Turdus torquatus Locally common in rocky habitats above 1500 m a.s.l., e.g. near Shinaband, Mistan, Nalabin, Kialakhan, Orand.
b
161 Turdus merula Common. b
162 Turdus philomelos Common in the forest belt. b
163 Turdus viscivorus Regularly seen in the forest belt, especially at the lower and upper treeline and the park-like forest stage. A family with 3 juveniles were found near Mamedoba at the lower tree line.
b
164 Oriolus oriolus Occurred in forest habitats. b
165 Sturnus vulgaris Common breeder of the riparian forest. Occurred also in montane meadow belt and Caspian lowland.
b
166 Pastor roseus Several migrating flocks seen in the beginning of May. Occurred also elsewhere, but disappeared at the end of May.
n
167 Prunella collaris Rare breeder in rocky habitats of altitudes above 2000 m a.s.l., e.g. Mistan, Pirasora.
b
168 Prunella ocularis
The specimen of the species was shot in June 1880 at the Kyziurdi mountain (Radde 1886a) and since then never again seen in the region (Patrikeev 2004). Dr. H. Müller rediscovered this species on 17 May in a rocky cliff with Juniperus spec. northeast of Mistan, where it was also seen on 22 June and 5 July by K. Gauger, J.Etzold and the author.
b
169 Prunella modularis
Rarely seen in the forest belt. Occurred also in riparian forest near Mistan.
b
170 Motacilla alba Regularly seen in villages, Caspian lowland and riparian forest. b 171 Motacilla citreola A bird on migration on 9 May near Dshangemiran. n
172 Motacilla flava feldegg
A locally common breeder of the lowland, especially around Boladi, Sirabil and Sarchuvar.
b
- Motacilla f. flava A pair was observed on 17 May near Pirasora. n 173 Motacilla cinerea Regularly seen along rivers of the forest belt. b
174 Anthus campestris Several territories found in Zuvand in 1400-2300 m.a.s.l in montane semi-desert.
b
175 Anthus trivialis Occurred in April in migrating flocks of about 5 birds. Singing males were observed since May e.g. around Dshangemiran, Shonadzhola or Pirasora.
b
176 Anthus pratensis 1 migrating bird near Aliabad on 4 April. n 177 Anthus cervinus 1 migrating bird near Ashagy Bilnia on 10 May. n
178 Anthus spinoletta 8 singing males and a food carrying bird was seen at 2000-2300 m.a.s.l. on a sub-alpine meadow between Razgov and Mistan.
b
179 Emberiza cia Common in rocky habitats. b
180 Emberiza hortulana
Common in rocky habitats around 2000 m a.s.l. Occurred during migration also elsewhere.
b
181 Emberiza melanocephala
Abundant in the montane meadow belt and also present in drier areas of the Caspian lowland. Arrived on 24 April from its
b
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Appendices
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wintering grounds.
182 Emberiza calandra
Common in the montane meadow belt and Caspian lowland. Occurred also along the riparian forest.
b
183 Fringilla coelebs Common and widespread species of the forest habitats. Occurred also in the lowlands along afforstations.
b
184 Carpodacus erythrinus
Regularly seen in the riparian forest above 1400 m a.s.l., but occurred also elsewhere.
b
185 Carduelis chloris Common below 1000 m a.s.l., above a rare sight. b 186 Carduelis spinus Regularly seen within the forest belt. b
187 Carduelis carduelis Common in the whole Talish mountains. b
188 Carduelis cannabina
Occurred above 1200 m a.s.l. near Lerik and was common in Zuvand upland.
b
189 Serinus pusillus Locally common in rocky habitats with a few shrubs, e.g. near Nalabin, Khozavi, Orand.
b
190 Pyrrhula pyrrhula Only a few observations within the forest belt. b
191 Coccothraustes coccothraustes Common in the forest belt. b
192 Rhodopechys sanguineus
1 pair at rocky cliffs northeast of Mistan and another pair, including a singing male, on a rocky outcrop north of Pirasora on 15 and 29 May, respectively.
b
193 Bucanetes githagineus
Fist seen on 8 May in Zuvand upland. Occurred regularly with up to 4 individuals near Divagach, Pirasora and Kialakhan. A fledged juvenile was seen in July near Divagach (C. Völlm pers. comm.).
b
194 Passer domesticus Common in villages, especially in the Caspian lowland with a large flock of 500 individuals near Hirkan village on 2 July.
b
195 Passer hispaniolensis 3 birds near Divagach on 22 June (K. Gauger pers. comm.). (b)
196 Passer montanus Only a total of 18 birds seen in the Caspian lowland. b 197 Petronia petronia Locally common breeder in rocks, e.g. west of Orand, Khozavi. b
Annex 2: Nesting guilds of selected bird species based on literature review (Flade 1994, Andretzke et al. 2005, Glutz von Blotzheim & Bauer 1991, Glutz von Blotzheim & Bauer 1994, Glutz von Blotzheim & Bauer 1998, Urquhart, & Bowley 2002, Kirwan et al. 2008, Alström & Mild 2003, Patrikeev 2004).
Bird species Nesting guild Bird species Nesting guild
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Appendices
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5+6+7+8+9 Lullula arborea 0.84 0.000999***
1+2+3+4+5+7 Parus major 0.922 0.000999***
Annex 5: Statistical analysis of the site parameters of the NMDS including all transects (Figure 21). ALTI = altitude, DATE = date, EXPO = slope exposition (after Parker 1982), SLO= slope steepness (after Parker 1982), HH = Herb layer height, HC = Herb layer cover, SH = Shrub layer height, SC = Shrub layer cover, LTH = Lower tree layer height, LTC = Lower tree layer cover, UTH = Upper tree layer height, UTC = Upper tree layer cover.
Site parameters r² Pr(>r)
ALTI 0.7225 0.0001***
SLOP 0.4568 0.0001***
UTH 0.7977 0.0001***
UTC 0.5126 0.0001***
LTH 0.6683 0.0001***
LTC 0.4452 0.0001***
SH 0.2757 0.0001***
HC 0.4185 0.0001***
SC 0.1629 0.0004***
HH 0.1579 0.0006***
DATE 0.0979 0.0109*
EXPO 0.0543 0.09119
Annex 6: Statistical analysis of the site parameters of the NMDS including all forest transects (Figure 22). ALTI = altitude, DATE = date, EXPO = slope exposition (after Parker 1982), SLO= slope steepness (after Parker 1982), HH = Herb layer height, HC = Herb layer cover, SH = Shrub layer height, SC = Shrub layer cover, LTH = Lower tree layer height, LTC = Lower tree layer cover, UTH = Upper tree layer height, UTC = Upper tree layer cover.
Site parameters r² Pr(>r)
UTH 0.7847 0.0001***
UTC 0.4377 0.0001***
LTH 0.5234 0.0001***
SC 0.3934 0.0001***
ALTI 0.3213 0.0007***
LTC 0.2197 0.005199**
DATE 0.1995 0.010799*
HH 0.1323 0.055594
SH 0.1348 0.058194
HC 0.1127 0.082692
EXPO 0.0654 0.258474
SLO 0.0414 0.431857
Appendices The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation
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Annex 7: Relative abundance values (territory/km) of each bird species per landscape type.
The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation Appendices
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Erklärung zur Diplomarbeit
Hiermit erkläre ich, die vorliegende Diplomarbeit mit dem Thema: „The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation“ selbstständig verfasst und keine anderen Hilfsmittel als die angegebenen verwendet zu haben. Aus anderen Werken in Wortlaut oder Sinngehalt entnommene Inhalte sind durch Quellenverweis, auch für Sekundärliteratur, kenntlich gemacht.
Greifswald, den 30.06.2010
Michael Heiß
Annex 3: Bird community table of the nine breeding bird communities including parameter values and species abundance per transect (territory/km).
To Diploma Thesis ‘The breeding bird communities of the Talish mountains (Azerbaijan) and their response to forest degradation’ by Michael Heiß