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Lesser Spotted Eagle Clanga pomarina Brehm, 1831 (Aves: Accipitridae) in

Dadia-Lefkimi-Soufli National Park, Greece: Population Trends and Spatial
Use in Respect to Continuous Land...

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 ACTA ZOOLOGICA BULGARICA
Research Article Acta zool. bulg., Suppl. 14, 2019: 7-14

Lesser Spotted Eagle Clanga pomarina Brehm, 1831

(Aves: Accipitridae) in Dadia-Lefkimi-Soufli National Park,
Greece: Population Trends and Spatial Use in Respect
to Continuous Landscape Changes during the Last 35 Years

Kostas Poirazidis1,2*, Vasileios Bontzorlos3, Stefan Schindler4,5 & Dimitris Vasilakis6

1
Department of Environmental Sciences, Ionian University, GR-29100, Panagoula, Zakynthos, Greece; E-mail: [email protected]
2
WWF Greece, 21 Lempesi Street, GR-11743, Athens, Greece
3
Department of Forestry and Management of Natural Environment, University of Applied Sciences of Thessaly, Mavromichali
str., 43100, PO Box 254,Karditsa, Greece; E-mail: [email protected]
4
Environment Agency Austria, Spittelauer Lände 5, 1090 Vienna, Austria; E-mail: [email protected]
5
Department of Botany and Biodiversity Research, University of Vienna, Rennweg 14, 1030 Vienna, Austria
6
Directorate of Evros Region Forestry Service, Hellenic Republic Decentralized Administration of Macedonia–Thrace, Kanari 12,
GR 68100, Alexadroupolis, Evros, Greece, [email protected]

Abstract: Lesser Spotted Eagle is a medium sized raptor with a stable population within its breeding range.
Nonetheless, its favourable habitats (a mosaic of forest and low-vegetation forest-openings, often with
water elements such as small ponds and flowing streams), are suffering from land abandonment as a result
of abrupt socio-economic changes related to rural depopulation and the decline of traditional agro-silvo-
pastoral practices. The species’ breeding population in Dadia National Park (Dadia NP) of approximately
20 pairs, is an important stronghold at the southern fringe of its European breeding range. The breeding
territories and their spatial distribution trends were studied in Dadia NP for 35 years (1979 to 2012). The
overall interannual trend was not statistically significant for the population, with a minimum value of 17
pairs (2001 and 2012) to a maximum of 22 (2005). Despite this long-term stability though, territories’
distribution in Dadia NP has changed remarkably. During the 1970s, a more uniform and clearly territo-
rial distribution had been observed in the western part of Dadia NP, the stronghold of the species. It was
an area characterized predominantly by forests, mixed with small-extension open landscapes, an optimal
foraging and breeding landscape for the species. In contrast, in the 21st Century the population has shifted
progressively to the eastern forest lowlands of Dadia NP adjacent to agricultural land, demonstrating a
clustered distribution with reduced territory distances in an agro-forest area, not considered optimal for
Lesser Spotted Eagle. This noticeable spatial change could be related to a decline in forest openings and
habitat mosaics to the western part, as a result of land abandonment. Thus, the most suitable region for the
species, decreased from 66% in 1979 to 54% in recent years, and specifically in Zone A of the National
Park the most radical change was observed (from 80% to 53%). In conclusion, increased landscape ho-
mogeneity and reforestation of open areas could be major threats to the viability of Lesser Spotted Eagle.
This has forced the species to shift its breeding territories already towards more humanized areas, in a
more clustered distribution and expose them to human induced risks. Conservation policies should strive
to increase heterogeneity at the landscape level and decrease the ecological impact of land abandonment.

Key words: Clanga pomarina, Dadia, Greece, landscape change, Maxent, niche shift, space use change

Corresponding author
*

7
Poirazidis K., V. Bontzorlos, S. Schindler & D. Vasilakis

Introduction (Natrix natrix Linnaeus, 1758) in its diet (42.3%,

Vlachos & Papageorgiou 1996). In Dadia NP, the
Lesser Spotted Eagle Clanga pomarina Brehm, eagle avoids north-facing slopes for nesting, al-
1831 is a medium sized raptor with a stable pop- though such sites provide protection from high sum-
ulation in most of its breeding range. Although it mer temperatures during the breeding season, which
is a species of Least Concern according to IUCN extends into July–August (Poirazidis et al. 2007).
(BirdLife International 2016), the Lesser A general deduction may be that this is an optimi-
Spotted Eagle is a priority species for conservation zation mechanism to avoid cold weather conditions
in the EU. During recent decades a negative trend that sometimes occur in the early breeding season
has been observed in many parts of its EU range (Kostrzewa & Kostrzewa 1990). Genetic analy-
(BirdLife International 2015)and a large-scale ses showed that the most common Baltic haplotype
action was drafted in a respective European Action was present also in the Dadia population of Lesser
Plan (Meyburg et al. 2001). Greece is forming the Spotted Eagle, indicating that the Balkan Peninsula
southern limit of the species’ western Palaearctic acted as a refugium for the European population
breeding range, and is estimated to host a small during the last ice age, constituting Dadia popu-
population of about 70-90 breeding pairs with nega- lation a source population for the other European
tive trend. It is considered an endangered eagle ac- populations; the northern regions were re-colonized
cording to the Red Book of the Threatened Animals after the deglaciation 8000±1500 years ago (Väli
of Greece (Legakis & Maragou 2009). Dadia- 2004).
Lefkimi-Soufli National Park (Dadia NP hereafter), In respect to social and economic changes that
a Natura 2000 area located in north-eastern Greece have both direct and indirect effects on the environ-
and renown as a major European biodiversity hot- ment, the abandonment of traditional rural and ag-
spot for raptors, holds the biggest population of this ricultural practices and professions as well as the
species in the country (Poirazidis et al. 2010a). depopulation of rural areas have often been indi-
The Lesser Spotted Eagle is a forest dwell- cated to produce landscape homogenization with
ing raptor with a preference for forest edges, which negative impacts on biodiversity (Zakkak et al.
offer rich food resources at a remarkable distance 2014). In specific, radical socio-economic changes
from human presence and settlements (Väli et al. that took place in the 1960’s all over Greece, e.g. ru-
2004, Mirski 2009). Throughout its breeding range, ral depopulation, development of the tourist indus-
it prefers mainly broadleaved forests (Treinys 2004, try (Xystrakis et al. 2017), along with European
Dombrovski & Ivanovski 2005, Mirski 2009) and agricultural policies (Renwick et al. 2013), played
during its foraging activity the species mainly preys an important role in fostering land abandonment
in forest openings, natural grasslands, small open throughout the rural areas in the country. That fact
valleys, low-height riparian vegetation, wetlands produced a number of ecological consequences at
and agricultural areas (Τreinys 2004, Meyburg et various spatial scales, with land abandonment of
al. 2004, Vali et al. 2017, Mirski et al. 2010, Zub rural areas being the first, removing the element
et al. 2010). In Dadia NP, it is considered a raptor of traditional human actions, which shaped habi-
with a narrow ecological niche, specialized in pine tat mosaics rich in biodiversity. Abandonment was
forest ecosystems, adjacent to forest openings, usu- also associated with landscape homogenization
ally with ponds and flowing streams, which pro- and densification adjacent to extended patches of
vide an adequate and preferable foraging habitat dense scrublands and forests (Roura-Pascual et
(Poirazidis et al. 2007). In Slovakia, the Balkans al. 2005). Within that context, the 35-year study in
and the Carpathians Lesser Spotted Eagle breeds in Dadia NP also hypothesized that land abandonment
completely dry mountain woods at middle altitude created a non-optimal habitat for the Lesser Spotted
(300-800 m, over 500 m on average) (Meyburg Eagle.
1971, Demerdzhiev et al. 2019). In contrast, in The present work examines the species’ spatial
Dadia NP, it mainly breeds in forests lower than use in the light of continuous landscape changes in
300 m of altitude, characterized by mosaic habitats Dadia NP, which affect many raptor species in the
dominated by forest edges, small portions of ma- region (Poirazidis 2017). We analyzed the long-
ture forests and local streams, while it avoids pure term adaptation of the breeding territories of Lesser
broadleaved forests (Poirazidis et al.2007). Spotted Eagle to the increasing homogenization of
Nest adjacency to local streams reflects its forest landscape in Dadia NP for the period 1979–
preference for this particular foraging habitat, as 2012, and our work had three main aims: a) assess
indicated by the large proportion of Grass Snakes long–term population variability of the species and

8
 Lesser Spotted Eagle Clanga pomarina Brehm, 1831 (Aves: Accipitridae) in Dadia-Lefkimi-Soufli...

explore population trends, b) assess spatial changes thematic foci were derived from each Landsat satel-
of suitable habitat based mainly on vegetation char- lite image: a) thematic variable of vegetation den-
acteristics and forest density, c) describe the eco- sity in three classes: dense forest (80–100%), loose
logical tolerance of the species during this period in forest (40–80%), open forest (0–40%), based on a
respect to landscape changes through environmen- hybrid method using vegetation index (Normalized
tal niche models (ENMs hereafter). Difference Bare Soil Index) and unsupervised clas-
sification (ISO-Data) to determine forest densifica-
tion. Normalized Difference Bare Soil Index was
Materials and Methods calculated as: NDBSI= ((S1 – N))/((S1 + N)), where
In respect to the population monitoring of the spe- S1 is the shortwave infrared band, with a wave-
cies realized between 1979 and 2000, the population length between 1.55 and 1.75 μm, and N is near
estimations of the Lesser Spotted Eagle were based infrared band with wave length between 0.76 and
on the works of Hallman (1979), Vlachos (1989), 0.90 μm. For each thematic class, the percentage of
Adamakopoulos et al. (1995) and after 2000, the cover was calculated in four different spatial scales
Systematic Raptor Monitoring scheme (SRM here- (500, 1000, 1500, 2000 m) using a circular mov-
after), which was established in Dadia NP by WWF ing window, b) four vegetation indices were used:
Greece (Poirazidis et al. 2001) and set the basis NDVI, MSAVI, EVI, NDMI, all calculated in the
for continuous and detailed monitoring of all rap- four spatial scales above, and c) a digital elevation
tors in the region, which continues until today (see model (https://asterweb.jpl.nasa.gov/gdem.asp) was
Poirazidis 2017 for a detailed analysis of moni- also applied including topographical characteristics
toring results). To explore the population trends of like terrain roughness, inclination and aspect affect-
Lesser Spotted Eagle, we implemented the non-par- ing vegetation characteristics and fundamental nest-
ametric and distribution-free test of Mann-Kendall ing and foraging factors. All variables were calcu-
(MK). The test was applied to the number of annual lated in a 30 m spatial analysis.
territories to statistically determine whether there Preliminary modelling was performed for each
was a monotonic upward or downward (increases/ multiscale variable, in order to choose the optimal
decreases) trend over time (Baldwin et al. 2012). scale based on maximum univariate gain among the
MK tau and p values were calculated, using Kendall four scales per variable. The initial full model, using
package (McLeod 2015) in R programming lan- all variables (at the optimal scale for multiscale var-
guage (R Core Team 2017). iables), was pruned by selecting a subset of uncor-
We used the MaxEnt software (Phillips et al. related variables based both on Variance Inflation
2006) to create models of a potential suitable habitat Factor(VIF) criterion (VIF < 10) and pairwise cor-
based mainly on vegetation characteristics. MaxEnt relation (r < 0.8) using “usdm” package in R envi-
is a general-purpose machine learning technique es- ronment (Naimi 2015). Response curves of univari-
timating the target species probability distribution ate models were created for each variable partici-
(statistical model) of maximum entropy (closest to pating in the prune model, in order to examine how
uniform) based on constant restrictions (constraints) each one of them affects the species habitat selec-
that are placed by the processed data and by using tion. The suitability map with a scale that was firstly
presence-only data as a function of explanatory vari- continuous between 0 and 1, was transformed into
ables (Phillips et al. 2006). In the present study, the a binary file (presence – absence) according to the
entire landscape was represented by 10.000 random threshold “Minimum training presence”. According
points with a random seed that were used as pseudo- to the threshold, all training locations (centre of ter-
absences points. The MaxEnt parameterization (e.g. ritories) would be correctly classified, as our goal
convergence threshold, regularization parameter λ) was to predict the suitable area based on all esti-
was set at default values (Phillips & Dudik 2008, mated territories, indicating the least-suitable envi-
Elith et al. 2011). ronmental conditions (Radosavljevic & Anderson
The geometric centre of each estimated terri- 2014).
tory for the Lesser Spotted Eagle was used as in-
dependent presence for the years 1979 and 2012 to
explore long-term spatial changes of the suitable
Results
habitat in Dadia NP. The explanatory variables were For34 years, since the late 1970’s, when Lesser
based on Landsat images of the closest available Spotted Eagle monitoring was actually initiated in
date to the population survey (satellite images of Dadia NP, the species’ numbers appear to be gener-
August for the years 1985 and 2011). The following ally stable. Since the first estimation, eagle breeding

9
Poirazidis K., V. Bontzorlos, S. Schindler & D. Vasilakis

numbers were successively estimated by different from 66% in 1979 to 54% in recent years, whereas
researchers: Hallman (1979) recorded 19 territo- into Zone A, which is the strictly protected core of
ries, Vlachos (1989) recorded a range of 16–20 the Natura 2000 area in Dadia NP, the most radi-
territories, Adamakopoulos et al. (1995) estimated cal change was demonstrated (reduction from 80%
a range of 14–17 territories and, since 2000 until re- in 1979 to 53% in 2011) (Fig. 2). ENMs also indi-
cent years, the long-term SRM has been producing cate that the current breeding habitat of the eagle in
reliable indices about the species breeding popula- Dadia NP has lower elevation than in 1979, whereas
tion at similar levels for the years 2001-2005 and less adjacent dense forest appears in the species’
2012, ranged from 17 to 22 territories (Poirazidis surroundings in 2012, even within large radius of 2
2017). The change in the territories’ numbers may km (Fig. 3). In that context, it is also noted that the
demonstrate certain irregular fluctuations (Fig. 1); species had a clearly narrower ecological niche in
nonetheless, according to Mann Kendal test, which 1979 with a diverse mosaic of suitable habitat ac-
includes a total of eight years of estimation (1979, cording to MSAVI, whereas in 2012, within even
1989, 2001, 2002, 2003, 2004, 2005, 2012) it over- larger radius, much more open areas appeared in
all indicates a stable population (tau = 0.046, 2-sid- its territories, while the species no longer occupied
ed p value = 0.92786). dense forest (Fig. 3).
Apart from the arithmetic estimations for the
Lesser Spotted Eagle in Dadia NP during the last
25 years, which indicate population stability, there
Discussion
is a notable spatial change both in horizontal and Major changes have occurred during the last decades
vertical landscape use. It has been found that while in the landscape of Dadia NP (Triantakostantis et
only50% of the pairs bred in elevations below 100 al. 2006, Xofis & Poirazidis 2018), including in-
m in the 1970’s, this number has progressively in- tensification of agriculture, increased forest exploi-
creased since 2001. In addition, the horizontal spa- tation, abandonment of traditional agricultural prac-
tial use by the Lesser Spotted Eagle with regard tices and reduction of forest openings. It could be
to its breeding habitats was uniformly distributed argued that these habitat changes affecting the for-
in Dadia NP in the 1970’s. Since 2000, though the aging areas of Lesser Spotted Eagle would have af-
population has shifted to breed predominantly in the fected the species’ population as well. Nonetheless,
lowlands and at the eastern part of Dadia NP (Fig. the breeding population appears stable during the
2). last 25 years. On the other hand, an apparent change
The habitat parameters which created a suitable has been noted on the spatial distribution of its terri-
landscape for the Lesser Spotted Eagles breeding tories. An abandonment of “in-forest” breeding ter-
in the western part of Dadia NP decreased overall ritories has been recorded since the first monitoring

Fig. 1. Breeding densities of Lesser Spotted Eagle in Dadia NP since 1970’s expressed in numbers of occupied territories.

10
 Lesser Spotted Eagle Clanga pomarina Brehm, 1831 (Aves: Accipitridae) in Dadia-Lefkimi-Soufli...

2004, Treinys & Mozgeris 2010, Mirski 2009, Zub

et al. 2010), it must be stressed that after 2000 the
species was slowly “pushed” to completely translo-
cate all its breeding attempts exclusively in the east-
ern lowlands of Dadia NP, denoting prior existence
of available habitats within the core of Dadia NP.
Habitat heterogeneity has been recognized as
one of the main landscape characteristics determin-
ing biodiversity patterns (Tuanmu & Jetz 2015),
and the richness in forest openings and diversity in
forest composition throughout the entire extent of
Dadia NP, is the major characteristic which creates a
unique landscape formation that supports high lev-
els of biodiversity (Poirazidis et al. 2010b).
Already in 2001, many areas in the south-
western parts of Dadia NP showed low levels of
forest diversity (Schindler et al. 2008) but in the
next decade general transition from open to forest
land, in most of Dadia NP parts, has led to increased
forest ecosystem homogeneity (Xofis & Poirazidis
2018). That is most possibly the main driver be-
hind the allocation of Lesser Spotted Eagle’s breed-
ing territories exclusively in the eastern lowlands.
Although a thorough analysis of environmental
parameters to assess other possible drivers behind
Fig. 2. Binary habitat-suitability map demonstrating this differentiated spatial use has not been realized,
Lesser Spotted Eagle spatial changes between 1970’s and surrogate factors corroborate the hypothesis. For
2000’s. instance, a significant increase has been noted in
forest dwelling raptor species of Dadia NP such as
results of 1979 (Hallmann 1979), where half of the Northern Goshawk Accipiter gentilis Linnaeus,
the eagle population bred within “interior” forest 1758, probably due to environmental factors such
sites. In 1979, the Lesser Spotted Eagle population as forest homogenization (Poirazidis et al. 2007,
was recorded to breed in rich ecotones, which were Poirazidis 2017).
formed of numerous forest openings that could sup- In recent years, a notable decline of the eagle’s
port a forest-edge raptor dweller. Nowadays, all population due to the loss of suitable habitats has
breeding pairs have established their territories on been observed in other countries such as Lithuania
the periphery of the Dadia NP at altitudes exclusive- (Treinys et al. 2007) and Germany (Meyburg et
ly lower than 100 m, where the breeding habitats al. 2004). In respect to Dadia NP, the reduction
are now dominated by more humanized landscapes of the suitable habitat has forced the species to
with less availability of forest openings, but an ad- translocate its breeding in the eastern non-optimal
jacency to agricultural areas. landscapes but still no decline of the population
That spatial change may also function as in- has been observed. The decline in open and semi-
creasing pressure on the abundance of the Lesser open habitats in the interior of the forest since the
Spotted Eagle, in a future of further reduction of 1950s (Triantakonstantis et al. 2006) has caused
suitable habitats, which may push the population to a heterogeneity loss that may have also caused a
limit down (Väli et al. 2004). According to Lohmus decline in the abundance of reptiles and amphib-
& Väli (1999), semi-natural forest openings offer ians, which represent an important food source for
better feeding conditions than agricultural arable the Lesser Spotted Eagle in Dadia NP (Vlachos
lands, and function as optimal habitats under stress & Papageorgiou 1996) and strongly depend on
for the species. On the other hand, if the stress fac- landscape heterogeneity at scales ranging from 50
tors cease but optimal habitats decrease, the birds to 500 ha (Schindler et al. 2013). In comparison,
will gradually move to alternative habitats. Even though with the Short-toed Eagle Circaetus gallicus
though Lesser Spotted Eagle is a species with no- Gmelin, 1788, a species which also relies on for-
table preference for forest edges to nest (Treinys est openings to capture its prey, the latter appears

11
Poirazidis K., V. Bontzorlos, S. Schindler & D. Vasilakis

Fig. 3. Environmental niche model results with regard to Lesser Spotted Eagle spatial preferences in respect to eleva-
tion, MSAVI index, forest density and forest openings in 1979 and 2012 in Dadia NP.

12
 Lesser Spotted Eagle Clanga pomarina Brehm, 1831 (Aves: Accipitridae) in Dadia-Lefkimi-Soufli...

to be increasing in the region (Poirazidis 2017). ronment, Physical Planning and Public Works, Ministry of
Nonetheless, it is possible that the tolerance thresh- Agriculture & WWF – Greece, Athens. (In Greek)
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