Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus)
Исследования видового состава и плотности населения птиц проводились в 1999-2007 гг. Для количественного учета птиц использовался маршрутный метод с некоторыми элементами метода картирования гнездовых территорий на учетных площадках. Три типа пойменных лесов Беларуси были охвачены учетами: дубравы,...
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| Published in: | Бранта: Cборник научных трудов Азово-Черноморской орнитологической станции |
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| Date: | 2009 |
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Інститут зоології ім. І.І. Шмальгаузена НАН України
2009
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| Cite this: | Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) / V. Sakhvon // Бранта: Сборник научных трудов Азово-Черноморской орнитологической станции. — 2009. — Вип. 12. — С. 27-39. — Бібліогр.: 30 назв. — англ. |
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Digital Library of Periodicals of National Academy of Sciences of Ukraine| _version_ | 1860240386272788480 |
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| citation_txt | Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) / V. Sakhvon // Бранта: Сборник научных трудов Азово-Черноморской орнитологической станции. — 2009. — Вип. 12. — С. 27-39. — Бібліогр.: 30 назв. — англ. |
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| container_title | Бранта: Cборник научных трудов Азово-Черноморской орнитологической станции |
| description | Исследования видового состава и плотности населения птиц проводились в 1999-2007 гг. Для количественного учета птиц использовался маршрутный метод с некоторыми элементами метода картирования гнездовых территорий на учетных площадках. Три типа пойменных лесов Беларуси были охвачены учетами: дубравы, «южные» черноольховые леса и черноольшаники с примесью в древостое и подросте ели («северные»). В общем 48 видов воробьинообразных птиц было зарегистрировано на гнездовании в пойменных лесах, для 37 из них получены значения плотности на учетных площадках.
Дослідження видового складу та щільності населення птахів проводилися в 1999-2007 рр. Для кількісного обліку птахів використовувався маршрутний метод з деякими елементами методу картування гніздових територій на облікових ділянках. Три типи заплавних лісів Білорусі були охоплені обліками: діброви, «південні» чорновільхові ліси і чорновільшаники з домішкою у деревостої і підрості ялини («північні»). Загалом 48 видів горобціподібних птахів було зареєстровано на гніздуванні в заплавних лісах, для 37 з них отримані значення щільності на облікових ділянках.
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Бранта: Сборник научных трудов Азово-Черноморской орнитологической станции
Вып. 12. 2009. - Фаунистика. 27
УДК 598.2: 591.55 (253)
COMPOSITION AND DIVERSITY OF PASSERINE BIRD ASSEMBLAGES IN
THE FLOODPLAIN DECIDUOUS FORESTS DURING THE BREEDING SEASON
(BELARUS)
V. Sakhvon
Museum of Zoology, Belarusian State University
Key words: breeding passerine bird assemblage, fl oodplain black alder and oak forests, species
richness, overall density.
Видовой состав и разнообразие сообществ воро-
бьинобразных птиц во время гнездового сезона в
пойменных лиственных лесах Беларуси. - В. Сах-
вон. Музей зоологии, Беларусский государственный
университет
Исследования видового состава и плотности населе-
ния птиц проводились в 1999-2007 гг. Для количествен-
ного учета птиц использовался маршрутный метод с
некоторыми элементами метода картирования гнез-
довых территорий на учетных площадках. Три типа
пойменных лесов Беларуси были охвачены учетами: дубравы, «южные» черноольховые
леса и черноольшаники с примесью в древостое и подросте ели («северные»). В общем
48 видов воробьинообразных птиц было зарегистрировано на гнездовании в пойменных
лесах, для 37 из них получены значения плотности на учетных площадках. MDS-анализ
показал, что структура сообществ птиц была детерминирована структурным разноо-
бразием лесов. Общая плотность гнездования и в целом видовое богатство возрастали
от «северных» черноольховых лесов к черноольшаникам Полесья. Структура сообществ
птиц «южных» черноольшаников оказалась более схожа с сообществами птиц дубрав,
чем черноольховых лесов с примесью ели («северных»). По результатам BEST анализа
Phylloscopus trochilus, Troglodytes troglodytes, Turdus philomelos, Aegithalos caudatus и Stur-
nus vulgaris определили распределение сообществ птиц по группам на основании степе-
ни их сходства. Общая плотность гнездования птиц в пойменных лесах была высокой
главным образом из-за значительной плотности широкораспространенных видов, харак-
теризующихся высокой степенью пластичности в выборе мест для гнездования. Всего
18 видов (48%) гнездилось, хотя бы раз за все годы исследований, с плотностью равной
либо выше 0.5 пар/га. Группа доминантов представлена 17 видами и в среднем 11-13 ви-
дов доминировали на отдельной площадке. Участие доминантных видов в сообществе
птиц на отдельной площадке было высоким и составляло от 57.6% до 75.7%. Доля видов
птиц, гнездящихся открыто на деревьях либо в подлеске составляла 31.2-51.4%, видов-
дуплогнездников - 25.4-40.9% и наземногнездящихся видов - 13.6-31.4%.
Ключевые слова: сообщества гнездящихся воробьинообразных, пойменные черноольхо-
вые леса и дубравы, видовое богатство, общая плотность.
Sakhvon V.
Composition and diversity of passerine bird assemblages...28
Видовий склад та різноманіття угруповань горобцеподібних птахів під
час гніздового сезону в заплавних листяних лісах Білорусі. - В.Сахвон.
Музей зоології, Білоруський державний університет.
Дослідження видового складу та щільності населення птахів проводилися в
1999-2007 рр. Для кількісного обліку птахів використовувався маршрутний
метод з деякими елементами методу картування гніздових територій на
облікових ділянках. Три типи заплавних лісів Білорусі були охоплені обліками:
діброви, «південні» чорновільхові ліси і чорновільшаники з домішкою у
деревостої і підрості ялини («північні»). Загалом 48 видів горобціподібних
птахів було зареєстровано на гніздуванні в заплавних лісах, для 37 з них
отримані значення щільності на облікових ділянках. MDS-аналіз пока-
зав, що структура угруповань птахів була детермінована структурним
різноманіттям лісів. Загальна щільність гніздування і в цілому видове ба-
гатство зростали від «північних» чорновільхових лісів до чорновільшаників
Полісся. Структура угруповань птахів «південних» чорновільшаників вия-
вилася більш схожа з угрупованнями птахів дібров, ніж чорновільхових лісів
з домішкою ялини («північних»). За результатами аналізу BEST Phylloscopus
trochilus, Troglodytes troglodytes, Turdus philomelos, Aegithalos caudatus і
Sturnus vulgaris визначено розподіл угруповань птахів по групах на основі
ступеня їх схожості. Загальна щільність гніздування птахів у заплавних
лісах була високою головним чином через значну щільність широкорозпов-
сюджених видів, що характеризуються високим ступенем пластичності
у виборі місць для гніздування. Загалом 18 видів (48%) гніздилось хоча б
раз за всі роки досліджень, з щільністю рівною або вище 0.5 пар/га. Група
домінантів представлена 17 видами, і в середньому 11-13 видів домінували на
окремій ділянці. Участь домінантних видів в угрупованні птахів на окремій
ділянці була високою і становила від 57.6% до 75.7%. Частка видів птахів,
що гніздяться відкрито на деревах або у підліску становила 31.2-51.4%,
видів, що гніздяться в дуплах - 25.4-40.9%, і видів, що гніздяться наземно -
13.6-31.4%.
Ключові слова: угруповання горобцеподібних птахів, що гніздяться,
заплавні чорновільхові ліси й діброви, видове багатство, загальна щільність.
Introduction
There are many works concerning study of bird assemblages of deciduous forests in the
European temperate zone (Nilsson, Liberg 1984; Waliczky 1991, 1992; Hansson 2001; Kralj,
Radović 2005; Leito et al. 2006). But the majority of researched forests situates in uplands, e.g.
they are dry and transformed by human activity. Some of long-term researches are carried out in
never-managed, «climax» old-growth forests (including swampy stands) of Białowieża National
Park (Poland) (Tomiałojć, Wesołowski 1996, 2004; Wesołowski, Tomiałojć 1997; Tomiałojć et
al. 1984; Wesołowski et al. 2002, 2006).
The fl oodplain forests in Belarus consist of oak, alder and very rare ash stands. They
occupy fl oodplains of rivers and are fl ooded during the spring high water. These forests are
mostly distributed in the territory of Polesye lowland along the Pripyat River (Yurkevich et al.,
1977). In the recent past the fl oodplain forests were intensively cut down, that led to signifi cant
reduction of the areas and age of these types of forests. Now most of them are taken under
protection.
Бранта: Сборник научных трудов Азово-Черноморской орнитологической станции
Вып. 12. 2009. - Фаунистика. 29
The fl oodplain forests bear the minimum of anthropogenic infl uence due to their diffi cult
position to traverse and the spring high water level, which lasts a signifi cant part of year (late March
to mid May and sometimes even later). That is why they can be considered as stable ecosystems
important for conservation of bird diversity in Belarus, including some rare species.
The structure of bird communities is determined by processes operating at local scales,
such as the fl oristic structure of habitats (spatial confi guration, productivity and diversity of
vegetation) (Wiens 1989). The character of spring high water (duration, altitude, frequency) is an
important factor infl uencing bird assemblages in the fl oodplain forests, especially ground nesting
species besides the effect of vegetation structure.
The aim of this research was to determine the composition and structure of the passerine
bird assemblages of the fl oodplain deciduous forests during the breeding season and explore the
main habitat characteristics that affect the given parameters.
Study area
The Republic of Belarus is situated in the geographical center of Europe. In general the
relief is represented with lowlands; the highest point is 346 meters above sea level. The climate
is moderately continental with frequent cyclones. Physical and climatic conditions provide the
domination of forest (mainly coniferous) and wetland ecosystems in the territory. The average
annual rainfall is 600-700 mm, the average temperature is 7.4 ºС in the southwest and 4.4 ºС in the
northeast. The average temperature in January is between - 8 and - 4.5 ºС and in July is between
17-18.8 ºС. Snow cover lies for 75-125 days. In the territory of Belarus two geobotanical regions
meet: European broad-leaved and Eurasian coniferous forests (Podolyako 2000; Loginov 2002).
The total area of forests in Belarus is 35.5% of the whole territory. They are represented by
boreal coniferous – 50.4 % of the forests territory, sub-boreal broad-leaved coniferous – 13.1
%, oak-leaved and broad-leaved – 3.8 % with their participation continuing to descend, small-
leaved forests – the rest. There are main types of fl oodplain forests – oak Quercetum fl uvialis
(only 7% from all the quantity of oak forests) and alder Alnetum fl uvialis forests (about 11 %
from all forests) (Yurkevich et al. 1977).
Description of the census areas
Our researches in fl oodplain forests were based on dominant plant species of a forest
stand, a young-tree layer and a character of a bush layer. Three types of the fl oodplain forests
were covered: the fl oodplain oak Quercus robur forests, the «southern» and the «northern»
fl oodplain black alder Alnus glutinosa forests. «Northern» black alder forests are characterized
by presence in the young-tree layer and in the forest stand of Norway spruce Picea abies in
comparison with the «southern» stands. We tried to choose the most typical, homogeneous and
as big as possible internal sites of the forest, avoiding the infl uence of various factors (glades,
open areas (meadows, bogs) etc.). Only the main features of each plot are presented. The age of
these forests is approximate because of contradictions in the botanical literature concerning the
defi nition of the age of fl oodplain tree stands. All studied forests were old. In total 8 census areas
have been placed (fi g. 1). These areas are Important Bird Areas and are taken under protection
in Belarus.
«Southern» fl oodplain black alder forests:
1. Plot AZs - 52°46’ N 26°00’ E. The transect (census plot) length is 1500 m. It is located
in the fl oodplain of the Shchara River. It is characterized by the signifi cant water level (up to 0.6m
during some seasons). The upper canopy is composed of black alders (40-50 cm tree diameter
at breast-height, aged 70-90). The well-developed bush layer is composed of Padus racemosa,
Sakhvon V.
Composition and diversity of passerine bird assemblages...30
Frangula alnus, and some species of willow Salix
sp., Rubus idaeus, R. caesius and Ribes sp. A dry
oak belt with hornbeam Carpinus betulus in the
young-tree layer presents a plot that is typical for
the fl oodplain alder forests. The herb layer (height
is more than 1 m at the end of spring): Carex sp.,
ferns, nettle Urtica dioica and other water-resistant
plants - is well-developed. There are many fallen
trunks (logs) and uprooted trees. The hop Humulus
lupulus densely braids trees that are preferred for
nesting by many birds.
2. Plot ATs – 52°12’ N 27°30’ E. The transect
length is 1000 m. It is also located in the fl oodplain
of the Shchara River and within 10 km from the
previous one. In comparison with the previous plot
this one consists of a younger tree stand (30-40
cm in diameter, 60-80 years old). The water level
usually does not exceed a depth of 30 cm and water
can dries up in summer. The bush layer is less
developed. There are many uprooted trees.
3. Plot APs - 52°09’ N 27°40’ E. The transect
length is 500 m. The forest we are interested in lies
in the fl oodplain of the Pripyat River. It is 70-90
years old. In early spring the water level can exceed 70 cm in depth and the plot remains swampy
at some places throughout a year. The upper canopy is composed of black alder with a small
number of birches Betula sp. and some large oaks. The young trees practically are absent, as well
as uprooted trees. The bush layer is represented as separate willow bushes and Frangula alnus.
«Northern» fl oodplain black alder forests:
1. Plot ASn - 54°12’ N 28°06’ E. This plot is situated in the fl oodplain of the Gaina River.
The transect length is 1000 m. Old alder stands (25-40 cm in diameter, 80 years and more) with
some inclusions of Picea abies dominate and form the upper canopy. In the lower canopy spruce
dominates. The young tree layer is also represented almost exclusively by spruces (height 2-7 m).
The forest is about 300 m in width and spreads along the Gaina River on the one side and borders
on a spruce forest on the other side. Birch becomes dominant with alder at the swampiest sites
of forest. Some places are presented with solitary aspens Populus tremula. The well-developed
bush layer is composed of willow and Frangula alnus. The high herb layer (1-1.3 m high) is
represented as nettle, ferns, Carex sp. and other marsh plants. This is the swampiest plot. The
water level can be 20-30 cm in summer and up to 50 cm in early spring. There are many uprooted
trees.
2. Plot ABn - 53°21’ N 28°51’ E. The study plot is located in the fl oodplain of the Berezina
River. The transect length is 700 m. An old black alder forest (30-45 cm diameter, 80 years and
more) occupies the lowest places of the fl oodplain and is surrounded by older dry oak-spruce
stands. Solitary aspens and oaks are presented in some sites. The well-developed young tree
layer is represented only by spruces (2-5 m height). The bush layer is composed of bird cherries
and willows. The water level is insignifi cant and does not exceed 30-40 cm in spring and dries
up in summer. The high herb layer is typical for swampy forests.
Floodplain oak forests:
This territory (52°12’ N 27°32’ E) of old oak forests (over 100 year-old) is situated in
the fl oodplain of the Slutch River and is included in the republic protected site «Middle Pripyat
1
3
2
4
Russia
Россия
Ukraine
Украина
Lithuania
Литва
Latvia
Латвия
Fig. 1. Distribution of study areas in Belarus
Notes: 1 – Plots APs, FF-1, FF-2, FF-3; 2 – AZs and
ATs; 3 – ABn; 4 – ASn (for explanations look through
the text)
Рис. 1. Распределение территорий исследова-
ния в Беларуси
Примечания: 1 – Площадки APs, FF-1, FF-2, FF-3; 2 –
AZs и ATs; 3 – ABn; 4 – ASn (объяснение см. в тексте).
Бранта: Сборник научных трудов Азово-Черноморской орнитологической станции
Вып. 12. 2009. - Фаунистика. 31
Floodplain». Three census plots have been chosen in 15 km2 territory. The fl oodplain oak forest
is not uniform. There are spots with the old pine Pinus silvestris; lowest places consist of alder
stands. Much more rarely hornbeams are found on uplands. It is quite typical for the fl oodplain
forests of Polesye Region. In all plots the average diameter of oak trees at breast-height is 40-55
cm, some trees reach more than 70 cm.
1. Plot FF-1 – The transect length is 500 m. The relief is plain and the plot is the swampiest
(the water level can exceed 40 cm in spring) in comparison with other oak plots. Water dries up
only in summer. More alders are found farther from the river, so the oak forest is replaced by
swampy alder stands. The oak stands with some inclusions of ash Fraxinus excelsior and aspens
dominate and form the upper canopy. The well-developed bush layer is represented mainly by
Corylus avellana (5-7 m height) and Frangula alnus and willows in damp places. The high
herb layer is mainly composed of Urtica dioica, Carex sp., Aegopobium podagraria etc. and is
formed on the gaps, but under Corylus avellana – is almost not present. This plot is characterized
by abundance of gaps in comparison with the other study oak stands. The well-pronounced
young tree layer is formed by maples Acer sp., birches etc.
2. Plot FF-2 – The length of transect is 600 m. The relief is undulate. The oak uplands
which are not fi lled up with water in spring alternate with lowlands fi lled up with water that never
dries up. These lowlands are frequently covered with willows. The oak stands dominate and with
some quantity of ashes and aspens in lowlands form the upper canopy. There are many dead
trees, especially oaks and aspens. The well-developed bush layer is formed by Corylus avellana,
Euonymus europaea and Rubus species. When trees are covered with foliage it becomes dark in
the forest. The herb layer is oppressed.
3. Plot FF-3 – The length of transect is 500 m. The main characteristics are similar to the
previous plot. The upper canopy layer is very open due to separate old oak stands and the herb
layer is well-developed (more than 1 m high in summer). The young tree layer and the bush
layer are well identifi ed and represented by maples, Malus silvestris, Pyrus comminis, Sorbus
aucuparia, Rubus species, Corylus avellana and Frangula alnus.
The water level in all oak plots can exceed more than 60 cm in spring and can be more
than 1.5 m in lowlands.
Methods
Bird census. The data were collected using the Finnish line transect method (Järvinen,
Väisänen 1976) with some elements (additional) of the territory mapping technique (Wesołowski
et al. 2002; Tomiałojć, Wesołowski 2004). For census of birds we chose a constant transect of
the certain length and counted all birds in a certain belt (afterwards this area was considered as
a census plot and was used for the calculation of the species density). We used only three main
belts to count birds (120, 150 and 200 m). We used 60 m in both sides for Pyrrhula pyrrhula,
Carpodacus erythrinus, Coccothraustes coccothraustes, Certhia familiaris, Aegithalos caudatus
and for all of Parus tits, except for Parus major. We used 100 m in both sides for Anthus trivialis,
Troglodytes troglodytes, Locustella fl uviatilis, Sturnus vulgaris, Sitta europaea, Oriolus oriolus,
Garrulus glandarius and for all species of Turdidae, except for Erithacus rubecula. 150 m (75m
to both sides) wide belt was used to count all other species. We used 200 m to both sides for
census of Corvus corax.
The counts were carried out during breeding seasons of 2000 and 2007. The counts (from
3 up to 10 during a breeding season) were made between the end of March and the middle
of June. Counting was conducted early in the morning during the period of maximum bird
activity. To increase the accuracy of results the counts were made with long stops. Some visits
were performed in the evening for the dusk active bird’s registration. In addition we carried
Sakhvon V.
Composition and diversity of passerine bird assemblages...32
out detailed inspection of the census area within the limits of transects to search for nests (or
mapping of territories) of some not numerous species, species with large breeding territories
and «problematic» for counts species (Sitta europaea, Locustella fl uviatilis and all species of
Turdidae, except for Erithacus rubecula) (Tomiałojć, Lontkowski 1989). At the end of the
breeding season the general map of the breeding territories for each species was made. It was
used for the calculation of the overall density.
Data analysis. Besides counts of birds on the census plots we revealed composition of
bird species in a breeding season using repeated inspection of studied territories of fl oodplain
forests. Some species of fl oodplain forests are not registered on the studied plots during the
counting. Parus cristatus, Turdus pilaris, Phoenicurus phoenicurus, Sylvia curruca, Acrocephalus
palustris and Passer montanus were found
breeding once. Motacilla alba, Ficedula
parva, Parus cyanus, Emberiza citrinella
and Lanius excubitor are distributed
sporadically and are rare everywhere in the
fl oodplain forests. These species were not
used for the statistical analysis.
The calculation of density without
any corrections was carried out as a number
of pairs/ha. A territorial male, inhabited
nest or birds with attributes of breeding
were accepted for a pair. The territories of
bigamists or bachelor males were treated
as equivalent to those of monogamists.
The number of species was considered as a
basic measure of species richness. Species
components over 5% of assemblage were
considered as dominants. For analyses of
bird assemblage we distinguished among
nesting birds the following groups: ground
nesters, openly nesting on trees and shrubs,
hole-nesters. Erithacus rubecula and
Certhia familiaris was considered as hole-
nesters.
To test whether bird assemblages
form natural assemblages in accordance
with the type of forest, we used MDS-
analysis of square-root transformed density
data (Bray-Curtis coeffi cient of similarity;
50 restarts; minimum stress 0.001). The statistical signifi cance of difference between the bird
assemblages was then tested with ANOSIM procedure. To fi nd a subset of species that best
“explain” the results of MDS-analysis, we used the BEST procedure. All tests were performed
using PRIMER 6.0 software (Clarke, Gorley 2006).
Results
Overall, 37 species were registered breeding within all the census plots that make 77.1%
from 48 passerine species breeding in the fl oodplain forests (Table 1-3). The number of species
in plots varied from 15 (plot FF-2) to 28 (plots AZs and ATs) in a single breeding season (Table
FF-3
FF-2
FF-1
APs
ATs
AZs
ASn
ABn
Fig. 2. Results of multidimensional scaling of the
surveyed bird assemblages in different types
of fl oodplain forests
Note: The bird assemblages are represented by abbreviations
of researched types of forests (see descriptions of census
plots).
Рис. 2. Результаты многомерного шкалирования иссле-
дованных сообществ птиц в разных типах
пойменных лесов.
Примечание: сообщества птиц обозначены буквенными индексами
исследованных типов лесов (см описание учетных площадок).
Бранта: Сборник научных трудов Азово-Черноморской орнитологической станции
Вып. 12. 2009. - Фаунистика. 33
Species
Вид
FF-1, length 500 m
длина 500 м
FF-2, length 600 m
длина 600 м
FF-3, length 500 m
длина 500 м
Pairs / Пары Mean / Среднее 2003 2002
2004 2005 p/ha
пар/га % pairs
пары
p/ha
пар/га % pairs
пары
p/ha
пар/га %
Fringilla coelebs 15 14 1.93 14.5 21 2.33 19.3 14 1.87 14.3
Sylvia atricapilla 10 6 1.07 8 8 0.89 7.4 - - -
Phylloscopus collybita 10 6 1.07 8 7 0.78 6.4 8 1.07 8.2
Phylloscopus sibilatrix 8 8 1.07 8 12 1.33 11 11 1.47 11.3
Parus major 6 7 0.87 6.5 8 0.89 7.4 5 0.67 5.1
Turdus philomelos 8 7 0.75 5.6 6 0.5 4.1 3 0.3 2.3
Phylloscopus trochilus 6 5 0.73 5.5 - - - - - -
Erithacus rubecula 4 6 0.67 5 13 1.44 12 10 1.33 10.2
Parus caeruleus 3 5 0.67 5 5 0.69 5.7 3 0.5 3.8
Turdus merula 7 6 0.65 4.9 7 0.58 4.8 7 0.7 5.4
Certhia familiaris 2 4 0.5 3.8 - - - - - -
Luscinia luscinia 4 6 0.5 3.8 - - - 4 0.4 3.1
Sylvia borin - - - - - - - 3 0.4 3.1
Muscicapa striata 1 4 0.33 2.5 - - - - - -
Ficedula albicollis 3 2 0.33 2.5 5 0.56 4.6 - - -
Ficedula hypoleuca 2 3 0.33 2.5 7 0.78 6.4 13 1.73 13.3
Troglodytes troglodytes 3 3 0.3 2.3 2 0.17 1.4 1 0.1 0.8
Sturnus vulgaris 3 3 0.3 2.3 4 0.33 2.8 7 0.7 5.4
Anthus trivialis - - - - - - - 3 0.3 2.3
Hippolais icterina 1 2 0.2 1.5 - - - - - -
Locustella fl uviatilis 2 2 0.2 1.5 - - - - - -
Sitta europaea 2 2 0.2 1.5 3 0.25 2.1 4 0.4 3.1
Luscinia svecica 2 2 0.2 1.5 - - - 4 0.4 3.1
Coccothraustes coccothraustes 1 1 0.17 1.3 4 0.56 4.6 3 0.5 3.8
Spinus spinus - 2 0.13 1 - - - - - -
Oriolus oriolus 1 1 0.1 0.8 - - - - - -
Garrulus glandarius 1 - 0.05 0.4 - - - 2 0.2 1.5
Total (27 species)
Всего (27 видов) 105 107 13.32 100 112 12.08 100 105 13.03 100
Notes: «-» – species was absent, but in calculation of mean was accepted; The bold – dominants (constituting
≥ 5 % of assemblage).
Примечания: «-» вид отсутствовал, но учитывался при расчетах среднего значения. Жирным шрифтом указаны
доминанты (составляющие ≥ 5 % сообщества).
Table 1. The breeding passerine bird assemblages of the fl oodplain oak forests
Таблица 1. Сообщества гнездящихся воробьинообразных птиц в пойменных дубравах.
1, 2). The highest number of species was found in the fl oodplain black alder forests of Polesye
region – 33 species (Table 2). According to results of cluster analysis at a level of similarity about
70 % it is possible to allocate three basic assemblages of birds of: 1) fl oodplain oak forests; 2)
«southern» fl oodplain black alder forests; and 3) fl oodplain alder forests with presence of spruce
at a forest stand and young-tree layer («northern»). The difference between bird assemblages
was statistically signifi cant (P=0.004, ANOSIM) (Fig. 2). The presence of bird assemblage of
the oak plot FF-1 in group «southern» black alder forests was explained by habitat structure
(impurity of alders in a tree stand and high swamp). The following 5 species: Phylloscopus
trochilus, Troglodytes troglodytes, Turdus philomelos, Aegithalos caudatus and Sturnus vulgaris
were found to best explain the grouping of the bird assemblages of different types of forests
studied (r = 0.926, BEST procedure).
Sakhvon V.
Composition and diversity of passerine bird assemblages...34
Table 2. The breeding passerine bird assemblages of the «southern» fl oodplain black alder forests
Таблица 2. Сообщества гнездящихся воробьинообразных птиц в южных пойменных черноольховых лесах.
Notes: «+» – species was present, but number of pairs is unknown, in calculation of mean density was not accepted;
«-» – species was absent, in calculation of mean was accepted; Dominants are indicated in bold.
Примечания: «+» вид присутствовал, но количество пар неизвестно, при расчетах среднего значения не учитывался.
«-» - вид отстутствовал, но учитывался при расчетах среднего значения. Жирным шрифтом указаны доминанты.
Table 3. The breeding passerine bird assemblages of the «northern» fl oodplain alder forests
Таблица 3. Сообщества гнездящихся воробьинообразных птиц в «северных» пойменных ольховых лесах.
Species
Вид
AZs, length 1500 m
длина 1500 м
ATs, length 1000 m
длина 1000 м
APs, length 500 m
длина 500 м
Pairs / пары Mean / среднее 2001 2006
2000 2001 p/ha
пар/га % Pairs
пар
p/ha
пар/га % Pairs
пар
p/ha
пар/га %
1 2 3 4 5 6 7 8 9 10 11
Fringilla coelebs 52 47 2.2 15.9 19 1.26 7.9 13 1.73 12.6
Phylloscopus collybita 35 53 1.95 14.1 20 1.33 8.4 7 0.93 6.8
Erithacus rubecula 26 27 1.17 8.5 20 1.33 8.4 4 0.53 3.9
Phylloscopus trochilus 23 28 1.13 8.2 21 1.4 8.8 6 0.8 5.8
Parus major 23 18 0.91 6.6 6 0.4 2.5 7 0.93 6.8
Sylvia atricapilla 17 21 0.84 6.1 18 1.2 7.6 8 1.06 7.7
Troglodytes troglodytes 17 24 0.68 4.9 11 0.55 3.5 5 0.5 3.6
Turdus merula 25 14 0.65 4.7 23 1.15 7.2 4 0.4 2.9
Phylloscopus sibilatrix 11 17 0.62 4.5 20 1.33 8.4 8 1.06 7.7
Turdus philomelos 11 16 0.45 3.3 14 0.7 4.4 7 0.7 5.1
Pyrrhula pyrrhula 8 6 0.38 2.8 2 0.16 1 - - -
Hippolais icterina + 8 0.35 2.5 14 0.93 5.9 - - -
Parus palustris 6 6 0.33 2.4 4 0.33 2.1 1 0.16 1.2
Aegithalos caudatus 7 4 0.3 2.2 4 0.33 2.1 1 0.16 1.2
Coccothraustes
coccothraustes
8 3 0.3 2.2 - - - - - -
Sylvia borin 5 6 0.24 1.7 6 0.4 2.5 3 0.4 2.9
Certhia familiaris 4 4 0.22 1.6 3 0.25 1.6 2 0.33 2.4
Carpodacus erythrinus 1 5 0.16 1.2 - - - - - -
Sitta europaea 8 2 0.16 1.2 2 0.1 0.6 1 0.1 0.7
Sturnus vulgaris + 5 0.16 1.2 7 0.35 2.2 8 0.8 5.8
Luscinia svecica 4 + 0.13 0.9 3 0.15 0.9 - - -
Parus caeruleus 2 2 0.11 0.8 2 0.16 1 1 0.16 1.2
Turdus iliacus 4 2 0.1 0.7 8 0.4 2.5 - - -
Lanius collurio - - - - 9 0.6 3.8 - - -
Ficedula hypoleuca 2 2 0.08 0.6 5 0.33 2.1 4 0.53 3.9
Luscinia luscinia 2 2 0.06 0.4 5 0.25 1.6 7 0.7 5.1
Garrulus glandarius 3 1 0.06 0.4 2 0.1 0.6 1 0.1 0.7
Prunella modularis - 2 0.04 0.3 2 0.13 0.8 3 0.4 2.9
Oriolus oriolus 1 1 0.03 0.2 1 0.05 0.3 1 0.1 0.7
Anthus trivialis - - - - 4 0.2 1.3 5 0.5 3.6
Chloris chloris - - - - - - - 1 0.13 0.9
Ficedula albicollis - - - - - - - 3 0.4 2.9
Locustella fl uviatilis - - - - - - - 1 0.1 0.7
Total (33 species)
Всего (33 вида) 305 326 13.81 100 255 15.87 100 112 13.71 100
Species
Вид
ASn, length 1000 m / длина 1000 м ABn, length 700 m / длина 700 м
Pairs / Пары Mean / Среднее 2007
2006 2007 p/ha
пар/га % Pairs
пары
p/ha
пар/га %
1 2 3 4 5 6 7 8
Fringilla coelebs 17 13 1 12.6 10 0.95 11.7
Phylloscopus trochilus 14 9 0.77 9.6 - - -
Sylvia atricapilla 9 14 0.77 9.6 6 0.57 7
Phylloscopus collybita 10 12 0.73 9.2 7 0.67 8.2
Бранта: Сборник научных трудов Азово-Черноморской орнитологической станции
Вып. 12. 2009. - Фаунистика. 35
Continuation of Table 3.
1 2 3 4 5 6 7 8
Erithacus rubecula 7 11 0.6 7.5 15 1.43 17.6
Troglodytes troglodytes 10 10 0.5 6.3 10 0.71 8.8
Phylloscopus sibilatrix 6 5 0.37 4.6 3 0.29 3.5
Ficedula hypoleuca 4 7 0.37 4.6 1 0.1 1.2
Parus major 9 2 0.37 4.6 6 0.57 7
Turdus merula 7 7 0.35 4.4 6 0.43 5.3
Muscicapa striata 5 5 0.33 4.2 1 0.1 1.2
Turdus philomelos 10 2 0.3 3.8 9 0.64 7.9
Certhia familiaris 4 3 0.29 3.7 - - -
Prunella modularis 3 5 0.27 3.4 4 0.38 4.7
Parus caeruleus 1 3 0.17 2.1 2 0.24 2.9
Sitta europaea 4 1 0.13 1.6 2 0.14 1.8
Locustella fl uviatilis 4 1 0.13 1.6 - - -
Sylvia borin - 3 0.1 1.3 - - -
Parus palustris 1 1 0.08 1 - - -
Parus montanus 2 - 0.08 1 - - -
Turdus iliacus 3 - 0.08 0.9 1 0.07 0.9
Hippolais icterina 1 1 0.07 0.8 2 0.19 2.3
Spinus spinus 2 - 0.07 0.8 - - -
Garrulus glandarius 1 - 0.03 0.3 1 0.07 0.9
Corvus corax 1 1 0.03 0.3 1 0.04 0.4
Ficedula albicollis - - - - 4 0.38 4.7
Luscinia luscinia - - - - 2 0.14 1.8
Total (27 species)
Всего (27 видов) 135 116 7.95 100 93 8.11 100
Notes: «-» – species was absent, but in calculation of mean was accepted; The bold – dominants.
Примечания: вид отсутствовал, но учитывался при расчетах среднего значения. Жирным шрифтом указаны доминанты.
Table 4. The comparative characteristics of main structural parameters of bird assemblages of the
fl oodplain forests
Таблица 4. Сравнительная характеристика главных структурных параметров сообществ птиц поймен-
ных лесов.
Parameters
Показатели
Oak forests
Дубравы
«Southern» black alder forests
«южные» черноольшаники
«Northern»
black alder forests
“северные”
черноольшаники
FF-1 FF-2 FF-3 AZs ATs APs ASn ABn
Total assemblage
Все сообщество
Number of species
Количество видов 24 15 18 27.5±0.5 28 26 22.5±1.5 20
Overall density (pairs/ha)
Общая плотность (пар/га) 13.32±0.22 12.08 13.03 13.61±0.48 15.87 13.71 7.95±0.49 8.11
Cumulative participation of
dominants (%)
Общая доля доминантов (%)
62.5±4.9 75.7 73.2 69.2±3.1 62.6 63.5 64.8±0.9 73.7
Densities of nesting groups
Плотность гнездящихся групп
Ground nesters
Наземногнездящиеся виды 3.77±0.23 2.11 3.63 3.85±0.58 4.66 4.09 1.99±0.21 1.1
Openly nesting on trees and
shrubs
Виды, открытогнездящиеся на
деревьях и кустарниках
5.68±0.08 5.03 4.07 6.66±0.01 7.96 5.68 3.88±0.17 4.15
Hole-nesters
Дуплогнездники 3.87±0.53 4.94 5.33 3.1±0.11 3.25 3.94 2.08±0.12 2.86
Note: Mean values ± standard deviations in individual plots.
Примечание: Среднее значение ± стандартное отклонение по отдельным площадкам.
Sakhvon V.
Composition and diversity of passerine bird assemblages...36
The total bird density varied between 7.46 (plot ASn in 2007) and 15.87 pairs/ha (plot ATs)
(Table 2, 3). The mean density for all years was the lowest in the plots of «northern» black alder
forests despite of similar number of species in assemblages in comparison with bird assemblages
of other forests. The mean density of birds in the oak plots was high (Table 1) and similar with
assemblages of alder forests of Polesye but the participation of species was lower than in black
alder stands.
The group of dominants was composed of 17 species. The number of dominants was
11-13 species in each type of forests in mean. The composition of the group of dominants was
similar between all the plots and the number of species varied from 6 to 9. Only two species
(Fringilla coelebs and Phylloscopus collybita) were dominants in all the plots. Parus caeruleus
and Certhia familiaris were dominants only in the fl oodplain oak forests at once, Hippolais
icterina and Luscinia luscinia - in «southern» alder forests and Muscicapa striata – in «northern»
tree stands. The fl oodplain alder forests are characterized by presence of Troglodytes troglodytes
in the structure of dominants. Participation of dominant species in breeding assemblages of a
single plot made from 57.6% (plot FF-1, 2005) to 75.7% (plot FF-2) (Table 4).
From 31.2% (plot FF-3) to 51.4% (plot AZs, 2000) of species of bird assemblages bred
openly on trees and shrubs in a single plot, 25.4% (plot FF-1 in 2004) - 40.9% (plot FF-2) were
hole-nesters (Table 4). The group of the species nesting on the ground was represented by 12
species (19.6% of all number) and their participation in the population amounted to 13.6% (plot
ABn) - 31.4% (plot AZs in 2001).
Discussion
The effect of woodland structure and diversity of vegetation on bird assemblages has been
widely documented (Solonen 1996; Jokimäki, Huhta 1996; Elmberg, Edenius 1999; Sallabanks
et al. 2000; Laiolo 2002; Brotons et al. 2003). The type of forest stand and bushes composition
plays the main role in the organization of bird assemblage. The division of assemblages of
birds into three groups according to types of forests confi rmed the aforesaid. Five species made
the greatest contribution to this grouping. Phylloscopus trochilus, Troglodytes troglodytes and
Aegithalos caudatus undoubtedly preferred the fl ooded black alder forests. It is possible the last
species at the forest-edge. Turdus philomelos preferred drier and lighter oak forests, although
bred in high densities in fl ooded alder forests as well. The nesting of Sturnus vulgaris depended
on availability of holes and it nested at the forest-edges. The fl oodplain oak forests better
correspond to these conditions. Sturnus vulgaris also nested in high density in «southern» black
alder forests due to inclusions of solitary old oak trees.
Generally the bird assemblages of the fl oodplain forests were characterized by a high
species diversity and density. The bird assemblages of the studied groups of fl oodplain forests
were highly similar in species composition, total bird density and density of particular species.
The overall density and partly species richness were changing along a habitat gradient from the
highest in the fl oodplain black alder forests of Polesye Region to the lowest in «northern» alder
stands with impurity of spruce. The structure of bird assemblages of «southern» black alder
forests was similar rather with assemblages of oak stands than the «northern» ones. In spite of
the fact that the species richness of birds in «northern» black alder forests was similar to data
of fl oodplain forests of Polesye, the densities of some species - Fringilla coelebs, Phylloscopus
collybita, Phylloscopus sibilatrix, Parus major, Sylvia atricapilla, Turdus merula, Hippolais
icterina and Sylvia borin in the assemblages were smaller and overall density was low. The
tendency of increase of density of birds assemblages with movement from the north to the south
for different types of woodland landscapes was discussed (Novikov 1960). Alnus glutinosa is
situated in the area of optimum distribution in Polesye Region (Yurkevitch et al. 1977) and these
Бранта: Сборник научных трудов Азово-Черноморской орнитологической станции
Вып. 12. 2009. - Фаунистика. 37
forests with the black alder were characterized by a high habitat variety preferable for birds. This
factor was the main cause of observed differences in the structure of assemblages. For example,
seven types of nest locations of Turdus merula were recognized in the «southern» alder forests
and fi ve of them occurred frequently. While only fi ve types of nest locations were occurred in
«northern» alder stands and only two location types were common. The places for nesting of
some other groups of birds were limited in «northern» alder forests with impurity of spruce
too. Thus, the ground nesters (Phylloscopus trochilus, Phylloscopus sibilatrix and Erithacus
rubecula too (because of the limited number of holes)) built nests on dry hummocks near tree
trunks in fl ooded forests. Spruces grew together with alders on hummocks in «northern» forests.
Such hummocks were covered with needles and become unsuitable for nesting.
Discs of fl at root systems of uprooted trees represent a special unit intermediate between
the ground and the tree-layers (Tomiałojć et al. 1984) and provide preferable nesting sites for
some species. The abundance of Turdus merula, Pyrrhula pyrrhula and especially Troglodytes
troglodytes was positively related to the number of uprooted trees. In black alder forests, where
the discs were common, a high density of these species was observed. Turdus philomelos and
Prunella modularis placed the nests there as well. The nests of two species settled down in the
discs very often (usually Turdus merula was one of neighbors).
The character of the spring high water infl uenced the group of ground nesting birds.
The most changeable group of these species was Anthus trivialis, Phylloscopus sibilatrix and
Phylloscopus trochilus, and their presence and nesting terms directly depended on duration of
high water and the character of relief. Different from the previous species Phylloscopus collybita
can make nests above ground (up to 40 cm) in swampy ash-alder stands (Piotrowska, Wesołowski
1989). The density of ground nesters amounted to not more than 2 pairs/ha in the swampiest
«northern» plots and it was 2-2.7 times lower than in other types of forests. On the plot ABn the
relief was uniform, dry islands were almost absent and the number of ground nesting species
and their density was the lowest (3 species, 1.1 pairs/ha). Due to the presence of considerable
number of dry islands the character of spring high water infl uenced ground nesters in fl oodplain
oak forests areas than in black alder forests.
As for species composition of birds our data were similar to data of the deciduous tree stands
(ash-alder, alder-swamp, oak-lime-hornbeam forests) of Białowieża National Park (Tomiałojć et
al. 1984; Wesołowski et al. 2002; Tomiałojć, Wesołowski 2004; Wesołowski et al. 2006), but the
overall densities of breeding birds were a little higher, mainly due to high numbers of several
widespread species characterized by wide ranges in a choice of nest sites. So, 8 species bred with
the density exceeding 1 pairs/ha at least one time: Fringilla coelebs, Erithacus rubecula, Sylvia
atricapilla, Phylloscopus collybita, Phylloscopus trochilus, Ficedula hypoleuca, Phylloscopus
sibilatrix and Turdus merula. Only Fringilla coelebs and Erithacus rubecula bred with the same
density in «northern» alder forests. The densities of 24% of species in the «northern» black
alder forests and 80% - in the oak forests were equal or more than 0.5 pairs/ha. In fact the
number of these species was 18 (48%). The densities of Fringilla coelebs, Erithacus rubecula
and Phylloscopus collybita were more than 0.5 pair/ha during all the years and in all the plots.
High densities of these species can be explained by the fact that still there were some changes in
habitat structure, especially in oak forests. The dying of oak and aspen trees, formation of gaps,
rapid growth of young trees and bushes in the places contributed to maintenance or even increase
of number of some species, mainly forest-edge species. The infl uence of edges and gaps on
abundance of birds is widely considered in the papers (Tomiałojć et al. 1984; Báldi, Kisbenedek
1994; Fuller 2000; Sallabanks et al. 2000). The overall bird density reached the highest level
(14.9 pair/ha in ash-alder plot) in BNP in 2001. According to our data the overall bird density
was the highest in this year as well. It’s obviously those factors acting on a larger scale could
have been involved (Wesołowski et al. 2006).
Sakhvon V.
Composition and diversity of passerine bird assemblages...38
The short-term studies are not quite enough for estimation stability-variability
of densities of birds assemblages. We registered the considerable changes in numbers for
Phylloscopus collybita and Parus major in black alder stands. The additional long-term studies
outside the plots showed the fl uctuation of numbers for Ficedula albicollis and Lanius collurio
in these forests. The fl uctuation of numbers of these species happened without any changes in the
habitat structure advantageous for these species. Holmes et al. (1986) found that the species with
high densities were more variable in number than those occurring at low densities in a temperate
deciduous forest.
The bird assemblages of fl oodplain forests of Belarus were characterized by high
species diversity, richness and density. These parameters were made by woodland structural
complexity. Changes in hydrology due to fl ood control are one of the main factors of habitat
change in these forests and consequently, of changes in structure of bird assemblages.
Undoubtedly all fl oodplain forests must be taken under protection in Belarus and observations of
bird assemblages must be continued.
Acknowledgements
I thank Dr. Sergey Mastitsky for statistical advice.
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| id | nasplib_isofts_kiev_ua-123456789-65852 |
| institution | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| issn | 1994-1722 |
| language | English |
| last_indexed | 2025-12-07T18:29:30Z |
| publishDate | 2009 |
| publisher | Інститут зоології ім. І.І. Шмальгаузена НАН України |
| record_format | dspace |
| spelling | Sakhvon, V. 2014-07-04T05:24:41Z 2014-07-04T05:24:41Z 2009 Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) / V. Sakhvon // Бранта: Сборник научных трудов Азово-Черноморской орнитологической станции. — 2009. — Вип. 12. — С. 27-39. — Бібліогр.: 30 назв. — англ. 1994-1722 https://nasplib.isofts.kiev.ua/handle/123456789/65852 598.2: 591.55 (253) Исследования видового состава и плотности населения птиц проводились в 1999-2007 гг. Для количественного учета птиц использовался маршрутный метод с некоторыми элементами метода картирования гнездовых территорий на учетных площадках. Три типа пойменных лесов Беларуси были охвачены учетами: дубравы, «южные» черноольховые леса и черноольшаники с примесью в древостое и подросте ели («северные»). В общем 48 видов воробьинообразных птиц было зарегистрировано на гнездовании в пойменных лесах, для 37 из них получены значения плотности на учетных площадках. Дослідження видового складу та щільності населення птахів проводилися в 1999-2007 рр. Для кількісного обліку птахів використовувався маршрутний метод з деякими елементами методу картування гніздових територій на облікових ділянках. Три типи заплавних лісів Білорусі були охоплені обліками: діброви, «південні» чорновільхові ліси і чорновільшаники з домішкою у деревостої і підрості ялини («північні»). Загалом 48 видів горобціподібних птахів було зареєстровано на гніздуванні в заплавних лісах, для 37 з них отримані значення щільності на облікових ділянках. I thank Dr. Sergey Mastitsky for statistical advice. en Інститут зоології ім. І.І. Шмальгаузена НАН України Бранта: Cборник научных трудов Азово-Черноморской орнитологической станции Фаунистика Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) Видовой состав и разнообразие сообществ воробьинобразных птиц во время гнездового сезона в пойменных лиственных лесах Беларуси Видовий склад та різноманіття угруповань горобцеподібних птахів під час гніздового сезону в заплавних листяних лісах Білорусі Article published earlier |
| spellingShingle | Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) Sakhvon, V. Фаунистика |
| title | Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) |
| title_alt | Видовой состав и разнообразие сообществ воробьинобразных птиц во время гнездового сезона в пойменных лиственных лесах Беларуси Видовий склад та різноманіття угруповань горобцеподібних птахів під час гніздового сезону в заплавних листяних лісах Білорусі |
| title_full | Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) |
| title_fullStr | Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) |
| title_full_unstemmed | Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) |
| title_short | Composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (Belarus) |
| title_sort | composition and diversity of passerine bird assemblages in the fl oodplain deciduous forests during the breeding season (belarus) |
| topic | Фаунистика |
| topic_facet | Фаунистика |
| url | https://nasplib.isofts.kiev.ua/handle/123456789/65852 |
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