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Food composition and energy demand of the White Stork Ciconia ciconia breeding population. Literature survey and preliminary results from Poland


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This paper presents broad literature survey and preliminary results of study of food composition of breeding White Stork Ciconia ciconia population from Poland. The field- work was conducted in the farmland landscapes and river valleys at three areas near Strzelce Opolskie, Leszno and Ostrów Wiekopolski. During the standard nest visits in the nestling period we noted prey remains both vomited by chicks as a stress reaction or delivered at nest by adult birds. The above data on the White Stork's food composition in the discussed popu- lations are represented in tables 1-3. Energy value of the food was assessed only for the pop- ulation from southern Poland. Use of several method of data collection are discussed in the paper.
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Food composition and energy demand
of the White Stork Ciconia ciconia
breeding population. Literature survey
and preliminary results from Poland
Jakub Z. Kosicki1, Piotr Profus2, Paweł T. Dolata3& Marcin Tobółka1
1Departament of Behavioural Ecology, Adam Mickiewicz University, Umultowska 89,
61-614 Poznań, Poland, e-mail:
2Institute of Nature Conservation, Polish Academy of Sciences, Mickiewicza 33,
31-512 Kraków, Poland
3South Wielkopolska Group of Polish Society for the Protection of Birds, Wrocławska 60 A/7,
63-400 Ostrów Wielkopolski, Poland
ABSTRACT:This paper presents broad literature survey and preliminary results of study of
food composition of breeding White Stork Ciconia ciconia population from Poland. The field-
work was conducted in the farmland landscapes and river valleys at three areas near Strzelce
Opolskie, Leszno and Ostrów Wiekopolski. During the standard nest visits in the nestling
period we noted prey remains both vomited by chicks as a stress reaction or delivered at nest
by adult birds. The above data on the White Stork’s food composition in the discussed popu-
lations are represented in tables 1–3. Energy value of the food was assessed only for the pop-
ulation from southern Poland. Use of several method of data collection are discussed in the
KEY WORDS:food, foraging, energy consumption, food examination methods, White Stork,
Ciconia ciconia
The quality and quantity of food that parents provide to their chicks is the most im-
portant environmental factor influencing reproduction success of many bird spe-
cies (Martin 1987), including the White Stork Ciconia ciconia (Tryjanowski et al.
2005). Thus the availability of food resources in breeding areas and wintering
grounds is a key factor that regulates survival, number and condition of the White
Tryjanowski P., Sparks T.H. & Jerzak L. (eds.)
The White Stork in Poland: studies in biology, ecology and conservation
Bogucki Wydawnictwo Naukowe, Poznań 2006
Stork population (see Mrugasiewicz 1972, Dallinga & Schoenmakers 1984, 1987,
1989, Profus 1991, 2006a, Struwe & Thomsen 1991, Tryjanowski & Kuźniak 2002,
Schaub et al. 2005, Sæther et al. 2006).
Many hypotheses about the influence of food quantity and quality on White
Storks have not been verified in a credible way. This is because our knowledge
about the stork’s food composition and quality is still limited. In the majority of
papers there are no quantitative analyses of the contribution of particular species
and systematic groups in the food composition, and – what is even more vital – no
assessment of the energy significance of particular components of the White
Stork’s diet has been made (see Bauer & Glutz 1966, Creutz 1985, Làzaro 1986,
Pinowski et al. 1986, 1991, Lakeberg 1995, Struwe & Thomsen 1991, Antczak
et al. 2002).
Habitat changes observed in the past ten years in the agricultural environment
which are unfavourable for storks, for example, the increase of arable monocultures
(e.g. maize Zea mays and rape Brassica napus) at the cost of grassland, excessive mech-
anization and intensification of agriculture, may all account for the diminishing pop-
ulation of the White Stork. Apparently, as a result of such human activities, 16–24%
of stork breeding sites disappeared in the provinces of Silesia and Lubuskie (Jerzak
et al. 2006, Profus 2006b, c, Wuczyński 2006); whereas in other parts of Poland the
size of populations was observed to be stable or increase significantly (Guziak &
Jakubiec 2006). Therefore, research conducted on the composition and quality of
food can contribute to efficient protection of the White Stork species.
Results presented in this paper, both a literature survey and field data, aim at
filling in the gaps in our knowledge about the composition and energy values of the
stork’s food. This paper partially presents data and conclusions contained in
Profus’ (2006a) study, however, it is supplemented by new methods of gaining ma-
terials and field data collected in the region of Wielkopolska.
Methodological assumptions
Methods used in specifying the White Stork diet: a literature review
Stomach analysis and leather cap put on chicks’ beaks
The first attempts to describe diet composition were from stomach content analyses
of storks killed and of those found dead (Rörig 1903, Eckstein 1907, Steinbacher
1936, Stammer 1937, Putzig 1938, Radkiewicz 1984, Nachtigall et al. 1998).
Leather caps put on chicks’ beaks (Fig. 1) so they could not eat food brought to
the nest. The food brought by adult birds to the nest and spat out was collected for
analysis, and after specifying its animal content, returned for the chicks to eat
(Krapivnyj 1957, Körös 1992). For species protection and ethical reasons, both
methods are not used any more.
Direct observation of foraging and feeding birds
Data about the White Stork’s food composition mainly come from direct observa-
tions of foraging birds (Schüz 1940, Pinowski et al. 1986, 1991, Pinowska &
Pinowski 1989, Struwe & Thomsen 1991, Böhning-Gaese 1992, Ranner 1995,
170 Jakub Z. Kosicki, Piotr Profus, Paweł T. Dolata & Marcin Tobółka
Skov 1999), and observations made when chicks were fed (Berndt 1938). Using
this method of specifying food composition, it is necessary to realise its drawbacks.
Among the caught animals, it is easier for the observer to identify more sizeable
prey (moles, rodents, fish, tailless amphibians, reptiles, and leeches) (e.g.
Lakeberg 1995).
Permanent recording by a camera situated close to thenest is a new variation of
direct observation of feeding (Dolata 2006), which makes it possible to identify the
prey better than with the use of a telescope or binoculars, not only thanks to the
proximity to the nest but particularly the view from above into the centre of the
nest (i.e. where the parent spits out food for chicks), which cannot be seen when
observing from the ground.
Analysis of spat-out items
The most frequent method is spat-out item analysis (e.g. Putzig 1935, Drescher
1936, Szijj & Szijj 1955, Schierer 1962, Baudouin 1973, Sackl 1987, Làzaro 1982,
1986, Mužini & Rasajski 1992, Pinowska et al. 1991, Pinowski et al. 1991,
Antczak et al. 2002).
This method allows specifying the biomass of particular animals that were
eaten (e.g. Antczak et al. 2002). However, we need to consider its constraints be-
cause the stork greatly digests vertebrate cartilage and bones (Steinbacher 1936,
Szijj & Szijj 1955, Schulz 1998), which – for this reason – are recorded in the
spat-out bits to a much smaller degree than expected when analysing the stork’s
food. For instance, in Antczak’s et al. studies (2002) bones constituted only 8% of
the pellets, while a hair analysis showed that the remains of voles Microtus sp. only
amounted to 86.5% of the spat-out items. In this situation, identifying prey, i.e.
mammals, through the presence of its hair in the spat out items is a crucial method
(Dziurdzik 1973). Earthworms Lumbricidae poses yet another problem as their
identification must be performed more precisely than in the case of bones, and a
microscope must be used for analysis (Antczak et al. 2002).
Food composition and energy demand of the White Stork breeding population 171
Fig. 1. Schematic drawing leather cap put on chicks’ beak (after Krapivnyj 1957)
Energy value of food; specifying energy demand of individuals
and populations
Specifying caloric demand, being an element of the energy budget of an individual
or the population, is based on a theoretical model which evaluates the assimilated
energy quantity in the period between the arrival and the departure from the
breeding area, as well as the energy that is necessary for the young to grow and be-
come independent. This method, however, has serious constraints because obtain-
ing good results is only possible for birds raised in captivity (e.g. Kushlan 1977,
Keller & Visser 1999).
Specifying the energy value of food
The calorific content of animal food depends mainly on the fat content and not so
much on carbohydrates, proteins or minerals. The energy value of animal fat ex-
ceeds 39 kJ/g of fresh mass (Górecki 1967, Dolnik et al. 1982). The energy value of
other vital elements of the body (proteins and carbohydrates) is nearly half as
much (carbohydrates and animal proteins respectively: 17.58 and 23.66 kJ/g;
Dolnik et al. 1982). So a change in the animal’s fat content is responsible for a sub-
stantial change in the calorific value of the whole body.
There are three methods for calorific value measurements. The first one is the
value obtained directly from burning in the calorimeter, i.e. a top burning value.
also called the dry mass calorie value. The second is obtained from the calorific
value of the dry mass by means of subtracting the ash content, which is the
ash-free calorific value. The third is the biomass calorific value. This value – just
like the previous ones – is reached by taking into account the water content in the
initial sample (Górecki 1967).
Specifying food and energy demand of a breeding pair and a chick:
field methods
Double labelled water (DLW) is an isotope method of testing animal metabolism
(Nagy 1987, Nagy & Obst 1991, Nagy et al. 1999). This technique of establishing
field metabolic rate (FMR) also covers the cost of base metabolic rate (BMR),
thermoregulation and basic life functions. Food-energy needs of birds are deter-
mined by a number of independent factors, but the most important one is the mass
of the body (Nagy & Obst 1991). The energy demand of a breeding stork can be cal-
culated by a general equation:
FMR (kJ/d) = 10.9 M0.64
where M = live body mass of the animal in grams (Nagy 1987, Bozinovic & Medel
The body mass of breeding storks of both sexes comes from information (recal-
culated) contained in the literature (Sasvári & Hegyi 2001). Field metabolic rates
obtained for an average male and female are respectively: 2136 and 1976 kJ/day, so
for the pair: 4112 kJ. The birds’ daily energy requirement is in fact higher because
the bird is not able to use the entire energy derived from food. Assuming that the
bird derives 75% of the energy from food on average (Bezzel & Prinzinger 1990) –
172 Jakub Z. Kosicki, Piotr Profus, Paweł T. Dolata & Marcin Tobółka
specific data for fish and invertebrates respectively: 77.2% and 73.9% according to
Castro et al. (1989); for fish, amphibians and mammals respectively: 81%, 78%
and 74% (Dolnik et al. 1982) – gross energy disposal for a male amounts to 2847
kJ/day, and the female 2634 kJ/day. So energy requirement of a pair of storks is
5481 kJ/day.
Once we know the daily energy requirement of a pair of storks, it is possible to
calculate the daily food intake (DFI). The amount of food necessary to cover daily
energy requirement may be calculated by dividing DDE by the energy value of 1 g
of food biomass.
In the case of chicks, food and energy demand depends on the individual stage
of development as well as the body mass. Total metabolised energy (TME)
(kJ/chick) is the quantity of energy needed from birth till the bird is able to fly, and
it is proportional to the body mass of a mature individual of the given species
(Drent et al. 1992, Weathers 1996). To calculate total metabolised energy, an equa-
tion published by Weathers (1996) was used:
TME = 6.65 M0.85 ×tfl
A chick’s energy requirements (gross energy content of food consumed) is also
higher than the field metabolic rate (FMR), because not all the energy in food is
used. Thus it is also necessary to consider the degree to which energy is used from
food. In fact, this is not known for the stork’s chicks, but it is known that young
birds use energy more efficiently than adults, which is well proven, for example, for
owls (e.g. Ceska 1980). The daily maximum energy demand (max. DME) of a
White Stork’s chick can be calculated with the use of an equation by Weathers
DME = 11.7 M0.91 × tfl
where M is chick mass (in grams), and tfl defines the length of time the chicks re-
main in the nest (in days).
With 85% assimilated food, the value calculated per chick is 3450 kJ. Such en-
ergy is contained in 784 g of food of an average caloric value (4.4 kJ/g of live body
mass). Extreme values are: 3287 kJ for a 2934 gram chick, and 3750 kJ for a 3392
gram chick, which corresponds to a biomass of 747 and 852 g at a food density of
4.4 kJ/g of fresh mass.
The quantity of food that ensures breeding success for a pair of birds
In order to assess food quantity – that a pair of storks must find in the breeding
area to cover their food needs and to raise their offspring – the following conditions
must be fulfilled, according to Profus (2006a):
a) a pair remains in the breeding area for 144 days;
b) average energy value of the food amounts to 4.4 kJ/g of the biomass;
c) a pair of weaker condition (parents of smaller mass and those raising 1–2 off-
spring) consume 1141 g food/day, while strong pairs (parents of bigger mass
with 3–6 offspring) consume on average 1317 g of food a day;
Food composition and energy demand of the White Stork breeding population 173
d) one chick receives 34.3 kg of food from its parents from hatching to the first
flight, and after the first flight (for the next 10 days) the chick gets 3 kg of food,
while it finds 2.5 kg on its own.
From arrival till departure the food demand of parents was assessed at 179.4 kg,
while a pair without young must consume a total of 141.7 kg of food within 130
days. The potential food resources that storks may use have not yet been fully as-
sessed (see Profus 2006a).
Research on the White Stork’s food in Poland
The food composition of the White Stork in Poland has not been studied exten-
sively (see Profus 1985, Ptaszyk 1998).
The first studies on the Polish territory were conducted by German ornitholo-
gists in the area that belonged to Germany at the time. Steinbacher’s work (1936)
was performed on the basis of materials from the then Eastern Prussia (storks
from the present Kaliningrad Region, with only a few samples came from Warmia
and Masuria). Publications by Drescher (1936) and Stammer (1937) concerned
only a part of Silesia (in the past German, now Polish). Scarce information about
the species’ food can also be found in Czudek’s paper (1935) which relates to the
area of Pszczyna and Cieszyn (Silesia).
In later times only Mrugasiewicz (1972) mentions “numerous” observations of
the young spitting out food during a nest check in the valley of the Barycz river
(Milicz district). Between 1959 and 1968 he stated that the main food there was
the Common Frog Rana temporaria and probably the Swamp Frog Rana arvalis.
Quality and quantity of results improved only after studies carried out in 1976
in the Masurian Lakeland in North-Eastern Poland (Pinowski et al. 1986, 1991,
Pinowska & Pinowski 1989, Pinowska et al. 1991), and then in Southern Poland at
the end of the 20th century (Profus 2006a).
The only publication about the food of the non-breeding storks in Poland was
made by Antczak et al. (2002) on the basis of studies in the valley of the Barycz
river in the province of Wielkopolska (SW Poland), where also breeding birds were
observed. The results of the observations have been used in this paper (Table 2).
Field data
Methods of collecting material
Field observations were carried out during the whole breeding season. Most mate-
rial was collected when nests were checked, and dangerous plastic string removed,
as well as material which fell from nests with fledgling, and when ringing the
young in the second half of June and the beginning of July. The composition of the
stork’s food was identified on the basis of animals or their remains found during
nest checks. They were sometimes vomited by the young as a stress reaction to the
checking person (very rare in comparison with the Black Stork Ciconia nigra)(P.T.
Dolata unpubl.). Whole prey or their remains were also collected under nests
where they had fallen when the young had been fed, or they had been thrown out
due to problems with swallowing.
174 Jakub Z. Kosicki, Piotr Profus, Paweł T. Dolata & Marcin Tobółka
Food composition and energy demand of the White Stork breeding population 175
Table 1. Food composition of breeding White Stork population near Strzelce Opolskie
(Southern Poland)
No. of pray Mass (g) %biomass
Fishs Pisces
Brown Trout Salmo trutta m. fario 2 137.5 3.6
Chub Leuciscus cephalus 1 150 3
Gudgeon Gobio gobio 13 104 2.1
Carp Cyprinus carpio 19 845 17
Pike Esox lucius 1 400 8
Crucian Carassius carassius 1 300 6
Stickleback Gasterosteus aculeatus 2 2.5 0.05
Amphibians Amphibia
Fire-bellied Toad Bombina bombina 2 12 0.2
Grass Frog Rana temporaria 3 45 0.9
Edible Frog Rana esculenta 1 32 0.6
Frogs indeterminate Rana sp. 4 160 3.2
Reptails Reptilia
Sand Lizard Lacerta agilis 1 15 0.3
Grass Snake Natrix natrix 2 300 6
Birds Aves
Mallard Anas platyrhynchos (juv.) 1 150 3
White Stork Ciconia ciconia (juv.) 1 100 2
Mammals Mammalia
Mole Talpa europea 9 684 13.8
Common Shrew Sorex araneus 1 9 0.2
Common Vole Microtus arvalis/M. agrestis 22 488 9.8
Water Vole Arvicola terrestris 3 274 5.5
Hare Lepus europaeus (juv.) 2 260 5.2
Mouse Apodemus sp. 1 30 0.6
Invertebrate Invertebrata
Tapeworm Ligula intestinalis 2 2 0.04
Caracoles Planorbis sp. 1
Crayfish Astacus sp. 1
Horse Leech Haemopis sanquisunga 3 8 0.2
Cockchafer M. melolontha 3 2 0.04
Earthworms Lumbricidae several hundreds 455 9.3
Moreover, some data come from visual observations with the use of binoculars.
A supplementary method was to massage the necks of just fed chicks in order to
make them regurgitate their food.
These methods of collecting material, however, have many constraints because
the material obtained was not homogeneous. So the results, except those from
southern Poland, mainly show particular elements of the diet expressed as a per-
Composition of the White Stork’s food
This paper uses results of research on food from the following regions in Poland:
a) Southern Poland, an area rich in fish ponds near Strzelce Opolskie [50°31'N,
18°18'E] in Opolskie province, Pszczyna [49°58'N, 18°57'E] and Gliwice
[50°17'N, 18°40'E] in Śląskie province (for details see Profus 2006a).
b) An agricultural landscape of western Poland, near Leszno [51°51'N, 16°35'E].
In this area of 810 km2, arable fields are interspersed with meadows, pasture,
human settlements and small woods (for details see Kuźniak 1994). The White
Storks build nests mainly on roofs of buildings, trees and electric poles. During
176 Jakub Z. Kosicki, Piotr Profus, Paweł T. Dolata & Marcin Tobółka
Table 2. Food composition of breeding White Storks near Ostrów Wielkopolski and Jarocin
Fishs Pisces AB
Carp Cyprinus carpio 2
Pike Esox lucius 5
Crucian Carassius carassius 1
Gudgeon/Mullet Gobio gobio 1
Amphibians Amphibia 
Green Frogs Rana esculenta complex 4
Common Spadefoot Pelobates fuscus – froglets ca 45
Reptails Reptilia 
Grass Snake Natrix natrix 3
Mammals Mammalia 
Small indeterminate mammals Mammalia 3
Rodent indeterminate Rodentia 1
Mole Talpa europea 3
Water Vole Arvicola terrestis 1
Invertebrate Invertebrata
Earthworms Lumbricidae several hundreds
Leeches Hirudinea ca. 12
Colorado Beetle Chrysomela decemlineata several items
Bush-cricket Tettigonia viridissima ca. 10
Explanations: A – the food collected under the nest and in nests, B – the food vomit in stress reaction
recent years the number of nests located on poles increased rapidly (Kuźniak
The Obra river valley, Western Poland [52°07'N; 16°04'E], material collected
from 1997–2000. The study area covers 417 km2, where the proportions of dif-
ferent habitat are: arable field 49%, meadows and pastures 28%, forests 14%,
and 9% inhabited areas, roads, etc. (for further details see Kuźniak 1994).
c) Southern Wielkopolska: 17 sites in the Ostrów Wielkopolski district
[51°30'–51°49'N, 17°31'–18°11'E], and three sites in the Jarocin district
[51°56'–52°06'N, 17°27'–17°38'E]. This area consists of intensive arable land,
and even more extensively farmed valleys of rivers such as the Barycz (with
many fish ponds), the Prosna and the Warta. The following were collected:
three prey (only vertebrates) from three nests, 13 prey (only vertebrates) found
under eight nests, and 11 portions of food (5× vertebrates, 5× invertebrates,
1× both) from eight nests. They were either food spat out by parents for the
young or food vomited by the young during nest checks. The data were gathered
from 1994 to 2005.
Food composition and energy demand of the White Stork breeding population 177
Table 3. Food composition of breeding White Storks near Leszno
Fishs Pisces AB
Bream Abramis brama 2
Crucian Carassius carassius 1
Eel Anguilla anguilla 1
Stickleback Gasterosteus aculeatus 1
Amphibians Amphibia
Grass Frog Rana temporaria 2
Fire-bellied Toad Bombina bombina 1
Reptails Reptilia
Grass Snake Natrix natrix 1
Birds Aves
Pied Wagtail Motacilla alba 1
Skylark Alauda arvensis 2
Mammals Mammalia
Mole Talpa europea 5
Hare Lepus europaeus (juv.) 2 
Common Vole and/or Field Vole
Microtus arvalis/M. agrestis
Rat Rattus sp. 2 
Inwertebrate Invertebrata 
Earthworms Lumbricidae several items 1
Explanations – see Table 2
The above data on the White Stork’s food composition in the discussed popula-
tions are represented in Tables 1–3.
The energy value of the food was assessed only for the population from South-
ern Poland. The total weight of storks’ food was estimated at 4975 g (92 verte-
brates and several hundred invertebrates, mainly earthworms). It means that ver-
tebrates amounted to 90.4% of the food’s mass, invertebrates 9.6%. Fish
constituted the main part of food at 39% of the biomass. The biggest item was a
Pike Esox lucius which at about 400 g contained an estimated 1900 kJ of energy.
Smaller fish were also caught, for example, a Three-spined Stickleback Gasterosteus
aculeatus of 1.1 g mass and calorific value of 3.80 kJ/g of the biomass (Massias &
Becker 1990), however the energy value of an individual did not exceed 4.2 kJ. Vital
components of the stork’s diet were small mammals which amounted to 35.1% of
the prey biomass. In this group the most frequently caught animals were voles
Microtus sp. but also Moles Talpa europaea, especially in the areas of high ground wa-
ter level. Apparently, moles are easier to catch in wet areas.
The death of chicks as a result of choking on big prey
Tables 1–3 do not include information gathered from nests from 1994 to 2005
about chicks that choked while eating. In two cases the cause of death was a Mole,
in three cases a Grass Snake Natrix natrix. These data were gathered during studies
in the Ostrzeszów district (1), and mainly from the Ostrów Wielkopolski district
(4). Also, in the same area a Grass Snake (length 80 cm, weight about 200 g) was
found in a alive stork’s oesophagus. It had stuck in such a way that its ribs blocked
the oesophagus. Previously, in Polish literature it was only noted that such big ani-
mals were difficult to swallow (Jakubiec & Szymoński 2000), and, they were not
mentioned as a cause of chicks’ death in Poland (e.g. Jakubiec 1991, Profus &
Chromik 2001). Five cases from a small area suggests that such death is not rare.
The only way a bird can obtain energy is by expending metabolic energy while for-
aging for food items that contain appropriate amounts and forms of energy. Birds
must actively pursue, capture, and consume the food that they obtain their energy
from. Furthermore, in order to maintain a positive energy balance and hence suffi-
cient energy available for reproduction, birds must choose among food items that
vary in their energy and nutrient contents (Maurer 1996).
The White Stork is an opportunist as regards its food because it uses resources
that are most easily available, a notion which is proved by observations carried out
at various types of habitat (cf. Pinowski et al. 1991, Profus 2006). Calorific value of
the food shall be also considered as a significant factor. Studies performed in the
Obra river valley showed that there was a strong relationship between the quantity
of potential food, i.e. voles, and the White Stork brood size (Tryjanowski &
Kuźniak 2002). A positive correlation between the brood size and the quantity of
food was also noticed in other areas in Central Europe (Tantzen 1962, Profus &
Mielczarek 1981, Bairlein & Henneberg 2000).
178 Jakub Z. Kosicki, Piotr Profus, Paweł T. Dolata & Marcin Tobółka
Data analysis with respect to collection methods, and a comparison with check
results of spat-out-bits show that prey found under nests or that found uneaten in
nests (e.g. Table 2 A) cannot be reliably treated as the species’ food. Presumably is
because some of the prey were either spat out by storks or too large to be digested.
Fresh prey either brought for the young by parents or vomited by chicks in the nest
as a reaction to stress just before or during a nest check constitutes another group
of material that most probably appears to reflect the real food composition. Inten-
tional provocation of the birds to vomit, for example by giving them an emetic,
could be supplementary to the method. Such a method was used during research
on the Hooded Crow’s Corvus cornix food (Zduniak 2005).
Invertebrates play a really vital role in storks’ food (Pinowska & Pinowski 1989,
Pinowski et al. 1991). They are the basic component of the diet since storks arrive in
the breeding ground. The most frequent groups are predatory Water Beetles
Dytiscidae, Ground Beetles Carabidae, Scarab Beetles Scarabaeidae, Snout Beetles
Curculionidae, and the Orthoptera. Due to the long ontogenetic development of the
Orthoptera, they become the stork’s food in mid June (Pinowski et al. 1991). Another
numerous group of invertebrates in the stork’s diet are earthworms Lumbricidae.
They are the main food component just after arriving from the wintering grounds
(Hornberger 1967). Earthworms are caught throughout the whole season on pas-
tures, meadows, and alfalfa fields, and in late summer during ploughing. Research
carried out in Germany support these findings (Böhning-Gaese 1992).
In the Belorussian part of the Białowieża Forest storks mainly eat various spe-
cies of frogs (Krapivnyj 1957), such as the Common Frog Rana temporaria, the
Swamp Frog Rana arvalis and the Water Frog Rana esculenta complex, which alto-
gether amount to 60.6% of the prey biomass and meet about 53% of the chicks’
food and energy needs. Vertebrates constitute total of 72.5% of the caught bio-
mass, and meet about 64% of the energy demand of the chicks (compare Profus
1986). The remaining 27.5% of the prey biomass is made up of invertebrates, with
the Mole Cricket Gryllotalpa gryllotalpa being the main one. The Diving Beetle’s
larva Dytiscus marginalis (3.5% of the biomass) is a quite frequent item, which along
with remaining insect species amount to 10.4% of the food biomass, and about
14% of the energy the chicks need. Surprisingly, in the diet of storks in this area
there is a low contribution of earthworms at only 1% of the biomass, and a quite
high proportion of leeches (1.1% of the biomass). In July and August the main food
of chick are insects from the Acrididae and Tettigoniidea families (Krapivnyj 1957).
Additionally, studies from North-Eastern Poland (Mazuria) show that among
animals caught by storks invertebrates dominate at 75.3%, while vertebrates con-
stitute 24.7%. In the invertebrate group there are small insects (54.7%), earth-
worms (19.3%), snails Mollusca (0.6%) and big insects (0.7%); and in animals the
vertebrates such as rodents (2.7%), moles (0.15%) and fish (0.2%) (Pinowska &
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We thank J. Grześkowiak, P. Tryjanowski, M. Antczak and T.H. Sparks for their
comments on this manuscript. Thank also go to M. Antczak, J. Fijał, T. Ekiert,
Food composition and energy demand of the White Stork breeding population 179
J. Pietrowiak, S. Kuźniak and PwG OTOP members for their help in field work.The
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... In recent years, the Western European population of the white stork has recovered, as a result of changes in foraging behaviour 24 . As an opportunistic species, white storks use the most abundant and most easily obtained food 25 . Use of anthropogenic food sources available from landfills and slaughterhouses, which caused shortening or aborting migration to wintering grounds, contributed to population growth 13,24 . ...
... However, in one year of the study, 2016, the most important factors for nest-site selection apart from non-irrigated arable lands and pastures and meadows were weather conditions (climate water deficit, soil moisture, and precipitation), not distance to landfill. This shows that, whereas there may be a general preference for type of habitat, in particular years this may change in accordance with current weather conditions, that affect natural food availability 25,39 . Although it should be mentioned, that 10-km buffer might be insufficient to detect clear differences, while storks may sometimes perform further foraging trips 26 . ...
... Moreover, breeding effect, although known as the best proxy 19 , may be insufficient. Secondly, according to previous studies, abundant invertebrate prey is a critical food resource for younger nestlings 25,83 . Thus, despite occupation of nests closer to landfills, which provide alternative resources, the resulting waste food is insufficient to ensure a higher level of breeding success due to a lack of natural food resources for nestling survival at an earlier stage of development. ...
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Food wastes are among the factors with the greatest effects on animal populations. The white stork is among bird species that clearly profit from feeding at landfills, at least in Western Europe and North Africa. However, the rate and the consequences of this feeding are still unknown in the Central-Eastern European population, which differs from the western population not only in terms of migration routes but also in the greater availability of suitable natural breeding habitats due to less intensified agriculture. The aim of the study was to describe the use of landfills and its consequences in terms of probability of nest occupation and breeding effects in different regions of Poland. Although the most important factors influencing nest-site selection and breeding effect are still habitat quality and weather conditions, distance to landfills is important in selection of nest sites. White storks use landfills most intensively late in the breeding season, independently of the density of breeding pairs. The results suggest that the use of landfills is not currently essential in the Central-Eastern European population of the white stork, does not affect breeding effect, and may be more frequent in non-breeders. However, this phenomenon is still developing and requires continuous monitoring.
... Its primary foraging sites are meadows, river valleys, wetlands, and pastures (e.g. Schulz 1998;Tobolka et al. 2012Tobolka et al. , 2013, but it is an opportunist in terms of food, i.e. it uses the most easily acquired and the most abundant food (Kosicki et al. 2006). Opportunistic foraging birds may exhibit a lower level of neophobia towards new food items (Cambefort 1981) and/or foraging sites. ...
... In Algeria, the proximity of landfills was a significant predictor for breeding effect, except the one very dry year of the study (Djerdali et al. 2016b), which contradicts our results and may be due to differences in food composition between different population of white stork under different climate (compare, e.g. Kosicki et al. 2006 andChenchouni 2016). Nevertheless, a trend to nest closer to landfills is visible, also in population of storks from Central-Eastern Europe, where the foraging on landfills is a developing phenomenon . ...
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Choosing an appropriate nest site is essential for successful breeding. Changes in land use cause populations of many species to decline although some species adapt to anthropogenic changes. The white stork Ciconia ciconia commonly uses artificial nest sites. Recently, white storks from Western Europe have been using landfills as feeding sites; the beginnings of this process are being observed in Central-Eastern Europe. The study aimed to determine factors influencing the probability of nest occupation and breeding effect in a Central-Eastern European population of white storks. We used long-term data from Western Poland on breeding effect, nest occupation, the structure supporting the nest, the proximity of the nearest landfills, landfill area, and land cover. The probability of nest occupation was significantly dependent on habitat quality (based on the share of the preferred type of land cover), the structure supporting the nest, and landfill proximity within a specific year. The breeding effect was influenced by habitat quality and nesting structure. We demonstrate that the type of nesting structure is an important factor influencing both the probability of nest reoccupation and breeding effect. However, the significance of landfills appears to be growing, and in recent years, storks prefer occupying nests closer to landfills, which may have significant consequences for the population of the white stork.
... Wilharm et al. 2017;Wilharm and Skiebe 2019). Moreover, the white stork is a foraging opportunist (Kosicki et al. 2006), using anthropogenic sources of food such as landfill (Tortosa et al. 2002). Therefore, several pathogenic microorganisms have been found in storks (Olias et al. 2010(Olias et al. , 2011Szczepańska et al. 2015;Wilharm et al. 2017), rendering them suitable subjects for tests of the role of melanins in the control of microorganisms that colonise wild birds. ...
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Many organisms are characterised by strikingly contrasting black and white colouration, but the function of such contrasts has been inadequately studied. In the present paper, we tested the function of black and white contrasting plumage in white stork Ciconia ciconia chicks. We found greater abundance and diversity of microorganisms on black compared to adjacent white feathers. In addition, nest size was positively correlated with the abundance and diversity of microorganisms on white feathers. Flight initiation distance (FID), defined as the distance at which adult white storks took flight when approached by a human, was negatively correlated with most measurements of microorganism abundance. Breeding success was generally positively correlated with the abundance and diversity of microorganisms on black feathers. The feather growth rate was positively correlated with some and negatively correlated with other measurements of microbial abundance and diversity. Finally, chick growth was negatively correlated with the number of microbial species on black feathers and positively with the abundance and diversity of microorganisms on white feathers. These findings are consistent not only with the role of microorganisms in the maintenance of a benign microbial environment which differs between black and white feathers, but also with the hypothesis that several taxa of microorganisms found in black and white plumage are virulent, with negative effects on the fitness of their hosts.
... Aquatic and terrestrial beetles represent one of the most important classes of prey items for the White Stork Ciconia ciconia, a species closely related to the Oriental Stork (Kosicki et al. 2006). However, the biomass of beetles in the White Stork diet is not as high as that of other prey animals, such as voles; therefore, beetles serve as a supplementary food source to White Storks (Antczak et al. 2002). ...
By 1971, the breeding populations of the oriental stork Ciconia boyciana in Japan had become extinct in wild. After long-term conservation efforts and breeding projects, Oriental Storks have been continuously reintroduced to the Toyooka Basin, their final habitat, since 2005. The restoration of shallow wetlands in river areas and paddy fields is urgently needed in the Toyooka Basin, because these areas represent the major foraging habitats for reintroduced Oriental Storks. However, limited information is available on their feeding habits prior to extinction in the wild and, thus, it is difficult to plan specific restoration targets for these foraging habitats. In the present study, we used stable isotopic analysis to compare the nutritional status of the reintroduced populations with that of the extinct populations using preserved (stuffed) specimens. The δ15N values in the extinct populations were higher than those of the reintroduced populations. The diet proportions of the extinct populations were calculated to be equally comprised of five prey items (brackish water fish, freshwater fish, amphibians, insects, and crustaceans), while the contribution of insects was remarkably higher than that of the other prey items in the reintroduced populations. We suggest that these stable isotopic trends and diet proportions reflect a decline in the trophic level of the reintroduced populations owing to the reduced prevalence of high-nutrient prey animals caused by the loss of healthy wetlands after the 1960s in the Toyooka Basin. To increase the abundance and diversity of natural prey species available to the reintroduced Oriental Storks in the Toyooka Basin, more eco-friendly paddy fields and restored wetlands are required in the current paddy and river areas, which would secure the continuity of these wetlands from sea areas to paddy areas using fishways. However, the sampling size of the extinct populations was limited and information about historical baseline changes in prey items was insufficient. Therefore, new approaches using reintroduced populations are required to plan future reintroduction projects for Oriental Storks.
... According to Jakub et al (2006), the White Stork is an opportunist in terms of its food because it uses the resources that are most readily available, a notion proven by the observations made in different types of habitat. From a batch of 40 pelots, the insects were 97,27%. ...
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The study of diet variation of the white stork Ciconia ciconia, according to the biological cycle in the region of Tébessa is based on the decortication of the rejected pelots. In 2014, during the nesting period of this wader, rejection pelots under the nests in El-Merdja colony were regularly collected. In the study area, the White Stork consumes different ranges of prey, both invertebrate and vertebrate, where Gasteropods, Arachnids, Aves and insects were included. Its diet was different in number and depended on the different phases of the life cycle. During the pre-breeding period, the food of this bird was based on 3 orders of prey: The Orthoptera represented a peak of 39.67%, followed by the Coleoptera 35.53%, and finally the Dermaptera 24.79%. While during the breeding and rearing period, the total consumption of the Order Coleoptera was almost doubled by a maximum peak of 63.51%, the Orthoptera decreased to 8.11%, and the Dermaptera reached a percentage of 25.56% by showing a stability of consumption between the different periods of the biological cycle, as well as 2 new orders which appeared in a very minimal way: The Hymenoptera 0.71%, and the Neuroptera 2.15%. Maximum richness was observed at the young chick rearing period with 24 species, and the Shannon diversity index applied to prey species was high; This explained the high availability of many preys in the field by the need to feed the chicks, while the equitability index values was 0.76 which showed that the numbers of the consumed preys tended to be in equilibrium with each other.
... range in Europe (Mužinić & Rašajski 1992, Antczak et al. 2002, Tsachalidis & Goutner 2002. Several studies revealed that the White Stork feeds upon a wide range of prey including invertebrate and vertebrate species (Melendro et al. 1978, Antczak et al. 2002, Kosicki et al. 2006, Cheriak et al. 2014. Earthworms, orthopterans, coleopterans, and small mammals (predominantly voles in Eastern Europe) seem to be primary food resources throughout the breeding range of the White Stork. ...
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Limited quantitative data are available on food habits of the White Stork (Ciconia ciconia) in Mediterranean environments, particularly in ricefields where a relatively new food resource, the invasive Red Swamp Crayfish (Procambarus clarkii), is abundant. We studied the diet of the White Stork in a heterogeneous landscape (Central Portugal) in order to compare the importance of the Red Swamp Crayfish as a food resource in a dominant agricultural/ricefield area in relation to a predominant woodland/agricultural area. White Storks´diet was analysed spatially (two sites) and seasonally (winter, spring, summer) using pellets (n = 122) collected between December 2012 and July 2013. Overall, from 1570 prey items identified, crayfish was the second most frequent and abundant prey in the diet (frequency of occurrence, FO = 79.5%; numerical frequency , NF = 22.9%, respectively), only surpassed by coleopterans (FO = 94.3%; NF = 57.7%). However, in terms of consumed biomass (global PB) crayfish dominated the diet (PB = 44.0%), representing 1.8 times the consumed biomass of coleopterans (PB = 24.2%). Consumption of crayfish was higher in the site with highest abundance of ricefields (NF: 32.0% vs. 17.7%; PB: 51.3% vs. 38.4%). Although no significant seasonal variations were detected in terms of the number of crayfish consumed by storks, consumed crayfish biomass was significantly higher in summer in relation to other seasons. Our findings suggest that in Mediterranean heterogeneous areas the White Stork feeds upon a wide range of prey taxa though, when available, coleopterans along with Red Swamp Crayfish dominate the diet.
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White stork nestlings can provide quantitative data on the quality of the environment, as they are dependent on their parents that provide locally foraged food. Blood was sampled from the brachial vein (n = 109) and the sampling was performed in parallel with ringing during breeding season 2020 from five areas in eastern Croatia: Lonjsko polje, Jelas polje, Slavonski Brod-east, Podunavlje, and Donje Podravlje. In the present study, for the first time in Croatia, the following enzymatic biomarkers were assessed in white stork nestlings: activities of acetylcholinesterase (AChE), carboxylesterase (CES), glutathione S-transferase (GST), and glutathione reductase (GR), as well as nonenzymatic biomarkers: levels of glutathione (GSH) and reactive oxygen species (ROS). All endpoints were measured in two blood fractions: plasma and a postmitochondrial fraction (S9). Nestlings from Podunavlje and Donje Podravlje, areas known for intensive agriculture, showed lower AChE and CES activity when compared to the other investigated areas, indicating the presence of inhibitory xenobiotics. Higher oxidative stress was observed in Slavonski Brod-east, an area surrounded by metal and engineering industry, and Podunavlje compared to the other sampling areas. Hence, this study shows the impact of pollutants from the surrounding metal, petroleum, and agricultural industry might have on the biomarkers in white stork nestlings, which are often seen as early-warning signals.
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The monograph presents the results of studies of the current state of taxa and populations of rare and endangered animals, common in the Khanty-Mansi Autonomous Area. Particular attention is paid to the description of animals included in the Red Books. Recommendations on possible protection measures to preserve this group of vertebrate and invertebrate representatives of the fauna of the region are proposed. The book is intended for zoologists, environmentalists, environmental organizations, local historians, teachers and university students and anyone interested in the conservation of nature.
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Artykuł zawiera informacje o występowaniu i populacji bociana czarnego na świecie, w Polsce oraz w Biebrzańskim Parku Narodowym i „Ostoi Biebrzańskiej”, o środowisku gniazdowania, wybiórczości drzew gniazdowych, biologii lęgowej, żerowaniu i pokarmie, wędrówkach i zimowaniu oraz zagrożeniach i ochronie gatunku.
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The white stork Ciconia ciconia is a typical open-area species, foraging mainly in farmland and wetland areas. The main aim of this paper was to describe the foraging ecology of white storks inside untypical habitat, i.e. forests in Poland. Data on white stork feeding in forests were based on responses to questionnaires distributed to several national mailing lists with a total of 1700 (16% subscribers) and via emails to naturalists (mainly to white stork researchers). In total 63 observations, from the years 2000–2015, were collected, mainly from eastern Poland. In all cases, only a single adult individual was recorded inside the forest, with a mean (±SD) distance to the forest edge of 50 ± 102 m (n = 597) and 1315 ± 1015 m (n = 63) to the nearest white stork nest. Birds foraging inside forests were recorded from late May to mid-August, but the greatest numbers were seen during June. The main prey was a lizard, the slow-worm Anguis fragilis, with a maximum of 10 individuals collected by a stork during one foraging session. We discuss the origin of the observed foraging behaviour, noting that the species is flexible and opportunistic in terms of consumed food. The observed foraging is probably similar to the original behaviour of the species within primeval forest, although food opportunism helps the white stork to use new foraging areas, for example landfills.
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The influence of biotope on foraging efficiency of White Stork (Ciconia ciconia).The purpose of this paper is to assess habitat-related changes in foraging of the White Stork (Ciconia ciconia). the efficiency of their foraging, as measured by the proportion of effective attacks in the total number of attacks, and the frequency of capture, as measured by the number of individual prey species captured per unit time. The study was carried out near the town of Mikolajki (53°50’N, 21°30’E), Mazurian Lakeland in north-eastern Poland, from May 22 to August 25, 1976. During this period foraging storks were observed for 106 hours and 18 minutes. Mostly two pairs of foraging birds were observed from dawn to noon on one day and from noon to the dusk on the following day, using 7 x 50 binoculars. The dates and types of agricultural treatments were noted . It has been found that foraging time accounts for 59% of the daily activity of white storks. There were differences in the foraging time depending on the habitat and time of the season . Most frequently they foraged in habitats with short vegetation. The highest foraging efficiency was noted when the birds followed a plough and on already ploughed field with no plant cover). Most often storks attacked prey when foraging during ploughing. On the already ploughed field, the frequency of attacks was much lower . The highest number of failed attacks took place in a wetland . The foraging efficiency of young storks was much lower than that of their parents, reaching 25 and 68% respectively (p <0.001) when they foraged together on a barley stubble. Relatively large, non- identified animals (small mammals up to the size of a mole, anurous amphibians) were generally captured in ploughed fields, meadows, and pastures, least frequently in corn fields (Fig. 2). Anuran amphibians were caught most often in wet habitats, earthworms during ploughing, less frequently in meadows and pastures . The efficiency of foraging and the frequency of prey capture depends on the habitat type, which determines prey availability.
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In the Mazurian Lakeland (North - Eastern Poland), 669 pellets from 49 nests of the White Stork Ciconia ciconia were collected. In 31 % of these pellets to remnants of imaginal forms of Elateridae belonging to 12 species were found. Agriotes obscurus and Actenicerus sjaelandicus occured in the greates number. In 14 % of the pellets there were 27 species of Curculionidae. Strophosoma capitatum and Cleonus piger were most numerous. Orthoptera, almost exclusivelybGryllotalpa gryllotalpa, occured in 16% of the pellets while, Dermaptera (Forficula auricularia obly ) in 4 %.
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DIE AUSWAHL DES BRUTBIOTOPS BEIM FELDSPERLING (PASSER M. MONTANUS L.)* In dieser Arbeit werden samtliche Faktoren analysiert, die den Feldsperling das Nisten im Walde, vor allem weit von dessen Rändern, zu meiden veranlassen. Es wird auch festgestellt, dass der Einsiedlungsmechanismus wahrend der Herbstbalzzeit derselbe ist, wie wahrend der Fruhlingsbalz. WYBIÓRCZOŚĆ ŚRODOWISK LĘGOWYCH U MAZURKA (PASSER M. MONTANUS L.) Streszczenie Badania prowadzono na obszarze położonym między korytem Wisły a Puszczą Kampinoską około 15 km na NW od Warszawy (52°20* N, 20°50/ F), Na obszarze badanym powieszono w latach 1960—1963 ogółem 616 skrzynek lęgowych typu A Sokołowskiego tworząc z nich 6 kolonii leśnych i 4 kolonie wiejskie (fig. 1). Kolonie 1—IV i 26—IT były położone w lesie, ale w sąsiedztwie budynków odpowiednio 800 i 400—600 m od brzegu lasu. Kolonia 2a-W znajdowała się około 600—800 m od brzegu lasu, a 300 m od budynków i kolonii 2b—W. Kolonia 4—1^ znajdowała się 100—400 m od brzegu lasu, a kolonia'3—W' położona była aż 1000 m od brzegu lasu. Kolonia 6—W mieściła się w lesie łęgowym między wałami przeciwpowodziowymi a korytem Wisły. W latach 1960—1965 przeglądano w okresie lęgowym co najmniej raz na tydzień wszystkie skrzynki lęgowe i określano, jakie gatunki ptaków lub jakie inne zwierzęta gnieżdżą się w nich oraz w jakim stadium rozwojowym znajdują się lęgi ptaków. Młode ptaki obrączkowano w gniazdach obrączkami aluminiowymi Stacji Ornitologicznej Instytutu Zoologicznego PAN z indywidualnym numerem oraz kolorowymi obrączkami z celuloidu, odrębnym kolorem znacząc pisklęta z różnych kolonii. W okresie jesienno-zimowym kontrolowano nocą skrzynki, by stwierdzić,jakie ptaki w nich nocują. Za pomocą siatek japońskich odławiano mazurki, a w okresie zalotów jesiennych obserwowano, które mazurki (stare, młode) w koloniach 1—B', 2—U' i 3—W zajmują skrzynki lęgowe. Stwierdzono, że częstość zajęcia skrzynki przez mazurki nie zależała od tego, czy skrzynka wisiała na gołym pniu, czy była otoczona gałęziami (tab. I). Jednak skrzynki w miejscach, gdzie zwarcie koron było duże (75—100%), były rzadziej zajęte przez mazurki niż skrzynki wiszące na pojedynczych drzewach lub w terenie o małym zwarciu koron. Im głębiej w lesie położona była kolonia lęgowa, tym sto pi eh jej zajęcia przez ‘mazurki był mniejszy (fig. 2). Stopień zajęcia kolonii leśnych przez mazurki wahał się w różnych latach o wiele bardziej niż w koloniach wiejskich (fig. 2). W latach o niskim poziomie liczebności populacji mazurków było ich niewiele w koloniach leśnych (fig. 2). Dane te wskazują, że środowisko kolonii wiejskich jest optymalnym biotopem dla tego gatunku. W koloniach leśnych mazurki konkurowały z muchołówkami żałobnymi [Ficedula hypoleuca (Pall.)] i sikorkami bogatkami (Parus major L.), ale były w tej konkurencji gatunkiem dominującym. W koloniach wiejskich, w których skrzynki miały otwory wystarczająco duże dla wróbla domowego (Passer domesticus L.) (Kolonia 2—/)),ten ostatni zajmował wszystkie wolne skrzynki (fig. 6, 7, tab. V). W takiej kolonii procent zajętych przez mazurki skrzynek był o wiele mniejszy niż w innych koloniach wiejskich, gdzie skrzynki lęgowe były niedostępne dla wróbla domowego (fig. 2, tab. V). Zaloty mazurków w okresie jesiennym i wiosennym są bardzo podobne do siebie, również wybiórczość środowisk jest .analogiczna, a istniejące różnice są wywołane odmiennym stanem ilościowym populacji mazurków w jesieni i na wiosnę. W okresie zalotów jesiennych, kiedy to przypada szczyt roczny ilości mazurków aktywnych płciowo^ również i stopień zajęcia kolonii, położonych zwłaszcza w gorszych środowiskach, jest większy niż w okresie lęgowym (fig. 2, tab. X). W latach o wysokim poziomie ilościowym populacji mazurków zajmują one w jesieni w większym stopniu gorsze środowiska położone głębiej w lesie niż w latach o niskim stanie ilościowym. Gorsze środowiska są zajmowane przez ptaki młode (tab. IX). Ptaki młode, które w jesieni zajęły skrzynki lęgowe,są mniej z nimi związane niż ptaki stare i po redukcji zimowej populacji często przenoszą się na wiosnę do wolnych już, lepszych środowisk. ANSCHRIFT DES VERFASSERS: Dr. Jan Pinowski Okologisches Institut der Polnischen Akademie der Wissenschaften Warszawa, ul. Nowy Świat 72, Polen.
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We studied population size and productivity of White Storks Ciconia ciconia in 1983-2000 in 417 km2 of the Obra river valley in western Poland and its response to Common Vole Microtus arvalis density. The 33 to 60 nesting pairs fluctuated in close accordance with vole density. Similarly, the total number of fledglings produced in the population was strongly correlated with vole density. We suggest that voles are important prey because of their high calorific value. Storks arriving from their Africa wintering quarters probably evaluate the study area with respect to available prey (mainly voles) and subsequently decide whether to breed in the Obra river valley.
The sections on Field characters, Habitat, Distribution, Population, Movements, Food and Breeding have been rewritten and updated. New sections or sub-sections have been added on Abundance (population density), Breeding settlement, resettlement, site fidelity, Age structure and survival, and Conservation. The 5th International White Stork Census, carried out in 1994-95, was the most complete up to now, covering almost the entire breeding range of the species. A detailed description of the population status and development of the White Stork, including the most recent data available, is therefore one of the priorities of this account. The data are of particular interest, as since the mid-1980s the population trend of the species has changed, from rather constant declines in most countries to moderate and, in several countries, considerable increases. The section on Population now gives, country by country, a clear picture of the population status of the White Stork in its entire breeding range. It also presents an updated distribution map. Knowledge on White Stork migration has considerably improved during the last few years. This development has been strongly supported by the introduction of new research methods, particularly satellite and radar tracking. The updated account now provides a comprehensive overview of migration routes, migration behaviour, and ecology of the species during migration, based on the most recent findings. Population biology of local and regional White Stork populations has been the subject of a large number of research projects during the last few years. From these studies, data on abundance, population structure, settlement, age, and survival, which are of general interest concerning the overall population of the species, were extracted and are presented in new and updated sections of the account. The same applies to the subject of feeding ecology and breeding biology of the White Stork, the relevant sections having been updated accordingly. A new section deals with the conservation status of the White Stork. It suggests the separation of the overall population into different core and peripheral populations and describes factors responsible for particular population trends, as well as measures needed to conserve White Stork populations in the long term.
Relationships between the parental quality and reproductive performance have been studied in many birds, but not in Ciconiiformes so far. We hypothesised that parental condition of White Storks Ciconia ciconia affects both parental care and breeding success. To examine our hypothesis we assessed body mass of White Stork parents and their offspring and recorded breeding performance and parental activity in Hungary during 1992-97. Sex-related differences in parental care were found: females incubated longer than males, males compensated for reduced incubation by females, males delivered more food than females. Condition-related differences in parental effort were found, with heavier females laying more eggs and incubating longer than lighter females. Heavy males and females delivered more food to their offspring than light males and females. Parental condition was positively correlated to clutch size and provisioning rate (and therefore the survival of last hatched chicks to fledge), but hatching success was not related. Light parents raised small broods, but with chicks of relatively high body mass while heavy parents raised large broods of lighter chicks. In years with many cold days during the breeding period, White Storks produced fewer but heavier offspring than in mild years. Body mass of parents increased during the period of chick provisioning, suggesting that parents favour their own future survival in that period.
In the winters of 1993/94 and 1994/95 the daily energy expenditure (DEE) of Great Cormorants Phalacrocorax carbo sinensis was measured using the doubly labelled water technique (DLW). This was the first time the method has been used on a Phalacrocoracid species. DLW trials were carried out on 5 caged birds and on 5 free-ranging wild birds at Lake Chiemsee. The mean body mass of the captive birds (2079 g) was not significantly different from and that of wild birds (2122 g). There was no significant difference in the total body water (TBW) of the two Cormorant groups (55.9% in captive birds and 56.7% in free-ranging birds). Estimated DEE (± SD) averaged 1325 ± 130 kJ day-1 (n = 5) in the caged birds and 2094 ± 174 kJ day-1 (n = 5) in the free-ranging ones, a highly significant difference. To match their DEE, it was calculated that the Cormorants had to consume 341 g of fish per day under aviary conditions and 539 g in the wild.