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Opportunity makes a predator: Great Spotted Woodpecker predation on Tit broods depends on nest box design.

Authors:
Adam Kaliński at University of Lodz
Adam Kaliński
Joanna Skwarska at University of Lodz
Joanna Skwarska
Jaroslaw Wawrzyniak at University of Lodz
Jaroslaw Wawrzyniak
Jerzy Banbura at University of Lodz
Jerzy Banbura
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Brief report
Opportunity makes a predator: Great Spotted Woodpecker
predation on Tit broods depends on nest box design
Joanna A. Skwarska, Adam Kaliñski, Jaros³aw Wawrzyniak
& Jerzy Bañbura
J. A. Skwarska, A. Kaliñski, J. Wawrzyniak & J. Bañbura, University of Lodz, Department
of Experimental Zoology and Evolutionary Biology, Banacha 12/16, 90237 £ód, Po-
land. E-mail joaskw@biol.uni.lodz.pl
Received 5 January 2009, accepted 30 July 2009
Secondary cavity nesters are species that breed in
cavities made by other species. A variable but
marked proportion of their nests undergo preda-
tion, especially in areas where specialized and/or
opportunistic predatory animals are abundant. In-
deed, predation is the most common cause of nest
loss in primeval forests (Walankiewicz 2002a,
2002b, Weso³owski 2002, Thompson & Burhans
2004, Weatherhead & Blouin-Demers 2004,
Yamaguchi et al. 2005). Because of their special
construction, nest boxes may be less accessible to
predators than are natural cavities, especially if
they are equipped with protective devices, such as
metal plates surrounding the entrance hole
(Walankiewicz 1991). On the other hand, nest
boxes can be more easily spotted by predators and
humans. In addition, ornithologists often distrib-
ute them at regular distances apart, which may
make them still easier to find (Mänd et al. 2005).
Under these circumstances, wooden nest boxes
without protective devices may greatly suffer from
predation. Interestingly, while building their nests,
secondary cavity breeders seem to maintain the
distance between the cavity entrance and the nest
cup by modifying the nest size (Alabrudziñska et
al. 2003, Kosiñski & Ksit 2007, Mazgajski &
Rykowska 2008). The reason may be that the
greater this distance is, the more difficult it will be
for larger predators to reach eggs or nestlings in
the nest.
Supplying forested habitats with large num-
bers of nest boxes usually increases the abundance
of cavity-nesting birds in those habitats (von
Haartman 1957, Enemar & Sjöstrand 1979, Slags-
vold 1975, Alerstam 1985, Stañski et al. 2008).
This increase in potential prey density may attract
more predatory animals, thus increasing nest
losses and potentially leading to the establishment
of an ecological trap for secondary cavity nesters
(Mänd et al. 2005). Indeed, the rate of nest loss in
study sites of some projects had become so high
that counter-predator measures had to be under-
taken. For example, in the long-term study on
Great Tits (Parus major) in Wytham Wood, UK, it
was necessary to change the design of nest boxes
when the originally-used wooden boxes started to
be depredated by weasels (Mustela nivalis) (Dunn
1977).
Studies in primeval forest conditions have
shown that the Great Spotted Woodpecker (Den-
drocopos major) is an efficient avian predator of
nests located in tree cavities (Walankiewicz
2002b, Czeszczewik & Walankiewicz 2003). This
species is also a common destroyer of wooden nest
Ornis Fennica 86:109112. 2009
boxes (Mainwaring & Hartley 2008). However,
the scale of nest-box brood losses caused by
Woodpeckers in relation to nest-box construction
has not been experimentally studied so far. In our
study on Blue (Parus caeruleus) and Great Tits,
predation on nestlings by Great Spotted Wood-
peckers occasionally took place. The aim of this
note is to show that woodpecker predation on tit
nestlings is not random with respect to the nest-
box construction.
The present study was carried out between
2005 and 2008 as part of a long-term project on
secondary cavity breeders using wooden nest
boxes set up in a mixed deciduous forest near
£ód, central Poland (see Marciniak et al. 2007 for
details of the study area). We used two types of
nest boxes differing in the construction of the front
wall. In the first type (Type 1), the front wall was
placed between the side walls, with a 12-mm gap
between the walls on both sides (Fig. 1a). In the
other type (Type 2), the front wall extensively cov-
ered the edges of the side walls so that no gap was
left on the front side (Fig. 1b). Under the protocol
of the study, nest boxes were visited at least once a
week to establish the occupancy by tit species and
their stage of reproduction. We recorded signs of
predators having been present, with particular at-
tention to evidence for woodpecker predation
(Fig. 1c). Because the destroyed nest boxes were
replaced by new Type 2 boxes and events of suc-
cessful Great Spotted Woodpecker predation on tit
nestlings were infrequent, we pooled the data for
four successive years. We recorded a total of 297
nest-box broods of Blue and Great Tits; of these,
194 were in Type 1 (with front gaps; 65.3%) and
103 were in Type 2 (without front gaps; 34.7%)
nest boxes.
We tested the frequency with which nest boxes
were attacked by Great Spotted Woodpeckers in
relation to the distribution predicted from nest-
box-type availability using the Chi-square test of
the goodness of fit (Zar 1996).
We detected 12 cases of Great Spotted Wood-
pecker predation on Blue and Great Tit nestlings.
These all concerned broods in Type 1 nest boxes,
i.e., those with front gaps (Fig. 1a). In all of these
cases, the woodpecker had drilled an opening by
enlarging the original side gap (Fig. 1c) in order to
get at the tit nestlings. Twelve Type 1 nest boxes as
compared with zero Type 2 boxes is evidently bi-
ased toward the former, when compared to an ex-
pected random proportion that should be 7.84 ver-
sus 4.16, respectively (X2= 6.37; df = 1; p= 0.016).
Even if the rate of Great Spotted Woodpecker
predation on Blue and Great Tit broods in nest
boxes in the studied woodland was not high, it was
110 ORNIS FENNICA Vol. 86, 2009
Fig. 1.a–Type1nest box with a gap between the front and side walls.b–Type2nest box, without gaps.
c – A Type 1 nest box with a gap enlarged by Great Spotted Woodpecker.
significantly related to the nest-box type. Only the
nest boxes with side gaps between the front wall
and the side walls were destroyed by woodpeck-
ers. We did not record a single case of an unoccu-
pied nest box being destroyed by woodpeckers.
Moreover, no single nest box was robbed at the
stage of egg laying or incubation either. The type
of nest-box damage described in this note always
concerned occupied nest boxes at the nestling
stage of breeding.
We suggest that the clear effect of nest-box
type on predation rate resulted from movements of
tit nestlings being easily detectable and begging
calls being more audible to woodpeckers in the
boxes with gaps. The generally low number of
cases of woodpecker predation suggests that Great
Spotted Woodpeckers may not be selective in at-
tacking nest boxes in comparison with their ex-
ploitation of natural food sources, but they may ef-
ficiently take the advantage of an opportunity of
easily getting nestlings. A practical recommenda-
tion for conservation purposes or scientific studies
would be to only use nest boxes without front gaps
in order to limit woodpecker predation on nest-
lings.
Mainwaring and Hartley (2008) reported a
similar case of the destruction of wooden nest
boxes and predation on tit nestlings. In their case,
the opening, enabling access to tit nestlings, was
drilled from the crack between the side wall and
the rear wall. Mainwaring and Hartley (2008) pro-
posed covering the nest box with wire mesh to pre-
vent woodpecker predation, and experimentally
confirmed the efficiency of this method in de-
creasing woodpecker predation rate.
Although the diet of Great Spotted Woodpeck-
ers during the breading season basically consists
of insects and other invertebrates collected on
trees, the presence of nestling items in it seems also
a consistent feature of the species (Cramp 1985).
Nilsson (1984) found that the Great Spotted
Woodpecker may be responsible for as much as
48% of predation on tit nest boxes. Woodpeckers
have been shown to be important predators of
birds nesting in natural tree cavities as well (Wa-
lankiewicz 1991, 2002a, Weso³owski 2002). The
case described in this paper shows that the nestling
predation by at least one species of woodpeckers is
rather opportunistic, and probably a consequence
of normal searching for prey on tree-trunks.
Käpytikan tiaisiin kohdistama pesäsaalistus
riippuu pöntön rakenteesta
Käpytikan tiedetään toisinaan suurentavan pönttö-
jen liitoskohtia päästäkseen käsiksi poikasiin. Täs-
sä tutkimuksessa vertailtiin kahteen eri tavoin ra-
kennettuun tiaispönttötyyppiin kohdistuvia, tikko-
jen aiheuttamia pesäpoikastuhoja: tyypin 1 pön-
tössä oli etu- ja sivulautojen väliin jätetty 12 mm
rako, tyypin 2 pöntöissä rakoa ei ollut. Sini- ja tali-
tiaisen pesäpoikasiin kohdistunutta saalistusta tut-
kittiin 297 tiaispesyeellä, joista 12 joutui käpyti-
kan saalistamiksi. Kaikki 12 tapausta  joissa tikka
oli aina suurentanut pöntön rakosia  kohdistuivat
tyypin 1 pönttöihin; ero tyyppiin 2 (nolla tapausta)
on tilastollisesti merkitsevä. Tutkijat esittävät, että
tikat kykenevät pöntön rakosista helpommin ha-
vaitsemaan poikaset ja tarttumaan tilaisuuteen hel-
pohkon saaliin toivossa. Tällaisten tapausten vält-
tämiseksi pöntön rakennuksessa etu- ja sivu-
lautojen väliin ei tulisi jättää rakosia.
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112 ORNIS FENNICA Vol. 86, 2009
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... This likely reflects the ability of species such as woodpeckers, snakes and rodents to physically access nests in cavities (Kuitunen and Aleknonis 1992;Czeszczewik 2004). We included studies using nest boxes, which are generally designed to reduce predation rates (Skwarska et al. 2009) and may have lower predation rates than nests in natural holes (Kuitunen and Makinen 1993) but we do not believe this affected these trends. Approximately half of the studies of hole nesting species included in our review were on populations nesting in natural holes, and although they suffered higher predation rates by corvids (1.52% ± 1.15) than nest box studies (0.14% ± 0.14), the reported predation rates were much lower than corvid nest predation observed on open nesting species. ...
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Full-text available
  • Jun 2021
Una lámina de plástico fijada alrededor de los árboles puede impedir que las serpientes ataquen las cajas nido de las aves Se han diseñado varios artilugios para impedir que los mamíferos y las aves ataquen las cajas nido. A pesar de que las serpientes son uno de los depredadores más comunes de las aves que anidan en cavidades, siempre han sido difíciles de evitar. En el presente artículo probamos un método originalmente concebido para evitar que los mamíferos trepen a los árboles. Para impedir que las serpientes trepen a los árboles y ataquen las cajas nido, utilizamos una lámina de acetato transparente de 80 cm de altura para envolver los troncos de los árboles en los que se ubicaba una muestra de cajas nido (N = 40) utilizadas por carboneros y herrerillos. Las demás cajas nido (N = 74) se dejaron como control. La tasa de depredación en los nidos de control fue del 20 % y solo del 2 % en las cajas nidos experimentales. El método puede ser útil para aumentar el éxito reproductivo de las aves y, por lo tanto, para aumentar la eficacia de los recursos dirigidos a obtener datos científicos, y el éxito reproductor de las especies en peligro de extinción.
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... Consequently, snake-specific alarm calls may not have evolved in northern Europe. Instead, great spotted woodpeckers (Dendrocopos major) are one of the major predators of the eggs and nestlings of tits in Europe (Skwarska, Kalinski, Wawrzyniak, & Banbura, 2009), although in Japan, the same species of woodpecker lives in the same habitats as tits but does not attack their nests. Future comparative studies may reveal the socioecological factors that contribute to driving the evolution of referentiality and compositionality in bird calls, thereby providing a unique model for examining the principles and general rules which drive the evolution of linguistic capabilities. ...
Animal linguistics: Exploring referentiality and compositionality in bird calls
Article
Full-text available
  • Jan 2021
Establishing the theory of language evolution is an ongoing challenge in science. One profitable approach in this regard is to seek the origins of linguistic capabilities by comparing language with the vocal communication systems of closely related relatives (i.e., the great apes). However, several key capabilities of language appear to be absent in non‐human primates, which limits the range of studies, such as direct phylogenetic comparison. A further informative approach lies in identifying convergent features in phylogenetically distant animals and conducting comparative studies. This approach is particularly useful with respect to establishing general rules for the evolution of linguistic capabilities. In this article, I review recent findings on linguistic capabilities in a passerine bird species, the Japanese tit (Parus minor). Field experiments have revealed that Japanese tits produce unique alarm calls when encountering predatory snakes, which serve to enhance the visual attention of call receivers with respect to snake‐like objects. Moreover, tits often combine discrete types of meaningful calls into fixed‐ordered sequences according to an ordering rule, conveying a compositional message to receivers. These findings indicate that two core capabilities of language, namely, referentiality and compositionality, have independently evolved in the avian lineage. I describe how these linguistic capabilities can be examined under field conditions and discuss how such research may contribute to exploring the origins and evolution of language. Understanding the origins and evolution of language is an ongoing challenge in science. Recent field studies revealed that several key features of language (referentiality and compositionality) have independently evolved in the avian lineage, providing a unique opportunity to explore the principles and general rules of language evolution.
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Creating wildlife habitat using artificial structures: a review of their efficacy and potential use in solar farms
Article
  • May 2024
The biodiversity crisis is exacerbated by a growing human population modifying nearly three‐quarters of the Earth's land surface area for anthropogenic uses. Habitat loss and modification represent the largest threat to biodiversity and finding ways to offset species decline has been a significant undertaking for conservation. Landscape planning and conservation strategies can enhance habitat suitability for biodiversity in human‐modified landscapes. Artificial habitat structures such as artificial reefs, nest boxes, chainsaw hollows, artificial burrows, and artificial hibernacula have all been successfully implemented to improve species survival in human‐modified and fragmented landscapes. As the global shift towards renewable energy sources continues to rise, the development of photovoltaic systems is growing exponentially. Large‐scale renewable projects, such as photovoltaic solar farms have large space requirements and thus have the potential to displace local wildlife. We discuss the feasibility of ‘conservoltaic systems’ – photovoltaic systems that incorporate elements tailored specifically to enhance wildlife habitat suitability and species conservation. Artificial habitat structures can potentially lessen the impacts of industrial development (e.g., photovoltaic solar farms) through strategic landscape planning and an understanding of local biodiversity requirements to facilitate recolonization.
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Predator-Proofing Avian Nestboxes: A Review of Interventions, Opportunities, and Challenges
Article
Full-text available
  • Dec 2023
Nestboxes are commonly used to increase the number and quality of nest sites available to birds that usually use tree cavities and are considered an important conservation intervention. Although usually safer than natural cavities, birds nesting in simple, unmodified wooden nestboxes remain at risk of depredation. Accordingly, numerous design and placement modifications have been developed to ‘predator-proof’ nestboxes. These include: (1) adding metal plates around entrance holes to prevent enlargement; (2) affixing wire mesh to side panels; (3) deepening boxes to increase distance to nest cup; (4) creating external entrance ‘tunnels’ or internal wooden ledges; (5) using more robust construction materials; (6) developing photosensitive shutters to exclude nocturnal predators; (7) using baffles to block climbing mammals; and (8) regular replacement and relocation. However, the benefits and costs of these modifications are not always well understood. In this global review, we collate information on predator-proofing avian nestboxes designed for tree cavity-nesting birds to assess the efficacy of techniques for different predators (mammalian, avian, and reptilian) in different contexts. We critique the potential for modifications to have unintended consequences—including increasing nest building effort, altering microclimate, reducing provisioning rate, and elevating ectoparasite and microbial loads—to identify hidden costs. We conclude by highlighting remaining gaps in knowledge and providing guidance on optimal modifications in different contexts.
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Effects of Nest Characteristics on Breeding Success of Great Tits Parus major
Article
Full-text available
  • Dec 2003
In this study, nest characteristics (size and proportions of basic components) were not correlated with the timing of breeding. Clutch size was negatively correlated with total nest mass but positively correlated with the proportion of the mass of the lining in the total nest mass. Analyses of hatching and fledging success showed that the quantity and proportion of moss in the nest structure as well as the nest size influenced the performance of eggs and nestlings at the nest. We suggest that variation in nest size and composition may be due to several contradictory pressures associated with the need to keep the moisture and temperature in the nest relatively constant, to protect the brood from predation, and to control sanitary standards.
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Nest Predation as a Limiting Factor to the Breeding Population Size of the Collared Flycatcher Ficedula albicollis in the Białowieża National Park (NE Poland)
Article
Full-text available
  • Dec 2002
This study is based on an analysis of nest-histories of 652 Collared Flycatcher nests found in natural tree cavities in the Bialowiez̀a National Park in 1988-1999. The secondary-cavity-nester Collared Flycatcher constitutes migratory single-brooded population breeding in high density in the primeval oak-lime-hornbeam (Querco/Tillio-Carpinetum) stands. Nest predation was the main reason of the breeding losses (240 nests) accounting for 91% (82%-100%) of them. Local production of fledglings was affected by nest predation caused by rodents, mustelids and Great Spotted Woodpecker. In this study a link between forest rodent cycles and the Collared Flycatcher fluctuations in number was documented. Number of produced fledglings depended on both, positively the number of the Collared Flycatcher breeding pairs in year N and negatively on the Yellow-necked Mouse density in year N. Rate of nest destruction is related to the density of the Yellow-necked Mouse recorded in BNP, while independent on the Collared Flycatcher density (nest predation limiting but not regulating). The predation pressure in some years keeps the Collared Flycatcher density at a level well below that of the potential the habitat resources (nest-sites, food). Local breeding density was shaped by fledglings productivity (breeding success) of the previous year. Earlier hypotheses concerning the Collared Flycatcher and other birds population limitation were also discussed.
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Breeding Losses in the Collared Flycatcher Ficedula albicollis Caused by Nest Predators in the Białowieża National Park (Poland)
Article
Full-text available
  • Jul 2002
The proportion of different nest predators robbing nests of the Collared Flycatcher Ficedula albicollis population breeding in natural cavities in the Białowieża National Park was studied during 1988–1998. Robbed nests were examined for signs betraying the predator. Mustelids, (mostly Martes martes) destroyed no more than 25% of the Collared Flycatcher nests, while rodents (mostly Apodemus flavicollis), and woodpeckers (Picoides major) were together responsible for the destruction of c. 75% of all the Collared Flycatchers broods. Rodents destroyed significantly more nests with eggs, whereas woodpeckers and Martes martes destroyed mostly broods with nestlings.
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The Evolution of Nest-Site Selection among Hole-Nesting Birds: The Importance of Nest Predation and Competition
Article
  • Oct 1984
Predation was the major cause of total nest failure. Predation rates on nests in natural holes were higher than in nest-boxes for great tit Parus major and pied flycatcher Ficedula hypoleuca, while no significant difference was found for blue tit P. caeruleus and marsh tit P. palustris. In tits, woodpeckers were responsible for 17% of predation on nests in natural cavities but for 48% on nests in boxes. Of nests that were preyed upon, woodpeckers destroyed a lower proportion of those of great than of those of blue tit and marsh tit. Minimum nest entrance widths were correlated with the size of the species. Depths of nesting holes were generally similar for different species, but blue tit occupied shallower holes than did great tit. Starling Sturnus vulgaris, nuthatch Sitta europaea, and blue tit occupied holes higher up in the trees than did great tit and pied flycatcher. Marsh tits nested very low. Total rates of nest failure and predation were greater in low nests than in higher ones for starling, blue tit, and marsh tit. All 4 species that vary their nest heights in relation to density prefer to nest high. This indicates that there is competition for safe nest sites. Starling reduced the breeding success of nuthatch by taking over holes occupied by the latter. -from Author
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Adaptation in Hole-Nesting Birds
Article
  • Sep 1957
A number of analogous (convergent) adaptations occur in hole-nesting birds. Their main causes are the safe nesting sites and the keen competition (both intra- and interspecific) for them. Competition has caused certain characteristics in behavior, such as development of territorial behavior only after finding a suitable hole, fighting for a nesting-hole instead of for a territorial area, and the male's demonstration of the nesting-hole in courtship-display. The safety of the nesting site has caused other adaptations, such as frequent polygamy, hissing notes in incubating adults and in the young, lack of cryptic color in the eggs, large clutch size, and slow development of eggs and young. A long period of evolution is needed, however, before a species is able to take full advantage of the safety of the nesting site. In primary hole-nesters the adaptations are much more clearly developed than in secondary hole-nesters. Tits, for instance, have maintained spotted eggs, open-nesting Parrots white eggs. The Pied and Spotted Flycatchers have nearly identical growth curves in the young, in spite of the one having very safe and the other very unsafe nesting sites, and so on. The number of subspecies was found to show a positive correlation to clutch size (hence, hole-nesting birds tend to have larger number of subspecies). This may be due to stronger selection in species with more offspring, and/or the fact that species with large clutches are relatively often non-migratory.
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Predation by Weasels (Mustela nivalis) on Breeding Tits (Parus Spp.) in Relation to the Density of Tits and Rodents
Article
  • Jun 1977
(1) Annual and seasonal variations in predation rates by weasels (Mustela nivalis L.) on tits were studied in Wytham Wood, Oxfordshire. Four Parus species of tits breed in the nest boxes which have been provided for them in the wood since 1947. (2) Weasels are by far the most important cause of nest failure in the wood and on average 20.8% of nests were preyed upon by them each year from 1947 to 1975. (3) There were marked annual fluctuations in predation rate, varying from 0-50%. Predation was insignificant until 1957, two years after myxomatosis cleared the wood of rabbits. This caused widespread predation pressure on small rodents and may have led weasels to explore nest boxes as an alternative source of food. (4) The nesting density of tits and the density of rodents, measured the same summer, together accounted for over 60% of the annual variation in predation rates; both prey types appear to be equally important. The possible influence of weasel density on predation rates is also discussed. (5) Because predation on tit nests is proportional to nesting density, Coal Tits Parus ater L. apparently suffer less predation than the other species by starting to breed first when few other nest boxes contain eggs. The staggered laying also results in interspecific variation in the stages (laying, incubation, nestling) at which nests are most likely to be preyed upon. (6) Using the combined nests of Blue Tits and Great Tits as a `prey population', it is shown that a high nest density hastens the onset of predation but if rodent density is high this tends to retard the rate of predation on tit nests. (7) This effect of prey density on the timing of predation accounts for seasonal variations in predation rates between 1959-63, described by Perrins (1965), and explains the decline in predation on nestlings after 1963. The relationship found here may also help to explain why Krebs (1970) found that predation on nestlings was not density dependent. (8) The breeding strategy of the tits is discussed and found to be little defence against predation by weasels. Laying date may, however, affect the incidence of adult mortality but there is insufficient information on survival to calculate overall fitness.
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Competition between the Great Tit Parus major and the Pied Flycatcher Ficedula hypoleuca: An Experiment
Article
  • Jan 1975
Literature dealing with competition between the Great Tit and the Pied Flycatcher for nesting sites and food is reviewed, and some data from English and Finnish populations are re-analysed. A preference difference is shown to exist for nesting holes, but because there is a general lack of such places, the two species regularly compete for them. There is some overlap in hunting behaviour and in the food items brought to the young. The interspecific competition may influence breeding numbers and reproductive success, depending on the availability of holes, density of birds, habitat type, and the times of egg-laying. When the interval between the onset of breeding between the two species is short, the flycatcher is favoured.
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Effects of the Introduction of Pied Flycatchers Ficedula hypoleuca on the Composition of a Passerine Bird Community
Article
  • Jan 1972
A nest-box experiment was carried out in rich subalpine birch forests, Swedish Lapland. During a six-year period the passerine bird density in an area (23 hectares) with a surplus of nest-boxes was compared with the density in a control area (29 hectares) without boxes. A large number of the nest-boxes were occupied by one species, the Pied Flycatcher, increasing the bird density in the nest-box area by about 75 per cent. This immigration of the flycatcher did not cause a demonstrable change in the population numbers of the other species. The year to year fluctuations of the flycatcher numbers were in accord with the fluctuations of the other species taken together in the same area and also with the fluctuations of the bird numbers in the control area. The result of this investigation is compared with that of a similar experiment in Southern Sweden.
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