balá- & balá-ová 2012_pjoe

Michal BALÁŽ1*, Mária BALÁŽOVÁ1

1 Department of Biology and Ecology, Pedagogical faculty, Catholic University, Hrabovská cesta 1, 03401 Ružomberok, Slovakia, *e-mail: [email protected] (corresponding author)

DIVERSITY AND ABUNDANCE OF BIRD COMMUNITIES IN THREE MOUNTAIN FOREST STANDS:

EFFECT OF THE HABITAT HETEROGENEITY

POLISH JOURNAL OF ECOLOGY(Pol. J. Ecol.)

60 3 629–634 2012

Short research contribution

ABSTRACT: The structural composition of the forest habitats is considered to be one of the most important factors affecting the breeding bird community composition. Structurally more homogenous forests are usually characterized by lower number of bird species and their densities. This study presents results of a comparison of the breeding bird communities in three Carpath-ian mountain forests, and the effect of the forest composition on breeding bird species richness and density. There are the bird communities oc-curring in semi-natural forests: natural mixed forest (NMF) – spruce-beech forest situated ca. 1000 m a.s.l. and UFZ-spruce forest in the upper forest zone (UFZ) up to 1510 m a.s.l., and the managed spruce forest (MSF) at altitude which is similar to NMF. The territory mapping method was used to estimate the number of breeding species and their densities during years 2004–2006. The species richness as well as the density of breeding bird communities were found to be higher in the semi-natural habitats (33 breeding species; 64.6 breeding pairs 10–1 ha in NMF and 28 breeding species; 57.6 breeding pairs 10–1 ha in UFZ) than in the managed habitat (21 breed-ing species; 53.6 breeding pairs 10–1 ha). This was true even for the high-elevated study plot despite the fact that the altitude is usually nega-tively correlated with both the species richness and density.

KEY WORDS: forest structure, bird species richness, Carpathians, habitat quality

The structure of a bird community is highly influenced by composition and con-figuration of its habitat. The major impact of vegetation pattern on breeding bird assem-blages is considered to be one of the most im-portant issues of the avian ecology (MacAr-thur and MacArthur 1961, C ody 1985).

In woodlands, which are among the best studied habitats in respect to the bird richness (e.g. Hansson 1994, Edenius and Sjöberg 1997), the differences are often found be-tween the plots of different size or age of tree species as well as between the interior and ex-terior parts of the forest fragments. The high-er values of bird diversity and density are usu-ally found in natural forests in comparison with the managed ones (e.g. Saniga 1995, Bashta 2007). Generally the natural prime-val forests are considered as habitats with the highest species richness and guild diversity (e.g. Głowaciński and Profus 1992, San-iga 1995, B oncina 2000). In this study the breeding bird communities in two Carpath-ian primeval forest habitats in Western Tatras Mountains were analyzed and compared with breeding community in a managed spruce

journal 31 v02.indb 629journal 31 v02.indb 629 2012-10-12 08:43:362012-10-12 08:43:36

Michal Baláž, Mária Balážová

forest of the same area to test the effect of the forest composition on species richness and density of the bird breeding communities.

The research was carried on in the Car-pathian forest region (N Slovakia). The stud-ied localities were characterized by different forest structure. Two of them represented semi-natural plots (“natural mixed forest” – NMF and “upper forest zone” – UFZ), while the third one represented a managed forest (“managed spruce forest” – MSF). Although the distance between NMF and MSF was only about one km, their floristic and structural composition of the vegetation was largely dif-ferent (Table 1). NMF could be characterized as a primeval spruce-beech forest with rela-tively high tree species and structural diversi-ty situated at the altitude of about 1000 m a.s.l. UFZ study plot represents natural spruce community situated on the border of moun-tain and subalpine zone at the altitude up to 1510 m a.s.l. On the other hand, MSF repre-sents a typical even-aged managed forest with very low tree species diversity situated at the altitude very similar to NMF (Table 1).

The breeding bird communities of the selected plots were studied during the years 2004–2006. The combined version of the mapping method (Tomialojć 1980) was used to estimate population densities. Species diversity (H’) and evenness (J’) were mea-sured by Shannon-Wiener index of diversity:

where pi is proportion of i-species and S is number of species, and t-test was used to test the differences.

Similarity of species composition of bird communities was measured by qualitative (QS) and quantitative (CN ) Sörensen index:

where A and B are the numbers of species in samples A and B, respectively and C is the number of species shared by the two samples, jN is sum of the lower of the two abundances for species found in both communities, na is total number of individuals in community A and nb is total number of individuals in community B.

Rarefaction method was used for better display of species richness in all localities us-ing the calculator program (Brzustowski 2005) to perform calculations. Structural characteristics of the studied plots were mea-sured within a 30 m radius from 12 points evenly distributed throughout all study plots. The cover of vegetation layers is a mean of five measurements taken from five different points within a 30 m radius. Amount of fallen and standing dead wood was subjectively estimat-ed on a nine-degree scale. Vegetation charac-teristics of the study plots were compared us-ing ANOVA with Tukey’s post-hoc tests.

Structural composition of vegetation dif-fered among all three study plots, however, MSF showed the lowest similarity in struc-tural characteristics (Table 1). This study plot was characterized by the lowest tree species diversity as well as by the lowest vegetation cover in lower vertical layers.

In total, 48 bird species were found within the three study plots during the years

Table 1. Structural characteristics of three forest stands under study.Co_decid – cover of deciduous trees, Co_con – cover of coniferous trees, Co_0–0.3 – cover of the veg-etation layer 0–0.3 m above the ground, Co_0.3–1 – cover of the vegetation layer 0.3–1 m above the ground, Co_1–3 – cover of the vegetation layer 1–3 m above the ground, amount of fallen (Dw_fall) and standing death wood (Dw_st). NMF – nature mixed forest, MSF – managed spruce forest, UFZ – upper forest zone.Forest Altitude Co_decid Co_con Co_0–0.3 Co_0.3–1 Co_1–3 Dw_st Dw_fallNMF 1020-1090 44.9 26.1 0.8 0.5 0.3 2.3 4.3MSF 920-1030 1.4 91.4 0.2 0.1 0.1 0.6 1.1UFZ 1330-1510 2.1 50 0.6 0.5 0.5 1.6 3P <0.001 <0.001 0.002 0.011 0.019 0.011 0.007NMF vs MSF <0.001 <0.001 0.002 0.023 0.387 0.009 0.005NMF vs UFZ <0.001 0.119 0.502 0.998 0.198 0.041 0.309MSF vs UFZ 0.989 0.005 0.019 0.021 0.015 0.121 0.112

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Table 2. Characteristics of breeding bird communities in three study forest stands in Carpathians. Den-sity – average number of pairs per 10 ha., dom. – % dominance. S(50) and S(90) number of breeding species in equal abundance – 50 and 90 individuals; H’ – index of diversity and J’ – index of evenness. NMF – nature mixed forest, MSF – managed spruce forest, UFZ – upper forest zone; + pair breeding on the edge of study plot, its density is not included in the overall estimate; – non-breeding species.

Species NMF MSF UFZdensity dom. density dom. density dom.

Fringilla coelebs L. 17.8 28 15.5 29 8 14Periparus ater (L.) 10.3 16 9.1 17 4.7 8Erithacus rubecula (L.) 5.6 9 6.2 12 6.9 12Regulus regulus (L.) 4.4 7 5.7 11 2.8 5Sylvia atricapilla (L.) 3.8 6 1.4 3 5.1 9Phylloscopus trochilus (L.) + + 8.2 14Prunella modularis (L.) 0.6 <1 0.5 <1 5.9 10Troglodytes troglodytes (L.) 2.8 4 1.9 4 2 3Turdus torquatus L. 0.7 1 1.4 3 3.9 7Certhia familiaris L. 2.2 3 3.3 6 0.4 0.7Phylloscopus collybita (Viellot) 1.8 3 – 3.1 5Lophophanes cristatus (L.) 1.7 3 1.4 3 2.4 4Regulus ignicapillus (Temminck) 1.7 3 1.9 4 0.4 <1Turdus philomelos C. L. Brehm 2.4 4 1.4 3 +Phylloscopus sibilatrix (Bechstein) 1.7 3 1 2Pyrrhula pyrrhula (L.) 0.3 <1 1 2 0.8 1Sitta europaea L. 1.7 3 + 0.4 <1Ficedula parva (Bechstein) 2.1 3Turdus merula L. 0.3 <1 0.5 <1 0.8 1Columba oenas L. 1.4 2Anthus trivialis (L.) 1.2 2Picoides tridactylus (L.) 1 2 +Poecile montanus Baldenstein 0.5 <1 0.4 <1Dendrocopos major (L.) 0.8 1Columba palumbus L. + + 0.4 <1Dendrocopos leucotos (Bechstein) 0.3 <1 –Dryocopus martius (L.) 0.3 <1 – –Picus canus Gmellin 0.3 <1 – –Accipiter gentilis (L.) (1)+Strix uralensis Pallas (1)+Lyrurus tetrix (L.) (1)+Carduelis spinus (L.) + + +Garrulus glandarius (L.) + + –Phoenicurus phoenicurus (L.) +Cuculus canorus L. + – +Aquilla chrysaetos (L.) –Tetrastes bonasia (L.) + –Buteo buteo (L.) – –Corvus corax L. – – –Falco tinnunculus L. –Glaucidium passerinum (L.) +Scolopax rusticola L. + +Loxia curvirostra L. + +Ficedula albicollis (Temminck) –Muscicapa striata (Pallas) – –Aegithalos caudatus (L.) –Coccothraustes coccothraustes (L.) –Turdus viscivorus L. – –Total 64.6 53.6 57.6 S(50) 22.1 16.2 19.3S(90) 28.7 20.1 24.7H´ 2.61 2.17 2.72J´ 0.81 0.81 0.87

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2004–2006. The highest number of the species was observed in UFZ, while the lowest spe-cies richness was recorded in MSF (Table 2). Altogether, 39 bird species were found to breed directly in one of the study habitats at least and 8 species breed constantly in all of them. The highest number of breeding species and the highest density of breeding community were found in NMF. The lowest number of the breeding species and the low-est density was observed in MSF (Table 2). MSF was also characterized as a habitat with the lowest species richness by the rarefaction species-individual curves (Fig. 1). There was found no difference in biodiversity between NMF and UFZ (P> 0.05, t = 0.57, df = 126). On the other hand, MSF differed significantly from both NMF (P <0.05, t = 2.19, df = 108) and UFZ (P <0.01, t = 2.72, df = 105). The similarity of communities structure reached values in the range from 0.7 to 0.78 (qualita-tive index) and from 0.51 to 0.74 (quantita-tive index). The lowest similarity was found between MSF and UFZ and between MSF and NMF in qualitative and quantitative in-dex, respectively. There were nine species de-tected as breeding species only in NMF. Six of them: Strix uralensis Pallas, Tetrastes bonasia (L.), Picus canus Gmelin, Dryocopus martius (L.), Dendrocopos leucotos (Bechstein) and Ficedula parva (Bechstein) are species of Eu-ropean importance. (according to NATURA 2000 criteria). Four species were registered only in UFZ and three of them – Glaucidium passerinum (L.), Lyrurus tetrix (L.) and Phoe-nicurus phoenicurus (L.) are species of Euro-

pean importance. The spruce plantation in MSF hosted no exclusive breeding species in respect to NMF and UFZ. The comparison of foraging guilds among localities revealed that NMF is the most valuable habitat for bird di-versity. Although canopy and ground feeders were the most abundant groups in all studied forest habitats, NMF was characterized by the highest number of foraging specialist groups. Breeding species which belonged to the air-space foragers and predators were detected only in NMF. On the other hand, MSF as the habitat with the lowest structural heteroge-neity, and UFZ as the plot situated in higher altitude, showed lower diversity of foraging specialists (Fig. 2).

As expected, what is already known from literature (e.g. B oncina 2000, Tews et al. 2004), these results show strong effect of the forest composition on the breeding bird communities. The habitats more resembling the unaffected natural conditions host bird communities with higher diversity and den-sity than do managed and changed habitats. Species composition and density of breeding pairs in NMF corresponded to what had al-ready been known about breeding bird com-munities from other parts of the Carpathian mountain forests which are usually inhabit-ed by 30–50 species with the density 60–70 breeding pairs 10–1 ha (e.g. Kocian 1981, Kieś 1991, Kropi l 1996, Korňan 2004). Bird community in another study plot with the semi-natural habitat, UFZ, is affected by the occurrence of the plot in the upper for-est zone which itself could have had some ef-fect on the community composition. Usually only low numbers of bird species could be found to breed in the upper mountain zone of the central European mountains, with the density varying from 23 to 60 breeding pairs 10–1 ha (Kozlowski 1974, Toperc-er 1989, Głowaciński and Profus 1992, Śl izowski 1991, Winding et al. 1993). The density of the bird community in the plot with the managed forest habitat, MSF, was found to correspond to the most Carpath-ian spruce forest communities (ca. 30 to 50 breeding pairs 10–1 ha) (e.g. Kr išt ín 1990, Bashta 2007), and also to other Central Eu-ropean mountain communities (e.g. D yrcz 1973, F lousek 1989, Winding et al. 1993). On the other hand, the species richness was

NMF

MSF

UFZ

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101520253035

0 50 100Number of individuals

Num

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f spe

cies

Fig. 1. Rarefaction (species-individuals) curves for three study forest stands in Carpathian Mts: NMF – nature mixed forest, UFZ – upper forest zone, MSF – managed spruce forest.

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lower in comparison with most of recorded data for such a type of habitat (21 in compari-son to 13–50; D yrcz 1973, Kr išt ín 1990, Winding et al. 1993, Bashta 2007). This could presumably be attributed to the effect of the forest structure. This forest was char-acterized by underdeveloped shrub level and young even aged trees, attributes preventing the understory and interior species (espe-cially the hole-nesters) from occurrence in higher species number.

Taking all this together, species richness, number of breeding species, density of breed-ing pairs and the diversity of foraging guilds were highest in the semi-natural study plot NMF. Forests which are heterogeneous in their structure, are characterized by higher number of ecological niches and consequent-ly by higher bird species diversity and den-sity (B oncina 2000, Tews et al. 2004). In summary, the presented results show strong influence of habitat structural heterogeneity on bird species composition. The managed spruce monoculture characterized by low val-ues of most of the important parameters im-portant for breeding birds offers less suitable conditions than the upper forest zone and the primeval spruce-beech forest. As diversity and density of bird communities are generally highly dependent on vertical and horizontal diversity of the forest offering high variety of suitable microhabitats, both these character-

istics usually decrease with increasing habitat uniformity (B oncina 2000, Bashta 2007).

The fact that the recorded diversity and abundance was significantly higher in the study plots with semi-natural habitat than in the managed forest despite that one of the for-mer ones was situated at considerably higher altitude (300–600 m), suggests that the habi-tat composition and structure resembling the natural conditions could compensate the neg-ative effect of altitude on breeding bird com-munities. This builds a framework hypothesis which could be tested on a broader altitudi-nal range of habitats with different levels of change in their natural composition on the parameters of breeding bird communities.

ACKNOWLEDGEMENTS: We would like to thank David Jandzik, Martina Jurčovičová and two anonymous reviewers for their helpful com-ments. This study was partially supported by the grant VEGA 2/0068/10.

REFERENCES

Bashta A.T. 2007 – Influence of Norway spruce plantations on the bird communities of a mon-tane forest area in the Eastern Carpathians (Ukraine) – Ornit. Beobachter, 104: 209–216.

B oncina A. 2000 – Comparison of structure and biodiversity in the Rajhevan virgin forest remnant and managed forest in the Dinaric region of Slovenia – Global Ecol. Biogeogr. 9: 201–211.

Brzustowski J. 2005 – Rarefaction calculator. http://www.biology.ualberta.ca/jbrzusto/rar-efact.php. [cited 19.8.2011]

C ody M.L. 1985 – Habitat selection in birds – Academic Press, New York, 558 pp.

D yrcz A. 1973 – Ptaki Polskiej cześci Karko-noszy [Birds of the Polish part of Karkonosze mountains] – Ochr. przyrody, 38: 213–284 (in Polish, English summary)

Edenius L . , Sjöberg K. 1997 – Distribution of birds in natural landscape mosaics of old-growth forests in northern Sweden: relations to habitat area and landscape context –Ecog-raphy, 20: 425–431.

Flousek J. 1989 – Impact of industrial emissions on bird population breeding in mountain spruce forest in Central Europe – Ann. Zool. Fenn. 26: 255–263.

Głowaciński Z. , Profus P. 1992 – Structure and vertical distribution of the breeding bird communities in the Polish Tatra national park – Ochr. przyrody, 50: 65–94.

0

5

10

15

20

25

BF CF AF GF P

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cies

NMF

UFZ

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Fig. 2. Bird foraging guilds (BF – bark feeders, CF – canopy feeders, AF – airspace foragers, GF – ground feeders, P – predators) in three study forest stands in Carpathian Mts: NMF – nature mixed forest, UFZ – upper forest zone, MSF – managed spruce forest.

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Hansson L. 1994 – Vertebrate distributions rel-ative to clear-cut edges in a boreal forest land-scape – Landscape Ecol. 9: 105–115.

Kieś B. 1991 – Bird community in a natural beech wood of the lower mountain forest zone of mt. Babia góra – Acta Zool. Cracov. 34: 519–533.

Kocian Ľ. 1981 – Vtáčie ekologické spoločenstvá v Západných Tatrách – Roháčoch [Bird com-munities in Západné Tatry mountains – Roháče] – Biologia, Bratislava, 36: 633–641 (in Slovak, English summary).

Korňan M. 2004 – Structure of the breeding bird assemblage of a primaeval beech-fir for-est in the Šrámková National Nature Reserve, the Malá Fatra Mts – Biologia, Bratislava, 59: 219 – 231.

Kozlowski J. 1974 – Liczebność i rozmieszczenie ptaków v rezerwacie „Turbacz“ w Gorcach. [Density and distribution of birds in Turbacz reservation in Gorce mountains] – Ochr. Przy-rody, 39: 245–276 (in Polish, English summary).

Kr išt ín A. 1990 – Breeding bird communities in natural and cultivated spruce forests in the Poľana Mts (In: Bird Census and Atlas Work, Eds: K. Šťastný, V. Bejček) – Prague, pp. 299–302.

Kropi l R . 1996 – The breeding bird community of the West Carpathians fir-spruce-beech pri-meval forest (The Dobroč nature reservation) – Biologia, Bratislava, 51: 585–598.

MacArthur R .H. , MacArthur J.W. 1961 – On bird species diversity – Ecology, 42: 594–598.

Saniga M. 1995 – Breeding bird communi-ties of the fir-beech to the dwarfed-pines vegetation tiers in the Veľká Fatra and Malá Fatra mountains – Biologia, Bratislava, 50: 185–193.

Śl izowski J. 1991 – Bird community of a spruce forest in the upper mountain forest zone on Polica (Polish western Carpathians) – Acta Zool. Cracov. 34: 535–551.

Tews J. , Brose U. , Gr imm V. , Tie lborg-er K. , Wichmann M. , S chwager M. , Je ltsch F. 2004 – Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures – J. Bio-geogr. 31: 79–92.

Tomialojć E .L . 1980 – The combined version of the mapping method (In: Bird census work and nature conservation, Eds. H. Oelke) – Göttingen, pp. 92–106.

Topercer J. 1989 – Ornitocenózy Štátnej prírodnej rezervácie Skalná Alpa. [Ornithoce-noses of the Skalná Alpa State Nature Reserve] – Ochrana prírody, 10: 271–287 (in Slovak, English summary).

Winding N. , Werner S . , Stadles S . , S lot-ta-Bachmayr L . 1993 – Die Struktur von Vogelgemeinschaften am alpinen Höhen-gradienten: Quantitative Brutvogel-Bestand-saufnahmen in den Hohen Tauern (Öster-reichische Zentralalpen) – Wissenschafliche Mitteilungen aus dem Nationalpark Hohe Tauern, 1: 106–124.

Received after revision April 2012

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