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SPATIAL DISTRIBUTION OF CARABIDS
and seasonat activi ty Fig. 2), and showed negative association in abundance. In the case of three species rabus hortenSiS LINNAEUS l 58 tibi S intricati XLINNAEUS lJ6l, Carabus violaceus I INNAEUS. lJ58) we have mund no significant correlations bet ween their distribution and the en viron mentat variabies and
Occurre iace of Other carabids.
In our stud y the aggregated distribution patiern was typicat for the carabids: there were l9 cases where the statistical test was applicabie and l8 distributionpatierias out of l9 were significantly aggregated Table 2). Non-randona spatiat distributions were common ly reported for carabids LUFF l 986. NIEM EI A l 988a NIEM EI A et a l. l 992ab). The analyses of indicator species and the variation incarabid catches among traps and habitats Tabies i and 3) also fhow that the collected carabid species have a clear habitat and microsites preferences with specific en Viron mental conditions. The causes of the variation in catches among traps and habitats are unclear LUFF l 986. NIEM EI A et tit l986). One difficultyis that descriptions of the en vironment are rare ly accurate en Ough to Suppon inserences abolit determinants of spatiat distribution NIEMELA & SPENCE l 994). In the literature, mur not mutuatly exclusive factors are mentioned that might expla in the spatiat distribution os carabid beetles NIEME A l 988a. NIEM EI A et al. l 985. l 988. l 996) l) differe iaces in enuit onmental conditions se. g. habitat heterogenei ty, ωOd re SOurces, microclimate) 2) autecological characteristi cs of the species, 3) small- scale dispersal, and so) interspecific interactions. Our analyses s howed that a significant proportion of the variation in calchwas associaled with a particular kind of en viro iamental heterogenei ty reflected by the cover of leas litter, herbs, shrub s. canopy layer and by the distribution os ca- rabids' preys Table d). This may be a fairly generat patieria among carabicis be- cause clear prefere iaces for microsites defined by ground vegetation and litterhave also been detected et sewhere THIELE l9JJ. NIEMEL A l 990. NIEM EI A et til. l 992a). Multiple regression analyses s howed that canopy layer was a significant positive predictor for Ahax Parallelmi eclus PILLER et MITTER PACHER. lJ83). Ahax partillet meduS PILLER et MITTER PACHER. lJ83) is a habitat generalist Table l); but significantly more individuals were captured in the forest interior
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tive correlation belween the number of individuals and the degree of aggregationfor the eighteen studied species. This may be interpreted as more abundant SpecieS are more aggregated. It may al SO be regarded as a simple statistical rule. Indeed. it is evident that both the mean and the variance are dependent on den-Sity, even when the patieria is not randona. BARTI ETT l 936) suggested that therelative Variance was likely to be a linear function of the mean. Zoologi sis frequently attribute a similar statement to I AO sl 968) for the relationship of Mori sita index and the mean. The Mori sita index is closely related to the index os dispersion. TAYLOR et al. l9 8) studi ed the patiern and density relationship for ahuge variety of different species, and they concluded that the multiplicative law suggested by TAYLOR sl 96 l) provides a better fit between Vari ance and mean. This relations hip is usual ly mentioned as Taylor's power law SOUTHwOODl9 8). The observed relations hip during our study is clearly a variant of the Tay
lationShi p. On the studied spatiat scale the different habitats gras S, forest edge, forest interior) are not independent of each other and there appears to be movement bel-ween the adjacent habitats NIEMEL A l 988h. NIEM EI A and HALME l 992). Although the majority of the species silowed clear differences in abundance bel-ween the habitat types, most species were also mund ovisi de their optimat en vi- ronment Table l). Small- scale dispersal between habitat patches may be caused by density-dependent processes GRUM l9 l), and also by movement belween reproduction habitat and hibernation habitat sWALLIN l 986. ANDERSEN l99J).
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SPATIAL DISTRIBUTION OF CARABIDS
At the studied spatiat scate, the occurrence of other carabids was a significant predictor of the abundance of a particular dominant and subdominant species Table d). Most of the correlations were positi Ve, SuggeSting Simit Ar response to habitat properties. Previous works NIEM EI A et al. l 992h. NIEMELA &SPENCE l 994) also reported some negative relationships. These relation sit ipswere usual ly between species of clearly different body si Zes or seasonat acti- vities, or both, and are probab ly better attributed to different microhabitat prenereiaces than to effecis of interspecific interactions. In our study there was a negative relations hip hetween the spatiat distribution of the smali Synuchus vivialis PANZER. lJ9J) and the much larger CartibuS circensis HERBST. lJ84 which alsos howed maximum activities at different times of the season. For similar reason Sinterspecific competition is not a likely explanation for the other patrs of species s howing a significant negative relationship Cartibus arcensis HEREST. t 84 and
Interspecific competition may be suspected to contribute to the negative re
l99J. l 998a. h) suggest that negative interactions are not a generat feature of these two species. NIEMELA l 988a) and NIEM EI A and SPENCE l 994) also reported two carabid species with similar siZe and sea sonat activity which showed negative interaction in a forest habitat. while these species in other forest associations showed a high overlap in spatiat distribution and in seasonat activi tys Ni EMEI A et ah l 992h. NIEMELA & HALME l 992) or there were no correlationbetween the occurrence of the two species NIEM EI A et til. l 993). These facts suggest that the type of relationships belween the occurre iace of carabids may nothe a generat Dature. The interactions can vary among habitat types with different en Viron mentat attributes. Further manipulative and non- manipulative studies are neces Sary to prove the presence of competition belween MoloPS PiceuS PANTER lJ93) and Pterostichus hv eisteri HEER. l84l hecati se distributionat data spatiat ly non-overi apping distribution) are just the first step to evaluate the mechan is in os interspecific competition. Overali. Our resulis suggest that carabids are useful as an indicator group to asses the en viron mentat variation, as they show different habitat choices. Our
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phasi Ze the priori ty of abiotic factors in the determination of carabid distributionand in the organi sation. Our resuli sti esses that a synthesis is needed. whicli integrates competition with other abiotic and hiotic ecological factors. This synthesis would hel p a tot in undet standing communi ty organi sation os carabilis. but in this reSpeci, we are stili at the beginning NIEM EI A l 993).
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SPATIAL DISTRIBUTION OF CARABIDS
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In the bulky volumes of the 'Contributions to a Manual os Palaeca reticDiPtera' morphological, physiological. genetical, ecological and economicup-to-date knowledge of dipterous species smidges and flies). which have significant importance in genetics as modet organisms, in plant culti vation as pests or beneficiat parasitoids, in animal hushandry and human health asVectors of serious il inesses and whicli are important for ecosystem functi On. are treated. Morphological keys with excellent figures) for adulis and larvae. whicli hel p re aders with identification of dipterous pesis and parasi- toids are provide l. while readers in fie id os applied dipterology will findsultable en viron mental ly friendly methods against pests or biological controi methods, among Others. Time table of series: l99J - Volume 2 l 998 Volume 3 Higher Brachycera). l 999 - Volume l Generat and Applied Dipterology) and Appendix Volume. The 2nd volume contains 38 dipterous fami ly chapters by 23 speciali sis frona l2 countries. with l895 figures on 2 8 plates.
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Acta Mologica Academiae Scientiarum Hungaricae 46 I). m. I 26. 2000
DETECTION OF SPATIAL AUTO COR RELATIONAM ONG BIRD TERRITORI ES BASED ON LINETRANSECT CENSUS ES
Spatiat autocorrelations in the territories of mur marshland-nesting passerines Sedgewarhler Acrocmhialtis schoenobaenus. Reed Warhler A. scimaceus, Reed Bunting Emheriaci schoenicius and Savi's Warbier Locustella luscinioides) were demonstrated by correlograms computed by Moran 's I coefficient. The data set was based upon territory mapping of bree-ding hirds in His-Balaton Nature Reserve Hungary). For the Sedge warbier Seven consecutive line transect censu ses were also evaluated. Except for the first two transect censu Ses which were carried out at the beginning of the breeding season, autocorrelation analysis of the line transect censuses revealed similar patierias as the more accurate, but more time-consuming. territory mapping. Spatiat autocorrelation analysis of transect data revealed the mainseatures of the autocorrelation structure. the high value at any of the first three lags, hut noautocorrelation at lag 4 or 5. I conclude, that the line transect censu ses provide adequate datato deieci spatiat relationships in territorial hirds. Key wordso spatiat autocorrelation. Moran 'S I. line transect, hird census, mars hiarad. Acroce-ohaluS Sp.
tecting patierias of territori es of marshland-nesting hird species. There are many other is sues that can he addi essed by spatiat autocorrelation. Spatiat autocorrelation analysis is sui table to reveat relations hips among Sample Sites in a geographic Space, where arran gements of Sample si te S could he irregular se. g. ran