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IZVORNI I ZNANSTVENI ČLANCI – ORIGINAL SCIENTIFIC PAPERS Šumarski list br. 9–10, CXXXIV (2010), 475-486 UDK 630* 114.6 + 411 (001) VARIATIONS OFCARABID BEETLE AND ANTASSEMBLAGES, AND THEIR MORPHO-ECOLOGICALTRAITS WITHIN NATURAL TEMPERATE FORESTS IN MEDVEDNICA NATURE PARK RAZLIKE U SASTAVU I MORFOLOŠKO-EKOLOŠKIM ZNAČAJKAMA MRAVAI TRČAKAU PRIRODNIM ŠUMAMANAPODRUČJU PARKAPRIRODE MEDVEDNICA 11 2 Lucija ŠERIĆ JELASKA, Ana JEŠOVNIK, Sven D. JELASKA, 311 Aljoša PIRNAT , Mladen KUČINIĆ, Paula DURBEŠIĆ SUMMARY: The aim of this study was to investigate responses of ant and carabid assemblages and their morpho-ecological traits to habitat differences within natural temperate forests in Medvednica Nature Park. Toquantify habitat differences in examined areas, both structural heterogeneity of the vegetation and taxonomic diversity of plants were measured on six plots. Habitat complexity was quantified using four habitat characteristics within the site: tree canopy cover; shrub canopy cover; ground herbs and leaf litter cover. Ants and carabids were sampled using pitfall traps. Ant species richness and abundance, unlike carabid species richness were positively correlated with habitat complexity, especially with leaf litter cover on plots. The responses of insects morpho-ecological traits to habitat were recorded, with more large bodied carabids present in more complex site and higher abundance of opportunist ant species in more open sites with low complexity of vegetation. Higher dominance of certain carabid species at the lower plots then those on the top of the mountain, suggest competitive exclusion, confirming lower areas as more stable. Species adapted to colder climate, that inhabit higher elevations such as flightless forest specialist Cychrus caraboidesandCara- bus irregularis, and boreo-montane ant species Camponotus herculeanus, are less competent to colonize lower areas. Furthermore, they may not survive severe instability of their habitats, especially in a changing climate. Overall results suggest that conservation issues need to be focused on preserving stability and structural complexity of forest habitat in summit areas of the mountain. Key words: biodiversity, vegetation structure, litter, altitude, nature conservation, forest habitat INTRODUCTION – Uvod Mountain landscapes usually offer steep gradients perature, precipitation, etc.), what is reflected in the for a wide range of environmental parameters (i.e. tem-adaptation and distribution of different types of organisms and ecological communities that occupy different 1 Dr. sc.Lucija Šerić Jelaska, dipl. prof. Ana Ješovnik, prof. dr. sc. Mladen Kučinić, prof. dr. sc. Paula Durbešić, positions along these gradients (review inHodkinson Division of Biology (*Group for Systematic Zoology & Entomo 2005). Many of these environmental factors are multifa logy), Faculty of Science, University of Zagreb, Rooseveltov trg 6, ceted and interlinked in defining overall structural com 10000 Zagreb, Croatia, Phone: +385 1 4877711; Fax: +385 1 4826260, e-mail: slucija@zg.biol.pmf.hr plexity of insect habitats, which Hodkinson (2005) 2 Doc. dr. sc. Sven D. Jelaska, Division of Biology (*Group has found to decrease with increasing altitude.The latter forTerrestrial biodiversity), Faculty of Science, University affects not only the species richness but also the species of Zagreb, Marulićev trg 20, 10000 Zagreb, Croatia 3 composition of insect communities (Whittaker 1952 Mr. sc. Aljoša Pirnat, Groharjeva 18, SI-1241 Kamnik, Slovenia |
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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486 in Hodkinson 2005). The physical structure of the environment, mediated by plant communities, can inf luence the distribution and interactions of species (review in Lawton 1983). “Structural heterogeneity hypothesis” assumes that structurally complex habitats may provide more niches and environmental resources for exploitation and thus increase species diversity, although empirical support for this relationship is biased towards studies of vertebrates and habitats under anthropogenic influence (Tews et al. 2004). Lassau and Hochuli (2004) and Lassau et al. (2005) showed that habitat complexity in forests may affect the composition of ant and beetles assemblages.Brose (2003b) showed that effects of habitat heterogeneity on ground beetle assemblages were positive on the micro- and 2 meso-scale (0.25 and 500–1000 m, respectively), and 2 were not significant on a macro-scale (10 km). In “taxonomic diversity hypothesis” plant taxonomic diversity is positively correlated with the diversity of herbivore insects (Murdoch etal. 1972,Root1973), and thus with predator diversity (Hunter and Price 1992). The importance of floristic richness for the ground beetle abundance was also reported by Baguette( 1993). Ants and ground beetles have been widely recommended as environmental, ecological and biodiversity indicators (e.g. Altegrim et al. 1997, Andersen 1997, Niemelä 2000, Szyszko et al. 2000, Andersen et al. 2002,Antonova and Penev 2006, Pearce and Venier 2006, Šerić Jelaska et al. 2007,Šerić Jelaska and Durbešić 2009). They respond well to natural and anthropogenic disturbances. Microhabitat characteristics, i.e. litter type, organic matter content, insolation, temperature fluctuation, are important for providing hunting, foraging niches and for protection of beetles from predators, ovipositioning, larval development, over wintering etc. (Thiele 1977,Pearce and Venier 2006). Theuse of carabids morpho-ecological traits like wing morphology, body size or diet are recommended in the evaluation of habitat quality (Blake etal. 1994,Gutiérrez et al. 2004,Gobbi andFontaneto 2008).Brose (2003a) found that large carabids, as more preferable prey because of their higher nutritive value and reduced foraging time required, prefer dense vegetation plots as enemy-free spaces.According toAndersen et al. (2002), ant functional groups can be used as indicators of habitat quality and forest health under different management (Stephens and Wagner2006). Lassau andHochuli (2004) suggested measurement of habitat complexity as a surrogate for the diversity of a range of arthropods including ants.Very often, different taxonomic groups indicate certain areas as important for protection or nature conservation, showing mutual positive correlations. Based on the existence of these correlations, the use of “surrogate species groups” for estimation of overall biodiversity has been reported (Garson etal. 2002;Satersdal etal. 2003); being particularly useful where limited biodiversity data exist. Vascular plants have been reported bySatersdal et al. (2003) as a good surrogate group, which also works for estimation of ground beetle diversity. Although, some habitat preferences are known for many species of ants and ground beetles, it is not always evident which environmental factors are the most responsible for species assemblage and distribution. The aim of this study was to analyze ants and carabid species richness and abundance, carabids body size and ants functional groups, in different habitats within natural temperate mature forests, and to test whether higher habitat complexity and plant species richness support higher insect diversity along vertical gradient in Mt. Medvednica. MATERIALAND METHODS – Materijal i metode Study Area – Područje istraživanja Mount Medvednica is part of the Croatian continental karst, located north of Zagreb, the capital of Croatia. The great floristic diversity (Dobrović et al. 2006), due to the mosaic structure of forest communities, geographical position and geological structure make this area very interesting for biodiversity investigations. Dobrović etal. (2006) reported that the floristic richness and diversity of the mountain was higher than similar regions in Croatia and several European countries (Italy,Austria, Slovakia, Poland and Serbia). For protection of its valuable forest areas, the western part of Mt. 2 Medvednica (228 km) was proclaimed a nature park by the Nature ProtectionAct in1981. Thehighestpeak of the mountain, named Sljeme, is placed 1035 m above sea level. Despite protection, anthropogenic pressures on Med vednica are constantly increasing (road construction, recreation, ski trails, logging, etc.), mainly at its upper elevations, changing the habitat quality.This can change structural complexity of forests, especially where trees and shrubs form part of the natural landscape. Surveyed forests are natural temperate ones on brown acid soil with silicate parent rocks (Basch 1995). For this investigation, six plots (50 x 50 m) were selected along the central profile of Mt. Medvednica across a vertical gradient on both slopes of the mountain (Figure 1).They were placed in mature stands of five different forest communities, that account for 78% of the total forest cover of the nature park:Querco–Castaneetum sativaeHt. 1938, (plot 1);Luzulo–Fagetum sylvaticaeMausel 1937, (plot 2);Lamio orvale–Fage |
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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486 Figure 1 Position of the study area - Medvednica Nature Park (black polygon) and position of investigated plots in the Park Slika 1. Područje istraživanja i položaj istraživanih ploha na području Parka prirode Medvednica |
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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486 tum sylvaticae Ht. 1938, (plot 3), Festuco drymeiae-sured. Because of its circular nature, the terrain aspect Abietetumass. nova,Vukelić &Baričević2007, was transformed into a continuous north-south gradient (plots 4 and 6);Chrysanthemo macrophylli–Aceretum (northness) by calculating the cosine of aspect values pseudoplatani (Ht. 1938) Borh. 1962 (plot 5), (Table (Guisan etal. 1999,Jelaskaetal. 2003). 1). For each plot, altitude, slope and aspect were mea- Table 1 Ecological variables and their values for six plots Tablica 1.Vrijednosti ekoloških varijabli na šest istraživanih ploha Variable/plot (50x50 m) 1 2 3 4 5 6 community/zajednica* qc lf lo af cm af Organic matter content/udio org. tvari (%) 8.99 16.36 15.96 21.47 18.82 13.28 Number of plant species/br. biljnih vrsta 62 19 41 43 36 39 Altitude/visina (m) 400 550 660 970 810 660 Slope/nagib (o) 21 18 17 28 21 7 Aspect/aspekt (o) 145 272 102 325 354 335 Northness/sjevernost -0.819 0.035 -0.208 0.819 0.995 0.906 Canopy openness/otvorenost sklopa (%) 4.04 10.62 4.95 12.69 4.98 4.94 Leaf area index/indeks lisne površine 3.91 2.59 4.13 2.34 3.81 3.41 Soil type/tip tla Brown acid/kiselo smeđe * qc- Querco- Castaneetumsativae, lf- Luzulo- Fagetumsylvaticae, lo- Lamioorvale- Fagetumsylvaticae, af – Festuco drymeiae - Abietetum, cm- Chrysanthemomacrophylli–Aceretumpseudoplatani Habitat complexity was quantified using four habitat used them in our analyses.Three characters (shrubs cavariables within the site: tree canopy cover; shrubs ca-nopy cover; ground herb cover; amount of leaf litter) nopy cover; ground herb cover and amount of leaf litter. were visually quantified into four categories (from score Each variable was scored with 0, 1, 2 and 3 using an 0 to score 3: low contribution to site complexity (score ordinal scale, where increasing scores indicate greater 0), the highest contribution to site complexity (score 3). habitat complexity.This is modified version of the tech- Tree canopy cover was measured as Leaf Area Index niques used byLassau andHochuli (2004, 2008). (LAI) and classified into four categories with scores Compared to theLassau andHochuli (2004) sco from 0 to 3. Summing up the scores for all quantified ring method, there were no differences in two additional characters, habitat complexity range from 5 (lowest ha- variables i.e. soil moisture and portion of logs, rocks and bitat complexity) to 10 (maximum habitat complexity) debris between investigated plots, hence we haven’t per plot (Table 2). Table 2 Ecological variables used to derive measures of habitat complexityand their values for six plots Tablica 2.Ekološke varijable korištene za određivanje kompleksnosti structure staništa na šest istraživanih ploha Variable/plot (50x50 m) 1 2 3 4 5 6 Tree canopy cover (LAI)/sklop krošnji (LAI) 3 (3.91) 1 (2.59) 3 (4.13) 1 (2.34) 3 (3.81) 2 (3.41) Shrub canopy cover/pokrovnost grmlja 2 0 1 0 0 2 Ground flora cover/pokrovnost prizemnog bilja 2 3 2 3 3 1 Leaf litter cover/količina listinca 3 3 3 1 2 1 Habitat complexity scores/vrijednosti kompleksnosti staništa 10 7 9 5 8 6 In each of the six plots, the number of vascular plants species was recorded on successive visits during the season (March – October). Data on canopy openness and LAI, as indirect estimators of available under story light, were measured using hemispherical photographs (as described inJelaska 2004 andJelaska et al. 2006) and analyzed with Gap Light Analyzer (GLA) software (Frazer etal. 1999). Soil samples were collected twice during this investigation, at the beginning of the season in March 2001 and in May 2002. Four samples of the top 10 cm of soil were collected from each plot, after removal of the litter. Samples were diagonally placed across the plot, at 14 m intervals following general scheme in Scholz et al. 1994.To determine the percentage of organic matter, 10 g of soil was ashed at 400oC. All environmental variables are displayed inTable 1. |
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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486 Carabid Beetles andAnts Sampling – Uzorkovanje trčaka i mrava Carabid beetles and ants were collected in 16 pitfall traps on each plot, placed in a regular rectangular net with cell size of 10 x 10 meters placed five meters from the edges of the plot.Altogether, 96 pitfall traps were exposed through the investigation period.Traps were filled with ethanol (96%), acetic acid (9%) and water in equal proportions and emptied every two to three weeks from March 2001 toApril 2002. Carabids and ants were identified using specialized keys (Collingwood 1979, Agosti and Collingwood 1987, Trautner and Data Analyses – The total abundance and species richness of ants and carabids was determined for each habitat (i.e. plot).The Shannon-Wiener and Sorensen indices (Krebs 1989) were calculated for all plots to assess ground beetles and ants’diversity and similarity using Programs for Ecological methodologyVer.5.2 (15-III. 2000). Sorensen similarity values between plots were used in cluster analyses with the Euclidean distance as distance measure and single linkage as linkage rule for constructing RESULTS Atotal of 9288 beetles and 2958 ants belonging to 43 and 20 species, respectively, were trapped (Table 3). The number of captured species per plot varied from 17 to 27 for carabids, and from 5 to 14 for ants.Abax parallelepipedus, Abax paralellus, Aptinus bombarda,Carabus violaceusandCychrus attenuatuswere carabids found at all surveyed plots (Table 3). Only one ant species Myr- Geigenmüller 1987,Hůrka 1996,Freude et al. 2004 andSeifert 2007). AccordingtoAndersen et al. (2002) ants were classified in four functional groups. Based on personal measurements of carabid specimens and data from Hůrka (1996) and Freude et al. (2004), average body size for analyzed species were classified into three categories: (1) small (4–8 mm); (2) medium (8–21 mm); and (3) large (22–40 mm) using a size distribution graph. Analiza podataka dendrograms using STATISTICA 6.1 (StatSoft Inc. 2003).Wecompare habitat complexity scores, plant species richness and measured environmental variables (soil organic matter, slope, aspect and elevation) with data about richness and abundance of ants and carabids, ant functional groups and carabids body size distribution patches using Pearson Product Moment correlations in STATISTICA6.1 software. – Rezultati mica ruginodiswas found at all plots, unlikeCrematogaster schmidti,Camponotus herculeanus,Formica gagatesandFormica rufarecorded at single plot (Table 3). Ant species richness was negatively associated with carabid richness per plot (r = -0.86;p= 0.029), opposite to the trend for their abundance (r = 0.85; p=0.031). Table 3 Carabid and ant species with number of specimens per plot. Plot number corresponds to those in Figure 1 Tablica 3.Vrste i brojnost jedinki trčaka (Carabid) i mrava (Ant) na istraživanim plohama. Broj plohe odgovara onome na slici 1. Carabid species/Plots 1 2 3 4 5 6 Abax carinatus(Duftschmid 1812) 156 23 2 1 26 Abax ovalis(Duftschmid 1812) 1 3 37 1 Abax parallelepipedus(Piller & Mitterpacher 1783) 654 89 328 5 61 50 Abax parallelus(Duftschmid 1812) 498 2 283 1 145 136 Agonum gracilipes(Duftschmid 1812 1 Platynus scrobiculatus(Fabricius 1801) 1 19 Amara aeneaDe Geer 1774 3 2 1 1 Aptinus bombarda (Illiger 1800) 1108 4 11 3 213 45 Bembidion lampros (Herbst 1784) 1 Bembidion deletumAudinet-Serville 1821 1 Carabus convexusFabricius 1775 445 343 362 1 Carabus coriaceusL. 1758 34 23 82 2 10 Carabus gigasCreutzer 1799 16 5 1 Carabus intricatusL. 1761 132 49 78 1 3 Carabus irregularis Fabricius 1792 72 85 12 Carabus nemoralisO. F.Müller 1764 636 379 52 15 6 Carabus praecellensPalliardi 1825 18 3 2 24 35 Carabus ulrichiiGermar 1824 548 29 100 52 8 Carabus violaceusL. 1758 23 168 43 29 35 63 |
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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486 Cychrus attenuatus(Fabricius 1792) 43 221 70 44 99 43 Cychrus caraboides(Linnaeus 1758) 5 1 Dromius fenestratus (Fabricius 1794) 3 Harpalus atratusLatreille 1804 1 Harpalus marginellus Dejean 1829 2 Harpalus laevipesZetterstedt 1828 1 Harpalus dimidiatus(P. Rossi 1790) 1 Leistus piceusFrölich 1799 5 3 1 Leistus rufomarginatus (Duftschmid 1812) 3 1 1 62 67 Licinus hoffmannseggii(Panzer 1803) 2 23 83 3 Molops elatus(Fabricius 1801) 9 27 4 Molops piceus(Panzer 1793) 20 27 5 1 37 Nebria brevicollis (Fabricius 1792) 1 Notiophilus biguttatus(Fabricius 1779) 1 5 1 Notiophilus rufipesCurtis 1829 23 3 1 Platyderus rufus(Duftschmid 1812) 4 24 Pseudoophonus rufipes(De Geer 1774) 1 1 Pterostichus anthracinus (Illiger 1798) 1 Pterostichus fasciatopunctatus (Creutzer 1799) 2 11 Pterostichus oblongopunctatus (Fabricius 1787) 121 120 Pterostichus transversalis (Duftschmid 1812) 5 7 79 Stomis rostratus (Sturm in Duftschmid 1812) 1 1 Synuchus vivalis(Illiger 1798) 1 4 1 Trechus cardioderus Putzeys 1870 1 Number of species/broj vrsta 18 1721 2726 Number of specimens/broj jedinki Ant species/Plots 4350 1346 1449 374 1117 631 Aphaenogaster subterranea(Latreille 1798) 1 57 2 3 4 4 5 6 9 Camponotus fallax(Nylander 1856) 2 Camponotus herculeanus(Linnaeus 1758) 2 Camponotus ligniperda(Latreille 1802) 11 144 4 7 Crematogaster schmidti (Mayr 1853) 26 Dolichoderus quadripunctatus(Linnaeus 1771) 8 1 1 1 Formica fuscaLinnaeus 1758 14 18 1 Formica gagatesLatreille 1798 22 Formica polyctenaFörster 1850 154 1 Formica rufaLinnaeus 1758 2 Lasius brunneus(Latreille 1798) 14 65 11 2 91 Lasius citrinusEmery 1922 1 1 Lasius emarginatus(Olivier 1792) 629 180 4 1 3 Lasius fuliginosus(Latreille 1798) 1 3 1 Myrmecina graminicola(Latreille 1802) 28 16 Myrmica ruginodisNylander 1846 50 244 127 199 54 156 Myrmica sabuletiMeinert 1861 4 2 Prenolepis nitens(Mayr 1853) 135 11 Stenamma debile(Förster 1850) 6 10 17 35 Temnothorax crassispinus(Karavajev 1926) 134 44 165 2 20 Number of species/broj vrsta 14 12 11 5 5 10 Number of specimens/broj jedinki 1126 No significant correlations were found between plant richness and arthropod diversity. Habitat complexity scores were positively correlated with carabid and ant abundances and ant species richness, but negatively correlated with carabid species richness (Table 4). Of 761 478 209 66 318 the four environmental variables assessed by scoring method, shrub and leaf litter cover were positively correlated with ant species richness and carabid abundance. Large negative correlations were found between carabid richness and litter cover (-0.90,p= 0.015). |
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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486 Figure 2 Tree diagram of cluster analyses using Sorensen indices based on carabid species composition in investigated plots as similarity measure. Numbers on X axes denote investigated plots Slika 2. Dendrogram sličnosti istraživanih ploha koristeći Sorensen indeks ovisno o prisutnosti vrsta trčaka tijekom razdoblja uzorkovanja. Broj na osi X označava plohe Figure3 Tree diagram of cluster analyses using Sorensen indices based on ant species composition as similarity measure. Numbers on X axes denote investigated plots Slika 3. Dendrogram sličnosti istraživanih ploha koristeći Sorensen indeks ovisno o prisutnosti vrsta mrava tijekom razdoblja uzorkovanja. Broj na osi X označava plohe |
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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486 Table 4 Correlation coefficients between ant and carabid assemblages and habitat variables Tablica 4.Koeficijenti korelacije između sastava mrava i trčaka sa stanišnim čimbenicima Habitat variables /stanišne varijable Community features /osobine zoocenoza Habitat complexity score / kompleksnost staništa Plant richness / broj biljnih vrsta Altitude / visina Aspect (northness) Organic matter / organska tvar (%) Carabid richness /broj vrsta trčaka Carabid abundance /abundancija trčaka Carabid Shannon indeks /Shannon indeks trčaka r=-0.62 p=0.192 r=0.82 p=0.044 r=-0.25 p=0.637 r=0.04 p=0.946 r=0.66 p=0.157 r=0.15 p=0.780 r=0.84 p=0.035 r=-0.80 p=0.058 r=0.44 p=0.388 r=0.87 p=0.024 r=-0.85 p=0.031 r=0.73 p=0.102 r=0.57 p=0.239 r=-0.80 p=0.057 r=0.14 p=0.788 There was no correlation between the heterogeneity of vegetation structure and plant species richness. Habitat complexity was negatively correlated with altitude (r = -0.70;p=0.12) and with aspect (r = -0.77;p= 0.074). The same trend was found for ant richness, abundance and diversity indices for ants’ communities (Table 4). Carabid species richness was positively correlated with altitude and aspect, opposite to carabids abundance and ant assemblages (Table 4). Large and small bodied species showed opposite trends in their distribution across investigated sites. Large carabids were positively cor related with leaf litter cover (r= 0.81, p=0.05). Small carabids positively correspond with higher altitude (r= 0.89, p=0.018). The most abundant carabids wereA. parallelepipedus, A. paralellus, A. bombarda,C. nemoralisandC. ullrichi(Table 3) respectively, accounting for 37.16% of the total catch. The highest number of specimens, comprising 47.11% of all specimens, was captured in the most complex site (plot 1), though this plot recorded almost the lowest number of species (i.e. 18, as compared to the minimum of 17 on plot 2). On the contrary, the highest number of carabid species (27) was recorded on plot 4 (with the lowest habitat complexity score) but with the lowest abundance making only 4% of captured specimens. The highest number of ant species (14) and abundance (1126 specimens) was recorded on plot 1. The lowest number of ant species and their abundance was recorded on plots 4 and 5. 45% of them are woodland species and only 10% of sampled species prefer open habitats.The rest of them occur in both woodland and open habitats.Analyzing the functional groups, according toAndersen (1997), 65% of recorded species (13 species) are cold climate specialists (CCS), 3 species are opportunists (O), 3 of them are subordinate camponotini (SC) and 1 species belong to generalized Myrmecinae (GM). Presence of species representatives of all four groups were recorded on plot 1, only.M. ruginodis( opportunist) that was found on all plots had the highest abundance in the low complexity areas (plots 2 and 5).The abundance of opportunist ants was negatively correlated with canopy cover (r= -0.91, p=0.05). High correlations were found between ant abundance and richness with the amount of organic matter in the soil (Table 4). Organic matter content in the soil rise with higher altitude (r= 0.9, p=0.012). Cluster analyses of species composition on plots revealed differences in carabid assemblages between the southern (plots 1, 2, 3) and northern slopes (plots 4, 5, 6) of Mt. Medvednica (Figure 2). Some species such asCarabus convexus,Notiophilus rufipesandCarabus intricatuswere found exclusively on the southern slope of mountain, unlike Carabus irregularis,Notiophilus biguttatus, Molops elatus,Platyderus rufusand all of the collected species of the genusPterostichus, which were recorded exclusively on northern slope of the mountain. There was also a significant difference in number of collected ants on southern and northern slopes (2365 on south, 593 on north). Cluster analyses using Sorensen indices based on ants’species composition, separate the two highest plots, 4 and 5, from others (Figure 3).An ant speciesC. herculeanusis recorded for plot 4, which is 970 m high and has northern exposure.C. herculeanus has altitudinal range from 1000–1700 m, and boreal mountain distribution, with a very restricted area. It prefers coniferous mountain forests and can withstand very low temperatures, 38.5 oC below zero (Seifert2007). |
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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486 DISCUSSION– Rasprava The relationship between habitat features and diversity of ants and carabid beetles was analyzed.We recorded great differences in carabid species composition and abundance among plots, where carabid abundance, but not the richness, was significantly higher in habitats with higher complexity scores.Brose (2003a) showed that an experimental reduction of vegetation complexity reduced the activity-abundance of large carabid species. Our results supports “enemy-free space hypothesis” (Lawton,1983) that prey species have more chances of escaping from natural enemies in dense vegetations. We found that large carabids prefer dense vegetation plots covered with leaf litter as enemy-free space. Furthermore, carabid species richness was negatively correlated with ant species richness.The results of ant richness analyses differ from those of Lassau and Hochuli (2004), but not those analyzing functional groups where they also found the largest abundance of opportunist species in low complexity sites. Stephens and Wagner (2006) have found that species richness, diversity, and dominance were a less satisfactory measure of various forest management impacts on ants than functional group analysis. The highest number of ant species (14) and abundance (1126 specimens) was recorded on plot 1, and the lowest number of the species and their abundance on plots 4 and 5. This result was in accordance with their habitat preferences.Ants in general, with excep tion of a few cold-temperate species, are thermophilic animals, and function poorly below 20 oC and not at all below 10 oC (Hölldobler and Wilson, 1994). Greater abundance and species number in lower altitudes can be explained, in addition to temperature, with precipitations, thickness and volume of leaf litter and available food resources (Brühl etal. 1999). The highest Shannon-Wiener index for carabid diversity (Table 5) was recorded on plot 5, situated in the Chrysanthemo macrophylli–Aceretum pseudoplatani forest, with high concentration of soil organic matter as a consequence of the longer persistence of snow-cover and hence higher soil humidity and a shorter micro-organism activity period.The herbaceous layer of this forest is characterized by nitrophilous plant species e.g. Lunaria rediviva L., Urtica dioicaL., Corydalis solida (L.) Swartz, etc.The impact of leaf litter origin on carabids has also been reported byNiemeläetal. (1992), Koivula etal. (1999) andMagura etal. (2005), in which the latter two authors observed significant impact of leaf litter on certain species e.g. Pterostichus oblongopunctatus and C. caraboides. In this study, both above mentioned species were found only on plots 4 and 5, which had the highest soil organic matter content. Soils in cooler climates commonly have more organic matter because of slower decomposition rate (BotandBenites2005). Table 5 Shannon-Wiener (H’) indices, numbers of equally common species (N) and Smith & Wilson evenness (S&W) for investigated plots Tablica 5.Shannon-Wiener (H’) indeksi raznolikosti, broj zajedničkih vrsta (N) te Smith & Wilson jednolikost (S&W) trčaka (carab.) i mrava (ants) na istraživanim plohama Plots / Indices 1 carab. ants 2 carab. ants 3 carab. ants 4 carab. ants 5 carab. ants 6 carab. ants H’ 3.049 2.262 2.724 2.585 2.951 2.074 3.201 0.366 3.711 0.977 3.625 1.936 N 8.27 4.80 6.61 6 7.73 4.21 9.19 1.29 13.10 1.97 12.34 3.83 S&W 0.138 0.26 0.153 0.39 0.128 0.322 0.291 0.22 0.182 0.293 0.218 0.295 On plots 1, 2 and 3, smaller number of species was recorded, while plots 4, 5 and 6 had a higher number of species with a low abundance, indicating less structured arthropods communities without dominant species. Also, cluster analyses of species composition on six plots divided carabids based on aspect and ants species based on elevation of investigated plots. Ant and carabid diversity and species body size distribution highly correlate with altitude and aspect (expressed as northness). On warmer, more south exposed plots (plots 1, 2 and 3) smaller number of carabid species was recorded. On the contrary, plots on northern slopes (plots 4, 5 and 6) had a higher number of species with a smaller number of specimens. Salgado et al. (1997) reported similar findings in their research of deciduous oak forests, where 10 of 42 species represented 90% of the total catch on five plots. They recorded a smaller number of species on plots with higher abundance of specimens and those with smaller abundance and larger number of species, the latter having unstable climatic conditions. In this study, plot 1, with recorded capture of 47.11% of the total catch, is situated at the lowest altitude with southern exposure, thereby ensuring more stable and warmer climate conditions that enable formation of ground beetle communities with the dominant and co-dominant species present (e.g. A. bombarda, A. parallelus, A. parallelepipedus, C. ullrichi, C. nemoralis, C. convexusandC. intricatus; Table 3). The highest number of carabid species, but the lowest abundance was recorded at the highest altitude (plot 4) with the lowest habitat complexity score. Flightless forest specialist such as C. caraboides that preferred higher |
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L. Šerić Jelaska,A. Ješovnik, S. D. Jelaska,A. Pirnat, M. Kučinić, P. Durbešić: VARIATIONS OF CARABID ... Šumarski list br. 9–10, CXXXIV (2010), 475-486 elevations and lower temperatures andC. irregularisare distributed on top of the mountain with a small number of individuals on the northern plots. Geomorphologic and climatic conditions, large habitat complexity and very low disturbances in plot 1, contribute in forming more stable carabid and ant communities. Community composition can be used to indicate broader aspects of habitat quality and more general changes (i.e. degradation and recovery following stress or disturbances), (Hodkinson andJackson 2005). High abundance of carabids but low richness at the warmer plots could suggest competitive exclusion confirming that habitat stability may have unimodal effect on richness. The intermediate disturbance hypothesis predicts that diversity will be greatest when physical disturbances prevent competitively dominant species from excluding other species from the community.At the low level of disturbances, diversity is low because only the best competitors persist and competitive exclusion leads to species loss at either end of the disturbance continuum (Death andWinterbourn 1995).Stephens andWagner (2006) found that different ant functional groups were dominant under different levels of disturbance intensity. Data about species richness and habitat complexity within undisturbed forest systems with low direct anthropogenic impacts can be used as reference data for environmental monitoring of changes in temperate forests. Knowledge on how management of forests relates to forest carabids and ants diversity is poorly documented in Croatia.Wefound that ant functional groups and carabids body size analyses respond well to differences in habitat complexity.These results confirm the need for sustainable forest management that will preserve higher level of habitat complexity that provide more niches and environmental resources for exploitation and thus support dominance of larger carabids and high animal biomass. Furthermore, carabids and ants may serve as target groups in climate change risks assesment in mountain ecosystems. ACKNOWLEDGEMENTS – Zahvala Grateful thanks to Gregor Bračko for helping us in tian Ministry of Science, Education and Sports (Grants species determination.This study was financed by Croa-0119-123, 119-1193080-1206 and 119-0000000-3169). REFERENCES – Literatura Agosti,D., and C.A.Collingwood,1987: Aprovisional list of the Balkan ants with a list to the worker caste. II. Key to the worker caste, including the European species without the Iberian. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 60:193–261. Altegrim,O., K.Sjöberg,and J. P.Ball,1997: Forestry effects on a boreal ground beetle community in spring: Selective logging and clear- cutting compared. Entomol. Fennica 8: 19–26. Andersen, A.N., 1997: Using ants as bioindicators: Multiscale issues in ant community ecology. Conserv. Ecol. (online) 1 (1): 8 URL:http://www.consecol.org/vol1/iss1/art8. Andersen, A. N., B. D. Hoffmann, J. Müller, and A. D. Griffiths, 2002: Using ants as bioindicators in land management: simplifying assessment of ant community responses. J.Appl. Ecol. 39: 8–17. Antonova,V.,and Ly.Penev,2006: Change in the zoogeographical structure of ants (Hymenoptera: Formicidae) caused by urban pressure in the Sofia region (Bulgaria). Myrmecologische Nachrichten 8: 271–276. Basch,O., 1995: Geološka karta Medvednice (Geological map of Medvednica), 1:62500. Institut za geološka istraživanja (In: Geološki vodić Medvednice, Ed. Šikić K.). Institut za geološka istraživanja, Zagreb, INANaftaplin, Zagreb. Baguette, M., 1993: Habitat selection of carabid beetles in deciduous woodlands of southern Belgium. Pedobiologia 37: 365–387. Blake,S., G. N. Foster, M. D. Eyre,and M. L. Luff, 1994: Effects of habitat type and grassland management practices on the body size distribution of carabid beetles. Pedobiologia 38 (6): 502–512. Bot,A., and J. Benites, 2005: The importance of soil organic matter: Key to drought-resistant soil and sustained food production. FAO Soils Bulletin 80 - URL: http://www. fao.org/docrep/ 009/a0100e/a0100e0f.htm#bm15. Brose, U., 2003a: Bottom-up control of carabid beetle communities in early successional wetlands: mediated by vegetation structure or plant diversity? Oecologia 135: 407–413. Brose, U., 2003b: Regional diversity of temporary wetland Carabid beetle communities: a matter of landscape features or cultivation intensity?Agr. Ecosyst. Environ. 98: 163–167. Brühl, C.A., M. Mohamed,and K. E. Linsenmair, 1999:Altitudinal distribution of leaf litter ants along a transect in primary forest on Mount Kinabalu, Sabah, Malaysia. J. Trop. Ecol. 15: 265–277. Collingwood, C.A., 1979:The Formicidae (Hymenoptera) of Fennoscandia and Denmark. Fauna Entomologica Scandinavica 8: 9–175. |
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