Differentiation of Coleoptera (Carabidae & Tenebrionidae) communities in Mediterraneantype ecosystems from mountainous areas in the Peloponnese, Greece I. Anastasiou & A. Legakis Zoological Museum, Dept. of Biology, Univ. of Athens, Panepistimioupolis, Athens, Greece
Introduction Mountains of Greece well defined habitat types Phrygana Maquis Deciduous Forests Coniferous Forests Mixed Forests Alpine Vegetation Epigeic invertebrate fauna Palaeogeographic history of mountains Same differentiation
Area of Study The Peloponnese is a mountainous area in Southern Greece Invertebrate fauna: High diversity High levels of endemism 5 mountains were studied
Area of Study 2 different habitats were studied: Maquis and Phrygana Maquis Erymanthos Alt: 850 m Dominant Sp.: Quercus coccifera, Juniperus sp. Maquis Mainalo Alt: 700 m Dominant Sp.: Quercus coccifera, Juniperus sp. Maquis Taygetos Alt: 600 m Dominant Sp.: Quercus coccifera, Juniperus sp., Arbutus adrachne, A. unedo Phrygana Chelmos Alt: 750 m Dominant Sp.: Euphorbia sp., Sarcopoterium spinosum Callicotome villosa Maquis Chelmos Alt: 850 m Dominant Sp.: Quercus coccifera, Juniperus sp. Phrygana Taygetos Alt: 600 m Dominant Sp.: Euphorbia sp., Sarcopoterium spinosum Callicotome villosa Maquis Parnonas Alt: 600 m Dominant Sp.: Quercus coccifera, Juniperus sp.
The present study looks into the structure and differentiation of the communities of two of the major soil coleopteran families (Carabidae( & Tenebrionidae) ) in Mediterranean-type type ecosystems and assesses the effects of a number of environmental parameters
Methods In each mountain: Mediterranean habitat types Pitfall traps Carabidae & Tenebrionidae
Methods Sampling Period Sampling Frequency Sampling Method July 1995 June 1998 Seasonally Standard Pitfall Traps (plastic cups of 250ml containing about 50ml of ethylene glycol) 20 traps per biotope (distance between them 15m) laying on two transect lines (distance between lines 50m)
Methods Environmental parameters 1. Humidity Dry vs. fresh weight of soil 2. ph ph-meter 3. Soil organic content Combustion 4. Water capacity Water retainment in dry soil 5. Altitude Altimeter 6. Position on N-S axis 7. Position on W-E axis 8. Extent of open ground 9. Cover by shrubs 10. Cover by phrygana
PHRYGANA Biotopes MAQUIS Altitudinal Range: from 600 to 750 m. Coverage: Shrubs, Bushes, Soil, Rocks. Vegetation: Spartium junceum, Astragalus sp., Sarcopoterium spinosum, Asparagus acutifolius, Teucrium capitatum, Phlomis fruticosa, Callicotome villosa, Satureja sp., Cistus creticus, Euphorbia sp. Altitudinal Range: from 600 to 850 m. Coverage: Bushes, Shrubs, Soil, Rocks. Vegetation: Quercus coccifera, Brachypodium sp., Phillyrea latifolia, Cistus creticus, Dorycnium hirsutum, Hypericum empetrifolium, Cotinus coggygria, Arbutus adrachne, A. unedo, Teucrium sp.
Issues studied Presence of species Species diversity Community structure Relationships between environmental parameters and community structure
Presence of Species per Biotope Legend Widely Distributed Northern Maquis & Phrygana Southern Maquis & Phrygana Widely Distributed in Maquis Mountain Specific in Maquis Phrygana Chelmos Phrygana Taygetos
Carabidae (43 species) Number of species 20 18 16 14 12 10 8 6 4 2 0 16 14 13 19 11 9 9 Xma Ema Mma Pma Tma Xfr Tfr Xma Ema Mma Pma Tma Xfr Tfr Maquis: Number of species is higher on Chelmos, lower on Taygetos Phrygana: Number of species is higher on Taygetos
Shannon index H' (loge) Carabidae 3 2.5 2 1.5 1 0.5 0 2,15 2,56 2,01 1,99 1,60 1,08 1,17 Xma Ema Mma Pma Tma Xfr Tfr Xma Ema Mma Pma Tma Xfr Tfr Ev enness J' Diversity and evenness are high in phrygana Diversity and evenness are higher on Chelmos and Erymanthos 1 0.8 0.6 0.4 0.2 0 0,77 0,76 0,73 0,78 0,87 0,53 0,45 Xma Ema Mma Pma Tma Xfr Tfr Xma Ema Mma Pma Tma Xfr Tfr
Tenebrionidae (23 species) Number of species 16 14 12 10 8 6 4 2 0 14 5 5 5 5 6 4 Xma Ema Mma Pma Tma Xfr Tfr Xma Ema Mma Pma Tma Xfr Tfr Number of species is higher in phrygana
Shannon index H' (loge) Tenebrionidae 2.5 2 1.5 1 0.5 0 2,07 1,56 1,39 1,01 0,84 1,21 0,84 Xma Ema Mma Pma Tma Xfr Tfr Xma Ema Mma Pma Tma Xfr Tfr Evenness J' Diversity and evenness are higher on Parnonas and in the phrygana of Taygetos 1.2 1 0.8 0.6 0.4 0.2 0 0,97 0,75 0,78 0,78 0,63 0,60 0,52 Xma Ema Mma Pma Tma Xfr Tfr Xma Ema Mma Pma Tma Xfr Tfr
Differentiation between mountains Carabidae 2.2 2.0 1.8 Shannon index (log e) 1.6 1.4 1.2 1.0 0.8 0.6 0.4 Chelmos Erymanthos Mainalo Parnonas Taygetos ±1.96*Std. Err. ±1.00*Std. Err. Mean
Differentiation between mountains Tenebrionidae 2.2 1.8 Shannon index (log e) 1.4 1.0 0.6 0.2-0.2-0.6 Chelmos Erymanthos Mainalo Parnonas Taygetos ±1.96*Std. Err. ±1.00*Std. Err. Mean
Differentiation between habitats Carabidae 2.2 2.0 Shannon index (log e) 1.8 1.6 1.4 1.2 1.0 0.8 Maquis Phrygana ±1.96*Std. Err. ±1.00*Std. Err. Mean
Differentiation between habitats Tenebrionidae 1.8 1.6 1.4 Shannon index (log e) 1.2 1.0 0.8 0.6 0.4 0.2 Maquis Phrygana ±1.96*Std. Err. ±1.00*Std. Err. Mean
Differentiation between seasons Carabidae 2.0 1.8 Shannon index (log e) 1.6 1.4 1.2 1.0 0.8 0.6 Spring Summer Autumn ±1.96*Std. Err. ±1.00*Std. Err. Mean
Differentiation between seasons Tenebrionidae 2.0 1.8 Shannon index (log e) 1.6 1.4 1.2 1.0 0.8 0.6 Spring Summer Autumn ±1.96*Std. Err. ±1.00*Std. Err. Mean
Differentiation between mountains and seasons Carabidae & Tenebrionidae 2.0 1.5 Spring Summer Autumn F=3.58 p=0,008 Shannon index (log e) 1.0 0.5 0.0-0.5 Chelmos Erymanthos Mainalo Parnonas Taygetos
Differentiation between seasons and habitats Carabidae & Tenebrionidae 1.8 1.6 1.4 Maquis Phrygana F=4,99 p=0,003 Shannon index (log e) 1.2 1.0 0.8 0.6 0.4 0.2 0.0-0.2 Spring Summer Autumn
1 0.9 Community Structure Analysis of Similarities Ha-Mt 0.8 0.7 0.6 0.5 0.4 0.3 Mt Mt-Ha Mt-Se Ha Ha-Se Se Se-Mt Se-Ha 0.2 0.1 0 Mt Ha Se High differentiation between mountains Intermediate differentiation between habitats. However, communities in different habitats on the same mountain are highly differentiated Low seasonal differentiation
Community Structure 90 Analysis of Similarities 80 70 60 50 40 30 20 10 0 Chel-Erym Chel-M ain Chel-Parn Chel-Tayg Erym-Main Erym-Parn Erym-Tayg Main-Parn Main-Tayg Parn-Tayg Mts Mts across Habs Mts across Seas The differentiation between mountains is mainly due to the different structure of communities on Chelmos
14 12 10 8 6 4 2 0 Community Structure Analysis of Similarities Seas Seas across Mts Seas across Hab Sum-Aut Sum-Spr Aut-Spr The differentiation between seasons is mainly due to differences between summer and autumn and between spring and autumn
Environmental Parameters Xma Ema Mma Pma Tma Xfr Tfr Altitude (m) 850 850 700 600 600 750 600 North - South 38 02 44 38 00 51 37 36 26 37 21 25 37 10 01 38 06 76 36 58 26 East - West 22 07 50 21 55 21 22 21 41 22 39 25 22 10 46 22 16 88 22 25 15 Soil humidity (%) 13.42 18.26 15.66 12.96 10.61 5.03 4.58 ph 7.39 7.44 7.46 7.41 7.42 7.61 7.38 Water capacity (%) 64.14 62.72 63.80 73.45 62.20 53.19 58.72 Soil organic content (%) 14.83 13.88 11.49 18.93 12.12 5.66 9.56 Shrubs (%) 89.43 92.67 75.92 91.60 75.64 0.00 0.00 Phrygana (%) 0 0 0 0 0 77.2 75 Shadow (%) 85.68 86.87 61.37 66.26 74.38 0.00 0.00
Correlation of Decrease in Density Activity between Spring and Summer S with Environmental Parameters Parameter Altitude Humidity ph Water capacity Organic matter Shrub cover Spearman R -0.52-0.33 0.33-0.44-0.56-0.78 p-value 0.23 0.46 0.46 0.32 0.19 0.04
Canonical Correspondence Analysis of Relative Abundances vs. Environmental Parameters
Canonical Correspondence Analysis of Relative Abundances vs. Environmental Parameters Axes 1 2 3 4 Total inertia Eigenvalues:.626.543.462.413 2.784 Species-environment correlations:.992.992.983.946 Cumulative percentage variance of species data: 22.5 42.0 58.6 73.4 of species-environment relation: 27.5 51.3 71.5 89.7 Sum of all unconstrained eigenvalues 2.784 Sum of all canonical eigenvalues 2.280
Conclusions 1 Diversity of Carabidae is higher in the northern mountains and in phrygana Number of species of Tenebrionidae is higher in phrygana and especially for Taygetos Diversity of Tenebrionidae does not show significant differences between maquis and phrygana Diversity of Tenebrionidae is higher for the southern mountains Diversity in phrygana is markedly higher. This is more evident in Carabidae Although the total diversity does not seem to differ significantly among mountains, in reality there is a marked difference in seasonal variation in each mountain The biggest difference in diversity between maquis and phrygana was observed during spring
Conclusions 2 The analysis of the structure the communities revealed a differentiation between stations of the different mountains, especially following a N-S gradient Differences in structure were also observed between the habitat types Community structure changes gradually between spring and summer and abruptly between summer and autumn Vegetation cover and the plant species composition seem to affect the distribution and abundance of the species The highest values of density/activity were obtained during spring During spring the most important environmental parameters affecting the structure of the community seem to be humidity (which is strongly correlated to shadow), the N-S gradient, and the presence of phrygana The increase of temperature during summer was compensated by movement to the shadow under the shrubs. This seems to be the reason for an increased summer activity in several stations
Acknowledgements We wish to thank the following colleagues for their assistance during the project: Christos GEORGIADIS Costantinos SOTIROPOULOS