University of Athens University of Athens (CARABIDAE, TENEBRIONIDAE) (CARABIDAE, (CARABIDAE, TENEBRIONIDAE) (CARABIDAE, Univ. of Athens, GR-15784 Greece

Kapodistrian Kapodistrian

National & National & International Congress on the Zoogeography and Ecology of Greece and Adjacent Regions, RELATIONSHIPS BETWEEN THE DIVERSITY OF RELATIONSHIPS BETWEEN THE DIVERSITY OF th EPIGEAN COLEOPTERA EPIGEAN COLEOPTERA Thessaloniki, Greece, 22-25/5/2002 9

Ioannis ANASTASIOU, Anna PAPADOPOULOU & Anastasios LEGAKIS Zoological Museum, Dept. of Biology, AND HABITAT CHARACTERISTICS ON MT. TAYGETOS AND HABITAT CHARACTERISTICS ON MT. TAYGETOS IntroductionIntroduction • Phrygana • Maquis • Deciduous Forests • Mountains of Greece Æwell defined habitat types • Coniferous Forests • Mixed Forests • Alpine Vegetation

Palaeogeographic history • Epigeic invertebrate fauna Same Differentiation of Mountains

• Mt. Taygetos is the southernmost mountain of continental Greece with a large number of endemic taxa BUT it’s invertebrate fauna has not been studied sufficiently.

The present study looks into the structure of the communities of the two major soil coleopteran families (Carabidae, Tenebrionidae) and assesses the effects of a number of environmental parameters. MethodsMethods

Sampling Period March – November 1997

Sampling Frequency Seasonally

Sampling Method • Standard Pitfall Traps (plastic cups of 250ml containing about 50ml of ethylene glycol) • 30 traps per biotope (distance between them Æ 15m) laying on two transect lines (distance between lines Æ 50m) StudyStudy AreaArea

DF 1 • •AL 2 (Deciduous Forest) (Sub – Alpine)

MA 1 • •MF 1 (Maquis) (Mixed Coniferous Forest)

PF 2 • •FR 1 (Pinus Forest) (Phrygana)

PF 1 • •AL 1 (Pinus Forest) (Sub – Alpine) BiotopesBiotopes AL 1 AL 2

• Altitudinal Range: ≅1650 m. • Coverage: Rocks, Bushes, Shrubs, Bare Soil. • Vegetation: Juniperus communis , Astragalus sp., Poa sp., Daphne oleoides, Abies cephalonica, Ranunculus sp., Compositae, Gramineae.

• Altitudinal Range: ≅1800 m. • Coverage: Rocks, Bushes, Shrubs, Bare Soil. • Vegetation: Juniperus communis , Astragalus sp., Calycotome villosa , Echinops sp., Euphorbia sp., Daphne oleoides, Boraginaceae, Gramineae. BiotopesBiotopes

PF 1 PF 2

• Altitudinal Range: ≅1350 m. • Altitudinal Range: ≅1400 m. • Coverage: Trees, Leaf Litter, Shrubs. • Coverage: Trees, Leaf Litter, Shrubs. • Vegetation: Pinus nigra , Pteridium • Vegetation: Pinus nigra , Pteridium aquilinum. aquilinum. BiotopesBiotopes MF 1 DF 1

• Altitudinal Range: ≅1500 m. • Altitudinal Range: ≅1000 m. • Coverage: Trees, Leaf Litter, Shrubs. • Coverage: Trees, Leaf Litter, • Vegetation: Pinus nigra , Abies cephalonica , Shrubs. Pteridium aquilinum , Fragaria sp., Gramineae. • Vegetation: Quercus frainetto. BiotopesBiotopes FR 1 MA 1

• Altitudinal Range: ≅640 m. • Altitudinal Range: ≅800 m. • Coverage: Shrubs, Bushes, Soil, • Coverage: Bushes, Shrubs, Soil, Rocks. Rocks. • Vegetation: Spartium junceum , • Vegetation: Quercus coccifera, Astragalus sp., Sarcopoterium Brachypodium sp., Phillyrea latifolia , spinosum, Asparangus acutifolius , Cistus creticus , Dorycnium hirsutum , Teucrium capitatum , Phlomis fruticosa , Hypericum empetrifolium , Cotinus Calycotome villosa , Saturejia sp., Cistus coggygria, Arbutus adrachne, A. unedo, creticus, Euphorbia sp. Teucrium sp. ResultsResults

Number of Species

carabidae tenebrionidae

50 40 30 20 10 0 al1 al2 fr1 ma1 df1 mf1 pf1 pf2 AbundanceAbundance

Carabidae: Number of Individuals

pf1 pf2 al1 2% 9% 16% mf1 26%

al2 df1 41% 1% fr1 ma1 4% Tenebrionidae: Number of Individuals 1% pf1 df1 mf1 0,1% ma1 1,0%0,7% pf2 4,2% 0,2% fr1 20,3% al1 46,4%

al2 27,1% DistributionDistribution ofof SpeciesSpecies

Carabidae: Number of Species

ma1 df1 mf1 pf1 7% 6% 9% fr1 9% 15% pf2 13%

al2 al1 19% 22% Tenebrionidae: Number of Species

mf1 pf1 pf2 df1 al1 2% 4% 2% 11% 21% ma1 9%

al2 fr1 26% 25% al1 al2 fr1 ma1 df1 mf1 pf1 pf2 Calathus corax Platyderus graecus PresencePresence ofof SpeciesSpecies Molops spartanus Carabus preslii Microlestes luctuosus Cymindis lineata perper BiotopeBiotope Cymindis sinuata Carabus convexus Zabrus validus Tapinopterus rebellis Leistus parvicollis Harpalus sulphuripes Leistus spinibarbis Laemostenus peloponnesiacus Legend Nebria brevicollis Amara eurynota Ophonus (Metophonus) sp. Harpalus rufipalpis Widely Distributed Licinus oertzeni Cymindis miliaris Philorhizus crucifer Zabrus robustus Mainly in Open Biotopes Lebia trimaculata Cymindis axillaris Notiophilus interstitialis Zabrus aetolus hellenicus Sub-Alpine Acinopus baudii Ophonus cribricollis Masoreus wetterhallii Trechus austriacus Open Biotopes Dixus obscurus Ophonus subquadratus Harpalus attenuatus Leistus magnicollis Phrygana Aptinus lugubris Leistus rufomarginatus Calosoma sycophanta Carabus merlini Mainly in Forests Notiophilus rufipes Zabrus graecus Calosoma inquisitor Syntomus obscuroguttatus Pinus Forest Ophonus taygetanus Ophonus krueperi Tapinopterus duponcheli Laemostenus cimmerius Mixed Coniferous Forest Pachycarus cyaneus Calathus cinctus Myas chalybaeus Carabus coriaceus Various Harpalus honestus Harpalus Characteristic Carabidae Species al Calathus corax 34.1 Cymindis sinnuata 26.4 al2 Calathus corax 84.8 df Myas chalybaeus 45.5 Notiophilus rufipes 31.7 fr Calathus cinctus 24.6 Acinopus baudi hellenicus 21.1 Cymindis sinnuata 18.0 ma Carabus preslii 78.6 pf Leistus magnicollis 45.9 Average contribution of Calathus corax 38.2 each species to the pf2 Calathus corax 24.3 overall similarity of the Platyderus graecus 20.4 sample Aptinus lugubris 19.7 mf Aptinus lugubris 75.7 Characteristic Tenebrionidae Species

al Idastraniella taygetanus 49.1 al2 Calyptopsis caraboides 38.4 Idastraniella taygetanus 20.3 df Pedinus helopioides 100 fr Dailognatha quadricollis 30.7 Graecopachys quadricollis 26.9 Average contribution of each species to the ma Dendarus plicatulus 71.8 overall similarity of the pf - sample pf2 Laena sp. B 100 mf Laena sp. B 100 CarabidaeCarabidae diversitydiversity (pooled(pooled samples)samples) TenebrionidaeTenebrionidae diversitydiversity (pooled(pooled samples)samples) 6 sprin g -autumn pattern spring pattern 1,8 Carabus preslii (al1) Platyderus graecus (pf1) 5 1,6

1,4 4 1,2 1 3 0,8 2 0,6

0,4 1

0,2 0 0 spring summer autumn spring summer autumn

autumn pattern 12 summer pattern Calathus corax (al1) 1,2 Myas chalybaeus (df1)

10 1

8 0,8

6 0,6

4 0,4

2 spring summer autumn 0,2 spring summer autumn

0 0 patterns of phenology fam. Carabidae

100%

80% autumn 60% summer 40% spring 20% spring-autumn

0% al1al1 al2 fr1 fr1 ma1 df1df1 mf1 pf1 pf1 pf2 patterns of phenology fam. Tenebrionidae

100% 80% 60% autumn 40% summer spring 20% 0% al1 al2 fr1 ma1 df1 CarabidaeCarabidae (pooled(pooled samplessamples perper biotope)biotope) TenebrionidaeTenebrionidae (pooled(pooled samplessamples perper biotope)biotope) Acb Pac

+3.5 Maw

fr1

Mil

Cac Opc Zav Crc

shr Cls Lem ma1 Tra Crm Ler Tad lit Crp Plg Apl pf1 mf1 pf2 str Nor al2 Crv Cyl Lac Mos Cys Zaa s-r Lep Tar Cax Cya df1 al1 Has Les Myc Hah Zar alt Lap

Cym -2.0 -2.5 +2.5 Axes 1 2 3 4 Total inertia

Eigenvalues: .627 .341 .302 .144 1.800 Species-environment correlations: .997 .991 .996 .996 Cumulative percentage variance of species data: 34.8 53.8 70.6 78.6 of species-environment relation: 40.7 62.9 82.5 91.9

Sum of all unconstrained eigenvalues 1.800 Sum of all canonical eigenvalues 1.540 Summary of Monte Carlo test

Test of significance of first canonical axis: eigenvalue = .627 F-ratio = 1.070 P-value = .0110

Test of significance of all canonical axes : Trace = 1.540 F-ratio = 2.368 P-value = .0005 ConclusionsConclusions • Carabidae have a higher number of species than Tenebrionidae • Tenebrionidae are especially low in number of species in coniferous forests • Open habitats have more species of Carabidae and Tenebrionidae than the other biotopes

• Highest abundance of Carabidae was recorded in sub-alpine and mixed forest. The latter had a lower number of species

• Highest number of Tenebrionidae was recorded in sub-alpine and phrygana • In Tenebrionidae diversity was higher in open habitats; a pattern that was not obvious in Carabidae • Each habitat type seems to have a different composition of phenology patterns • Clustering of Carabidae samples indicates a major division between open and closed habitats • Clustering of Tenebrionidae indicates a major division between forest and non-forest habitats • The most important parameters that affect the Carabidae communities seem to be coverage of soil by pine litter, altitude and coverage of soil by other types of litter ACKNOWLEDGEMENTS

We wish to thank the following colleagues for their assistance during the project:

Kanella RADEA Eleni GOURGOU Dimitris TSAPARIS Christos GEORGIADIS