Spatialvariabilityofbee  co mmunities:fromlocalassemblages  tointeractionnetworks 

 ‡Ö”‹ƒ’”‡•‡–ƒ†ƒ’‡”ǣ

 AnnaTornéNoguera

 ‡”‘’–ƒ”ƒŽ‰”ƒ—†‡‘ –‘”ƒ



 „‡Ž˜‹•–Ǧ‹Ǧ’Žƒ—†‡Ž•†‹”‡ –‘”•ǣ  Dr.JordiBoschGras Dr.AnselmRodrigoDomínguez   Ǧ  





”‘‰”ƒƒ†‡‘ –‘”ƒ–‡ ‘Ž‘‰‹ƒ‡””‡•–”‡  Ȃ‹˜‡”•‹–ƒ–—–֐‘ƒ†‡ƒ” ‡Ž‘ƒ ƒ‹‰ʹͲͳͷ 

































‘”±‘‰—‡”ƒǡƒȋʹͲͳͷȌ  Spatialvariabilityofbeecommunities:fromlocalassemblagestointeractionnetworks  ‡›™‘”†•ǣ Antagonistic networks, bee community, beta diversity, body size, cavityǦnesting bees and wasps, exploitative competition, floral and nesting resources, local spatial variability, pollinator community,speciesturnover  “—‡•–ƒ–‡•‹Šƒ‡•–ƒ–ˆ‹ƒ­ƒ†ƒ‹–Œƒ­ƒ–Žƒ„‡ ƒ’”‡†‘ –‘”ƒŽ†‡FormacióndePersonalInvestigador ȋ  ȌǦʹͲͳͲǦͲͶʹͷʹͲ‹Žƒ„‡ ƒǦ Ǧͳ͵ǦͲ͹Ͳ͵ͺ†‡ŽMinisteriodeCienciaeInnovaciónƒ–”ƒ˜±•†‡Ž ’”‘Œ‡ –‡   ʹͲͲͻǦͳʹ͸Ͷ͸ǡ ’‡Ž ’”‘Œ‡ –‡ ‘•‘Ž‹†‡”Ǧ‘–‡• ʹͲͲͺǦͲͲͲͶͲ †‡Ž MinisteriodeEducaciónyCiencia‹’‡”ˆ‘•’”‘˜‹‡–•†‡Žƒ‹’—–ƒ ‹×†‡ƒ” ‡Ž‘ƒƒ–”ƒ˜±•†‡Žƒ”  †‡Ž ƒ””ƒˆǤ 

 













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‡”›ƒ˜‹†Š‘”‡ƒ—









Agraïments/Acknowledgements

””‹„ƒ–•ƒƒ“—‡•–’—–ǡǯƒ‰”ƒ†ƒ”‹ƒ†‘ƒ”Ž‡•‰” ‹‡•ƒ–‘–Š‘“—‹ǡ†ǯ—ƒƒ‡”ƒ‘ƒŽ–”ƒǡŠƒ ˆ‘”ƒ–’ƒ”–†ǯƒ“—‡•–ƒ–‡•‹‹Šƒˆ‡–’‘••‹„Ž‡“—‡ƒ‹šÖ–‹”±•‡†ƒ˜ƒ–Ǥ˜—‹ƒ“—‡•–ƒ–‡•‹±•—ƒ ”‡ƒŽ‹–ƒ–‰” ‹‡•ƒ–‘–•‡ŽŽ•Ǥ

”‹‡”†‡–‘–ǡǯƒ‰”ƒ†ƒ”‹ƒƒ‰”ƒ‹”‘Ž–À••‹ƒŽ•‡—•†‹”‡ –‘”•ǡ ‡Ž ‘”†‹ ‹ Žǯ•‡Žǡ ‡Ž ˆ‡– †ǯŠƒ˜‡”Ǧ‡†‘ƒ–Žǯ‘’‘”–—‹–ƒ–†‡ˆ‡”—ƒ–‡•‹†‘ –‘”ƒŽǤ‘Ž–‡•‰” ‹‡•’‡”ƒˆ‡”Ǧ‡ ”±‹š‡” †—”ƒ––‘–•ƒ“—‡•–•ƒ›• ‘ƒ ‹‡–Àˆ‹ ƒ‹ ‘ƒ’‡”•‘ƒǤ‘–‡Ž“—‡ǯŠ‡—‡•‡›ƒ–‹Š‡ ƒ’”°•ǡƒ ƒ†°‹ ƒ‡–‹’‡”•‘ƒŽǡ–±—˜ƒŽ‘”‹ˆ‹‹–Ǥ ƒ•‡–—ƒ‡–ƒ’ƒ‘Ž–‹–‡•ƒ‡Žƒ“—ƒŽ ǯŠ‡—†‘ƒ–‘Ž–À••‹ǡ‹—‡•–‹ •‹ ‡”ƒ‡–‘Ž–ƒ‰”ƒÃ†ƒǤ‡ƒŽ‡–ƒ“—‡•–ƒ–‡•‹‘Šƒ—”‹ƒ •‡–’‘••‹„Ž‡•‡•‡˜‘•ƒŽ–”‡•Ǥ

ƒ„±˜—ŽŽƒ‰”ƒ‹”‘Ž–‡•’‡ ‹ƒŽ‡–ƒŽǯ ‡Ž‡ƒ–‘–ƒŽƒˆ‡‹ƒǡ‹’Ž‹ ƒ ‹×ǡ ƒŒ—†ƒǡ ”‹—”‡• ‹ ƒ‹•–ƒ– ƒŽ ŽŽƒ”‰ †‡ –‘–ƒ Žƒ –‡•‹Ǥ ƒ •‡– — ’Žƒ‡” ‘’ƒ”–‹” –ƒ–À••‹‡• Š‘”‡• †‡ ƒ’ ‹ Žƒ„‘”ƒ–‘”‹ƒ„–—ǡ‹Ž‹—ƒ„ƒ–ƒŽŽ‡•ǡ‹•‡•‡’‡”†”‡ƒ‹‡Ž„‘Š—‘”Ǥ ƒ••‡–—ƒ’‡­ƒ ‹’”‡• ‹†‹„Ž‡‡ƒ“—‡•–ƒ–‡•‹Ǥ

‘Ž–‡•‰” ‹‡•–ƒ„±ƒŽƒ˜‹ǡ’‡”Žƒ•‡˜ƒ ‘–”‹„— ‹×‹ƒ’‘”–ƒ ‹‘•‡ƒ“—‡•–ƒ–‡•‹ǡ‹ƒŽ‡”‰‹‘ǡ ’‡”–‘–ƒŽǯƒŒ—†ƒƒ ƒ’‹ƒŽŽƒ„‘”ƒ–‘”‹ǡ’‡”Žƒ•‡˜ƒ’ƒ ‹° ‹ƒ‡•‡›ƒ–Ǧ‡–ƒš‘‘‹ƒ‹’‡”Žƒ •‡˜ƒƒ‹•–ƒ–‹ ‘’ƒ›‘‹ƒ†—”ƒ––‘–ƒ“—‡•–’‡”À‘†‡Ǥ

–‘–•‡Ž•‡•–—†‹ƒ–•“—‡ǡ†ǯƒŽ‰—ƒƒ‡”ƒ‘ƒŽ–”ƒǡŠ‡— ‘–”‹„—Ö‡Žƒ”‡ƒŽ‹–œƒ ‹×†ǯƒ“—‡•–ƒ –‡•‹ǣƒ”ƒǡ”‹ǡ‡” ‡ǡƒ”–ƒǡƒ”–ƒǡ ƒǡ ‘•‡’ǡ ”‡‡ǡ ”‡‡ǡ‘„‡”–‘ǡƒ” ‘ǡƒ—”ƒǡ ƒ˜‹†ǡ–‘‹‘ǡŽ„‡”–ǡ‡—•ǡ ‘ƒ‘Žǡƒǡƒ”‹ƒŽ„ƒǡ±‘ǡŽ„ƒǡ‘Žǡ •ƒ„‡ŽǤǤǤƒ–‘–•‡ŽŽ•ǡ ‘Ž–‡•‰” ‹‡•Ǥ

‘Ž–‡•‰” ‹‡•–ƒ„±ƒ–‘–ƒŽƒ‰‡–†‡Ž”‡ƒˆǣ„‡ ƒ”‹•ǡ–° ‹ •ǡ ‹˜‡•–‹‰ƒ†‘”•ǡ ’‡”•‘ƒŽ †‡ •—’‘”–ǤǤǤ ’‡” ˆ‡” †‡Ž ŽŽ‘  †‡ ˆ‡‹ƒ — ŽŽ‘  –ƒ ƒ‰”ƒ†ƒ„Ž‡ǡ ‘ ‘’ƒ”–‹” ƒŽ‡‰”‹‡• ‹ †‡•‡•’‡”ƒ ‹‘•ǡ ‘ ƒ‹ •‡ —• ƒ ƒ„ƒ Žǯ‡‡”‰‹ƒǤ  ‘Ž– ‡•’‡ ‹ƒŽ‡– ƒŽ• ‘’ƒ›• †‡Ž ‡— ƒ–‹ †‡•’ƒ–š‰”ƒǡ’Ž‡†ǯ‹˜‡–•‹–”ƒ•–‘•‹˜‡”•‡„Žƒ–•ȋ’”‘–‘ƒ ƒ„ƒ”ž‡Ž•—ˆ”‹‹‡–‘ǨȌǡ‹ †‡Ž‡—‘—†‡•’ƒ–š’‡–‹–ǡ’Ž‡†‡ ‘†‹•‹‘Ž–•”‹—”‡•Ǥ

”‡ƒŽŽ›™ƒ––‘–Šƒ”‘ˆǤŽò–Š‰‡ˆ‘”Š‘•–‹‰‡‹–Š‡Ǧƒ”•–ƒ†–Ǥ –™ƒ•ƒƒƒœ‹‰ ‡š’‡”‹‡ ‡ –‘ „‡ –Š‡”‡Ǥ ƒ› –Šƒ• –‘ ƒŽŽ › ‘ŽŽ‡ƒ‰—‡• ˆ”‘ –Š‡  ‘Ǧ‡–™‘”• ‰”‘—’ǡ ‡•’‡ ‹ƒŽŽ›Š”‹•–‹ƒˆ‘”Š‡”ˆ”‹‡†•Š‹’ǡˆ‘”„‡‹‰–Š‡”‡ǡˆ‘”ƒŽŽ –Š‡ •Šƒ”‡† ‘‡–• ‹ –Š‡  v 

‘ˆˆ‹ ‡ƒ†‡•’‡ ‹ƒŽŽ›‘—–‘ˆ‹–Ǣƒ†ƒŽ•‘‡˜‹ǡ†”‹ƒƒ† —Ž‡ˆ‘”ƒ‹‰›•–ƒ›•‘‹ ‡ƒ† Š‡Ž’‹‰‡™‹–Š‡˜‡”›–Š‹‰ ‡‡†‡†ǤŠƒ•–‘› Ž‹„‹‰’ƒ”–‡”•ƒ†ˆ”‹‡†•ǡ‡•’‡ ‹ƒŽŽ› ƒ”–‹ƒ†•‡ Š‘ǡˆ‘”–Š‡‰”‡ƒ––‹‡•‹ƒ†‘—––Š‡Ž‡––‡”œ‡–”—Ǥ‘›„‡•–”‘‘‹‡•ǣ †”‹ƒǡ‘”ƒǡ ”‡†ƒ†ƒ”–‹ƒǡ–Šƒ•ˆ‘”ƒ‹‰›Ž‹ˆ‡•‘‡ƒ•› ƒ† ˆ—Ǩ Ž•‘ǡ ƒ› –Šƒ•–‘–Š‡ ‘— Š‹‡•ˆ‘”–Š‡‰”‡ƒ–•—’‡”Ǧ‹–‡”ƒ–‹‘ƒŽ‡˜‡‹‰•ǡƒ†ǡ‘ˆ ‘—”•‡ǡ–‘–Š‡ —„ ”‡™ˆ‘”ƒŽŽ–Š‡‰”‡ƒ–Š‹Žƒ”‹‘—•„‹œƒ””‡‹‰Š–•Ǥ

ƒ„±ǯƒ‰”ƒ†ƒ”‹ƒƒ‰”ƒ‹”‡Ž•—’‘”–‹ƒ‹•–ƒ–†‡–‘–ƒŽƒ‰‡–†‡ƒŽǯ”‘ŽƒǣƒŽ‡–Àǡ”‹ǡ”‹ ǡ—‹ǡŽ•ƒ‹‡˜‹Ǥ ” ‹‡•’‡”–‘–•‡Ž•‘‡–• ‘’ƒ”–‹–•‹‹ˆ‹‹–ƒ–†ǯŠ‹•–Ö”‹‡••—””‡ƒŽ‹•–‡• ˜‹• —†‡•‡—’‹•‘Žƒ‰”ƒ˜‡–ƒ–˜ƒ ƒ’ƒŽǯ‡•“—‡””‡Ǥ‹—”‡ȋͻƒ›•ǨȌƒ„˜‘•ƒŽ–”‡•Šƒ•‡– ‹’ƒ‰ƒ„Ž‡Ǩ ” ‹‡•–ƒ„±ƒŽƒ‰‡–†‡ƒ‹š‘ǣƒ—ǡ‘‹ǡƒ–Ž‹ƒǡƒ†ƒǡƒ‡Ž‹ †”ƒ ’‡” ‘’ƒ”–‹”Žƒ†ƒ””‡”ƒ‡–ƒ’ƒ†ǯƒ“—‡•–ƒ–‡•‹ƒ„‹Ǥ ƒ–‘–•‡Ž•ƒ‹ •‹ƒ‹‰—‡•ǡ ‘’ƒ›• ‹ ‘’ƒ›‡•“—‡‡•–‹‘‹“—‡Š‡—‹’”°•‡‹—ƒ’ƒ”–†‡˜‘•ƒŽ–”‡•ǡ “—‡ ǯŠ‡— †‘ƒ– •—’‘”–‹ƒŽ‡‰”‹ƒǡ ‡”˜‡•‡•‹‡š —”•‹‘•ǡƒˆ‡ –‡‹ƒ‘”Ǥ–‘–•˜‘•ƒŽ–”‡•ǡ‘Ž–‡•‰” ‹‡•Ǥ

‹ƒŽ‡– ǯƒ‰”ƒ†ƒ”‹ƒ ƒ‰”ƒ‹”‘Ž–•‹ ‡”ƒ‡– ƒŽƒ‡˜ƒˆƒÀŽ‹ƒ–‘–‡Ž•—’‘”–“—‡ǯŠƒ †‘ƒ–ǡ‘Ž–‡•’‡ ‹ƒŽ‡–ƒŽ•‡—•’ƒ”‡•ǡ’‡”’‘•ƒ”ǦǯŠ‘ˆ ‹Ž“—ƒ±•†‹ˆÀ ‹Ž‡”ƒǤŽƒ‡˜ƒ ‰‡”ƒƒǡ’‡”‡• ‘Ž–ƒ”Ǧ‡‹‡–‡†”‡ǯǡƒŽƒ‡˜ƒ˜‹ƒǡ’‡”‡• ‘Ž–ƒ”Ǧ‡–‘–‹‘‡–‡†”‡ǯǡ‹ ƒŽƒ’‡–‹–ƒ„”‹Žǡ’‡”†‘ƒ”Ǧ‡–ƒ–ƒƒŽ‡‰”‹ƒ‹‡‡”‰‹ƒ•‡•‡•ƒ„‡”ǦŠ‘Ǥ







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Tableofcontents

AGRAÏMENTS/ACKNOWLEDGEMENTS...... V

TABLEOFCONTENTS...... VII

ABSTRACT...... 1

RESUM...... 3

GENERALINTRODUCTION...... 5

       ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤ͹

     ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤͺ

       ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤͻ

   ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤͳͳ

      ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤͳʹ

CHAPTER1.DETERMINANTSOFSPATIALDISTRIBUTIONINABEECOMMUNITY:NESTING

RESOURCES,FLOWERRESOURCES,ANDBODYSIZE...... 23

ͳǤͳǤ  ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤʹͷ

ͳǤʹǤ   ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤʹͺ

1.2.1. Studyarea...... 28

1.2.2. Beesampling...... 29

1.2.3. Flowerresources...... 29

1.2.4. Nestingsubstrates...... 30

1.2.5. Statisticalanalysis...... 30

ͳǤ͵Ǥ ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤ͵ʹ

1.3.1. Beecommunity...... 32

1.3.2. Spatialdistributionofflowerandnestingresources...... 32

1.3.3. Beespatialdistribution...... 33

1.3.4. Relationshipbetweenresourcesandbeespatialdistribution...... 36

 vii 

ͳǤͶǤ   ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤ͵ͻ

CHAPTER2.COLLATERALEFFECTSOFBEEKEEPING:IMPACTSONPOLLENǦNECTAR

RESOURCESANDWILDBEECOMMUNITIES...... 51

ʹǤͳǤ  ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤͷ͵

ʹǤʹǤ   ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤͷ͸

2.2.1. Studyarea...... 56

2.2.2. Flowerresourcesurveys...... 57

2.2.3. Pollinatorvisitationrates...... 58

2.2.4. Beecommunity...... 59

2.2.5. Flowerabundance...... 59

2.2.6. Statisticalanalysis...... 60

ʹǤ͵Ǥ ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤ͸ʹ

2.3.1. Pollenandnectarconsumption...... 62

2.3.2. Beecommunity...... 64

ʹǤͶǤ   ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤ͸ͷ

CHAPTER3.WITHINHABITATEǦDIVERSITYINAHOSTǦPARASITOIDINTERACTION

NETWORK...... 75

͵ǤͳǤ  ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤ͹͹

͵ǤʹǤ   ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤͺͲ

3.2.1. Studyarea...... 80

3.2.2. Surveys...... 80

3.2.3. Statisticalanalysis...... 81

͵Ǥ͵Ǥ ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤͺ͵

3.3.1. Hostcommunity...... 84

3.3.2. Parasitoidcommunity...... 84

3.3.3. Interactions...... 86

 viii 

3.3.4. RelationshipbetweenspeciesandinteractionEǦdiversity...... 87

3.3.5. DistancedecayofEǦdiversity...... 87

͵ǤͶǤ   ǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤǤͺͺ

GENERALCONCLUSIONS...... 97

APPENDIXA...... 101

APPENDIXB...... 111





 ix



Abstract

”‰ƒ‹•• ƒ”‡ Š‡–‡”‘‰‡‡‘—•Ž› †‹•–”‹„—–‡† ˆ‘ŽŽ‘™‹‰ •’ƒ–‹ƒŽ ’ƒ––‡”•ǡ™Š‹ Šƒ”‡–Š‡ ‘•‡“—‡ ‡ ‘ˆ ƒ› „‹‘–‹  ƒ† ƒ„‹‘–‹  ˆƒ –‘”•Ǥ ‡ ‘ˆ –Š‡ ƒ‹ ‰‘ƒŽ• ‹ ‡ ‘Ž‘‰› ‹• –‘ —†‡”•–ƒ† Š‘™ •’‡ ‹‡• ‹–‡”ƒ –‹‘• ™‹–Š ‘–Š‡” ‘”‰ƒ‹•• ƒ† ™‹–Š –Š‡ ‡˜‹”‘‡– †‡–‡”‹‡–Š‡•‡‘„•‡”˜‡†•’‡ ‹‡•†‹•–”‹„—–‹‘’ƒ––‡”•Ǥ –Š‹•–Š‡•‹•ǡ™‡ƒƒŽ›œ‡–Š‡ˆƒ –‘”• ‡š’Žƒ‹‹‰ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ„‹Ž‹–› ‘ˆ „‡‡ ‘—‹–‹‡• ƒ† ƒ••‘ ‹ƒ–‡† ‹–‡”ƒ –‹‘ ‡–™‘”• ƒ– Ž‘ ƒŽ • ƒŽ‡Ǥ  Š‡ •–—†› ™ƒ• ‘†— –‡† ‹ –Š‡ ƒ””ƒˆ ƒ–—”ƒŽ ƒ”ǡ ‹ ƒ ͵ʹʹƒ”‡ƒ Š‘‘‰‡‡‘—•Ž› ‘ —’‹‡† „› ‡†‹–‡””ƒ‡ƒ • ”—„Žƒ†ǡ ƒ† Žƒ ‹‰ •–”‘‰ ‰”ƒ†‹‡–• ‘” ‡ ‘Ž‘‰‹ ƒŽ „ƒ””‹‡”•Ǥ ‡ ƒ—•‡ „‡‡• ”‡Ž› ‘ ˆŽ‘”ƒŽ ƒ† ‡•–‹‰ ”‡•‘—” ‡• –‘ •—”˜‹˜‡ ƒ† ”‡’”‘†— ‡ǡ–Š‡•‡ƒ”‡‡š’‡ –‡†–‘‹ˆŽ—‡ ‡–Š‡Ž‘ ƒŽ•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ–Š‡„‡‡ ‘—‹–›Ǥ –Š‡ˆ‹”•– Šƒ’–‡”ǡ™‡‡š’Ž‘”‡Š‘™–Š‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆˆŽ‘™‡”ƒ†‡•–‹‰”‡•‘—” ‡• †‡–‡”‹‡•Ž‘ ƒŽ•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‹–Š‡„‡‡ ‘—‹–›Ǥ‡ ˆ‘—†ƒ Ž‡ƒ”‰‡‘‰”ƒ’Š‹ ƒŽ ’ƒ––‡”‘ˆ•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‹„‡‡ ‘—‹–› ‘’‘•‹–‹‘ǤŠ‹•’ƒ––‡”™ƒ•’ƒ”–Ž›†”‹˜‡ „›ˆŽ‘”ƒŽ”‡•‘—” ‡•™‹–Šƒ‡‰Ž‹‰‹„Ž‡ ‘–”‹„—–‹‘‘ˆ‡•–‹‰•—„•–”ƒ–‡ƒ˜ƒ‹Žƒ„‹Ž‹–›ǡƒ†™ƒ• •–”‘‰Ž›”‡Žƒ–‡†–‘„‘†›•‹œ‡ǣ•ƒŽŽ„‡‡•’‡ ‹‡•ȋδͷͷ‰Ȍ†‹•’Žƒ›‡†•–”‘‰•’ƒ–‹ƒŽ’ƒ––‡”•ǡ ™Š‹Ž‡ Žƒ”‰‡ •’‡ ‹‡• ȋε͹Ͳ‰Ȍ –‡†‡† –‘ „‡ ‘”‡ ‡˜‡Ž› †‹•–”‹„—–‡†ƒ ”‘••–Š‡ƒ”‡ƒǤ‡‡ •’ƒ–‹ƒŽ†‹•–”‹„—–‹‘ƒ›ƒŽ•‘„‡ƒˆˆ‡ –‡†„›’‡”–—”„ƒ–‹‘”‡‰‹‡•ǤŠ‡•–—†›ƒ”‡ƒ ‘–ƒ‹• •‡˜‡”ƒŽƒ’‹ƒ”‹‡•ǡƒˆˆ‘”†‹‰ƒ‘’’‘”–—‹–›–‘‡š’Ž‘”‡–Š‡’‘–‡–‹ƒŽ‡ˆˆ‡ –•‘ˆ„‡‡‡‡’‹‰‘ ™‹Ž†„‡‡ ‘—‹–‹‡•ȋŠƒ’–‡”ʹȌǤ ‘‡›„‡‡•ȋApismelliferaȌƒ”‡Š‹‰ŠŽ›‡ˆˆ‹ ‹‡–ˆ‘”ƒ‰‡”• ƒ†Šƒ˜‡–Š‡ƒ„‹Ž‹–›–‘”‡ ”—‹–‘–Š‡”ˆ‘”ƒ‰‡”•–‘–Š‡‘•–”‡™ƒ”†‹‰ ˆŽ‘™‡” ’ƒ– Š‡•ǡ ’‘–‡–‹ƒŽŽ›‡•–ƒ„Ž‹•Š‹‰ƒ ‘’‡–‹–‹˜‡‡ˆˆ‡ –‘‘–Š‡”’‘ŽŽ‹ƒ–‘”•Ǥ‡ˆ‘—†–Šƒ–Š‘‡›„‡‡• ™‡”‡–Š‡ƒ‹ ‘–”‹„—–‘”•–‘’‘ŽŽ‡Ȁ‡ –ƒ”†‡’Ž‡–‹‘‘ˆ–Š‡–™‘ƒ‹ˆŽ‘™‡”‹‰’Žƒ–•‹ –Š‡•–—†›ƒ”‡ƒǤ‡ƒŽ•‘ˆ‘—†–Šƒ––Š‡„‡‡ ‘—‹–›™ƒ•‘†‹ˆ‹‡† ‹ ƒ”‡ƒ• Ž‘•‡ –‘ ƒ’‹ƒ”‹‡•ǡ ™Š‡”‡ Š‘‡› „‡‡ †‡•‹–‹‡• ™‡”‡ Š‹‰Š‡”Ǥ ƒ”‰‡Ǧ•‹œ‡† „‡‡ •’‡ ‹‡•ǡ ™‹–Š ‰”‡ƒ–‡” ‡‡”‰‡–‹  ”‡“—‹”‡‡–•ǡ ƒ† –Š‡”‡ˆ‘”‡ ‘”‡ Ž‹‡Ž› –‘ „‡ ƒˆˆ‡ –‡†„›Ž‘™ˆ‘‘†”‡•‘—” ‡ ƒ˜ƒ‹Žƒ„‹Ž‹–›ǡ ™‡”‡ Ž‡•• ƒ„—†ƒ– ‹ ƒ”‡ƒ• Ž‘•‡ –‘ ƒ’‹ƒ”‹‡•Ǥ Š‡•‡ •’ƒ–‹ƒŽ Šƒ‰‡• ‹ ‘—‹–›•–”— –—”‡ƒ”‡„‘—†–‘ƒˆˆ‡ ––Š‡‹†‡–‹–›ƒ†–Š‡‡–™‘” •–”— –—”‡ ‘ˆ –Š‡ ‹–‡”ƒ –‹‘• „‡‡• ‡•–ƒ„Ž‹•Š ™‹–Š ‘–Š‡” ‘”‰ƒ‹•• ‘•‡“—‡–Ž›ǡ ‹–Š‡–Š‹”† Šƒ’–‡”ǡ™‡ •–—†› –Š‡ Ž‘ ƒŽ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ„‹Ž‹–› ‘ˆ –Š‡ ‘—‹–› ‘ˆ ƒ˜‹–›Ǧ‡•–‹‰ „‡‡• ƒ† ™ƒ•’• –‘‰‡–Š‡” ™‹–Š –Š‡‹” ‡•–• ƒ••‘ ‹ƒ–‡• ȋ’ƒ”ƒ•‹–‘‹†•ǡ Ž‡’–‘’ƒ”ƒ•‹–‡•ǡ ’”‡†ƒ–‘”• ƒ† • ƒ˜‡‰‡”•Ȍǡƒ•™‡ŽŽƒ•–Š‡‹”‹–‡”ƒ –‹‘•Ǥ‡ƒƒŽ›œ‡–Š‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡ ‘—‹–› ƒ† ‹–‡”ƒ –‹‘ EǦ†‹˜‡”•‹–›ǡ ƒ† ‡š’Ž‘”‡ –Š‡ •‘—” ‡• ‘ˆ –Š‡ ‘„•‡”˜‡† •’ƒ–‹ƒŽ ˜ƒ”‹ƒ„‹Ž‹–›Ǥ

 1 

’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‘–Š ‘—‹–‹‡•ȋŠ‘•–•ƒ†’ƒ”ƒ•‹–‡•Ȍ™ƒ•Š‹‰Šƒ†ƒ‹Ž›†”‹˜‡„› •’‡ ‹‡•–—”‘˜‡”ǡ™‹–Šƒ˜‡”›Ž‘™‹ˆŽ—‡ ‡‘ˆ‡•–‡†‡••ǡ‡ƒ‹‰–Šƒ–Ž‘ ƒŽ ‘—‹–‹‡• ™‡”‡Š‹‰ŠŽ›‹†‹‘•› ”ƒ–‹ Ǥ –‡”ƒ –‹‘EǦ†‹˜‡”•‹–›™ƒ•ƒŽ•‘˜‡”›Š‹‰Šƒ†‘•–Ž›†—‡–‘–Š‡ Š‹‰Š•’‡ ‹‡•–—”‘˜‡”ǡ™‹–Šƒ˜‡”›Ž‘™ ‘–”‹„—–‹‘‘ˆ‹–‡”ƒ –‹‘”‡™‹”‹‰Ǥ ‘–Š‡”™‘”†•ǡ •’‡ ‹‡•–‡†‡†–‘‹–‡”ƒ –•‹‹Žƒ”Ž›ƒ ”‘••’Ž‘–•Ǥ˜‡”ƒŽŽǡ–Š‹•–Š‡•‹•†‡‘•–”ƒ–‡•–Šƒ–„‡‡ ‘—‹–‹‡•ƒ†–Š‡‹”ƒ–ƒ‰‘‹•–‹ ‹–‡”ƒ –‹‘‡–™‘”•ƒ›˜ƒ”›ƒ–ƒ˜‡”›•ƒŽŽ• ƒŽ‡ƒ† ƒ”‡ Š‹‰ŠŽ› ‘†‹–‹‘‡† „› Ž‘ ƒŽ ˆƒ –‘”•ǡ ƒ† –Šƒ– „‡‡ ‘—‹–‹‡• ƒ› „‡ ƒˆˆ‡ –‡† „› ‹–‡•‹˜‡„‡‡‡‡’‹‰ǡƒ™‹†‡•’”‡ƒ†Š—ƒƒ –‹˜‹–›—•—ƒŽŽ›ƒ••—‡†–‘„‡„‡‡ˆ‹ ‹ƒŽǤ







 2 

Resum

Ž• ‘”‰ƒ‹•‡• ‡• –”‘„‡ †‹•–”‹„—Ö• Š‡–‡”‘‰°‹ƒ‡– •‡‰—‹– ’ƒ–”‘• ‡•’ƒ ‹ƒŽ•ǡ ‡Ž• “—ƒŽ• •× ‘•‡“ò° ‹ƒ†‡˜ƒ”‹•ˆƒ –‘”•„‹Ö–‹ •‹ƒ„‹Ö–‹ •Ǥ†‡Ž•’”‹ ‹’ƒŽ•‘„Œ‡ –‹—•†‡Žǯ‡ ‘Ž‘‰‹ƒ ±• ‡–‡†”‡ ‘ Ž‡• ‹–‡”ƒ ‹‘• †‡ Ž‡• ‡•’° ‹‡• ƒ„ ƒŽ–”‡• ‘”‰ƒ‹•‡• ‹ ƒ„ Žǯƒ„‹‡– †‡–‡”‹‡ ‡Ž• •‡—• ’ƒ–”‘• †‡ †‹•–”‹„— ‹×Ǥ  ƒ“—‡•–ƒ –‡•‹ ƒƒŽ‹–œ‡‡Ž•ˆƒ –‘”•“—‡ ‡š’Ž‹“—‡ Žƒ ˜ƒ”‹ƒ„‹Ž‹–ƒ– ‡•’ƒ‹ƒŽ †‡ Ž‡• ‘—‹–ƒ–• †ǯƒ„‡ŽŽ‡• ‹ †‡ Ž‡• šƒ”š‡• †ǯ‹–‡”ƒ ‹‘• ƒ••‘ ‹ƒ†‡•ƒ‡• ƒŽƒŽ‘ ƒŽǤǯ‡•–—†‹•ǯŠƒ†—–ƒ–‡”‡ƒŽƒ” †‡Ž ƒ””ƒˆǡ‡—ƒ”‡ƒ†‡͵ʹʹ ƒ„ —ƒ ˜‡‰‡–ƒ ‹× ƒ”„—•–‹˜ƒ ‡†‹–‡””‹ƒ Š‘‘‰°‹ƒǡ •‡•‡ „ƒ””‡”‡• ‡ ‘ŽÖ‰‹“—‡• ‹ ‰”ƒ†‹‡–•ƒ” ƒ–•Ǥ–°•“—‡Ž‡•ƒ„‡ŽŽ‡•†‡’‡‡†‡Ž•”‡ —”•‘•ˆŽ‘”ƒŽ•‹†‡‹†‹ˆ‹ ƒ ‹×’‡”–ƒŽ †‡•‘„”‡˜‹—”‡‹”‡’”‘†—‹”Ǧ•‡ǡ•ǯ‡•’‡”ƒ“—‡ƒ“—‡•–•‹ˆŽ—‡ ‹ÃŽƒ˜ƒ”‹ƒ„‹Ž‹–ƒ–‡•’ƒ ‹ƒŽŽ‘ ƒŽ†‡ Žƒ ‘—‹–ƒ– †ǯƒ„‡ŽŽ‡•Ǥ  ‡Ž ’”‹‡” ƒ’À–‘Ž ‡š’Ž‘”‡ ‘ Žƒ †‹•–”‹„— ‹× ‡•’ƒ ‹ƒŽ †‡Ž• ”‡ —”•‘• ˆŽ‘”ƒŽ• ‹ †‡ ‹†‹ˆ‹ ƒ ‹× †‡–‡”‹ƒ Žƒ ˜ƒ”‹ƒ„‹Ž‹–ƒ– ‡•’ƒ‹ƒŽ Ž‘ ƒŽ †‡ Žƒ ‘—‹–ƒ– †ǯƒ„‡ŽŽ‡•Ǥ ‡–”‘„ƒ–—’ƒ–”׉‡‘‰”ˆ‹  Žƒ”†‡˜ƒ”‹ƒ„‹Ž‹–ƒ–‡•’ƒ‹ƒŽ‡Žƒ ‘’‘•‹ ‹×†‡Žƒ ‘—‹–ƒ–†ǯƒ„‡ŽŽ‡•Ǥ“—‡•–’ƒ–”ׇ•†‡—’ƒ” ‹ƒŽ‡–ƒŽ•”‡ —”•‘• ˆŽ‘”ƒŽ•ǡ ƒ„ —ƒ ‘–”‹„— ‹× †‡ Žƒ †‹•’‘‹„‹Ž‹–ƒ– †‡ •—„•–”ƒ–• †‡ ‹†‹ˆ‹ ƒ ‹× “—ƒ•‹ ‹•‹‰‹ˆ‹ ƒ–ǡ ‹ ‡•– ‡•–”‡–ƒ‡– ”‡Žƒ ‹‘ƒ†ƒ ƒ„ Žƒ ‹†ƒ ‘”’‘”ƒŽǣ Ž‡• ‡•’° ‹‡• †ǯƒ„‡ŽŽƒ ’‡–‹–‡• ȋδͷͷ‰Ȍ ‘•–”‡ˆ‘”–•’ƒ–”‘•‡•’ƒ ‹ƒŽ•ǡ‡–”‡“—‡Ž‡•‡•’° ‹‡•‰”ƒ•ȋε͹Ͳ‰Ȍ–‡†‡‹š‡ƒ–”‘„ƒ”Ǧ •‡†‹•–”‹„—Ƈ•±•—‹ˆ‘”‡‡–‡Žǯ”‡ƒ†ǯ‡•–—†‹Ǥƒ†‹•–”‹„— ‹×‡•’ƒ ‹ƒŽ†‡Ž‡•ƒ„‡ŽŽ‡• –ƒ„±’‘–˜‡—”‡ǯ•ƒˆ‡ –ƒ†ƒ’‡””°‰‹•†‡’‡”–‘”„ƒ ‹‘•Ǥǯ”‡ƒ†ǯ‡•–—†‹ ‘–±˜ƒ”‹•ƒ’‹ƒ”‹•ǡ ‘ˆ‡”‹–Ǧ‘• Žƒ ‘’‘”–—‹–ƒ– †ǯ‡š’Ž‘”ƒ” ‡Ž• ‡ˆ‡ –‡• ’‘–‡ ‹ƒŽ• †‡ Žǯƒ’‹ —Ž–—”ƒ ‡ ‘—‹–ƒ–• †ǯƒ„‡ŽŽ‡••ƒŽ˜ƒ–‰‡•ȋƒ’À–‘ŽʹȌǤǯƒ„‡ŽŽƒ†‡Žƒ‡ŽȋApismelliferaȌ±•—ƒ‡•’° ‹‡‘Ž–‡ˆ‹ ‹‡– ƒŽǯŠ‘”ƒ†‡„—• ƒ”‡Œƒ”‹ǡƒ±•ǡ–±ŽǯŠƒ„‹Ž‹–ƒ–†‡”‡ Ž—–ƒ”ƒŽ–”‡•ƒ„‡ŽŽ‡•†‡Žƒ‡Ž‡‡Ž•ŽŽ‘ • ƒ„ ±• †‹•’‘‹„‹Ž‹–ƒ– †ǯƒŽ‹‡–ǡ ‡•–ƒ„Ž‹– ’‘–‡ ‹ƒŽ‡– — ‡ˆ‡ –‡ ‘’‡–‹–‹— •‘„”‡ ‡Ž• ƒŽ–”‡•’‘ŽȉŽ‹‹–œƒ†‘”•Ǥ ‡–”‘„ƒ–“—‡Ž‡•ƒ„‡ŽŽ‡•†‡Žƒ‡Ž•×Ž‡•“—‡ ‘–”‹„—‡‹š‡†ǯ—ƒ ƒ‡”ƒ±••‹‰‹ˆ‹ ƒ–‹˜ƒƒŽƒ†‹•‹— ‹×†‡Ž’‘ŽȉŽ‡‹‡Ž° –ƒ”†‡Ž‡•†—‡•’Žƒ–‡•±• ‹’‘”–ƒ–•ƒŽǯ”‡ƒ†ǯ‡•–—†‹Ǥƒ„±Š‡–”‘„ƒ–“—‡Žƒ ‘—‹–ƒ–†ǯƒ„‡ŽŽ‡•‡•˜‡—‘†‹ˆ‹ ƒ†ƒ ‡ ”‡‡• ’”‘’‡”‡• ƒŽ• ƒ’‹ƒ”‹•ǡ ‘ Ž‡• †‡•‹–ƒ–• †ǯƒ„‡ŽŽ‡• †‡ Žƒ‡Ž•×±•‡Ž‡˜ƒ†‡•Ǥ‡• ‡•’° ‹‡•†ǯƒ„‡ŽŽ‡•‰”ƒ•ǡƒ„ƒŒ‘”•”‡“—‡”‹‡–•‡‡”‰°–‹ •‹ǡ ‘•‡“ò‡–‡–ǡƒ„±• ’”‘„ƒ„‹Ž‹–ƒ–•†‡˜‡—”‡ǯ•ƒˆ‡ –ƒ†‡•’‡”—ƒ†‹•’‘‹„‹Ž‹–ƒ–„ƒ‹šƒ†‡”‡ —”•‘•ƒŽ‹‡–ƒ”‹•ǡ•× ‡›•ƒ„—†ƒ–•‡”‡‡•’”‘’‡”‡•ƒŽ•ƒ’‹ƒ”‹•Ǥ“—‡•–• ƒ˜‹•‡•’ƒ ‹ƒŽ•‡Žǯ‡•–”— –—”ƒ†‡ Ž‡• ‘—‹–ƒ–•ƒˆ‡ –ƒ”ƒŽƒ‹†‡–‹–ƒ–‹Žǯ‡•–”— –—”ƒ†‡Ž‡•‹–‡”ƒ ‹‘•†‡Ž‡•šƒ”š‡•“—‡Ž‡• ƒ„‡ŽŽ‡•‡•–ƒ„Ž‡‹š‡ƒ„ƒŽ–”‡•‘”‰ƒ‹•‡•Ǥ ‘•‡“ò° ‹ƒǡ‡‡Ž–‡” ‡” ƒ’À–‘Žǡ‡•–—†‹‡Žƒ

 3 

˜ƒ”‹ƒ„‹Ž‹–ƒ–‡•’ƒ ‹ƒŽƒ‡• ƒŽƒŽ‘ ƒŽ†ǯ—ƒ ‘—‹–ƒ–†ǯƒ„‡ŽŽ‡•‹˜‡•’‡•‹†‹ˆ‹ ƒ–•‡ ƒ˜‹–ƒ–• ’”‡‡•–ƒ„Ž‡”–‡• ‹ †‡ ŽŽ—” ˆƒ—ƒ ƒ••‘ ‹ƒ†ƒ ȋ’ƒ”ƒ•‹–‘‹†•ǡ Ž‡’–‘’ƒ”•‹–•ǡ ’”‡†ƒ†‘”• ‹ ƒ””‘›ƒ‹”‡•Ȍǡƒ‹šÀ ‘†‡Ž‡••‡˜‡•‹–‡”ƒ ‹‘•ǤƒŽ‹–œ‡Žƒ”‡Žƒ ‹×‡–”‡ŽƒEǦ†‹˜‡”•‹–ƒ–†‡ Ž‡• ‘—‹–ƒ–•‹†‡Ž‡•‹–‡”ƒ ‹‘•‹‡š’Ž‘”‡Ž‡• ƒ—•‡•†‡Žƒ ˜ƒ”‹ƒ„‹Ž‹–ƒ– ‡•’ƒ ‹ƒŽ ‘„•‡”˜ƒ†ƒǤƒ˜ƒ”‹ƒ„‹Ž‹–ƒ–‡•’ƒ ‹ƒŽ†ǯƒ„†—‡• ‘—‹–ƒ–•ȋŠ‘•–‡•‹’ƒ”ƒ•‹–‘‹†•Ȍ±•‡Ž‡˜ƒ†ƒ‹ †‡‰—†ƒ’”‹ ‹’ƒŽ‡–ƒŽ”‡ ƒ˜‹†ǯ‡•’° ‹‡•ǡƒ„—ƒ‹ˆŽ—° ‹ƒ‘Ž–„ƒ‹šƒ†‡Žǯƒ‹—ƒ‡–ǡ ‹†‹ ƒ– “—‡ Ž‡• ‘—‹–ƒ–• Ž‘ ƒŽ• •× ƒŽ–ƒ‡– ‹†‹‘•‹ ”–‹“—‡•Ǥ ƒ EǦ†‹˜‡”•‹–ƒ– †ǯ‹–‡”ƒ ‹‘•–ƒ„±±•‘Ž–‡Ž‡˜ƒ†ƒ‹†‡‰—†ƒ’”‹ ‹’ƒŽ‡–ƒŽ”‡ ƒ˜‹†ǯ‡•’° ‹‡•ǡƒ„—ƒ ‘–”‹„— ‹× ‘Ž– „ƒ‹šƒ †‡Ž ”‡ ƒ„Ž‡Œƒ– †ǯ‹–‡”ƒ ‹‘•Ǥ  ƒŽ–”‡•’ƒ”ƒ—Ž‡•ǡŽ‡•‡•’° ‹‡• –‡†‡‹š‡ƒ‹–‡”ƒ ‹‘ƒ”†‡ƒ‡”ƒ•‹‹Žƒ”‡–‘–‡•Ž‡•’ƒ” ‡ŽȉŽ‡•Ǥ‰‡‡”ƒŽǡƒ“—‡•–ƒ–‡•‹ †‡‘•–”ƒ“—‡Ž‡• ‘—‹–ƒ–•†ǯƒ„‡ŽŽ‡•‹Ž‡••‡˜‡•šƒ”š‡•†ǯ‹–‡”ƒ ‹‘•ƒ–ƒ‰‘‹•–‡•’‘†‡ ˜ƒ”‹ƒ”ƒ—ƒ‡• ƒŽƒ‡•’ƒ‹ƒŽ‘Ž–’‡–‹–ƒ‹‡•˜‡—‡ƒŽ–ƒ‡– ‘†‹ ‹‘ƒ†‡•’‡Ž•ˆƒ –‘”•Ž‘ ƒŽ•ǡ‹ “—‡Ž‡• ‘—‹–ƒ–•†ǯƒ„‡ŽŽ‡•’‘†‡˜‡—”‡ǯ•ƒˆ‡ –ƒ†‡•’‡”Žǯƒ’‹ —Ž–—”ƒ‹–‡•‹˜ƒǡ—ƒƒ –‹˜‹–ƒ– Š—ƒƒ‰‡‡”ƒŽ‹–œƒ†ƒ“—‡Šƒ„‹–—ƒŽ‡–±• ‘•‹†‡”ƒ†ƒ ‘ƒ„‡‡ˆ‹ ‹‘•ƒǤ







 4 

   

Generalintroduction













  ‡‡”ƒŽ‹–”‘†— –‹‘

Introduction

 ‘Ž‘‰› ƒ„‡†‡ˆ‹‡†ƒ•–Š‡•–—†›‘ˆ‘”‰ƒ‹••ƒ†–Š‡‹”‹–‡”ƒ –‹‘• ™‹–Š –Š‡‹” ‡˜‹”‘‡– ƒ† ™‹–Š ‘–Š‡” ‘”‰ƒ‹•• –Šƒ– †‡–‡”‹‡ ƒ„—†ƒ ‡ ƒ† †‹•–”‹„—–‹‘ ‘ˆ •’‡ ‹‡• ȋ”‡„• ‡– ƒŽǤ ͳͻͻͶȌǤ ‘—–Ž‡•• •–—†‹‡• Šƒ˜‡ „‡‡ †‘‡ –‘ —†‡”•–ƒ† Š‘™ •— Š ‹–‡”ƒ –‹‘••Šƒ’‡–Š‡‘„•‡”˜‡†’ƒ––‡”•‘ˆ†‹˜‡”•‹–›ƒ††‹•–”‹„—–‹‘ ‘ˆ •’‡ ‹‡• ȋ‡Ǥ‰Ǥ ƒ”™‹ ͳͺͷͻǢ —– Š‹•‘ ͳͻͷͻǢ ƒ ”–Š—” ͳͻ͹ʹǢ ƒ› ͳͻͺͺȌǤ ƒ–—”‡‹•Š‹‰ŠŽ› Š‡–‡”‘‰‡‡‘—•ǡ‡ƒ‹‰–Šƒ–ƒ‹ƒŽ•ƒ†’Žƒ–•ƒ”‡†‹•–”‹„—–‡†—‡˜‡Ž›ƒ ”‘••–Š‡•’ƒ ‡ǡ „‡ ‘‹‰•›•–‡•‘ˆ‘”‰ƒ‹œ‡†Š‡–‡”‘‰‡‡‹–›ȋƒ”‰ƒŽ‡ˆͳͻ͸ͺȌǤŠ‡‘„•‡”˜‡†Š‡–‡”‘‰‡‡‹–› ‹ –Š‡ •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‘ˆ •’‡ ‹‡• ‹• ƒ ‘•‡“—‡ ‡ ‘ˆ –Š‡ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ„‹Ž‹–› ‘ˆ ƒ„‹‘–‹  ˆƒ –‘”•ƒ†„‹‘–‹ ‹–‡”ƒ –‹‘•ǡ‹ ‘„‹ƒ–‹‘™‹–Š’”‘ ‡••‡•‘ˆ‡š–‹ –‹‘ƒ† ‘Ž‘‹œƒ–‹‘ ȋ–”‘‰‡–ƒŽǤͳͻͺͶǡ‹ Ž‡ˆ•ƒ† ŠŽ—–‡”ͳͻͻ͵ǡ‡‰‘‡–ƒŽǤʹͲͲ͸ȌǤŠ—•ǡ•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘ ’ƒ––‡”• ’”‘˜‹†‡ ‹’‘”–ƒ– ‹ˆ‘”ƒ–‹‘ ‘– ‘Ž› ƒ„‘—– —””‡– ‘†‹–‹‘• „—– ƒŽ•‘ ƒ„‘—– ’ƒ•–’”‘ ‡••‡•ƒ–ƒ‰‹˜‡•‹–‡Ǥ

–Šƒ•„‡‡™‡ŽŽ‡•–ƒ„Ž‹•Š‡†–Šƒ–‡˜‹”‘‡–ƒŽŠ‡–‡”‘‰‡‡‹–›‹• ”— ‹ƒŽ–‘ƒ‹–ƒ‹•’‡ ‹‡• †‹˜‡”•‹–›ȋ‡Ǥ‰Ǥƒ ”–Š—”ƒ†ƒ ”–Š—”ͳͻ͸ͳǡ‹ƒƒͳͻ͸͸ǡ‡‹„—ŽŽ‡–ƒŽǤʹͲͲͲȌǡƒ†–Šƒ– Š‘‘‰‡‹œƒ–‹‘ ‘ˆ –Š‡ ‡˜‹”‘‡– ƒ Ž‡ƒ† –‘ „‹‘–‹  ‹’‘˜‡”‹•Š‡– ƒ† ‘•‡“—‡– Š‘‘‰‡‹œƒ–‹‘ ‘ˆ –Š‡ „‹‘–ƒ ȋ ‹‡› ƒ† ‘ ™‘‘† ʹͲͲͳǡ ƒŠ‡Ž ʹͲͲʹǡ Š”—•Š ‡– ƒŽǤ ʹͲͲ͸ȌǤ —”–Š‡”‘”‡ǡ‹ ”‡ƒ•‡†„‹‘†‹˜‡”•‹–›—•—ƒŽŽ›’”‘‘–‡•ˆ— –‹‘ƒŽ†‹˜‡”•‹–›ǡ™Š‹ ŠŠƒ• „‡‡’”‘˜‡–‘—Ž–‹ƒ–‡Ž›‰—ƒ”ƒ–‡‡‡ ‘•›•–‡•‡”˜‹ ‡•ȋÇƴƒœƒ†ƒ„‹†‘ʹͲͲͳǡƒ†‘––‡‡– ƒŽǤʹͲͳͳǡ ƒ‰‹ ‡–ƒŽǤʹͲͳͷȌǤ ‘™‡˜‡”ǡ‰Ž‘„ƒŽ„‹‘†‹˜‡”•‹–›‹•–Š”‡ƒ–‡‡†„›—Ž–‹’Ž‡ˆƒ –‘”•ǡ ’‘–‡–‹ƒŽŽ› Œ‡‘’ƒ”†‹œ‹‰ ‡ ‘•›•–‡ •‡”˜‹ ‡•Ǥ ˆ‘”–—ƒ–‡Ž›ǡ ‹ˆ‘”ƒ–‹‘ ƒ„‘—– •’‡ ‹‡• †‹•–”‹„—–‹‘ƒ†•’ƒ–‹ƒŽ†›ƒ‹ •‹•Žƒ ‹‰ˆ‘”‘•–•’‡ ‹‡•ƒ†‘•–”‡‰‹‘•‘ˆ–Š‡™‘”Ž†Ǥ



Localvariabilityofcommunitycompositionandstructure

•†‡ˆ‹‡†„›ƒ”‰ƒŽ‡ˆȋͳͻͻ͵Ȍǡ„‹‘•’Š‡”‡‹•‘–‘Ž› ‘’‘•‡†„›‹†‹˜‹†—ƒŽ•ǡ™Š‹ Šƒ”‡–Š‡ ‘”‡”•–‘‡•‘ˆ‡ ‘•›•–‡•ǡ„—–ƒŽ•‘„›–Š‡’Š›•‹ ƒŽ‡˜‹”‘‡–Ǥ‘’—Žƒ–‹‘•‡‡†–‘ˆ‹† ƒ†‡“—ƒ–‡ƒ„‹‘–‹ ƒ†„‹‘–‹ ”‡•‘—” ‡•–‘„‡ƒ„Ž‡–‘•—•–ƒ‹–Š‡•‡Ž˜‡•ƒ–ƒ‰‹˜‡•‹–‡Ǥ‡‡ ˆ”‘–Š‹•’‡”•’‡ –‹˜‡ǡƒ‰‘‘†Ǧ“—ƒŽ‹–›Šƒ„‹–ƒ–•Š‘—Ž†’”‘˜‹†‡‡‘—‰Š”‡•‘—” ‡•ˆ‘”•—”˜‹˜ƒŽ ƒ†”‡’”‘†— –‹‘ǡƒ†Šƒ„‹–ƒ– “—ƒŽ‹–›‹•‘™–‘‹ˆŽ—‡ ‡ •’‡ ‹‡•†‹˜‡”•‹–›ȋ‡”‰ ͳͻͻ͹ǡ ‡––•–‡‹ ƒ†  Š‹† ͳͻͻͻǡ Š‘ƒ• ‡– ƒŽǤ ʹͲͲͳȌǤ Š‡”‡ˆ‘”‡ǡ ”‡•‘—” ‡ ƒ˜ƒ‹Žƒ„‹Ž‹–› ƒ†  7 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•   

†‹•–”‹„—–‹‘ƒ”‡‡š’‡ –‡†–‘’Žƒ›ƒ˜‡”›‹’‘”–ƒ–”‘Ž‡‹†‡–‡”‹‹‰•’‡ ‹‡•†‹˜‡”•‹–›ƒ† ‘’‘•‹–‹‘ȋƒ ”–Š—”ͳͻ͸ͷǡ ƒ‡‡–ƒŽǤʹͲͲʹǡ ‘Š‡–ƒŽǤʹͲͲ͹ȌǤ

’‡ ‹‡• ˆ— –‹‘ƒŽ –”ƒ‹–• ƒ”‡ ƒŽ•‘ ‡š’‡ –‡† –‘ •Šƒ’‡ „‹‘†‹˜‡”•‹–› ’ƒ––‡”• ȋŠ‹’Ž‡› ‡– ƒŽǤ ʹͲͲ͸ǡ ”‡‡‡–ƒŽǤʹͲͲͺǡ”ƒˆ–‡–ƒŽǤʹͲͲͺȌǤ•’‡ ‹ƒŽŽ›‹’‘”–ƒ–‹•–Š‡—•‡‘ˆ•’ƒ ‡„›‘„‹Ž‡ ‘”‰ƒ‹••ǡ™Š‹ Š‹•—•—ƒŽŽ›”‡Žƒ–‡†–‘„‘†›•‹œ‡ȋ™‹Šƒ”–‡–ƒŽǤͳͻͺͺǡ‡Ž–ƒ†—”‡ͳͻͻͻǡ ”‡‡Ž‡ƒˆ‡–ƒŽǤʹͲͲ͹ȌǤ‘„‹Ž‹–›‹•‹’‘”–ƒ–‘–‘Ž›‹”‡Žƒ–‹‘–‘ˆ‘”ƒ‰‹‰”ƒ‰‡•ǡ„—–ƒŽ•‘ ‹–‡”•‘ˆ†‹•’‡”•ƒŽƒ„‹Ž‹–›ǡ™Š‹ Š†‡–‡”‹‡•–Š‡ ƒ’ƒ ‹–›–‘ ‘Ž‘‹œ‡‡™ƒ”‡ƒ•ȋ‹ŽŽ•‘ ͳͻͻ͵ǡ‡‡†‡–ƒŽǤʹͲͲͲǡ—ŽŽ‘ ‡–ƒŽǤʹͲͲʹǡŽƒ‹•ƒ ‡‡–ƒŽǤʹͲͲͺȌǤ

‘ ƒŽ ‘—‹–›•–”— –—”‡ ƒƒŽ•‘„‡ƒˆˆ‡ –‡†„›Š‹•–‘”‹ ƒŽ‡˜‡–•ȋ‹ Ž‡ˆ•ͳͻͺ͹ǡ —ƒ‹ ʹͲͳͲȌǤ‘•‡“—‡–Ž›ǡ —””‡– ‘—‹–›ƒ••‡„Žƒ‰‡•ƒ”‡–Š‡”‡•—Ž–‘ˆ•’‡ ‹‡•”‡•’‘•‡•–‘ ’ƒ•–’”‘ ‡••‡••— Šƒ•‹‹‰”ƒ–‹‘Ȁ‡‹‰”ƒ–‹‘ƒ†‡š–‹ –‹‘‡˜‡–• ‘” Š‹•–‘”‹ ƒŽ ’‘’—Žƒ–‹‘†›ƒ‹ •Ǥ



Effectsofperturbationsonlocalcommunities

ƒ–—”‡‹• ‘•–ƒ–Ž›ƒˆˆ‡ –‡†„›ƒ–—”ƒŽƒ†ƒ–Š”‘’‘‰‡‹ †‹•–—”„ƒ ‡•Ǥ•‘‡‘ˆ–Š‡ƒ‹ †”‹˜‡”• ‘ˆ Šƒ„‹–ƒ– ‘†‹ˆ‹ ƒ–‹‘ǡ †‹•–—”„ƒ ‡• ƒ› Šƒ˜‡ ’”‘ˆ‘—† ‡ˆˆ‡ –• ‘ ‘—‹–› †›ƒ‹ •ǡ•‘‡–‹‡•Ž‡ƒ†‹‰–‘”ƒ†‹ ƒŽ Šƒ‰‡•ȋ›•–”ڏ‡–ƒŽǤʹͲͲͲǡ ƒ •‘ƒ†˜‡”’‡  ʹͲͲͲǡ ƒ”†‘•‘ ‡– ƒŽǤ ʹͲͲͺȌǤ — Š †‹•–—”„ƒ ‡• ƒ”‡ ‘•‹†‡”‡† –‘ „‡ ƒ ’”‹ƒ”› ƒ—•‡ ‘ˆ •’ƒ–‹ƒŽŠ‡–‡”‘‰‡‡‹–›‹ƒ–—”ƒŽ ‘—‹–‹‡•ȋ‘—•ƒͳͻͺͶȌǤ ‘”‹•–ƒ ‡ǡ™‹Ž†ˆ‹”‡‹•‘™ –‘’Žƒ›ƒ‹’‘”–ƒ–”‘Ž‡‹–Š‡•–”— –—”‡‘ˆ™‘”Ž†‡ ‘•›•–‡•ȋ‘† ƒ† ‡‡Ž‡› ʹͲͲͷȌǤ Š‹Ž‡•‘‡•’‡ ‹‡•ƒ›„‡ˆƒ˜‘”‡†„›ˆ‹”‡ǡ‘–Š‡”•’‡ ‹‡•ƒ”‡‡‰ƒ–‹˜‡Ž›ƒˆˆ‡ –‡†ƒ†ƒ› ‡˜‡—†‡”‰‘Ž‘ ƒŽ‡š–‹ –‹‘ȋ•“—‡‡–ƒŽǤʹͲͲ͵ǡ‡”‡•‡–ƒŽǤʹͲͲ͵ǡ”ƒ‡–ƒŽǤʹͲͲ͸ȌǤ’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‘ˆ ‘—‹–‹‡• ™‹ŽŽ „‡ ƒŽ•‘ †‡–‡”‹‡† „› –Š‡ Š‡–‡”‘‰‡‡‹–› ‘ˆ ’‡”–—”„ƒ–‹‘ ‹–‡•‹–›ƒ†ˆ”‡“—‡ ›ƒ†–Š‡Š‡–‡”‘‰‡‡‹–›‘ˆ’‘•–Ǧ’‡”–—”„ƒ–‹‘’ƒ––‡”•ȋ—”‡”‡–ƒŽǤ ͳͻͻͺǡ‘†”‹‰‘‡–ƒŽǤʹͲͲͶȌǤ

–Š”‘’‘‰‡‹  ’‡”–—”„ƒ–‹‘•ǡ ‹ Ž—†‹‰ Ž‹ƒ–‡ Šƒ‰‡ǡ Žƒ† —•‡ Šƒ‰‡ ƒ† ƒ› „‹‘–‹  Šƒ‰‡••— Šƒ•„‹‘Ž‘‰‹ ƒŽ‹˜ƒ•‹‘•ǡƒ”‡–Š‡ƒŒ‘” ƒ—•‡‘ˆŠƒ„‹–ƒ–‘†‹ˆ‹ ƒ–‹‘ȋ‹–‘—•‡ ‡–ƒŽǤͳͻͻ͹ȌǤƒ†—•‡ Šƒ‰‡ǡ’”‘„ƒ„Ž›–Š‡‘•– ‘‘ƒŽ–‡”ƒ–‹‘‘ˆƒ–Š”‘’‘‰‡‹ ‘”‹‰‹ǡ ƒ›‡–ƒ‹Ž’”‘ˆ‘—† Šƒ‰‡•‹ ‘—‹–› ‘’‘•‹–‹‘ƒ†•–”— –—”‡–Š”‘—‰Š„‹‘–‹ ƒ† ƒ„‹‘–‹   ˆƒ –‘”• ȋ‘™‹‰ ‡– ƒŽǤ ͳͻͻͻǡ ‘••‹‘ ‡– ƒŽǤ ʹͲͲͷǡ ‡”‡‹”ƒ ‡– ƒŽǤ ʹͲͳͲȌǡ ƒ† ƒ  8   ‡‡”ƒŽ‹–”‘†— –‹‘

’‘–‡–‹ƒŽŽ› ƒŽ–‡” ‡ ‘•›•–‡ •‡”˜‹ ‡• ȋ”‡‡ ‡– ƒŽǤ ʹͲͲ͹ȌǤ ‘” ‡šƒ’Ž‡ǡ —”„ƒ‹œƒ–‹‘ Šƒ• „‡‡ ’”‘˜‡ –‘ „‡ –Š‡ ƒ—•‡ ‘ˆ •‡˜‡”ƒŽ ‘–‘”‹‘—• ‡š–‹ –‹‘• ‘ˆ ƒ–‹˜‡ •’‡ ‹‡•ǡ Ž‡ƒ†‹‰ –‘ „‹‘–‹ Š‘‘‰‡‹œƒ–‹‘ǡƒ†•–”‘‰Ž›‡Šƒ ‹‰–Š‡‡•–ƒ„Ž‹•Š‡–‘ˆ‘ƒ–‹˜‡•’‡ ‹‡•ȋŽƒ‹” ʹͲͲͳǡƒ”œŽ—ˆˆʹͲͲͳǡ ‹‡›ʹͲͲ͸ǡƒ— Šƒ”†‡–ƒŽǤʹͲͲ͸ǡ †‘ƒŽ†‡–ƒŽǤʹͲͲͺǡŠ‘ Šƒ–‡– ƒŽǤʹͲͳͲȌǤŽ‹ƒ–‡ Šƒ‰‡‹•ƒ‘–Š‡”‹’‘”–ƒ–†”‹˜‡”‘ˆ ‘—‹–›‘†‹ˆ‹ ƒ–‹‘Ǥ‡‘ˆ‹–• ƒ‹‰‡‡”ƒŽ‡ˆˆ‡ –•‹•’‘Ž‡™ƒ”†ƒ†ƒŽ–‹–—†‹ƒŽ•Š‹ˆ–‹•’‡ ‹‡•†‹•–”‹„—–‹‘•ȋƒ”‡•ƒ‡– ƒŽǤͳͻͻͻǡ ‹ Ž‹‰‡–ƒŽǤʹͲͲ͸ǡŠ‘ƒ•ʹͲͳͲǡŠ‡‡–ƒŽǤʹͲͳͳȌǤ

‹ƒŽŽ›ǡ–Š‡‹–”‘†— –‹‘‘ˆ‡š‘–‹ •’‡ ‹‡•‹•Šƒ˜‹‰‹’‘”–ƒ–‹’ƒ –•‘ƒ–‹˜‡ ‘—‹–› •–”— –—”‡•ȋ‡˜‹‡‡–ƒŽǤʹͲͲ͵ǡŠ”‡ˆ‡Ž†ʹͲͳͲȌǤ  ‘Ž‘‰‹ ƒŽ ‹˜ƒ•‹‘• ƒ ƒ””› ‹’‘”–ƒ– ‘•‡“—‡ ‡•ˆ‘” ‘—‹–‹‡•ȋ‘”–‡”ƒ†ƒ˜‹‰ƒ‘ͳͻͻͲǡ‘†‰‡ͳͻͻ͵ǡ ”‘•Š‘ŽœʹͲͲʹȌ ƒ† Šƒ˜‡ ’”‘ˆ‘—† ‡ˆˆ‡ –• ‘ ‹–‡”ƒ –‹‘ ‡–™‘”• ȋ”ƒ˜‡•‡– ƒ† ‹ Šƒ”†•‘ ʹͲͲ͸ǡ ƒ”–‘‡—•‡–ƒŽǤʹͲͲͺȌǤ‹‹Žƒ”Ž›–‘–Š‡‹–”‘†— –‹‘‘ˆ‹˜ƒ•‹˜‡•’‡ ‹‡•ǡ–Š‡‹–”‘†— –‹‘‘ˆ Žƒ”‰‡ †‡•‹–‹‡• ‘ˆ †‘‡•–‹  ƒ‹ƒŽ• •— Š ƒ• •Š‡‡’ ƒ† ƒ––Ž‡ ƒ› ƒŽ•‘ Šƒ˜‡ ’”‘ˆ‘—† ‹’ƒ –•‘ƒ–‹˜‡ ‘—‹–‹‡•–Š”‘—‰Š ‘’‡–‹–‹‘ˆ‘”ˆ‘‘†”‡•‘—” ‡•ȋ–‡™ƒ”–‡–ƒŽǤʹͲͲʹǡ ƒŽ†‹‡–ƒŽǤʹͲͲͶǡ‘—‰‡–ƒŽǤʹͲͲͷȌǤ

’‡ ‹‡•ƒ† ‘—‹–‹‡•ƒ”‡ǡ–‘ƒ‰”‡ƒ–‡”‘”Ž‡••‡”†‡‰”‡‡ǡ•‡•‹„Ž‡ –‘ ƒŽŽ •— Š Šƒ„‹–ƒ– ‘†‹ˆ‹ ƒ–‹‘•ǡ™Š‹ ŠŽ‡ƒ†–‘ Šƒ‰‡•‹•’‡ ‹‡•†‹•–”‹„—–‹‘•ƒ† ‘—‹–›•–”— –—”‡ƒ† ‘’‘•‹–‹‘Ǥ‘•‡“—‡–Ž›ǡ•’‡ ‹‡•ƒ† ‘—‹–›•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›ȋEǦ†‹˜‡”•‹–›Ȍ ƒ„‡ƒ ”‡Ž‡˜ƒ–‡ƒ•—”‡ˆ‘” ‘•‡”˜ƒ–‹‘’—”’‘•‡•ȋ‘†‹–‡–ƒŽǤʹͲͲʹȌǤ



Localspatialvariabilityofinteractions

’‡ ‹‡•‹–‡”ƒ –™‹–Š‡ƒ Š‘–Š‡”ǡˆ‘”‹‰ ‘’Ž‡š‡–™‘”•Ǥ –‡”ƒ –‹‘•’Žƒ›ƒ‡••‡–‹ƒŽ ”‘Ž‡‹•’‡ ‹‡•†‹•–”‹„—–‹‘–”‘—‰Šǡˆ‘”‡šƒ’Ž‡ǡ ‘’‡–‹–‹‘ǡ’”‡†ƒ–‹‘ƒ††‹ˆˆ‡”‡––›’‡• ‘ˆ—–—ƒŽ‹•ȋ‡‰‘‡–ƒŽǤʹͲͲ͸ȌǤ‡˜‡”ƒŽ‘†‡Ž•ǡ•— Šƒ•–Š‡’”‡†ƒ–‘”Ǧ’”‡›‘–ƒǦ‘Ž–‡””ƒ ‘†‡Žǡƒ†–Š‡Š‘•–Ǧ’ƒ”ƒ•‹–‡‹ Š‘Ž•‘Ǧƒ‹Ž‡›‘†‡ŽŠƒ˜‡„‡‡’”‘’‘•‡† –‘ —†‡”•–ƒ† ƒ† ’”‡†‹ – –Š‡ ‘—– ‘‡ ‘ˆ ‹–‡”ƒ –‹‘• ‘ ’‘’—Žƒ–‹‘• ƒ† ‘—‹–‹‡•Ǥ–Š‡”‘†‡Ž• Šƒ˜‡„‡‡’”‘’‘•‡†–‘‡š’Žƒ‹–Š‡—†‡”Ž›‹‰‡ Šƒ‹••–Šƒ–—Ž–‹ƒ–‡Ž› ƒ—•‡•’‡ ‹‡•–‘ ‹–‡”ƒ –Ǥ ‘” ‹•–ƒ ‡ǡ –Š‡ ‡—–”ƒŽ‹–› Š›’‘–Š‡•‹• ’‘•–—Žƒ–‡• –Šƒ– ‹†‹˜‹†—ƒŽ• ‹–‡”ƒ – ”ƒ†‘Ž›ǡ •‘ –Šƒ– –Š‡ ˆ”‡“—‡ › ‘ˆ ‹–‡”ƒ –‹‘• ƒ† –Š‡ —„‡” ‘ˆ ‹–‡”ƒ –‹‰ •’‡ ‹‡• †‡’‡†• ‘ –Š‡ ƒ„—†ƒ ‡ ‘ˆ ‡ƒ Š •’‡ ‹‡• ȋŽò–Š‰‡ ‡– ƒŽǤ ʹͲͲͺǡ žœ“—‡œ ‡– ƒŽǤ ʹͲͲͻ„ȌǤ

 9 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•   

„—†ƒ–•’‡ ‹‡•Šƒ˜‡ƒŠ‹‰Š‡”’”‘„ƒ„‹Ž‹–›–‘‡š’‡”‹‡ ‡ƒ”ƒ†‘‡ ‘—–‡”ƒ†–Š‡”‡ˆ‘”‡ ƒ”‡‘”‡’”‘‡–‘‹–‡”ƒ –Ǥ ƒ–ƒ‰‘‹•–‹ ‡–™‘”•ǡŠ‘•–Ȁ’”‡›ƒ„—†ƒ ‡Šƒ•ƒŽ•‘„‡‡ ’”‘’‘•‡† ƒ• ƒ ‡ Šƒ‹• –‘ ‡š’Žƒ‹ ‡–™‘” ’ƒ––‡”• –Š”‘—‰Š –Š‡ Ž‡ƒ”‹‰ ƒ„‹Ž‹–‹‡• ‘ˆ ’ƒ”ƒ•‹–‡•Ȁ’”‡†ƒ–‘”• ȋ •Š‹‹ ƒ† Š‹ƒ†ƒ ʹͲͳʹȌǤ ‹ƒŽŽ›ǡ –”ƒ‹– ƒ– Š‹‰ ȋ‹ Ž—†‹‰ ‘”’Š‘Ž‘‰‹ ƒŽƒ†’Š‡‘Ž‘‰‹ ƒŽ–”ƒ‹–•Ȍ„‡–™‡‡‹–‡”ƒ –‹‰•’‡ ‹‡•‹•ƒ‡••‡–‹ƒŽ ‘†‹–‹‘ ˆ‘”–Š‡”‡ƒŽ‹œƒ–‹‘‘ˆƒ‹–‡”ƒ –‹‘ȋ ‘”†ƒ‘‡–ƒŽǤʹͲͲ͵ȌǤ

Š‡Š‡–‡”‘‰‡‡‘—•†‹•–”‹„—–‹‘‘ˆ‘”‰ƒ‹••Šƒ•ƒ‡ˆˆ‡ –‘–Š‡‹†‡–‹–›ƒ†•–”‡‰–Š‘ˆ –Š‡‹”‹–‡”ƒ –‹‘•ȋ‰”ƒ™ƒŽ‡–ƒŽǤʹͲͲ͸ȌǡŽ‡ƒ†‹‰–‘Š‹‰Š˜ƒ”‹ƒ„‹Ž‹–›‹‡–™‘”•–”— –—”‡ƒ† ‹–‡”ƒ –‹‘ ‘’‘•‹–‹‘ƒ ”‘•••’ƒ ‡ȋŽ‡•‡ƒ† ‘”†ƒ‘ʹͲͲʹǡžœ“—‡œ‡–ƒŽǤʹͲͲͻ„ȌǤ ‘” ‹•–ƒ ‡ǡ‹–Šƒ•„‡‡†‡‘•–”ƒ–‡†–Šƒ––Š‡•–”— –—”‡‘ˆ’Žƒ–Ǧƒ‹ƒŽ—–—ƒŽ‹•–‹ ‡–™‘”• ‹•Žƒ”‰‡Ž›•Šƒ’‡†„›•’‡ ‹‡•†‹•–”‹„—–‹‘ȋ‘”ƒŽ‡•ƒ†žœ“—‡œʹͲͲͺǡžœ“—‡œ‡–ƒŽǤʹͲͲͻƒǡ —”Ž‡ƒ†Žƒ” אʹͲͳͳȌǤ

‡ ƒ—•‡Ž‘ ƒŽˆƒ –‘”•ƒ› Šƒ‰‡ƒ ”‘•••’ƒ ‡ǡ–Š‡•ƒ‡•’‡ ‹‡•ƒ›‹–‡”ƒ –†‹ˆˆ‡”‡–Ž›ƒ– †‹ˆˆ‡”‡– •‹–‡•Ǥ ’‘”–ƒ–Ž›ǡ •‘‡ ˆƒ –‘”• ƒ› †‹ˆˆ‡”‡–Ž› ƒˆˆ‡ ––Š‡–™‘–”‘’Š‹ Ž‡˜‡Ž• ‹˜‘Ž˜‡†‹ƒ‹–‡”ƒ –‹‘ǡ‹ˆŽ—‡ ‹‰‘–‘Ž›–Š‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆƒ‰‹˜‡•’‡ ‹‡•ǡ„—– ƒŽ•‘‹–•ƒ„‹Ž‹–›–‘‹–‡”ƒ –™‹–Š ‡”–ƒ‹’ƒ”–‡”•Ǥ•–Š‡–”‘’Š‹  Ž‡˜‡Ž ‹ ”‡ƒ•‡•ǡ ‹– ‹• ‹ ”‡ƒ•‹‰Ž› †‹ˆˆ‹ —Ž– ‹• –‘ ’”‡†‹ – •’‡ ‹‡• †‹•–”‹„—–‹‘ „‡ ƒ—•‡ ‘”‡ ˆƒ –‘”• ‹–‡”˜‡‡ ‹ •Šƒ’‹‰•— Š†‹•–”‹„—–‹‘Ǥ

  •—ǡ ‹–‡”ƒ –‹‘ ‡–™‘”• ƒ”‡ ’ƒ”–‹ —Žƒ”Ž› •‡•‹–‹˜‡ –‘ Šƒ„‹–ƒ– Šƒ‰‡ „‡ ƒ—•‡ –Š‡› †‡’‡†‘ƒ ‘„‹ƒ–‹‘‘ˆˆƒ –‘”•†‹ˆˆ‡”‡–Ž›ƒˆˆ‡ –‹‰–Š‡˜ƒ”‹‘—•–”‘’Š‹ Ž‡˜‡Ž•‹˜‘Ž˜‡†Ǥ ‘”–Š‹•”‡ƒ•‘ǡ‡ ‘Ž‘‰‹ ƒŽ‡–™‘”• ƒ„‡”‡‰ƒ”†‡†ƒ•’‘–‡–‹ƒŽ„‹‘‹†‹ ƒ–‘”•‘ˆ‡ ‘Ž‘‰‹ ƒŽ Šƒ‰‡Ǥ ‘” ‹•–ƒ ‡ǡ ƒ ‹’‘˜‡”‹•Š‡† ‡–™‘” •–”— –—”‡ ™‘—Ž† ”‡ˆŽ‡ – „‹‘–‹  Š‘‘‰‡‹œƒ–‹‘ȋŽ„”‡ Š–‡–ƒŽǤʹͲͲ͹ȌǤ –‡”ƒ –‹‘•Šƒ˜‡„‡‡ˆ‘—†–‘„‡‘”‡•‡•‹–‹˜‡–‘ Šƒ„‹–ƒ– †‹•–—”„ƒ ‡ „‡ ƒ—•‡ –Š‡› ƒ› „‡ Ž‘•– „‡ˆ‘”‡ ‘—‹–‹‡••Š‘™ƒ›•‹‰•‘ˆ ƒŽ–‡”ƒ–‹‘ ȋ›Ž‹ƒƒ‹• ‡– ƒŽǤ ʹͲͲ͹ȌǤ ‘‡–Š‡Ž‡••ǡ ‡–™‘”• Šƒ˜‡ •‡Ž†‘ „‡‡ —•‡† ƒ• „‹‘‹†‹ ƒ–‘”•Ǥ

Šƒ‰‡•‹‡–™‘”‘”‰ƒ‹œƒ–‹‘ƒ›—Ž–‹ƒ–‡Ž›ƒˆˆ‡ –‡ ‘•›•–‡•‡”˜‹ ‡•Ǥ ‘”‡šƒ’Ž‡ǡ‹ˆ ‡š–‹ –‹‘”‹•‹•ƒ••‘ ‹ƒ–‡†–‘’ƒ”–‹ —Žƒ”„‹‘Ž‘‰‹ ƒŽƒ––”‹„—–‡•ǡ ‡”–ƒ‹‡ ‘Ž‘‰‹ ƒŽˆ— –‹‘• ƒ›„‡Ž‘•–‹‹’‘˜‡”‹•Š‡†‡–™‘”•ȋ‡‘––‡–ƒŽǤʹͲͲͶǡ ‘–ƒ‹‡‡–ƒŽǤʹͲͲ͸ȌǤ ‘”–Š‹• ”‡ƒ•‘ǡ –Š‡ ‘ ‡’– ‘ˆ ‹–‡”ƒ –‹‘ ‘•‡”˜ƒ–‹‘ Šƒ• „‡‡ ’”‘’‘•‡†ƒ•ƒ‡š–‡•‹‘‘ˆ •’‡ ‹‡• ‘•‡”˜ƒ–‹‘ȋ‡ƒ”•‡–ƒŽǤͳͻͻͺȌǤ  10   ‡‡”ƒŽ‹–”‘†— –‹‘

Ecologicalimportanceofbees

’’”‘š‹ƒ–‡Ž›ͺ͹ǤͷΨ‘ˆ–Š‡‡š‹•–‹‰ƒ‰‹‘•’‡”•”‡Ž›‘ƒ‹ƒŽ’‘ŽŽ‹ƒ–‹‘ȋŽŽ‡”–‘‡–ƒŽǤ ʹͲͳͳȌǤ‘‰–Š‡˜ƒ”‹‘—•‰”‘—’•‘ˆ’‘ŽŽ‹ƒ–‘”•ǡ„‡‡•ƒ”‡—†‘—„–‡†Ž›–Š‡‘•–‹’‘”–ƒ–ǡ „‘–Š‹–‡”•‘ˆˆŽ‘™‡”˜‹•‹–ƒ–‹‘”ƒ–‡•ƒ†’‘ŽŽ‹ƒ–‹‰‡ˆˆ‹ ‹‡ ›ǡ„‡‹‰–Š‡ƒ‹’‘ŽŽ‹ƒ–‘”• ‹‘•–‡ ‘•›•–‡•Ǥ‘—‰ŠŽ›ǡʹͲǡͲͲͲ„‡‡•’‡ ‹‡•ƒ”‡ —””‡–Ž›†‡• ”‹„‡†™‘”Ž†™‹†‡ȋ• Š‡” ƒ†‹ ‡”‹‰ʹͲͳͳȌǤŠ‡›ƒ”‡’”‡•‡–‹‡˜‡”› ‘–‹‡–‡š ‡’––ƒ”–‹ ƒǡ‹ƒ™‹†‡”ƒ‰‡ ‘ˆ†‹ˆˆ‡”‡–Šƒ„‹–ƒ–•ȋ‹ Š‡‡”ʹͲͲͲȌǤ•’‘ŽŽ‹ƒ–‘”•ǡ–Š‡›’”‘˜‹†‡ƒ‡••‡–‹ƒŽ‡ ‘•›•–‡ •‡”˜‹ ‡ȋ‘•‡›ƒ†ƒ—‰ŠƒʹͲͲ͸ȌǤ‡‡•’‡ ‹‡•†‹˜‡”•‹–›‹•–Š‡”‡ˆ‘”‡ ”— ‹ƒŽˆ‘”‡ ‘•›•–‡ ˆ— –‹‘‹‰ǡ„‘–Š‹ƒ–—”ƒŽƒ†ƒ‰”‹ —Ž–—”ƒŽ•›•–‡•ǤǤ

—”˜‡›•„ƒ•‡†‘Š‹•–‘”‹ ƒŽ†ƒ–ƒ†‡‘•–”ƒ–‡–Šƒ–„‡‡†‹˜‡”•‹–›‹•†‡ Ž‹‹‰‹•‡˜‡”ƒŽ’ƒ”–• ‘ˆ–Š‡™‘”Ž†ȋ‹‡•‡‹Œ‡”‡–ƒŽǤʹͲͲ͸ǡ‘––•‡–ƒŽǤʹͲͳͲǡʹͲͳͶǡƒ‡”‘‡–ƒŽǤʹͲͳͳȌǤ —”‘’‡ǡ –Š‡”‡ ƒ”‡ ƒŽ‘•– ʹǡͲͲͲ ™‹Ž† „‡‡ •’‡ ‹‡• ͻǤʹΨ ‘ˆ ™Š‹ Š ƒ”‡ ‘™ –‘ „‡ –Š”‡ƒ–‡‡† ™‹–Š ‡š–‹ –‹‘ǡƒ†ͷ͸Ǥ͹Ψ ƒ‘–„‡‡˜ƒŽ—ƒ–‡††—‡–‘Žƒ ‘ˆ‘™Ž‡†‰‡ƒ†ˆ—†‹‰ȋ‹‡–‘‡–ƒŽǤ ʹͲͳͶȌǤŠ—•ǡ–Š‡’‘’—Žƒ–‹‘•–ƒ–—•ƒ†˜‹ƒ„‹Ž‹–›‘ˆ–Š‡˜ƒ•–ƒŒ‘”‹–›‘ˆ„‡‡•’‡ ‹‡•‹—”‘’‡ ƒ”‡Žƒ”‰‡Ž›—‘™Ǥ

‡‡ ‘—‹–‹‡•Šƒ˜‡ƒ ‘„‹ƒ–‹‘‘ˆ Šƒ”ƒ –‡”‹•–‹ •–Šƒ–ƒ‡–Š‡‹†‡ƒŽˆ‘”–Š‡•–—†›‘ˆ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ„‹Ž‹–› ‘ˆ ‘—‹–‹‡• ƒ† –Š‡‹” ‹–‡”ƒ –‹‘•Ǥ ‹”•–‘ˆƒŽŽǡ„‡‡•Šƒ˜‡˜‡”› ‘–”ƒ•–‡† ˆ— –‹‘ƒŽ –”ƒ‹–•ǡ •— Š ƒ• „‘†› •‹œ‡ǡ ’”‘„‘• ‹• Ž‡‰–Šǡ ƒ† •‘ ‹ƒŽ‹–› ȋ‹ Š‡‡” ʹͲͲͲȌǤ‹ˆˆ‡”‡ ‡•‹„‘†›•‹œ‡‹’Ž›†‹ˆˆ‡”‡ ‡•‹‡‡”‰‡–‹ ”‡“—‹”‡‡–•ǡˆ‘”ƒ‰‹‰”ƒ‰‡• ƒ††‹•’‡”•ƒŽƒ„‹Ž‹–›ǡƒ††‹ˆˆ‡”‡ ‡•‹’”‘„‘• ‹•Ž‡‰–Š‹’Ž›†‹ˆˆ‡”‡ ‡•‹ƒ ‡••‹„‹Ž‹–›–‘ †‹ˆˆ‡”‡––›’‡•‘ˆˆŽ‘™‡”•Ǥ‹•’ƒ”‹–›‹•‘ ‹ƒŽ‹–›‹”‡Žƒ–‹‘–‘ ‘Ž‘› •‹œ‡ ƒ† –Š‡”‘”‡‰—Žƒ–‹‘ ƒŽ•‘ ‹’Ž› ‹’‘”–ƒ– †‹ˆˆ‡”‡ ‡• ‹ ‡‡”‰‡–‹  ”‡“—‹”‡‡–•Ǥ  ‡‡ ‘—‹–‹‡•‹ Ž—†‡˜ƒ”‹‘—•Ž‡˜‡Ž•‘ˆˆ‡‡†‹‰•’‡ ‹ƒŽ‹œƒ–‹‘ǡˆ”‘ƒ””‘™’‘ŽŽ‡•’‡ ‹ƒŽ‹•–• –‘ ™‹†‡ ‰‡‡”ƒŽ‹•–•Ǥ ‹ƒŽŽ›ǡ „‡‡ ‘—‹–‹‡• †‹•’Žƒ› ƒ ”ƒ‰‡ ‘ˆ ‡•–‹‰ Šƒ„‹–•ǡ ‹ Ž—†‹‰ ‰”‘—†‡•–‡”•ǡ ƒ˜‹–›‡•–‡”•ƒ†•’‡ ‹‡•„—‹Ž†‹‰‡š’‘•‡†‡•–•Ǥ ƒ††‹–‹‘ǡ•‘‡•’‡ ‹‡• ”‡“—‹”‡ ‡š–‡”ƒŽ ƒ–‡”‹ƒŽ•ǡ ‘ˆ–‡ ‘ˆ ’Žƒ– ‘”‹‰‹ǡ –‘ „—‹Ž† –Š‡‹” ‡•–•Ǥ Š—•ǡ †‹ˆˆ‡”‡– „‡‡ •’‡ ‹‡• ™‹–Š‹ ƒ ‘—‹–› ƒ”‡ Ž‹‡Ž› –‘ „‡ •‡•‹–‹˜‡ –‘ †‹ˆˆ‡”‡– ˆƒ –‘”•ǡ ‘” ”‡•’‘† †‹ˆˆ‡”‡–Ž›–‘–Š‡•ƒ‡ˆƒ –‘”•ȋ‡•–”‹ Šͳͻͻ͸Ȍǡƒ†–‘‹–‡”ƒ –™‹–Š†‹ˆˆ‡”‡–ˆŽ‘™‡”•’‡ ‹‡• ƒ ‘”†‹‰–‘–Š‡‹”ˆ— –‹‘ƒŽ–”ƒ‹–•Ǥ

ƒ††‹–‹‘–‘‹–‡”ƒ –‹‰™‹–ŠˆŽ‘™‡”•ǡ„‡‡•ƒŽ•‘‹–‡”ƒ –™‹–Šƒ˜ƒ”‹‡–›‘ˆƒ–—”ƒŽ‡‡‹‡•ǡ ‹ Ž—†‹‰’”‡†ƒ–‘”•ǡ’ƒ”ƒ•‹–‘‹†•ǡ Ž‡’–‘’ƒ”ƒ•‹–‡•ƒ†‡•–• ƒ˜ƒ‰‡”•ǤŠ‡•‡˜ƒ”‹‘—•‰”‘—’•  11 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•   

‘ˆ‡‡‹‡•ƒ›†‹ˆˆ‡”‹–Š‡‹”Ž‡˜‡Ž‘ˆ•’‡ ‹ƒŽ‹œƒ–‹‘ǡƒ†ƒ›”‡•’‘†–‘ƒ•—‹–‡‘ˆ ƒ††‹–‹‘ƒŽˆƒ –‘”•‹ƒ††‹–‹‘–‘„‡‡ ‘—‹–›•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘Ǥ



Objectivesofthisthesis

Š‹•†‹••‡”–ƒ–‹‘•–—†‹‡•–Š‡ˆƒ –‘”•–Šƒ–‹ˆŽ—‡ ‡–Š‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆƒ„‡‡ ‘—‹–› ƒ† ‹–• ‹–‡”ƒ –‹‘ ‡–™‘” ™‹–Š ƒ–—”ƒŽ ‡‡‹‡• ‹ –Š‡ ƒ””ƒˆ ƒ–—”ƒŽ ƒ” ȋƒ” ‡Ž‘ƒȌǤ Š‡•–—†›ƒ”‡ƒ‹•ƒ ‘–‹—‘—•‡†‹–‡””ƒ‡ƒ• ”—„Žƒ†ǡ™‹–Š˜‡”›•‹‹Žƒ”˜‡‰‡–ƒ–‹‘–›’‡ǡ ‰‡‘Ž‘‰›ƒ†’‡”–—”„ƒ–‹‘Š‹•–‘”›ǤŠ‡Žƒ ‘ˆ’Š›•‹ ƒŽ‘”‡ ‘Ž‘‰‹ ƒŽ„ƒ””‹‡”•ƒ†‰”ƒ†‹‡–•‹ –Š‡•–—†›ƒ”‡ƒƒˆˆ‘”†•—•™‹–Šƒ‘’’‘”–—‹–›–‘ƒƒŽ›œ‡–Š‡‹–”‹•‹ •’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ –Š‡ ‘—‹–›ƒ†‹–•‹–‡”ƒ –‹‘‡–™‘”™‹–Š‘—–‘‹•‡ˆƒ –‘”•†—‡–‘ Šƒ‰‡•‹Šƒ„‹–ƒ– –›’‡Ǥ Š‡ ƒ‹ ‘„Œ‡ –‹˜‡ ‘ˆ –Š‹• –Š‡•‹• ‹• –‘ ‹˜‡•–‹‰ƒ–‡ Š‘™ Ž‘ ƒŽ ”‡•‘—” ‡• ƒ† Š—ƒ ’‡”–—”„ƒ–‹‘ƒ›ƒˆˆ‡ ––Š‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆƒ„‡‡ ‘—‹–›ƒ–ƒŽ‘ ƒŽ• ƒŽ‡ǡƒ†Š‘™ ‘—‹–›•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‹ˆŽ—‡ ‡•–Š‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ‹–•‹–‡”ƒ –‹‘‡–™‘”Ǥ

Chapter1ƒƒŽ›•‡•–Š‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆƒ™‹Ž†„‡‡ ‘—‹–›ƒ–ƒŽ‘ ƒŽ• ƒŽ‡ǡƒ†Š‘™ –Š‡ ‘—‹–›”‡•’‘†•–‘–Š‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆˆŽ‘”ƒŽƒ†‡•–‹‰”‡•‘—” ‡•Ǥ‡ ƒ—•‡ „‡‡•†‡’‡†‘„‘–Š”‡•‘—” ‡–›’‡•–‘•—”˜‹˜‡ƒ†”‡’”‘†— ‡ǡ•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆ„‡‡•‹• ‡š’‡ –‡†–‘ˆ‘ŽŽ‘™•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆˆŽ‘™‡”•ƒ†Ȁ‘”‡•–‹‰•‹–‡•ȋ‹ Ž‡ˆ•ͳͻͺ͹ȌǤ –Šƒ• „‡‡™‡ŽŽ‡•–ƒ„Ž‹•Š‡†–Šƒ–ˆ‘”ƒ‰‹‰”ƒ‰‡•‘ˆ„‡‡•ƒ”‡†‹”‡ –Ž›”‡Žƒ–‡†–‘•’‡ ‹‡•„‘†›•‹œ‡ ȋ ƒ–Šƒƒ†• Šƒ”–‡ʹͲͲʹǡ ”‡‡Ž‡ƒˆ‡–ƒŽǤʹͲͲ͹ǡ —±†‘–‡–ƒŽǤʹͲͲͻǡ—”„— Š‡‡–ƒŽǤ ʹͲͳͲȌǤŠ—•ǡ™‡ƒ††‹–‹‘ƒŽŽ›‡š’Ž‘”‡–Š‡”‘Ž‡‘ˆ„‘†›•‹œ‡‘–Š‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆ–Š‡ „‡‡ ‘—‹–›ǤŠ‹Ž‡Žƒ†• ƒ’‡Ǧ• ƒŽ‡ˆƒ –‘”•ƒˆˆ‡ –‹‰„‡‡ ‘—‹–‹‡•Šƒ˜‡„‡‡ƒ’Ž› •–—†‹‡†ǡŽ‹––Ž‡‹•‘™ƒ„‘—–Ž‘ ƒŽǦ• ƒŽ‡ˆƒ –‘”•‹ˆŽ—‡ ‹‰–Š‡Ǥ Š‡”‡ˆ‘”‡ǡ –Š‡ ˆ‹”•– Šƒ’–‡”‘ˆ–Š‹•–Š‡•‹•‹•‹–‡†‡†–‘‡š’Ž‘”‡•— Š”‡Žƒ–‹‘•Š‹’•™‹–Š–Š”‡‡•’‡ ‹ˆ‹ ‘„Œ‡ –‹˜‡•ǣ ȋͳȌ‘ƒƒŽ›œ‡ˆŽ‘”ƒŽƒ†‡•–‹‰”‡•‘—” ‡ ‘’‘•‹–‹‘Š‡–‡”‘‰‡‡‹–›ƒ––Š‡Šƒ„‹–ƒ–• ƒŽ‡ǡ ȋʹȌ–‘ƒƒŽ›œ‡–Š‡†‹•–”‹„—–‹‘‘ˆ„‡‡ ‘’‘•‹–‹‘ƒ ”‘••–Š‡Šƒ„‹–ƒ–ƒ†ȋ͵Ȍ–‘‡•–ƒ„Ž‹•Š ™Š‡–Š‡”•’‡ ‹‡•™‹–Š†‹ˆˆ‡”‡–„‘†›•‹œ‡•”‡•’‘††‹ˆˆ‡”‡–Ž›–‘Ž‘ ƒŽ”‡•‘—” ‡ƒ˜ƒ‹Žƒ„‹Ž‹–› ƒ†•Š‘™†‹ˆˆ‡”‡–’ƒ––‡”•‘ˆ•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘Ǥ

ƒ˜‹‰•‡‡–Š‡‡ˆˆ‡ –‘ˆˆŽ‘™‡”ƒ†‡•–‹‰”‡•‘—” ‡•‘„‡‡ ‘—‹–› ‘’‘•‹–‹‘ƒ† •–”— –—”‡ǡChapter2•–—†‹‡•–Š‡”‡•’‘•‡‘ˆ–Š‡•ƒ‡ ‘—‹–›–‘ƒŠ—ƒ’‡”–—”„ƒ–‹‘ǡ ƒ‡Ž› „‡‡‡‡’‹‰Ǥ ‡‡‡‡’‹‰ ƒ –‹˜‹–‹‡• ”‡•—Ž– ‹ –Š‡ ‹–”‘†— –‹‘ ‘ˆ Žƒ”‰‡ ƒ‘—–• ‘ˆ

 12   ‡‡”ƒŽ‹–”‘†— –‹‘

Š‘‡›„‡‡ȋApismellifera)‹†‹˜‹†—ƒŽ™‘”‡”•ˆ‘”ƒ‰‹‰ˆ‘”‡ –ƒ”ƒ†’‘ŽŽ‡Ǥ—‡–‘–Š‡‹” Š‹‰ŠŽ› ‡ˆˆ‹ ‹‡– ˆ‘”ƒ‰‹‰ „‡Šƒ˜‹‘” ȋ˜‘ ”‹• Š ͳͻ͸͹ǡ ‡‡Ž‡› ͳͻͺͷǡ‹ Š–‡”ƒ†‡”ƒƒ–› ʹͲͲ͵Ȍǡ ƒƒ‰‡† Š‘‡› „‡‡• ƒ› ’‘•‡ ƒ ’”‘„Ž‡ –‘ –Š‡ ƒ–‹˜‡ „‡‡ ‘—‹–› ™Š‡ ”‡•‘—” ‡•ƒ”‡• ƒ” ‡ǡŽ‘™‡”‹‰’‘ŽŽ‡ƒ†‡ –ƒ”ƒ˜ƒ‹Žƒ„‹Ž‹–›Ǥ‹Ž†„‡‡•’‡ ‹‡•‘ˆŽƒ”‰‡„‘†› •‹œ‡ƒ›„‡–Š‡‘•–ƒˆˆ‡ –‡††—‡–‘–Š‡‹”Š‹‰Š‡”ˆ‡‡†‹‰”‡“—‹”‡‡–•ȋòŽŽ‡”‡–ƒŽǤʹͲͲ͸ȌǤ Šƒ’–‡”ʹǡ™‡‡ƒ•—”‡’‘ŽŽ‡Ǧ‡ –ƒ” ‘•—’–‹‘‹’Ž‘–•’”‘‰”‡••‹˜‡Ž›†‹•–ƒ–ˆ”‘ ƒ’‹ƒ”‹‡• ƒ† ”‡Žƒ–‡ –Š‹• ‘•—’–‹‘ –‘ Š‘‡› „‡‡ ƒ† ™‹Ž† „‡‡ ˆ‘”ƒ‰‹‰ ƒ –‹˜‹–›Ǥ —” ‘„Œ‡ –‹˜‡‹•–‘—†‡”•–ƒ†–Š‡‹’ƒ –‘ˆŠ‘‡›„‡‡ˆŽ‘™‡”˜‹•‹–ƒ–‹‘‘’‘ŽŽ‡ƒ†‡ –ƒ” ‘•—’–‹‘ƒ†–Š‡‡ˆˆ‡ –‘ˆ„‡‡‡‡’‹‰‘–Š‡ƒ„—†ƒ ‡ǡ”‹ Š‡••ƒ† ‘’‘•‹–‹‘‘ˆ –Š‡Ž‘ ƒŽ™‹Ž†„‡‡ ‘—‹–›Ǥ

‹ƒŽŽ›ǡ Chapter 3•–—†‹‡•–Š‡Ž‘ ƒŽ•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ–Š‡ ƒ˜‹–›Ǧ‡•–‹‰„‡‡ ƒ† ™ƒ•’ ‘—‹–›ǡ –Š‡‹” ‡•–Ǧƒ••‘ ‹ƒ–‡† ˆƒ—ƒ ȋ’ƒ”ƒ•‹–‘‹†•ǡ Ž‡’–‘’ƒ”ƒ•‹–‡•ǡ ’”‡†ƒ–‘”• ƒ† • ƒ˜‡‰‡”•Ȍǡƒ†–Š‡”‡•—Ž–‹‰‹–‡”ƒ –‹‘‡–™‘”ǤŠ‹• Šƒ’–‡” ƒŽ•‘ ƒ††”‡••‡• –Š‡ ”‡Žƒ–‹‘•Š‹’ „‡–™‡‡ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ–‹‘ ‘ˆ –Š‡ –™‘ –”‘’Š‹  Ž‡˜‡Ž• ƒ† ™‹–Š –Š‡ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ–‹‘‘ˆ–Š‡‹–‡”ƒ –‹‘•Ǥ’‡ ‹‡•EǦ†‹˜‡”•‹–›’ƒ––‡”• ƒ„‡†‹˜‹†‡†‹–‘–™‘ƒ††‹–‹˜‡ ‘’‘‡–•ǣ•’‡ ‹‡•–—”‘˜‡”ƒ†•’‡ ‹‡•Ž‘••ȋƒ•‡Ž‰ƒʹͲͳͲȌǤƒŽ‘‰‘—•Ž›ǡ‹–‡”ƒ –‹‘EǦ †‹˜‡”•‹–›’ƒ––‡”•ƒ›„‡†‹˜‹†‡†‹–‘ƒ ‘’‘‡–‘ˆ•’‡ ‹‡•–—”‘˜‡”ƒ†ƒ ‘’‘‡–‘ˆ ”‡™‹”‹‰ ȋ•ƒ‡ •’‡ ‹‡• ‹–‡”ƒ –‹‰ †‹ˆˆ‡”‡–Ž› ‹ †‹ˆˆ‡”‡– Ž‘ ƒŽ‹–‹‡•Ȍ ȋ‘‹•‘– ‡– ƒŽǤ ʹͲͳʹȌǤ Š‹•Žƒ•– Šƒ’–‡”‡˜ƒŽ—ƒ–‡•–Š‡•‡•‘—” ‡•‘ˆ˜ƒ”‹ƒ–‹‘ƒ†–Š‡‹” ‘–”‹„—–‹‘–‘–Š‡‘„•‡”˜‡† •’ƒ–‹ƒŽ ’ƒ––‡”•Ǥ  ’ƒ”–‹ —Žƒ”ǡ ™‡ Šƒ˜‡ –Š‡ ˆ‘ŽŽ‘™‹‰ ‘„Œ‡ –‹˜‡• ȋͳȌ –‘ ƒƒŽ›œ‡ –Š‡ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ–‹‘‘ˆƒ ‘—‹–›‘ˆ ƒ˜‹–›Ǧ‡•–‹‰„‡‡•ƒ†™ƒ•’•ȋŠ‡ ‡ˆ‘”–ŠŠ‘•–•Ȍƒ†–Š‡‹”‡•– ƒ••‘ ‹ƒ–‡•ȋŠ‡ ‡ˆ‘”–Š’ƒ”ƒ•‹–‘‹†•Ȍƒ ”‘••ƒ ‘–‹—‘—•Šƒ„‹–ƒ–ǡȋʹȌ–‘•–—†›EǦ†‹˜‡”•‹–›‘ˆ Š‘•–Ǧ’ƒ”ƒ•‹–‘‹†‹–‡”ƒ –‹‘•ƒ†ȋ͵Ȍ–‘‡š’Ž‘”‡–Š‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡Š‘•–ǡ’ƒ”ƒ•‹–‘‹†ƒ† ‹–‡”ƒ –‹‘EǦ†‹˜‡”•‹–›ƒ†–‘‡šƒ‹‡–Š‡‹”†‹•–ƒ ‡†‡ ƒ›Ǥ



References‰”ƒ™ƒŽǡǤǤ‡–ƒŽǤʹͲͲ͸Ǥ ‘—‹–›Š‡–‡”‘‰‡‡‹–›ƒ†–Š‡‡˜‘Ž—–‹‘‘ˆ‹–‡”ƒ –‹‘• „‡–™‡‡’Žƒ–•ƒ†‹•‡ –Š‡”„‹˜‘”‡•ǤǦǤ‡˜Ǥ‹‘ŽǤͺͳǣ͵ͶͻȂ͵͹͸Ǥ

Ž„”‡ Š–ǡ Ǥ ‡– ƒŽǤ ʹͲͲ͹Ǥ –‡”ƒ –‹‘ †‹˜‡”•‹–› ™‹–Š‹ “—ƒ–‹ˆ‹‡† ‹•‡ – ˆ‘‘† ™‡„• ‹ ”‡•–‘”‡† ƒ† ƒ†Œƒ ‡–‹–‡•‹˜‡Ž›ƒƒ‰‡†‡ƒ†‘™•ǤǦ Ǥ‹Ǥ ‘ŽǤ͹͸ǣͳͲͳͷȂͳͲʹͷǤ

 13 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•   

”ƒǡǤ‡–ƒŽǤʹͲͲ͸Ǥ‘•–Ǧˆ‹”‡”‡ ‘˜‡”›‘ˆ‡†‹–‡””ƒ‡ƒ‰”‘—†ƒ– ‘—‹–‹‡•ˆ‘ŽŽ‘™•˜‡‰‡–ƒ–‹‘ ƒ††”›‡••‰”ƒ†‹‡–•ǤǦ Ǥ‹‘‰‡‘‰”Ǥ͵͵ǣͳʹͶ͸ȂͳʹͷͺǤ

• Š‡”ǡ Ǥ Ǥ ƒ† ‹ ‡”‹‰ǡ Ǥ ʹͲͳͳǤ ‡‡ •’‡ ‹‡• ‰—‹†‡ ȋ ›‡‘’–‡”ƒǣ ’‘‹†‡ƒǣ –Š‘’Š‘”ƒȌǤ Ǧ ‹• ‘˜‡”‹ˆ‡ǣ™™™Ǥ†‹• ‘˜‡”Ž‹ˆ‡Ǥ‘”‰Ǥ

ƒŽ†‹ǡ Ǥ ‡– ƒŽǤ ʹͲͲͶǤ ‹‰Š ’‘–‡–‹ƒŽ ˆ‘” ‘’‡–‹–‹‘ „‡–™‡‡ ‰—ƒƒ ‘•ƒ†•Š‡‡’‹ƒ–ƒ‰‘‹ƒǤǦ Ǥ ‹Ž†ŽǤƒƒ‰‡Ǥ͸ͺǣͻʹͶȂͻ͵ͺǤ

ƒ”–‘‡—•ǡ Ǥ ‡– ƒŽǤ ʹͲͲͺǤ ‘–”ƒ•–‹‰ ‡ˆˆ‡ –• ‘ˆ ‹˜ƒ•‹˜‡ ’Žƒ–• ‹ ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘” ‡–™‘”•Ǥ Ǧ ‡ ‘Ž‘‰‹ƒͳͷͷǣ͹͸ͳȂ͹͹ͲǤ

ƒ•‡Ž‰ƒǡǤʹͲͳͲǤƒ”–‹–‹‘‹‰–Š‡–—”‘˜‡”ƒ†‡•–‡†‡•• ‘’‘‡–•‘ˆ„‡–ƒ†‹˜‡”•‹–›ǤǦ Ž‘„Ǥ ‘ŽǤ ‹‘‰‡‘‰”Ǥͳͻǣͳ͵ͶȂͳͶ͵Ǥ

‡‰‘ǡǤ‡–ƒŽǤʹͲͲ͸Ǥ ‘Ž‘‰›ǣˆ”‘‹†‹˜‹†—ƒŽ•–‘‡ ‘•›•–‡•ǤǦŽƒ ™‡ŽŽ—„Ž‹•Š‹‰Ǥ

‡”‰ǡ%Ǥͳͻͻ͹Ǥ‹˜‡”•‹–›ƒ†ƒ„—†ƒ ‡‘ˆ„‹”†•‹”‡Žƒ–‹‘–‘ˆ‘”‡•–ˆ”ƒ‰‡–ƒ–‹‘ǡŠƒ„‹–ƒ–“—ƒŽ‹–› ƒ†Š‡–‡”‘‰‡‡‹–›ǤǦ‹”†–—†›ͶͶǣ͵ͷͷȂ͵͸͸Ǥ

‹‡•‡‹Œ‡”ǡ ǤǤ‡–ƒŽǤʹͲͲ͸Ǥƒ”ƒŽŽ‡Ž†‡ Ž‹‡•‹’‘ŽŽ‹ƒ–‘”•ƒ†‹•‡ –Ǧ’‘ŽŽ‹ƒ–‡†’Žƒ–•‹”‹–ƒ‹ƒ† –Š‡‡–Š‡”Žƒ†•ǤǦ ‹‡ ‡͵ͳ͵ǣ͵ͷͳȂͶǤ

Žƒ‹”ǡǤʹͲͲͳǤ‹”†•ƒ†„—––‡”ˆŽ‹‡•ƒŽ‘‰—”„ƒ‰”ƒ†‹‡–•‹–™‘‡ ‘”‡‰‹‘•‘ˆ–Š‡‹–‡†–ƒ–‡•ǣ‹• —”„ƒ‹œƒ–‹‘ ”‡ƒ–‹‰ƒŠ‘‘‰‡‡‘—•ˆƒ—ƒǫǦ ǣ‹‘–‹ Š‘‘‰‡‹œƒ–‹‘Ǥ’‹‰‡”ǡ’’Ǥ͵͵Ȃͷ͸Ǥ

Žò–Š‰‡ǡ Ǥ ‡– ƒŽǤ ʹͲͲͺǤ Šƒ– †‘ ‹–‡”ƒ –‹‘ ‡–™‘” ‡–”‹ • –‡ŽŽ —• ƒ„‘—– •’‡ ‹ƒŽ‹œƒ–‹‘ ƒ† „‹‘Ž‘‰‹ ƒŽ–”ƒ‹–•ǫǦ ‘Ž‘‰›ͺͻǣ͵͵ͺ͹ȂͻͻǤ

‘†ǡǤ Ǥƒ†‡‡Ž‡›ǡ ǤǤʹͲͲͷǤ ‹”‡ƒ•ƒ‰Ž‘„ƒŽDzŠ‡”„‹˜‘”‡dzǣŠ‡‡ ‘Ž‘‰›ƒ†‡˜‘Ž—–‹‘‘ˆˆŽƒƒ„Ž‡ ‡ ‘•›•–‡•ǤǦ”‡†• ‘ŽǤ˜‘ŽǤʹͲǣ͵ͺ͹Ȃ͵ͻͶǤ

‘••‹‘ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͷǤ ‘‹Ž ‹ ”‘„‹ƒŽ ‘—‹–› ”‡•’‘•‡ –‘ Žƒ† —•‡ Šƒ‰‡ ‹ ƒ ƒ‰”‹ —Ž–—”ƒŽ Žƒ†• ƒ’‡‘ˆ™‡•–‡”‡›ƒǤǦ‹ ”‘„Ǥ ‘ŽǤͶͻǣͷͲȂ͸ʹǤ

—ŽŽ‘ ǡ ǤǤ‡–ƒŽǤʹͲͲʹǤŽƒ–†‹•’‡”•ƒŽƒ† ‘Ž‘‹œƒ–‹‘’”‘ ‡••‡•ƒ–Ž‘ ƒŽƒ†Žƒ†• ƒ’‡• ƒŽ‡•ǤǦ ǣ ‹•’‡”•ƒŽ ‘Ž‘‰›ǣͶʹ†›’‘•‹—‘ˆ–Š‡”‹–‹•Š ‘Ž‘‰‹ ƒŽ‘ ‹‡–›Ǥ’’Ǥʹ͹ͻȂ͵ͲʹǤ

 14   ‡‡”ƒŽ‹–”‘†— –‹‘

—”Ž‡ǡǤǤƒ†Žƒ” אǡǤʹͲͳͳǤŠ‡ˆ—–—”‡‘ˆ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘”†‹˜‡”•‹–›ǣ—†‡”•–ƒ†‹‰‹–‡”ƒ –‹‘ ‡–™‘”•ƒ ”‘••–‹‡ǡ•’ƒ ‡ǡƒ†‰Ž‘„ƒŽ Šƒ‰‡ǤǦǤ Ǥ‘–ǤͻͺǣͷʹͺȂ͵ͺǤ

ƒ†‘––‡ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͳͳǤ ‡›‘† •’‡ ‹‡•ǣ — –‹‘ƒŽ †‹˜‡”•‹–› ƒ† –Š‡ ƒ‹–‡ƒ ‡ ‘ˆ ‡ ‘Ž‘‰‹ ƒŽ ’”‘ ‡••‡•ƒ†•‡”˜‹ ‡•ǤǦ Ǥ’’ŽǤ ‘ŽǤͶͺǣͳͲ͹ͻȂͳͲͺ͹Ǥ

ƒ‡”‘ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͳͳǤ ƒ––‡”• ‘ˆ ™‹†‡•’”‡ƒ† †‡ Ž‹‡ ‹ ‘”–Š‡”‹ ƒ „—„Ž‡ „‡‡•Ǥ Ǧ ”‘ Ǥ ƒ–ŽǤ ƒ†Ǥ ‹ǤǤǤǤͳͲͺǣ͸͸ʹȂ͸͸͹Ǥ

ƒ”†‘•‘ǡǤ Ǥ‡–ƒŽǤʹͲͲͺǤŠ‡‹’ƒ –‘ˆ‡š–”‡‡ˆŽ‘‘†‹‰‡˜‡–•ƒ†ƒ–Š”‘’‘‰‡‹ •–”‡••‘”•‘–Š‡ ƒ ”‘„‡–Š‹  ‘—‹–‹‡•ǯ†›ƒ‹ •ǤǦ•–—ƒ”Ǥ‘ƒ•–ǤŠ‡Žˆ ‹Ǥ͹͸ǣͷͷ͵Ȃͷ͸ͷǤ

Š‡ǡ ǤǦǤ‡–ƒŽǤʹͲͳͳǤƒ’‹†”ƒ‰‡•Š‹ˆ–•‘ˆ•’‡ ‹‡•‘ˆ Ž‹ƒ–‡™ƒ”‹‰ǤǦ ‹‡ ‡͵͵͵ǣͳͲʹͶȂͳͲʹ͸Ǥ

‘†‹–ǡǤ‡–ƒŽǤʹͲͲʹǤ‡–ƒǦ†‹˜‡”•‹–›‹–”‘’‹ ƒŽˆ‘”‡•––”‡‡•ǤǦ ‹‡ ‡ʹͻͷǣ͸͸͸Ȃ͸͸ͻǤ

ƒ”™‹ǡ Ǥ ͳͺͷͻǤ  –Š‡ ‘”‹‰‹ ‘ˆ •’‡ ‹‡• „› ‡ƒ• ‘ˆ ƒ–—”ƒŽ •‡Ž‡ –‹‘ǡ ‘” –Š‡ ’”‡•‡”˜ƒ–‹‘ ‘ˆ ˆƒ˜‘—”‡†”ƒ ‡•‹–Š‡•–”—‰‰Ž‡ˆ‘”Ž‹ˆ‡ǤǦ ‘Š—””ƒ›Ǥ

ǃœǡƴ Ǥ ƒ† ƒ„‹†‘ǡ Ǥ ʹͲͲͳǤ ‹˜‡ Žƒ †‹ˆˆ±”‡ ‡ǣ ’Žƒ– ˆ— –‹‘ƒŽ †‹˜‡”•‹–› ƒ––‡”• –‘ ‡ ‘•›•–‡ ’”‘ ‡••‡•ǤǦ”‡†• ‘ŽǤ˜‘ŽǤͳ͸ǣ͸Ͷ͸Ȃ͸ͷͷǤ

‘™‹‰ǡ Ǥ Ǥ ‡– ƒŽǤ ͳͻͻͻǤ Š‡ ‹’ƒ – ‘ˆ ƒ ‡Ž‡”ƒ–‹‰ Žƒ†Ǧ—•‡ Šƒ‰‡‘–Š‡Ǧ› Ž‡‘ˆ–”‘’‹ ƒŽ ƒ“—ƒ–‹ ‡ ‘•›•–‡•ǣ—””‡– ‘†‹–‹‘•ƒ†’”‘Œ‡ –‡† Šƒ‰‡•ǤǦ‹‘‰‡‘ Š‡‹•–”›Ͷ͸ǣͳͲͻȂͳͶͺǤ

Š”‡ˆ‡Ž†ǡ Ǥ ǤʹͲͳͲǤ ‘•›•–‡ ‘•‡“—‡ ‡•‘ˆ„‹‘Ž‘‰‹ ƒŽ‹˜ƒ•‹‘•ǤǦ—Ǥ‡˜Ǥ ‘ŽǤ˜‘ŽǤ›•–ǤͶͳǣ ͷͻȂͺͲǤ

•“—‡ǡǤǤ‡–ƒŽǤʹͲͲ͵Ǥˆˆ‡ –•‘ˆ†‡•‡”–™‹Ž†ˆ‹”‡•‘†‡•‡”–”‘”–‘‹•‡ȋGopherusagassiziiȌƒ†‘–Š‡” •ƒŽŽ˜‡”–‡„”ƒ–‡•ǤǦ‘—–Š™‡•–Ǥƒ–ǤͶͺǣͳͲ͵ȂͳͳͳǤ

‘–ƒ‹‡ǡ Ǥ ‡– ƒŽǤ ʹͲͲ͸Ǥ — –‹‘ƒŽ †‹˜‡”•‹–› ‘ˆ ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘” ‹–‡”ƒ –‹‘ ™‡„• ‡Šƒ ‡• –Š‡ ’‡”•‹•–‡ ‡‘ˆ’Žƒ– ‘—‹–‹‡•ǤǦŽ‘‹‘ŽǤͶǣͳʹͻȂ͵ͷǤ

—ƒ‹ǡǤʹͲͳͲǤ‘—‹–›ƒ••‡„Ž›†›ƒ‹ •‹•’ƒ ‡ǤǦ ǣ‡”Š‘‡ˆǡ ǤǤƒ†‘”‹ǡǤ Ǥȋ‡†•Ȍǡ ‘—‹–›‡ ‘Ž‘‰›ǣ’”‘ ‡••‡•ǡ‘†‡Ž•ǡƒ†ƒ’’Ž‹ ƒ–‹‘•Ǥšˆ‘”†‹˜‡”•‹–›”‡••ǡ’’ǤͶͷȂͷͶǤ

ƒ‰‹ ǡǤ‡–ƒŽǤʹͲͳͷǤ — –‹‘ƒŽ‹†‡–‹–›ƒ††‹˜‡”•‹–›‘ˆƒ‹ƒŽ•’”‡†‹ –‡ ‘•›•–‡ˆ— –‹‘‹‰„‡––‡” –Šƒ•’‡ ‹‡•Ǧ„ƒ•‡†‹†‹ ‡•ǤǦ”‘ ǤǤ‘ Ǥ‹‘ŽǤ ‹ǤʹͺʹǣͳȂͺǤ

 15 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•   

ƒ–ŠƒǡǤƒ†• Šƒ”–‡ǡǤʹͲͲʹǤ ‘”ƒ‰‹‰”ƒ‰‡•‘ˆ•‘Ž‹–ƒ”›„‡‡•ǤǦ Ǥ‹Ǥ ‘ŽǤ͹ͳǣ͹ͷ͹Ȃ͹͸ͶǤ

”‡‡ǡ ǤǤ‡–ƒŽǤʹͲͲͺǤ‹ ”‘„‹ƒŽ„‹‘‰‡‘‰”ƒ’Š›ǣˆ”‘–ƒš‘‘›–‘–”ƒ‹–•ǤǦ ‹‡ ‡͵ʹͲǣͳͲ͵ͻȂͳͲͶ͵Ǥ

”‡‡Ž‡ƒˆǡǤǤ‡–ƒŽǤʹͲͲ͹Ǥ‡‡ˆ‘”ƒ‰‹‰”ƒ‰‡•ƒ†–Š‡‹””‡Žƒ–‹‘•Š‹’–‘„‘†›•‹œ‡ǤǦ‡ ‘Ž‘‰‹ƒͳͷ͵ǣ ͷͺͻȂͻ͸Ǥ

”‘•Š‘ŽœǡǤʹͲͲʹǤ ‘Ž‘‰‹ ƒŽƒ†‡˜‘Ž—–‹‘ƒ”› ‘•‡“—‡ ‡•‘ˆ ‘ƒ•–ƒŽ‹˜ƒ•‹‘•ǤǦ”‡†• ‘ŽǤ˜‘ŽǤ ͳ͹ǣʹʹȂʹ͹Ǥ

—±†‘–ǡ Ǥ ‡– ƒŽǤ ʹͲͲͻǤ ‡Žƒ–‹‘•Š‹’ „‡–™‡‡ „‘†› •‹œ‡ ƒ† Š‘‹‰ ƒ„‹Ž‹–› ‹ –Š‡ ‰‡—• •‹ƒ ȋ ›‡‘’–‡”ƒǢ‡‰ƒ Š‹Ž‹†ƒ‡ȌǤǦ ‘ŽǤ–‘‘ŽǤ͵ͶǣͳͷͺȂͳ͸ͳǤ

‹ Ž‹‰ǡ Ǥ ‡– ƒŽǤ ʹͲͲ͸Ǥ Š‡ †‹•–”‹„—–‹‘• ‘ˆ ƒ ™‹†‡ ”ƒ‰‡ ‘ˆ –ƒš‘‘‹  ‰”‘—’• ƒ”‡ ‡š’ƒ†‹‰ ’‘Ž‡™ƒ”†•ǤǦ Ž‘„ǤŠƒ‰Ǥ‹‘ŽǤͳʹǣͶͷͲȂͶͷͷǤ

—– Š‹•‘ǡ ǤǤͳͻͷͻǤ ‘ƒ‰‡–‘ƒ–ƒ‘•ƒŽ‹ƒ‘”™Š›ƒ”‡–Š‡”‡•‘ƒ›‹†•‘ˆƒ‹ƒŽ•ǫǦǤƒ–Ǥ ͻ͵ǣͳͶͷȂͳͷͻǤ

•Š‹‹ǡ Ǥ ƒ† Š‹ƒ†ƒǡ Ǥ ʹͲͳʹǤ ‡ƒ”‹‰ ’”‡†ƒ–‘” ’”‘‘–‡• ‘‡š‹•–‡ ‡ ‘ˆ ’”‡› •’‡ ‹‡• ‹ Š‘•– Ȃ ’ƒ”ƒ•‹–‘‹†•›•–‡•ǤǦ”‘ Ǥƒ–ŽǤ ƒ†Ǥ ‹ǤǤǤǤͳͲͻǣͷͳͳͻǤ

ƒ •‘ǡǤǤƒ†˜‡”’‡ ǡ ǤǤʹͲͲͲǤ‡•’‘•‡•‘ˆ’Žƒ–’‘’—Žƒ–‹‘•ƒ† ‘—‹–‹‡•–‘ ‡˜‹”‘‡–ƒŽ Šƒ‰‡•‘ˆ–Š‡Žƒ–‡—ƒ–‡”ƒ”›ǤǦƒŽ‡‘„‹‘Ž‘‰›ʹ͸ǣͳͻͶȂʹʹͲǤ

‘ŠǡǤ‡–ƒŽǤʹͲͲ͹Ǥ‘‹Ž—–”‹‡–•‹ˆŽ—‡ ‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘•‘ˆ–”‘’‹ ƒŽ–”‡‡•’‡ ‹‡•ǤǦ”‘ Ǥƒ–ŽǤ  ƒ†Ǥ ‹ǤǤǤǤͳͲͶǣͺ͸ͶȂͻǤ

‘”†ƒ‘ǡǤ‡–ƒŽǤʹͲͲ͵Ǥ ˜ƒ”‹ƒ–’”‘’‡”–‹‡•‹ ‘‡˜‘Ž—–‹‘ƒ”›‡–™‘”•‘ˆ’Žƒ–Ǧƒ‹ƒŽ‹–‡”ƒ –‹‘•ǤǦ  ‘ŽǤ‡––Ǥ͸ǣ͸ͻȂͺͳǤ

‡ƒ”•ǡǤǤ‡–ƒŽǤͳͻͻͺǤ†ƒ‰‡”‡†—–—ƒŽ‹••ǣŠ‡ ‘•‡”˜ƒ–‹‘‘ˆ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘”‹–‡”ƒ –‹‘•ǤǦ —Ǥ‡˜Ǥ ‘ŽǤ›•–Ǥʹͻǣͺ͵ȂͳͳʹǤ

‡Ž–ǡǤƒ†—”‡ǡǤƒͳͻͻͻǤ‡”‰‡–‹  ‘•–”ƒ‹–•ƒ†–Š‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡„‘†›•‹œ‡ƒ† Š‘‡”ƒ‰‡ƒ”‡ƒ‹ƒƒŽ•ǤǦ ‘Ž‘‰›ͺͲǣ͵͵͹Ȃ͵ͶͲǤ

”ƒˆ–ǡǤ ǤǤ‡–ƒŽǤʹͲͲͺǤ — –‹‘ƒŽ–”ƒ‹–•ƒ†‹ Š‡Ǧ„ƒ•‡†–”‡‡ ‘—‹–›ƒ••‡„Ž›‹ƒƒœ‘‹ƒ ˆ‘”‡•–ǤǦ ‹‡ ‡͵ʹʹǣͷͺͲȂͷͺʹǤ

 16   ‡‡”ƒŽ‹–”‘†— –‹‘

”‡„•ǡ Ǥ ‡– ƒŽǤ ͳͻͻͶǤ  ‘Ž‘‰›ǣ –Š‡ ‡š’‡”‹‡–ƒŽ ƒƒŽ›•‹• ‘ˆ †‹•–”‹„—–‹‘ ƒ† ƒ„—†ƒ ‡Ǥ Ǧ ƒ”’‡” ‘ŽŽ‹•‘ŽŽ‡‰‡—„Ž‹•Š‡”•Ǥ

”‡‡ǡ Ǥ ‡– ƒŽǤ ʹͲͲ͹Ǥ ‘ŽŽ‹ƒ–‹‘ƒ† ‘–Š‡” ‡ ‘•›•–‡ •‡”˜‹ ‡•’”‘†— ‡† „› ‘„‹Ž‡ ‘”‰ƒ‹••ǣƒ ‘ ‡’–—ƒŽˆ”ƒ‡™‘”ˆ‘”–Š‡‡ˆˆ‡ –•‘ˆŽƒ†Ǧ—•‡ Šƒ‰‡ǤǦ ‘ŽǤ‡––ǤͳͲǣʹͻͻȂ͵ͳͶǤ

‡˜‹‡ǡ ǤǤ‡–ƒŽǤʹͲͲ͵Ǥ‡ Šƒ‹••—†‡”Ž›‹‰–Š‡‹’ƒ –•‘ˆ‡š‘–‹ ’Žƒ–‹˜ƒ•‹‘•ǤǦ”‘ Ǥ‹‘ŽǤ ‹Ǥ ʹ͹Ͳǣ͹͹ͷȂͺͳǤ

‘†‰‡ǡǤǤͳͻͻ͵Ǥ ˜ƒ•‹‘•ǣ‡••‘•ˆ‘” ‘Ž‘‰›ǤǦ”‡†• ‘ŽǤ˜‘ŽǤͺǣͳ͵͵Ȃͳ͵͹Ǥ

‘•‡›ǡ ǤǤƒ†ƒ—‰ŠƒǡǤʹͲͲ͸ǤŠ‡‡ ‘‘‹ ˜ƒŽ—‡‘ˆ‡ ‘Ž‘‰‹ ƒŽ•‡”˜‹ ‡•’”‘˜‹†‡†„›‹•‡ –•ǤǦ ‹‘• ‹‡ ‡ͷ͸ǣ͵ͳͳǤ

ƒ ”–Š—”ǡǤͳͻ͸ͷǤƒ––‡”•‘ˆ•’‡ ‹‡•†‹˜‡”•‹–›ǤǦ‹‘ŽǤ‡˜ǤͶͲǣͷͳͲȂͷ͵͵Ǥ

ƒ ”–Š—”ǡ Ǥ Ǥ ͳͻ͹ʹǤ ‡‘‰”ƒ’Š‹ ƒŽ ‡ ‘Ž‘‰›ǣ ’ƒ––‡”• ‹ –Š‡ †‹•–”‹„—–‹‘‘ˆ•’‡ ‹‡•ǤǦ”‹ ‡–‘ ‹˜‡”•‹–›”‡••Ǥ

ƒ ”–Š—”ǡǤƒ†ƒ ”–Š—”ǡ Ǥͳͻ͸ͳǤ„‹”†•’‡ ‹‡•†‹˜‡”•‹–›ǤǦ ‘Ž‘‰›ͶʹǣͷͻͶȂͷͻͺǤ

ƒ”‰ƒŽ‡ˆǡǤͳͻ͸ͺǤ‡”•’‡ –‹˜‡•‹‡ ‘Ž‘‰‹ ƒŽ–Š‡‘”›ǤǦ‹˜‡”•‹–›‘ˆŠ‹ ƒ‰‘”‡••Ǥ

ƒ”‰ƒŽ‡ˆǡǤͳͻͻ͵Ǥ‡‘”Àƒ†‡Ž‘••‹•–‡ƒ•‡ ‘Ž×‰‹ ‘•ǤǦ‹˜‡”•‹–ƒ–†‡ƒ” ‡Ž‘ƒǤ

ƒ”œŽ—ˆˆǡ Ǥ ʹͲͲͳǤ ‘”Ž†™‹†‡ —”„ƒ‹œƒ–‹‘ ƒ† ‹–• ‡ˆˆ‡ –• ‘ „‹”†•ǤǦ ǣƒ”œŽ—ˆˆǡ ǤǤ‡–ƒŽǤȋ‡†•Ȍǡ ˜‹ƒ‡ ‘Ž‘‰›ƒ† ‘•‡”˜ƒ–‹‘‹ƒ—”„ƒ‹œ‹‰™‘”Ž†Ǥ’‹‰‡”ǡ’’ǤͳͻȂͶ͹Ǥ

ƒ›ǡǤǤͳͻͺͺǤ ‘™ƒ›•’‡ ‹‡•ƒ”‡–Š‡”‡‘ƒ”–ŠǫǦ ‹‡ ‡ʹͶͳǣͳͶͶͳȂͳͶͶͻǤ

 †‘ƒŽ†ǡǤ Ǥ‡–ƒŽǤʹͲͲͺǤŠ‡‹’Ž‹ ƒ–‹‘•‘ˆ —””‡–ƒ†ˆ—–—”‡—”„ƒ‹œƒ–‹‘ˆ‘”‰Ž‘„ƒŽ’”‘–‡ –‡† ƒ”‡ƒ•ƒ†„‹‘†‹˜‡”•‹–› ‘•‡”˜ƒ–‹‘ǤǦ‹‘ŽǤ‘•‡”˜ǤͳͶͳǣͳ͸ͻͷȂͳ͹Ͳ͵Ǥ

 ƒ‡ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲʹǤ ‡•‘—” ‡Ǧ„ƒ•‡† ‹ Š‡• ’”‘˜‹†‡ ƒ „ƒ•‹• ˆ‘” ’Žƒ– •’‡ ‹‡• †‹˜‡”•‹–› ƒ† †‘‹ƒ ‡‹ƒ” –‹ –—†”ƒǤǦƒ–—”‡Ͷͳͷǣ͸ͺȂ͹ͳǤ

 ‹‡›ǡǤǤʹͲͲ͸Ǥ”„ƒ‹œƒ–‹‘ƒ•ƒƒŒ‘” ƒ—•‡‘ˆ„‹‘–‹ Š‘‘‰‡‹œƒ–‹‘ǤǦ‹‘ŽǤ‘•‡”˜Ǥͳʹ͹ǣ ʹͶ͹Ȃʹ͸ͲǤ

 17 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•   

 ‹‡›ǡǤǤƒ†‘ ™‘‘†ǡ ǤǤʹͲͲͳǤ‹‘–‹ Š‘‘‰‡‹œƒ–‹‘ǣƒ•‡“—‡–‹ƒŽƒ†•‡Ž‡ –‹˜‡’”‘ ‡••ǤǦ ǣ‘ ™‘‘†ǡ ǤǤƒ† ‹‡›ǡǤǤȋ‡†•Ȍǡ‹‘–‹ Š‘‘‰‡‹œƒ–‹‘Ǥ’”‹‰‡”ǡ’’ǤͳȂͳ͹Ǥ

‡‘––ǡ Ǥ‡–ƒŽǤʹͲͲͶǤ‘Ž‡”ƒ ‡‘ˆ’‘ŽŽ‹ƒ–‹‘‡–™‘”•–‘•’‡ ‹‡•‡š–‹ –‹‘•ǤǦ”‘ ǤǤ‘ Ǥ‹‘ŽǤ  ‹Ǥʹ͹ͳǣʹ͸ͲͷȂʹ͸ͳͳǤ

‹ Š‡‡”ǡǤǤʹͲͲͲǤŠ‡„‡‡•‘ˆ–Š‡™‘”Ž†ǤǦ ‘Š• ‘’‹•‹˜‡”•‹–›”‡••Ǥ

‘”ƒŽ‡•ǡ Ǥ Ǥ ƒ† žœ“—‡œǡ Ǥ Ǥ ʹͲͲͺǤ Š‡ ‡ˆˆ‡ – ‘ˆ •’ƒ ‡ ‹ ’Žƒ–Ǧƒ‹ƒŽ —–—ƒŽ‹•–‹  ‡–™‘”•ǣ ‹•‹‰Š–•ˆ”‘ƒ•‹—Žƒ–‹‘•–—†›ǤǦ‹‘•ͳͳ͹ǣͳ͵͸ʹȂͳ͵͹ͲǤ

òŽŽ‡”ǡ Ǥ ‡– ƒŽǤ ʹͲͲ͸Ǥ —ƒ–‹–ƒ–‹˜‡ ’‘ŽŽ‡ ”‡“—‹”‡‡–• ‘ˆ •‘Ž‹–ƒ”› „‡‡•ǣ ’Ž‹ ƒ–‹‘• ˆ‘” „‡‡ ‘•‡”˜ƒ–‹‘ƒ†–Š‡‡˜‘Ž—–‹‘‘ˆ„‡‡ȂˆŽ‘™‡””‡Žƒ–‹‘•Š‹’•ǤǦ‹‘ŽǤ‘•‡”˜Ǥͳ͵Ͳǣ͸ͲͶȂ͸ͳͷǤ

‹‡–‘ǡǤ‡–ƒŽǤʹͲͳͶǤ—”‘’‡ƒ‡†‹•–‘ˆ‡‡•ǤǦ—„Ž‹ ƒ–‹‘ˆˆ‹ ‡‘ˆ–Š‡—”‘’‡ƒ‹‘Ǥ

›•–”ڏǡǤ‡–ƒŽǤʹͲͲͲǤ‘”ƒŽ”‡‡ˆ†‹•–—”„ƒ ‡ƒ†”‡•‹Ž‹‡ ‡‹ƒŠ—ƒǦ†‘‹ƒ–‡†‡˜‹”‘‡–ǤǦ ”‡†• ‘ŽǤ˜‘ŽǤͳͷǣͶͳ͵ȂͶͳ͹Ǥ

Ž‡•‡ǡ Ǥ Ǥ ƒ† ‘”†ƒ‘ǡ Ǥ ʹͲͲʹǤ ‡‘‰”ƒ’Š‹  ’ƒ––‡”• ‹ ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘” —–—ƒŽ‹•–‹  ‡–™‘”•Ǥ Ǧ  ‘Ž‘‰›ͺ͵ǣʹͶͳ͸ȂʹͶʹͶǤ

ŽŽ‡”–‘ǡ Ǥ‡–ƒŽǤʹͲͳͳǤ ‘™ƒ›ˆŽ‘™‡”‹‰’Žƒ–•ƒ”‡’‘ŽŽ‹ƒ–‡†„›ƒ‹ƒŽ•ǫǦ‹‘•ͳʹͲǣ͵ʹͳȂ͵ʹ͸Ǥ

ƒ”‡•ƒǡǤ‡–ƒŽǤͳͻͻͻǤ‘Ž‡™ƒ”†•Š‹ˆ–•‹‰‡‘‰”ƒ’Š‹ ƒŽ”ƒ‰‡•‘ˆ„—––‡”ˆŽ›•’‡ ‹‡•ƒ••‘ ‹ƒ–‡†™‹–Š ”‡‰‹‘ƒŽ™ƒ”‹‰Ǥ͵ͻͻǣͷ͹ͻȂͷͺ͵Ǥ

ƒ— Šƒ”†ǡǤ‡–ƒŽǤʹͲͲ͸Ǥ—Ž–‹’Ž‡‡ˆˆ‡ –•‘ˆ—”„ƒ‹œƒ–‹‘‘–Š‡„‹‘†‹˜‡”•‹–›‘ˆ†‡˜‡Ž‘’‹‰ ‘—–”‹‡•ǣ Š‡ ƒ•‡‘ˆƒˆƒ•–Ǧ‰”‘™‹‰‡–”‘’‘Ž‹–ƒƒ”‡ƒȋ‘ ‡’ ‹×ǡŠ‹Ž‡ȌǤǦ‹‘ŽǤ‘•‡”˜Ǥͳʹ͹ǣʹ͹ʹȂʹͺͳǤ

‡”‡‹”ƒǡ ǤǤ‡–ƒŽǤʹͲͳͲǤ ‡ƒ”‹‘•ˆ‘”‰Ž‘„ƒŽ„‹‘†‹˜‡”•‹–›‹–Š‡ʹͳ•– ‡–—”›ǤǦ ‹‡ ‡͵͵ͲǣͳͶͻ͸Ȃ ͳͷͲͳǤ

‡”‡•ǡǤǤ‡–ƒŽǤʹͲͲ͵Ǥ‡”–‡„”ƒ–‡”‡•’‘•‡•–‘•—”ˆƒ ‡™‹Ž†ˆ‹”‡•‹ƒ ‡–”ƒŽƒœ‘‹ƒˆ‘”‡•–ǤǦ”›š ͵͹ǣͻ͹ȂͳͲͻǤ

‹ƒƒǡǤǤͳͻ͸͸Ǥƒ–‹–—†‹ƒŽ‰”ƒ†‹‡–•‹•’‡ ‹‡•†‹˜‡”•‹–›ǣƒ”‡˜‹‡™‘ˆ ‘ ‡’–•ǤǦǤƒ–ǤͳͲͲǣ͵͵Ȃ Ͷ͸Ǥ

 18   ‡‡”ƒŽ‹–”‘†— –‹‘

Žƒ‹•ƒ ‡ǡ Ǥ ‡– ƒŽǤ ʹͲͲͺǤ ‘Ž‘‹œƒ–‹‘ ‘ˆ ƒ ‹ˆ‹  ‹•Žƒ†• „› ’ƒ”ƒ•‹–‡• ‘ˆ Ž‘™ †‹•’‡”•ƒŽ ƒ„‹Ž‹–›ǣ Š›Ž‘‰‡‘‰”ƒ’Š› ‘ˆ –™‘ ‘‘‰‡‡ƒ •’‡ ‹‡• ’ƒ”ƒ•‹–‹œ‹‰ „—––‡”ˆŽ›ˆ‹•Š‡• ‹ –Š‡ ‘—–Š ƒ ‹ˆ‹   ‡ƒǤǦ Ǥ‹‘‰‡‘‰”Ǥ͵ͷǣ͹͸Ȃͺ͹Ǥ

‘‹•‘–ǡǤ‡–ƒŽǤʹͲͳʹǤŠ‡†‹••‹‹Žƒ”‹–›‘ˆ•’‡ ‹‡•‹–‡”ƒ –‹‘‡–™‘”•ǤǦ ‘ŽǤ‡––Ǥͳͷǣͳ͵ͷ͵Ȃ͸ͳǤ

‘”–‡”ǡ Ǥ Ǥ ƒ† ƒ˜‹‰ƒ‘ǡ Ǥ Ǥ ͳͻͻͲǤ ˜ƒ•‹‘ ‘ˆ ’‘Ž›‰›‡ ˆ‹”‡ ƒ–• †‡ ‹ƒ–‡• ƒ–‹˜‡ ƒ–• ƒ† †‹•”—’–•ƒ”–Š”‘’‘† ‘—‹–›ǤǦ ‘Ž‘‰›͹ͳǣʹͲͻͷȂʹͳͲ͸Ǥ

‘––•ǡǤ Ǥ‡–ƒŽǤʹͲͳͲǤ Ž‘„ƒŽ’‘ŽŽ‹ƒ–‘”†‡ Ž‹‡•ǣ–”‡†•ǡ‹’ƒ –•ƒ††”‹˜‡”•ǤǦ”‡†• ‘ŽǤ˜‘ŽǤʹͷǣ ͵ͶͷȂͷ͵Ǥ

‘––•ǡǤ‡–ƒŽǤʹͲͳͶǤ–ƒ–—•ƒ†”‡†•‘ˆ—”‘’‡ƒ‘ŽŽ‹ƒ–‘”•Ǥ‡›ˆ‹†‹‰•‘ˆ–Š‡’”‘Œ‡ –ǤǦ ‡•‘ˆ–—„Ž‹•Š‡”•Ǥ

ƒŠ‡Žǡ Ǥ ǤʹͲͲʹǤ ‘‘‰‡‹œƒ–‹‘‘ˆˆ”‡•Š™ƒ–‡”ˆƒ—ƒ•ǤǦ—Ǥ‡˜Ǥ ‘ŽǤ›•–Ǥ͵͵ǣʹͻͳȂ͵ͳͷǤ

‡‡†ǡǤǤ‡–ƒŽǤʹͲͲͲǤŠ‡”‘Ž‡‘ˆ†‹•’‡”•ƒŽƒ††‹•–—”„ƒ ‡‹†‡–‡”‹‹‰•’ƒ–‹ƒŽŠ‡–‡”‘‰‡‡‹–›‹ •‡†‡–ƒ”›‘”‰ƒ‹••ǤǦ ‘Ž‘‰›ͺͳǣʹͲͳͳȂʹͲʹ͸Ǥ

‹ Š–‡”ǡǤƒ†‡”ƒƒ–›ǡ ǤʹͲͲ͵Ǥ ‘‡›„‡‡ˆ‘”ƒ‰‹‰„‡Šƒ˜‹‘”ǤǦ ǣŽ‘‰‰‡”ǡǤ Ǥƒ†ƒ•—ƒ™ƒǡǤ ȋ‡†•Ȍǡš’Ž‘”‹‰ƒ‹ƒŽ„‡Šƒ˜‹‘”‹Žƒ„‘”ƒ–‘”›ƒ†ˆ‹‡Ž†ǣƒŠ›’‘–Š‡•‹•Ǧ–‡•–‹‰ƒ’’”‘ƒ Š–‘–Š‡ †‡˜‡Ž‘’‡–ǡ ƒ—•ƒ–‹‘ǡˆ— –‹‘ǡƒ†‡˜‘Ž—–‹‘‘ˆƒ‹ƒŽ„‡Šƒ˜‹‘”ǤŽ•‡˜‹‡”ǡ’’Ǥͳ͵͵Ǥ

‹ Ž‡ˆ•ǡǤǤͳͻͺ͹Ǥ‘—‹–›†‹˜‡”•‹–›ǣ”‡Žƒ–‹˜‡”‘Ž‡•‘ˆŽ‘ ƒŽƒ†”‡‰‹‘ƒŽ’”‘ ‡••‡•ǤǦ ‹‡ ‡ʹ͵ͷǣ ͳ͸͹Ȃͳ͹ͳǤ

‹ Ž‡ˆ•ǡ Ǥ Ǥ ƒ†  ŠŽ—–‡”ǡ Ǥ ͳͻͻ͵Ǥ ’‡ ‹‡• †‹˜‡”•‹–› ‹ ‡ ‘Ž‘‰‹ ƒŽ ‘—‹–‹‡•ǣ Š‹•–‘”‹ ƒŽ ƒ† ‰‡‘‰”ƒ’Š‹ ƒŽ’‡”•’‡ –‹˜‡•ǤǦ‹˜‡”•‹–›‘ˆŠ‹ ƒ‰‘”‡••Ǥ

‘†”‹‰‘ǡǤ‡–ƒŽǤʹͲͲͶǤ‹”‡ –”‡‰‡‡”ƒ–‹‘‹•‘––Š‡‘Ž›”‡•’‘•‡‘ˆ‡†‹–‡””ƒ‡ƒˆ‘”‡•–•–‘Žƒ”‰‡ ˆ‹”‡•ǤǦ ‘Ž‘‰›ͺͷǣ͹ͳ͸Ȃ͹ʹͻǤ

‡‡Ž‡›ǡǤǤͳͻͺͷǤ ‘‡›„‡‡‡ ‘Ž‘‰›ǣƒ•–—†›‘ˆƒ†ƒ’–ƒ–‹‘‹•‘ ‹ƒŽŽ‹ˆ‡ǤǦ”‹ ‡–‘‹˜‡”•‹–›”‡••Ǥ

Š‹’Ž‡›ǡǤ‡–ƒŽǤʹͲͲ͸Ǥ ”‘’Žƒ––”ƒ‹–•–‘’Žƒ– ‘—‹–‹‡•ǣƒ•–ƒ–‹•–‹ ƒŽ‡ Šƒ‹•–‹ ƒ’’”‘ƒ Š–‘ „‹‘†‹˜‡”•‹–›ǤǦ ‹‡ ‡͵ͳͶǣͺͳʹȂͺͳͶǤ

 19 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•   

Š‘ Šƒ–ǡǤ‡–ƒŽǤʹͲͳͲǤ ˜ƒ•‹‘ǡ ‘’‡–‹–‹‘ǡƒ†„‹‘†‹˜‡”•‹–›Ž‘••‹—”„ƒ‡ ‘•›•–‡•ǤǦ‹‘• ‹‡ ‡ ͸ͲǣͳͻͻȂʹͲͺǤ

‘—•ƒǡǤǤͳͻͺͶǤŠ‡”‘Ž‡‘ˆ†‹•–—”„ƒ ‡ǤǦ—Ǥ‡˜Ǥ ‘ŽǤ›•–Ǥͳͷǣ͵ͷ͵Ȃ͵ͻͳǤ

–‡™ƒ”–ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲʹǤ ‡’‘”‘•’ƒ–‹ƒŽ †‹•–”‹„—–‹‘• ‘ˆ ‡Žǡ —Ž‡ †‡‡”ǡ ƒ† ƒ––Ž‡ǣ ”‡•‘—” ‡ ’ƒ”–‹–‹‘‹‰ƒ† ‘’‡–‹–‹˜‡†‹•’Žƒ ‡‡–ǤǦ ǤƒƒŽǤͺ͵ǣʹʹͻȂʹͶͶǤ

–”‘‰ǡǤǤ‡–ƒŽǤͳͻͺͶǤ •‡ –•‘’Žƒ–•Ǥ‘—‹–›’ƒ––‡”•ƒ†‡ Šƒ‹••ǤǦŽƒ ™‡ŽŽ ‹‡–‹ˆ‹  —„Ž‹ ƒ–‘•Ǥ

™‹Šƒ”–ǡǤ‡–ƒŽǤͳͻͺͺǤ‡Žƒ–‹‰„‘†›•‹œ‡–‘–Š‡”ƒ–‡‘ˆŠ‘‡”ƒ‰‡—•‡‹ƒƒŽ•ǤǦ ‘Ž‘‰›ǣ͵ͻ͵Ȃ ͵ͻͻǤ

Š‘ƒ•ǡǤǤʹͲͳͲǤŽ‹ƒ–‡ǡ Ž‹ƒ–‡ Šƒ‰‡ƒ†”ƒ‰‡„‘—†ƒ”‹‡•ǤǦ‹˜‡”•Ǥ‹•–”‹„Ǥͳ͸ǣͶͺͺȂͶͻͷǤ

Š‘ƒ•ǡ ǤǤ‡–ƒŽǤʹͲͲͳǤŠ‡“—ƒŽ‹–›ƒ†‹•‘Žƒ–‹‘‘ˆŠƒ„‹–ƒ–’ƒ– Š‡•„‘–Š†‡–‡”‹‡™Š‡”‡„—––‡”ˆŽ‹‡• ’‡”•‹•–‹ˆ”ƒ‰‡–‡†Žƒ†• ƒ’‡•ǤǦ”‘ ǤǤ‘ Ǥ‹‘ŽǤ ‹Ǥʹ͸ͺǣͳ͹ͻͳȂͳ͹ͻ͸Ǥ

Š”—•ŠǡǤ Ǥ‡–ƒŽǤʹͲͲ͸Ǥ”‡†‹ –‹‰–Š‡‡ˆˆ‡ –•‘ˆŠƒ„‹–ƒ–Š‘‘‰‡‹œƒ–‹‘‘ƒ”‹‡„‹‘†‹˜‡”•‹–›ǤǦ  ‘ŽǤ’’ŽǤͳ͸ǣͳ͸͵͸Ȃͳ͸ͶʹǤ

”ƒ˜‡•‡–ǡ Ǥ ƒ† ‹ Šƒ”†•‘ǡ Ǥ Ǥ ʹͲͲ͸Ǥ ‹‘Ž‘‰‹ ƒŽ ‹˜ƒ•‹‘• ƒ• †‹•”—’–‘”• ‘ˆ ’Žƒ– ”‡’”‘†— –‹˜‡ —–—ƒŽ‹••ǤǦ”‡†• ‘ŽǤ˜‘ŽǤʹͳǣʹͲͺȂͳ͸Ǥ

—”‡”ǡ Ǥ Ǥ ‡– ƒŽǤ ͳͻͻͺǤ ƒ –‘”• ‹ˆŽ—‡ ‹‰ •— ‡••‹‘ǣ Ž‡••‘• ˆ”‘ Žƒ”‰‡ǡ ‹ˆ”‡“—‡– ƒ–—”ƒŽ †‹•–—”„ƒ ‡•ǤǦ ‘•›•–‡•ͳǣͷͳͳȂͷʹ͵Ǥ

›Ž‹ƒƒ‹•ǡ ǤǤ‡–ƒŽǤʹͲͲ͹Ǥ ƒ„‹–ƒ–‘†‹ˆ‹ ƒ–‹‘ƒŽ–‡”•–Š‡•–”— –—”‡‘ˆ–”‘’‹ ƒŽŠ‘•–Ǧ’ƒ”ƒ•‹–‘‹†ˆ‘‘† ™‡„•ǤǦƒ–—”‡ͶͶͷǣʹͲʹȂʹͲͷǤ

žœ“—‡œǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͻƒǤ ˜ƒŽ—ƒ–‹‰ —Ž–‹’Ž‡ †‡–‡”‹ƒ–• ‘ˆ –Š‡ •–”— –—”‡ ‘ˆ ’Žƒ–Ǧƒ‹ƒŽ —–—ƒŽ‹•–‹ ‡–™‘”•ǤǦ ‘Ž‘‰›ͻͲǣʹͲ͵ͻȂʹͲͶ͸Ǥ

žœ“—‡œǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͻ„Ǥ ‹–‹‰ ’ƒ––‡” ƒ† ’”‘ ‡•• ‹ ’Žƒ–Ǧƒ‹ƒŽ —–—ƒŽ‹•–‹  ‡–™‘”•ǣ ƒ ”‡˜‹‡™ǤǦǤ‘–ǤͳͲ͵ǣͳͶͶͷȂͷ͹Ǥ

‹–‘—•‡ǡǤǤ‡–ƒŽǤͳͻͻ͹Ǥ —ƒ†‘‹ƒ–‹‘‘ˆƒ”–Šǯ•‡ ‘•›•–‡•ǤǦ ‹‡ ‡ʹ͹͹ǣͶͻͶȂͶͻͻǤ

 20   ‡‡”ƒŽ‹–”‘†— –‹‘

‘ ”‹• Šǡ Ǥ ͳͻ͸͹Ǥ Š‡ †ƒ ‡ Žƒ‰—ƒ‰‡ ƒ† ‘”‹‡–ƒ–‹‘ ‘ˆ „‡‡•Ǥ Ǧ ƒ”˜ƒ”† ‹˜‡”•‹–› ”‡••Ǥ ƒ„”‹†‰‡ǡǤ

‡‹„—ŽŽǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͲǤ ‹˜‡”•‹–› ‘ˆ „—––‡”ˆŽ‹‡• ‹ –Š‡ ƒ‰”‹ —Ž–—”ƒŽ Žƒ†• ƒ’‡ǣ –Š‡ ”‘Ž‡ ‘ˆ ˆƒ”‹‰ •›•–‡ƒ†Žƒ†• ƒ’‡Š‡–‡”‘‰‡‡‹–›ǤǦ ‘‰”ƒ’Š›ȋ‘’ǤȌǤʹ͵ǣ͹Ͷ͵Ȃ͹ͷͲǤ

‡•–”‹ Šǡ Ǥ ͳͻͻ͸Ǥ ƒ„‹–ƒ– ”‡“—‹”‡‡–• ‘ˆ ‡–”ƒŽ —”‘’‡ƒ „‡‡• ƒ† –Š‡ ’”‘„Ž‡• ‘ˆ ’ƒ”–‹ƒŽ Šƒ„‹–ƒ–•Ǥ Ǧ ǣ ƒ–Š‡•‘ǡ Ǥ ‡– ƒŽǤ ȋ‡†•Ȍǡ‹‡ƒ ‘ ‹‡–› ›’‘•‹— ‡”‹‡•Ǥ ‘ŽǤ ͳͺǤ  ƒ†‡‹  ”‡••‹‹–‡†ǡ’’ǤͳȂͳ͸Ǥ

‡––•–‡‹ǡǤƒ† Š‹†ǡǤͳͻͻͻǤ‘•‡”˜ƒ–‹‘‘ˆƒ”–Š”‘’‘††‹˜‡”•‹–›‹‘–ƒ‡™‡–Žƒ†•ǣˆˆ‡ – ‘ˆƒŽ–‹–—†‡ǡŠƒ„‹–ƒ–“—ƒŽ‹–›ƒ†Šƒ„‹–ƒ–ˆ”ƒ‰‡–ƒ–‹‘‘„—––‡”ˆŽ‹‡•ƒ†‰”ƒ••Š‘’’‡”•ǤǦ Ǥ’’ŽǤ  ‘ŽǤ͵͸ǣ͵͸͵Ȃ͵͹͵Ǥ

‹ŽŽ•‘ǡǤ Ǥͳͻͻ͵Ǥ‹•’‡”•ƒŽ‘†‡ǡ•‡‡†•Šƒ†‘™•ǡƒ† ‘Ž‘‹œƒ–‹‘’ƒ––‡”•ǤǦ‡‰‡–ƒ–‹‘ͳͲ͹ǦͳͲͺǣ ʹ͸ͳȂʹͺͲǤ

‘—‰ǡǤǤ‡–ƒŽǤʹͲͲͷǤ‘’‡–‹–‹‘ƒ† ‘’‡•ƒ–‹‘ƒ‘‰ ƒ––Ž‡ǡœ‡„”ƒ•ǡƒ†‡Ž‡’Šƒ–•‹ƒ•‡‹Ǧ ƒ”‹†•ƒ˜ƒƒ‹ƒ‹‹’‹ƒǡ‡›ƒǤǦ‹‘ŽǤ‘•‡”˜Ǥͳʹʹǣ͵ͷͳȂ͵ͷͻǤ

—”„— Š‡ǡǤ‡–ƒŽǤʹͲͳͲǤƒš‹—ˆ‘”ƒ‰‹‰”ƒ‰‡•‹•‘Ž‹–ƒ”›„‡‡•ǣ‘Ž›ˆ‡™‹†‹˜‹†—ƒŽ•Šƒ˜‡–Š‡ ƒ’ƒ„‹Ž‹–›–‘ ‘˜‡”Ž‘‰ˆ‘”ƒ‰‹‰†‹•–ƒ ‡•ǤǦ‹‘ŽǤ‘•‡”˜ǤͳͶ͵ǣ͸͸ͻȂ͸͹͸Ǥ





 21



   Šƒ’–‡”ͳ

Determinantsofspatialdistribution inabeecommunity:nesting resources,flowerresources,andbody size





















TornéǦNoguera,A;Rodrigo,A;Arnan,X;Osorio,S;BarrilǦGraells,H;CorreiadaRochaǦFilho,

L;Bosch,J.(2014)PLoSONE9(5):e97255.doi:10.1371/journal.pone.0097255



Šƒ’–‡”ͳ

Summary †‡”•–ƒ†‹‰ „‹‘†‹˜‡”•‹–› †‹•–”‹„—–‹‘ ‹• ƒ ’”‹ƒ”› ‰‘ƒŽ ‘ˆ ‘—‹–› ‡ ‘Ž‘‰›Ǥ – ƒ Žƒ†• ƒ’‡ • ƒŽ‡ǡ „‡‡ ‘—‹–‹‡• ƒ”‡ ƒˆˆ‡ –‡† „› Šƒ„‹–ƒ– ‘’‘•‹–‹‘ǡ ƒ–Š”‘’‘‰‡‹  Žƒ† —•‡ǡ ƒ† ˆ”ƒ‰‡–ƒ–‹‘Ǥ ‘™‡˜‡”ǡ Ž‹––Ž‡ ‹ˆ‘”ƒ–‹‘ ‹• ƒ˜ƒ‹Žƒ„Ž‡ ‘ Ž‘ ƒŽǦ• ƒŽ‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆ„‡‡ ‘—‹–‹‡•™‹–Š‹Šƒ„‹–ƒ–•–Šƒ–ƒ”‡—‹ˆ‘”ƒ––Š‡ Žƒ†• ƒ’‡• ƒŽ‡Ǥ‡•–—†‹‡†ƒ„‡‡ ‘—‹–›ƒŽ‘‰™‹–ŠˆŽ‘”ƒŽƒ†‡•–‹‰”‡•‘—” ‡•‘˜‡” ƒ ͵ʹ ʹ ƒ”‡ƒ ‘ˆ —‹–‡””—’–‡† ‡†‹–‡””ƒ‡ƒ • ”—„Žƒ†Ǥ —” ‘„Œ‡ –‹˜‡• ™‡”‡ ȋ‹Ȍ –‘ ƒƒŽ›œ‡ˆŽ‘”ƒŽƒ†‡•–‹‰”‡•‘—” ‡ ‘’‘•‹–‹‘ƒ––Š‡Šƒ„‹–ƒ–• ƒŽ‡Ǥ‡ƒ•™Š‡–Š‡”–Š‡•‡ ”‡•‘—” ‡• ˆ‘ŽŽ‘™ ƒ ‰‡‘‰”ƒ’Š‹ ƒŽ ’ƒ––‡” ƒ ”‘•• –Š‡ • ”—„Žƒ† ƒ– „‡‡Ǧˆ‘”ƒ‰‹‰ ”‡Ž‡˜ƒ– †‹•–ƒ ‡•Ǣ ȋ‹‹Ȍ –‘ ƒƒŽ›œ‡ –Š‡ †‹•–”‹„—–‹‘ ‘ˆ „‡‡ ‘’‘•‹–‹‘ ƒ ”‘•• –Š‡ • ”—„Žƒ†Ǥ ‡‡• „‡‹‰Š‹‰ŠŽ›‘„‹Ž‡‘”‰ƒ‹••ǡ™‡ƒ•™Š‡–Š‡”„‡‡ ‘’‘•‹–‹‘•Š‘™•ƒŠ‘‘‰‡‡‘—• †‹•–”‹„—–‹‘ ‘” ‡Ž•‡ ˜ƒ”‹‡• •’ƒ–‹ƒŽŽ›Ǥ ˆ •‘ǡ ™‡ ƒ• ™Š‡–Š‡” –Š‹• ˜ƒ”‹ƒ–‹‘ ‹• ‹””‡‰—Žƒ” ‘” ˆ‘ŽŽ‘™•ƒ‰‡‘‰”ƒ’Š‹ ƒŽ’ƒ––‡”ƒ†™Š‡–Š‡”„‡‡•”‡•’‘†’”‹ƒ”‹Ž›–‘ˆŽ‘™‡”‘”–‘‡•–‹‰ ”‡•‘—” ‡•Ǣ ƒ† ȋ‹‹‹Ȍ –‘ ‡•–ƒ„Ž‹•Š ™Š‡–Š‡” „‘†› •‹œ‡ ‹ˆŽ—‡ ‡• –Š‡ ”‡•’‘•‡ –‘ Ž‘ ƒŽ ”‡•‘—” ‡ ƒ˜ƒ‹Žƒ„‹Ž‹–› ƒ† —Ž–‹ƒ–‡Ž› •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘Ǥ ‡ ‘„–ƒ‹‡† ͸ͷͺͲ •’‡ ‹‡• „‡Ž‘‰‹‰–‘ͻͺ•’‡ ‹‡•Ǥ‡•’‹–‡„‡‡‘„‹Ž‹–›ƒ†–Š‡ƒ„•‡ ‡‘ˆ‡˜‹”‘‡–ƒŽ„ƒ””‹‡”•ǡ ‘—”„‡‡ ‘—‹–›•Š‘™•ƒ Ž‡ƒ”‰‡‘‰”ƒ’Š‹ ƒŽ’ƒ––‡”ǤŠ‹•’ƒ––‡”‹•‘•–Ž›ƒ––”‹„—–ƒ„Ž‡ –‘Š‡–‡”‘‰‡‡‘—•†‹•–”‹„—–‹‘‘ˆ•ƒŽŽȋδͷͷ‰Ȍ•’‡ ‹‡•ȋ™‹–Š’”‡•—‡†•ƒŽŽ‡”ˆ‘”ƒ‰‹‰ ”ƒ‰‡•Ȍǡƒ†‹•‘•–Ž›‡š’Žƒ‹‡†„›ˆŽ‘™‡””‡•‘—” ‡•”ƒ–Š‡”–Šƒ‡•–‹‰•—„•–”ƒ–‡•Ǥ˜‡ –Š‡ǡƒŽƒ”‰‡’”‘’‘”–‹‘ȋͷͶǤͺΨȌ‘ˆ•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›”‡ƒ‹•—‡š’Žƒ‹‡†„›ˆŽ‘™‡”‘” ‡•–‹‰ ”‡•‘—” ‡•Ǥ ‡ ‘ Ž—†‡ –Šƒ– „‡‡ ‘—‹–‹‡• ƒ”‡ •–”‘‰Ž› ‘†‹–‹‘‡† „› Ž‘ ƒŽ ‡ˆˆ‡ –•ƒ†ƒ›‡šŠ‹„‹–•’ƒ–‹ƒŽŠ‡–‡”‘‰‡‡‹–›’ƒ––‡”•ƒ–ƒ• ƒŽ‡ƒ•Ž‘™ƒ•ͷͲͲǦͳͲͲͲ‹ ’ƒ– Š‡• ‘ˆ Š‘‘‰‡‡‘—• Šƒ„‹–ƒ–Ǥ Š‡•‡ ”‡•—Ž–• Šƒ˜‡ ‹’‘”–ƒ– ‹’Ž‹ ƒ–‹‘• ˆ‘” Ž‘ ƒŽ ’‘ŽŽ‹ƒ–‹‘†›ƒ‹ •ƒ†•’ƒ–‹ƒŽ˜ƒ”‹ƒ–‹‘‘ˆ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘”‡–™‘”•Ǥ   1.1. Introduction

”‘ƒ•–”‹ –Ž›–Š‡‘”‡–‹ ƒŽ’‡”•’‡ –‹˜‡ǡƒ ‘—‹–›ƒ›„‡†‡ˆ‹‡†ƒ•–Š‡ƒ••‡„Žƒ‰‡‘ˆ •’‡ ‹‡• ‘ —’›‹‰ ƒ ƒ”‡ƒ ™‹–Š‹ ™Š‹ Š ƒŽŽ ‹†‹˜‹†—ƒŽ• ƒ”‡ ‡“—ƒŽŽ›Ž‹‡Ž›–‘‹–‡”ƒ –ǡ–Š—• Š‹†‡”‹‰ •’ƒ–‹ƒŽ Š‡–‡”‘‰‡‡‹–› ‹ †‹•–”‹„—–‹‘ ‘” ƒ„—†ƒ ‡ ȋ ‘Ž›‘ƒ ‡– ƒŽǤ ʹͲͲͷȌǤ ‘™‡˜‡”ǡ™‡Ž‹˜‡‹ƒŠ‹‰ŠŽ›Š‡–‡”‘‰‡‡‘—•™‘”Ž†ǡƒ†‡˜‡–Š‡‘•–—‹ˆ‘”Šƒ„‹–ƒ–••Š‘™ ‹’‘”–ƒ–Ž‡˜‡Ž•‘ˆ•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‹‡˜‹”‘‡–ƒŽ ‘†‹–‹‘•ƒ–‘‡• ƒŽ‡‘”ƒ‘–Š‡”Ǥ ”‘ ƒ ‘”‡ †‡–‡”‹‹•–‹  ’‡”•’‡ –‹˜‡ǡ •’‡ ‹‡• ‘’‘•‹–‹‘ ‹• ‡š’‡ –‡† –‘ „‡ Ž‘•‡Ž› ”‡Žƒ–‡† –‘ –Š‹• ™‹–Š‹ǦŠƒ„‹–ƒ– Š‡–‡”‘‰‡‡‹–›ǡ ˆ‘” ‡šƒ’Ž‡ ‹ ”‡•‘—” ‡ ƒ˜ƒ‹Žƒ„‹Ž‹–› ȋ‹ Ž‡ˆ• ͳͻͺ͹ȌǤ ‘™‡˜‡”ǡ –Š‡ ‡ˆˆ‡ –• ‘ˆ ”‡•‘—” ‡ †‹•–”‹„—–‹‘ ‘ ‘—‹–› ‘’‘•‹–‹‘ ƒ› „‡ †‹ˆˆ‹ —Ž– –‘ ’”‡†‹ – ˆ‘” •‡˜‡”ƒŽ ”‡ƒ•‘•Ǥ ‹”•–ǡ †‹ˆˆ‡”‡– •’‡ ‹‡• ƒ› ”‡•’‘† –‘ ”‡•‘—” ‡ †‹•–”‹„—–‹‘ƒ–†‹ˆˆ‡”‡–• ƒŽ‡•Ǥƒ”‰‡•’‡ ‹‡•ǡ™‹–Š‰”‡ƒ–‡”ˆ‘‘†”‡“—‹”‡‡–•ƒ†‰”‡ƒ–‡”

 25 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

‘„‹Ž‹–›ƒ”‡‡š’‡ –‡†–‘”‡•’‘†–‘”‡•‘—” ‡†‹•–”‹„—–‹‘ƒ–Žƒ”‰‡” • ƒŽ‡• ȋ ‘ŽŽ‹‰ ͳͻͻʹȌǤ ƒŽŽ•’‡ ‹‡•ǡ‘–Š‡‘–Š‡”Šƒ†ǡƒ›„‡ƒ„Ž‡–‘•ƒ–‹•ˆ›–Š‡‹”‡‡†•™‹–Š‹ƒ•ƒŽŽƒ”‡ƒƒ† –Š‡”‡ˆ‘”‡„‡‘”‡•‡•‹–‹˜‡–‘Ž‘ ƒŽ • ƒŽ‡ˆƒ –‘”•Ǥ‡ ‘†ǡƒ‰‹˜‡•’‡ ‹‡•ƒ›†‡’‡†‘ ˜ƒ”‹‘—•”‡•‘—” ‡•™‹–Š†‹ˆˆ‡”‹‰†‹•–”‹„—–‹‘’ƒ––‡”•ǡƒ†–Š—•”‡•’‘†–‘‡ƒ Š”‡•‘—” ‡ƒ– ƒ†‹ˆˆ‡”‡–• ƒŽ‡ȋ‡•–”‹ Šͳͻͻ͸ȌǤ‘ ƒŽ ‘—‹–›•–”— –—”‡‹•ˆ—”–Š‡”•Šƒ’‡†„›•’‡ ‹‡•ǯ ˆ— –‹‘ƒŽ–”ƒ‹–•ǡ•— Šƒ•†‹•’‡”•ƒŽƒ„‹Ž‹–›ǡƒ†„›‹–‡”ƒ –‹‘•„‡–™‡‡•’‡ ‹‡•”‡•—Ž–‹‰‹ ‡‹–Š‡”ƒ˜‘‹†ƒ ‡‘”ƒ––”ƒ –‹‘ȋ‡•‡–ƒ”‹–•‡–ƒŽǤʹͲͲͷȌǤ ‹ƒŽŽ›ǡ ‘—‹–›•–”— –—”‡ƒ›„‡ Š‹•–‘”‹ ƒŽŽ› ‘–‹‰‡–ǡ •‘ –Šƒ– ‡˜‡ —†‡” •‹‹Žƒ” ‡˜‹”‘‡–ƒŽ ‘†‹–‹‘•ǡ †‹ˆˆ‡”‡– •’‡ ‹‡•ƒ••‡„Žƒ‰‡•ƒ›ƒ”‹•‡ƒ•ƒ”‡•—Ž–‘ˆ†‹ˆˆ‡”‡–‹‹‰”ƒ–‹‘Š‹•–‘”›‘”†‹•–—”„ƒ ‡ ‡˜‡–•ȋ —ƒ‹ʹͲͳͲȌǤ

–Š‹••–—†›™‡ƒƒŽ›œ‡–Š‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆƒ„‡‡ ‘—‹–› ƒ• ™‡ŽŽ ƒ• –Š‡ †‹•–”‹„—–‹‘ ‘ˆ –Š‡ ‡•–‹‰ ƒ† ˆŽ‘”ƒŽ ”‡•‘—” ‡• ‘ ™Š‹ Š „‡‡• †‡’‡†Ǥ ‘•– „‡‡ •’‡ ‹‡• „—‹Ž†‡•–•ƒ†’”‘˜‹•‹‘–Š‡™‹–Š’‘ŽŽ‡ƒ†‡ –ƒ”ƒ•ˆ‘‘†ˆ‘”–Š‡‹”Žƒ”˜ƒ‡Ǥ ‡ƒ„‡‡ Šƒ•‡•–ƒ„Ž‹•Š‡†ƒ–ƒ‡•–‹‰•‹–‡ǡ‹– ‘†— –•”‡’‡ƒ–‡†’‘ŽŽ‡Ǧ‡ –ƒ” ˆ‘”ƒ‰‹‰ –”‹’•ǡ –Š—• „‡ ‘‹‰ƒ ‡–”ƒŽ’Žƒ ‡ˆ‘”ƒ‰‡”Ǥ‡ ƒ—•‡†‹ˆˆ‡”‡–•’‡ ‹‡•—•‡†‹ˆˆ‡”‡–‡•–‹‰•—„•–”ƒ–‡• ƒ†ˆƒ˜‘—”†‹ˆˆ‡”‡–’‘ŽŽ‡•‘—” ‡•ǡ„‡‡†‹˜‡”•‹–›‹•‡š’‡ –‡†–‘„‡Š‹‰Š‡”‹ƒ”‡ƒ•Š‘•–‹‰ƒ ˜ƒ”‹‡–›‘ˆ‡•–‹‰ƒ†ˆŽ‘”ƒŽ”‡•‘—” ‡•ȋ‘—Ž•–‘ƒ† ‘‘†‡ŽŽʹͲͳͳȌǤ‘ŽŽ‡•’‡ ‹ƒŽ‹œƒ–‹‘‹ „‡‡•”ƒ‰‡•ˆ”‘’‘Ž›Ž‡ –›ȋ•’‡ ‹‡• ‘ŽŽ‡ –‹‰’‘ŽŽ‡ˆ”‘ƒ›—”‡Žƒ–‡†’Žƒ–ˆƒ‹Ž‹‡•Ȍǡ–‘ ‘Ž‹‰‘Ž‡ –› ȋ ‘ŽŽ‡ –‹‰ ’‘ŽŽ‡ ˆ”‘ ƒ •‹‰Ž‡ ’Žƒ– ˆƒ‹Ž›Ȍǡ ƒ† ‘‘Ž‡ –› ȋ ‘ŽŽ‡ –‹‰ ’‘ŽŽ‡ ˆ”‘ƒ•‹‰Ž‡’Žƒ–‰‡—•ȌǤ•ˆ‘”‡•–‹‰•—„•–”ƒ–‡•ǡ‘•–„‡‡•’‡ ‹‡•‡š ƒ˜ƒ–‡–Š‡‹”‡•–• —†‡”‰”‘—†ǡ„—–•‘‡†‘•‘‹†‡ƒ†™‘‘†‘”‹•‘ˆ–Ǧ’‹–Š•–‡•Ǥ–Š‡”•’‡ ‹‡•‡š’Ž‘‹– †‹ˆˆ‡”‡––›’‡•‘ˆ’”‡Ǧ‡š‹•–‹‰ ƒ˜‹–‹‡•ǡƒ†ƒ•ƒŽŽ‡”—„‡”„—‹Ž†‡š’‘•‡†‡•–•ƒ––ƒ Š‡† –‘”‘ •‘”–‘–Š‡˜‡‰‡–ƒ–‹‘Ǥ ‹ƒŽŽ›ǡ•‘‡„‡‡•’‡ ‹‡•ƒ”‡ Ž‡’–‘’ƒ”ƒ•‹–‹ ǡŽƒ›‹‰–Š‡‹”‡‰‰• ‹‡•–•‘ˆ‘–Š‡”„‡‡•’‡ ‹‡•ǡ—•—ƒŽŽ›‘ˆƒ‰‹˜‡‰‡—•Ǥ—„‡”‘ˆ•–—†‹‡•Šƒ˜‡†‘ —‡–‡† –Š‡ ‹ˆŽ—‡ ‡ ‘ˆ ˆŽ‘™‡” ”‡•‘—” ‡• ‘ –Š‡ •–”— –—”‡ ‘ˆ „‡‡ ‘—‹–‹‡• ȋ ƒ–Šƒ ‡– ƒŽǤ ͳͻͻͶǡ‘––•‡–ƒŽǤʹͲͲ͵ǡʹͲͲͶǡ Šƒˆˆ‡”•‡–ƒŽǤʹͲͲͺǡ ”—†‡Ž‡–ƒŽǤʹͲͳͲǡ ”ò†‡–ƒŽǤʹͲͳͲǡ ƒ•–ƒ‰‡›”‘Žƒ† ƒ –‡ŽʹͲͳʹǡ„‡Ž‹‰‡–ƒŽǤʹͲͳʹȌǤ ‡™‡”•–—†‹‡•Šƒ˜‡ƒ††”‡••‡†–Š‡”‘Ž‡‘ˆ ‡•–‹‰•—„•–”ƒ–‡•ȋŽ–œ‡–ƒŽǤʹͲͲ͵ǡ‘––•‡–ƒŽǤʹͲͲ͵ǡʹͲͲͷǡ ”—†‡Ž‡–ƒŽǤʹͲͳͲǡ—””ƒ›‡–ƒŽǤ ʹͲͳʹȌǡ ƒ† ‡•–ƒ„Ž‹•Š‹‰ –Š‡ ”‡Žƒ–‹˜‡ ‹’‘”–ƒ ‡ ‘ˆ ˆŽ‘™‡” versus‡•–‹‰”‡•‘—” ‡•Šƒ• „‡ ‘‡ƒ‡›–‘’‹ ‹„‡‡‡ ‘Ž‘‰›”‡•‡ƒ” Šȋ–‡ˆˆƒǦ‡™‡–‡”ƒ†‡•–’ŠƒŽʹͲͲͺȌǤŠ‹Ž‡ ƒ––ƒ Š‹‰ƒ‰”‡ƒ–‡”™‡‹‰Š––‘ˆŽ‘™‡””‡•‘—” ‡•ǡ–Š‡”‡˜‹‡™‘ˆ‘—Ž•–‘ƒ† ‘‘†‡ŽŽȋʹͲͳͳȌ ‡’Šƒ•‹œ‡• –Š‡ ‡‡† –‘ ‘•‹†‡” „‘–Š –›’‡• ‘ˆ ”‡•‘—” ‡•ǡ ’ƒ”–Ž›„‡ ƒ—•‡‘ˆ–Š‡•’ƒ–‹ƒŽ

 26 Šƒ’–‡”ͳ

‘’Ž‡š‹–›‘ˆ”‡•‘—” ‡†‹•–”‹„—–‹‘ƒ†’ƒ”–Ž›„‡ ƒ—•‡‡•–‹‰•—„•–”ƒ–‡†‹˜‡”•‹–›‹•‘ˆ–‡ ‘””‡Žƒ–‡†™‹–Š’Žƒ–†‹˜‡”•‹–›Ǥ

‡‡• ƒ”‡ ƒ„Ž‡ –‘ ˆŽ› Ž‘‰ †‹•–ƒ ‡• ƒ† –Š‡”‡ˆ‘”‡ Šƒ˜‡ –Š‡ ƒ’ƒ ‹–› –‘ ”‡ƒ†‹Ž› ‘Ž‘‹œ‡ •—‹–ƒ„Ž‡•‹–‡•Ǥ‡˜‡”ƒŽ•–—†‹‡•Šƒ˜‡‡•–‹ƒ–‡†„‡‡ˆ‘”ƒ‰‹‰”ƒ‰‡•–Š”‘—‰Š–Š‡—•‡‘ˆ˜ƒ”‹‘—• –‡ Š‹“—‡•ǡ ‹ Ž—†‹‰ ‡ƒ•—”‡• ‘ˆ –”‹’ †—”ƒ–‹‘ǡ ‡š’‡”‹‡–• ‘ˆ Š‘‹‰ ƒ„‹Ž‹–›ǡ Šƒ”‘‹  ”ƒ†ƒ”–”ƒ ‹‰ǡƒ”Ǧ”‡ ƒ’–—”‡‡š’‡”‹‡–•ƒ†‰‡‡–‹ ƒƒŽ›•‹•‘ˆˆ‘”ƒ‰‹‰„‡‡•ȋ•„‘”‡ ‡–ƒŽǤͳͻͻͻǡƒŽ–Š‡”Ǧ ‡ŽŽ™‹‰ƒ† ”ƒŽʹͲͲͲǡ ƒ–Šƒƒ†• Šƒ”–‡ʹͲͲʹǡ‡•–’ŠƒŽ‡– ƒŽǤ ʹͲͲ͸ǡ ”‡‡Ž‡ƒˆ ‡– ƒŽǤ ʹͲͲ͹ǡ —±†‘– ‡– ƒŽǤ ʹͲͲͻǡ ‘—Ž•‘ ʹͲͳͲǡ —”„— Š‡ ‡– ƒŽǤ ʹͲͳͲǡ ƒ”˜‡ŽŽ‡–ƒŽǤʹͲͳʹȌǤŠ‡•‡•–—†‹‡•‹†‹ ƒ–‡–Šƒ–‘•–•’‡ ‹‡•ˆ‘”ƒ‰‡™‹–Š‹ƒˆ‡™Š—†”‡† ‡–‡”• ˆ”‘ –Š‡‹” ‡•– „—– •‘‡ ƒ› ˆŽ› –Š‘—•ƒ†• ‘ˆ ‡–‡”•Ǥ Š‡•‡ •–—†‹‡• ƒŽ•‘ •Š‘™ ƒ ‘•‹•–‡–’‘•‹–‹˜‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡„‘†›•‹œ‡ƒ†‡•–‹ƒ–‡†ˆ‘”ƒ‰‹‰†‹•–ƒ ‡Ǥ‡ƒ› –Š—•‡š’‡ –•’‡ ‹‡•‘ˆ†‹ˆˆ‡”‡–„‘†›•‹œ‡•–‘”‡•’‘††‹ˆˆ‡”‡–Ž› –‘ •’ƒ–‹ƒŽ ”‡•‘—” ‡ †‹•–”‹„—–‹‘Ǥ

”‡˜‹‘—••–—†‹‡•Šƒ˜‡•Š‘™†‹ˆˆ‡”‡ ‡•‹„‡‡ ‘—‹–› ‘’‘•‹–‹‘ƒ–Žƒ†• ƒ’‡• ƒŽ‡• ƒ† ‹ ”‡Žƒ–‹‘ –‘ Šƒ„‹–ƒ– ‘’‘•‹–‹‘ǡ ƒ–Š”‘’‘‰‡‹  Žƒ† —•‡ Šƒ‰‡ ƒ† ˆ”ƒ‰‡–ƒ–‹‘ ȋ–‡ˆˆƒǦ‡™‡–‡”‡–ƒŽǤʹͲͲʹǡ”‘•‹‡–ƒŽǤʹͲͲͺǡ‹ ‡––•‡–ƒŽǤ ʹͲͲͺǡ ‹ˆ”‡‡ ‡– ƒŽǤ ʹͲͲͻǡ —””ƒ› ‡– ƒŽǤ ʹͲͳʹǡ ‹ƒƒ ‡– ƒŽǤ ʹͲͳʹȌǤ ‘™‡˜‡”ǡ ™‡ ‘™ ‘ˆ ‘ •–—†‹‡• ‡š’Ž‘”‹‰ –Š‡ †‹•–”‹„—–‹‘ ‘ˆ ƒ ‡–‹”‡ „‡‡ ‘—‹–› ƒ– ƒ Ž‘ ƒŽ • ƒŽ‡ ™‹–Š‹ƒŠƒ„‹–ƒ––Šƒ–ƒ›„‡ ‘•‹†‡”‡† Š‘‘‰‡‡‘—• ƒ– ƒ Žƒ†• ƒ’‡ • ƒŽ‡Ǥ Š‹• • ƒŽ‡ ‹• ‹’‘”–ƒ– „‡ ƒ—•‡ ‘•– ‹†‹˜‹†—ƒŽ„‡‡‘˜‡‡–•’”‘„ƒ„Ž›‘ —”ƒ––Š‹•• ƒŽ‡Ǥ—”•–—†›™ƒ• ‘†— –‡†‹ƒƒ”‡ƒ ‘˜‡”‡†„› ‘–‹‰—‘—•‡†‹–‡””ƒ‡ƒ• ”—„Žƒ†ǡ™‹–Š—‹ˆ‘” Ž‹ƒ–‹  ‘†‹–‹‘•ƒ†‘ ‡ ‘Ž‘‰‹ ƒŽ ‘” ’Š›•‹ ƒŽ „ƒ””‹‡”•Ǥ ‡ ƒ—•‡ „‡‡• ƒ”‡ Š‹‰ŠŽ› ‘„‹Ž‡ǡ ‘‡ ‹‰Š– ‡š’‡ – ™‹–Š‹Ǧ Šƒ„‹–ƒ–†‹ˆˆ‡”‡ ‡•‹„‡‡†‹•–”‹„—–‹‘–‘„‡•ƒŽŽǤ ‘™‡˜‡”ǡƒˆ‡™•–—†‹‡•Šƒ˜‡•Š‘™–Šƒ– ’‘ŽŽ‹ƒ–‘”ƒ••‡„Žƒ‰‡•˜‹•‹–‹‰˜ƒ”‹‘—•’Žƒ–Ǧ•’‡ ‹‡•ƒ›˜ƒ”›ƒ–• ƒŽ‡•‘ˆŠ—†”‡†•‘”‡˜‡ –‡•‘ˆ‡–‡”•ȋ ‡””‡”ƒͳͻͺͺǡ‹ Ž‡›‡–ƒŽǤͳͻͻͻǡ ƒ‘˜•ý‡–ƒŽǤʹͲͳ͵ȌǤ

‘•–‘†‡Ž•‘ ‘—‹–›ƒ••‡„Ž›†›ƒ‹ •ƒ••—‡–Šƒ–‡˜‹”‘‡–ƒŽ ‘†‹–‹‘•ƒ”‡ Š‘‘‰‡‡‘—•ƒ ”‘••ƒ’ƒ– Š‘ˆ—‹ˆ‘”Šƒ„‹–ƒ–ǡƒŽ–Š‘—‰Š–Š‹•ƒ••—’–‹‘‹• Ž‡ƒ”Ž›‘–‡– ‹ƒ›•›•–‡•ȋ —ƒ‹ʹͲͳͲȌǤ—”ˆ‹”•–‘„Œ‡ –‹˜‡‹•–‘ƒƒŽ›œ‡ˆŽ‘”ƒŽƒ†‡•–‹‰”‡•‘—” ‡ ‘’‘•‹–‹‘Š‡–‡”‘‰‡‡‹–›ƒ––Š‡Šƒ„‹–ƒ–• ƒŽ‡Ǥ‡ƒ•™Š‡–Š‡”–Š‹• Š‡–‡”‘‰‡‡‹–› ‹• ‹””‡‰—Žƒ” ‘” ‡Ž•‡ ˆ‘ŽŽ‘™• ƒ ‰‡‘‰”ƒ’Š‹ ƒŽ ’ƒ––‡” ƒ ”‘•• –Š‡ • ”—„Žƒ† ƒ– „‡‡Ǧˆ‘”ƒ‰‹‰ ”‡Ž‡˜ƒ–†‹•–ƒ ‡•Ǥ—”•‡ ‘†‘„Œ‡ –‹˜‡‹•–‘ƒƒŽ›œ‡–Š‡†‹•–”‹„—–‹‘‘ˆ„‡‡ ‘’‘•‹–‹‘

 27 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

ƒ ”‘•• –Š‡ Šƒ„‹–ƒ–Ǥ  Š‘‘‰‡‡‘—• †‹•–”‹„—–‹‘ ™‘—Ž† „‡ ‹ ƒ‰”‡‡‡– ™‹–Š –Š‡ ƒ„‘˜‡Ǧ ‡–‹‘‡†–Š‡‘”‡–‹ ƒŽ†‡ˆ‹‹–‹‘‘ˆ ‘—‹–›ȋ ‘Ž›‘ƒ‡–ƒŽǤʹͲͲͷȌǡƒ†™‘—Ž†”‡ˆŽ‡ –Š‹‰Š Ž‡˜‡Ž• ‘ˆ ‘‡ –‹‘ ƒ‘‰ ’Ž‘–•ǡ ‡‹–Š‡” –Š”‘—‰Š ˆ‘”ƒ‰‹‰ ‘˜‡‡–•ǡ –Š”‘—‰Š †‹•’‡”•ƒŽ ”ƒ–‡•ǡ‘”„‘–ŠǤ ‹˜‡–Š‡•‹œ‡‘ˆ–Š‡ƒ”‡ƒ•ƒ’Ž‡†ȋͷǤͶ„›͸ǤʹȌƒ†–Š‡Š‹‰Š†‡‰”‡‡‘ˆ ‘„‹Ž‹–›†‹•’Žƒ›‡†„›„‡‡•ǡ™‡ƒ••—‡–Šƒ–ƒ›„‡‡•’‡ ‹‡•‹•ƒ„Ž‡–‘ ‘Ž‘‹œ‡ƒ•—‹–ƒ„Ž‡’Ž‘– ‹‘—”•–—†›ƒ”‡ƒ‘˜‡”‘‡‘”ƒˆ‡™‰‡‡”ƒ–‹‘•ǤŽ–‡”ƒ–‹˜‡Ž›ǡ„‡‡ ‘’‘•‹–‹‘‹‰Š–•Š‘™ ƒŠ‡–‡”‘‰‡‡‘—•†‹•–”‹„—–‹‘‹ˆ„‡‡ˆ‘”ƒ‰‹‰ƒ”‡ƒ•™‡”‡•ƒŽŽƒ†„‡‡†‹•–”‹„—–‹‘ Ž‘•‡Ž› –”ƒ ‡† •’ƒ–‹ƒŽ ˜ƒ”‹ƒ–‹‘ ‹ ”‡•‘—” ‡ ƒ˜ƒ‹Žƒ„‹Ž‹–› ƒ– –Š‡ Ž‘ ƒŽ• ƒŽ‡Ǥ ˆ–Š‡Žƒ––‡”ǡ™‡ƒ• ™Š‡–Š‡” „‡‡• ”‡•’‘† ’”‹ƒ”‹Ž› –‘ ˆŽ‘™‡” ‘” –‘ ‡•–‹‰ ”‡•‘—” ‡ †‹•–”‹„—–‹‘Ǥ —” „‡‡ ‘—‹–›‹•”‹ Šȋͻͺ•’‡ ‹‡•Ȍƒ†‡ ‘’ƒ••‡•ƒ™‹†‡”ƒ‰‡‘ˆ„‘†›•‹œ‡•ƒ†–Š‡”‡ˆ‘”‡ ’”‡•—‡†‡‡”‰‡–‹ ”‡“—‹”‡‡–•ƒ†‘„‹Ž‹–›Ǥ—”–Š‹”†‘„Œ‡ –‹˜‡‹•–‘‡•–ƒ„Ž‹•Š™Š‡–Š‡” •’‡ ‹‡•™‹–Š†‹ˆˆ‡”‡–„‘†›•‹œ‡•”‡•’‘††‹ˆˆ‡”‡–Ž›–‘Ž‘ ƒŽ”‡•‘—” ‡ƒ˜ƒ‹Žƒ„‹Ž‹–›ƒ†•Š‘™ †‹ˆˆ‡”‡–’ƒ––‡”•‘ˆ•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘Ǥ



1.2. MaterialsandMethods

1.2.1. Studyarea

Š‡•–—†›™ƒ• ‘†— –‡†‹–Š‡ƒ–—”ƒŽƒ”‘ˆ ƒ””ƒˆȋƒ” ‡Ž‘ƒǡ’ƒ‹ȌǤ‡•‡Ž‡ –‡† ʹͳ’Ž‘–•‘ˆͶͲšͶͲ†‹•–”‹„—–‡†‘”‡‘”Ž‡••”‡‰—Žƒ”Ž›ƒ ”‘••–Š‡’ƒ”ǡ‡ ‘’ƒ••‹‰ƒ ‘˜‡”ƒŽŽƒ”‡ƒ‘ˆ͵ʹʹǤ‹•–ƒ ‡•„‡–™‡‡‡ƒ”‡•–’Ž‘–•”ƒ‰‡†ˆ”‘ͷͺͷ–‘ͳ͵ͷͶǤŠ‡ –™‘‘•–†‹•–ƒ–’Ž‘–•™‡”‡͸Ǥʹƒ’ƒ”–ǤŽ‘–•”ƒ‰‡†‹ƒŽ–‹–—†‡ˆ”‘ʹͷͷ–‘ͷͶͷǡƒ† –Š‡‹”†‹•–ƒ ‡–‘–Š‡ ‘ƒ•–”ƒ‰‡†ˆ”‘ͳͷͲͲ–‘͸ͺͲͲǤ–ƒŽƒ†• ƒ’‡• ƒŽ‡ǡ–Š‡•–—†›ƒ”‡ƒ ƒ„‡ ‘•‹†‡”‡†Š‘‘‰‡‡‘—•ǤŠ‡ʹͳ•‡Ž‡ –‡†’Ž‘–••Šƒ”‡–Š‡•ƒ‡˜‡‰‡–ƒ–‹‘–›’‡ǡ•‘‹Ž –›’‡ ƒ† ”‡ ‡– †‹•–—”„ƒ ‡ Š‹•–‘”›Ǥ Š›•‹ ƒŽ ‘” ‡˜‹”‘‡–ƒŽ „ƒ””‹‡”• ƒ”‡ Žƒ ‹‰ ƒ† –Š‡”‡ƒ”‡‘•‹‰‹ˆ‹ ƒ– Ž‹ƒ–‹ ‰”ƒ†‹‡–•ǤŠ‡’ƒ”‹•Ž‘ ƒ–‡† ‘ ƒ ƒ”•–‹  ƒ••‹ˆ ‘ˆ Ž‹‡•–‘‡ ƒ† †‘Ž‘•–‘‡Ǥ Š‹• •‘‹Ž –›’‡ ˆƒ˜‘—”• †”ƒ‹ƒ‰‡ǡ –Š—• Š‹†‡”‹‰ ™ƒ–‡” •–‘”ƒ‰‡Ǥ –”‡ƒ„‡†•ƒ”‡Žƒ ‹‰ƒ†‘‡‘ˆ–Š‡’Ž‘–•‹•Ž‘ ƒ–‡†ƒ––Š‡„‘––‘‘ˆƒ˜ƒŽŽ‡›ǤŠ‡ƒ”‡ƒ‹• ‘ —’‹‡†„›ƒ‡†‹–‡””ƒ‡ƒ• ”—„Žƒ†ǤŽƒ– ‘’‘•‹–‹‘˜ƒ”‹‡•Ž‘ ƒŽŽ›ˆ”‘’Ž‘––‘’Ž‘–ǡ „—– ‹• ƒŽ™ƒ›• Žƒ”‰‡Ž› †‘‹ƒ–‡† „› Quercus cocciferaǡ Pistacia lentiscusǡ Rosmarinus officinalisƒ†Thymusvulgaris.

  28 Šƒ’–‡”ͳ

1.2.2. Beesampling

‡ ‘†— –‡† ͺ •—”˜‡›• ȋ‘‡ ‡˜‡”› –™‘ ™‡‡•Ȍ ˆ”‘ ‹† ƒ” Š –‘ Žƒ–‡ —‡ ʹͲͳͲǡ –Š—• ‡ ‘’ƒ••‹‰–Š‡ƒ‹ˆŽ‘™‡”‹‰’‡”‹‘†‘ˆ–Š‡• ”—„Žƒ†ȋˆŽ‘™‡”•ƒ”‡˜‡”›• ƒ” ‡‹ —Ž› ƒ† —‰—•–ȌǤ ‘ ƒ˜‘‹† –Š‡ ‹ˆŽ—‡ ‡ ‘ˆ ™‡ƒ–Š‡” ‘†‹–‹‘•ǡ •—”˜‡›• ™‡”‡ ‘†— –‡† •‹—Ž–ƒ‡‘—•Ž› ‹ ƒŽŽ ’Ž‘–•Ǥ  ‡ƒ Š •—”˜‡› ™‡ ’Žƒ ‡† ͸ •ƒ’Ž‹‰ •–ƒ–‹‘• ‹ –™‘ ’ƒ”ƒŽŽ‡Ž ”‘™•ǡ ™‹–Š ƒ †‹•–ƒ ‡ ‘ˆ ͳͲ  „‡–™‡‡ •–ƒ–‹‘•Ǥ ‘ŽŽ‘™‹‰ ‡•–’ŠƒŽ ‡– ƒŽǤ ȋʹͲͲͺȌǡ ‡ƒ Š •–ƒ–‹‘ ™ƒ• ‘’‘•‡† ‘ˆ ƒ ‡–ƒŽ „ƒ” Š‘Ž†‹‰ ͵ ’ƒ –”ƒ’• ȋͳͷǦ Ǧ†‹ƒ‡–‡”’Žƒ•–‹ „‘™Ž• ’ƒ‹–‡†›‡ŽŽ‘™ǡ™Š‹–‡ƒ†„Ž—‡ǡ”‡•’‡ –‹˜‡Ž›ǡ™‹–ŠǦ”‡ˆŽ‡ –‹‰’ƒ‹–ȌǤ”ƒ’•™‡”‡Ž‘ ƒ–‡† ƒ–ʹͲǦͶͲ ƒ„‘˜‡‰”‘—†Ž‡˜‡Žƒ†ƒ’’”‘š‹ƒ–‡Ž›ͷͲ ƒ™ƒ›ˆ”‘–Š‡‡ƒ”‡•–ˆŽ‘™‡”‹‰ ’Žƒ–Ǥ ‡ˆ‘”‡ ͻǣ͵Ͳ ‘ ‡ƒ Š •ƒ’Ž‹‰ †ƒ›ǡ –”ƒ’• ™‡”‡ ˆ‹ŽŽ‡† ™‹–Š ™ƒ–‡” ‘–ƒ‹‹‰ ƒ •ƒŽŽ ƒ‘—– ‘ˆ †‡–‡”‰‡– ƒ† ‘ŽŽ‡ –‡† ƒˆ–‡” ͳͺǣͲͲǡ –Š—• ‘˜‡”‹‰ ‘•– ‘ˆ –Š‡ †ƒ‹Ž› ƒ –‹˜‹–› ’‡”‹‘†Ǥ ƒ –”ƒ’’‹‰ Šƒ• „‡‡ •Š‘™ –‘ —†‡”‡•–‹ƒ–‡ „‡‡ ”‹ Š‡•• ƒ† –‘ ’”‘˜‹†‡ ƒ ‹ ‘’Ž‡–‡ ‡ƒ•—”‡ ‘ˆ ˆŽ‘™‡” ˜‹•‹–ƒ–‹‘ ‘’ƒ”‡† –‘ ‡––‹‰ ‘ˆ ˆŽ‘™‡” ˜‹•‹–‹‰ ‹•‡ –• ȋ‡•–’ŠƒŽ‡–ƒŽǤʹͲͲͺǡ‘’‹ ‡–ƒŽǤʹͲͳ͵ȌǤ ‘™‡˜‡”ǡ‘—”ƒ‹ ‘ ‡”•™‡”‡–‘•ƒ’Ž‡ƒŽŽʹͳ ’Ž‘–••‹—Ž–ƒ‡‘—•Ž›ǡ–‘ƒ˜‘‹† ‘ŽŽ‡ –‘”„‹ƒ•ǡƒ†–‘ƒ’’Ž›–Š‡•ƒ‡•ƒ’Ž‹‰‡ˆˆ‘”––‘‡ƒ Š ’Ž‘–Ǥ —” ‰‘ƒŽ ™ƒ• –‘ Šƒ”ƒ –‡”‹œ‡ –Š‡ „‡‡ ‘—‹–›ǡ ”ƒ–Š‡” –Šƒ •ƒ’Ž‡ „‡‡ǦˆŽ‘™‡” ‹–‡”ƒ –‹‘•Ǥ

ƒ’–—”‡†•’‡ ‹‡•™‡”‡†”‹‡†ƒ†’‹‡†ˆ‘”‹†‡–‹ˆ‹ ƒ–‹‘‹–Š‡Žƒ„‘”ƒ–‘”›Ǥ ”‘–Š‡•‡ •ƒ’Ž‡• ™‡ ‘„–ƒ‹‡† ‡ƒ•—”‡• ‘ˆ •’‡ ‹‡• ”‹ Š‡•• ȋ—„‡” ‘ˆ •’‡ ‹‡• ƒ’–—”‡†Ȍǡ ƒ„—†ƒ ‡ȋ—„‡”‘ˆ‹†‹˜‹†—ƒŽ• ƒ’–—”‡†Ȍƒ† ‘’‘•‹–‹‘ȋƒ„—†ƒ ‡‘ˆ‡ƒ Š•’‡ ‹‡•Ȍ ˆ‘”‡ƒ Š’Ž‘–Ǥ ”‡•Š„‘†›™‡‹‰Š–•™‡”‡‘„–ƒ‹‡†ˆ”‘‡––‡†•’‡ ‹‡•ǤŽŽ•’‡ ‹‡•™‡”‡ ™‡‹‰Š‡† ƒ ˆ‡™ Š‘—”• ƒˆ–‡” „‡‹‰ ƒ’–—”‡† ƒ†ǡ ‹ƒ•— Š ƒ• ’‘••‹„Ž‡ǡ ™‡ ‡ƒ•—”‡† ‘”‡ –Šƒ‘‡•’‡ ‹‡’‡”•’‡ ‹‡•ȋ‡ƒα͹ǤͶǢ”ƒ‰‡αͳǦͷʹȌǤ‡—•‡ˆ‡ƒŽ‡™‡‹‰Š–‹ƒŽŽ ƒƒŽ›•‡•Ǥ



1.2.3. Flowerresources

‘‡•–‹ƒ–‡ˆŽ‘™‡””‹ Š‡••™‡ ‘—–‡†ƒŽŽˆŽ‘™‡”•’‡ ‹‡•ƒŽ‘‰–™‘ͶͲšͳ–”ƒ•‡ –• ƒ””ƒ‰‡†ƒ•ƒ ‡–”‡†‹–Š‡ ‡–”‡‘ˆ–Š‡•ƒ’Ž‹‰•–ƒ–‹‘‰”‹†Ǥ Š‹• ™ƒ• †‘‡ –Š”‡‡ –‹‡•ǡ‹‹†’”‹Žǡ‹†ƒ›ƒ†‹† —‡Ǥ ƒ††‹–‹‘ǡ™‡‡•–‹ƒ–‡†ˆŽ‘™‡”†‡•‹–›‘ˆ–Š‡ ƒ‹ ˆŽ‘™‡”‹‰ •’‡ ‹‡• ȋR. officinalisǡ T. vulgarisǡ Dorycnium pentaphyllum, Cistus albidusǡ

 29 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

Cistussalvifoliusƒ†CistusmonspeliensisȌ‹‡ƒ Š’Ž‘–ǤŠ‡•‡•’‡ ‹‡•”‡’”‡•‡–͹ͲǦͻͲΨ‘ˆ –Š‡ ˆŽ‘™‡”• ’”‘†— ‡† ‹ –Š‡ •–—†› ƒ”‡ƒ ȋ—’—„Ž‹•Š‡† †ƒ–ƒ ˆ”‘ ™‡‡Ž› ˆŽ‘™‡” ‘—–• ‹ –”ƒ•‡ –•ƒ–ͳʹ†‹ˆˆ‡”‡–•‹–‡•ƒ ”‘••–Š‡’ƒ”ȌǤ‡ˆ‹”•– ƒŽ —Žƒ–‡†–Š‡˜‘Ž—‡‘ˆ‡ƒ ŠˆŽ‘™‡” ’ƒ– Š ‹ –Š‡ –”ƒ•‡ –• „› ‡ƒ•—”‹‰ –™‘ ’‡”’‡†‹ —Žƒ” ™‹†–Š• ƒ† –Š‡ Š‡‹‰Š–Ǥ Š‡ǡ –‘ ‡•–ƒ„Ž‹•Šƒ”‡Žƒ–‹‘•Š‹’„‡–™‡‡’ƒ– Š˜‘Ž—‡ƒ†—„‡”‘ˆˆŽ‘™‡”•ǡ™‡ ‘—–‡†ƒŽŽ‘’‡ ˆŽ‘™‡”•‹ƒ•—„•ƒ’Ž‡‘ˆ’ƒ– Š‡•ȋαͷͻǦʹʹ͸’‡”•’‡ ‹‡•Ȍƒ–’‡ƒ„Ž‘‘ȋ‹‡ƒ” ”‡‰”‡••‹‘ǣ”ʹαͲǤ͵͸ǦͲǤ͸͵ǡ’αͲǤͲͲͳǦͲǤͲͳͷȌǤŠ‡–Š”‡‡Cistus•’‡ ‹‡•™‡”‡• ƒ” ‡ ‘’ƒ”‡† –‘–Š‡‘–Š‡”•’‡ ‹‡•ƒ†–Š‡‹”„Ž‘‘‹‰’‡”‹‘†•‘˜‡”Žƒ’’‡†™‹†‡Ž›ǤŠ‡”‡ˆ‘”‡ǡ™‡Ž—’‡† –‘‰‡–Š‡”–Š‡–Š”‡‡•’‡ ‹‡•‹ƒ•‹‰Ž‡˜ƒ”‹ƒ„Ž‡ȋCistusˆŽ‘™‡”•ȌǤ ƒƒ––‡’––‘–‡ƒ•‡ƒ’ƒ”– –Š‡‡ˆˆ‡ –•‘ˆ’‘ŽŽ‡ƒ†‡ –ƒ”™‡—•‡†‡ƒ•—”‡•‘ˆ’‘ŽŽ‡ƒ† ‡ –ƒ” ’”‘†— –‹‘ ’‡” ˆŽ‘™‡”‘ˆ‡ƒ Š•’‡ ‹‡•ȋ—’—„Ž‹•Š‡††ƒ–ƒȌ–‘‡•–‹ƒ–‡’‘ŽŽ‡ƒ†‡ –ƒ”†‡•‹–›‹‡ƒ Š’Ž‘–Ǥ ‘™‡˜‡”ǡ–Š‡•‡–™‘˜ƒ”‹ƒ„Ž‡•™‡”‡Š‹‰ŠŽ› ‘””‡Žƒ–‡†ȋ”αͲǤͻ͸ǡ’δͲǤͲͲͲͳȌǡƒ†–Š‡›™‡”‡ ƒŽ•‘ ‘””‡Žƒ–‡†–‘ˆŽ‘™‡”†‡•‹–›ȋ”αͲǤͺʹǡ’δͲǤͲͲͲͳƒ†”αͲǤ͹͹ǡ’δͲǤͲͲͲͳǡ”‡•’‡ –‹˜‡Ž›ȌǤ Š‡”‡ˆ‘”‡ǡ™‡—•‡ˆŽ‘™‡”†‡•‹–›‹ƒŽŽƒƒŽ›•‡•Ǥ



1.2.4. Nestingsubstrates

‡ —•‡† –Š‡ ƒ„‘˜‡Ǧ‡–‹‘‡† –”ƒ•‡ –• –‘ ‡ƒ•—”‡ ƒ˜ƒ‹Žƒ„‹Ž‹–› ‘ˆ ‡•–‹‰ •—„•–”ƒ–‡•Ǥ  ‡˜‡”›ʹ‘ˆ–”ƒ•‡ –™‡’Žƒ ‡†ƒ™‹”‡‰”‹††‡Ž‹‹–‹‰͵ʹ ‡ŽŽ•ȋ‡ƒ Š‡ƒ•—”‹‰ͲǤͲ͵ͳʹȌǡ ƒ†‡ƒ Š ‡ŽŽ™ƒ•• ‘”‡†ƒ• ‘–ƒ‹‹‰‘‡‘”‘’‘–‡–‹ƒŽ‡•–‹‰•—„•–”ƒ–‡•Ǥ‡—•‡†–Š‡ ˆ‘ŽŽ‘™‹‰‡•–‹‰•—„•–”ƒ–‡˜ƒ”‹ƒ„Ž‡•ǣΨ„ƒ”‡•‘‹ŽǡΨ„ƒ”‡•‘‹Ž™‹–Š•–‘‡•ǡ’”‡•‡ ‡‘ˆ†‡ƒ† ™‘‘†ǡ—„‡”‘ˆŠ‘Ž‡•‹”‘ •ǡ—„‡”‘ˆ˜ƒ ƒ–•ƒ‹Ž•Š‡ŽŽ•ǡΨQuercuscoccifera ‘˜‡”ǡ ƒ†ΨAmpelodesmosmauritanica ‘˜‡”ǤQuercuscoccifera™ƒ•‹ Ž—†‡†„‡ ƒ—•‡™‡‘ˆ–‡ ‘„•‡”˜‡† Bombus terrestris „—„Ž‡„‡‡• ‡•–‹‰ ƒ– –Š‡‹” „ƒ•‡Ǥ Ampelodesmos mauritanica ™ƒ•‹ Ž—†‡†„‡ ƒ—•‡‹–’”‘†— ‡••‘ˆ–Ǧ’‹–Šƒ†Š‘ŽŽ‘™•–‡•–Šƒ–‹‰Š–„‡—•‡†„›•‘‡ „‡‡•’‡ ‹‡•‹–Š‡‰‡‡”ƒCeratina,Heriades,Protosmiaƒ†Hoplitis.



1.2.5. Statisticalanalysis

ŽŽ ˆŽ‘™‡” ”‡•‘—” ‡ ˜ƒ”‹ƒ„Ž‡• ™‡”‡ •“—ƒ”‡Ǧ”‘‘– –”ƒ•ˆ‘”‡† –‘ ‹’”‘˜‡ ‘”ƒŽ‹–› ƒ† Š‘‘• ‡†ƒ•–‹ ‹–›Ǥ ‡•–‹‰ ”‡•‘—” ‡ ˜ƒ”‹ƒ„Ž‡• ™‡”‡ Ž‘‰ –”ƒ•ˆ‘”‡†ǡ ‡š ‡’– Q. coccifera

 30 Šƒ’–‡”ͳ

‘˜‡”ǡ ™Š‹ Š ™ƒ• •“—ƒ”‡Ǧ”‘‘– –”ƒ•ˆ‘”‡†Ǥ ƒ”‡ •‘‹Ž ƒ† „ƒ”‡ •‘‹Ž ™‹–Š •–‘‡• ™‡”‡ •‹‰‹ˆ‹ ƒ–Ž› ‘””‡Žƒ–‡†ȋ”αͲǤ͸͹ǡ’αͲǤͲͲͳȌƒ†–Š—•™‡Ž—’‡†–Š‡–‘‰‡–Š‡”‹ƒ•‹‰Ž‡ ˜ƒ”‹ƒ„Ž‡ ȋ„ƒ”‡ •‘‹Ž ‘˜‡”ȌǤ Š‡ ”‡ƒ‹‹‰ ”‡•‘—” ‡ ˜ƒ”‹ƒ„Ž‡• ™‡”‡ ‘– •‹‰‹ˆ‹ ƒ–Ž› ‘””‡Žƒ–‡†Ǥ

‡ —•‡† ‘”ƒǯ•  ‘””‡Ž‘‰”ƒ• –‘ ‡š’Ž‘”‡ •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‘ˆ ˆŽ‘™‡” ”‹ Š‡••ǡ ˆŽ‘™‡” †‡•‹–›‘ˆ‡ƒ Š•ƒ’Ž‡†•’‡ ‹‡•ǡ‘˜‡”ƒŽŽˆŽ‘™‡”ƒ„—†ƒ ‡ǡ ‘˜‡”‘ˆ‡ƒ Š‡•–‹‰•—„•–”ƒ–‡ǡ „‡‡ •’‡ ‹‡• ”‹ Š‡••ǡ ‘˜‡”ƒŽŽ „‡‡ ƒ„—†ƒ ‡ǡ ƒ† „‡‡ ƒ„—†ƒ ‡ ‘ˆ‡ƒ Š‘ˆ–Š‡ͳͻ‘•– ƒ„—†ƒ–•’‡ ‹‡•ȋ–Š‘•‡”‡’”‡•‡–‹‰‘”‡–ŠƒͲǤͷΨ‘ˆ–Š‡–‘–ƒŽ‹†‹˜‹†—ƒŽ• ƒ’–—”‡†ȌǤ ‘” –Š‡˜ƒ”‹ƒ„Ž‡Dz’”‡•‡ ‡‘ˆ†‡ƒ†™‘‘†dz™‡—•‡†–Š‡„‹ƒ”› ‘‹Ǧ‘—– ‘””‡Ž‘‰”ƒǤ ‘ ‡š’Ž‘”‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆ„‡‡ ‘—‹–› ‘’‘•‹–‹‘ǡ™‡—•‡†ƒƒ–‡Žǯ• ‘””‡Ž‘‰”ƒ ‘„–ƒ‹‡†ˆ”‘ƒƒ–”‹š‘ˆ‰‡‘‰”ƒ’Š‹ ƒŽ†‹•–ƒ ‡•ƒ†ƒƒ–”‹š‘ˆ •‹‹Žƒ”‹–› ȋޔ‡•‡ǯ• ‹†‡šȌ ‘ˆ „‡‡ •’‡ ‹‡• ‘’‘•‹–‹‘Ǥ Š‡ —„‡” ‘ˆ ‹–‡”˜ƒŽ• ‹ ƒŽŽ ‘””‡Ž‘‰”ƒ• ™ƒ• ƒŽ —Žƒ–‡†„ƒ•‡†‘–—”‰‡ǯ•”—Ž‡Ǥ‹‰‹ˆ‹ ƒ ‡‘ˆ‡ƒ Š ‘””‡Ž‘‰”ƒ™ƒ•–‡•–‡†–Š”‘—‰Š͵ͲͲ ’‡”—–ƒ–‹‘•ƒ†’Ǧ˜ƒŽ—‡•™‡”‡ƒ’’Ž‹‡†ƒ’”‘‰”‡••‹˜‡‘ˆ‡””‘‹ ‘””‡ –‹‘Ǥ ‘ ˆ—”–Š‡” ‡š’Ž‘”‡ •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‘ˆ „‡‡ ‘’‘•‹–‹‘ǡ ™‡ ”— ƒ Ž—•–‡” ƒƒŽ›•‹• –‘ ‰”‘—’ ’Ž‘–• ƒ ‘”†‹‰–‘„‡‡ ‘’‘•‹–‹‘•‹‹Žƒ”‹–›—•‹‰ Ž‹ƒ‰‡”—Ž‡ƒ†— Ž‹†‡ƒ†‹•–ƒ ‡•ǡ ƒ† ”‡’”‡•‡–‡† –Š‡ ”‡•—Ž–‹‰ ‰”‘—’• ‘ ƒ ƒ’ ‘ˆ –Š‡ •–—†› ƒ”‡ƒǤ Š‡•‡ ƒƒŽ›•‡• ™‡”‡ ‘†— –‡†™‹–Š–Š‡•–ƒ–‹•–‹ ƒŽ’ƒ ƒ‰‡’‡‹ȋ‡˜‡Ž‘’‡–‘”‡‡ƒʹͲͳͲȌƒ†–Š‡ •‘ˆ–™ƒ”‡˜ǤͶǤͲȋƒ‰‡Ž‡–ƒŽǤʹͲͳͲȌǤ

Š‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡„‡‡•’‡ ‹‡•”‹ Š‡••ƒ†ˆŽ‘™‡””‹ Š‡••ǡ„‡–™‡‡„‡‡ƒ„—†ƒ ‡ ƒ†‘˜‡”ƒŽŽˆŽ‘™‡”ƒ„—†ƒ ‡ǡƒ†„‡–™‡‡„‡‡ƒ„—†ƒ ‡ƒ†„‡‡”‹ Š‡••™ƒ•ƒƒŽ›œ‡† ™‹–Š•‹’Ž‡Ž‹‡ƒ””‡‰”‡••‹‘ǤŠ‡ ‘–”‹„—–‹‘‘ˆˆŽ‘™‡”ȋˆŽ‘™‡”†‡•‹–›‘ˆR.officinalisǡT. vulgarisǡD.pentaphyllumƒ†CistusȌƒ†‡•–‹‰”‡•‘—” ‡ȋ’”‡•‡ ‡‘ˆ†‡ƒ†™‘‘†ǡΨ„ƒ”‡ •‘‹Žǡ—„‡”‘ˆŠ‘Ž‡•‹”‘ •ƒ†—„‡”‘ˆ˜ƒ ƒ–•ƒ‹Ž•Š‡ŽŽ•Ȍ˜ƒ”‹ƒ„Ž‡•–‘„‡‡•’‡ ‹‡• ”‹ Š‡•• ƒ† „‡‡ ƒ„—†ƒ ‡ ™ƒ• ƒƒŽ›œ‡† ™‹–Š ‰‡‡”ƒŽ Ž‹‡ƒ” ‘†‡Ž•Ǥ Quercus coccifera ‘˜‡” ƒ† A. mauritanica ‘˜‡” ™‡”‡ ‘– ‹ Ž—†‡† ‹ –Š‡•‡ ƒƒŽ›•‡• „‡ ƒ—•‡ ‘—” ‡†—†ƒ ›ƒŽ›•‹•ȋ•‡‡„‡Ž‘™Ȍ ‘—Ž†‘–ˆ‹†ƒ›•’‡ ‹‡•ƒ••‘ ‹ƒ–‡†–‘–Š‡•‡•—„•–”ƒ–‡•Ǥ ‡ •‡Ž‡ –‡† –Š‡ ‘•– ’ƒ”•‹‘‹‘—• ‘†‡Ž „ƒ•‡† ‘ ƒ‹‡ǯ• ˆ‘”ƒ–‹‘ ”‹–‡”‹‘ ȋ Ȍ —•‹‰ –Š‡ •–‡’   ˆ— –‹‘ ™‹–Š ˆ‘”™ƒ”† ƒ† „ƒ ™ƒ”† ‡Ž‹‹ƒ–‹‘ ‹’Ž‡‡–‡† ‹ –Š‡  Ž‹„”ƒ”› ȋ‡ƒ„Ž‡• ƒ† ‹’Ž‡› ͳͻͻͻȌ ‘ˆ –Š‡  •‘ˆ–™ƒ”‡ ȋ ‡˜‡Ž‘’‡– ‘”‡ ‡ƒ ʹͲͳͲȌǤ‹ ‡‡‹–Š‡”„‡‡•’‡ ‹‡•”‹ Š‡••‘”ƒ„—†ƒ ‡™‡”‡ƒ—–‘ ‘””‡Žƒ–‡†ȋ•‡‡”‡•—Ž–•Ȍǡ ™‡†‹†‘–‹ Ž—†‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ„Ž‡•‹–Š‡•‡ƒƒŽ›•‡•Ǥ

 31 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

‘‡•–ƒ„Ž‹•Š–Š‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡–Š‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆ„‡‡ ‘’‘•‹–‹‘ƒ†ˆŽ‘™‡” ƒ† ‡•–‹‰ ”‡•‘—” ‡• ™‡ ‘†— –‡† ƒ ‘”†‹ƒ–‹‘ ƒƒŽ›•‹•Ǥ ‡ ˆ‹”•–”—ƒ†‡–”‡†‡† ‘””‡•’‘†‡ ‡ƒƒŽ›•‹•ȋȌ–‘†‡–‡”‹‡™Š‡–Š‡”‘—”†ƒ–ƒŠƒ†ƒ—‹‘†ƒŽ‘”ƒŽ‹‡ƒ” ”‡•’‘•‡ȋ‡’惐†e‹Žƒ—‡”ʹͲͲ͵ȌǤŠ‡”‡•—Ž–•‘ˆ–Š‹•ƒƒŽ›•‹••Š‘™‡†–Šƒ–‘—”†ƒ–ƒ™‡”‡ •—ˆˆ‹ ‹‡–Ž›Š‘‘‰‡‡‘—•ƒ† ‘ˆ‘”‡†–‘ƒ‘†‡Ž™‹–ŠƒŽ‹‡ƒ””‡•’‘•‡Ǥ‡–Š—•ƒ’’Ž‹‡† ƒ‡†—†ƒ ›ƒŽ›•‹•ȋȌǤ‡—•‡†–Š‡•‘ˆ–™ƒ”‡ƒ‘ ‘˜ǤͶǤͷ–‘†‘–Š‡•‡ƒƒŽ›•‡•ȋ‡” ”ƒƒƒ†e‹Žƒ—‡”ʹͲͲʹȌǤ‡ ƒ—•‡„‘†›™‡‹‰Š– Ž‡ƒ”Ž› ‘†‹–‹‘‡†„‡‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘ǡ ™‡”—–™‘•ǡ‘‡‹ Ž—†‹‰‘Ž›•ƒŽŽ•’‡ ‹‡•ȋˆ”‡•Š„‘†›™‡‹‰Š– δ ͷͷ ‰Ȍ ƒ† –Š‡ ‘–Š‡”‹ Ž—†‹‰‘Ž›Žƒ”‰‡•’‡ ‹‡•ȋε͹Ͳ‰ȌǤ „‘–ŠƒƒŽ›•‹•ǡ•’‡ ‹‡•ƒ„—†ƒ ‡†ƒ–ƒ™‡”‡ •“—ƒ”‡Ǧ”‘‘––”ƒ•ˆ‘”‡†ƒ† ‡–”‡†Ǥ‡ ƒ—•‡™‡†‹†‘–™ƒ––‘ƒ––ƒ Š–‘‘— Š™‡‹‰Š––‘ ”ƒ”‡•’‡ ‹‡•ȋ–Š‡ƒŒ‘”‹–›Ȍ™‡†‹†‘–•–ƒ†ƒ”†‹œ‡ƒ„—†ƒ ‡†ƒ–ƒǤ ˜‹‡™‘ˆ–Š‡”‡•—Ž–• ‘„–ƒ‹‡†‹–Š‡ Ž—•–‡”ƒƒŽ›•‹•ǡ‰‡‘‰”ƒ’Š‹ ƒŽ ‘‘”†‹ƒ–‡•™‡”‡‹–”‘†— ‡†ƒ• ‘˜ƒ”‹ƒ„Ž‡•Ǥ ‡•‘—” ‡˜ƒ”‹ƒ„Ž‡•™‡”‡ƒ—–‘ƒ–‹ ƒŽŽ›•‡Ž‡ –‡†™‹–Š–Š‡forward‘’–‹‘ǡƒ†•‹‰‹ˆ‹ ƒ ‡‘ˆ ‡ƒ Š ˜ƒ”‹ƒ„Ž‡ ƒ† •‹‰‹ˆ‹ ƒ ‡ ‘ˆ –Š‡ ‘˜‡”ƒŽŽ ‘†‡Ž ™‡”‡ –‡•–‡† ™‹–Š ‘–‡ ƒ”Ž‘ •‹—Žƒ–‹‘•—†‡””‡†— ‡†‘†‡ŽȋͶͻͻ’‡”—–ƒ–‹‘•ȌǤ



1.3. Results

1.3.1. Beecommunity

‡ ƒ’–—”‡† ͸ͷͺͲ •’‡ ‹‡• ‘””‡•’‘†‹‰ –‘ ͻͺ •’‡ ‹‡• ‹ ˆ‹˜‡ ˆƒ‹Ž‹‡•ǣ ’‹†ƒ‡ ȋʹ͹ •’‡ ‹‡•Ȍǡ‡‰ƒ Š‹Ž‹†ƒ‡ȋʹ͸Ȍǡ†”‡‹†ƒ‡ȋʹ͵Ȍǡ ƒŽ‹ –‹†ƒ‡ȋͳͺȌƒ†‘ŽŽ‡–‹†ƒ‡ȋͶȌȋƒ„Ž‡ͳȌǤ ‹‡–‡‡•’‡ ‹‡•”‡’”‡•‡–‡†ͻ͵ǤʹΨ‘ˆ–Š‡•’‡ ‹‡• ƒ’–—”‡†ǡƒ†͵Ͳ‘ˆ–Š‡”‡ƒ‹‹‰͹ͻ •’‡ ‹‡•™‡”‡•‹‰Ž‡–‘•ǤLasioglossumsubhirtum™ƒ•–Š‡‘•–ƒ„—†ƒ–•’‡ ‹‡•ȋʹ͹ǤͳΨ‘ˆ –‘–ƒŽ •’‡ ‹‡•Ȍǡ ˆ‘ŽŽ‘™‡† „› Andrena djelfensis ȋͳͶǤͳΨȌǤ Ž‘– •’‡ ‹‡• ”‹ Š‡•• ”ƒ‰‡† „‡–™‡‡ ʹͶ ƒ† ͶͶǡ ƒ† ƒ„—†ƒ ‡ „‡–™‡‡ ʹͲ͹ ƒ† ͷͷͻǤ Š‡ ”‡Žƒ–‹‘•Š‹’ „‡–™‡‡ „‡‡ •’‡ ‹‡•”‹ Š‡••ƒ†ƒ„—†ƒ ‡ˆƒ‹Ž‡†•‹‰‹ˆ‹ ƒ ‡ȋ”ʹαͲǤͳͷǢ’αͲǤͲͻȌǤ



1.3.2. Spatialdistributionofflowerandnestingresources

‘–Š ˆŽ‘™‡” †‡•‹–› ȋʹ͹ –‘ ʹ͸ͷ ˆŽ‘™‡”•ȀʹȌ ƒ† •’‡ ‹‡• ”‹ Š‡•• ȋͷ –‘ ʹ͹Ȍ ˜ƒ”‹‡† ™‹†‡Ž› ƒ ”‘••’Ž‘–•ȋƒ„Ž‡ʹƒ†͵ȌǤ Ž‘™‡”ƒ„—†ƒ ‡ƒ†”‹ Š‡••™‡”‡‘–”‡Žƒ–‡†ȋ”ʹαͲǤͲ͹Ǣ  32 Šƒ’–‡”ͳ

’εͲǤʹͷȌǤ Ž‘™‡” ƒ„—†ƒ ‡ †‹† ‘– •Š‘™ •’ƒ–‹ƒŽ ƒ—–‘ ‘””‡Žƒ–‹‘ ȋ α ǦͲǤͲʹͶǡ ’α ͲǤͷͳȌǤ •–‡ƒ†ǡˆŽ‘™‡”•’‡ ‹‡•”‹ Š‡••™ƒ••‹‰‹ˆ‹ ƒ–Ž›ƒ—–‘ ‘””‡Žƒ–‡†ȋ αͲǤͳͺ͸ǡ’δͲǤͲͲͲͳȌǡ™‹–Š ƒ ‰”ƒ†‹‡– ‘ˆ ’‘•‹–‹˜‡ ƒ—–‘ ‘””‡Žƒ–‹‘ ƒ– •Š‘”– †‹•–ƒ ‡• ȋδ ͳͲͲͲȌ’”‘‰”‡••‹˜‡Ž›Ž‘•‹‰ •‹‰‹ˆ‹ ƒ ‡ƒ–Ž‘‰‡”†‹•–ƒ ‡•ǤŠ‡‘Ž›ˆŽ‘™‡”•’‡ ‹‡•™‹–Šƒ•‹‰‹ˆ‹ ƒ–‘”ƒǯ• ™ƒ•T. vulgarisȋ αͲǤͲͶͻǡ’αͲǤͲͳͷȌȋ ‹‰ǤͳǤͳȌǤŠ‡ƒ••‘ ‹ƒ–‡† ‘””‡Ž‘‰”ƒƒ‰ƒ‹•Š‘™‡†ƒ‰”ƒ†‹‡– ‘ˆ’‘•‹–‹˜‡ƒ—–‘ ‘””‡Žƒ–‹‘ƒ–•Š‘”–†‹•–ƒ ‡ Žƒ••‡•™‹–Šƒ’”‘‰”‡••‹˜‡Ž‘••‘ˆ•‹‰‹ˆ‹ ƒ ‡Ǥ Rosmarinusofficinalis™ƒ•‘”‡‘”Ž‡••‡˜‡Ž›†‹•–”‹„—–‡†–Š”‘—‰Š‘—––Š‡’ƒ”ǡ™Š‡”‡ƒ•D. pentaphyllum™ƒ•‘•–ƒ„—†ƒ–‹–Š‡‘”–ŠǦ™‡•–‡”‡†‰‡ǤCistus•’’ǤˆŽ‘™‡”†‡•‹–›™ƒ• Ž‘™ ‘’ƒ”‡†–‘–Š‡‘–Š‡”•’‡ ‹‡•ǡƒ†˜ƒ”‹‡†ˆ”‘’Ž‘––‘’Ž‘–•Š‘™‹‰‘ Ž‡ƒ”’ƒ––‡” ȋ ‹‰ǤͳǤͳǢƒ„Ž‡ʹȌǤ

‡•–‹‰ •—„•–”ƒ–‡ ‘’‘•‹–‹‘ ƒŽ•‘ ˜ƒ”‹‡† ™‹†‡Ž› ƒ ”‘•• ’Ž‘–• ȋ ‹‰ǤͳǤʹȌǤƒ”‡•‘‹Žƒ†Q. coccifera ‘˜‡”™‡”‡–Š‡‘Ž›–™‘‡•–‹‰•—„•–”ƒ–‡•’”‡•‡–‹ƒŽŽ’Ž‘–•Ǥ ‘™‡˜‡”ǡƒŽŽ’Ž‘–• ‡š ‡’–‘‡‘ˆˆ‡”‡†ƒ–Ž‡ƒ•–Ͷ‘ˆ–Š‡͸‡•–‹‰”‡•‘—” ‡•ǤŠ‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆ‡•–‹‰ •—„•–”ƒ–‡• ™ƒ• Š‹‰ŠŽ› Š‡–‡”‘‰‡‡‘—• ȋ ‹‰Ǥ ͳǤʹǡ ƒ„Ž‡ ʹȌǤ ‘‡ ‘ˆ–Š‡‡•–‹‰•—„•–”ƒ–‡• •Š‘™‡†ƒ†‹• ‡”ƒ„Ž‡•’ƒ–‹ƒŽ’ƒ––‡”ǡ‡š ‡’–ˆ‘”Š‘Ž‡•‹”‘ • ȋ α ͲǤͲͶͷǡ ’α ͲǤͲʹȌǡ ƒ‰ƒ‹ •Š‘™‹‰†‡ ”‡ƒ•‹‰’‘•‹–‹˜‡ƒ—–‘ ‘””‡Žƒ–‹‘™‹–Š‹ ”‡ƒ•‹‰†‹•–ƒ ‡Ǥ



1.3.3. Beespatialdistribution

‡‹–Š‡”„‡‡ƒ„—†ƒ ‡ȋ αǦͲǤͲͷǡ’αͲǤͻͻȌ‘”•’‡ ‹‡•”‹ Š‡••ȋ αͲǤͲͲʹǡ’αͲǤͳ͹Ȍ•Š‘™‡† •’ƒ–‹ƒŽ ƒ—–‘ ‘””‡Žƒ–‹‘Ǥ •–‡ƒ†ǡ „‡‡ ‘’‘•‹–‹‘ †‹† •Š‘™ •‹‰‹ˆ‹ ƒ– ƒ—–‘ ‘””‡Žƒ–‹‘ ȋƒ–‡Ž”αͲǤʹ͹Ǣ’αͲǤͲͲ͵ȌǤŠ‡™‡ƒƒŽ›œ‡†–Š‡ͳͻ‘•–ƒ„—†ƒ–•’‡ ‹‡••‡’ƒ”ƒ–‡Ž›ǡ™‡ ˆ‘—†•’ƒ–‹ƒŽƒ—–‘ ‘””‡Žƒ–‹‘ˆ‘”ͻ‘ˆ–Š‡ȋƒ„Ž‡ͳǤͳȌǤ‹‰‹ˆ‹ ƒ–ƒ—–‘ ‘””‡Žƒ–‹‘‘ —””‡† ‘•–Ž›ƒ–†‹•–ƒ ‡•δͻͷͲǤ ’‘”–ƒ–Ž›ǡ•’‡ ‹‡••Š‘™‹‰•‹‰‹ˆ‹ ƒ–ƒ—–‘ ‘””‡Žƒ–‹‘Šƒ† Ž‘™‡” „‘†› ™‡‹‰Š– ȋ‡ƒ ά ǣ ʹͲ ά ͳ͵ ‰Ǣ α ͻȌ –Šƒ –Š‘•‡ ™‹–Š ‘ •‹‰‹ˆ‹ ƒ– ƒ—–‘ ‘””‡Žƒ–‹‘ȋͳͲͲά͹ʹ‰ǢαͳͲȌȋƒ„Ž‡ͳǤͳǢƒǦŠ‹–‡›ǣαǦʹǤͺͷͺǢ’αͲǤͲͲͶȌǤŠ‡ Ž—•–‡”ƒƒŽ›•‹•‘ˆ–Š‡’Ž‘–•„ƒ•‡†‘„‡‡ ‘’‘•‹–‹‘•‹‹Žƒ”‹–›”‡•—Ž–‡†‹ˆ‹˜‡‰”‘—’•ƒ† ”‡˜‡ƒŽ‡†ƒ Ž‡ƒ”‰‡‘‰”ƒ’Š‹ ƒŽ’ƒ––‡”ȋ ‹‰ǤͳǤ͵ȌǤ –‡”‡•–‹‰Ž›ǡ –Š‡ –™‘ ‘•– ƒ„—†ƒ– •’‡ ‹‡•ǡ Lasioglossum subhirtumƒ†Andrena djelfensisǡ •Š‘™‡† ’ƒ”–‹ƒŽŽ› •‡‰”‡‰ƒ–‡† †‹•–”‹„—–‹‘•ǤLasioglossumsubhirtum™ƒ•†‘‹ƒ–‹–Š‡ ‡–”ƒŽƒ†™‡•–‡”ƒ”‡ƒ•‘ˆ–Š‡ ’ƒ”ǡ™Š‡”‡ƒ•A.djelfensis™ƒ•†‘‹ƒ–‘–Š‡‡ƒ•–‡”•‹†‡Ǥ„—†ƒ ‡‘ˆ–Š‡•‡–™‘

 33 























Figure1.1 ƒ’‘ˆ–Š‡ ƒ””ƒˆƒ”•Š‘™‹‰–Š‡†‡•‹–›‘ˆˆŽ‘™‡””‡•‘—” ‡•ȋ—„‡”‘ˆˆŽ‘™‡”•ȀʹȌ‹‡ƒ Š’Ž‘–ȋαʹͳȌǤ

 



























 Figure1.2 ƒ’‘ˆ–Š‡ ƒ””ƒˆƒ”•Š‘™‹‰–Š‡ƒ„—†ƒ ‡‘ˆ‡•–‹‰”‡•‘—” ‡•‹‡ƒ Š’Ž‘–ȋαʹͳȌǤ ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

•’‡ ‹‡••Š‘™‡†ƒ•‹‰‹ˆ‹ ƒ–‡‰ƒ–‹˜‡ ‘””‡Žƒ–‹‘ȋ”•αǦͲǤ͸ʹǢ’αͲǤͲͲ͵ȌǤ –Š‡”•’‡ ‹‡•ƒŽ•‘ •Š‘™‡† ƒ ‰‡‘‰”ƒ’Š‹ ƒŽ ’ƒ––‡”Ǥ Lasioglossum malachurum ™ƒ• ‘•– ƒ„—†ƒ– ‹ –Š‡  •‹†‡ǡLasioglossumbimaculatum‹–Š‡ƒ†ǡƒ†Lasioglossumalbocinctum‹–Š‡Ǥ Panurgusdentipes™ƒ•ƒ„—†ƒ–‘Ž›‹’Ž‘–ͳǡ™‹–Šƒ„‡‡ ‘’‘•‹–‹‘ƒ”‡†Ž›†‹ˆˆ‡”‡– ˆ”‘–Šƒ–‘ˆƒŽŽ‘–Š‡”’Ž‘–•Ǥ–Š‡‘–Š‡”Šƒ†ǡ•’‡ ‹‡••— Šƒ•RhodanthidiumsticticumǡApis melliferaǡ Andrena nigroaeneaƒ†Bombus terrestris•Š‘™‡†ƒ— Š‘”‡Š‘‘‰‡‡‘—• †‹•–”‹„—–‹‘–Š”‘—‰Š‘—––Š‡’ƒ”Ǥ‡ ƒŽ —Žƒ–‡†–Š‡ ‘‡ˆˆ‹ ‹‡–‘ˆ˜ƒ”‹ƒ–‹‘ȋαʹͳ’Ž‘–•Ȍ‘ˆ –Š‡ƒ„—†ƒ ‡‘ˆ–Š‡ͳͻƒ‹•’‡ ‹‡•ƒ•ƒ‡ƒ•—”‡‘ˆ–Š‡‹”†‡‰”‡‡‘ˆ•’ƒ–‹ƒŽŠ‡–‡”‘‰‡‡‹–›Ǥ ’‡ ‹‡• ™‹–Š Š‹‰Š‡” ‘‡ˆˆ‹ ‹‡–• ‘ˆ ˜ƒ”‹ƒ–‹‘ ȋεͲǤͻͷȌ Šƒ† Ž‘™‡”„‘†›™‡‹‰Š–ȋ‡ƒάǣ ʹ͹ǤͶάʹͷǤͺ‰ǢαͳͳȌ–Šƒ–Š‘•‡™‹–ŠŽ‘™‡”ȋδͲǤͻͲȌ ‘‡ˆˆ‹ ‹‡–•‘ˆ˜ƒ”‹ƒ–‹‘ȋͳͲǤͻά͹͸ǤͶ‰Ǣ αͺȌȋƒ„Ž‡ͳǤͳǢƒǦŠ‹–‡›ǣαʹǤ͹͸͸Ǣ’αͲǤͲͲ͸Ȍǡ ‘””‘„‘”ƒ–‹‰–Š‡ ‘ Ž—•‹‘–Šƒ– –Š‡‘„•‡”˜‡†•’ƒ–‹ƒŽ’ƒ––‡”™ƒ•‘•–Ž›†—‡–‘•ƒŽŽ•’‡ ‹‡•Ǥ



1.3.4. Relationshipbetweenresourcesandbeespatialdistribution

‡‡•’‡ ‹‡•”‹ Š‡••™ƒ•‘–”‡Žƒ–‡†–‘ˆŽ‘™‡”•’‡ ‹‡•”‹ Š‡••ȋ”ʹαͲǤͲͷǢ’αͲǤ͵ʹȌǤ ‘™‡˜‡”ǡ –Š‹•Žƒ ‘ˆ”‡Žƒ–‹‘•Š‹’™ƒ• ƒ—•‡†„›’Ž‘–ͳȋ™‹–Š–Š‡Š‹‰Š‡•–„‡‡”‹ Š‡••ƒ†ƒ”ƒ–Š‡” —‹“—‡„‡‡ ‘’‘•‹–‹‘Ȍ•–”‘‰Ž›†‡˜‹ƒ–‹‰ˆ”‘–Š‡‰‡‡”ƒŽ–”‡†•Š‘™„›–Š‡”‡•–‘ˆ–Š‡ ’Ž‘–•Ǥ š Ž—•‹‘ ‘ˆ –Š‹• ’Ž‘– ™‘—Ž† ƒ—•‡ –Š‡ ˆŽ‘™‡”Ǧ„‡‡ ”‹ Š‡•• ”‡Žƒ–‹‘•Š‹’ –‘ „‡ ‘‡ •‹‰‹ˆ‹ ƒ–ȋ”ʹαͲǤʹͷǢ’αͲǤͲʹȌǤŠ‡•‡Ž‡ –‡† ‡š’Žƒ‹‹‰„‡‡”‹ Š‡••‹ Ž—†‡†‘‡•–‹‰ •—„•–”ƒ–‡˜ƒ”‹ƒ„Ž‡•ǡƒ†‘Ž›‘‡ˆŽ‘™‡”˜ƒ”‹ƒ„Ž‡ȋCistusˆŽ‘™‡”ƒ„—†ƒ ‡Ȍǡ„—–™‹–Šƒ‘Ǧ •‹‰‹ˆ‹ ƒ–’Ǧ˜ƒŽ—‡ȋ”ʹαͲǤͲ͹ǡ’αͲǤͳʹʹǢƒ„Ž‡ͶȌǤ‡‡ƒ„—†ƒ ‡™ƒ•‘–”‡Žƒ–‡†–‘‘˜‡”ƒŽŽ ˆŽ‘™‡” ƒ„—†ƒ ‡ ȋ”ʹαͲǤͲͷǢ’εͲǤ͵ȌǤŠ‡„‡•–‘†‡Ž‡š’Žƒ‹‹‰„‡‡ƒ„—†ƒ ‡‹ Ž—†‡† ƒ„—†ƒ ‡ ‘ˆ Cistusƒ†T. vulgarisˆŽ‘™‡”•ȋ”ʹαͲǤ͵ʹȌǤ ‘™‡˜‡”ǡ ‘Ž› ƒ„—†ƒ ‡ ‘ˆ Cistus ˆŽ‘™‡”•™ƒ••‹‰‹ˆ‹ ƒ–ȋ’αͲǤͲͳ͵Ǣƒ„—†ƒ ‡‘ˆT.vulgarisˆŽ‘™‡”•ǡ’αͲǤͳ͸ͻȌǤ•™‹–Š„‡‡ ”‹ Š‡••ǡ„‡‡ƒ„—†ƒ ‡™ƒ•‘–”‡Žƒ–‡†–‘‡•–‹‰•—„•–”ƒ–‡ƒ˜ƒ‹Žƒ„‹Ž‹–›ȋƒ„Ž‡ͶȌǤ

Š‡‘ˆ•ƒŽŽ•’‡ ‹‡•ȋδͷͷ‰Ȍ‹†‹ ƒ–‡•–Šƒ––Š‡•’ƒ–‹ƒŽ†‹•–”‹„—–‹‘‘ˆ„‡‡ ‘’‘•‹–‹‘ ‹• Ž‡ƒ”Ž›ƒ••‘ ‹ƒ–‡†–‘ˆŽ‘™‡””‡•‘—” ‡•ƒ†‘Ž›™‡ƒŽ›–‘‡•–‹‰”‡•‘—” ‡•ȋ ‹‰ǤͳǤͶȌǤ™‘ ˆŽ‘™‡” ˜ƒ”‹ƒ„Ž‡• ™‡”‡ •‹‰‹ˆ‹ ƒ– ‹ –Š‡ ‘†‡Žǣ T. vulgaris ȋ‘–”‹„—–‹‘ –‘ –Š‡ ‘†‡Žα ͳͳǤ͹ΨǢ ’αͲǤͲͳȌ ƒ† Cistus •’’Ǥ ȋ‘–”‹„—–‹‘ –‘ –Š‡ ‘†‡Žα ͻǤͺΨǢ ’αͲǤͲͲ͸Ȍ.Š‡‘†‡Ž ‹ Ž—†‹‰ ƒŽŽ ˜ƒ”‹ƒ„Ž‡• ™ƒ• •‹‰‹ˆ‹ ƒ– ȋ’αͲǤͲʹȌ ƒ† ‡š’Žƒ‹‡† ͶͷǤʹΨ ‘ˆ –Š‡ ‘„•‡”˜‡† ˜ƒ”‹ƒ ‡ȋƒ„Ž‡ͳǤʹȌǤŠ‡ˆ‹”•–ƒš‹•‡š’Žƒ‹‡†ʹͷǤͶΨ‘ˆ–Š‡˜ƒ”‹ƒ ‡ƒ†™ƒ•†‡ˆ‹‡†„›  36 























 Figure1.3 ƒ’‘ˆ–Š‡ ƒ””ƒˆƒ”•Š‘™‹‰–Š‡ƒ„—†ƒ ‡‘ˆ–Š‡ͳͻ‘•–ƒ„—†ƒ–„‡‡•’‡ ‹‡•ȋ”‡’”‡•‡–‹‰‘”‡–ŠƒͲǤͷΨ‘ˆ–Š‡•’‡ ‹‡••ƒ’Ž‡†Ȍ ‹‡ƒ Š’Ž‘–ȋαʹͳȌǤŽ‘–•‰”‘—’‡†„ƒ•‡†‘„‡‡ ‘’‘•‹–‹‘ƒ ‘”†‹‰–‘ Ž—•–‡”ƒƒŽ›•‹•Ǥ 

1 Table1.1ƒ”ƒ‡–‡”•‘ˆ–Š‡ͳͻ‘•–ƒ„—†ƒ–„‡‡•’‡ ‹‡•‹–Š‡ ƒ””ƒˆ ‘—‹–›Ǥ

CVof Bodyweight Nesting Pollen  Abundance Moran'sI P abundance (mg) substrate specialization Sociality Lasioglossumgriseolum ͹ͳ ͲǤͲʹͶ 0.04 ͳǤͳʹ ͵Ǥͺ ‘‹Ž ‘Ž›Ž‡ –‹ ǫ Lasioglossumtransitorium ͳͶ͹ ǦͲǤͲͷ ͲǤͻ ͲǤͻͻ ͹Ǥ͸ ‘‹Ž ‘Ž›Ž‡ –‹ ǫ Lasioglossummediterraneum Ͷ͸ ͲǤͲ͵Ͷ 0.03 ͳǤʹͶ ͳʹǤͶ ‘‹Ž ‘Ž›Ž‡ –‹ ǫ ǫ Lasioglossummalachurum ʹʹʹ ͲǤͲͺ 0.001 ͳǤʹʹ ͳ͵Ǥ͵ ‘‹Ž ‘Ž›Ž‡ –‹ ‘ ‹ƒŽ Andrenadjelfensis ͻʹ͸ ͲǤͳ͵ͳ 0 ͳǤͲ͵ ͳͷǤͲ ‘‹Ž ‘Ž›Ž‡ –‹ ǫ ‘Ž‹–ƒ”› Lasioglossumsubhirtum ͳ͹ͺͲ ͲǤͲͶͷ 0.01 ͲǤ͹͵ ͳ͸ǤͶ ‘‹Ž ‘Ž›Ž‡ –‹ ǫ Panurgusdentipes ͳͳ͹ ǦͲǤͲʹ͸ 0.03 ʹǤͷͻ ͳͺǤͻ ‘‹Ž Ž‹‰‘Ž‡ –‹ ‘Ž‹–ƒ”› Lasioglossumbimaculatum ʹͲʹ ͲǤͲͺ͵ 0 ͳǤʹͳ ʹͲǤͲ ‘‹Ž ‘Ž›Ž‡ –‹ ǫ ‘Ž‹–ƒ”› Osmiarufohirta ͳʹʹ ͲǤͳͲͻ 0 ͳǤͳ͸ ʹ͹ǤͶ ƒ‹Ž•Š‡ŽŽ• ‘Ž›Ž‡ –‹ ‘Ž‹–ƒ”› Andrenasenecionis ͺ͸ ǦͲǤͲ͵ͺ ͲǤ͹ ͲǤͺͻ ͶͲǤͲ ‘‹Ž Ž‹‰‘Ž‡ –‹ ǫ ‘Ž‹–ƒ”› Halictussimplex ͹ͳ ǦͲǤͲ͹͹ ͲǤͷ ͳǤͳ͵ ͶͲǤͲ ‘‹Ž ‘Ž›Ž‡ –‹ ‘Ž‹–ƒ”›ǫ Lasioglossumalbocinctum ʹͲͺ ͲǤͲ͵Ͷ 0.01 ͳǤʹ ͶͻǤͶ ‘‹Ž ‘Ž›Ž‡ –‹ ‘Ž‹–ƒ”› Rhodanthidiumseptemdentatum ͳʹͳ ǦͲǤͲͲ͸ ͲǤ͵ ͲǤͷ ͺͷǤͻ ƒ‹Ž•Š‡ŽŽ• ‘Ž›Ž‡ –‹ ‘Ž‹–ƒ”› Andrenanigroaenea ʹʹͺ ǦͲǤͲʹͶ ͲǤͷ ͲǤͶͳ ͺ͸Ǥͷ ‘‹Ž ‘Ž›Ž‡ –‹ ‘Ž‹–ƒ”› Halictusscabiosae ͵ͺ ǦͲǤͲʹ ͲǤ ͵ ͳǤ͸Ͷ ͻ͵Ǥ͸ ‘‹Ž ‘Ž›Ž‡ –‹ ‘ ‹ƒŽ Apismellifera ͷʹͺ ͲǤͲͳͳ ͲǤͳ ͲǤ͵͹ ͻ͹ǤͶ ƒ”‰‡ ƒ˜‹–‹‡• ‘Ž›Ž‡ –‹ ‘ ‹ƒŽ Rhodanthidiumsticticum ͺͳ͹ ǦͲǤͲͳͳ ͲǤʹ ͲǤͺ͹ ͳͲ͵Ǥʹ ƒ‹Ž•Š‡ŽŽ• ‘Ž›Ž‡ –‹ ‘Ž‹–ƒ”› Anthophoradispar Ͷ͸ ǦͲǤͲʹͶ ͲǤ ͷ ͲǤͺ ͳͺͺǤͶ ‘‹Ž ‘Ž›Ž‡ –‹ ǫ ‘Ž‹–ƒ”› Bombusterrestris ͵ͷͶ ǦͲǤͲͳͷ ͲǤͶ ͲǤͶ͹ ʹͷͲǤͷ ƒ”‰‡ ƒ˜‹–‹‡• ‘Ž›Ž‡ –‹ ‘ ‹ƒŽ 2 „—†ƒ ‡ȋ—„‡”‘ˆ•’‡ ‹‡• ƒ’–—”‡†Ȍǡ‘”ƒǯ• ȋ•‹‰‹ˆ‹ ƒ–pǦ˜ƒŽ—‡•‹„‘Ž†Ȍǡ ‘‡ˆˆ‹ ‹‡–‘ˆ˜ƒ”‹ƒ–‹‘‘ˆƒ„—†ƒ ‡ȋαʹͳ’Ž‘–•Ȍǡˆ”‡•Šˆ‡ƒŽ‡„‘†›™‡‹‰Š–ǡ‘ˆ–Š‡ͳͻ‘•–ƒ„—†ƒ–•’‡ ‹‡•‹–Š‡ 3 ƒ””ƒˆ„‡‡ ‘—‹–›Ǥ’‡ ‹‡•‘”†‡”‡†„›‹ ”‡ƒ•‹‰™‡‹‰Š–Ǥ

 Šƒ’–‡”ͳ

T.vulgarisˆŽ‘™‡”•ƒ†—„‡”‘ˆŠ‘Ž‡•‹”‘ •‘–Š‡‘‡Šƒ†ǡƒ†„›Cistus•’’ǤˆŽ‘™‡”• ‘–Š‡‘–Š‡”Šƒ†ȋ ‹‰ǤͳǤͶȌǤŠ‡•‡ ‘†ƒš‹•‡š’Žƒ‹‡†‘Ž›ͷǤ͵Ψ‘ˆ–Š‡˜ƒ”‹ƒ ‡.–Š‡ ‘–Š‡”Šƒ†ǡ–Š‡‘†‡Ž‘ˆŽƒ”‰‡•’‡ ‹‡•™ƒ•‘Ǧ•‹‰‹ˆ‹ ƒ–Ǥ Š‡ ‘˜‡”ƒŽŽ ˜ƒ”‹ƒ ‡ ‡š’Žƒ‹‡†™ƒ•Ž‘™‡”ȋ͵ͺǤͻΨǢƒ„Ž‡ͳǤʹȌǡƒ†‘˜ƒ”‹ƒ„Ž‡•‡–‡”‡†–Š‡‘†‡ŽǤ



1.4. Discussion

Š‡ ƒ””ƒˆ„‡‡ ‘—‹–›•Š‘™•ƒ Ž‡ƒ”•’ƒ–‹ƒŽ’ƒ––‡”ƒ––Š‡Šƒ„‹–ƒ–• ƒŽ‡ǡ™‹–Š†‹ˆˆ‡”‡– •’‡ ‹‡•†‘‹ƒ–‹‰‹†‹ˆˆ‡”‡–’Ž‘–••‡’ƒ”ƒ–‡†„›ƒ•ˆ‡™ƒ•ͷͲͲǦͳͲͲͲǤŠ‹•’ƒ––‡”‹•†—‡ –‘•ƒŽŽǦ•‹œ‡†•’‡ ‹‡•ȋδͷͷ‰Ȍǡ™‹–ŠŽƒ”‰‡”•’‡ ‹‡••Š‘™‹‰ƒ‘”‡‘”Ž‡••Š‘‘‰‡‡‘—• †‹•–”‹„—–‹‘Ǥ  Ž‹‡Ž› ‡š’Žƒƒ–‹‘ ˆ‘” –Š‹• ‘—– ‘‡ ‹• –Šƒ– ‘—” ‹–‡”Ǧ’Ž‘– †‹•–ƒ ‡ ™ƒ• •—ˆˆ‹ ‹‡––‘ƒ ‘‘†ƒ–‡–Š‡ˆ‘”ƒ‰‹‰ƒ”‡ƒ•‘ˆ•ƒŽŽ„‡‡•„—–‘––Š‘•‡‘ˆŽƒ”‰‡•’‡ ‹‡•Ǥ ’‘•‹–‹˜‡ ”‡Žƒ–‹‘•Š‹’ „‡–™‡‡ „‘†› •‹œ‡ ƒ† ˆ‘”ƒ‰‹‰ ƒ”‡ƒ• Šƒ• „‡‡ ™‡ŽŽ ‡•–ƒ„Ž‹•Š‡† ȋ ƒ–Šƒƒ†• Šƒ”–‡ʹͲͲʹǡ ”‡‡Ž‡ƒˆ‡–ƒŽǤʹͲͲ͹ǡ —±†‘–‡–ƒŽǤʹͲͲͻǡ—”„— Š‡‡–ƒŽǤ ʹͲͳͲȌǤ Š‡ ‡–Š‘†• —•‡† ‹ –Š‡•‡ ƒ† ‘–Š‡” ”‡Žƒ–‡† •–—†‹‡•ǡ Š‘™‡˜‡”ǡ –‡† –‘ ’”‘˜‹†‡ ‡•–‹ƒ–‡•‘ˆ‡‹–Š‡”‹‹—‘”ƒš‹—ˆ‘”ƒ‰‹‰”ƒ‰‡•Ǥ –—ƒŽˆ‘”ƒ‰‹‰†‹•–ƒ ‡•Šƒ˜‡ „‡‡•Š‘™–‘ Šƒ‰‡‹–‹‡ƒ†•’ƒ ‡„ƒ•‡†‘”‡•‘—” ‡ƒ˜ƒ‹Žƒ„‹Ž‹–›ȋ‹•• Š‡”ƒ†‡‡Ž‡› ͳͻͺʹǡ‡‡ƒƒ†ƒ–‹‡•ʹͲͲͲǡ‹ ‡•ƒ†‘• ŠʹͲͲͲǡŠƒ––ƒ Šƒ”›ƒ ‡– ƒŽǤ ʹͲͲʹǡ ‘—Ž•‘ ʹͲͳͲǡ —”„— Š‡ ‡– ƒŽǤ ʹͲͳͲǡ ƒ”˜‡ŽŽ ‡– ƒŽǤ ʹͲͳʹǡ ‹–Š‡–ƒŽǤʹͲͳʹȌǤ—””‡•—Ž–• ’”‘˜‹†‡‹†‹”‡ –‡˜‹†‡ ‡–Šƒ–ǡ‹ƒ–—”ƒŽŠƒ„‹–ƒ–•™‹–Šƒ„—†ƒ–ˆŽ‘™‡””‡•‘—” ‡•ǡ•’‡ ‹‡• •ƒŽŽ‡”–Šƒͷͷ‰–‡†–‘ˆ‘”ƒ‰‡™‹–Š‹ƒ”ƒ†‹—•‘ˆʹͷͲǦͷͲͲǤ—‡–‘–Š‡‹”Ž‘™ˆ‘‘† ”‡“—‹”‡‡–• ȋòŽŽ‡” ‡– ƒŽǤ ʹͲͲ͸Ȍǡ •ƒŽŽ •’‡ ‹‡• ƒ› „‡ ƒ„Ž‡ –‘ ‘„–ƒ‹ •—ˆˆ‹ ‹‡– ’‘ŽŽ‡Ǧ ‡ –ƒ””‡•‘—” ‡•™‹–Š‹ƒ•ƒŽŽˆ‘”ƒ‰‹‰”ƒ†‹—•Ǥ ’ƒ”ƒŽŽ‡Ž•–—†‹‡•‹‘—”•–—†›ƒ”‡ƒ™‡Šƒ˜‡ ‘„•‡”˜‡†Lasioglossumtransitoriumˆ‡ƒŽ‡•ȋ„‘†›•‹œ‡ǣ͹Ǥ͸‰Ȍ ‘’Ž‡–‹‰‡–‹”‡ˆ‘”ƒ‰‹‰ „‘—–•‘•‹‰Ž‡R.officinalis’Žƒ–•ȋ™Š‹ Šƒ›†‹•’Žƒ›Š—†”‡†•‘ˆ‘’‡ˆŽ‘™‡”•ȌŽ‘ ƒ–‡† ™‹–Š‹ͷͲ ‘ˆ–Š‡‹”‡•–Ǥ

ƒ††‹–‹‘–‘ˆ‘”ƒ‰‹‰ˆŽ‹‰Š–•ǡ‘—”’Ž‘–• ‘—Ž†„‡Ž‹‡†„›†‹•’‡”•ƒŽ ‘˜‡‡–•Ǥ ‘ ‘—” ‘™Ž‡†‰‡ǡ ‹ˆ‘”ƒ–‹‘ ‘ „‡‡ †‹•’‡”•ƒŽ †‹•–ƒ ‡• ‹• ‘•–Ž› Žƒ ‹‰ǡ „—– •‘‡ ‡˜‹†‡ ‡ •—‰‰‡•–•†‹•’‡”•ƒŽ†‹•–ƒ ‡•‘ˆƒ–Ž‡ƒ•–ƒˆ‡™Ǥƒ”‡†Osmiacornutaˆ‡ƒŽ‡•ȋƒ•‘Ž‹–ƒ”› •’‡ ‹‡••Ž‹‰Š–Ž›Žƒ”‰‡”–ŠƒApismelliferaȌŠƒ˜‡„‡‡ˆ‘—†‡•–‹‰ʹƒ™ƒ›ˆ”‘–Š‡‹” ”‡Ž‡ƒ•‡ •‹–‡ ȋ‘• Š ƒ† ‹ ‡• ʹͲͲ͸ȌǤ Š‡”‡ ‹• ƒŽ•‘ ‡˜‹†‡ ‡ –Šƒ– Bombus•’‡ ‹‡•ƒ› †‹•’‡”•‡ƒ•— Šƒ•͵ǦͳͲȋ ‘—Ž•‘ʹͲͳͲȌǤ˜‡ƒ••—‹‰•ƒŽŽ‡”†‹•’‡”•ƒŽ†‹•–ƒ ‡•  39 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡• 

ˆ‘” •ƒŽŽ‡” „‡‡•ǡ ƒ†‰‹˜‡–Š‡ Žƒ  ‘ˆ ’Š›•‹ ƒŽ ƒ† ‡˜‹”‘‡–ƒŽ „ƒ””‹‡”• ‹ –Š‡ ƒ””ƒˆ • ”—„Žƒ†ǡƒ›•’‡ ‹‡••Š‘—Ž†„‡ƒ„Ž‡–‘ ‘˜‡”–Š‡Ž‹‹–•‘ˆ‘—”•–—†›ƒ”‡ƒ‘˜‡”‘‡‘”ƒˆ‡™ ‰‡‡”ƒ–‹‘•ǤŠ‡”‡ˆ‘”‡ǡ–Š‡ˆƒ ––Šƒ–‘—”„‡‡ ‘—‹–›•Š‘™••— Šƒ Ž‡ƒ”•’ƒ–‹ƒŽ’ƒ––‡” •—‰‰‡•–• ƒ •–”‘‰ ‹ˆŽ—‡ ‡ ‘ˆ ‡˜‹”‘‡–ƒŽ ‘†‹–‹‘• ƒ– ƒ ˜‡”› Ž‘ ƒŽ • ƒŽ‡ǡ ƒ– Ž‡ƒ•– ˆ‘” •ƒŽŽ•’‡ ‹‡•Ǥ





 Figure1.4‹’Ž‘–‘ˆ‘†‡Ž”‡Žƒ–‹‰•ƒŽŽ„‡‡•’‡ ‹‡•ȋδͷͷ‰Ȍ–‘ˆŽ‘™‡” ƒ† ‡•–‹‰ ”‡•‘—” ‡•Ǥ ””‘™•”‡’”‡•‡–”‡•‘—” ‡• ȋˆŽ‘™‡”• ‹ Ž‘™‡” ƒ•‡ǡ  ‡•–‹‰•—„•–”ƒ–‡•‹—’’‡” ƒ•‡Ȍǡƒ†—„‡”•„‡‡•’‡ ‹‡•Ǥ ‘”•’‡ ‹‡•ƒ‡• •‡‡ƒ„Ž‡ͳǤ

  ‡•–‹‰”‡•‘—” ‡••Š‘™ƒ‹””‡‰—Žƒ”‘•ƒ‹ †‹•–”‹„—–‹‘ƒ ”‘••–Š‡’ƒ”ǤŠ‡›ƒ”‡‘–‰‘‘† ’”‡†‹ –‘”•‘ˆ„‡‡ƒ„—†ƒ ‡ƒ†”‹ Š‡••ǡƒ†‘Ž›ƒ ‘—–ˆ‘”ƒ•ƒŽŽ’ƒ”–‘ˆ–Š‡‡š’Žƒ‹‡† ˜ƒ”‹ƒ ‡‘ˆ„‡‡ ‘’‘•‹–‹‘Ǥ ‘—” ‘—‹–›ǡ‘•–•’‡ ‹‡•ȋ͸ʹΨǡ‹ Ž—†‹‰ͳ͵‘ˆ–Š‡ͳͻ ‘•–ƒ„—†ƒ–Ȍ‡•–—†‡”‰”‘—†‘”ƒ”‡ Ž‡’–‘’ƒ”ƒ•‹–‹ ‘•’‡ ‹‡•‡•–‹‰——†‡”‰”‘—†Ǥ– –Š‡ •ƒ‡ –‹‡ǡ ’ƒ– Š‡• ‘ˆ „ƒ”‡ •‘‹Ž ƒ”‡ ƒ„—†ƒ– ƒ† ™‹†‡Ž› †‹•–”‹„—–‡† ƒ ”‘•• –Š‡ ’ƒ”ǡ

 40 Šƒ’–‡”ͳ

•—‰‰‡•–‹‰–Šƒ––Š‡›ƒ›‘–„‡ƒŽ‹‹–‹‰”‡•‘—” ‡Ǥ’‡ ‹‡•™‹–Š‘”‡•’‡ ‹ƒŽ‹œ‡†‡•–‹‰ Šƒ„‹–•ƒ›„‡‘”‡ ‘†‹–‹‘‡†„›‡•–‹‰•—„•–”ƒ–‡ƒ˜ƒ‹Žƒ„‹Ž‹–›Ǥ ‘”‡šƒ’Ž‡ǡƒ„—†ƒ ‡ ‘ˆ O. rufohirta ™ƒ• ƒ”‰‹ƒŽŽ› ƒ••‘ ‹ƒ–‡† –‘ ƒ„—†ƒ ‡ ‘ˆ ˜ƒ ƒ– •ƒ‹Ž •Š‡ŽŽ• ȋ”α ͲǤͶͳǡ ’α ͲǤͲ͸ȌǤ

 Table 1.2 ——Žƒ–‹˜‡ ˜ƒ”‹ƒ ‡ ‡š’Žƒ‹‡† „›  ‘†‡Ž• ”‡Žƒ–‹‰ ˆŽ‘™‡” ƒ† ‡•–‹‰ ”‡•‘—” ‡•–‘„‡‡•’‡ ‹‡• ‘’‘•‹–‹‘Ǥ

  š‹•ͳ š‹•ʹ š‹•͵ š‹•Ͷ ‘–ƒŽƒ”‹ƒ ‡ Smallspecies(n=62)   ——Žƒ–‹˜‡’‡” ‡–ƒ‰‡‘ˆ•’‡ ‹‡•˜ƒ”‹ƒ ‡ ͵ͺǤʹ Ͷ͸Ǥ͵ ͷʹǤͲ ͷ͸Ǥͷ  ——Žƒ–‹˜‡’‡” ‡–ƒ‰‡‘ˆ•’‡ ‹‡•Ǧ‡˜‹”‘‡–˜ƒ”‹ƒ ‡ ͷ͸Ǥͳ ͸͹Ǥͺ ͹͸Ǥʹ ͺʹǤͺ  —‘ˆ ƒ‘‹ ƒŽ‡‹‰‡˜ƒŽ—‡•  ͲǤͶͷʹ Largespecies(n=36)   ——Žƒ–‹˜‡’‡” ‡–ƒ‰‡‘ˆ•’‡ ‹‡•˜ƒ”‹ƒ ‡ ͳͷǤ͵ ʹ͵Ǥͺ ͵ͳǤ͸ ͵͸Ǥͺ  ——Žƒ–‹˜‡’‡” ‡–ƒ‰‡‘ˆ•’‡ ‹‡•Ǧ‡˜‹”‘‡–˜ƒ”‹ƒ ‡ ͵ͲǤ͹ Ͷ͹Ǥ͹ ͸͵Ǥ͵ ͹͵Ǥͺ  —‘ˆ ƒ‘‹ ƒŽ‡‹‰‡˜ƒŽ—‡•  ͲǤ͵ͺͻ 

 Ž‘™‡” ”‡•‘—” ‡• ƒŽ•‘ •Š‘™ Š‡–‡”‘‰‡‡‹–› ƒ ”‘•• –Š‡ ’ƒ”ǡ „—– ‹ ‘’ƒ”‹•‘ –‘ ‡•–‹‰ ”‡•‘—” ‡•ǡ–Š‡‹”†‹•–”‹„—–‹‘•Š‘™•‘”‡‘ˆƒ‰‡‘‰”ƒ’Š‹ ƒŽ’ƒ––‡”Ǥ Ž‘™‡”• Ž‡ƒ”Ž›’Žƒ›ƒ ‰”‡ƒ–‡” ”‘Ž‡ –Šƒ ‡•–‹‰ •—„•–”ƒ–‡• ‹ •–”— –—”‹‰ –Š‡ •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‘ˆ ‘—” „‡‡ ‘—‹–›ǡƒ ‘—–‹‰ˆ‘”ƒ‰‘‘†’ƒ”–‘ˆ–Š‡‡š’Žƒ‹‡†˜ƒ”‹ƒ ‡ ‹ ƒ„—†ƒ ‡ ƒ† ‘’‘•‹–‹‘ǤŠ‹•‘—– ‘‡‹•‹ƒ‰”‡‡‡–™‹–Š–Š‡ˆ‡™•–—†‹‡• ‘•‹†‡”‹‰„‘–Š–›’‡•‘ˆ ”‡•‘—” ‡•ȋ‘––•‡–ƒŽǤʹͲͲ͵ǡ‘—Ž•–‘ƒ† ‘‘†‡ŽŽʹͲͳͳȌǤ –‹•‹’‘”–ƒ––‘‘–‡–Šƒ––Š‡•‡ ”‡•—Ž–••Š‘—Ž†‘–‡ ‡••ƒ”‹Ž›„‡‹–‡”’”‡–‡†‹–‡”•‘ˆ‡˜‘Ž—–‹‘ƒ”›’‘ŽŽ‡•’‡ ‹ƒŽ‹œƒ–‹‘Ǥ ‘” ‹•–ƒ ‡ǡ ƒ„—†ƒ ‡ ‘ˆ L. subhirtumǡ –Š‡ ‘•– ƒ„—†ƒ– •’‡ ‹‡•ǡ ™ƒ• ’‘•‹–‹˜‡Ž› ‘””‡Žƒ–‡†–‘T.vulgarisˆŽ‘™‡”†‡•‹–›ȋ”αͲǤ͹Ͷǡ’αͲǤͲͲͲͳȌǤ ‘™‡˜‡”ǡL.subhirtum‹• Ž‡ƒ”Ž› ’‘Ž›Ž‡ –‹ ȋ‡•–”‹ ŠͳͻͻͲȌǡƒ†‹ ƒ””ƒˆ™‡Šƒ˜‡‘„•‡”˜‡†ˆ‡ƒŽ‡•‘ˆ–Š‹••’‡ ‹‡•ȋαͶͷȌ ˆ‘”ƒ‰‹‰ ‘ ͳ͵ ’Žƒ– •’‡ ‹‡• „‡Ž‘‰‹‰ –‘ ͹ ’Žƒ– ˆƒ‹Ž‹‡•Ǥ –Š‡” •–”‘‰ ƒ••‘ ‹ƒ–‹‘• ‹˜‘Ž˜‹‰ ’‘Ž›Ž‡ –‹  •’‡ ‹‡• ‹ Ž—†‡ Lasioglossum albocinctum ™‹–Š D. penthaphyllumȋ”α ͲǤ͸͵ǡ ’α ͲǤͲͲʹȌ ƒ† L. transitorium ™‹–Š Cistus •’’Ǥ ȋ”α ͲǤͷ͵ǡ ’α ͲǤͲͳȌǤ Ž‹‰‘Ž‡ –‹  •’‡ ‹‡• ƒ‡—’ƒ‹’‘”–ƒ–ˆ”ƒ –‹‘‘ˆ‘—”„‡‡ ‘—‹–›ȋʹͳ‘Ž‹‰‘Ž‡ –‹ •’‡ ‹‡•ǡͶͷ’‘Ž›Ž‡ –‹ ǡ ͳ͵ Ž‡’–‘’ƒ”ƒ•‹–‹ ǡ ƒ† ͳͻ —‘™Ȍǡ „—– ‘Ž› ‘‡ ’‘•‹–‹˜‡Ž› ‘™ ‘Ž‹‰‘Ž‡‰‡ǡ –Š‡  41 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

•–‡”ƒ ‡ƒ‡ •’‡ ‹ƒŽ‹•– Panurgus dentipesǡ ™ƒ• ƒ‘‰ –Š‡ ͳͻ ‘•– ƒ„—†ƒ– •’‡ ‹‡•Ǥ Š‡ ”‡ƒ‹‹‰‘Ž‹‰‘Ž‡ –‹ •’‡ ‹‡•™‡”‡”ƒ”‡ǡ‘ˆ–‡”‡’”‡•‡–‡†„›‘‡‘”ƒˆ‡™‹†‹˜‹†—ƒŽ•ǡƒ† ‘•–Ž›˜‹•‹–‹‰‘Ǧƒ„—†ƒ–’Žƒ–•‹–Š‡•–‡”ƒ ‡ƒ‡ǡ”ƒ••‹ ƒ ‡ƒ‡ǡ ƒ— —Žƒ ‡ƒ‡ ƒ† ‘”ƒ‰‹ƒ ‡ƒ‡Ǥ ‡˜‡”ƒŽ •–—†‹‡• Šƒ˜‡ ˆ‘—† ƒ ’‘•‹–‹˜‡ ”‡Žƒ–‹‘•Š‹’ „‡–™‡‡ ˆŽ‘™‡” ƒ† „‡‡ •’‡ ‹‡•”‹ Š‡••ȋ‘––•‡–ƒŽǤʹͲͲ͵ǡ„‡Ž‹‰‡–ƒŽǤʹͲͲͺǡ™ƒ‹•‡”ƒ† ‡†”‹šʹͲͲͺǡ ”—†‡Ž ‡–ƒŽǤʹͲͳͲǡ ”ò†‡–ƒŽǤʹͲͳͲȌǤ  ƒ””ƒˆǡ–Š‹•”‡Žƒ–‹‘•Š‹’™ƒ• ‘Ǧ•‹‰‹ˆ‹ ƒ– „—–ǡ ƒ• ‡–‹‘‡†ǡ–Š‹•™ƒ• ƒ—•‡†„›ƒ•‹‰Ž‡•‹–‡ȋ’Ž‘–ͳȌ†‹•’Žƒ›‹‰ƒ—‹“—‡„‡‡ ‘’‘•‹–‹‘ƒ† •–”‘‰Ž›†‡˜‹ƒ–‹‰ˆ”‘–Š‡‰‡‡”ƒŽ–”‡†Ǥ

‘–™‹–Š•–ƒ†‹‰ –Š‡ •‹‰‹ˆ‹ ƒ– ‡ˆˆ‡ –• ‘ˆ ˆŽ‘™‡” ”‡•‘—” ‡•ǡ ƒ• — Š ƒ• ͷͶǤͺΨ ‘ˆ –Š‡ ˜ƒ”‹ƒ ‡ ‹ •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‘ˆ –Š‡ ƒ””ƒˆ „‡‡ ‘—‹–› ”‡ƒ‹• —‡š’Žƒ‹‡†Ǥ  ƒ††‹–‹‘ –‘ ”‡•‘—” ‡ †‹•–”‹„—–‹‘ǡ ‘—‹–› ƒ••‡„Ž› †›ƒ‹ • †‡’‡† ‘ ‹‹‰”ƒ–‹‘ ‡˜‡–• ƒ† ‹–‡”ƒ –‹‘• „‡–™‡‡ •’‡ ‹‡•Ǥ ‹‰”ƒ–‹‘ Š‹•–‘”› ȋˆ‘” ‡šƒ’Ž‡ǡ –Š‡ ‘”†‡” ‘ˆ •’‡ ‹‡•ƒ””‹˜ƒŽƒ–ƒ•‹–‡Ȍƒ›•–”‘‰Ž›‹ˆŽ—‡ ‡–Š‡ˆ‹ƒŽ‘—– ‘‡ ‹ –‡”• ‘ˆ •’‡ ‹‡• ‘’‘•‹–‹‘ ȋ —ƒ‹ ʹͲͳͲȌǤ ‡ ƒ—•‡ ‘—” ’Ž‘–• ƒ”‡ Ž‘ ƒ–‡† ƒ ”‘••ƒƒ”‡ƒ‘ˆ ‘–‹‰—‘—• Šƒ„‹–ƒ–‹–‹•ˆƒ‹”–‘ƒ••—‡Š‹‰ŠŽ‡˜‡Ž•‘ˆ†‹•’‡”•ƒŽƒ‘‰’ƒ– Š‡•ǡ ™Š‹ Š ™‘—Ž† –‡† –‘ Š‘‘‰‡‹œ‡ „‡‡ †‹•–”‹„—–‹‘Ǥ ‘™‡˜‡”ǡ ‹‹‰”ƒ–‹‘ ‡˜‡–• ˆ”‘ ‘—–•‹†‡ –Š‡ Šƒ„‹–ƒ– ȋ —ƒ‹ ʹͲͳͲȌ ƒ† Ž‘ ƒŽ †‹ˆˆ‡”‡ ‡• ‹ ƒ–—”ƒŽ ‘”–ƒŽ‹–› ˆƒ –‘”• •— Š ƒ• ’”‡†ƒ–‹‘ ƒ† ’ƒ”ƒ•‹–‹•ǡ ƒ• ™‡ŽŽ ƒ• ‘’‡–‹–‹˜‡ ‹–‡”ƒ –‹‘• ƒ‘‰ „‡‡ •’‡ ‹‡• ȋ‘—Ž•–‘ ƒ† ‘‘†‡ŽŽ ʹͲͳͳȌ ƒ› ‘–”‹„—–‡ –‘ –Š‡ ƒ‹–‡ƒ ‡ ‘ˆ Ž‘ ƒŽ †‹ˆˆ‡”‡ ‡• ‹ ‘—‹–› ‘’‘•‹–‹‘Ǥ ‡ˆ‘—†ƒ‡‰ƒ–‹˜‡ƒ••‘ ‹ƒ–‹‘„‡–™‡‡–Š‡–™‘‘•–ƒ„—†ƒ–•’‡ ‹‡•ǡ Lasioglossum subhirtumƒ†Andrenadjelfensisǡ™Š‘•‡ˆŽ‹‰Š–’‡”‹‘†•‘˜‡”Žƒ’™‹†‡Ž›ǡ„—–™‡†‘‘–Šƒ˜‡ –Š‡ ‡ ‡••ƒ”› ‹ˆ‘”ƒ–‹‘ –‘ ‡•–ƒ„Ž‹•Š ™Š‡–Š‡” –Š‹• ’ƒ––‡” ‹‰Š– „‡ ƒ––”‹„—–‡† –‘ ‘’‡–‹–‹‘Ǥ‘–Š‡”ˆƒ –‘”–Šƒ– ‘—Ž†’ƒ”–‹ƒŽŽ›‡š’Žƒ‹–Š‡‰‡‘‰”ƒ’Š‹ ƒŽ’ƒ––‡”‘„•‡”˜‡†‹• ’Š‹Ž‘’ƒ–”›ǤŠ‡–‡†‡ ›‘ˆˆ‡ƒŽ‡•–‘‡•–ƒ––Š‡‹”ƒ–ƒŽ‡•–‹‰ •‹–‡ Šƒ• „‡‡ •Š‘™ ‹ •‘‡ „‡‡ •’‡ ‹‡• ƒ† ‘—Ž† ‘–”‹„—–‡ –‘ –Š‡ ‹ ”‡ƒ•‡ ‘ˆ Ž‘ ƒŽ „‡‡ †‡•‹–› ˆ‘ŽŽ‘™‹‰ ‘Ž‘‹œƒ–‹‘ ‘ˆ ƒ ‰‹˜‡ ’ƒ– Š ȋƒ‡‰ƒ ͳͻͻͲǡ –‘‹‹ ‡– ƒŽǤ ʹͲͲͲȌǤ –Š‡” —‡ƒ•—”‡† ‡˜‹”‘‡–ƒŽˆƒ –‘”••— Šƒ•–‘’‘ Ž‹ƒ–‹ ˜ƒ”‹ƒ–‹‘ ‘—Ž†ƒŽ•‘ ‘–”‹„—–‡–‘–Š‡‘„•‡”˜‡† „‡‡ ‘’‘•‹–‹‘’ƒ––‡”Ǥƒ‹Ž›ƒš‹—ƒ†‹‹—–‡’‡”ƒ–—”‡•ƒ›˜ƒ”›ƒ•— Šƒ• ͺ ͑ ƒ‘‰ ‹ ”‘•‹–‡• †‹•–ƒ– ‘Ž› ˆ‡™ Š—†”‡† ‡–‡”• ˆ”‘ ‡ƒ Š‘–Š‡”ȋ ‡”Ž›‡–ƒŽǤ ʹͲͳͲȌǤ‘‡•–—†‹‡•Šƒ˜‡ˆ‘—†’‘ŽŽ‹ƒ–‘” ‘’‘•‹–‹‘‘ˆ‹†‹˜‹†—ƒŽ’Žƒ–•–‘„‡Š‹‰ŠŽ› ‹ˆŽ—‡ ‡†„›•ƒŽŽǦ• ƒŽ‡˜ƒ”‹ƒ–‹‘‹‹ ”‘ Ž‹ƒ–‹ ˆƒ –‘”••— Šƒ••‘Žƒ”‹””ƒ†‹ƒ ‡ǡ•Šƒ†‹‰ ƒ†•‘‹Ž™‡–‡••ȋ ‡””‡”ƒͳͻͻͷǡ ƒ‘˜•ý‡–ƒŽǤʹͲͳ͵ȌǤ ƒ††‹–‹‘–‘–”›‹‰–‘‡Ž— ‹†ƒ–‡–Š‡ ˆƒ –‘”•”‡•’‘•‹„Ž‡ˆ‘”–Š‡—‡š’Žƒ‹‡†•’ƒ–‹ƒŽ˜ƒ”‹ƒ–‹‘‘„•‡”˜‡†ǡ‹–™‘—Ž†„‡‹’‘”–ƒ––‘  42 Šƒ’–‡”ͳ

‡•–ƒ„Ž‹•Š™Š‡–Š‡”–Š‡‘„•‡”˜‡†’ƒ––‡”‹••–ƒ„Ž‡‹–‹‡Ǥ‡†‘‘–‡š’‡ –‡•–‹‰•—„•–”ƒ–‡ ƒ˜ƒ‹Žƒ„‹Ž‹–›–‘˜ƒ”›— Šˆ”‘‘‡›‡ƒ”–‘–Š‡‡š–ǡ„—–„Ž‘‘‹‰‹–‡•‹–›‹•™‡ŽŽ‘™ ˆŽ— –—ƒ–‡™‹†‡Ž›ˆ”‘›‡ƒ”–‘›‡ƒ”ȋ%‰”‡ͳͻͺͺǡ””‘›‘ͳͻͻͲǡ ‘—›‡ƒ† —‹”‡ͳͻͻͳȌǡ ’‘–‡–‹ƒŽŽ›ƒˆˆ‡ –‹‰„‡‡ˆ‘”ƒ‰‹‰ƒ”‡ƒ•Ǥ

—”•–—†›†‡‘•–”ƒ–‡•–Šƒ–„‡‡ ‘—‹–‹‡•ƒ›†‹•’Žƒ› Ž‡ƒ”’ƒ––‡”• ‘ˆ •’ƒ–‹ƒŽ Š‡–‡”‘‰‡‡‹–› ƒ– ƒ ”‡Žƒ–‹˜‡Ž› •ƒŽŽ • ƒŽ‡ ȋͷͲͲǦͳͲͲͲ Ȍ ‹ ƒ”‡ƒ• ‘ˆ ‘–‹‰—‘—• •—‹–ƒ„Ž‡ Šƒ„‹–ƒ–ƒ†‹–Š‡ƒ„•‡ ‡‘ˆŽ‘ ƒŽ„ƒ””‹‡”•Ǥ ’‘”–ƒ–Ž›ǡ–Š‡‘„•‡”˜‡†Š‡–‡”‘‰‡‡‹–›‹•‘– ‹””‡‰—Žƒ”ǡ „—– ˆ‘ŽŽ‘™• ƒ ‰‡‘‰”ƒ’Š‹ ƒŽ ’ƒ––‡”ǡ ƒ† ‹• ‘Ž› ’ƒ”–Ž› ‡š’Žƒ‹‡† „› ˆŽ‘™‡” ƒ˜ƒ‹Žƒ„‹Ž‹–›Ǥ Š‹• ”‡•—Ž– ‹• ”‡ƒ”ƒ„Ž‡ „‡ ƒ—•‡ „‡‡• ƒ”‡ Š‹‰ŠŽ› ‘„‹Ž‡ ‘”‰ƒ‹•• ȋ„‘–Š ‹ –‡”• ‘ˆ ˆ‘”ƒ‰‹‰ ƒ† †‹•’‡”•ƒŽȌǡ ƒ† –Š‡”‡ˆ‘”‡ ‘‡ ‹‰Š– ‡š’‡ – ƒ ‘”‡ Š‘‘‰‡‡‘—• †‹•–”‹„—–‹‘Ǥ ‡ ƒ—•‡ †‹ˆˆ‡”‡– „‡‡ •’‡ ‹‡• Šƒ˜‡ †‹ˆˆ‡”‡– ˆŽ‘™‡”’”‡ˆ‡”‡ ‡•ƒ††‹ˆˆ‡”‹ –Š‡‹” ’‘ŽŽ‹ƒ–‹‰ ƒ„‹Ž‹–‹‡•ǡ ‘—” ”‡•—Ž–• Šƒ˜‡ ‹’‘”–ƒ– ‹’Ž‹ ƒ–‹‘• ˆ‘” Ž‘ ƒŽ ’‘ŽŽ‹ƒ–‹‘ †›ƒ‹ •Ǥ ‡˜‡”ƒŽ •–—†‹‡• Šƒ˜‡ ˆ‘—† †‹ˆˆ‡”‡ ‡• ‹ ”‡’”‘†— –‹˜‡ •— ‡•• ƒ‘‰ ’‘’—Žƒ–‹‘•˜‹•‹–‡†„›†‹ˆˆ‡”‡–’‘ŽŽ‹ƒ–‘”•ȋ”‹ ‡‡–ƒŽǤʹͲͲͷǡ ”—‡– ƒ† ™‡‡– ʹͲͲ͸ǡ ׏‡œ‡–ƒŽǤʹͲͲ͹ȌǤ—”•–—†›•—‰‰‡•–•–Šƒ–†‹ˆˆ‡”‡ ‡•‹’‘ŽŽ‹ƒ–‹‘Ž‡˜‡Ž•ƒ›ƒŽ•‘‘ —” ™‹–Š‹ ƒ ’Žƒ– ’‘’—Žƒ–‹‘ ƒ• ƒ ”‡•—Ž– ‘ˆ Š‡–‡”‘‰‡‡‘—• Ž‘ ƒŽ ’‘ŽŽ‹ƒ–‘” †‹•–”‹„—–‹‘Ǥ —” ”‡•—Ž–•ƒŽ•‘Šƒ˜‡‹’‘”–ƒ– ‘•‡“—‡ ‡•ˆ‘”–Š‡•–—†›‘ˆ•’ƒ–‹ƒŽ ˜ƒ”‹ƒ–‹‘ ‘ˆ ’Žƒ–Ǧ ’‘ŽŽ‹ƒ–‘”‡–™‘”•ȋ‘”ƒŽ‡•ƒ†žœ“—‡œʹͲͲͺǡ ƒ‘˜•ý‡–ƒŽǤʹͲͳ͵Ȍǡƒ•‘˜‡”ƒŽŽ’‘ŽŽ‹ƒ–‘” ‘—‹–› ‘’‘•‹–‹‘ƒ›„‡ Šƒ‰‹‰ƒ–•ƒŽŽ‡”• ƒŽ‡•–Šƒ’”‡˜‹‘—•Ž›–Š‘—‰Š–Ǥ



Acknowledgements

‡ƒ”‡˜‡”›‰”ƒ–‡ˆ—Ž–‘Ǥžœƒ”‘ǡǤǤ‡“—‡•‡•ǡǤ‡˜‘Ž–וǡ ǤǤ‹ŽŽ‡ŽŽƒ•ǡǤƒ•ǡǤ‹‡”ƒǡ Ǥ‹Žƒ†±•ǡǤŽƒ˜‹ƒǡǤג‡œǡǤ ‘œžŽ‡œǡƒ†Ǥ—Ó‘œˆ‘” –Š‡‹” Š‡Ž’ ™‹–Š ˆ‹‡Ž† ƒ† Žƒ„‘”ƒ–‘”›™‘”Ǥ‡–ŠƒǤƒ‹••‹°”‡ǡǤ”‘‡‰‡ǡƒ† Ǥ Ǥƒ‡ˆ‘”–‹’•‘–Š‡—•‡‘ˆ’ƒ –”ƒ’•ǡƒ†•‡˜‡”ƒŽ•’‡ ‹ƒŽ‹•–•ȋǤ‘‰—•ŠǡǤƒ•–”‘ǡ Ǥ Ǥƒ–Š‡ǡǤòŽŽ‡”ǡ Ǥ Ǥ”–‹œǡǤƒ—Ž›ǡ Ǥ”ƒœǡǤ‹• ŠǡǤ Š‡— ŠŽǡƒ†Ǥ Š™ƒ”œȌˆ‘”–Š‡‹”Š‡Ž’™‹–Š„‡‡‹†‡–‹ˆ‹ ƒ–‹‘Ǥ‡ƒ”‡ ƒŽ•‘–Šƒˆ—Ž–‘‹’—–ƒ ‹×†‡ƒ” ‡Ž‘ƒˆ‘”’‡”‹••‹‘–‘™‘”‹ ƒ””ƒˆƒ–—”ƒŽƒ”ǡƒ† –‘–Š‡•–ƒˆˆ‘ˆ–Š‡’ƒ”ˆ‘”ˆƒ ‹Ž‹–ƒ–‹‰‘—”™‘”Ǥ



 43 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

References

 ‡”Ž›ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͳͲǤ Š‡ ‰‡‘‰”ƒ’Š› ‘ˆ Ž‹ƒ–‡ Šƒ‰‡ǣ ’Ž‹ ƒ–‹‘• ˆ‘” ‘•‡”˜ƒ–‹‘ „‹‘‰‡‘‰”ƒ’Š›ǤǦ‹˜‡”•Ǥ‹•–”‹„Ǥͳ͸ǣͶ͹͸ȂͶͺ͹Ǥ

%‰”‡ǡ ǤͳͻͺͺǤ‡–™‡‡Ǧ›‡ƒ”˜ƒ”‹ƒ–‹‘‹ˆŽ‘™‡”‹‰ƒ†ˆ”—‹–•‡–‹ˆ”‘•–Ǧ’”‘‡ƒ†ˆ”‘•–Ǧ•Š‡Ž–‡”‡† ’‘’—Žƒ–‹‘•‘ˆ†‹‘‡ ‹‘—•RubuschamaemorusǤǦ‡ ‘Ž‘‰‹ƒ͹͸ǣͳ͹ͷȂͳͺ͵Ǥ

–‘‹‹ǡǤ‡–ƒŽǤʹͲͲͲǤŠ‹Ž‘’ƒ–”›‹–Š‡‡‘–”‘’‹ ƒŽ‰”‘—†Ǧ‡•–‹‰•‘Ž‹–ƒ”›†‹‰‰‡”„‡‡ǡDiadasina distinctaȋ ‘Ž„‡”‰ǡͳͻͲ͵Ȍȋ ›‡‘’–‡”ƒǣ’‹†ƒ‡Ȍƒ–ƒ‡•–‹‰•‹–‡‹•‘—–Š‡””ƒœ‹ŽǤǦ‡˜Ǥ –‘ŽǤʹǣͳͳͳȂͳͳͻǤ

””‘›‘ǡ ǤͳͻͻͲǤ‹–‘• Ž‹ƒ–‹ ‘•›†‡ˆŽ‘”ƒ ‹×‡ƒ–‘””ƒŽ‡•†‡Ž†‡•’ƒÓƒǤǦƒ‰ƒ• ƒŽ‹ƒͳ͸ǣʹͷȂ ͷͲǤ

‡‡ƒǡǤƒ†ƒ–‹‡•ǡ ǤǤǤʹͲͲͲǤ‘‰Ǧ”ƒ‰‡ˆ‘”ƒ‰‹‰„›–Š‡Š‘‡›Ǧ„‡‡ǡApismelliferaǤǦ — –Ǥ ‘ŽǤͳͶǣͶͻͲȂͶͻ͸Ǥ

Šƒ––ƒ Šƒ”›ƒǡǤ‡–ƒŽǤʹͲͲʹǤ”‡”‘ƒ†•ƒ†”ƒ‹Ž”‘ƒ†•„ƒ””‹‡”•–‘„—„Ž‡„‡‡‘˜‡‡–‹ƒ–‡’‡”ƒ–‡ •—„—”„ƒ ‘•‡”˜ƒ–‹‘ƒ”‡ƒǫǦ‹‘ŽǤ‘•‡”˜ǤͳͲͻǣ͵͹ȂͶͷǤ

‘• Šǡ Ǥ ƒ† ‹ ‡•ǡ Ǥ ʹͲͲ͸Ǥ ‡Žƒ–‹‘•Š‹’ „‡–™‡‡ „‘†› •‹œ‡ǡ ’”‘˜‹•‹‘‹‰ ”ƒ–‡ǡ Ž‘‰‡˜‹–› ƒ† ”‡’”‘†— –‹˜‡•— ‡••‹ˆ‡ƒŽ‡•‘ˆ–Š‡•‘Ž‹–ƒ”›„‡‡OsmiacornutaǤǦ‡Šƒ˜Ǥ ‘ŽǤ‘ ‹‘„‹‘ŽǤ͸Ͳǣ ʹ͸Ȃ͵͵Ǥ

”‘•‹ǡǤ Ǥ‡–ƒŽǤʹͲͲͺǤŠ‡‡ˆˆ‡ –•‘ˆˆ‘”‡•–ˆ”ƒ‰‡–ƒ–‹‘‘„‡‡ ‘—‹–‹‡•‹–”‘’‹ ƒŽ ‘—–”›•‹†‡Ǥ Ǧ Ǥ’’ŽǤ ‘ŽǤͶͷǣ͹͹͵Ȃ͹ͺ͵Ǥ

”—‡–ǡ Ǥ ƒ† ™‡‡–ǡ Ǥ Ǥ ʹͲͲ͸Ǥ ’ƒ – ‘ˆ ‹•‡ – ’‘ŽŽ‹ƒ–‘” ‰”‘—’ ƒ† ˆŽ‘”ƒŽ †‹•’Žƒ› •‹œ‡ ‘ ‘—– ”‘••‹‰”ƒ–‡ǤǦ˜‘Ž—–‹‘͸Ͳǣʹ͵ͶȂʹͶ͸Ǥ

ƒ”˜‡ŽŽǡǤ‡–ƒŽǤʹͲͳʹǤ‘Ž‡ —Žƒ”ƒ†•’ƒ–‹ƒŽƒƒŽ›•‡•”‡˜‡ƒŽ Ž‹•„‡–™‡‡ ‘Ž‘›Ǧ•’‡ ‹ˆ‹ ˆ‘”ƒ‰‹‰ †‹•–ƒ ‡ƒ†Žƒ†• ƒ’‡ǦŽ‡˜‡Ž”‡•‘—” ‡ƒ˜ƒ‹Žƒ„‹Ž‹–›‹–™‘„—„Ž‡„‡‡•’‡ ‹‡•ǤǦ‹‘•ͳʹͳǣ͹͵ͶȂ ͹ͶʹǤ

ƒ•–ƒ‰‡›”‘Žǡ Ǥ ƒ† ƒ –‡Žǡ Ǥ ʹͲͳʹǤ ”ƒ˜‡Ž‹‰ ’Žƒ–Ǧƒ‹ƒŽ †‹˜‡”•‹–› ”‡Žƒ–‹‘•Š‹’•ǣ  ‡–ƒǦ ”‡‰”‡••‹‘ƒƒŽ›•‹•ǤǦ ‘Ž‘‰›ͻ͵ǣʹͳͳͷȂʹͳʹͶǤ

 44 Šƒ’–‡”ͳ

„‡Ž‹‰ǡǤ‡–ƒŽǤʹͲͲͺǤ ‘™†‘‡•’Žƒ–”‹ Š‡••ƒˆˆ‡ –’‘ŽŽ‹ƒ–‘””‹ Š‡••ƒ†–‡’‘”ƒŽ•–ƒ„‹Ž‹–›‘ˆ ˆŽ‘™‡”˜‹•‹–•ǫǦ‹‘•ͳͳ͹ǣͳͺͲͺȂͳͺͳͷǤ

„‡Ž‹‰ǡ Ǥ ‡– ƒŽǤ ʹͲͳʹǤ —Ž–‹–”‘’Š‹  ‡ˆˆ‡ –• ‘ˆ ‡š’‡”‹‡–ƒŽ Šƒ‰‡• ‹ ’Žƒ– †‹˜‡”•‹–› ‘ ƒ˜‹–›Ǧ ‡•–‹‰„‡‡•ǡ™ƒ•’•ǡƒ†–Š‡‹”’ƒ”ƒ•‹–‘‹†•ǤǦ‡ ‘Ž‘‰‹ƒͳ͸ͻǣͶͷ͵ȂͶ͸ͷǤ

Ž–œǡ Ǥ ‡– ƒŽǤ ʹͲͲ͵Ǥ ‡•–‹‰ ƒ† ‡•– –”‡‡• ‘ˆ •–‹‰Ž‡•• „‡‡• ȋ’‹†ƒ‡ǣ ‡Ž‹’‘‹‹Ȍ ‹ Ž‘™Žƒ† †‹’–‡”‘ ƒ”’ ˆ‘”‡•–• ‹ ƒ„ƒŠǡ ƒŽƒ›•‹ƒǡ ™‹–Š ‹’Ž‹ ƒ–‹‘• ˆ‘” ˆ‘”‡•– ƒƒ‰‡‡–Ǥ Ǧ ‘”Ǥ  ‘ŽǤ ƒƒ‰‡Ǥͳ͹ʹǣ͵ͲͳȂ͵ͳ͵Ǥ

”ò†ǡ Ǥ‡–ƒŽǤʹͲͳͲǤ‘ŽŽ‹ƒ–‘”†‹˜‡”•‹–›ƒ†•’‡ ‹ƒŽ‹œƒ–‹‘‹”‡Žƒ–‹‘–‘ˆŽ‘™‡”†‹˜‡”•‹–›ǤǦ‹‘•ͳͳͻǣ ͳͷͺͳȂͳͷͻͲǤ

—ƒ‹ǡǤʹͲͳͲǤ‘—‹–›ƒ••‡„Ž›†›ƒ‹ •‹•’ƒ ‡ǤǦ ǣ‡”Š‘‡ˆǡ ǤǤƒ†‘”‹ǡǤ Ǥȋ‡†•Ȍǡ ‘—‹–›‡ ‘Ž‘‰›ǣ’”‘ ‡••‡•ǡ‘†‡Ž•ǡƒ†ƒ’’Ž‹ ƒ–‹‘•Ǥšˆ‘”†‹˜‡”•‹–›”‡••ǡ’’ǤͶͷȂͷͶǤ

ƒ–ŠƒǡǤƒ†• Šƒ”–‡ǡǤʹͲͲʹǤ ‘”ƒ‰‹‰”ƒ‰‡•‘ˆ•‘Ž‹–ƒ”›„‡‡•ǤǦ Ǥ‹Ǥ ‘ŽǤ͹ͳǣ͹ͷ͹Ȃ͹͸ͶǤ

ƒ–Šƒǡ Ǥ ‡– ƒŽǤ ͳͻͻͶǤ ”ƒ’Ǧ‡•–‹‰ „‡‡• ƒ† ™ƒ•’• ‘Ž‘‹œ‹‰ •‡–Ǧƒ•‹†‡ ˆ‹‡Ž†•ǣ •— ‡••‹‘ ƒ† „‘†›•‹œ‡ǡƒƒ‰‡‡–„› —––‹‰ƒ†•‘™‹‰ǤǦ‡ ‘Ž‘‰‹ƒͻͺǣͺȂͳͶǤ

׏‡œǡ ǤǤ‡–ƒŽǤʹͲͲ͹Ǥ‘ŽŽ‹ƒ–‘”†‹˜‡”•‹–›ƒˆˆ‡ –•’Žƒ–”‡’”‘†— –‹‘ƒ†”‡ ”—‹–‡–ǣ–Š‡–”ƒ†‡‘ˆˆ• ‘ˆ‰‡‡”ƒŽ‹œƒ–‹‘ǤǦ‡ ‘Ž‘‰‹ƒͳͷ͵ǣͷͻ͹Ȃ͸ͲͷǤ

‘—Ž•‘ǡǤʹͲͳͲǤ—„Ž‡„‡‡•Ǥ‡Šƒ˜‹‘—”ǡ‡ ‘Ž‘‰›ƒ† ‘•‡”˜ƒ–‹‘ǤǦšˆ‘”†‹˜‡”•‹–›”‡••Ǥ

”‡‡Ž‡ƒˆǡǤǤ‡–ƒŽǤʹͲͲ͹Ǥ‡‡ˆ‘”ƒ‰‹‰”ƒ‰‡•ƒ†–Š‡‹””‡Žƒ–‹‘•Š‹’–‘„‘†›•‹œ‡ǤǦ‡ ‘Ž‘‰‹ƒͳͷ͵ǣ ͷͺͻȂͻ͸Ǥ

”—†‡Žǡ Ǥ ‡– ƒŽǤ ʹͲͳͲǤ Ž‘”ƒŽ ƒ† ‡•–‹‰ ”‡•‘—” ‡•ǡ Šƒ„‹–ƒ– •–”— –—”‡ǡ ƒ† ˆ‹”‡ ‹ˆŽ—‡ ‡ „‡‡ †‹•–”‹„—–‹‘ƒ ”‘••ƒ‘’‡Ǧˆ‘”‡•–‰”ƒ†‹‡–ǤǦ ‘ŽǤ’’ŽǤʹͲǣͳ͸͹ͺȂͻʹǤ

—±†‘–ǡ Ǥ ‡– ƒŽǤ ʹͲͲͻǤ ‡Žƒ–‹‘•Š‹’ „‡–™‡‡ „‘†› •‹œ‡ ƒ† Š‘‹‰ ƒ„‹Ž‹–› ‹ –Š‡ ‰‡—• •‹ƒ ȋ ›‡‘’–‡”ƒǢ‡‰ƒ Š‹Ž‹†ƒ‡ȌǤǦ ‘ŽǤ–‘‘ŽǤ͵ͶǣͳͷͺȂͳ͸ͳǤ

‡””‡”ƒǡǤǤͳͻͺͺǤƒ”‹ƒ–‹‘‹—–—ƒŽ‹••ǣ–Š‡•’ƒ–‹‘Ǧ–‡’‘”ƒŽ‘•ƒ‹ ‘ˆƒ’‘ŽŽ‹ƒ–‘”ƒ••‡„Žƒ‰‡ǤǦ ‹‘ŽǤ Ǥ‹Ǥ‘ ͵ͷǣͻͷȂͳʹͷǤ

 45 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

‡””‡”ƒǡǤǤͳͻͻͷǤ‹ ”‘ Ž‹ƒ–‡ƒ†‹†‹˜‹†—ƒŽ˜ƒ”‹ƒ–‹‘‹’‘ŽŽ‹ƒ–‘”•ǣ Ž‘™‡”‹‰’Žƒ–•ƒ”‡‘”‡ –Šƒ–Š‡‹”ˆŽ‘™‡”•ǤǦ ‘Ž‘‰›͹͸ǣͳͷͳ͸ȂͳͷʹͶǤ

‘ŽŽ‹‰ǡǤǤͳͻͻʹǤ”‘••Ǧ• ƒŽ‡‘”’Š‘Ž‘‰›ǡ‰‡‘‡–”›ǡƒ††›ƒ‹ •‘ˆ‡ ‘•›•–‡•ǤǦ ‘ŽǤ‘‘‰”Ǥ ͸ʹǣͶͶ͹Ǥ

‘Ž›‘ƒǡǤ‡–ƒŽǤʹͲͲͷǤ‡–ƒ ‘—‹–‹‡•ǣ’ƒ–‹ƒŽ†›ƒ‹ •ƒ† ‡ ‘Ž‘‰‹ ƒŽ ‘—‹–‹‡•ǤǦŠ‹ ƒ‰‘ ”‡••Ǥ

‘—›‡ǡǤǤƒ† —‹”‡ǡǤǤͳͻͻͳǤˆˆ‡ –•‘ˆ•‘™’ƒ ‘–‹‹‰ƒ†ƒ„—†ƒ ‡‘ˆˆŽ‘™‡”‹‰‹ Delphinium nelsoniiȋƒ— —Žƒ ‡ƒ‡Ȍǣ‹’Ž‹ ƒ–‹‘•ˆ‘” Ž‹ƒ–‡ Šƒ‰‡ǤǦǤ Ǥ‘–Ǥ ͹ͺǣ ͻͻ͹Ȃ ͳͲͲͳǤ

ƒ‘˜•ýǡǤ‡–ƒŽǤʹͲͳ͵Ǥ‘•’‡ ‹ˆ‹ ƒ†Š‡–‡”‘•’‡ ‹ˆ‹ ’Žƒ–†‡•‹–‹‡•ƒ–•ƒŽŽǦ• ƒŽ‡ ƒ†”‹˜‡’Žƒ–Ǧ ’‘ŽŽ‹ƒ–‘”‹–‡”ƒ –‹‘•ǤǦ‘‡ͺǣ‡͹͹͵͸ͳǤ

™ƒ‹•‡”ǡǤǤƒ† ‡†”‹šǡǤǤʹͲͲͺǤ‹˜‡”•‹–›ƒ†ƒ„—†ƒ ‡‘ˆ„‡‡•ȋ ›‡‘’–‡”ƒǣ’‹ˆ‘”‡•Ȍ‹ ƒ–‹˜‡ƒ†”—†‡”ƒŽ‰”ƒ••Žƒ†•‘ˆƒ‰”‹ —Ž–—”ƒŽŽ›†‘‹ƒ–‡†Žƒ†• ƒ’‡•ǤǦ‰”‹ Ǥ ‘•›•–Ǥ˜‹”‘Ǥ ͳʹͶǣʹͲͲȂʹͲͶǤ

‡’æǡ Ǥƒ†e‹Žƒ—‡”ǡǤʹͲͲ͵Ǥ—Ž–‹˜ƒ”‹ƒ–‡ƒŽ›•‹•‘ˆ ‘Ž‘‰‹ ƒŽƒ–ƒ—•‹‰ǤǦƒ„”‹†‰‡ ‹˜‡”•‹–›”‡••Ǥ

‹ Ž‡›ǡǤǤ‡–ƒŽǤͳͻͻͻǤ’ƒ–‹ƒŽ’”‡†‹ –ƒ„‹Ž‹–›ƒ†”‡•‘—” ‡•’‡ ‹ƒŽ‹œƒ–‹‘‘ˆ„‡‡•ȋ ›‡‘’–‡”ƒǣ ’‘‹†‡ƒȌƒ–ƒ•—’‡”ƒ„—†ƒ–ǡ™‹†‡•’”‡ƒ†”‡•‘—” ‡ǤǦ‹‘ŽǤ Ǥ‹Ǥ‘ ǤǣͳͳͻȂͳͶ͹Ǥ

‘”ƒŽ‡•ǡ Ǥ Ǥ ƒ† žœ“—‡œǡ Ǥ Ǥ ʹͲͲͺǤ Š‡ ‡ˆˆ‡ – ‘ˆ •’ƒ ‡ ‹ ’Žƒ–Ǧƒ‹ƒŽ —–—ƒŽ‹•–‹  ‡–™‘”•ǣ ‹•‹‰Š–•ˆ”‘ƒ•‹—Žƒ–‹‘•–—†›ǤǦ‹‘•ͳͳ͹ǣͳ͵͸ʹȂͳ͵͹ͲǤ

òŽŽ‡”ǡ Ǥ ‡– ƒŽǤ ʹͲͲ͸Ǥ —ƒ–‹–ƒ–‹˜‡ ’‘ŽŽ‡ ”‡“—‹”‡‡–• ‘ˆ •‘Ž‹–ƒ”› „‡‡•ǣ ’Ž‹ ƒ–‹‘• ˆ‘” „‡‡ ‘•‡”˜ƒ–‹‘ƒ†–Š‡‡˜‘Ž—–‹‘‘ˆ„‡‡ȂˆŽ‘™‡””‡Žƒ–‹‘•Š‹’•ǤǦ‹‘ŽǤ‘•‡”˜Ǥͳ͵Ͳǣ͸ͲͶȂ͸ͳͷǤ

—””ƒ›ǡǤǤ‡–ƒŽǤʹͲͳʹǤ‘ ƒŽǦ• ƒŽ‡ˆƒ –‘”••–”— –—”‡™‹Ž†„‡‡ ‘—‹–‹‡•‹’”‘–‡ –‡†ƒ”‡ƒ•ǤǦ Ǥ ’’ŽǤ ‘ŽǤͶͻǣͻͻͺȂͳͲͲͺǤ

•„‘”‡ǡ ǤǤ‡–ƒŽǤͳͻͻͻǤŽƒ†• ƒ’‡Ǧ• ƒŽ‡‘ˆ„—„Ž‡„‡‡ˆ‘”ƒ‰‹‰•–—†›”ƒ‰‡ƒ† ‘•–ƒ ›—•‹‰ Šƒ”‘‹ ”ƒ†ƒ”ǤǦ Ǥ’’ŽǤ ‘ŽǤ͵͸ǣͷͳͻȂͷ͵͵Ǥ

 46 Šƒ’–‡”ͳ

‘’‹ ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͳ͵Ǥ ˜ƒŽ—ƒ–‹‘ ‘ˆ ‘‘ ‡–Š‘†• ˆ‘” •ƒ’Ž‹‰ ‹˜‡”–‡„”ƒ–‡ ’‘ŽŽ‹ƒ–‘” ƒ••‡„Žƒ‰‡•ǣ‡–•ƒ’Ž‹‰‘—–Ǧ’‡”ˆ‘”’ƒ–”ƒ’•ǤǦ‘‡ͺǣ‡͸͸͸͸ͷǤ

‘––•ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲ͵Ǥ ‹‹‰ „‡‡• ƒ† ˆŽ‘™‡”•ǣ ‘™ †‘ ˆŽ‘”ƒŽ ‘—‹–‹‡• •–”— –—”‡ ’‘ŽŽ‹ƒ–‘” ‘—‹–‹‡•ǫǦ ‘Ž‘‰›ͺͶǣʹ͸ʹͺȂʹ͸ͶʹǤ

‘––•ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͶǤ ‡ –ƒ” ”‡•‘—” ‡ †‹˜‡”•‹–› ‘”‰ƒ‹•‡• ˆŽ‘™‡”Ǧ˜‹•‹–‘” ‘—‹–› •–”— –—”‡Ǥ Ǧ –‘‘ŽǤš’Ǥ’’ŽǤͳͳ͵ǣͳͲ͵ȂͳͲ͹Ǥ

‘––•ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͷǤ ‘Ž‡ ‘ˆ ‡•–‹‰ ”‡•‘—” ‡• ‹ ‘”‰ƒ‹•‹‰ †‹˜‡”•‡ „‡‡ ‘—‹–‹‡• ‹ ƒ ‡†‹–‡””ƒ‡ƒŽƒ†• ƒ’‡ǤǦ ‘ŽǤ–‘‘ŽǤ͵Ͳǣ͹ͺȂͺͷǤ

”‹ ‡ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͷǤ ‡’‘”ƒŽ ƒ† •’ƒ–‹ƒŽ ˜ƒ”‹ƒ–‹‘ ‘ˆ ƒ ‘–ƒ‡ Š‡”„ǣ ƒ •‡˜‡Ǧ›‡ƒ” •–—†›Ǥ Ǧ  ‘Ž‘‰›ͺ͸ǣʹͳͲ͸Ȃʹͳͳ͸Ǥ

‡˜‡Ž‘’‡–‘”‡‡ƒʹͲͳͲǤǣŽƒ‰—ƒ‰‡ƒ†‡˜‹”‘‡–ˆ‘”•–ƒ–‹•–‹ ƒŽ ‘’—–‹‰Ǥ‹’”‡••Ǥ

ƒ‰‡Žǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͳͲǤ ǣ ƒ ‘’”‡Š‡•‹˜‡ ƒ’’Ž‹ ƒ–‹‘ ˆ‘” ’ƒ–‹ƒŽ ƒŽ›•‹• ‹ ƒ ”‘‡ ‘Ž‘‰›Ǥ Ǧ  ‘‰”ƒ’Š›ȋ‘’ǤȌǤ͵͵ǣͶ͸ȂͷͲǤ

‡•‡–ƒ”‹–•ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͷǤ ƒ„‹–ƒ– •‡Ž‡ –‹‘ǡ •’‡ ‹‡• ‹–‡”ƒ –‹‘•ǡƒ†’”‘ ‡••‡•‘ˆ ‘—‹–› ƒ••‡„Ž› ‹ ‘’Ž‡š Žƒ†• ƒ’‡•Ǥ Ǧ ǣ ‘Ž›‘ƒǡ Ǥ ‡– ƒŽǤ ȋ‡†•Ȍǡ ‡–ƒ ‘—‹–‹‡•ǣ •’ƒ–‹ƒŽ †›ƒ‹ •ƒ†‡ ‘Ž‘‰‹ ƒŽ ‘—‹–‹‡•Ǥ‹˜‡”•‹–›‘ˆŠ‹ ƒ‰‘”‡••ǡ’’Ǥ͵͹ͶȂ͵ͻͺǤ

‹ ‡––•ǡǤ Ǥ‡–ƒŽǤʹͲͲͺǤƒ†• ƒ’‡‡ˆˆ‡ –•‘ ”‘’’‘ŽŽ‹ƒ–‹‘•‡”˜‹ ‡•ǣƒ”‡–Š‡”‡‰‡‡”ƒŽ’ƒ––‡”•ǫǦ  ‘ŽǤ‡––ǤͳͳǣͶͻͻȂͷͳͷǤ

‹ Ž‡ˆ•ǡǤǤͳͻͺ͹Ǥ‘—‹–›†‹˜‡”•‹–›ǣ”‡Žƒ–‹˜‡”‘Ž‡•‘ˆŽ‘ ƒŽƒ†”‡‰‹‘ƒŽ’”‘ ‡••‡•ǤǦ ‹‡ ‡ʹ͵ͷǣ ͳ͸͹Ȃͳ͹ͳǤ

‘—Ž•–‘ǡ Ǥ Ǥ ƒ† ‘‘†‡ŽŽǡ Ǥ ʹͲͳͳǤ Š‡ ”‘Ž‡ ‘ˆ ”‡•‘—” ‡• ƒ†”‹••‹”‡‰—Žƒ–‹‰™‹Ž†„‡‡ ’‘’—Žƒ–‹‘•ǤǦ—Ǥ‡˜Ǥ–‘‘ŽǤͷ͸ǣʹͻ͵Ȃ͵ͳʹǤ

 Šƒˆˆ‡”•ǡǤǤ‡–ƒŽǤʹͲͲͺǤ”–Š”‘’‘†ƒ••‡„Žƒ‰‡•ƒ”‡„‡•–’”‡†‹ –‡†„›’Žƒ–•’‡ ‹‡• ‘’‘•‹–‹‘ǤǦ  ‘Ž‘‰›ͺͻǣ͹ͺʹȂ͹ͻͶǤ

‹–ŠǡǤǤ‡–ƒŽǤʹͲͳʹǤ‘ŽŽ‡—•‡„›‡‰ƒŽ‘’–ƒ•™‡ƒ–„‡‡•‹”‡Žƒ–‹‘–‘”‡•‘—” ‡ƒ˜ƒ‹Žƒ„‹Ž‹–›‹ƒ –”‘’‹ ƒŽˆ‘”‡•–ǤǦ ‘ŽǤ–‘‘ŽǤ͵͹ǣ͵ͲͻȂ͵ͳ͹Ǥ

 47 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

–‡ˆˆƒǦ‡™‡–‡”ǡ Ǥƒ†‡•–’ŠƒŽǡǤʹͲͲͺǤŠ‡‹–‡”’Žƒ›‘ˆ’‘ŽŽ‹ƒ–‘”†‹˜‡”•‹–›ǡ’‘ŽŽ‹ƒ–‹‘•‡”˜‹ ‡• ƒ†Žƒ†• ƒ’‡ Šƒ‰‡ǤǦ Ǥ’’ŽǤ ‘ŽǤͶͷǣ͹͵͹Ȃ͹ͶͳǤ

–‡ˆˆƒǦ‡™‡–‡”ǡ Ǥ ‡– ƒŽǤ ʹͲͲʹǤ  ƒŽ‡Ǧ†‡’‡†‡– ‡ˆˆ‡ –• ‘ˆ Žƒ†• ƒ’‡ ‘–‡š– ‘ –Š”‡‡ ’‘ŽŽ‹ƒ–‘” ‰—‹Ž†•ǤǦ ‘Ž‘‰›ͺ͵ǣͳͶʹͳȂͳͶ͵ʹǤ

‡””ƒƒǡǤ Ǥ Ǥƒ†e‹Žƒ—‡”ǡǤʹͲͲʹǤƒ‘ ‘ˆ‘”‹†‘™•˜‡”•‹‘ͶǤͷǤ‹‘‡–”‹•ǦŽƒ–‡•‡ƒ” Š –‡”ƒ–‹‘ƒŽǤ

‡ƒ„Ž‡•ǡǤǤƒ†‹’Ž‡›ǡǤǤͳͻͻͻǤ‘†‡”ƒ’’Ž‹‡†•–ƒ–‹•–‹ •™‹–ŠǦǤǦ’”‹‰‡”ǡ‡™‘”Ǥ

‹ƒƒǡǤ Ǥ‡–ƒŽǤʹͲͳʹǤ ‘™™‡ŽŽ†‘™‡—†‡”•–ƒ†Žƒ†• ƒ’‡‡ˆˆ‡ –•‘’‘ŽŽ‹ƒ–‘”•ƒ†’‘ŽŽ‹ƒ–‹‘ •‡”˜‹ ‡•ǫǦ Ǥ‘ŽŽ‹ƒ–Ǥ ‘ŽǤ͹ǣ͵ͳȂͶͳǤ

‹ ‡•ǡ Ǥ ƒ† ‘• Šǡ Ǥ ʹͲͲͲǤ ‡•– •‹–‡ ‘”‹‡–ƒ–‹‘ ƒ† ”‡Ž‘ ƒ–‹‘ ‘ˆ ’‘’—Žƒ–‹‘• ‘ˆ –Š‡ ‘” Šƒ”† ’‘ŽŽ‹ƒ–‘”Osmiacornutaȋ ›‡‘’–‡”ƒǣ‡‰ƒ Š‹Ž‹†ƒ‡ȌǤǦ˜‹”‘Ǥ–‘‘ŽǤʹͻǣ͸ͻȂ͹ͷǤ

‹•• Š‡”ǡ Ǥ Ǥ ƒ† ‡‡Ž‡›ǡ Ǥ Ǥ ͳͻͺʹǤ ‘”ƒ‰‹‰ •–”ƒ–‡‰› ‘ˆ Š‘‡›„‡‡ ‘Ž‘‹‡• ‹ ƒ –‡’‡”ƒ–‡ †‡ ‹†—‘—•ˆ‘”‡•–ǤǦ ‘Ž‘‰›͸͵ǣͳ͹ͻͲȂͳͺͲͳǤ

ƒŽ–Š‡”Ǧ ‡ŽŽ™‹‰ǡǤƒ† ”ƒŽǡǤʹͲͲͲǤ ‘”ƒ‰‹‰†‹•–ƒ ‡•‘ˆBombusmuscorum,Bombuslapidariusǡ ƒ†Bombusterrestrisȋ ›‡‘’–‡”ƒǡ’‹†ƒ‡ȌǤǦ Ǥ •‡ –‡Šƒ˜Ǥͳ͵ǣʹ͵ͻȂʹͶ͹Ǥ

‡•–’ŠƒŽǡǤ‡–ƒŽǤʹͲͲ͸Ǥ ‘”ƒ‰‹‰–”‹’†—”ƒ–‹‘‘ˆ„—„Ž‡„‡‡•‹”‡Žƒ–‹‘–‘Žƒ†• ƒ’‡Ǧ™‹†‡”‡•‘—” ‡ ƒ˜ƒ‹Žƒ„‹Ž‹–›ǤǦ ‘ŽǤ–‘‘ŽǤ͵ͳǣ͵ͺͻȂ͵ͻͶǤ

‡•–’ŠƒŽǡǤ‡–ƒŽǤʹͲͲͺǤ‡ƒ•—”‹‰„‡‡†‹˜‡”•‹–›‹†‹ˆˆ‡”‡–—”‘’‡ƒŠƒ„‹–ƒ–•ƒ†„‹‘‰‡‘‰”ƒ’Š‹ ƒŽ ”‡‰‹‘•ǤǦ ‘ŽǤ‘‘‰”Ǥ͹ͺǣ͸ͷ͵Ȃ͸͹ͳǤ

‡•–”‹ ŠǡǤͳͻͻͲǤ‹‡‹Ž†„‹‡‡ƒ†‡Ǧò”––‡„‡”‰•ǣ†‹‡ ƒ––—‰‡—†”–‡ǤǦǤŽ‡”Ǥ

‡•–”‹ Šǡ Ǥ ͳͻͻ͸Ǥ ƒ„‹–ƒ– ”‡“—‹”‡‡–• ‘ˆ ‡–”ƒŽ —”‘’‡ƒ „‡‡• ƒ† –Š‡ ’”‘„Ž‡• ‘ˆ ’ƒ”–‹ƒŽ Šƒ„‹–ƒ–•Ǥ Ǧ ǣ ƒ–Š‡•‘ǡ Ǥ ‡– ƒŽǤ ȋ‡†•Ȍǡ‹‡ƒ ‘ ‹‡–› ›’‘•‹— ‡”‹‡•Ǥ ‘ŽǤ ͳͺǤ  ƒ†‡‹  ”‡••‹‹–‡†ǡ’’ǤͳȂͳ͸Ǥ

‹ˆ”‡‡ǡǤ‡–ƒŽǤʹͲͲͻǤ‡–ƒǦƒƒŽ›•‹•‘ˆ„‡‡•ǯ”‡•’‘•‡•–‘ƒ–Š”‘’‘‰‡‹ †‹•–—”„ƒ ‡ǤǦ ‘Ž‘‰›ͻͲǣ ʹͲ͸ͺȂʹͲ͹͸Ǥ

 48 Šƒ’–‡”ͳ

ƒ‡‰ƒǡǤͳͻͻͲǤŠ‹Ž‘’ƒ–”›ƒ†‡•–ˆ‘—†‹‰‹ƒ’”‹‹–‹˜‡Ž›•‘ ‹ƒŽ„‡‡ǡHalictusrubicundusǤǦ‡Šƒ˜Ǥ  ‘ŽǤ‘ ‹‘„‹‘ŽǤʹ͹ǣ͵͹ȂͶʹǤ

—”„— Š‡ǡǤ‡–ƒŽǤʹͲͳͲǤƒš‹—ˆ‘”ƒ‰‹‰”ƒ‰‡•‹•‘Ž‹–ƒ”›„‡‡•ǣ‘Ž›ˆ‡™‹†‹˜‹†—ƒŽ•Šƒ˜‡–Š‡ ƒ’ƒ„‹Ž‹–›–‘ ‘˜‡”Ž‘‰ˆ‘”ƒ‰‹‰†‹•–ƒ ‡•ǤǦ‹‘ŽǤ‘•‡”˜ǤͳͶ͵ǣ͸͸ͻȂ͸͹͸Ǥ







 49



   Šƒ’–‡”ʹ

Collateraleffectsofbeekeeping: impactsonpollenǦnectarresources andwildbeecommunities























TornéǦNoguera,A;Rodrigo,A;Osorio,S;Bosch,J.(2015)

 ‡’–‡†—†‡”‹‘””‡˜‹•‹‘•‹BasicandAppliedEcology



Šƒ’–‡”ʹ

Summary—‡–‘–Š‡ ‘–”‹„—–‹‘‘ˆŠ‘‡›„‡‡•ȋ’‹•‡ŽŽ‹ˆ‡”ƒȌ–‘™‹Ž†ˆŽ‘™‡”ƒ† ”‘’ ’‘ŽŽ‹ƒ–‹‘ǡ„‡‡‡‡’‹‰Šƒ•–”ƒ†‹–‹‘ƒŽŽ›„‡‡ ‘•‹†‡”‡†ƒ•—•–ƒ‹ƒ„Ž‡’”ƒ –‹ ‡Ǥ ‘™‡˜‡”ǡ Š‹‰ŠŠ‘‡›„‡‡†‡•‹–‹‡•ƒ›Šƒ˜‡ƒ‹’ƒ –‘Ž‘ ƒŽ’‘ŽŽ‡ƒ†‡ –ƒ”ƒ˜ƒ‹Žƒ„Ž‹–›ǡ™Š‹ Š ‹–—”ƒ›‡‰ƒ–‹˜‡Ž›ƒˆˆ‡ –‘–Š‡”’‘ŽŽ‹ƒ–‘”•ǤŠ‹•‹•‡šƒ ‡”„ƒ–‡†„›–Š‡ƒ„‹Ž‹–›‘ˆŠ‘‡› „‡‡•–‘”‡ ”—‹–ˆ‘”ƒ‰‡”•–‘Š‹‰ŠŽ›”‡™ƒ”†‹‰ˆŽ‘™‡”’ƒ– Š‡•Ǥ‡‡ƒ•—”‡†ˆŽ‘”ƒŽ”‡•‘—” ‡ ‘•—’–‹‘‹”‘•‡ƒ”›ȋ‘•ƒ”‹—•‘ˆˆ‹ ‹ƒŽ‹•Ȍƒ†–Š›‡ȋŠ›—• ˜—Ž‰ƒ”‹•Ȍ ‹ ʹͳ ’Ž‘–•Ž‘ ƒ–‡†ƒ–†‹ˆˆ‡”‡–†‹•–ƒ ‡•ˆ”‘ƒ’‹ƒ”‹‡•‹–Š‡• ”—„Žƒ†‘ˆ ƒ””ƒˆƒ–—”ƒŽƒ” ȋƒ” ‡Ž‘ƒȌǡƒ†”‡Žƒ–‡†–Š‡•‡‡ƒ•—”‡•–‘˜‹•‹–ƒ–‹‘”ƒ–‡•‘ˆŠ‘‡› „‡‡•ǡ „—„Ž‡„‡‡• ȋ‘„—•–‡””‡•–”‹•Ȍƒ†‘–Š‡”’‘ŽŽ‹ƒ–‘”•Ǥ –Š‡•ƒ‡’Ž‘–•ǡ™‡‡ƒ•—”‡†ˆŽ‘™‡”†‡•‹–› ƒ†—•‡†’ƒ–”ƒ’•–‘ Šƒ”ƒ –‡”‹œ‡–Š‡™‹Ž†„‡‡ ‘—‹–›Ǥ Ž‘™‡””‡•‘—” ‡ ‘•—’–‹‘ ™ƒ•Žƒ”‰‡Ž›‡š’Žƒ‹‡†„›Š‘‡›„‡‡˜‹•‹–ƒ–‹‘ƒ†‘Ž›ƒ”‰‹ƒŽŽ› „› „—„Ž‡„‡‡ ˜‹•‹–ƒ–‹‘Ǥˆ–‡”ƒ ‘—–‹‰ˆ‘”ˆŽ‘™‡”†‡•‹–›ǡ’Ž‘–• Ž‘•‡–‘ƒ’‹ƒ”‹‡•Šƒ†Ž‘™‡”™‹Ž†„‡‡ „‹‘ƒ••ǤŠ‹•™ƒ•†—‡–‘ƒŽ‘™‡”ƒ„—†ƒ ‡‘ˆŽƒ”‰‡„‡‡•’‡ ‹‡•ǡ–Š‘•‡‘”‡Ž‹‡Ž›–‘„‡ ƒˆˆ‡ –‡†„›Š‘‡›„‡‡ ‘’‡–‹–‹‘Ǥ‡ ‘ Ž—†‡–Šƒ–Š‘‡›„‡‡•ƒ”‡–Š‡ƒ‹ ‘–”‹„—–‘”• –‘’‘ŽŽ‡Ȁ‡ –ƒ” ‘•—’–‹‘‘ˆ–Š‡–™‘ƒ‹ˆŽ‘™‡”‹‰’Žƒ–•‹–Š‡• ”—„Žƒ†ǡƒ†–Šƒ– ƒ––Š‡†‡•‹–‹‡• —””‡–Ž›‘ —””‹‰‹–Š‡’ƒ”ȋ͵ǤͷŠ‹˜‡•ȀʹȌ–Š‡™‹Ž†„‡‡ ‘—‹–› ‹•„‡‹‰ƒˆˆ‡ –‡†Ǥ—”•–—†›•—’’‘”–•–Š‡Š›’‘–Š‡•‹•–Šƒ–Š‹‰ŠŠ‘‡› „‡‡ †‡•‹–‹‡• ƒ› Šƒ˜‡ƒ‹’ƒ –‘‘–Š‡”’‘ŽŽ‹ƒ–‘”•˜‹ƒ ‘’‡–‹–‹‘ˆ‘”ˆŽ‘™‡””‡•‘—” ‡•Ǥ 

2.1. Introduction

Š‡ ‹–”‘†— –‹‘ ‘ˆ Žƒ”‰‡ ’‘’—Žƒ–‹‘• ‘ˆ Š‹‰ŠŽ› ‘’‡–‹–‹˜‡ •’‡ ‹‡• ‹–‘ ƒ ‡™ ƒ”‡ƒ ƒ› ƒˆˆ‡ – ”‡•‹†‡– ’‘’—Žƒ–‹‘• —Ž–‹ƒ–‡Ž› ”‡•—Ž–‹‰ ‹ Šƒ‰‡• ‹ –Š‡ •–”— –—”‡ ‘ˆ ƒ–‹˜‡ ‘—‹–‹‡• ȋ‡˜‹‡ ‡– ƒŽǤ ʹͲͲ͵ǡ Š”‡ˆ‡Ž† ʹͲͳͲȌǤ Š‹• ƒ› ‘ —” ™Š‡ ‡š‘–‹  •’‡ ‹‡•ǡ ‹–”‘†— ‡† ‡‹–Š‡” ƒ ‹†‡–ƒŽŽ› ‘” ‹–‡–‹‘ƒŽŽ›ǡ –—” ‹˜ƒ•‹˜‡ ƒ† ‘’‡–‡ ˆ‘” Ž‹‹–‡† ”‡•‘—” ‡•™‹–ŠŽ‘ ƒŽ•’‡ ‹‡•‘ —’›‹‰ƒ•‹‹Žƒ”‹ Š‡ȋ‡–”‡ƒ†ƒ•‡ͳͻͻ͸ǡ›‡”•ʹͲͲͲȌǤ ƒ”ƒ†‹‰ƒ–‹  ‡šƒ’Ž‡• ‘ˆ ‡š‘–‹  ƒ‹ƒŽ •’‡ ‹‡• ‘—– ‘’‡–‹‰ Ž‘ ƒŽ•’‡ ‹‡•ƒ”‡–Š‡ ”‰‡–‹‡ ƒ– ȋ —ƒ ƒ† ‘”†‘ ͳͻͻ͸ǡ ‘Ž™ƒ› ͳͻͻͻȌ ƒ† –Š‡ •‹ƒ ƒ”’ȋ ”‘•‡–ƒŽǤ ʹͲͲ͹ǡ ƒ’•‘ ‡– ƒŽǤ ʹͲͲͺȌǤ  ƒ††‹–‹‘ –‘ ‡š‘–‹  •’‡ ‹‡•ǡ †‘‡•–‹ ƒ–‡† •’‡ ‹‡• ƒ› ƒŽ•‘ ƒˆˆ‡ –”‡•‹†‡–•’‡ ‹‡•Ǥ Ž‡ƒ”‡šƒ’Ž‡‹•–Š‡’”‡•‡ ‡‘ˆ ƒ––Ž‡‘”•Š‡‡’‹ƒ–—”ƒŽ‘”•‡‹Ǧ ƒ–—”ƒŽƒ”‡ƒ•ǡ’‘–‡–‹ƒŽŽ› ‘’‡–‹‰™‹–ŠŽƒ”‰‡Š‡”„‹˜‘”‡•ˆ‘”’ƒ•–—”‡ȋ–‡™ƒ”–‡–ƒŽǤʹͲͲʹǡ ‘—‰‡–ƒŽǤʹͲͲͷȌǤ‘”‡‘˜‡”ǡ†‘‡•–‹ ƒ–‡†ƒ‹ƒŽ•„‡‡ˆ‹–ˆ”‘ Š—ƒ ƒ••‹•–ƒ ‡ǡ ‹ Ž—†‹‰’”‘–‡ –‹‘ƒ‰ƒ‹•–’”‡†ƒ–‘”•ƒ†˜‡–‡”‹ƒ”› ƒ”‡Ǥ

‘‰†‘‡•–‹ ƒ–‡†ƒ‹ƒŽ•ǡ–Š‡—”‘’‡ƒŠ‘‡›„‡‡ȋApismelliferaȌ‹•—†‘—„–‡†Ž›‘‡ ‘ˆ–Š‡‘•–‰Ž‘„ƒŽŽ›•’”‡ƒ†Ǥƒ–‹˜‡–‘—”ƒ•‹ƒƒ†ˆ”‹ ƒ‹–Šƒ• „‡‡ ‹–”‘†— ‡† ‹–‘ ƒŽŽ

 53 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

‘–‹‡–• ‡š ‡’– –ƒ” –‹ ƒ ˆ‘” –Š‡ ’”‘†— –‹‘ ‘ˆ Š‘‡› ƒ† ‘–Š‡” Š‹˜‡ ’”‘†— –• ȋ”ƒ‡ ͳͻͻͲȌǤ˜‡‹ˆ”‡ ‡–•–—†‹‡•Šƒ˜‡•Š‘™™‹Ž†’‘ŽŽ‹ƒ–‘”•–‘„‡‘”‡‡ˆˆ‡ –‹˜‡’‘ŽŽ‹ƒ–‘”• –Šƒ Š‘‡› „‡‡• ȋ ƒ”‹„ƒŽ†‹ ‡– ƒŽǤ ʹͲͳ͵Ȍǡ Š‘‡› „‡‡• ƒ”‡ ”‘—–‹‡Ž›”‡’‘”–‡†–‘’”‘˜‹†‡ƒ ‹’‘”–ƒ–‡ ‘•›•–‡•‡”˜‹ ‡‹–‡”•‘ˆ™‹Ž†ˆŽ‘™‡”ƒ† ”‘’’‘ŽŽ‹ƒ–‹‘ǤŠ—•ǡ„‡‡‡‡’‹‰ Šƒ•–”ƒ†‹–‹‘ƒŽŽ›„‡‡ ‘•‹†‡”‡†ƒ„‡‡ˆ‹ ‹ƒŽ’”ƒ –‹ ‡ǡƒ†‹–••—•–ƒ‹ƒ„‹Ž‹–›Šƒ•„‡‡–ƒ‡ ˆ‘” ‰”ƒ–‡†Ǥ Š‹• ‹• ”‡ˆŽ‡ –‡† ‹ –Š‡ —””‡– Žƒ  ‘ˆ •’‡ ‹ˆ‹  Ž‡‰‹•Žƒ–‹‘ ‹ ‘•– ‘—–”‹‡• ™‘”Ž†™‹†‡ǡ ™Š‡”‡„› „‡‡‡‡’‹‰ ‹• ‘•‹†‡”‡† –‘ „‡ „‡‡ˆ‹ ‹ƒŽ ƒ† ‹• —•—ƒŽŽ› ƒŽŽ‘™‡† ‹ ƒ–—”‡”‡•‡”˜‡•ƒ†‘–Š‡”–›’‡•‘ˆ’”‘–‡ –‡†ƒ”‡ƒ•ǡ‹ Ž—†‹‰•‘‡ƒ–‹‘ƒŽƒ”•Ǥ ƒ› ƒ•‡•ǡ „‡‡‡‡’‹‰ ‹ –Š‡•‡ ƒ”‡ƒ• ‹• ‘– ‘Ž› ƒŽŽ‘™‡† „—– ‡˜‡ ’”‘‘–‡†ƒ•ƒ–”ƒ†‹–‹‘ƒŽǡ •—•–ƒ‹ƒ„Ž‡ƒ –‹˜‹–›ȋ‹ˆ‘”ƒ–‹‘‘„–ƒ‹‡†ˆ”‘ƒ–—”ƒŽ’ƒ”ƒ†™‹Ž†Ž‹ˆ‡ƒƒ‰‡”•ˆ”‘ͺ —”‘’‡ƒ ‘—–”‹‡•ǡ•‡‡ƒ ‘™Ž‡†‰‡‡–•ȌǤ –‹•–Š‡”‡ˆ‘”‡‘–•—”’”‹•‹‰–Šƒ–A.mellifera ‹• ”‘—–‹‡Ž› ”‡’‘”–‡† ƒ• ‘‡ ‘ˆ –Š‡ †‘‹ƒ– •’‡ ‹‡• ‹ ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘” ‡–™‘”• ™‘”Ž†™‹†‡ǡ‡˜‡‹•–—†‹‡• ‘†— –‡†‹ƒ–—”ƒŽŠƒ„‹–ƒ–•ȋ‡Ǥ‰Ǥ ‘”—’‡–ƒŽǤʹͲͲͺǡ‘• Š‡–ƒŽǤ ʹͲͲͻǡƒ‹•‡”Ǧ—„—”›‡–ƒŽǤʹͲͲͻǡƒŽ‹†‘‡–ƒŽǤʹͲͳͶǢ•‡‡ƒ˜‹ŽƒƬƒ”†Ž‡ʹͲͲͺˆ‘”ƒ”ƒ”‡ ‡š ‡’–‹‘ȌǤ ‘™‡˜‡”ǡƒ•‹‘–Š‡”•‹†•‘ˆƒ‹ƒŽŠ—•„ƒ†”›ǡŽƒ”‰‡ƒ’‹ƒ”‹‡•”‡•—Ž–‹‰‹Š‹‰Š †‡•‹–‹‡•‘ˆˆ‘”ƒ‰‡”•ƒ›Šƒ˜‡ƒ‹’ƒ –‘Ž‘ ƒŽˆ‘‘†”‡•‘—” ‡•ȋ’‘ŽŽ‡ƒ†‡ –ƒ”‹–Š‹• ƒ•‡Ȍǡ ™Š‹ Š ‹ –—” ƒ› ‡‰ƒ–‹˜‡Ž› ƒˆˆ‡ – ‘–Š‡” ˆŽ‘™‡”Ǧ˜‹•‹–‹‰ ‹•‡ –•Ǥ ‘‡› „‡‡• Šƒ˜‡ Š‹‰Š‡‡”‰‡–‹ ”‡“—‹”‡‡–•„‡ ƒ—•‡–Š‡›Ž‹˜‡‹Žƒ”‰‡ ‘Ž‘‹‡• ‘’”‹•‹‰ –‡• ‘ˆ –Š‘—•ƒ†•‘ˆ‹†‹˜‹†—ƒŽ•ƒ†„‡ ƒ—•‡–Š‡›ƒ‹–ƒ‹‡Ž‡˜ƒ–‡†Š‹˜‡–‡’‡”ƒ–—”‡•‡˜‡†—”‹‰ –Š‡ ™‹–‡” ȋ‡‡Ž‡› ͳͻͺͷȌǤ —‡ –‘ –Š‹• Š‹‰Š ‡‡”‰‡–‹  ”‡“—‹”‡‡–•ǡ –Š‡‹” ˆ‘”ƒ‰‹‰ ”ƒ‰‡• •’ƒ•‡˜‡”ƒŽ‹Ž‘‡–”‡•ȋ‹•• Š‡”ƒ†‡‡Ž‡›ͳͻͺʹȌǤ ƒ††‹–‹‘ǡŠ‘‡›„‡‡•Šƒ˜‡–Š‡ƒ„‹Ž‹–› ȋ—‹“—‡–‘–Š‡ƒ†•‘‡•–‹‰Ž‡••„‡‡•Ȍ–‘ ‘—‹ ƒ–‡–Š‡Ž‘ ƒ–‹‘‘ˆˆŽ‘™‡””‡•‘—” ‡• –‘ ‡•– ƒ–‡•ǡ –Š—• ‘ ‡–”ƒ–‹‰ Žƒ”‰‡ —„‡”• ‘ˆ ˆ‘”ƒ‰‡”• ‹ Š‹‰ŠŽ›”‡™ƒ”†‹‰’ƒ– Š‡• ȋ˜‘ ”‹• Šͳͻ͸͹ȌǤŠ—•ǡŠ‘‡›„‡‡•ƒ”‡Š‹‰ŠŽ›‡ˆˆ‹ ‹‡–’‘ŽŽ‡Ǧ‡ –ƒ”ˆ‘”ƒ‰‡”•ƒ†ǡ™Š‡ ’”‡•‡– ‹ Žƒ”‰‡ †‡•‹–‹‡•ǡ ƒ› ’‘–‡–‹ƒŽŽ› ”‡ƒ–‡ ƒ ‘’‡–‹–‹‘ • ‡ƒ”‹‘ ™‹–Š ‘–Š‡” ’‘ŽŽ‹ƒ–‘”•Ǥ

‘’‡–‹–‹‘ƒ›–ƒ‡’Žƒ ‡–Š”‘—‰Š‹–‡”ˆ‡”‡ ‡‘”–Š”‘—‰Š”‡•‘—” ‡‡š’Ž‘‹–ƒ–‹‘ȋ‹Žƒ ͳͻͺʹȌǤ –‡”ˆ‡”‡ ‡ ‘’‡–‹–‹‘ ‘ —”• †‹”‡ –Ž› „‡–™‡‡ ‹†‹˜‹†—ƒŽ• –Š”‘—‰Š ƒ‰‰”‡••‹˜‡ ‡ ‘—–‡”•ȋ‡Ǥ‰ǤǡŠ‘‡›„‡‡• Šƒ•‹‰‘–Š‡”’‘ŽŽ‹ƒ–‘”•‘—–‘ˆƒˆŽ‘™‡”‘”ˆŽ‘™‡”’ƒ– ŠȌǤ— Š ƒ‰‰”‡••‹˜‡‹–‡”ƒ –‹‘•Šƒ˜‡•‘‡–‹‡•„‡‡‘„•‡”˜‡†ȋ‡Ǥ‰Ǥ‹—•Ǧ‡†‘‡–ƒŽǤʹͲͲͷȌǡ„—– –Š‡ˆƒ ––Šƒ–‘•–•–—†‹‡•†‘‘–”‡’‘”–ƒ‰‰”‡••‹˜‡‡ ‘—–‡”•‹†‹ ƒ–‡•–Šƒ––Š‡›ƒ”‡‘– ‘‘ȋ‡Ǥ‰Ǥ‘—„‹ͳͻ͹ͺǡ —†‡™‡œƒ†Ž‡‹ʹͲͳ͵ȌǤˆ–‡”•‡˜‡”ƒŽ›‡ƒ”•‘ˆˆ‹‡Ž†™‘”ǡ™‡ ƒƒ••‡”––Šƒ–•— Š‹–‡”ƒ –‹‘•ƒ”‡˜‡”›”ƒ”‡‹‘—”•–—†›ƒ”‡ƒǤš’Ž‘‹–ƒ–‹˜‡ ‘’‡–‹–‹‘  54 Šƒ’–‡”ʹ

‘ —”•‹†‹”‡ –Ž›„‡–™‡‡‹†‹˜‹†—ƒŽ•–Š”‘—‰ŠƒŽ‹‹–‹‰”‡•‘—” ‡•— Šƒ•ˆ‘‘†‘”‡•–‹‰ •‹–‡•Ǥ ‘’‡–‹–‹‘ ˆ‘” ‡•–‹‰ ”‡•‘—” ‡• ƒ „‡ ”—Ž‡† ‘—– „‡ ƒ—•‡ ™‹Ž† „‡‡• ‹ –‡’‡”ƒ–‡ œ‘‡•†‘‘–‡•–‹–Š‡‹†‘ˆŽƒ”‰‡ ƒ˜‹–‹‡•—•‡†„›Š‘‡›„‡‡•ǡƒ†„‡ ƒ—•‡ƒƒ‰‡† ˆ‡”ƒŽ ‘Ž‘‹‡• ƒ”‡ ˜‡”› ”ƒ”‡ ‹ ‘—” •–—†› ƒ”‡ƒǡ ƒ• ‹ ‘•– ‘ˆ —”‘’‡ ȋ ƒˆˆ± ‡– ƒŽǤ ʹͲͲͻȌǤ ‘’‡–‹–‹‘ ˆ‘” ˆŽ‘™‡” ”‡•‘—” ‡• ‹• — Š ‘”‡ Ž‹‡Ž› –‘ ‘ —” „‡ ƒ—•‡ Š‘‡› „‡‡• ƒ”‡ Š‹‰ŠŽ›‰‡‡”ƒŽ‹•–‹’‘ŽŽ‡ƒ†‡ –ƒ”—•‡ǡƒ†–Š‡‹”†‹‡–™‹†‡Ž›‘˜‡”Žƒ’•™‹–Š–Šƒ–‘ˆ‘–Š‡” ˆŽ‘™‡”Ǧ˜‹•‹–‹‰•’‡ ‹‡•Ǥ

ƒ”‹‘—• •–—†‹‡• Šƒ˜‡ ‡š’Ž‘”‡† ’‘–‡–‹ƒŽ ƒ†˜‡”•‡ ‡ˆˆ‡ –• ‘ˆ Š‘‡› „‡‡• ‘ Ž‘ ƒŽ ’‘ŽŽ‹ƒ–‘” ‘—‹–‹‡•Ǥ ‘™‡˜‡”ǡ–‘†‡‘•–”ƒ–‡ƒ ‘’‡–‹–‹‘• ‡ƒ”‹‘‹–‹•‡š–”‡‡Ž›†‹ˆˆ‹ —Ž–ǡ‘ ƒ ‘—– ‘ˆ –Š‡ Žƒ”‰‡ ˆ‘”ƒ‰‹‰ ”ƒ‰‡• ‘ˆ Š‘‡› „‡‡• ȋ•‡˜‡”ƒŽ Ȍ ȋ‡‡Ž‡› ͳͻͺͷǡ ‘—Ž•‘ ʹͲͲ͵Ȍǡ ‘„‹‡†™‹–Š–Š‡‹”ƒ„‹Ž‹–›–‘ ‘—‹ ƒ–‡–Š‡Ž‘ ƒ–‹‘‘ˆ”‹ ŠˆŽ‘™‡”’ƒ– Š‡•ǡ–Š—• ƒŽŽ‘™‹‰ ‘Ž‘‹‡•–‘ƒ†Œ—•––Š‡‹”ˆ‘”ƒ‰‹‰ƒ”‡ƒ•ƒ†ˆŽ‘™‡” Š‘‹ ‡•ƒ•’‘ŽŽ‡Ǧ‡ –ƒ”•–ƒ†‹‰ ”‘’•˜ƒ”›–Š”‘—‰Š–‹‡ȋ‹•• Š‡”ƒ†‡‡Ž‡›ͳͻͺʹȌǤ

‘” –Š‹• ”‡ƒ•‘ǡ ‘•– •–—†‹‡• Šƒ˜‡ •‘ ˆƒ” ˆ‘ —•‡† ‘ ‹†‹”‡ – ‡˜‹†‡ ‡• ‘ˆ ‘’‡–‹–‹‘ „‡–™‡‡ Š‘‡› „‡‡• ƒ† ™‹Ž† „‡‡•ǡ •— Š ƒ• ”‡•‘—” ‡ ‘˜‡”Žƒ’ ȋ–‡ˆˆƒǦ‡™‡–‡” ƒ† • Šƒ”–‡ʹͲͲͲȌǡƒ†‹ Šƒ‰‡•‹’‘ŽŽ‡Ǧ‡ –ƒ””‡•‘—” ‡—•‡ȋ ‘”—’ƒ†‡‘––ʹͲͲͷǡ ƒŽ‹†‘ ‡– ƒŽǤ ʹͲͳͶȌǡ ˆ‘”ƒ‰‹‰ ƒ –‹˜‹–› ȋŠ‘•‘ ʹͲͲͶȌ ƒ† ˜‹•‹–ƒ–‹‘ ”ƒ–‡• ȋ‘—„‹ ͳͻ͹ͺǡ —†‡™‡œ ƒ† Ž‡‹ ʹͲͳ͵Ȍ ‘ˆ ™‹Ž† ’‘ŽŽ‹ƒ–‘”• ‘ˆ”‘–‡† ™‹–Š †‹ˆˆ‡”‡–Š‘‡›„‡‡ • ‡ƒ”‹‘•Ǥ–Š‡”•–—†‹‡•Šƒ˜‡‡ƒ•—”‡† Šƒ‰‡•‹’‘’—Žƒ–‹‘ƒ„—†ƒ ‡ƒ†”‹ Š‡••‘ˆ ™‹Ž† „‡‡• —†‡” †‹ˆˆ‡”‡– Š‘‡› „‡‡ †‡•‹–‹‡• ȋ‘—„‹ ͳͻ͹ͺǡ –‡ˆˆƒǦ‡™‡–‡” ƒ† • Šƒ”–‡ʹͲͲͲǡ‘—„‹ƒ†‘Ž†ƒʹͲͲͳǡ ‘”—’ƒ†‡‘––ʹͲͲͷȌǤ ‡™‡”•–—†‹‡•Šƒ˜‡ Ž‘‘‡† ˆ‘” ‘”‡ †‹”‡ – ‡˜‹†‡ ‡ ‘ˆ ‘’‡–‹–‹‘ǡ •— Š ƒ• Šƒ‰‡• ‹ ”‡’”‘†— –‹˜‡ •— ‡•• ȋ–‡ˆˆƒǦ‡™‡–‡” ƒ† • Šƒ”–‡ ʹͲͲͲǡ Š‘•‘ ʹͲͲͶǡ ‘—Ž•‘ ƒ† ’ƒ””‘™ ʹͲͲͻǡ Ž„‰ƒ‹‡–ƒŽǤʹͲͳͶȌǡƒ†–Š‡‘—– ‘‡•‘ˆ–Š‡•‡•–—†‹‡•ƒ”‡‘– ‘•‹•–‡–Ǥ‘‡Šƒ˜‡ˆ‘—† ‡‰ƒ–‹˜‡ ‡ˆˆ‡ –• ‘ˆ Š‘‡› „‡‡• ȋŠ‘•‘ ʹͲͲͶǡ ‘—Ž•‘ ƒ† ’ƒ””‘™ ʹͲͲͻȌ ™Š‹Ž‡ ‘–Š‡”• Šƒ˜‡‘–ȋ–‡ˆˆƒǦ‡™‡–‡”ƒ†• Šƒ”–‡ʹͲͲͲǡ‘—„‹ƒ†‘Ž†ƒʹͲͲͳȌǤ

‘” ‡š’Ž‘‹–ƒ–‹˜‡ ‘’‡–‹–‹‘ –‘ ‘ —”ǡ ˆŽ‘”ƒŽ ”‡•‘—” ‡• •Š‘—Ž† „‡ Ž‹‹–‹‰Ǥ —”’”‹•‹‰Ž›ǡ Š‘™‡˜‡”ǡ‘•–—†›Šƒ•Š‹–Š‡”–‘‡ƒ•—”‡†–Š‡‡ˆˆ‡ –•‘ˆŠ‘‡›„‡‡ƒ„—†ƒ ‡‘’‘ŽŽ‡ƒ† ‡ –ƒ”ƒ˜ƒ‹Žƒ„‹Ž‹–›ǤŠ‹•‹•‹’‘”–ƒ–„‡ ƒ—•‡™‡ —””‡–Ž›†‘‘–‘™–Š‡ƒ‰‹–—†‡‘ˆ–Š‡ ‹’ƒ –‘ˆŠ‘‡›„‡‡•‘ˆŽ‘™‡””‡•‘—” ‡• ‘’ƒ”‡†–‘”‡•‹†‡–’‘ŽŽ‹ƒ–‘”•Ǥ –Š‹••–—†›™‡ ƒ††”‡••–Š‡’‘–‡–‹ƒŽ‡ˆˆ‡ –•‘ˆ„‡‡‡‡’‹‰‘™‹Ž†„‡‡ ‘—‹–‹‡•‹ƒ‡˜‹”‘‡–ƒŽŽ›

 55 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

’”‘–‡ –‡†ƒ–—”ƒŽƒ”‡ƒǤ—”‘„Œ‡ –‹˜‡‹•–‘•–—†›–Š‡‹’ƒ –‘ˆŠ‘‡›„‡‡ˆŽ‘™‡”˜‹•‹–ƒ–‹‘‘ ’‘ŽŽ‡ƒ†‡ –ƒ” ‘•—’–‹‘ƒ†–Š‡‡ˆˆ‡ –‘ˆ„‡‡‡‡’‹‰‘–Š‡ƒ„—†ƒ ‡ǡ”‹ Š‡••ƒ† ‘’‘•‹–‹‘‘ˆ–Š‡Ž‘ ƒŽ™‹Ž†„‡‡ ‘—‹–›Ǥ‡ ƒ—•‡Š‘‡›„‡‡•ƒ”‡˜‡”›ƒ„—†ƒ–ƒ† ‰‹˜‡ –Š‡‹” ƒ„‹Ž‹–› –‘ ”‡ ”—‹– Žƒ”‰‡ —„‡”• ‘ˆ ˆ‘”ƒ‰‡”• –‘ –Š‡ ‘•– ”‡™ƒ”†‹‰ ˆŽ‘™‡” ’ƒ– Š‡•ǡ™‡Šƒ˜‡–Š”‡‡Š›’‘–Š‡•‡•ǣȋͳȌ ‘‡›„‡‡•™‹ŽŽ„‡–Š‡ƒ‹ ‘–”‹„—–‘”•–‘ˆŽ‘™‡” ”‡•‘—” ‡†‡’Ž‡–‹‘Ǥ‡–Š‡”‡ˆ‘”‡‡š’‡ –’‘ŽŽ‡ƒ†‡ –ƒ”ƒ˜ƒ‹Žƒ„‹Ž‹–›–‘‘–Š‡”’‘ŽŽ‹ƒ–‘”•–‘ „‡ Ž‘™‡” ‹ ƒ”‡ƒ• Ž‘•‡ –‘ ƒ’‹ƒ”‹‡•Ǣ ȋʹȌ Š‡ •–”— –—”‡ ‘ˆ –Š‡ ™‹Ž† „‡‡ ‘—‹–› ™‹ŽŽ „‡ ‘†‹ˆ‹‡† „› Š‹‰Š Š‘‡› „‡‡ †‡•‹–‹‡•Ǥ ‡ ‡š’‡ – ™‹Ž† „‡‡ ”‹ Š‡•• ƒ† ƒ„—†ƒ ‡ –‘ „‡ Ž‘™‡” Ž‘•‡–‘ƒ’‹ƒ”‹‡•Ǣȋ͵Ȍ‘‰™‹Ž†„‡‡•ǡ™‡‡š’‡ –Žƒ”‰‡•’‡ ‹‡•ȋ™‹–ŠŠ‹‰Š‡”ˆ‡‡†‹‰ ”‡“—‹”‡‡–•ǢòŽŽ‡”‡–ƒŽǤǡʹͲͲ͸Ȍǡ–‘„‡‘•–ƒˆˆ‡ –‡†Ǥ



2.2. Materialsandmethods

2.2.1. Studyarea

Š‹••–—†›™ƒ• ‘†— –‡†‹–Š‡ƒ–—”ƒŽƒ”‘ˆ‡Ž ƒ””ƒˆȋƒ” ‡Ž‘ƒǡƒ–ƒŽ‘‹ƒǡ’ƒ‹Ȍǡƒ ‡†‹–‡””ƒ‡ƒ • ”—„Žƒ† †‘‹ƒ–‡† „› Quercus cocciferaǡ Pistacia lentiscusǡ Rosmarinus officinalisƒ†ThymusvulgarisǤŠ‡ƒ–—”ƒŽƒ”‘ˆ‡Ž ƒ””ƒˆ‹• Žƒ••‹ˆ‹‡†ƒ• ƒ–‡‰‘”›‘ˆ–Š‡ –‡”ƒ–‹‘ƒŽ‹‘ˆ‘”‘•‡”˜ƒ–‹‘‘ˆƒ–—”‡ȋ Ȍȋ—†Ž‡›ʹͲͲͺȌǡ™Š‹ Š‹ Ž—†‡•–Š‡ ƒŒ‘”‹–›ȋ͸ʹΨȌ‘ˆ–Š‡‡˜‹”‘‡–ƒŽŽ›’”‘–‡ –‡†Žƒ†‹–Š‡‡†‹–‡””ƒ‡ƒ”‡‰‹‘ȋג‡œ ”ƒ–‡–ƒŽǤʹͲͲ͹ȌǤƒ–‡‰‘”›’ƒ”•ƒ”‡†‡ˆ‹‡†ƒ•’”‘–‡ –‡†ƒ”‡ƒ• ™‹–Š ƒ ‹’‘”–ƒ– „‹‘Ž‘‰‹ ƒŽǡ ‡ ‘Ž‘‰‹ ƒŽǡ —Ž–—”ƒŽ ƒ† ’‹ –—”‡•“—‡ ˜ƒŽ—‡ „ƒ•‡† ‘ –Š‡ ‹–‡”ƒ –‹‘ „‡–™‡‡ Š—ƒ ’‘’—Žƒ–‹‘• ƒ† –Š‡ ‡˜‹”‘‡– ˜‹ƒ –”ƒ†‹–‹‘ƒŽ ƒƒ‰‡‡– ’”ƒ –‹ ‡•Ǥ Š—•ǡ –Š‡ ’”‘–‡ –‹‘ ‘ˆ •— Š ’‡‘’Ž‡Ǧƒ–—”‡ ‹–‡”ƒ –‹‘• ‹• –Š‡ ƒ‹ ‘•‡”˜ƒ–‹‘ ‘„Œ‡ –‹˜‡ ȋ—†Ž‡› ʹͲͲͺȌǤ ƒ–ƒŽ‘‹ƒǡ —””‡–’‘Ž‹ ‹‡•”‡‰—Žƒ–‹‰‡˜‹”‘‡–ƒŽ‹’ƒ –•‘ˆŠ—ƒƒ –‹˜‹–‹‡• †‘‘–‡–‹‘„‡‡‡‡’‹‰ȋƒ™ʹͲȀʹͲͲͻǡ ͷͷʹͶȌǤƒ–Š‡”ǡ„‡‡‡‡’‹‰‹• ‘•‹†‡”‡† ƒ‹‘ —‘—•ƒ –‹˜‹–›ƒ†A. melifera ‹• †‡ Žƒ”‡† ƒ Dz•’‡ ‹‡• ‘ˆ •’‡ ‹ƒŽ ‹–‡”‡•–dz ȋ‡ ”‡‡ ͳͳͲȀʹͲͲ͵ǡ ͵ͺ͹ͲȌǤ

—”•–—†›ƒ”‡ƒ‹•‡–‹”‡Ž›Ž‘ ƒ–‡†‹–Š‡’ƒ”ǡ‡ ‘’ƒ••‹‰ƒ•—”ˆƒ ‡‘ˆ͵ʹʹǤ‡•‡Ž‡ –‡† ʹͳ ’Ž‘–• ‘ˆ ͶͲ  š ͶͲ  †‹•–”‹„—–‡† ”‡‰—Žƒ”Ž› ƒ ”‘•• –Š‡ •–—†› ƒ”‡ƒǤ ‹•–ƒ ‡• „‡–™‡‡ ‡ƒ”‡•– ’Ž‘–• ”ƒ‰‡† ˆ”‘ ͷͺͷ –‘ ͳ͵ͷͶ Ǥ ƒ•‡†  ‘ –Š‡ ‹ˆ‘”ƒ–‹‘ ’”‘˜‹†‡† „› –Š‡ —–‘‘‘—• ‘˜‡”‡–ƒ†•—„•‡“—‡–Ž›˜‡”‹ˆ‹‡†insituǡ™‡’‹’‘‹–‡†ʹͳƒ’‹ƒ”‹‡• Ž‘•‡  56 Šƒ’–‡”ʹ

–‘–Š‡•–—†›ƒ”‡ƒˆ‘”ƒ–‘–ƒŽ‘ˆͶ͹ͷŠ‹˜‡•Ǥ‹‹—†‹•–ƒ ‡‘ˆ‘—”’Ž‘–•–‘–Š‡‡ƒ”‡•– ƒ’‹ƒ”›”ƒ‰‡†ˆ”‘ʹ͸ʹ–‘ͷͳʹʹǤ



2.2.2. Flowerresourcesurveys

‘•–—†›ˆŽ‘™‡””‡•‘—” ‡ ‘•—’–‹‘ǡ™‡™‘”‡†‘”‘•‡ƒ”›ȋRosmarinusofficinalisȌƒ† –Š›‡ȋThymusvulgarisȌǤŠ‡•‡–™‘•’‡ ‹‡•ƒ”‡ǡ„›ˆƒ”ǡ–Š‡‘•–ƒ„—†ƒ–‡–‘‘’Š‹Ž‘—• •’‡ ‹‡•‹–Š‡•–—†›ƒ”‡ƒǡ’”‘†— ‹‰͹ͲǦͻͲΨ‘ˆ–Š‡ˆŽ‘™‡”•‹–Š‡• ”—„Žƒ†ȋ‘• Š‡–ƒŽǤ •—„‹––‡†ǡ Ž‘ ‡– ƒŽǤ •—„‹––‡†Ǣ ƒ† —’—„Ž‹•Š‡† †ƒ–ƒ ˆ”‘ ͳʹ •‹–‡• ™‹–Š‹ –Š‡ ’ƒ”ȌǤ  ƒ††‹–‹‘ǡ –Š‡ –™‘ •’‡ ‹‡• ƒ”‡ ˜‡”› ƒ––”ƒ –‹˜‡ –‘ Š‘‡› „‡‡• ƒ†ƒ”‡ ‘•‹†‡”‡†Š‹‰ŠŽ› †‡•‹”ƒ„Ž‡ˆ‘”Š‘‡›’”‘†— –‹‘ȋƒ„”ƒʹͲͲͺȌǤŽŽ•—”˜‡›•™‡”‡ ‘†— –‡†‹ʹͲͳͳ—†‡” ˆƒ‹”™‡ƒ–Š‡”Ǥ



Pollen

‘•‡ƒ”›’‘ŽŽ‡•—”˜‡›•™‡”‡ ‘†— –‡†‹ƒ” Šǡ™Š‡–Š‡•’‡ ‹‡•™ƒ•‹ˆ—ŽŽ„Ž‘‘Ǥ  ‡ƒ Š’Ž‘–ǡ™‡•‡Ž‡ –‡†„‡–™‡‡ʹͲƒ†͵Ͳ’Žƒ–•‘™Š‹ Š™‡ƒ”‡†ͺ”‡ ‡–Ž›Ǧ‘’‡‡† ˆŽ‘™‡”• †‹•–”‹„—–‡† ”ƒ†‘Ž› ™‹–Š‹ –Š‡ ’Žƒ– ȋ™‹–Š ˆ”‡•Šǡ ˆ—ŽŽ› ’‘ŽŽ‡ǦŽ‘ƒ†‡† •–ƒ‡•ȌǤ ‡ˆ‘”‡–Š‡‘•‡–‘ˆ’‘ŽŽ‹ƒ–‘”ƒ –‹˜‹–›ȋͻǣͲͲȌǡ™‡ ‘ŽŽ‡ –‡†Ͷƒ”‡†ˆŽ‘™‡”•’‡”’Žƒ–ǡƒ† •–‘”‡†–Š‡–‘‰‡–Š‡”‹ƒ˜‹ƒŽˆ‹ŽŽ‡†™‹–Š‡–Šƒ‘Ž͹ͲΨǤˆ–‡”ͳͺǣͲͲǡ™Š‡ˆ‘”ƒ‰‹‰ƒ –‹˜‹–› Šƒ† ‡ƒ•‡†ǡ –Š‡ ”‡ƒ‹‹‰ Ͷ ˆŽ‘™‡”• ™‡”‡ ‘ŽŽ‡ –‡† ƒ† ’”‡•‡”˜‡† ˆ‘ŽŽ‘™‹‰ –Š‡ •ƒ‡ ’”‘ ‡†—”‡Ǥ

Š›‡ ’‘ŽŽ‡ •—”˜‡›• ™‡”‡ ‘†— –‡† ‹ ’”‹Žǡ †—”‹‰ ’‡ƒ „Ž‘‘ ‘ˆ –Š‹• •’‡ ‹‡•Ǥ ‡ •‡Ž‡ –‡†„‡–™‡‡ʹͲƒ†͵Ͳ–Š›‡’Žƒ–•’‡”’Ž‘–ƒ†ƒ”‡†Ͷ”‡ ‡–Ž›Ǧ‘’‡‡†ˆŽ‘™‡”•‹ ‡ƒ Š‘ˆ–Š‡ǡˆ‘ŽŽ‘™‹‰–Š‡•ƒ‡ ”‹–‡”‹ƒƒ•ˆ‘””‘•‡ƒ”›Ǥ‡ˆ‘”‡ͻǣͲͲ™‡ ‘ŽŽ‡ –‡†–Š‡–™‘ •–ƒ‡•‘ˆ‘‡•‹†‡ȋŽ‡ˆ–‘””‹‰Š–Ȍ‘ˆ‡ƒ ŠˆŽ‘™‡”ƒ†•–‘”‡†–Š‡–‘‰‡–Š‡”‹ƒ˜‹ƒŽˆ‹ŽŽ‡† ™‹–Š‡–Šƒ‘Ž͹ͲΨǤˆ–‡”ͳͺǣͲͲ™‡ ‘ŽŽ‡ –‡†–Š‡–™‘”‡ƒ‹‹‰•–ƒ‡•‘ˆ‡ƒ ŠˆŽ‘™‡”Ǥ

–Š‡Žƒ„‘”ƒ–‘”›ǡ˜‹ƒŽ•™‹–Š•–ƒ‡•™‡”‡•‘‹ ƒ–‡†ˆ‘”ͳͲ‹—–‡•‹ƒ—Ž–”ƒ•‘‹ „ƒ–Š–‘ †‹•Ž‘†‰‡’‘ŽŽ‡‰”ƒ‹•ˆ”‘–Š‡ƒ–Š‡”•Ǥˆ–‡”™ƒ”†•ǡ‡ƒ Šƒ–Š‡”™ƒ•‹•’‡ –‡†—†‡”–Š‡ •–‡”‡‘‹ ”‘• ‘’‡ ƒ† ’‘ŽŽ‡ ‰”ƒ‹• •–‹ŽŽ ƒ†Š‡”‹‰ –‘ –Š‡ ƒ–Š‡”• ™‡”‡ ƒ—ƒŽŽ› †‡–ƒ Š‡†

 57 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

™‹–Š–Š‡ƒ‹†‘ˆƒ‹•‡ –’‹Ǥƒ–‡”ǡ™‡–‘‘ͺ†”‘’•‘ˆʹǤͷɊŽ ‘ˆ –Š‡ ”‡•—Ž–‹‰ ’‘ŽŽ‡ •—•’‡•‹‘ ƒ† ‘—–‡† –Š‡ —„‡” ‘ˆ ’‘ŽŽ‡ ‰”ƒ‹• —†‡” ƒ •–‡”‡‘‹ ”‘• ‘’‡Ǥ ”‡˜‹‘—• –”‹ƒŽ••Š‘™‡†–Šƒ––Š‡—„‡”‘ˆ†”‘’•‡ ‡••ƒ”›–‘•–ƒ„‹Ž‹œ‡’‘ŽŽ‡ ‘—–•™ƒ•͸Ǥ‡–Š‡ ‡ƒ•—”‡†–Š‡”‡ƒ‹‹‰‡–Šƒ‘Ž˜‘Ž—‡‹–Š‡˜‹ƒŽǡƒ†‡•–‹ƒ–‡† –Š‡ –‘–ƒŽ —„‡” ‘ˆ ’‘ŽŽ‡‰”ƒ‹•‹‡ƒ Š•ƒ’Ž‡Ǥ ”‘–Š‡•‡†ƒ–ƒǡ™‡‡•–‹ƒ–‡†–Š‡—„‡”‘ˆ’‘ŽŽ‡‰”ƒ‹•’‡” ˆŽ‘™‡”‹–Š‡‘”‹‰ƒ†‹–Š‡‡˜‡‹‰ǡ™Š‹ Š™‡—•‡†–‘ ƒŽ —Žƒ–‡’‘ŽŽ‡ ‘•—’–‹‘Ǥ ˜‡”ƒŽŽǡ™‡•ƒ’Ž‡†ͶͲͲͷ”‘•‡ƒ”›ˆŽ‘™‡”•ƒ†ʹ͵͸͸–Š›‡ˆŽ‘™‡”•Ǥ



Nectar

‡ –ƒ” ‘•—’–‹‘ ‹• †‹ˆˆ‹ —Ž– –‘ ‡ƒ•—”‡ „‡ ƒ—•‡ ‡ –ƒ” •‡ ”‡–‹‘ ‹• ƒ ‘”‡ ‘” Ž‡•• ‘–‹—‘—• ’”‘ ‡•• ȋƒ ‹‹ ‡– ƒŽǤ ʹͲͲ͵Ȍǡ •‘ –Šƒ– ‘•—’–‹‘ ƒ› „‡ ‘’‡•ƒ–‡† „› •—„•‡“—‡– •‡ ”‡–‹‘Ǥ  •‘‡ ƒ•‡•ǡ •‡ ”‡–‹‘ ƒ› „‡ ‡˜‡ •–‹—Žƒ–‡† „› ‘•—’–‹‘ ȋƒ•–‡ŽŽƒ‘• ‡– ƒŽǤ ʹͲͲʹǡ ”†ƒ‘ ƒ† ”‡Žƒ• ʹͲͲͶȌǤ  ƒ††‹–‹‘ǡ ‡ –ƒ” •‡ ”‡–‹‘ ƒ „‡ ‘†‹–‹‘‡†„›™‡ƒ–Š‡” ‘†‹–‹‘•ȋ ƒ‘„•‡ƒ†”‹•–Œž••‘ͳͻͻͶǡ‡–ƒ‹†‘—ƒ†‡–• ͳͻͻ͸ȌǤ‡–Š‡”‡ˆ‘”‡†‡ ‹†‡†–‘‡ƒ•—”‡‡ –ƒ”•–ƒ†‹‰ ”‘’•ƒ––Š‡‡†‘ˆ–Š‡†ƒ›ƒ•ƒ •—””‘‰ƒ–‡ˆ‘”‡ –ƒ” ‘•—’–‹‘Ǥ

Š›‡ ˆŽ‘™‡”• ’”‘†— ‡ ˜‡”› •ƒŽŽ ƒ‘—–• ‘ˆ ‡ –ƒ” ȋ”ƒ ‡– ƒŽǤ ʹͲͳͶȌǡ ™Š‹ Š ƒ› „‡ ‘‡†‹ˆˆ‹ —Ž––‘‡š–”ƒ –ǡ‡•’‡ ‹ƒŽŽ›‹™ƒ”†”›†ƒ›•Ǥ ‘”–Š‹•”‡ƒ•‘ǡ‡ –ƒ”•—”˜‡›•™‡”‡ ‘Ž› ‘†— –‡†‘”‘•‡ƒ”›Ǥ––Š‡‡†‘ˆ‡ƒ Š•ƒ’Ž‹‰†ƒ›ǡ™‡—•‡†ͳǦɊŽ ƒ’‹ŽŽƒ”›–—„‡• –‘‡ƒ•—”‡ –Š‡˜‘Ž—‡ ‘ˆ ‡ –ƒ””‡ƒ‹‹‰‹–Š‡ ˆŽ‘™‡”•Ǥ Š‹•™ƒ•†‘‡ ‘ ‘•– ‘ˆ–Š‡ ˆŽ‘™‡”• —•‡† ‹ –Š‡ ‡˜‡‹‰ ’‘ŽŽ‡ •—”˜‡›•Ǥ ‡ ‡ƒ•—”‡† ‡ –ƒ” •–ƒ†‹‰ ”‘’• ‹ ͳ͸ʹͺ ”‘•‡ƒ”›ˆŽ‘™‡”•Ǥ



2.2.3. Pollinatorvisitationrates

‘ ”‡Žƒ–‡ ’‘ŽŽ‡ ƒ† ‡ –ƒ” ‘•—’–‹‘ –‘ ’‘ŽŽ‹ƒ–‘” ƒ –‹˜‹–›ǡ ™‡ ‘†— –‡† ’‘ŽŽ‹ƒ–‘” •—”˜‡›•„‡–™‡‡ͻǣͲͲƒ†ͳͺǣͲͲ‹‡ƒ Š’Ž‘–‘–Š‡•ƒ‡†ƒ›‹™Š‹ Š’‘ŽŽ‡ƒ†‡ –ƒ” ‡ƒ•—”‡•™‡”‡–ƒ‡Ǥ–‡ƒ Š’Ž‘–ǡ ™‡•‡Ž‡ –‡† ͳͲ”‘•‡ƒ”›ƒ† ͳͲ–Š›‡’Žƒ–•ǤŠ‡•‡ ’Žƒ–•™‡”‡‘––Š‡•ƒ‡—•‡†‹’‘ŽŽ‡Ȁ‡ –ƒ”•—”˜‡›•–‘ƒ˜‘‹†’‘–‡–‹ƒŽƒ ‹†‡–ƒŽ ‘–ƒ – ™‹–ŠˆŽ‘™‡”•ƒ”‡†ˆ‘”’‘ŽŽ‡Ǧ‡ –ƒ”‡ƒ•—”‡•Ǥ‡ƒ Šƒ”‡†’Žƒ– ™‡ ‘†— –‡† ƒ  58 Šƒ’–‡”ʹ

—„‡”‘ˆ’‘ŽŽ‹ƒ–‘” ‘—–•ȋ‡ƒαͳͲǡ”ƒ‰‡αͷǦͳͷȌ–Š”‘—‰Š‘—––Š‡†ƒ›Ǥ ‡ƒ Š ‘—––Š‡ •‡Ž‡ –‡† ’Žƒ– ™ƒ• ‘„•‡”˜‡† ˆ‘” ʹ ‹—–‡• ƒ† ƒŽŽ ’‘ŽŽ‹ƒ–‘”• ‘–ƒ –‹‰ ˆŽ‘™‡”• ™‡”‡ ”‡ ‘”†‡†Ǥ‘–ƒŽ‘„•‡”˜ƒ–‹‘–‹‡™ƒ•͹ʹŠͶͺ‹ˆ‘””‘•‡ƒ”›ƒ†͹͸Š͵Ͷ‹ˆ‘”–Š›‡Ǥ ––Š‡‡†‘ˆ–Š‡†ƒ›ǡ™‡ ‘—–‡†–Š‡—„‡”‘ˆ‘’‡ˆŽ‘™‡”•‹‡ƒ Š’Žƒ–ǤApismellifera ƒ†–Š‡„—„Ž‡„‡‡Bombusterrestris™‡”‡ǡ„›ˆƒ”ǡ–Š‡–™‘‘•–ˆ”‡“—‡–•’‡ ‹‡•˜‹•‹–‹‰–Š‡ –™‘’Žƒ–•’‡ ‹‡•ǤŠ‡”‡ˆ‘”‡ǡ™‡‰”‘—’‡†’‘ŽŽ‹ƒ–‘”•‹–‘–Š”‡‡ ƒ–‡‰‘”‹‡•ǣA.melliferaǡB. terrestrisǡƒ†‘–Š‡”’‘ŽŽ‹ƒ–‘”•ȋ‘•–Ž›‘–Š‡”„‡‡•ǡƒŽ‘‰™‹–Š•‘‡†‹’–‡”ƒ• ƒ† ƒ ˆ‡™ Ž‡’‹†‘’–‡”ƒ•ƒ† ‘Ž‡‘’–‡”ƒ•ȌǤ‹•‹–ƒ–‹‘”ƒ–‡•‘ˆ‡ƒ Š’‘ŽŽ‹ƒ–‘”‰”‘—’™‡”‡ ƒŽ —Žƒ–‡†ƒ• –Š‡—„‡”‘ˆ ‘–ƒ –•’‡”‹—–‡ƒ†’‡”ͳͲͲͲˆŽ‘™‡”•Ǥ



2.2.4. Beecommunity

‘ƒ••‡••„‡‡ ‘—‹–›•–”— –—”‡ƒ† ‘’‘•‹–‹‘ǡ™‡’Žƒ ‡†͸•ƒ’Ž‹‰•–ƒ–‹‘•‹‡ƒ Š ’Ž‘–Ǥ ƒ Š •ƒ’Ž‹‰ •–ƒ–‹‘ ‘•‹•–‡† ‘ˆ ƒ ‡–ƒŽ „ƒ” Š‘Ž†‹‰ ͵ ’ƒ –”ƒ’• ’ƒ‹–‡† ›‡ŽŽ‘™ǡ ™Š‹–‡ ƒ† „Ž—‡ ”‡•’‡ –‹˜‡Ž›ǡ ‘‡ ‡–‡” ƒ„‘˜‡ –Š‡ ‰”‘—† ȋ‡•–’ŠƒŽ ‡– ƒŽǤ ʹͲͲͺȌǤ ‡ ‘†— –‡†ͺ„‹™‡‡Ž›•—”˜‡›•ˆ”‘‹†ƒ” Š–‘Žƒ–‡ —‡ʹͲͳͲǡ‹™Š‹ Š–”ƒ’•™‡”‡•‡–—’ „‡ˆ‘”‡ͻǣ͵Ͳƒ† ‘ŽŽ‡ –‡†ƒˆ–‡”ͳͺǣͲͲǤŽŽ’Ž‘–•™‡”‡•ƒ’Ž‡†‘–Š‡•ƒ‡ͺ†ƒ›•ȋ•‡‡‘”±Ǧ ‘‰—‡”ƒ‡–ƒŽǤȋʹͲͳͶȌˆ‘”†‡–ƒ‹Ž•ȌǤ‡ ƒ’–—”‡†͸ͷͺͲ„‡‡•’‡ ‹‡•ǡ™Š‹ Š™‡”‡†”‹‡†ƒ† ’‹‡† ˆ‘” ‹†‡–‹ˆ‹ ƒ–‹‘Ǥ  ƒ††‹–‹‘ǡ ™‡ ‡––‡† ƒ† ™‡‹‰Š‡† ƒˆ‡™‹†‹˜‹†—ƒŽ•‘ˆ‡ƒ Š •’‡ ‹‡•Ȁ•‡š –‘ ‘„–ƒ‹ ‡ƒ•—”‡• ‘ˆ ˆ”‡•Š „‘†› ™‡‹‰Š– ȋαͳǦͷʹ •’‡ ‹‡•’‡”•’‡ ‹‡•ȌǤ ’‡ ‹‡•™‡”‡•—„•‡“—‡–Ž› Žƒ••‡†ƒ••ƒŽŽȋ„‘†›™‡‹‰Š–δͷͷ‰Ȍ‘”Žƒ”‰‡ȋε͹Ͳ‰Ȍȋ•‡‡ ‘”±Ǧ‘‰—‡”ƒ‡–ƒŽǤȋʹͲͳͶȌˆ‘”ƒ‘”‡†‡–ƒ‹Ž‡†‡š’Žƒƒ–‹‘ȌǤ



2.2.5. Flowerabundance

‘‡•–‹ƒ–‡ˆŽ‘™‡”ƒ„—†ƒ ‡‹‡ƒ Š’Ž‘–ǡ™‡ ‘•‹†‡”‡†–Š‡ƒ‹ˆŽ‘™‡”‹‰•’‡ ‹‡•‹–Š‡ • ”—„Žƒ†ȋR.officinalisǡT.vulgarisǡDorycniumpentaphyllumǡCistusalbidusǡCistussalvifolius ƒ†CistusmonspeliensisȌǡ™Š‹ Š–‘‰‡–Š‡”ƒ ‘—–ˆ‘”ε͹ͲΨ‘ˆ–Š‡ˆŽ‘™‡”•‹‡ƒ Š’Ž‘–Ǥ‡ ‡ƒ•—”‡†–™‘’‡”’‡†‹ —Žƒ”™‹†–Š•ƒ†–Š‡Š‡‹‰Š–‘ˆ‡ƒ ŠˆŽ‘™‡”’ƒ– Š‹–™‘ͶͲšͳ ’‡”’‡†‹ —Žƒ”–”ƒ•‡ –• ‡–‡”‡†‹–Š‡‹††Ž‡‘ˆ–Š‡’Ž‘–ǤŠ‡™‡‡•–‹ƒ–‡†–Š‡—„‡” ‘ˆˆŽ‘™‡”•‘ˆ‡ƒ Š•’‡ ‹‡•„ƒ•‡†‘’”‡˜‹‘—•Ž›‡•–ƒ„Ž‹•Š‡†‡“—ƒ–‹‘•”‡Žƒ–‹‰’ƒ– Š˜‘Ž—‡

 59 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

ƒ†—„‡”‘ˆ‘’‡ˆŽ‘™‡”•ƒ–’‡ƒ„Ž‘‘ȋ”ʹαͲǤ͵͸ȂͲǤ͸͵ǡ ’αͲǤͲͲͳȂͲǤͲͳͷȌ ȋ•‡‡ ‘”±Ǧ ‘‰—‡”ƒ‡–ƒŽǤǡʹͲͳͶȌǤ‡ ƒ—•‡–Š‡–Š”‡‡Cistus•’‡ ‹‡•™‡”‡— ŠŽ‡••ƒ„—†ƒ––Šƒ–Š‡ ‘–Š‡”•’‡ ‹‡•ǡƒ†–Š‡‹”ˆŽ‘™‡”‹‰’‡”‹‘†•Žƒ”‰‡Ž›‘˜‡”Žƒ’ǡ™‡Ž—’‡†–‘‰‡–Š‡”–Š‡•‡–Š”‡‡ •’‡ ‹‡•‹–‘ƒ•‹‰Ž‡ ƒ–‡‰‘”›ȋCistusƒ„—†ƒ ‡ȌǤ



2.2.6. Statisticalanalysis

Visitationratesandpollen/nectarconsumption

”‡Ž‹‹ƒ”›ƒƒŽ›•‡••Š‘™‡†‘ ‘””‡Žƒ–‹‘„‡–™‡‡‡š’Žƒƒ–‘”›˜ƒ”‹ƒ„Ž‡•ȋ˜‹•‹–ƒ–‹‘”ƒ–‡• ‘ˆ –Š‡ †‹ˆˆ‡”‡– ’‘ŽŽ‹ƒ–‘” ‰”‘—’•ȌǤ  ’‘ŽŽ‡ ƒƒŽ›•‹•ǡ Š‘‡› „‡‡ ˜‹•‹–ƒ–‹‘ ”ƒ–‡ ƒ† ‘–Š‡” ’‘ŽŽ‹ƒ–‘”• ˜‹•‹–ƒ–‹‘ ”ƒ–‡ ™‡”‡ Ž‘‰Ǧ–”ƒ•ˆ‘”‡† „‡ ƒ—•‡ –Š‡”‡ ™ƒ• ƒ Ž‘‰ƒ”‹–Š‹  ”‡Žƒ–‹‘•Š‹’„‡–™‡‡–Š‡•‡˜ƒ”‹ƒ„Ž‡•ƒ†’‘ŽŽ‡ ‘•—’–‹‘Ǥ‡‹‹–‹ƒŽŽ›ˆ‹–ƒ‰‡‡”ƒŽ‹œ‡† Ž‹‡ƒ”‘†‡Žȋ Ȍƒ••—‹‰ƒ„‹‘‹ƒŽ‡””‘”†‹•–”‹„—–‹‘ȋƒ†‡“—ƒ–‡ˆ‘”’”‘’‘”–‹‘†ƒ–ƒ •— Šƒ•’‘ŽŽ‡ ‘•—’–‹‘Ȍǡ™‹–ŠA.mellifera˜‹•‹–ƒ–‹‘”ƒ–‡ǡBombusspp.˜‹•‹–ƒ–‹‘”ƒ–‡ƒ† ‘–Š‡” ’‘ŽŽ‹ƒ–‘”• ˜‹•‹–ƒ–‹‘ ”ƒ–‡ ƒ• ’”‡†‹ –‹˜‡ ˜ƒ”‹ƒ„Ž‡•Ǥ ‘™‡˜‡”ǡ–Š‡‘†‡Ž•Š‘™‡† ‘˜‡”†‹•’‡”•‹‘ǤŠ‡”‡ˆ‘”‡ǡ™‡ˆ‹ƒŽŽ›‘’–‡†ˆ‘”ƒ“—ƒ•‹„‹‘‹ƒŽ Ǥ‡–Š‡ ‘’ƒ”‡†–Š‡ •ƒ–—”ƒ–‡† ‘†‡Ž ™‹–Š –Š‡ ˜ƒ”‹‘—• ‘Ǧ•ƒ–—”ƒ–‡† ‘†‡Ž• ƒ† Š‘•‡–Š‡„‡•–‘‡—•‹‰  ȋƒ•   ƒ‘– „‡ ƒŽ —Žƒ–‡† ˆ‘” “—ƒ•‹ ‘†‡Ž ˆƒ‹Ž‹‡•ȌǤ ‹ƒŽŽ›ǡ ™‡ Š‡ ‡† ˆ‘” ‘”ƒŽ‹–› ƒ† Š‘‘• ‡†ƒ•–‹ ‹–› ‘ˆ –Š‡ ”‡•‹†—ƒŽ•Ǥ ‡ —•‡ ’•‡—†‘Ǧʹ ƒ• ƒ ‡ƒ•—”‡ ‘ˆ –Š‡ ‰‘‘†‡••Ǧ‘ˆǦˆ‹–Ǥ

‡ –ƒ”ƒƒŽ›•‡•ǡ™‡Ž‘‰Ǧ–”ƒ•ˆ‘”‡†–Š‡‡š’Žƒƒ–‘”›˜ƒ”‹ƒ„Ž‡Dz‘–Š‡”’‘ŽŽ‹ƒ–‘”•˜‹•‹–ƒ–‹‘ ”ƒ–‡dz „‡ ƒ—•‡ ‹– •Š‘™‡† ƒ Ž‘‰ƒ”‹–Š‹  ”‡Žƒ–‹‘•Š‹’ ™‹–Š –Š‡ ”‡•’‘•‡ ˜ƒ”‹ƒ„Ž‡Ǥ ‡ ˆ‹– ƒ ‰‡‡”ƒŽ‹œ‡† Ž‹‡ƒ” ‘†‡Ž ȋ Ȍ ™‹–Š ƒ ƒ—••‹ƒ ‡””‘” †‹•–”‹„—–‹‘ǡ ™‹–Š ‡ –ƒ” •–ƒ†‹‰ ”‘’ ƒ• –Š‡ ”‡•’‘•‡ ˜ƒ”‹ƒ„Ž‡ ƒ† ˜‹•‹–ƒ–‹‘ ”ƒ–‡ ‘ˆ –Š‡ ˜ƒ”‹‘—• ’‘ŽŽ‹ƒ–‘” ‰”‘—’• ƒ• ’”‡†‹ –‹˜‡ ˜ƒ”‹ƒ„Ž‡•Ǥ ‡ •‡Ž‡ –‡† –Š‡ „‡•– ‘†‡Ž —•‹‰ –Š‡ •‡ ‘†Ǧ‘”†‡” ƒ‹‡̵• ˆ‘”ƒ–‹‘”‹–‡”‹‘ȋ  Ȍǡƒ†‡“—ƒ–‡ˆ‘”•ƒŽŽ•ƒ’Ž‡•Ǥ



 

 60 Šƒ’–‡”ʹ

Beecommunity

‡—•‡†‹”ƒ‘•‘ˆ–™ƒ”‡ȋ‘•ʹͲͳͶȌ–‘‡•–ƒ„Ž‹•Š–Š‡Ž‹‡ƒ”†‹•–ƒ ‡‘ˆ‡ƒ Š’Ž‘––‘–Š‡ ‡ƒ”‡•– ƒ’‹ƒ”› ‹ –Š‡ ƒ”‡ƒǤ Š‹• ‡ƒ•—”‡ǡ ™Š‹ Š ‹• ‘‘Ž› —•‡† ‹ Š‘‡› „‡‡ •–—†‹‡• ȋ–‡ˆˆƒǦ‡™‡–‡” ƒ† • Šƒ”–‡ ʹͲͲͲǡ Š‘•‘ ʹͲͲͶǡ —†‡™‡œ ƒ† Ž‡‹ ʹͲͳ͵ǡ Ž„‰ƒ‹ ‡– ƒŽǤ ʹͲͳͶȌǡ ™ƒ• ‡‰ƒ–‹˜‡Ž› ‘””‡Žƒ–‡† –‘ Š‘‡› „‡‡ ˜‹•‹–ƒ–‹‘ ”ƒ–‡• ȋ”ʹα ͶʹǤʹͷǡ ’αͲǤͲͲͻȌǡƒ†–‘Š‘‡›„‡‡ƒ„—†ƒ ‡‹–Š‡’ƒ–”ƒ’•ȋŽ‘‰ƒ”‹–Š‹ ”‡Žƒ–‹‘•Š‹’ǡ”ʹαͶͻǤ͹͵ǡ ’αͲǤͲͲͲͶȌǤ





Figure 2.1 ƒ”–‹ƒŽ ”‡‰”‡••‹‘ ’Ž‘–• •Š‘™‹‰ –Š‡ ‘–”‹„—–‹‘ ‘ˆ Š‘‡› „‡‡ ƒ† „—„Ž‡„‡‡  ˜‹•‹–ƒ–‹‘”ƒ–‡•–‘”‘•‡ƒ”›ƒ†–Š›‡’‘ŽŽ‡ ‘•—’–‹‘‹ʹͳ’Ž‘–•ǡ‘ ‡–Š‡‡ˆˆ‡ –‘ˆ‘–Š‡” ‡š’Žƒƒ–‘”›˜ƒ”‹ƒ„Ž‡•‡–‡”‹‰–Š‡ •Šƒ•„‡‡”‡‘˜‡†ȋ„—„Ž‡„‡‡˜‹•‹–ƒ–‹‘”ƒ–‡•‹ȋȌ  ƒ†ȋȌǢŠ‘‡›„‡‡˜‹•‹–ƒ–‹‘”ƒ–‡•‹ȋȌƒ†ȋȌȌǤ 

‘‡˜ƒŽ—ƒ–‡–Š‡’‘–‡–‹ƒŽ”‡Žƒ–‹‘•Š‹’„‡–™‡‡†‹•–ƒ ‡–‘–Š‡‡ƒ”‡•–ƒ’‹ƒ”›ƒ†™‹Ž†„‡‡ ‘—‹–›•–”— –—”‡ǡ™‡”— ‘†‡Ž•ˆ‘”™‹Ž†„‡‡ƒ„—†ƒ ‡ǡ™‹Ž†„‡‡”‹ Š‡••ƒ†  61 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

™‹Ž†„‡‡„‹‘ƒ••Ǥ‡ ƒ—•‡™‹Ž†„‡‡ ‘—‹–›•–”— –—”‡ƒ›ƒŽ•‘„‡‹ˆŽ—‡ ‡†„›ˆŽ‘™‡” ƒ˜ƒ‹Žƒ„‹Ž‹–› ȋ‘”±Ǧ‘‰—‡”ƒ ‡– ƒŽǤ ʹͲͳͶȌǡ ™‡ ‹ Ž—†‡† ˆŽ‘™‡” ƒ„—†ƒ ‡ ‘ˆ T. vulgarisǡ R. officinalisǡ D. pentaphyllumƒ†Cistus ƒ• ’”‡†‹ –‘” ˜ƒ”‹ƒ„Ž‡•Ǥ ‡ †‹† ‘– ‹ Ž—†‡ ‡•–‹‰ •—„•–”ƒ–‡ƒ˜ƒ‹Žƒ„‹Ž‹–›‹–Š‡ƒƒŽ›•‹•„‡ ƒ—•‡™‡‘™ˆ”‘’”‡˜‹‘—••–—†‹‡•–Šƒ––Š‹•‹•‘– ƒ ‰‘‘† ’”‡†‹ –‘” ‘ˆ „‡‡ ‘—‹–› •–”— –—”‡ ƒ† ‘’‘•‹–‹‘ ‹ –Š‡ •–—†› ƒ”‡ƒ ȋ‘”±Ǧ ‘‰—‡”ƒ‡–ƒŽǤʹͲͳͶȌǤ

‡‡„‹‘ƒ••™ƒ•ƒƒŽ›œ‡†™‹–Šƒ ™‹–Šƒ ƒ—••‹ƒ†‹•–”‹„—–‹‘Ǥ ‘”„‡‡ƒ„—†ƒ ‡ƒ† „‡‡”‹ Š‡••‘†‡Ž•ǡ™‡ Š‘•‡ƒ ™‹–Šƒ‘‹••‘‡””‘”†‹•–”‹„—–‹‘ǡƒ†‡“—ƒ–‡ˆ‘” ‘—– †ƒ–ƒǤ ‘™‡˜‡”ǡ„‘–Š‘†‡Ž••Š‘™‡†‘˜‡”†‹•’‡”•‹‘ǡƒ†–Š—•™‡‘’–‡†ˆ‘”‘†‡Ž•™‹–Šƒ ‡‰ƒ–‹˜‡ „‹‘‹ƒŽ †‹•–”‹„—–‹‘Ǥ  ƒŽŽ –Š”‡‡ ƒƒŽ›•‡•ǡ ™‡ •‡Ž‡ –‡† –Š‡ „‡•– ‘†‡Ž ™‹–Š –Š‡    ”‹–‡”‹‘Ǥ‡•–‘†‡Ž•™‡”‡Žƒ–‡” Š‡ ‡†ˆ‘”‘”ƒŽ‹–›ƒ†Š‘‘• ‡†ƒ•–‹ ‹–›Ǥ‡ ƒ—•‡ Žƒ”‰‡ „‡‡• ‹‰Š– ”‡•’‘† †‹ˆˆ‡”‡–Ž› ˆ”‘ •ƒŽŽ „‡‡• †—‡ –‘ –Š‡‹” Š‹‰Š‡” ˆ‡‡†‹‰ ”‡“—‹”‡‡–•ȋòŽŽ‡”‡–ƒŽǤʹͲͲ͸Ȍǡ™‡”—ƒ††‹–‹‘ƒŽƒƒŽ›•‡••‡’ƒ”ƒ–‡Ž›ˆ‘”•ƒŽŽȋδͷͷ‰Ȍ ƒ† Žƒ”‰‡ ȋε͹Ͳ ‰Ȍ „‡‡•Ǥ Š‡ „‡•– ‘†‡Ž ‡š’Žƒ‹‹‰ ™‹Ž† „‡‡ ”‹ Š‡•• •Š‘™‡† Š‡–‡”‘• ‡†ƒ•–‹ ‹–›ǤŠ—•ǡ™‡—•‡†Š‹–‡ǯ•Š‡–‡”‘• ‡†ƒ•–‹ ‹–›Ǧ ‘””‡ –‡† ‘˜ƒ”‹ƒ ‡ƒ–”‹ ‡• –‘ƒ‡‹ˆ‡”‡ ‡Ǥ

ŽŽƒƒŽ›•‡•™‡”‡ ‘’—–‡†™‹–Šȋ‘”‡‡ƒʹͲͳͶȌǤ



2.3. Results

2.3.1. Pollenandnectarconsumption

Apismelliferaƒ†B.terrestrisƒ ‘—–‡†ˆ‘”–Š‡ƒŒ‘”‹–›‘ˆ˜‹•‹–•–‘„‘–Š”‘•‡ƒ”›ȋ͸ͳǤʹ ƒ†͵ͲǤͳΨǡ”‡•’‡ –‹˜‡Ž›Ȍƒ†–Š›‡ȋ͵ͻǤͷƒ†͵ͶǤͺΨȌǤ‹•‹–•‘ˆ‘–Š‡”’‘ŽŽ‹ƒ–‘”•ƒ‘—–‡† –‘ͺǤ͹ƒ†ʹͷǤ͹Ψ‘ˆ–Š‡˜‹•‹–•–‘”‘•‡ƒ”›ƒ†–Š›‡ǡ”‡•’‡ –‹˜‡Ž›Ǥ ‘‡›„‡‡ˆŽ‘™‡”˜‹•‹–• ƒ†„—„Ž‡„‡‡ˆŽ‘™‡”˜‹•‹–•™‡”‡‘– ‘””‡Žƒ–‡†ȋ”‘•‡ƒ”›ǣ”ʹαͲǤͲͶǡ’αͲǤʹ͹Ǣ–Š›‡ǣ”ʹα ͲǤͲͷǡ’αͲǤͳͻȌǤ

‡ƒά—„‡”‘ˆ’‘ŽŽ‡‰”ƒ‹•‹‡™Ž›Ǧ‘’‡‡†”‘•‡ƒ”›ˆŽ‘™‡”•™ƒ•ͷͳͺͷάͳͷͷͻǡ ƒ†–Š‡•‡—„‡”•†‡ ”‡ƒ•‡†–‘ͳͺ͵ͳάͳͷͳ͹„›–Š‡‡†‘ˆ–Š‡†ƒ›Ǥ‘ŽŽ‡ ‘•—’–‹‘‹ ‘—” ’Ž‘–• ”ƒ‰‡† ˆ”‘ ʹͷǤͳΨ –‘ ͻͲǤͳΨ ȋ‡ƒ ά  α ͸ͷǤ͸ ά ͳͺǤͶȌǤŠ‡„‡•–‘†‡Žˆ‘” ”‘•‡ƒ”› ’‘ŽŽ‡ ‘•—’–‹‘ ȋ’•‡—†‘Ǧʹα ͲǤͷͶȌ ‹ Ž—†‡† A. mellifera ˜‹•‹–ƒ–‹‘ ”ƒ–‡  62 Šƒ’–‡”ʹ

ȋ’αͲǤͲͲͶȌ ƒ†ǡ ƒ”‰‹ƒŽŽ›ǡ B. terrestris˜‹•‹–ƒ–‹‘”ƒ–‡ȋ’αͲǤͲ͸Ȍȋ ‹‰ǤʹǤͳȌǤŠ‡‘†‡Ž ‹ Ž—†‹‰‘Ž›A.mellifera˜‹•‹–ƒ–‹‘”ƒ–‡ȋ’αͲǤͲͲͳǢ’•‡—†‘ǦʹαͲǤͶͶȌ™ƒ•‘Ž›ƒ”‰‹ƒŽŽ› †‹ˆˆ‡”‡–ˆ”‘–Š‡•ƒ–—”ƒ–‡†‘†‡Žȋ αͶǤͳͻǡ’αͲǤͲ͸ȌǤ





Figure 2.2ƒ”–‹ƒŽ”‡‰”‡••‹‘’Ž‘–••Š‘™‹‰–Š‡ ‘–”‹„—–‹‘‘ˆŠ‘‡›„‡‡ƒ†„—„Ž‡„‡‡ ˜‹•‹–ƒ–‹‘”ƒ–‡•–‘”‘•‡ƒ”›‡ –ƒ”•–ƒ†‹‰ ”‘’•‹ʹͳ’Ž‘–•ǡ ‘ ‡ –Š‡ ‡ˆˆ‡ – ‘ˆ ‘–Š‡” ‡š’Žƒƒ–‘”›˜ƒ”‹ƒ„Ž‡•‡–‡”‹‰–Š‡ •Šƒ•„‡‡”‡‘˜‡†ȋ„—„Ž‡„‡‡˜‹•‹–ƒ–‹‘”ƒ–‡•‹ȋȌǢ Š‘‡›„‡‡˜‹•‹–ƒ–‹‘”ƒ–‡•‹ȋȌȌǤ   Š›‡ˆŽ‘™‡”• ‘–ƒ‹‡†ͳʹʹͲά͹͵͹’‘ŽŽ‡‰”ƒ‹•‹–Š‡‘”‹‰ƒ†ͷ͹͹άͶ͵ͻƒ––Š‡‡† ‘ˆ–Š‡†ƒ›ǤŠ›‡’‘ŽŽ‡ ‘•—’–‹‘‹–Š‡˜ƒ”‹‘—•’Ž‘–•”ƒ‰‡† „‡–™‡‡ ͳͻǤʹΨ ƒ† ͹͸ǤͷΨȋ‡ƒάαͷͶǤ͵άͳͷǤͻȌǤŠ‡„‡•–‘†‡Žˆ‘”–Š›‡’‘ŽŽ‡ ‘•—’–‹‘ȋ’•‡—†‘Ǧ ʹα ͲǤͶʹȌ ‹ Ž—†‡† A. mellifera ˜‹•‹–ƒ–‹‘ ”ƒ–‡ ȋ’αͲǤͲͲʹȌ ƒ† B. terrestris ˜‹•‹–ƒ–‹‘ ”ƒ–‡ ȋ’αͲǤͲͶȌȋ ‹‰ǤʹǤͳȌǤ

‘•‡ƒ”›‡ –ƒ”•–ƒ†‹‰ ”‘’•‹–Š‡ʹͳ’Ž‘–•”ƒ‰‡†ˆ”‘Ͳ–‘ ͸Ǥ͵ͳ ɊȀˆŽ‘™‡” ȋͲǤʹ͸ ά ͲǤ͵ͻȌǤ Š‡ „‡•– ‘†‡Ž ‡š’Žƒ‹‹‰ ”‘•‡ƒ”› ‡ –ƒ” Ž‡˜‡Ž• ȋ”ʹα ͲǤͶʹȌ ‹ Ž—†‡† A. mellifera ˜‹•‹–ƒ–‹‘ ”ƒ–‡ ȋ’αͲǤͲͶȌ ƒ†ǡ ƒ”‰‹ƒŽŽ›ǡ B. terrestris ˜‹•‹–ƒ–‹‘ ”ƒ–‡ ȋ’αͲǤͲͷȌ ȋ ‹‰Ǥ ʹǤʹȌǤ • ™‹–Š ”‘•‡ƒ”› ’‘ŽŽ‡ ‘•—’–‹‘ǡ –Š‡ ‘†‡Ž ‹ Ž—†‹‰ ‘Ž› A. mellifera ˜‹•‹–ƒ–‹‘ ”ƒ–‡ ȋ’αͲǤͲʹ͹ǡ”ʹαͲǤʹ͸Ȍ™ƒ•‘Ž›ƒ”‰‹ƒŽŽ›Ž‡••‡š’Ž‹ ƒ–‹˜‡–Šƒ–Š‡•ƒ–—”ƒ–‡†‘†‡Žȋ αͶǤͳͻǡ ’αͲǤͲ͸ȌǤ



 63 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

2.3.2. Beecommunity

ƒ–”ƒ’•—”˜‡›•›‹‡Ž†‡†͸ͷͺͲ„‡‡•’‡ ‹‡• ‘””‡•’‘†‹‰–‘ͻͺ•’‡ ‹‡•Ǥ‹š–›Ǧ–”‡‡‘ˆ–Š‡ ‘ǦApis•’‡ ‹‡•™‡”‡•ƒŽŽȋˆ”‡•Š„‘†›™‡‹‰Š–δͷͷ‰Ȍƒ†͵Ͷ™‡”‡Žƒ”‰‡ȋε͹Ͳ‰ȌǤ





Figure2.3ƒ”–‹ƒŽ”‡‰”‡••‹‘’Ž‘–••Š‘™‹‰–Š‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡†‹•–ƒ ‡–‘–Š‡‡ƒ”‡•– ƒ’‹ƒ”›ƒ†Žƒ”‰‡„‡‡ƒ„—†ƒ ‡ƒ†„‹‘ƒ••‹ʹͳ’Ž‘–•ǡ‘ ‡–Š‡‡ˆˆ‡ –‘ˆ‘–Š‡”‡š’Žƒƒ–‘”› ˜ƒ”‹ƒ„Ž‡• ‡–‡”‹‰ –Š‡ • Šƒ• „‡‡ ”‡‘˜‡† ȋCistusˆŽ‘™‡”ƒ„—†ƒ ‡‹ȋȌƒ†ȋȌǢ †‹•–ƒ ‡–‘–Š‡‡ƒ”‡•–ƒ’‹ƒ”›‹ȋȌƒ†ȋȌȌǤ   ‘˜ƒ”‹ƒ„Ž‡•‡–‡”‡†–Š‡‘†‡Ž–‘‡š’Žƒ‹™‹Ž†„‡‡”‹ Š‡••ȋƒ„Ž‡ʹǤͳȌǡƒ†•‹‹Žƒ””‡•—Ž–• ™‡”‡‘„–ƒ‹‡†™Š‡•ƒŽŽƒ†Žƒ”‰‡„‡‡•™‡”‡ ƒƒŽ›œ‡†•‡’ƒ”ƒ–‡Ž›ȋƒ„Ž‡ʹǤͳȌǤŠ‡„‡•– ‘†‡Žˆ‘”„‡‡ƒ„—†ƒ ‡ȋ’•‡—†‘ǦʹαͲǤͶͺȌ‹ Ž—†‡†CistusˆŽ‘™‡”ƒ„—†ƒ ‡ȋ’αͲǤͲͲʹȌ ƒ†T.vulgarisˆŽ‘™‡”ƒ„—†ƒ ‡ȋ’αͲǤͲͲͶȌǤ‹‹Žƒ””‡•—Ž–•™‡”‡‘„–ƒ‹‡†™Š‡‘Ž›•ƒŽŽ „‡‡• ™‡”‡ –ƒ‡ ‹–‘ ƒ ‘—– ȋ’•‡—†‘Ǧʹα ͲǤͶͳǢ Cistusƒ„—†ƒ ‡ȋ’αͲǤͲͲͺȌǢT. vulgaris

 64 Šƒ’–‡”ʹ

ƒ„—†ƒ ‡ȋ’αͲǤͲ͵ȌȌǤ •–‡ƒ†ǡ–Š‡„‡•–ˆ‹–‘†‡Žˆ‘”Žƒ”‰‡„‡‡•ȋ’•‡—†‘ǦʹαͲǤͷͲȌ‹ Ž—†‡† †‹•–ƒ ‡–‘–Š‡‡ƒ”‡•– ƒ’‹ƒ”›ȋ’αͲǤͲʹȌ ƒ†ǡƒ”‰‹ƒŽŽ›ǡ Cistusƒ„—†ƒ ‡ȋ’αͲǤͲ͸Ȍȋ ‹‰Ǥ ʹǤ͵Ǣƒ„Ž‡ʹǤͳȌǤ‘„‡ ‘•‡”˜ƒ–‹˜‡ǡ™‡”‡Ǧ”—–Š‡Žƒ––‡”ƒƒŽ›•‹•™‹–Š‘—–͵’‘••‹„Ž‡‡˜‡”ƒ‰‡ ’‘‹–•ȋ‘‘ǯ•αͲǤͷ–‘ͳȌǡƒ†‘„–ƒ‹‡†•‹‹Žƒ””‡•—Ž–•™‹–Š ƒ Ž‘™‡” ‰‘‘†‡••Ǧ‘ˆǦˆ‹– ȋ’•‡—†‘ǦʹαͲǤʹͺǡ†‹•–ƒ ‡–‘ƒ’‹ƒ”›’αͲǤͲʹǡCistusƒ„—†ƒ ‡’αͲǤͲ͸ȌǤŠ‡„‡•–™‹Ž†„‡‡ „‹‘ƒ••‘†‡Žȋ’•‡—†‘ǦʹαͲǤͷ͸Ȍ‹ Ž—†‡†CistusˆŽ‘™‡”ƒ„—†ƒ ‡ȋ’αͲǤͲͲʹȌƒŽ‘‰™‹–Š †‹•–ƒ ‡–‘–Š‡‡ƒ”‡•–ƒ’‹ƒ”›ȋ’αͲǤͲʹȌǤŠ‡„‡•–‘†‡Žˆ‘”•ƒŽŽ„‡‡•ȋ’•‡—†‘ǦʹαͲǤʹ͹Ȍ ‹ Ž—†‡†‘Ž›Cistusƒ„—†ƒ ‡ȋ’αͲǤͲʹȌȋƒ„Ž‡ʹǤͳȌǤ‘˜‡”•‡Ž›ǡ–Š‡„‡•–‘†‡Žˆ‘”Žƒ”‰‡ „‡‡•ȋ’•‡—†‘ǦʹαͲǤͷͶȌ‹ Ž—†‡††‹•–ƒ ‡–‘–Š‡‡ƒ”‡•–ƒ’‹ƒ”›ȋ’αͲǤͲͲ͹Ȍƒ†ǡƒ”‰‹ƒŽŽ›ǡ Cistusƒ„—†ƒ ‡ȋ’αͲǤͲ͸Ȍȋ ‹‰ǤʹǤ͵ȌǤ



2.4. Discussion

‘‡›„‡‡• ‘—–—„‡”‡† –Š‡ ‘•– ˆ”‡“—‡– ™‹Ž† „‡‡ ȋ–Š‡ „—„Ž‡„‡‡ B. terrestrisȌ ‘ ”‘•‡ƒ”›ƒ†–Š›‡ˆŽ‘™‡”•ǡ–Š‡–™‘ƒ‹ˆŽ‘™‡”‹‰’Žƒ–•‹–Š‡•–—†›ƒ”‡ƒǤŽŽ™‘”‡”•‘ˆ –Š‡•‡–™‘„‡‡•’‡ ‹‡• ‘ŽŽ‡ –‡†‡ –ƒ”ǡƒ†•‘‡‘ˆ–Š‡ƒŽ•‘ ‘ŽŽ‡ –‡†’‘ŽŽ‡Ǥ—””‡•—Ž–• †‡‘•–”ƒ–‡ –Šƒ– Š‘‡› „‡‡• ™‡”‡ǡ „› ˆƒ”ǡ –Š‡ ƒ‹ •’‡ ‹‡• ‘–”‹„—–‹‰ –‘ ’‘ŽŽ‡ ƒ† ‡ –ƒ” ‘•—’–‹‘ǤŠ‡ ‘–”‹„—–‹‘‘ˆB.terrestris™ƒ•— ŠŽ‘™‡”ǡƒ†‘–Š‡”’‘ŽŽ‹ƒ–‘”• ’Žƒ›‡†ƒ‘Ǧ†‡–‡ –ƒ„Ž‡”‘Ž‡‹ˆŽ‘™‡””‡•‘—” ‡ ‘•—’–‹‘Ǥ‡ƒ’‘ŽŽ‡ ‘•—’–‹‘’‡” ’Ž‘–™ƒ••Ž‹‰Š–Ž›Š‹‰Š‡”ˆ‘””‘•‡ƒ”›ȋ‡ƒα͸ͷǤ͸Ψǡ”ƒ‰‡αʹͷǤͳǦͻͲǤͳΨȌ–Šƒˆ‘”–Š›‡ ȋ‡ƒα ͷͶǤ͵Ψǡ ”ƒ‰‡α ͳͻǤʹ Ǧ ͹͸ǤͷΨȌǡ „—– –‘ ƒ ‰”‡ƒ–‡” ‘” Ž‡••‡” ‡š–‡–ǡ ƒŽŽ ’Ž‘–• Šƒ† ‘•‹†‡”ƒ„Ž‡ƒ‘—–•‘ˆ’‘ŽŽ‡ƒ†‡ –ƒ”ƒ˜ƒ‹Žƒ„Ž‡ƒ––Š‡‡†‘ˆ–Š‡†ƒ›ǤŠ‹•ƒ›•—‰‰‡•– –Šƒ– ˆŽ‘™‡” ”‡•‘—” ‡• ™‡”‡ ‘– ƒ Ž‹‹–‹‰ ˆƒ –‘” ˆ‘” –Š‡ „‡‡ ‘—‹–›Ǥ ‘™‡˜‡”ǡ –Š‡ ‡‡”‰‡–‹  ‰ƒ‹ ‘„–ƒ‹‡† ˆ”‘ ˆŽ‘™‡”• ™‹–Š ’‘ŽŽ‡Ǧ‡ –ƒ” Ž‡˜‡Ž• „‡Ž‘™ ƒ ‡”–ƒ‹ –Š”‡•Š‘Ž† ƒ› „‡ ‹•—ˆˆ‹ ‹‡– –‘ ‘’‡•ƒ–‡ ˆ‘”ƒ‰‹‰ ‘•–•ǡ ‡•’‡ ‹ƒŽŽ› ˆ‘” Žƒ”‰‡ „‡‡•ǡ ™‹–Š Š‹‰Š‡” ‡‡”‰‡–‹  †‡ƒ†• ȋ ‡‹”‹ Š ͳͻ͹ͷȌǤ ‡‡• Šƒ˜‡ „‡‡ •Š‘™ –‘ ‘˜‡ƒ™ƒ›ˆ”‘Ž‡•• ”‡™ƒ”†‹‰’ƒ– Š‡•ȋ ‡‹”‹ Šͳͻ͹ͻȌǤ—”’‘ŽŽ‡Ǧ‡ –ƒ”•—”˜‡›•™‡”‡ ‘†— –‡††—”‹‰’‡ƒ „Ž‘‘‘ˆ–Š‡–™‘ƒ‹ˆŽ‘™‡”•’‡ ‹‡•‹–Š‡•–—†›ƒ”‡ƒǤ›–Š‡‡†‘ˆ’”‹ŽǡˆŽ‘™‡””‡•‘—” ‡• „‡ ‘‡— Š• ƒ” ‡”‹–Š‡ƒ”ǡƒ†‘˜‡”ƒŽŽ˜‹•‹–ƒ–‹‘”ƒ–‡•ƒ”‡— ŠŠ‹‰Š‡”ȋ‘• Š‡–ƒŽǤ ʹͲͲͻǡ ‹Ž‡ŽŽƒ‡–ƒŽǤʹͲͳ͵ǡ Ž‘‡–ƒŽǤ•—„‹––‡†ȌǤ‘•‡“—‡–Ž›ǡ™‡‡š’‡ ––Š‡’‘–‡–‹ƒŽ‡ˆˆ‡ –• ‘ˆ‹–‡•‹˜‡Š‘‡›„‡‡ˆ‘”ƒ‰‹‰–‘„‡‰”‡ƒ–‡”Žƒ–‡‹Žƒ–‡Ǧ•’”‹‰ƒ†•—‡”Ǥ

 65 

Table2.1‡•—Ž–•‘ˆ •ƒƒŽ›œ‹‰™‹Ž†„‡‡”‹ Š‡••ǡƒ„—†ƒ ‡ƒ†„‹‘ƒ••‹ʹͳ’Ž‘–•ƒ•ƒˆ— –‹‘‘ˆ†‹•–ƒ ‡–‘–Š‡‡ƒ”‡•–ƒ’‹ƒ”›ǡƒ†ƒ„—†ƒ ‡‘ˆCistusǡ Thymusvulgaris,Rosmarinusofficinalisƒ†DorycniumpentaphyllumˆŽ‘™‡”•ǤǦ˜ƒŽ—‡•ƒ”‡‘Ž›‰‹˜‡ˆ‘”˜ƒ”‹ƒ„Ž‡•‡–‡”‹‰–Š‡‘†‡Ž•Ǥ•‡—†‘Ǧʹ‹•’”‘˜‹†‡†ˆ‘” ‡ƒ Š‘†‡Žƒ•ƒ‡ƒ•—”‡‘ˆ‰‘‘†‡••Ǧ‘ˆǦˆ‹–Ǥ

‡•’‘•‡˜ƒ”‹ƒ„Ž‡ š’Žƒƒ–‘”›˜ƒ”‹ƒ„Ž‡• •‡—†‘Ǧʹ   ‹•–ƒ ‡–‘ƒ’‹ƒ”› Cistus T.vulgaris R.officinalis D.pentaphyllum ‹Ž†„‡‡”‹ Š‡•• ƒ”‰‡•’‡ ‹‡•ͳ • • • • • ǦǦ  ƒŽŽ•’‡ ‹‡•ʹ • • • • • ǦǦ  ŽŽ•’‡ ‹‡• • • • • • ǦǦ ‹Ž†„‡‡ƒ„—†ƒ ‡ ƒ”‰‡•’‡ ‹‡•ͳ ’αͲǤͲͳͻ ’αͲǤͲ͸ͳ • • • ͲǤͷͲ  ƒŽŽ•’‡ ‹‡•ʹ • ’αͲǤͲͲͺ ’αͲǤͲ͵Ͳ • • ͲǤͶͳ  ŽŽ•’‡ ‹‡• • ’αͲǤͲͲʹ ’αͲǤͲͶʹ • • ͲǤͶͺ ‹Ž†„‡‡„‹‘ƒ•• ƒ”‰‡•’‡ ‹‡•ͳ ’αͲǤͲͲ͹ ’αͲǤͲͷͻ • • • ͲǤͷͶ  ƒŽŽ•’‡ ‹‡•ʹ • ’αͲǤͲͳ͸ • • • ͲǤʹ͹  ŽŽ•’‡ ‹‡• ’αͲǤͲͳ͹ ’αͲǤͲͳ͸ • • • ͲǤͷ͸

ͳ‘†›™‡‹‰Š–ε͹Ͳ‰ǣʹ‘†›™‡‹‰Š–δͷͷ‰Ǥ 







 Šƒ’–‡”ʹ

—” •–—†› ƒŽ•‘ •Š‘™• –Šƒ– ™‹Ž† „‡‡ ‘—‹–› ‹• ƒˆˆ‡ –‡† ƒ† ‘†‹ˆ‹‡† ‹ ƒ”‡ƒ• Ž‘•‡ –‘ ƒ’‹ƒ”‹‡•ǡ ™‹–Š ƒ Ž‘™‡” ‘˜‡”ƒŽŽ ™‹Ž† „‡‡ „‹‘ƒ•• ‡†‹ƒ–‡† „› ƒ Ž‘™‡” ƒ„—†ƒ ‡ ‘ˆ Žƒ”‰‡ „‡‡•Ǥ ƒŽŽ „‡‡• ”‡“—‹”‡ Ž‡•• ‡‡”‰› –‘ ˆŽ› ƒ† •—•–ƒ‹ ˆ‘”ƒ‰‹‰ ƒ† ‡•–‹‰ ƒ –‹˜‹–‹‡• ȋ ‡‹”‹ Šͳͻ͹ͷȌǤ ƒ††‹–‹‘ǡ•ƒŽŽ„‡‡•”‡“—‹”‡•ƒŽŽ‡”’‘ŽŽ‡Ȁ‡ –ƒ”ƒ‘—–•–‘’”‘†— ‡ ƒ‘ˆˆ•’”‹‰ȋòŽŽ‡”‡–ƒŽǤʹͲͲ͸ȌǤŠ—•ǡ’‘ŽŽ‡ƒ†‡ –ƒ”•–ƒ†‹‰ ”‘’•‹ƒ”‡ƒ• Ž‘•‡–‘–Š‡ ƒ’‹ƒ”‹‡• ƒ› „‡ •—ˆˆ‹ ‹‡– ˆ‘” •ƒŽŽ „‡‡• „—– ‘– ˆ‘” Žƒ”‰‡ „‡‡•Ǥ ˆ •‘ǡ Žƒ”‰‡ „‡‡• ƒ› „‡ ˆ‘” ‡†–‘‡•–•‘‡™Š‡”‡‡Ž•‡‘”™‹†‡–Š‡‹”ˆ‘”ƒ‰‹‰”ƒ‰‡•ǡ™Š‹ Šƒ”‡™‡ŽŽ‘™–‘„‡ ’‘•‹–‹˜‡Ž›”‡Žƒ–‡†–‘„‘†›•‹œ‡ȋ ƒ–Šƒƒ†• Šƒ”–‡ʹͲͲʹǡ ”‡‡Ž‡ƒˆ‡–ƒŽǤʹͲͲ͹ȌǤ• ˆ‘”•ƒŽŽ„‡‡•ǡ‡˜‡‹ˆ–Š‡‹”ƒ„—†ƒ ‡†‹†‘–†‹‹‹•Š Ž‘•‡–‘ƒ’‹ƒ”‹‡•ǡ–Š‡‹”ˆ‹–‡••‹‰Š– •–‹ŽŽ „‡ ƒˆˆ‡ –‡† „› –Š‡ Ž‘™‡” ’‘ŽŽ‡Ȁ‡ –ƒ” •–ƒ†‹‰ ”‘’•Ǥ – –Š‡ ‹–”ƒǦ•’‡ ‹ˆ‹  Ž‡˜‡Žǡ „‡‡ ƒ†—Ž– „‘†› •‹œ‡ ‹• †‹”‡ –Ž› ”‡Žƒ–‡† –‘ –Š‡ ƒ‘—– ‘ˆ ’‘ŽŽ‡Ǧ‡ –ƒ” ‘•—‡† „› –Š‡ Žƒ”˜ƒ ȋ‹„‡‹”‘ͳͻͻͶǡ‘• Šƒ†‹ ‡•ʹͲͲʹȌǡƒ†•‘‡•–—†‹‡•Šƒ˜‡•Š‘™ ”‡†— –‹‘• ‹ ‘ˆˆ•’”‹‰ „‘†› •‹œ‡ ‹ ’‘’—Žƒ–‹‘• ˆŽ›‹‰ ‹ ƒ”‡ƒ• ™‹–Š Ž‘™ Ž‡˜‡Ž• ‘ˆ ˆŽ‘™‡” ”‡•‘—” ‡• ȋ‡–‡”•‘ ƒ† ‘‹–„‡”‰ ʹͲͲ͸ǡ ‘• Š ʹͲͲͺȌǤ ˆˆ•’”‹‰ ƒŽŽ‘ ƒ–‡† •ƒŽŽ‡” ’‘ŽŽ‡Ȁ‡ –ƒ” ’”‘˜‹•‹‘•ƒ”‡‘”‡Ž‹‡Ž›–‘†‹‡†—”‹‰†‡˜‡Ž‘’‡–ȋ‘• ŠʹͲͲͺȌƒ††—”‹‰™‹–‡”‹‰ ȋ‡’‡†‹‘ƒ†‘” Š‹‘ͳͻͺʹǡ‘• Šƒ†‡’ʹͲͲͶȌǤƒŽŽ‡”ˆ‡ƒŽ‡•ƒ”‡ƒŽ•‘Ž‡••Ž‹‡Ž›–‘ ˆ‘—†ƒ‡•–ȋ‡’‡†‹‘ƒ†‘” Š‹‘ͳͻͺʹǡ‘• Šƒ†‹ ‡•ʹͲͲ͸ȌǤ –Š‡” •–—†‹‡• Šƒ˜‡ ˆ‘—†–Šƒ–„—„Ž‡„‡‡ ‘Ž‘‹‡•’”‘†— ‡•ƒŽŽ‡”™‘”‡”•‹ƒ”‡ƒ•™‹–Šƒƒ‰‡†Š‘‡›„‡‡•ǡ ’”‘„ƒ„Ž› †—‡ –‘ ’‘ŽŽ‡Ȁ‡ –ƒ” • ƒ” ‹–› ȋ ‘—Ž•‘ ƒ† ’ƒ””‘™ ʹͲͲͻǡŽ„‰ƒ‹‡–ƒŽǤʹͲͳͶȌǤ ‹Ž† „‡‡• ˆ‘”ƒ‰‹‰ ‹ ƒ”‡ƒ• ™‹–Š Ž‘™ Ž‡˜‡Ž• ‘ˆ ”‡•‘—” ‡• ƒ› „‡ˆ‘” ‡†–‘ƒ‡Ž‘‰‡” ˆ‘”ƒ‰‹‰–”‹’•–‘‰ƒ–Š‡”ƒ’‘ŽŽ‡Ȁ‡ –ƒ”Ž‘ƒ†Ǥ •‘Ž‹–ƒ”›„‡‡•ǡ‡•–•ƒ”‡Ž‡ˆ–—‰—ƒ”†‡†™Š‡ –Š‡‡•–‹‰ˆ‡ƒŽ‡‹•ˆ‘”ƒ‰‹‰ƒ†Ž‘‰ˆ‘”ƒ‰‹‰–”‹’•ȋ‡‹†‡ŽƒʹͲͲ͸Ȍƒ†Ž‘™”‡•‘—” ‡ Ž‡˜‡Ž•ȋ ‘‘†‡ŽŽʹͲͲ͵ȌŠƒ˜‡„‡‡•Š‘™–‘”‡•—Ž–‹‹ ”‡ƒ•‡† Ž‡’–‘’ƒ”ƒ•‹–‹•‹•‘Ž‹–ƒ”› „‡‡•Ǥ ”‡˜‹‘—• •–—†‹‡• ‹˜‡•–‹‰ƒ–‹‰ –Š‡ ’‘–‡–‹ƒŽ ‹’ƒ – ‘ˆ Š‘‡› „‡‡• ‘ ™‹Ž† „‡‡ ‘—‹–‹‡• Šƒ˜‡ ƒŽ•‘ ˆ‘—† ™‹Ž† „‡‡ ƒ„—†ƒ ‡ –‘ „‡ Ž‘™‡” ‡ƒ” ƒ’‹ƒ”‹‡• ȋ ‘”—’ ƒ† ‡‘–– ʹͲͲͷǡ Š‘•‘ ʹͲͲ͸Ȍǡ „—–‘–Š‡”•Šƒ˜‡‘–ȋ–‡ˆˆƒǦ‡™‡–‡” ƒ† • Šƒ”–‡ ʹͲͲͲǡ ‘—„‹ ƒ† ‘Ž†ƒ ʹͲͲͳȌǤ  –Š‡ ‘–Š‡” Šƒ†ǡ ƒ† ‹ ƒ‰”‡‡‡– ™‹–Š ‘–Š‡” •–—†‹‡• ȋ–‡ˆˆƒǦ‡™‡–‡” ƒ† • Šƒ”–‡ ʹͲͲͲǡ ‘—„‹ ƒ† ‘Ž†ƒ ʹͲͲͳǡ ‘”—’ ƒ† ‡‘–– ʹͲͲͷȌǡ„‡‡”‹ Š‡••™ƒ•‘–‹ˆŽ—‡ ‡†„›’”‘š‹‹–›–‘ƒ’‹ƒ”‹‡•‹‘—”•–—†›Ǥ

ƒ††‹–‹‘–‘Š‘‡›„‡‡†‡•‹–›ǡ„‡‡ƒ„—†ƒ ‡ƒ†„‹‘ƒ••ƒ› ƒŽ•‘ „‡ ‹ˆŽ—‡ ‡† „› ˆŽ‘™‡”ƒ„—†ƒ ‡ƒ††‹•–”‹„—–‹‘Ǥ—”‘†‡Ž••Š‘™–Šƒ–CistusˆŽ‘™‡”•Šƒ˜‡ƒ‹’‘”–ƒ– ”‘Ž‡•–”— –—”‹‰–Š‡ ƒ””ƒˆ„‡‡ ‘—‹–›ǤŠ‡–Š”‡‡Cistus•’‡ ‹‡•‘ —””‹‰‹–Š‡’ƒ” „Ž‘‘ ‹ ’”‹Žǡ ƒ– ƒ –‹‡ ™Š‡ ™‹Ž† „‡‡ ƒ„—†ƒ ‡ ƒ† †‹˜‡”•‹–›ƒ”‡Š‹‰Šǡƒ†ˆŽ‘™‡”  67 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

”‡•‘—” ‡• •Š‘™ ƒ •–”‘‰ †‡ Ž‹‡ ƒˆ–‡” –Š‡ „Ž‘‘‹‰ ’‡”‹‘† ‘ˆR. officinalisƒ†T. vulgaris ȋ‘• Š‡–ƒŽǤʹͲͲͻǡ ‹Ž‡ŽŽƒ‡–ƒŽǤʹͲͳ͵ǡ Ž‘‡–ƒŽǤ•—„‹––‡†ȌǤ–Š‡”’Žƒ–•„Ž‘‘‹‰ƒ––Š‹•–‹‡ ƒ”‡ ‡‹–Š‡” ˜‡”› • ƒ” ‡ ȋGladiolus illyricusǡ Orobanche latisquamaȌǡ ‘” ’”‘†— ‡ •ƒŽŽ‡” ƒ‘—–•‘ˆ’‘ŽŽ‡ƒ†‡ –ƒ”ȋD.penthaphyllumȌȋ Ž‘‡–ƒŽǤ•—„‹––‡†ȌǤ”‡˜‹‘—••–—†‹‡•‹ –Š‡•ƒ‡ƒ”‡ƒŠƒ˜‡•Š‘™–Šƒ–C. albidusƒ†C. salvifolius ‘•–‹–—–‡ ƒ Š—„ ‹ –Š‡ ƒ””ƒˆ ’‘ŽŽ‹ƒ–‹‘ ‡–™‘”ǡ ƒ––”ƒ –‹‰ Š‹‰Š‡” —„‡”• ‘ˆ ’‘ŽŽ‹ƒ–‘” •’‡ ‹‡• ƒ† ”‡ ‡‹˜‹‰ Š‹‰Š‡” ˆŽ‘™‡”˜‹•‹–ƒ–‹‘”ƒ–‡•–Šƒƒ›‘–Š‡”’Žƒ–•’‡ ‹‡•ȋ‘• Š‡–ƒŽǤʹͲͲͻǡƒ†•—„‹––‡†ȌǤ

—”•–—†›’”‘˜‹†‡•‡˜‹†‡ ‡–‘•—’’‘”––Š‡Š›’‘–Š‡•‹•–Šƒ–Š‹‰Š †‡•‹–‹‡• ‘ˆ ƒƒ‰‡† Š‘‡›„‡‡•Šƒ˜‡ƒ‡‰ƒ–‹˜‡‹’ƒ –‘™‹Ž†„‡‡ ‘—‹–‹‡•Ǥ—””‡•—Ž–•’‘‹––‘’‘ŽŽ‡Ǧ ‡ –ƒ”†‡’Ž‡–‹‘ƒ•ƒ‡ Šƒ‹•‡š’Žƒ‹‹‰–Š‹•‡‰ƒ–‹˜‡‹’ƒ –Ǥ‘‘—”‘™Ž‡†‰‡ǡ–Š‹•‹• –Š‡ ˆ‹”•– –‹‡ ˆŽ‘™‡” ”‡•‘—” ‡ ‘•—’–‹‘ Šƒ• „‡‡ ‡ƒ•—”‡† ‹•–—†‹‡•‡š’Ž‘”‹‰–Š‡ ’‘–‡–‹ƒŽ ‡ˆˆ‡ –• ‘ˆ ƒƒ‰‡† Š‘‡› „‡‡• ‘ ™‹Ž† ’‘ŽŽ‹ƒ–‘”•Ǥ ‘ ‘ˆ‹” ‘” ”‡ˆ—–‡ –Š‹• Š›’‘–Š‡•‹•ǡˆ—–—”‡•–—†‹‡••Š‘—Ž†‹ Ž—†‡Ž‘‰Ǧ–‡”‘‹–‘”‹‰‘ˆ™‹Ž†„‡‡’‘’—Žƒ–‹‘•ƒ† †‹”‡ –‡ƒ•—”‡•‘ˆˆ‹–‡••Ǥ ”‘ƒƒ’’Ž‹‡†’‡”•’‡ –‹˜‡ǡ†‡ ‹•‹‘•‘–Š‡—„‡”‘ˆŠ‹˜‡• ƒŽŽ‘™‡†‹ƒ‡˜‹”‘‡–ƒŽŽ›’”‘–‡ –‡†ƒ”‡ƒ•Š‘—Ž†„‡„ƒ•‡†‘–Š‡ ƒ””›‹‰ ƒ’ƒ ‹–›‘ˆ –Š‡ˆŽ‘™‡” ‘—‹–› ƒ––Š‡Žƒ†• ƒ’‡Ž‡˜‡ŽǤ ‘™‡˜‡”ǡ–‘’”‘˜‹†‡ƒ”ƒ‰‡‘ˆƒ’’”‘’”‹ƒ–‡ Š‹˜‡†‡•‹–‹‡•‹•‡š–”‡‡Ž›†‹ˆˆ‹ —Ž–ˆ‘”•‡˜‡”ƒŽ”‡ƒ•‘•Ǥ ‹”•–ǡ‡˜‡‹ƒƒ–—”ƒŽŠƒ„‹–ƒ–•— Š ƒ• –Š‡ ƒ””ƒˆ ƒ–—”ƒŽ ƒ”ǡ ˆŽ‘™‡” •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‹• ˆƒ” ˆ”‘ Š‘‘‰‡‡‘—• ȋ‘”±Ǧ ‘‰—‡”ƒ ‡– ƒŽǤ ʹͲͳͶȌǤ ‡ ‘†ǡ ƒ˜ƒ‹Žƒ„‹Ž‹–› ‘ˆ ˆŽ‘™‡” ”‡•‘—” ‡• Šƒ‰‡• †”ƒƒ–‹ ƒŽŽ› –Š”‘—‰Š‘—––Š‡•‡ƒ•‘ƒ†ˆ”‘›‡ƒ”–‘›‡ƒ”ȋ Ž‘‡–ƒŽǤ•—„‹––‡†ȌǤŠ‹”†ǡˆ‘”ƒ‰‹‰”ƒ‰‡•‘ˆ Š‘‡›„‡‡••’ƒ•‡˜‡”ƒŽ‹Ž‘‡–‡”•ȋ‹•• Š‡”ƒ†‡‡Ž‡›ͳͻͺʹȌǤ ‘—”–Šǡ”‡•‘—” ‡†‡’Ž‡–‹‘ ƒ› ƒŽ•‘ †‡’‡† ‘ –Š‡ ƒ„—†ƒ ‡ ‘ˆ ”‡•‹†‡– ’‘ŽŽ‹ƒ–‘” ’‘’—Žƒ–‹‘•Ǥ ‘‡–Š‡Ž‡••ǡ ‘—” •–—†› •—‰‰‡•–• –Šƒ– ƒ– †‡•‹–‹‡• ‘˜‡” ͵Ǥͷ Š‹˜‡• ’‡” ʹȋͶ͹ͷŠ‹˜‡•Ȁͳ͵ͶʹȌǡ ™‹Ž† „‡‡ ‘—‹–‹‡•ƒ”‡Ž‹‡Ž›–‘„‡ƒˆˆ‡ –‡†‹‘—”•–—†›ƒ”‡ƒǤ



Epilogue

Š‡ ƒ””ƒˆƒ–—”ƒŽƒ”‹•’ƒ”–‹ƒŽŽ›Ž‘ ƒ–‡†‹–Š‡—‹ ‹’ƒŽ‹–›‘ˆŽ‹˜‡ŽŽƒǤ ƒ›ʹͲͳʹǡ–Š‡ ‹–› ‘— ‹Ž†‹• —••‡†ƒ’‡–‹–‹‘–‘‹•–ƒŽŽ͵ͷ͹‡™Š‘‡›„‡‡Š‹˜‡•‹–Š‡ƒ”ǤŠ‡ ‘— ‹Ž ‡šƒ‹‡†ƒ”‡’‘”– ‘‹••‹‘‡†„›–Š‡„‘ƒ”†‘ˆ†‹”‡ –‘”•‘ˆ–Š‡ƒ” ƒ—–‹‘‹‰ƒ„‘—––Š‡ ’‘–‡–‹ƒŽ‡ˆˆ‡ –•‘ˆ‹–‡•‹˜‡„‡‡‡‡’‹‰‘‘–Š‡”’‘ŽŽ‹ƒ–‘”•ǤŠ‡ ‘— ‹Žˆ‹ƒŽŽ›ƒ’’”‘˜‡†

 68 Šƒ’–‡”ʹ

–Š‡ ‹•–ƒŽŽƒ–‹‘ ‘ˆ –Š‡ ͵ͷ͹ Š‹˜‡• „ƒ•‡† ‘ —””‡– Ž‡‰‹•Žƒ–‹‘ ‘•‹†‡”‹‰ „‡‡‡‡’‹‰ ƒ ̶‹‘ —‘—•ƒ –‹˜‹–›̶Ǥ



Aknowledgements

‡ƒ”‡–Šƒˆ—Ž–‘ Ǥƒ””‹ŽǦ ”ƒ‡ŽŽ•ǡ Ǥƒ›ƒǡǤ‹‡”ƒǡǤ‡˜‡”–±ǡǤ— —”—ŽŽǡǤǤ‡“—‡•‡•ǡ Ǥ‘ ‡Žǡ Ǥ”ƒ–ǡǤžœƒ”‘ǡǤ‡˜‘Ž–וǡ ǤǤ‹Ž‡ŽŽƒ•ǡǤƒ•ǡǤ‹Žƒ†±•ǡǤŽƒ˜‹ƒǡǤג‡œǡ Ǥ ‘œžŽ‡œǡƒ†Ǥ—Ó‘œˆ‘”–Š‡‹”Š‡Ž’™‹–Šˆ‹‡Ž†ƒ†Žƒ„‘”ƒ–‘”›™‘”ǡƒ†–‘Ǥ ‡”ž†‡œ ˆ‘”Š‹••–ƒ–‹•–‹ ƒŽƒ†˜‹ ‡ǤǤ—‰‰ƒȋ ǡ ”‡Žƒ†Ȍǡ Ǥ ƒŽŽ‡”ȋ Š™‡‹œ‡”‹• Š‡ƒ–‹‘ƒŽ’ƒ”ǡ ™‹–œ‡”Žƒ†Ȍǡ Ǥ ‡‹‘‡ ȋ‡–•¡ŠƒŽŽ‹–—•ǡ ‹Žƒ†Ȍǡ Ǥ ‘”ƒ‹• ȋ‹†‘• ƒ–‹‘ƒŽ ƒ”ǡ ”‡‡ ‡Ȍǡ Ǥ ƒ‡ ȋ ‘”†„”—•˜‡”‡–ǡ ™‡†‡Ȍǡ Ǥ ‡ŽŽ‡ ȋƒ” ‘ ƒœ‹‘ƒŽ‡ †‡ŽŽǯ•’”‘‘–‡ǡ –ƒŽ›Ȍǡ Ǥ‘””‡–×ȋ‹’—–ƒ ‹×†‡ƒ” ‡Ž‘ƒǡ’ƒ‹Ȍƒ† ǤǤ‹ ‡–‡ȋ  ǡ‘”–—‰ƒŽȌ‹†Ž› ’”‘˜‹†‡†‹ˆ‘”ƒ–‹‘‘„‡‡‡‡’‹‰Ž‡‰‹•Žƒ–‹‘‹–Š‡‹””‡•’‡ –‹˜‡ ‘—–”‹‡•ǤŠ‹••–—†›™ƒ• •—’’‘”–‡†„›–Š‡’ƒ‹•Š‹‹•–”›‘ˆ ‹‡ ‡ƒ† ‘˜ƒ–‹‘ȋ  ǡ’”‘Œ‡ –• ʹͲͲͷǦ ͲͲͶͻͳǡ ʹͲͲͻǦͳʹ͸Ͷ͸ƒ†ʹͲͲͺǦͲͲͶͲǢ  ˆ‡ŽŽ‘™•Š‹’ȋǦʹͲͳͲǦͲͶʹͷʹͲȌ–‘ǤǦǤȌǡ ƒ†„›‹’—–ƒ ‹×†‡ƒ” ‡Ž‘ƒǤǤǤ™ƒ••—’’‘”–‡†„›ƒ ˆ‡ŽŽ‘™•Š‹’ȋʹͲͳʹ ͲͺͲͶͺͶȌ ˆ”‘ ‡‡”ƒŽ‹–ƒ–†‡ƒ–ƒŽ—›ƒǤ

References

”ƒǡǤ‡–ƒŽǤʹͲͳͶǤ ‡ƒŽ‡”‡’”‘†— –‹˜‡•— ‡••‹‰›‘†‹‘‡ ‹‘—•Thymusvulgarisǣ‘ŽŽ‡˜‡”•—• —–”‹‡–Ž‹‹–ƒ–‹‘ƒ†’‘ŽŽ‹ƒ–‘”ˆ‘”ƒ‰‹‰„‡Šƒ˜‹‘—”ǤǦ‘–Ǥ Ǥ‹Ǥ‘ Ǥͳ͹ͷǣ͵ͻͷȂͶͲͺǤ

‘• Šǡ ǤʹͲͲͺǤ”‘†— –‹‘‘ˆ—†‡”•‹œ‡†‘ˆˆ•’”‹‰‹ƒ•‘Ž‹–ƒ”›„‡‡ǤǦ‹Ǥ‡Šƒ˜Ǥ͹ͷǣͺͲͻȂͺͳ͸Ǥ

‘• Šǡ Ǥƒ†‹ ‡•ǡǤʹͲͲʹǤ‘†›•‹œ‡ƒ•ƒ‡•–‹ƒ–‘”‘ˆ’”‘†— –‹‘ ‘•–•‹ƒ•‘Ž‹–ƒ”›„‡‡ǤǦ ‘ŽǤ –‘‘ŽǤʹ͹ǣͳʹͻȂͳ͵͹Ǥ

‘• Šǡ Ǥƒ†‡’ǡǤǤʹͲͲͶǤˆˆ‡ –‘ˆ’”‡Ǧ™‹–‡”‹‰ƒ†™‹–‡”‹‰–‡’‡”ƒ–—”‡”‡‰‹‡•‘™‡‹‰Š– Ž‘••ǡ•—”˜‹˜ƒŽǡƒ†‡‡”‰‡ ‡–‹‡‹–Š‡ƒ•‘„‡‡Osmia cornutaȋ ›‡‘’–‡”ƒǣ ‡‰ƒ Š‹Ž‹†ƒ‡ȌǤǦ’‹†‘Ž‘‰‹‡͵ͷǣͶ͸ͻȂͶ͹ͻǤ

 69 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

‘• Šǡ Ǥ ƒ† ‹ ‡•ǡ Ǥ ʹͲͲ͸Ǥ ‡Žƒ–‹‘•Š‹’ „‡–™‡‡ „‘†› •‹œ‡ǡ ’”‘˜‹•‹‘‹‰ ”ƒ–‡ǡ Ž‘‰‡˜‹–› ƒ† ”‡’”‘†— –‹˜‡•— ‡••‹ˆ‡ƒŽ‡•‘ˆ–Š‡•‘Ž‹–ƒ”›„‡‡OsmiacornutaǤǦ‡Šƒ˜Ǥ ‘ŽǤ‘ ‹‘„‹‘ŽǤ͸Ͳǣ ʹ͸Ȃ͵͵Ǥ

‘• Šǡ Ǥ‡–ƒŽǤʹͲͲͻǤŽƒ–Ǧ’‘ŽŽ‹ƒ–‘”‡–™‘”•ǣƒ††‹‰–Š‡’‘ŽŽ‹ƒ–‘”ǯ•’‡”•’‡ –‹˜‡ǤǦ ‘ŽǤ‡––Ǥͳʹǣ ͶͲͻȂͳͻǤ

›‡”•ǡ Ǥ Ǥ ʹͲͲͲǤ ‘’‡–‹–‹‘ „‡–™‡‡ –™‘ ‡•–—ƒ”‹‡ •ƒ‹Ž•ǣ ‹’Ž‹ ƒ–‹‘• ˆ‘” ‹˜ƒ•‹‘• ‘ˆ ‡š‘–‹  •’‡ ‹‡•ǤǦ ‘Ž‘‰›ͺͳǣͳʹʹͷȂͳʹ͵ͻǤ

ƒ„”ƒǡ ǤʹͲͲͺǤ Ž‘”ƒ‡ŽȉŽÀˆ‡”ƒ†‡ƒ–ƒŽ—›ƒǤǦ ǣ’‹ —Ž–—”ƒ‹’”‘†— ‹×†‡‡Žƒƒ–ƒŽ—›ƒǤ‘••‹‡” –° ‹ Ǥ ‡‡”ƒŽ‹–ƒ–†‡ƒ–ƒŽ—›ƒǤ‡’ƒ”–ƒ‡–†ǯ‰”‹ —Ž–—”ƒǡŽ‹‡–ƒ ‹×‹ ‹×—”ƒŽǡ’’ǤʹͲǤ

ƒ•–‡ŽŽƒ‘•ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲʹǤ ›ƒ‹  ‡ –ƒ” ”‡’Ž‡‹•Š‡– ‹ˆŽ‘™‡”•‘ˆ‡•–‡‘ ȋ ”‘’Š—Žƒ”‹ƒ ‡ƒ‡ȌǤǦǤ Ǥ‘–ǤͺͻǣͳͳͳȂͳͳͺǤ

”ƒ‡ǡǤͳͻͻͲǤ‡‡•ƒ†„‡‡‡‡’‹‰ǣ• ‹‡ ‡ǡ’”ƒ –‹ ‡ƒ†™‘”Ž†”‡•‘—” ‡•ǤǦ ‡‹‡ƒ‡™‡•Ǥ

ƒ˜‹Žƒǡ Ǥ Ǥ ƒ† ƒ”†Ž‡ǡ Ǥ Ǥ ʹͲͲͺǤ ƒ”‹ƒ–‹‘ ‹ ƒ–‹˜‡ ’‘ŽŽ‹ƒ–‘”• ‹ –Š‡ ƒ„•‡ ‡ ‘ˆ Š‘‡›„‡‡•ǣ ‹’Ž‹ ƒ–‹‘• ˆ‘” ”‡’”‘†— –‹˜‡ •— ‡•• ‘ˆ ƒ —•–”ƒŽ‹ƒ ‰‡‡”ƒŽ‹•–Ǧ’‘ŽŽ‹ƒ–‡† Š‡”„ Trachymene incisaȋ’‹ƒ ‡ƒ‡ȌǤǦ‘–Ǥ Ǥ‹Ǥ‘ Ǥͳͷ͸ǣͶ͹ͻȂͶͻͲǤ

—†Ž‡›ǡǤʹͲͲͺǤ —‹†‡Ž‹‡•ˆ‘”ƒ’’Ž›‹‰’”‘–‡ –‡†ƒ”‡ƒƒƒ‰‡‡– ƒ–‡‰‘”‹‡• ȋ —†Ž‡›ǡ †ǤȌǤ Ǧ Ǥ

Š”‡ˆ‡Ž†ǡ Ǥ ǤʹͲͳͲǤ ‘•›•–‡ ‘•‡“—‡ ‡•‘ˆ„‹‘Ž‘‰‹ ƒŽ‹˜ƒ•‹‘•ǤǦ—Ǥ‡˜Ǥ ‘ŽǤ˜‘ŽǤ›•–ǤͶͳǣ ͷͻȂͺͲǤ

Ž„‰ƒ‹ǡ Ǥ ‡– ƒŽǤ ʹͲͳͶǤ Š‡ ‡ˆˆ‡ – ‘ˆ ’”‘š‹‹–› –‘ ƒ Š‘‡›„‡‡ ƒ’‹ƒ”› ‘ „—„Ž‡„‡‡ ‘Ž‘› ˆ‹–‡••ǡ †‡˜‡Ž‘’‡–ǡƒ†’‡”ˆ‘”ƒ ‡ǤǦ’‹†‘Ž‘‰‹‡ͶͷǣͷͲͶȂͷͳ͵Ǥ

‹Ž‡ŽŽƒǡ Ǥ‡–ƒŽǤʹͲͳ͵Ǥ Ž‘”ƒŽƒ†˜‡”–‹•‡‡–• ‡–‹ƒ Šƒ‰‹‰’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘”•ƒ”‡–ǤǦ ‹Ǥ‡’Ǥ͵ǣ ͵Ͷ͵ͶǤ

‘”—’ǡ Ǥ Ǥ ƒ† ‡‘––ǡ Ǥ ʹͲͲͷǤ Š‡ ”‡Žƒ–‹‘•Š‹’ „‡–™‡‡ –Š‡ƒ„—†ƒ ‡•‘ˆ„—„Ž‡„‡‡•ƒ† Š‘‡›„‡‡•‹ƒƒ–‹˜‡Šƒ„‹–ƒ–ǤǦ ‘ŽǤ–‘‘ŽǤ͵ͲǣͶ͹Ȃͷ͹Ǥ

‘”—’ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͺǤ Š‡ ”‡•–‘”ƒ–‹‘ ‘ˆ ‡ ‘Ž‘‰‹ ƒŽ ‹–‡”ƒ –‹‘•ǣ ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘” ‡–™‘”• ‘ ƒ ‹‡–ƒ†”‡•–‘”‡†Š‡ƒ–ŠŽƒ†•ǤǦ Ǥ’’ŽǤ ‘ŽǤͶͷǣ͹ͶʹȂ͹ͷʹǤ

 70 Šƒ’–‡”ʹ

ƒ”‹„ƒŽ†‹ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͳ͵Ǥ ‹Ž† ’‘ŽŽ‹ƒ–‘”• ‡Šƒ ‡ ˆ”—‹– •‡–‘ˆ ”‘’•”‡‰ƒ”†Ž‡••‘ˆŠ‘‡›„‡‡ ƒ„—†ƒ ‡ǤǦ ‹‡ ‡͵͵ͻǣͳ͸ͲͺȂͳͳǤ

ƒ–ŠƒǡǤƒ†• Šƒ”–‡ǡǤʹͲͲʹǤ ‘”ƒ‰‹‰”ƒ‰‡•‘ˆ•‘Ž‹–ƒ”›„‡‡•ǤǦ Ǥ‹Ǥ ‘ŽǤ͹ͳǣ͹ͷ͹Ȃ͹͸ͶǤ

‘‘†‡ŽŽǡ Ǥ ʹͲͲ͵Ǥ ‘‘† ƒ˜ƒ‹Žƒ„‹Ž‹–› ƒˆˆ‡ –• Osmia pumilaȋ ›‡‘’–‡”ƒǣ‡‰ƒ Š‹Ž‹†ƒ‡Ȍˆ‘”ƒ‰‹‰ǡ ”‡’”‘†— –‹‘ǡƒ†„”‘‘†’ƒ”ƒ•‹–‹•ǤǦ‡ ‘Ž‘‰‹ƒͳ͵ͶǣͷͳͺȂʹ͹Ǥ

‘—Ž•‘ǡǤʹͲͲ͵Ǥˆˆ‡ –•‘ˆ‹–”‘†— ‡†„‡‡•‘ƒ–‹˜‡‡ ‘•›•–‡•ǤǦ—Ǥ‡˜Ǥ ‘ŽǤ˜‘ŽǤ›•–Ǥ͵ͶǣͳȂ ʹ͸Ǥ

‘—Ž•‘ǡǤƒ†’ƒ””‘™ǡǤǤʹͲͲͻǤ˜‹†‡ ‡ˆ‘” ‘’‡–‹–‹‘„‡–™‡‡Š‘‡›„‡‡•ƒ†„—„Ž‡„‡‡•Ǣ ‡ˆˆ‡ –•‘„—„Ž‡„‡‡™‘”‡”•‹œ‡ǤǦ Ǥ •‡ –‘•‡”˜Ǥͳ͵ǣͳ͹͹ȂͳͺͳǤ

”‡‡Ž‡ƒˆǡǤǤ‡–ƒŽǤʹͲͲ͹Ǥ‡‡ˆ‘”ƒ‰‹‰”ƒ‰‡•ƒ†–Š‡‹””‡Žƒ–‹‘•Š‹’–‘„‘†›•‹œ‡ǤǦ‡ ‘Ž‘‰‹ƒͳͷ͵ǣ ͷͺͻȂͻ͸Ǥ

‡‹”‹ ŠǡǤͳͻ͹ͷǤ‡”‰‡–‹ •‘ˆ’‘ŽŽ‹ƒ–‹‘ǤǦ—Ǥ‡˜Ǥ ‘ŽǤ›•–Ǥ͸ǣͳ͵ͻȂͳ͹ͲǤ

‡‹”‹ Šǡ Ǥ ͳͻ͹ͻǤ ‡•‘—” ‡ Š‡–‡”‘‰‡‡‹–› ƒ† ’ƒ––‡”• ‘ˆ ‘˜‡‡– ‹ ˆ‘”ƒ‰‹‰ „—„Ž‡„‡‡•Ǥ Ǧ ‡ ‘Ž‘‰‹ƒͶͲǣʹ͵ͷȂʹͶͷǤ

‘Ž™ƒ›ǡ Ǥ Ǥ ͳͻͻͻǤ ‘’‡–‹–‹˜‡ ‡ Šƒ‹•• —†‡”Ž›‹‰ –Š‡ †‹•’Žƒ ‡‡– ‘ˆ ƒ–‹˜‡ ƒ–• „› –Š‡ ‹˜ƒ•‹˜‡”‰‡–‹‡ƒ–ǤǦ ‘Ž‘‰›ͺͲǣʹ͵ͺȂʹͷͳǤ

—†‡™‡œǡǤƒ†Ž‡‹ǡǤǦǤʹͲͳ͵Ǥ‘’‡–‹–‹‘„‡–™‡‡Š‘‡›„‡‡•ƒ†™‹Ž†„‡‡•ƒ†–Š‡”‘Ž‡‘ˆ ‡•–‹‰”‡•‘—” ‡•‹ƒƒ–—”‡”‡•‡”˜‡ǤǦ Ǥ •‡ –‘•‡”˜Ǥͳ͹ǣͳʹ͹ͷȂͳʹͺ͵Ǥ

—ƒǡǤ Ǥƒ† ‘”†‘ǡǤǤͳͻͻ͸Ǥš’Ž‘‹–ƒ–‹‘ƒ†‹–‡”ˆ‡”‡ ‡ ‘’‡–‹–‹‘„‡–™‡‡–Š‡‹˜ƒ•‹˜‡ ”‰‡–‹‡ƒ–ǡLinepithemahumileǡƒ†ƒ–‹˜‡ƒ–•’‡ ‹‡•ǤǦ‡ ‘Ž‘‰‹ƒͳͲͷǣͶͲͷȂͶͳʹǤ

”‘•ǡǤǤ‡–ƒŽǤʹͲͲ͹Ǥ‡†— ‡† ‘†‹–‹‘ˆƒ –‘”‘ˆ–™‘ƒ–‹˜‡ˆ‹•Š•’‡ ‹‡• ‘‹ ‹†‡–™‹–Š‹˜ƒ•‹‘‘ˆ ‘Ǧƒ–‹˜‡•‹ƒ ƒ”’•‹–Š‡ ŽŽ‹‘‹•‹˜‡”ǡǤǤǤ •–Š‹•‡˜‹†‡ ‡ˆ‘” ‘’‡–‹–‹‘ƒ†”‡†— ‡† ˆ‹–‡••ǫǦ Ǥ ‹•Š‹‘ŽǤ͹ͳǣʹͷͺȂʹ͹͵Ǥ

ƒˆˆ±ǡǤ‡–ƒŽǤʹͲͲͻǤ•–‹ƒ–‹‰–Š‡†‡•‹–›‘ˆŠ‘‡›„‡‡ ‘Ž‘‹‡•ƒ ”‘••–Š‡‹”ƒ–—”ƒŽ”ƒ‰‡–‘ˆ‹ŽŽ–Š‡ ‰ƒ’‹’‘ŽŽ‹ƒ–‘”†‡ Ž‹‡ ‡•—•‡•ǤǦ‘•‡”˜Ǥ‹‘ŽǤʹͶǣͷͺ͵Ȃͻ͵Ǥ

 71 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

ƒ‘„•‡ǡ Ǥ Ǥ ƒ† ”‹•–Œž••‘ǡ Ǥ Ǥ ͳͻͻͶǤ ˆŽ—‡ ‡ ‘ˆ –‡’‡”ƒ–—”‡ ƒ† ˆŽ‘”‡– ƒ‰‡ ‘ ‡ –ƒ” •‡ ”‡–‹‘‹TrifoliumrepensǤǦǤ‘–Ǥ͹Ͷǣ͵ʹ͹Ȃ͵͵ͶǤ

ƒ‹•‡”Ǧ—„—”›ǡǤǤ‡–ƒŽǤʹͲͲͻǤ‘—‹–›•–”— –—”‡‘ˆ’‘ŽŽ‹ƒ–‹‘™‡„•‘ˆƒ—”‹–‹ƒŠ‡ƒ–ŠŽƒ† Šƒ„‹–ƒ–•ǤǦ‡”•’‡ –ǤŽƒ– ‘ŽǤ˜‘ŽǤ›•–ǤͳͳǣʹͶͳȂʹͷͶǤ

‡˜‹‡ǡ ǤǤ‡–ƒŽǤʹͲͲ͵Ǥ‡ Šƒ‹••—†‡”Ž›‹‰–Š‡‹’ƒ –•‘ˆ‡š‘–‹ ’Žƒ–‹˜ƒ•‹‘•ǤǦ”‘ Ǥ‹‘ŽǤ ‹Ǥ ʹ͹Ͳǣ͹͹ͷȂͺͳǤ

ג‡œ”ƒ–ǡǤ‡–ƒŽǤʹͲͲ͹Ǥ•‡‘ˆ ’”‘–‡ –‡†ƒ”‡ƒ•ƒƒ‰‡‡– ƒ–‡‰‘”‹‡•‹–Š‡‡†‹–‡””ƒ‡ƒ ”‡‰‹‘ȋג‡œ”ƒ–ǡ‘•‡›±•ǡƒ†‘‰—‡”ƒǡ†•ǤȌǤǦ‘•‡Œ‡”Àƒ†‡‡†‹‘„‹‡–‡‘ˆ —–ƒ†‡†ƒŽ— Àƒǡ‡˜‹ŽŽƒǡ’ƒ‹ƒ† ǡ Žƒ†ǡ™‹–œ‡”Žƒ†ƒ†ƒŽƒ‰ƒǡ’ƒ‹Ǥ

òŽŽ‡”ǡ Ǥ ‡– ƒŽǤ ʹͲͲ͸Ǥ —ƒ–‹–ƒ–‹˜‡ ’‘ŽŽ‡ ”‡“—‹”‡‡–• ‘ˆ •‘Ž‹–ƒ”› „‡‡•ǣ ’Ž‹ ƒ–‹‘• ˆ‘” „‡‡ ‘•‡”˜ƒ–‹‘ƒ†–Š‡‡˜‘Ž—–‹‘‘ˆ„‡‡ȂˆŽ‘™‡””‡Žƒ–‹‘•Š‹’•ǤǦ‹‘ŽǤ‘•‡”˜Ǥͳ͵Ͳǣ͸ͲͶȂ͸ͳͷǤ

”†ƒ‘ǡ Ǥ ƒ† ”‡Žƒ•ǡ Ǥ Ǥ ʹͲͲͶǤ ‡‡”‘—•ǦŽ‹‡ ˆŽ‘™‡”•ǣ ‡ –ƒ”’”‘†— –‹‘‹–™‘‡’‹’Š›–‹  „”‘‡Ž‹ƒ†•ƒ†ƒ‡–ƒǦƒƒŽ›•‹•‘ˆ”‡‘˜ƒŽ‡ˆˆ‡ –•ǤǦ‡ ‘Ž‘‰‹ƒͳͶͲǣͶͻͷȂͷͲͷǤ

ƒ ‹‹ǡǤ‡–ƒŽǤʹͲͲ͵Ǥ‡ –ƒ”„‹‘†‹˜‡”•‹–›ǣ•Š‘”–”‡˜‹‡™ǤǦŽƒ–›•–Ǥ˜‘ŽǤʹ͵ͺǣ͹ȂʹͳǤ

‡–ƒ‹†‘—ǡǤƒ†‡–•ǡǤͳͻͻ͸Ǥ‘‡•–‡’‡”ƒ–—”‡•–”‡••‹†— ‡‡ –ƒ”•‡ ”‡–‹‘‹‡†‹–‡””ƒ‡ƒ ’Žƒ–•ǫǦ‡™Š›–‘ŽǤͳ͵͵ǣͷͳ͵ȂͷͳͺǤ

‡–‡”•‘ǡ Ǥ Ǥƒ†‘‹–„‡”‰ǡǤǤʹͲͲ͸Ǥ ’ƒ –•‘ˆˆŽ‹‰Š–†‹•–ƒ ‡‘•‡š”ƒ–‹‘ƒ†”‡•‘—” ‡ƒŽŽ‘ ƒ–‹‘ –‘‘ˆˆ•’”‹‰‹–Š‡Ž‡ƒˆ —––‡”„‡‡ǡMegachilerotundataǤǦ‡Šƒ˜Ǥ ‘ŽǤ‘ ‹‘„‹‘ŽǤͷͻǣͷͺͻȂͷͻ͸Ǥ

‡–”‡ǡ Ǥ ƒ† ƒ•‡ǡ Ǥ Ǥ ͳͻͻ͸Ǥ  ‡š’‡”‹‡–ƒŽ †‡‘•–”ƒ–‹‘ ‘ˆ ‡š’Ž‘‹–ƒ–‹‘ ‘’‡–‹–‹‘ ‹ ƒ ‘‰‘‹‰‹˜ƒ•‹‘ǤǦ ‘Ž‘‰›͹͹ǣͳͳͺȂͳ͵ʹǤ

‹—•Ǧ‡†‘ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͷǤ Ž‘”ƒŽ ”‡•‘—” ‡ —•‡ ƒ† ‹–‡”ƒ –‹‘• „‡–™‡‡ Apis melliferaƒ† ƒ–‹˜‡„‡‡•‹ — —”„‹– ”‘’•‹— ƒ–žǡ±š‹ ‘ǤǦƒǤ–‘‘ŽǤͳ͵͹ǣͶͶͳȂͶͶͻǤ

‘•ǡǤʹͲͳͶǤ‹”ƒ‘Ǥ ‡‘‰”ƒ’Š‹  ˆ‘”ƒ–‹‘›•–‡ƒ†‡‘–‡‡•‹‰‘ˆ–™ƒ”‡ǤǦ ‡‘‰”Ǥ ˆǤ ›•–Ǥ‡‘–‡‡•Ǥ‘ˆ–™Ǥ

‘”‡‡ƒʹͲͳͶǤǣŽƒ‰—ƒ‰‡ƒ†‡˜‹”‘‡–ˆ‘”•–ƒ–‹•–‹ ƒŽ ‘’—–‹‰Ǥ

 72 Šƒ’–‡”ʹ

‹„‡‹”‘ǡ Ǥ Ǥ ͳͻͻͶǤ ”‘™–Š ‹ „—„Ž‡ „‡‡ Žƒ”˜ƒ‡ǣ ”‡Žƒ–‹‘ „‡–™‡‡ †‡˜‡Ž‘’‡– –‹‡ǡ ƒ••ǡ ƒ† ƒ‘—–‘ˆ’‘ŽŽ‡‹‰‡•–‡†ǤǦƒǤ Ǥ‘‘ŽǤ͹ʹǣͳͻ͹ͺȂͳͻͺͷǤ

‘—„‹ǡǤǤͳͻ͹ͺǤ‘’‡–‹–‹˜‡‹–‡”ƒ –‹‘•„‡–™‡‡‡‘–”‘’‹ ƒŽ’‘ŽŽ‹ƒ–‘”•ƒ†ƒˆ”‹ ƒ‹œ‡†Š‘‡› „‡‡•ǤǦ ‹‡ ‡ʹͲͳǣͳͲ͵ͲȂͳͲ͵ʹǤ

‘—„‹ǡ Ǥ Ǥ ƒ† ‘Ž†ƒǡ Ǥ ʹͲͲͳǤ ‘ ‘’‡–‹‰ Š‘‡› „‡‡• ƒ––‡”ǫ ›ƒ‹ •ƒ†ƒ„—†ƒ ‡ ‘ˆ ƒ–‹˜‡„‡‡•„‡ˆ‘”‡ƒ†ƒˆ–‡”Š‘‡›„‡‡‹˜ƒ•‹‘ǤǦ‘’—ŽǤ ‘ŽǤͶ͵ǣͷ͵Ȃ͸ʹǤ

ƒ’•‘ǡǤ Ǥ‡–ƒŽǤʹͲͲͺǤ‹‡–‘˜‡”Žƒ’ƒ‘‰–™‘•‹ƒ ƒ”’ƒ†–Š”‡‡ƒ–‹˜‡ˆ‹•Š‡•‹„ƒ ™ƒ–‡” Žƒ‡•‘–Š‡ ŽŽ‹‘‹•ƒ†‹••‹••‹’’‹”‹˜‡”•ǤǦ‹‘ŽǤ ˜ƒ•‹‘•ͳͳǣͶͺ͵ȂͶͻ͸Ǥ

‡‡Ž‡›ǡǤǤͳͻͺͷǤ ‘‡›„‡‡‡ ‘Ž‘‰›ǣƒ•–—†›‘ˆƒ†ƒ’–ƒ–‹‘‹•‘ ‹ƒŽŽ‹ˆ‡ǤǦ”‹ ‡–‘‹˜‡”•‹–›”‡••Ǥ

‡‹†‡Žƒǡ Ǥ ʹͲͲ͸Ǥ ’‡Ǧ ‡ŽŽ ’ƒ”ƒ•‹–‹• •Šƒ’‡• ƒ–‡”ƒŽ ‹˜‡•–‡– ’ƒ––‡”• ‹ –Š‡ ‡† ƒ•‘ „‡‡OsmiarufaǤǦ‡Šƒ˜Ǥ ‘ŽǤͳ͹ǣͺ͵ͻȂͺͶͺǤ

–‡ˆˆƒǦ‡™‡–‡”ǡ Ǥ ƒ† • Šƒ”–‡ǡ Ǥ ʹͲͲͲǤ ‡•‘—” ‡ ‘˜‡”Žƒ’ ƒ† ’‘••‹„Ž‡ ‘’‡–‹–‹‘ „‡–™‡‡ Š‘‡›„‡‡•ƒ†™‹Ž†„‡‡•‹ ‡–”ƒŽ—”‘’‡ǤǦ‡ ‘Ž‘‰‹ƒͳʹʹǣʹͺͺȂʹͻ͸Ǥ

–‡™ƒ”–ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲʹǤ ‡’‘”‘•’ƒ–‹ƒŽ †‹•–”‹„—–‹‘• ‘ˆ ‡Žǡ —Ž‡ †‡‡”ǡ ƒ† ƒ––Ž‡ǣ ”‡•‘—” ‡ ’ƒ”–‹–‹‘‹‰ƒ† ‘’‡–‹–‹˜‡†‹•’Žƒ ‡‡–ǤǦ ǤƒƒŽǤͺ͵ǣʹʹͻȂʹͶͶǤ

‡’‡†‹‘ǡǤ Ǥƒ†‘” Š‹‘ǡǤ ǤͳͻͺʹǤŠ‡‘–›’‹ ˜ƒ”‹ƒ„‹Ž‹–›‹‡•–‹‰•— ‡••ƒ‘‰Osmialignaria ’”‘’‹“—ƒˆ‡ƒŽ‡•‹ƒ‰Žƒ••Š‘—•‡‡˜‹”‘‡–ǣȋ ›‡‘’–‡”ƒǣ‡‰ƒ Š‹Ž‹†ƒ‡ȌǤǦ ‘ŽǤ–‘‘ŽǤ ͹ǣͶͷ͵ȂͶ͸ʹǤ

Š‘•‘ǡǤǤʹͲͲͶǤ‘’‡–‹–‹˜‡‹–‡”ƒ –‹‘•„‡–™‡‡–Š‡‹˜ƒ•‹˜‡—”‘’‡ƒŠ‘‡›„‡‡ƒ†ƒ–‹˜‡ „—„Ž‡„‡‡•ǤǦ ‘Ž‘‰›ͺͷǣͶͷͺȂͶ͹ͲǤ

Š‘•‘ǡ Ǥ Ǥ ʹͲͲ͸Ǥ ‡–‡ –‹‰ –Š‡ ‡ˆˆ‡ –• ‘ˆ ‹–”‘†— ‡† •’‡ ‹‡•ǣƒ ƒ•‡•–—†›‘ˆ ‘’‡–‹–‹‘ „‡–™‡‡’‹•ƒ†‘„—•ǤǦ‹‘•ͳͳͶǣͶͲ͹ȂͶͳͺǤ

‹ŽƒǡǤͳͻͺʹǤ‡•‘—” ‡ ‘’‡–‹–‹‘ƒ† ‘—‹–›•–”— –—”‡ǤǦ”‹ ‡–‘‹˜‡”•‹–›”‡••Ǥ

‘”±Ǧ‘‰—‡”ƒǡ Ǥ ‡– ƒŽǤ ʹͲͳͶǤ ‡–‡”‹ƒ–• ‘ˆ •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‹ ƒ „‡‡ ‘—‹–›ǣ ‡•–‹‰ ”‡•‘—” ‡•ǡˆŽ‘™‡””‡•‘—” ‡•ǡƒ†„‘†›•‹œ‡ǤǦ‘‡ͻǣ‡ͻ͹ʹͷͷǤ

 73 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

ƒŽ‹†‘ǡǤ‡–ƒŽǤʹͲͳͶǤ ’ƒ –‘†‡Žƒ‹–”‘†— ‹×†‡Žƒƒ„‡Œƒ†‘±•–‹ ƒȋApismelliferaǡ’‹†ƒ‡Ȍ‡‡Ž ƒ”“—‡ƒ ‹‘ƒŽ†‡Ž‡‹†‡ȋ‡‡”‹ˆ‡ǡ •Žƒ•ƒƒ”‹ƒ•ȌǤǦ ‘•‹•–‡ƒ•ʹ͵ǣͷͺȂ͸͸Ǥ

‹•• Š‡”ǡ Ǥ Ǥ ƒ† ‡‡Ž‡›ǡ Ǥ Ǥ ͳͻͺʹǤ ‘”ƒ‰‹‰ •–”ƒ–‡‰› ‘ˆ Š‘‡›„‡‡ ‘Ž‘‹‡• ‹ ƒ –‡’‡”ƒ–‡ †‡ ‹†—‘—•ˆ‘”‡•–ǤǦ ‘Ž‘‰›͸͵ǣͳ͹ͻͲȂͳͺͲͳǤ

‘ ”‹• Šǡ Ǥ ͳͻ͸͹Ǥ Š‡ †ƒ ‡ Žƒ‰—ƒ‰‡ ƒ† ‘”‹‡–ƒ–‹‘ ‘ˆ „‡‡•Ǥ Ǧ ƒ”˜ƒ”† ‹˜‡”•‹–› ”‡••Ǥ ƒ„”‹†‰‡ǡǤ

‡•–’ŠƒŽǡǤ‡–ƒŽǤʹͲͲͺǤ‡ƒ•—”‹‰„‡‡†‹˜‡”•‹–›‹†‹ˆˆ‡”‡–—”‘’‡ƒŠƒ„‹–ƒ–•ƒ†„‹‘‰‡‘‰”ƒ’Š‹ ƒŽ ”‡‰‹‘•ǤǦ ‘ŽǤ‘‘‰”Ǥ͹ͺǣ͸ͷ͵Ȃ͸͹ͳǤ

‘—‰ǡǤǤ‡–ƒŽǤʹͲͲͷǤ‘’‡–‹–‹‘ƒ† ‘’‡•ƒ–‹‘ƒ‘‰ ƒ––Ž‡ǡœ‡„”ƒ•ǡƒ†‡Ž‡’Šƒ–•‹ƒ•‡‹Ǧ ƒ”‹†•ƒ˜ƒƒ‹ƒ‹‹’‹ƒǡ‡›ƒǤǦ‹‘ŽǤ‘•‡”˜Ǥͳʹʹǣ͵ͷͳȂ͵ͷͻǤ



 74 

   Šƒ’–‡”͵

WithinhabitatEǦdiversityinahostǦ parasitoidinteractionnetwork

























TornéǦNoguera,A;Rodrigo,A;Bosch,J.(2015)

’”‡’ƒ”ƒ–‹‘



Šƒ’–‡”͵

Summary ’‡ ‹‡•ƒ••‡„Žƒ‰‡•ƒ†–Š‡‹”‹–‡”ƒ –‹‘•˜ƒ”›–Š”‘—‰Š•’ƒ ‡ǡ‰‡‡”ƒ–‹‰ †‹˜‡”•‹–›’ƒ––‡”•ƒ–†‹ˆˆ‡”‡–•’ƒ–‹ƒŽ• ƒŽ‡•Ǥ ‡ ‘Ž‘‰›ǡȾǦ†‹˜‡”•‹–›‹•ƒ ‘‘‡ƒ•—”‡ ‘ˆ•’‡ ‹‡•–—”‘˜‡”„‡–™‡‡Ž‘ ƒŽ‹–‹‡•ǡ„—–‹–Šƒ•”ƒ”‡Ž›„‡‡—•‡†–‘‡š’Ž‘”‡‹–‡”ƒ –‹‘ –—”‘˜‡”Ǥ‡ ‡–†‡˜‡Ž‘’‡–‘ˆȾǦ†‹˜‡”•‹–›‹†‡š‡•’”‘˜‹†‡•ƒ‡™ˆ”ƒ‡™‘”–‘ƒƒŽ›œ‡ •’ƒ–‹ƒŽ˜ƒ”‹ƒ–‹‘‘ˆ‡ ‘Ž‘‰‹ ƒŽ‡–™‘”•ƒ†–‘‹†‡–‹ˆ›–Š‡•‘—” ‡•‘ˆ–Š‹•˜ƒ”‹ƒ–‹‘Ǥ ‡”‡ǡ ™‡•–—†›–Š‡Ž‘ ƒŽǦ• ƒŽ‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ–‹‘‘ˆƒ ƒ˜‹–›Ǧ‡•–‹‰„‡‡ ƒ† ™ƒ•’ ‘—‹–› ȋŠ‘•–•Ȍǡ–Š‡‹”‡•–ƒ••‘ ‹ƒ–‡•ȋ’ƒ”ƒ•‹–‘‹†•Ȍǡƒ†–Š‡”‡•—Ž–‹‰ƒ–ƒ‰‘‹•–‹ ‡–™‘”‘˜‡”ƒ Š‘‘‰‡‡‘—•Šƒ„‹–ƒ–Ǥ‘‘„–ƒ‹„‡‡Ȁ™ƒ•’‡•–•™‡’Žƒ ‡†‡•–Ǧ–”ƒ’•ƒ–ʹͷ•‹–‡•‘˜‡”ƒ͵ʹ ʹƒ”‡ƒǤ‹–‡•™‡”‡•‡’ƒ”ƒ–‡†„›ͷͲͲǦͳͲͲͲǤ ‘•–•ƒ†–Š‡‹”’ƒ”ƒ•‹–‘‹†•™‡”‡”‡ƒ”‡†‹ –Š‡ Žƒ„‘”ƒ–‘”›Ǥ ‡ ‘„–ƒ‹‡† ͳͷͶͳ ‡•–• ȋͶͻͷͶ ‡ŽŽ•Ȍ „‡Ž‘‰‹‰ –‘ͶͳŠ‘•–•’‡ ‹‡•ƒ† ‘–ƒ‹‹‰ʹ͸’ƒ”ƒ•‹–‘‹†•’‡ ‹‡•ǤŠ‡‘•–ƒ„—†ƒ–Š‘•–•’‡ ‹‡•–‡†‡†–‘Šƒ˜‡Š‹‰Š‡” ’ƒ”ƒ•‹–‹•”ƒ–‡ǤEǦ†‹˜‡”•‹–›™ƒ•Š‹‰Šˆ‘”„‘–ŠŠ‘•–•ƒ†’ƒ”ƒ•‹–‡•ǡƒ†–Š‡ƒ‹†”‹˜‡”‘ˆ ˜ƒ”‹ƒ„‹Ž‹–›™ƒ••’‡ ‹‡•–—”‘˜‡”ǡ™‹–Šƒ˜‡”›‹‘” ‘–”‹„—–‹‘‘ˆ‘”†‡”‡†•’‡ ‹‡•Ž‘•• ȋ‡•–‡†‡••ȌǤŠƒ–‹•ǡŽ‘ ƒŽ•’‡ ‹‡•”‹ Š‡••–‡†‡†–‘„‡•‹‹Žƒ” ƒ ”‘•• –Š‡ •–—†› ƒ”‡ƒ ‘—‹–› ‘’‘•‹–‹‘ –‡†‡† –‘ †‹ˆˆ‡” „‡–™‡‡ •‹–‡•Ǥ –‡”ƒ –‹‘ȾǦ†‹˜‡”•‹–›™ƒ•ƒŽ•‘ Š‹‰Šǡƒ†‘•–Ž›†—‡–‘•’‡ ‹‡•–—”‘˜‡”™‹–ŠƒŽ‘™ ‘–”‹„—–‹‘‘ˆ”‡™‹”‹‰ȋ•ƒ‡•’‡ ‹‡• ‹–‡”ƒ –‹‰†‹ˆˆ‡”‡–Ž›ƒ–†‹ˆˆ‡”‡–•‹–‡•ȌǤ •—ǡƒŽ–Š‘—‰Š•’‡ ‹‡• ‘’‘•‹–‹‘™ƒ•”ƒ–Š‡” ‹†‹‘•› ”ƒ–‹  –‘ ‡ƒ Š •‹–‡ǡ •’‡ ‹‡• ‹–‡”ƒ –‡† •‹‹Žƒ”Ž› ƒ ”‘••†‹ˆˆ‡”‡–•‹–‡•Ǥ ‘•–ȾǦ †‹˜‡”•‹–›‹ ”‡ƒ•‡†™‹–Š‰‡‘‰”ƒ’Š‹ ƒŽ†‹•–ƒ ‡ǡ„—–’ƒ”ƒ•‹–‘‹†ƒ†‹–‡”ƒ –‹‘ȾǦ†‹˜‡”•‹–› †‹†‘–Ǥ—””‡•—Ž–•ƒ††‹–‹‘ƒŽŽ›‹†‹ ƒ–‡–Šƒ–‹–‡”ƒ –‹‘ȾǦ†‹˜‡”•‹–›‹•„‡––‡”‡š’Žƒ‹‡†„› Š‘•–ȾǦ†‹˜‡”•‹–›–Šƒ„›’ƒ”ƒ•‹–‘‹†ȾǦ†‹˜‡”•‹–›ǡ’”‘„ƒ„Ž›†—‡–‘–Š‡Š‹‰Š‡””ƒ‰‡‘ˆŠ‘•–ȾǦ †‹˜‡”•‹–›Ǥ‡†‹• —••–Š‡‹’‘”–ƒ ‡‘ˆ‹†‡–‹ˆ›‹‰–Š‡•‘—” ‡•‘ˆ˜ƒ”‹ƒ–‹‘–‘—†‡”•–ƒ† –Š‡ †”‹˜‡”• ‘ˆ –Š‡ ‘„•‡”˜‡† Š‡–‡”‘‰‡‡‹–›Ǥ ‡ ƒ—•‡ ‘—‹–‹‡• ƒ† –Š‡‹” ‹–‡”ƒ –‹‘• ƒ›˜ƒ”›ƒ–ƒ˜‡”›Ž‘ ƒŽ• ƒŽ‡ȋ–Š‹••–—†›Ȍǡ™‡ƒŽ•‘‡’Šƒ•‹œ‡–Š‡ ‡‡† –‘ •ƒ’Ž‡ ƒ •—ˆˆ‹ ‹‡–Ž›Žƒ”‰‡—„‡”‘ˆ•‹–‡•”‡‰ƒ”†Ž‡••‘ˆ–Š‡ ƒ—•‡‘ˆ–Š‡‘„•‡”˜‡†˜ƒ”‹ƒ–‹‘Ǥ

3.1. Introduction

‹˜‡”•‹–›’ƒ––‡”•™‡ ƒ —””‡–Ž›‘„•‡”˜‡‹ƒ–—”‡ƒ”‡–Š‡‘—– ‘‡‘ˆ—Ž–‹’Ž‡„‹‘–‹ ƒ† ƒ„‹‘–‹ ˆƒ –‘”•ǡƒ†ƒ”‡ˆŽ‡ –‹‘‘ˆ–Š‡‹–‡”ƒ –‹‘•„‡–™‡‡–Š‡Ǥ ‘ —†‡”•–ƒ† •— Š ’ƒ––‡”•ƒ†–Š‡‹”—†‡”Ž›‹‰’”‘ ‡••‡•‹•‘ˆƒŒ‘”‹’‘”–ƒ ‡‹ ‘—‹–›‡ ‘Ž‘‰›Ǥ ‘” ƒ› †‡ ƒ†‡•ǡ ‡ ‘Ž‘‰‹•–• Šƒ˜‡ •–—†‹‡† •’‡ ‹‡• †‹˜‡”•‹–› ƒ– • ƒŽ‡•”ƒ‰‹‰ˆ”‘•‡˜‡”ƒŽ ‡–‡”•–‘ƒ›‹Ž‘‡–‡”•ǤŠ‹––ƒ‡”ȋͳͻ͸ͲȌ™ƒ•–Š‡ˆ‹”•––‘’”‘’‘•‡ ƒ ’ƒ”–‹–‹‘‹‰ ‘ˆ †‹˜‡”•‹–› ƒ ”‘•• ͵ †‹ˆˆ‡”‡– •’ƒ–‹ƒŽ • ƒŽ‡•ǣ ȽǦ†‹˜‡”•‹–›ǡ ™Š‹ Š ‡ƒ•—”‡• –Š‡ ”‡Žƒ–‹‘•Š‹’ „‡–™‡‡–Š‡—„‡”‘ˆ•’‡ ‹‡•ƒ†‹†‹˜‹†—ƒŽ•ȋ‹Ǥ‡Ǥ†‹˜‡”•‹–›Ȍ‹ƒ’ƒ”–‹ —Žƒ”Ž‘ ƒŽ‹–›ȋ ‹•Š‡” ‡– ƒŽǤ ͳͻͶ͵Ȍǡ JǦ†‹˜‡”•‹–›ǡ ƒ ‡ƒ•—”‡ ‘ˆ ”‡‰‹‘ƒŽ †‹˜‡”•‹–›ǡ ƒ† ȾǦ†‹˜‡”•‹–›ǡ ƒ‡ƒ•—”‡‘ˆ †‹˜‡”•‹–›–—”‘˜‡”„‡–™‡‡Ž‘ ƒŽ‹–‹‡•ǤȾǦ†‹˜‡”•‹–›’”‘˜‹†‡•ƒ‡ƒ•—”‡‘ˆ ‘—‹–›•’ƒ–‹ƒŽ  77 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

˜ƒ”‹ƒ„‹Ž‹–›ǡ”‡ˆŽ‡ –‹‰Š‹•–‘”‹ ƒŽ’”‘ ‡••‡•ƒ†”‡˜‡ƒŽ‹‰‹ˆ‘”ƒ–‹‘‘’‘’—Žƒ–‹‘†›ƒ‹ • ƒ† •’‡ ‹‡• ”‡•’‘•‡• –‘ Šƒ„‹–ƒ– ‘†‹ˆ‹ ƒ–‹‘• •— Š ƒ• ‡˜‹”‘‡–ƒŽ ‰”ƒ†‹‡–• ƒ† ’‡”–—”„ƒ–‹‘•Ǥ ‘”–Š‹•”‡ƒ•‘ǡȾǦ†‹˜‡”•‹–›Šƒ•„‡ ‘‡ƒ”‡Ž‡˜ƒ– ‡ƒ•—”‡ ‹ „‹‘Ž‘‰‹ ƒŽ ‘•‡”˜ƒ–‹‘ ȋ‘†‹– ‡– ƒŽǤ ʹͲͲʹȌǤ ȾǦ†‹˜‡”•‹–› ƒ „‡ ’ƒ”–‹–‹‘‡† ‹–‘ –™‘ ƒ††‹–‹˜‡ ‘’‘‡–•ǣ •’ƒ–‹ƒŽ –—”‘˜‡”ǡ –Šƒ– ‹•ǡ †‹••‹‹Žƒ”‹–› †—‡ –‘ •’‡ ‹‡• ”‡’Žƒ ‡‡–ǡ ƒ† ‡•–‡†‡••‘ˆƒ••‡„Žƒ‰‡•ǡ–Šƒ–‹•ǡ†‹••‹‹Žƒ”‹–›†—‡–‘‘”†‡”‡†•’‡ ‹‡•Ž‘••ȋƒ•‡Ž‰ƒʹͲͳͲȌǤ ‘™‹‰ –Š‡ ”‡Žƒ–‹˜‡ ‹’‘”–ƒ ‡ ‘ˆ –Š‡•‡ –™‘ ‘’‘‡–• ‹• ‡••‡–‹ƒŽ –‘ —†‡”•–ƒ† –Š‡ ƒ—•‡•‘ˆ–Š‡‘„•‡”˜‡†•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›Ǥ

‡ ƒ—•‡ •’‡ ‹‡• ƒ”‡ ‘– ‹•‘Žƒ–‡† „—– ‹‡”•‡† ‹ ‘’Ž‡š ‡–™‘”• ‘‡ –‹‰ –Š‡ †‹”‡ –Ž› ƒ† ‹†‹”‡ –Ž› ™‹–Š ‘–Š‡” •’‡ ‹‡•ǡ •’ƒ–‹ƒŽ Š‡–‡”‘‰‡‡‹–› ‹ •’‡ ‹‡• ‘—‹–› •–”— –—”‡ ‹• ‡š’‡ –‡† –‘ ’”‘ˆ‘—†Ž› ƒˆˆ‡ – ‡–™‘” •–”— –—”‡ ȋŽ‡•‡ ƒ† ‘”†ƒ‘ ʹͲͲʹǡ žœ“—‡œ ‡– ƒŽǤ ʹͲͲͻ„ȌǤ ‡ ‡–Ž›ǡ •‘‡ •–—†‹‡• Šƒ˜‡ ‡‹–Š‡” –Š‡‘”‡–‹ ƒŽŽ› ‘” ‡’‹”‹ ƒŽŽ› ƒ††”‡••‡† •— Š ”‡Žƒ–‹‘•Š‹’ ȋ—”Ž‡ ƒ† Žƒ” א ʹͲͳͳȌǤ ‘” ‡šƒ’Ž‡ǡ •’ƒ–‹ƒŽ ƒ‰‰”‡‰ƒ–‹‘ ƒ† ‹†‡–‹–› ‘ˆ ’Žƒ–•ǡ –‘‰‡–Š‡” ™‹–Š ƒ‹ƒŽ ‘„‹Ž‹–›ǡ Šƒ• „‡‡•Š‘™–‘Šƒ˜‡ƒ•–”‘‰ ‹ˆŽ—‡ ‡‘‹†‡–‹–›ǡ•–”‡‰–Š ƒ††‹•–”‹„—–‹‘‘ˆ‹–‡”ƒ –‹‘•‹’Žƒ–Ǧƒ‹ƒŽ—–—ƒŽ‹•–‹  ‡–™‘”•ȋ‘”ƒŽ‡•ƒ†žœ“—‡œʹͲͲͺȌǤ –Šƒ•ƒŽ•‘„‡‡†‡‘•–”ƒ–‡† –Šƒ– ‡–™‘” •–”— –—”ƒŽ ’ƒ––‡”• ƒ”‡ Žƒ”‰‡Ž› •Šƒ’‡† „› ”‡Žƒ–‹˜‡ •’‡ ‹‡• ƒ„—†ƒ ‡ ƒ† •’ƒ–‹‘–‡’‘”ƒŽ ’ƒ––‡”•‘ˆ‹–‡”ƒ –‹‰•’‡ ‹‡•ȋžœ“—‡œ‡–ƒŽǤʹͲͲͻƒȌǤ ‹ƒŽŽ›ǡ‡˜‡‹ˆ†‡ ƒ›‘ˆ•‹‹Žƒ”‹–›‹ •’‡ ‹‡• ‘’‘•‹–‹‘ ™‹–Š †‹•–ƒ ‡ Šƒ• „‡‡ ™‡ŽŽ ‡•–ƒ„Ž‹•Š‡† ȋ‡‘Žƒƒ†Š‹–‡ͳͻͻͻǡ ‘‹‹‡ ‡– ƒŽǤ ʹͲͲ͹Ȍǡ Ž‹––Ž‡ ‹• ‘™ ƒ„‘—– †‡ ƒ› ‘ˆ •‹‹Žƒ”‹–› ‘ˆ ‹–‡”ƒ –‹‘• „‡–™‡‡ •’‡ ‹‡•ǡƒ†Š‘™–Š‡–™‘†‡ ƒ›’ƒ––‡”•ƒ”‡”‡Žƒ–‡†Ǥ‡ ƒ—•‡‹–‡”ƒ –‹‘•ƒ”‡‹ˆŽ—‡ ‡†„› –Š‡˜ƒ”‹ƒ„‹Ž‹–›‘ˆ–Š‡–™‘–”‘’Š‹ Ž‡˜‡Ž•’Ž—•–Š‡‹”‹Š‡”‡–˜ƒ”‹ƒ„‹Ž‹–› ȋ‹Ǥ‡Ǥ •ƒ‡ •’‡ ‹‡• ‹–‡”ƒ –‹‰ †‹ˆˆ‡”‡–Ž›Ȍǡ ‹–‡”ƒ –‹‘• ƒ”‡ ‡š’‡ –‡† –‘ †‹•’Žƒ› ‘”‡ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ–‹‘ –Šƒ •’‡ ‹‡•Ǥ

Ž–Š‘—‰Š ȾǦ†‹˜‡”•‹–› ‹• ™‹†‡Ž› —•‡† ƒ‘‰ ‡ ‘Ž‘‰‹•–• –‘ ‡š’Ž‘”‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ–‹‘‘ˆ ‘—‹–‹‡•ǡ‹–Šƒ•„‡‡•‡Ž†‘—•‡†–‘‡š’Ž‘”‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ–‹‘‘ˆ‹–‡”ƒ –‹‘•Ǥ‘‹•‘–ƒ† ‘ŽŽ‡ƒ‰—‡• ȋʹͲͳʹȌ ’”‘’‘•‡† ƒ ‡™ †‹••‹‹Žƒ”‹–› ‹†‡š –‘ ‡š’Ž‘”‡ †‹ˆˆ‡”‡ ‡• „‡–™‡‡ ‹–‡”ƒ –‹‘ ‡–™‘”• ƒ ”‘•• •’ƒ ‡Ǥ ‘ ƒ††‹–‹‘ƒŽŽ› †‡Ž˜‡ ‹–‘ –Š‡ •‘—” ‡ ‘ˆ ˜ƒ”‹ƒ„‹Ž‹–› „‡–™‡‡ ‡–™‘”•ǡ –Š‡› •—„†‹˜‹†‡† ȾǦ†‹˜‡”•‹–› ‘ˆ ‹–‡”ƒ –‹‘• ‹–‘ ƒ ‘’‘‡– †—‡ –‘ •’‡ ‹‡•–—”‘˜‡”ƒ†ƒ ‘’‘‡–†—‡–‘‹–‡”ƒ –‹‘–—”‘˜‡”„‡–™‡‡‰‹˜‡•’‡ ‹‡•ǡ–Šƒ– ‹•ǡ •Šƒ”‡† •’‡ ‹‡• ‹–‡”ƒ –‹‰ †‹ˆˆ‡”‡–Ž› ȋ”‡™‹”‹‰ȌǤ Š‡› ƒŽ•‘ ’”‘’‘•‡† ‘’ƒ”‹‰ ‹–‡”ƒ –‹‘• „‡–™‡‡ ‘˜‡”Žƒ’’‹‰ •’‡ ‹‡• ‘ˆ ‡ƒ Š Ž‘ ƒŽ ™‡„ –‘ –Š‡‹” ‘—–‡”’ƒ”–• ‹ –Š‡

 78 Šƒ’–‡”͵

”‡‰‹‘ƒŽ™‡„–‘†‡–‡”‹‡‹ˆ‹–‡”ƒ –‹‘•ˆ‘—†ƒ––Š‡”‡‰‹‘ƒŽŽ‡˜‡Žƒ”‡ƒŽ•‘ˆ‘—†ƒ––Š‡ Ž‘ ƒŽŽ‡˜‡Žǡ‹†‹ ƒ–‹‰–Šƒ–•’‡ ‹‡•‹–‡”ƒ –•‹‹Žƒ”Ž›ƒ ”‘••–Š‡”‡‰‹‘ǤŠ‹•ƒ’’”‘ƒ ŠƒŽŽ‘™• ‡•–ƒ„Ž‹•Š‹‰™Š‡–Š‡”–Š‡”‡‰‹‘ƒŽ™‡„‘Ž›”‡ˆŽ‡ –•–Š‡”‡‰‹‘ƒŽǡ„—–‘––Š‡Ž‘ ƒŽ„‡Šƒ˜‹‘” ‘ˆ •’‡ ‹‡• ‹–‡”ƒ –‹‘• ‘”ǡ ‘ –Š‡ ‘–”ƒ”›ǡ ‹– ”‡ˆŽ‡ –• „‘–Š –Š‡ ”‡‰‹‘ƒŽ ƒ† –Š‡ Ž‘ ƒŽ „‡Šƒ˜‹‘”ǤŠ‡ˆ‘”‡”‘—– ‘‡™‘—Ž†ƒ’’Ž›‹ˆ†‹ˆˆ‡”‡ ‡•ƒ‘‰Ž‘ ƒŽ™‡„•ƒ”‡‘•–Ž›†—‡ –‘ ‹–‡”ƒ –‹‘ ”‡™‹”‹‰ǡ –Šƒ– ‹•ǡ ‹ˆ •’‡ ‹‡• ‹–‡”ƒ – †‹ˆˆ‡”‡–Ž› ƒ ”‘•• ’Ž‘–•Ǥ ‘˜‡”•‡Ž›ǡ –Š‡ •‡ ‘†‘—– ‘‡™‘—Ž†ƒ’’Ž›–‘•‹–—ƒ–‹‘•‹™Š‹ Š•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–› ‘ˆ ‹–‡”ƒ –‹‘• ‹• ‘•–Ž›†—‡–‘•’‡ ‹‡•–—”‘˜‡”ǡƒ† ‘‘•’‡ ‹‡•‹†‹ˆˆ‡”‡–Ž‘ ƒŽ™‡„•–‡†–‘‹–‡”ƒ – •‹‹Žƒ”Ž›Ǥ

‡ ‡–Ž›ǡ•‘‡•–—†‹‡•Šƒ˜‡‡’‹”‹ ƒŽŽ›ƒ††”‡••‡†EǦ†‹˜‡”•‹–›‘ˆ‹–‡”ƒ –‹‘•ƒ–‹†‹˜‹†—ƒŽǦ ȋ—’‘–‡–ƒŽǤʹͲͳͶȌƒ† ‘—‹–›Ž‡˜‡Ž•‹ƒ–—”ƒŽȋ‘‹•‘–‡–ƒŽǤʹͲͳʹǡž––‹Ž‘‡–ƒŽǤʹͲͳ͵ǡ ƒ”•–‡•‡ ‡– ƒŽǤ ʹͲͳͶǡ ‹ƒ‘‘ ƒ† —”Ž‡ ʹͲͳͶǡ ”ÞŒ‡Ž•‰ƒƒ”† ‡– ƒŽǤ ʹͲͳͷȌ ƒ† ƒ‰”‹ —Ž–—”ƒŽ ȋ‘”ˆ‘Ž ‡– ƒŽǤ ʹͲͳͶȌ Šƒ„‹–ƒ–•Ǥ  •‘‡ ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘” ‡–™‘”•ǡ •’‡ ‹‡• –—”‘˜‡”Šƒ•„‡‡•Š‘™–‘„‡–Š‡ƒ‹ ƒ—•‡‘ˆ‹–‡”ƒ –‹‘–—”‘˜‡” ȋ‹ƒ‘‘ ƒ† —”Ž‡ ʹͲͳͶǡ ”ÞŒ‡Ž•‰ƒƒ”† ‡– ƒŽǤ ʹͲͳͷȌǡ „—– ‘–Š‡”• Šƒ˜‡ ›‹‡Ž†‡† ƒ •Ž‹‰Š–Ž› ‰”‡ƒ–‡” ‘–”‹„—–‹‘‘ˆ”‡™‹”‹‰ȋƒ”•–‡•‡‡–ƒŽǤʹͲͳͶȌǤ ƒ††‹–‹‘ǡ•‘‡•–—†‹‡•ȋ”ÞŒ‡Ž•‰ƒƒ”†‡– ƒŽǤʹͲͳͷȌŠƒ˜‡ˆ‘—†’‘ŽŽ‹ƒ–‘”–—”‘˜‡”–‘„‡–Š‡ƒŒ‘” ‘–”‹„—–‘”–‘‹–‡”ƒ –‹‘–—”‘˜‡”ǡ ™Š‹Ž‡‘–Š‡”•ȋ‹ƒ‘‘ƒ†—”Ž‡ʹͲͳͶȌƒ––”‹„—–‡ƒ’”‡†‘‹ƒ–”‘Ž‡–‘’Žƒ––—”‘˜‡”Ǥ

‡”‡ǡ™‡™‘”™‹–Šƒ ‘—‹–›‘ˆ ƒ˜‹–›Ǧ‡•–‹‰„‡‡•ƒ†™ƒ•’•ȋŠ‡ ‡ˆ‘”–ŠŠ‘•–•Ȍƒ† –Š‡‹” ‡•– ƒ••‘ ‹ƒ–‡•ǡ ‹ Ž—†‹‰ ’ƒ”ƒ•‹–‘‹†•ǡ Ž‡’–‘’ƒ”ƒ•‹–‡•ǡ ’”‡†ƒ–‘”• ƒ† • ƒ˜‡‰‡”• ȋŠ‡ ‡ˆ‘”–Š’ƒ”ƒ•‹–‘‹†•Ȍƒ†•–—†›–Š‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ–‹‘‘ˆ–Š‡–™‘ ‘—‹–‹‡•ǡƒ•™‡ŽŽƒ• –Š‡‹” ‹–‡”ƒ –‹‘•Ǥ ”‡˜‹‘—• •–—†‹‡• ™‘”‹‰ ™‹–Š ƒ˜‹–›Ǧ‡•–‹‰„‡‡•ƒ†™ƒ•’•ƒƒŽ›œ‡ †‹ˆˆ‡”‡ ‡• „‡–™‡‡ Šƒ„‹–ƒ–• ‘” ƒŽ‘‰ ƒ ‡˜‹”‘‡–ƒŽ ‰”ƒ†‹‡– ȋŽ„”‡ Š– ‡– ƒŽǤ ʹͲͲ͹ǡ ›Ž‹ƒƒ‹• ‡– ƒŽǤ ʹͲͲ͹ǡ •‘”‹‘ ‡– ƒŽǤ ʹͲͳͷȌǤ ‘˜‡”•‡Ž›ǡ ‘—” •–—†› ™ƒ• ‘†— –‡† ƒ ”‘•• ƒ ‘–‹—‘—•Šƒ„‹–ƒ–ƒ†ƒ††”‡••‡••’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›ƒ–ƒŽ‘ ƒŽ• ƒŽ‡ȋ†‹•–ƒ ‡„‡–™‡‡’Ž‘–• ƒǤ ͳͲͲͲ ȌǤ ‘–™‹–Š•–ƒ†‹‰ –Š‡ Žƒ  ‘ˆ ‡˜‹”‘‡–ƒŽ ‰”ƒ†‹‡–• ‘” ’Š›•‹ ƒŽ „ƒ””‹‡”•ǡ ’”‡˜‹‘—••–—†‹‡•‹–Š‡•ƒ‡•–—†›ƒ”‡ƒŠƒ˜‡ˆ‘—†ƒ”‡†‰‡‘‰”ƒ’Š‹ ƒŽ’ƒ––‡”•‘ˆ•’ƒ–‹ƒŽ Š‡–‡”‘‰‡‡‹–›‹„‡‡•’‡ ‹‡• ‘’‘•‹–‹‘ȋ‘”±Ǧ‘‰—‡”ƒ‡–ƒŽǤʹͲͳͶȌǤ—”ƒ‹‹•–Š‡”‡ˆ‘”‡ –‘•–—†›–Š‡‹–”‹•‹ ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ–Š‡Š‘•–Ǧ’ƒ”ƒ•‹–‘‹†‡–™‘”ǡ”ƒ–Š‡”–Šƒ–‘‡•–ƒ„Ž‹•ŠŠ‘™ †‹ˆˆ‡”‡–‡˜‹”‘‡–ƒŽˆƒ –‘”•ƒ›ƒˆˆ‡ –‹–‡”ƒ –‹‘‹†‡–‹–›ƒ†‡–™‘”•–”— –—”‡Ǥ

‡Šƒ˜‡–Š‡ˆ‘ŽŽ‘™‹‰‘„Œ‡ –‹˜‡•ǣ

 79 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

ͳǦ ‘ƒƒŽ›œ‡–Š‡•’ƒ–‹ƒŽ˜ƒ”‹ƒ–‹‘‘ˆ–Š‡Š‘•–ƒ†’ƒ”ƒ•‹–‘‹† ‘—‹–‹‡• ƒ ”‘•• ƒ ‘–‹—‘—•Šƒ„‹–ƒ–Ǥ‡ƒ•Š‘™ ‘—‹–› ‘’‘•‹–‹‘˜ƒ”‹‡•Ž‘ ƒŽŽ›ƒ†™Š‡–Š‡” –Š‹•˜ƒ”‹ƒ–‹‘ˆ‘ŽŽ‘™•ƒ•’ƒ–‹ƒŽ’ƒ––‡”Ǥ‡ƒŽ•‘ƒ•Š‘™–Š‡–™‘ ‘—‹–‹‡•ƒ”‡ ”‡Žƒ–‡†Ǥ ʹǦ ‘•–—†›EǦ†‹˜‡”•‹–›‘ˆŠ‘•–Ǧ’ƒ”ƒ•‹–‘‹†‹–‡”ƒ –‹‘•Ǥ‡ƒ•™Š‡–Š‡”–Š‡ƒ‹†”‹˜‡” ‘ˆ‹–‡”ƒ –‹‘–—”‘˜‡”‹••’‡ ‹‡•–—”‘˜‡”‘””‡™‹”‹‰Ǥ ͵Ǧ ‘‡š’Ž‘”‡–Š‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡Š‘•–ǡ’ƒ”ƒ•‹–‘‹†ƒ†‹–‡”ƒ –‹‘EǦ†‹˜‡”•‹–›ƒ† –‘‡šƒ‹‡–Š‡‹”†‹•–ƒ ‡†‡ ƒ›Ǥ‡ƒ•™Š‡–Š‡”Š‘•–ƒ†’ƒ”ƒ•‹–‘‹†ȾǦ†‹˜‡”•‹–›ƒ”‡ ‰‘‘† ’”‡†‹ –‘”• ‘ˆ ‹–‡”ƒ –‹‘ ȾǦ†‹˜‡”•‹–›Ǥ ‡ ƒŽ•‘ ƒ• ™Š‡–Š‡”†‹•–ƒ ‡†‡ ƒ› ’ƒ––‡”•ƒ”‡•‹‹Žƒ”ˆ‘”–Š‡–Š”‡‡‰”‘—’•Ǥ



3.2. Methodsandmaterials

3.2.1. Studyarea

Š‡•–—†›™ƒ• ‘†— –‡†‹–Š‡ ƒ””ƒˆƒ–—”ƒŽƒ”ȋƒ” ‡Ž‘ƒǡ’ƒ‹Ȍǡƒ‡†‹–‡””ƒ‡ƒ • ”—„Žƒ† †‘‹ƒ–‡† „› Quercus cocciferaǡ Pistacia lentiscusǡ Rosmarinus officinalisƒ† ThymusvulgarisǤ—”•–—†›ƒ”‡ƒǡ‡–‹”‡Ž›Ž‘ ƒ–‡†‹–Š‡’ƒ”ǡ‡ ‘’ƒ••‡•ƒ•—”ˆƒ ‡‘ˆ͵ʹ ʹǤ  ‡ •‡Ž‡ –‡† ʹͷ ’Ž‘–• †‹•–”‹„—–‡† ‘”‡ ‘” Ž‡•• ”‡‰—Žƒ”Ž› ƒ ”‘•• –Š‡ •–—†› ƒ”‡ƒǤ ‹•–ƒ ‡•„‡–™‡‡‡ƒ”‡•–’Ž‘–•”ƒ‰‡†ˆ”‘ͷͺͷ–‘ͳ͵ͷͶǤ



3.2.2. Surveys

‡ƒ Š’Ž‘–™‡’Žƒ ‡†ƒ–”ƒ’Ǧ‡•–‹‰•–ƒ–‹‘ˆƒ ‹‰Ǥƒ Š•–ƒ–‹‘ ‘–ƒ‹‡†•‡˜‡†”‹ŽŽ‡† ™‘‘†„Ž‘ •™‹–Š‹•‡”–‡†’ƒ’‡”–—„‡•Ǥƒ Š™‘‘†„Ž‘ ƒ ‘‘†ƒ–‡†ʹͷ–—„‡•‘ˆƒ‰‹˜‡ †‹ƒ‡–‡”ȋʹǡ͵ǡͶǡͷǡ͸ǡ͹‘”ͺȌǡ”‡•—Ž–‹‰‹ͳ͹ͷ‡•–‹‰ ƒ˜‹–‹‡•’‡”•–ƒ–‹‘Ǥƒ’‡”–—„‡ Ž‡‰–Š™ƒ•ͷ ˆ‘”–Š‡ʹƒ†͵†‹ƒ‡–‡”•ƒ†ͳͷ ˆ‘”–Š‡”‡•–Ǥƒ Š‡•–‹‰•–ƒ–‹‘ Šƒ†͹–”ƒ’Ǧ‡•–•ǡ‘‡‘ˆ‡ƒ Š†‹ƒ‡–‡”Ǥ‡•–‹‰•–ƒ–‹‘•™‡”‡ Š‡ ‡†‡˜‡”›ʹ™‡‡•ƒ† –—„‡• ‘–ƒ‹‹‰ ‘’Ž‡–‡† ‡•–• ™‡”‡ ’—ŽŽ‡† ‘—– ƒ† ”‡’Žƒ ‡† ™‹–Š ‡’–› ‘‡•ǡ •‘ –Šƒ– –Š‡”‡™‡”‡‡•–‹‰ ƒ˜‹–‹‡•‘ˆƒŽŽ†‹ƒ‡–‡”•ƒ˜ƒ‹Žƒ„Ž‡ƒ–ƒŽŽ–‹‡•Ǥ‡•–‹‰•–ƒ–‹‘•”‡ƒ‹‡†

 80 Šƒ’–‡”͵

‹–Š‡ˆ‹‡Ž†ˆ”‘ ‡„”—ƒ”›–‘ –‘„‡”ǡ‹ʹͲͳͳƒ†ʹͲͳ͵Ǥƒ–ƒ‘ˆ–Š‡–™‘›‡ƒ”•ƒ”‡’‘‘Ž‡† –‘‰‡–Š‡”‹–Š‡ƒƒŽ›•‡•Ǥ

–Š‡Žƒ„‘”ƒ–‘”›ǡ‡•–•™‡”‡‹†‹˜‹†—ƒŽŽ›’Žƒ ‡†‹–‡•––—„‡•ƒ†‡’–‹ƒ–‡’‡”ƒ–—”‡ Šƒ„‡” •‹—Žƒ–‹‰ ‘–ŠŽ› ƒ„‹‡– –Š‡”‘’‡”‹‘†• ‘ˆ –Š‡ •–—†› •‹–‡Ǥ •’”‹‰ƒ† •—‡”ǡ ‡•–• ™‡”‡ †‹••‡ –‡† ƒ† –Š‡‹” ‘–‡–• ”‡ ‘”†‡†Ǥ ‘•–•ƒ†’ƒ”ƒ•‹–‘‹†•™‡”‡ ”‡ƒ”‡†ƒ†‹†‡–‹ˆ‹‡†Ǥ



3.2.3. Statisticalanalysis

Hostandparasitoidcommunities

‘ Šƒ”ƒ –‡”‹œ‡ ‘—‹–› •–”— –—”‡ ƒ– ‡ƒ Š ‡•–‹‰ •–ƒ–‹‘ǡ ™‡ —•‡† –Š‡ ˜ƒ”‹ƒ„Ž‡• Š‘•– ƒ„—†ƒ ‡ ȋ—„‡” ‘ˆ Š‘•– ‡ŽŽ•Ȍǡ Š‘•– ”‹ Š‡•• ȋ—„‡” ‘ˆ Š‘•–•’‡ ‹‡•Ȍǡ’ƒ”ƒ•‹–‘‹† ƒ„—†ƒ ‡ȋ—„‡”‘ˆ ‡ŽŽ•’ƒ”ƒ•‹–‹œ‡†Ȍǡ’ƒ”ƒ•‹–‘‹†”‹ Š‡••ȋ—„‡”‘ˆ’ƒ”ƒ•‹–‘‹†•’‡ ‹‡•Ȍ ƒ†’ƒ”ƒ•‹–‹•”ƒ–‡ȋ؊‘•– ‡ŽŽ•’ƒ”ƒ•‹–‹œ‡†ȌǤ

‡—•‡†”ƒ”‡ˆƒ –‹‘ —”˜‡•–‘†‡–‡”‹‡–Š‡ ‘’Ž‡–‡‡••‘ˆ‘—”•ƒ’Ž‹‰‘ˆ–Š‡Š‘•–ƒ† ’ƒ”ƒ•‹–‘‹† ‘—‹–›Ǥ ƒ”‡ˆƒ –‹‘• ™‡”‡ ‘†— –‡† ™‹–Š ˜‡‰ƒ ʹǤͲǦͳͲȋ•ƒ‡‡–ƒŽǤ ʹͲͳͷȌˆ‘”ȋ‘”‡‡ƒʹͲͳͶȌǤ

‡‡ƒ•—”‡†–Š‡ ‘””‡Žƒ–‹‘„‡–™‡‡Š‘•–ƒ„—†ƒ ‡ƒ†”‹ Š‡••ȋˆ‘”ƒŽŽŠ‘•–•ƒ†ˆ‘” ’ƒ”ƒ•‹–‹œ‡† Š‘•–• •‡’ƒ”ƒ–‡Ž›Ȍǡ ƒ† „‡–™‡‡ ’ƒ”ƒ•‹–‘‹† ƒ„—†ƒ ‡ ƒ† ”‹ Š‡••Ǥ •‹‰ –Š‡ •ƒ‡’”‘ ‡†—”‡ǡ™‡ƒŽ•‘‡ƒ•—”‡†–Š‡ ‘””‡Žƒ–‹‘„‡–™‡‡Š‘•–ȋƒŽŽŠ‘•–•Ȍƒ†’ƒ”ƒ•‹–‘‹† ƒ„—†ƒ ‡ƒ†”‹ Š‡••™‹–Šƒ’‡ƒ”ƒǯ•”Š‘Ǥ

‡ ƒŽ —Žƒ–‡†–Š‡‘˜‡”ƒŽŽEǦ†‹˜‡”•‹–›ȋƒŽŽǦ•‹–‡ ‘’ƒ”‹•‘Ȍ„ƒ•‡†‘ޔ‡•‡ǯ•†‹••‹‹Žƒ”‹–›

‹†‡šˆ‘”–Š‡Š‘•–ȋEǦ Ȍǡ’ƒ”ƒ•‹–‹œ‡†Š‘•–ȋEǦ Ȍƒ†’ƒ”ƒ•‹–‘‹†ȋEǦȌ ‘—‹–‹‡•ǡƒ†

„”‘‡ ‹– †‘™ ‹–‘ –Š‡ ‘’‘‡– ‘ˆ EǦ†‹˜‡”•‹–›†—‡–‘•’‡ ‹‡•–—”‘˜‡”ȋE Ȍǡ –Šƒ– ‹•ǡ

•’‡ ‹‡•”‡’Žƒ ‡‡–ǡƒ†–Š‡ ‘’‘‡–‘ˆ–Š‡EǦ†‹˜‡”•‹–›†—‡–‘‡•–‡†‡••ȋEȌǡ–Šƒ–‹•ǡ •’‡ ‹‡•Ž‘••ȋƒ•‡Ž‰ƒʹͲͳͲȌǤ ƒ††‹–‹‘ǡ–‘Šƒ˜‡ƒ‡ƒ•—”‡‘ˆ†‹˜‡”•‹–›ˆ‘”‡ƒ Š•‹–‡Ǧ–‘Ǧ•‹–‡

‘’ƒ”‹•‘ǡ™‡ ƒŽ —Žƒ–‡†EǦ†‹˜‡”•‹–›‘ˆ’ƒ‹”™‹•‡ ‘’ƒ”‹•‘•ˆ‘”ƒŽŽŠ‘•–•ȋE Ȍǡ’ƒ”ƒ•‹–‹œ‡†

Š‘•–•ȋE Ȍƒ†’ƒ”ƒ•‹–‘‹†•ȋEȌǤŠ‡•‡ƒƒŽ›•‡•™‡”‡ ‘†— –‡†™‹–Š„‡–ƒ’ƒ‹”ͳǤ͵ȋƒ•‡Ž‰ƒ ƒ†”‡ʹͲͳʹȌˆ‘”ȋ‘”‡‡ƒʹͲͳͶȌǤ‘””‡Žƒ–‹‘•„‡–™‡‡Š‘•–EǦ†‹˜‡”•‹–›ȋƒŽŽŠ‘•–•

 81 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

ȋE Ȍƒ†’ƒ”ƒ•‹–‹œ‡†Š‘•–•ȋE Ȍ•‡’ƒ”ƒ–‡Ž›Ȍƒ†’ƒ”ƒ•‹–‘‹†EǦ†‹˜‡”•‹–›ȋEȌ™‡”‡–‡•–‡†™‹–Š ’‡ƒ”ƒǯ• ‘””‡Žƒ–‹‘Ǥ ‘ ƒƒŽ›œ‡ †‹•–ƒ ‡ †‡ ƒ› ‘ˆ –Š‡ †‹ˆˆ‡”‡– ‘’‘‡–• ‘ˆ EǦ †‹˜‡”•‹–›ǡ ™‡ ƒŽ —Žƒ–‡† ‘””‡Žƒ–‹‘• „‡–™‡‡ ‰‡‘‰”ƒ’Š‹ ƒŽ †‹•–ƒ ‡ƒ†Š‘•–EǦ†‹˜‡”•‹–›

ȋE Ȍǡ ’ƒ”ƒ•‹–‹œ‡† Š‘•– EǦ†‹˜‡”•‹–› ȋE Ȍ ƒ† ’ƒ”ƒ•‹–‘‹† EǦ†‹˜‡”•‹–› ȋEȌ ™‹–Š ’‡ƒ”ƒǯ• ‘””‡Žƒ–‹‘Ǥ



Interactions

‡–‡•–‡†™Š‡–Š‡”’ƒ”ƒ•‹–‹•”ƒ–‡™ƒ•”‡Žƒ–‡†–‘Š‘•–ƒ„—†ƒ ‡™‹–Šƒ ‘†‡ŽǤ  ‘†‡Ž™ƒ•ƒŽ•‘—•‡†–‘ƒƒŽ›œ‡–Š‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡–Š‡ƒ„—†ƒ ‡‘ˆ‡ƒ Š•’‡ ‹‡•ƒ† ‹–•’ƒ”ƒ•‹–‹•”ƒ–‡Ǥ

•™‹–ŠŠ‘•–ƒ†’ƒ”ƒ•‹–‘‹† ‘—‹–‹‡•ǡ™‡–‡•–‡†–Š‡“—ƒŽ‹–›‘ˆ‘—”‹–‡”ƒ –‹‘•ƒ’Ž‹‰ ™‹–Š”ƒ”‡ˆƒ –‹‘ —”˜‡•—•‹‰˜‡‰ƒʹǤͲǦͳͲȋ•ƒ‡‡–ƒŽǤʹͲͳͷȌˆ‘”ȋ‘”‡‡ƒʹͲͳͶȌǤ

‡„—‹Ž–ƒ‹–‡”ƒ –‹‘‡–™‘”ˆ‘”‡ƒ Š’Ž‘–ȋʹͷŽ‘ ƒŽ™‡„•Ȍǡƒ†ƒ”‡‰‹‘ƒŽ™‡„’‘‘Ž‹‰ –Š‡ †ƒ–ƒ ˆ”‘ ƒŽŽ ’Ž‘–•Ǥ ‘” –Š‡ ”‡‰‹‘ƒŽ ™‡„ǡ ™‡ ƒŽ —Žƒ–‡† ‰‡‡”ƒŽ‹–› ȋ™‡‹‰Š–‡† ‡ƒ —„‡”‘ˆŠ‘•–•’‡”’ƒ”ƒ•‹–‘‹†Ȍǡ˜—Ž‡”ƒ„‹Ž‹–›ȋ™‡‹‰Š–‡†‡ƒ—„‡”‘ˆ’ƒ”ƒ•‹–‘‹†•’‡”

Š‘•–Ȍǡ ‹–‡”ƒ –‹‘ ‡˜‡‡•• ȋŠƒ‘ǯ• ‡˜‡‡••Ȍ ƒ† ʹǯ ȋƒ ‡ƒ•—”‡ ‘ˆ ‘˜‡”ƒŽŽ ‡–™‘” •’‡ ‹ƒŽ‹œƒ–‹‘ǢŽò–Š‰‡‡–ƒŽǤʹͲͲ͸ȌǤŠ‡ǡ–‘‘„–ƒ‹ƒ‡ƒ•—”‡‘ˆ•‹‰‹ˆ‹ ƒ ‡ˆ‘”ƒŽŽ–Š‡•‡ ‡–”‹ •™‡”—ƒ—ŽŽ‘†‡ŽȋͳͲͲͲ‹–‡”ƒ–‹‘•Ȍ™Š‹ Šƒ‹–ƒ‹•ƒ”‰‹ƒŽ –‘–ƒŽ• ƒ† ‘‡ –ƒ ‡ ‘ˆ –Š‡ ‘”‹‰‹ƒŽ ‡–™‘” ȋžœ“—‡œ ‡– ƒŽǤ ʹͲͲ͹ȌǤ ŽŽ ‡–™‘” ƒƒŽ›•‹• ™‡”‡ ‘†— –‡†™‹–Š„‹’ƒ”–‹–‡ʹǤͲʹȋ‘”ƒ‡–ƒŽǤʹͲͲͺȌˆ‘”ȋ‘”‡‡ƒʹͲͳͶȌǤ

‡ ‡š’Ž‘”‡† Ž‘ ƒŽ ™‡„ †‹••‹‹Žƒ”‹–› „› ‡ƒ•—”‹‰ EǦ†‹˜‡”•‹–›‘ˆ‹–‡”ƒ –‹‘•ƒ‘‰Ž‘ ƒŽ

‡–™‘”• ȋEȌǤ ‘ŽŽ‘™‹‰ ‘‹•‘– ‡– ƒŽǤ ȋʹͲͳʹȌǡ –Š‹• †‹••‹‹Žƒ”‹–› ™ƒ• •—„†‹˜‹†‡† ‹–‘

†‹••‹‹Žƒ”‹–›†—‡–‘•’‡ ‹‡•–—”‘˜‡”ȋEȌƒ††‹••‹‹Žƒ”‹–›†—‡–‘‹–‡”ƒ –‹‘”‡™‹”‹‰ǡ–Šƒ–

‹•ǡ ‘‘ Š‘•– ƒ† ’ƒ”ƒ•‹–‘‹† •’‡ ‹‡• ‹–‡”ƒ –‹‰ †‹ˆˆ‡”‡–Ž› ‹†‹ˆˆ‡”‡–’Ž‘–•ȋEȌǤ

‘‡–‹‡•‹–™ƒ•‘–’‘••‹„Ž‡–‘ ƒŽ —Žƒ–‡E†—‡–‘–Š‡Žƒ ‘ˆ•Šƒ”‡†•’‡ ‹‡•„‡–™‡‡–™‘

’Ž‘–•Ǥ –Š‡•‡ ƒ•‡•ǡƒŽŽ‡–™‘”˜ƒ”‹ƒ„‹Ž‹–›ȋEȌ™ƒ•ƒ••‹‰‡†–‘•’‡ ‹‡•–—”‘˜‡”ȋEȌǤ 

ƒ††‹–‹‘ǡ ƒ† ƒ‰ƒ‹ ˆ‘ŽŽ‘™‹‰ ‘‹•‘– ‡– ƒŽǤ ȋʹͲͳʹȌǡ ™‡ ƒŽ —Žƒ–‡† Eǯǡ ƒ ‡ƒ•—”‡ ‘ˆ †‹••‹‹Žƒ”‹–›‘ˆ‹–‡”ƒ –‹‘•„‡–™‡‡Ž‘ ƒŽ™‡„•ƒ†–Š‡‹” ‘—–‡”’ƒ”–•‹–Š‡”‡‰‹‘ƒŽ™‡„Ǥ ŽŽ–Š‡•‡‹†‡š‡•™‡”‡ ƒŽ —Žƒ–‡†™‹–Š„‡–ƒŽ‹’ƒ ƒ‰‡ˆ‘”ȋ‘”‡‡ƒʹͲͳͶȌǤ

 82  Šƒ’–‡”͵

Š‡”‡Žƒ–‹‘•Š‹’‘ˆ‹–‡”ƒ –‹‘EǦ†‹˜‡”•‹–›ȋEȌ™‹–Š’ƒ”ƒ•‹–‹œ‡†Š‘•–EǦ†‹˜‡”•‹–›ȋE Ȍƒ†

’ƒ”ƒ•‹–‘‹†EǦ†‹˜‡”•‹–›ȋEȌ™ƒ•–‡•–‡†™‹–ŠƒŽ‹‡ƒ”‘†‡ŽǤ‹ ‡„‘–Š‡š’Žƒƒ–‘”›˜ƒ”‹ƒ„Ž‡• ™‡”‡•Ž‹‰Š–Ž› ‘””‡Žƒ–‡†ȋ”αͲǤ͵͹Ȍǡ™‡ ƒŽ —Žƒ–‡†–Š‡˜ƒ”‹ƒ ‡‹ˆŽƒ–‹‘ˆƒ –‘”–‘ƒ‡•—”‡ ‘Ž‹‡ƒ”‹–› ™ƒ•Ž‘™ȋ α ͳǤͳ͹ȌǤ ‡” ‡–ƒ‰‡ ‘ˆ˜ƒ”‹ƒ ‡‡š’Žƒ‹‡† „› –Š‡ ‘†‡Ž ™ƒ• ƒŽ —Žƒ–‡†™‹–Špmvd‡–”‹ —•‹‰”‡Žƒ‹’‘’ƒ ƒ‰‡ȋ ”ڏ’‹‰ʹͲͲ͸Ȍˆ‘”ȋ‘”‡‡ƒ ʹͲͳͶȌǤ ‘ ƒƒŽ›œ‡ †‹•–ƒ ‡†‡ ƒ›‘ˆ–Š‡ †‹ˆˆ‡”‡– ‘’‘‡–• ‘ˆ ‡–™‘” EǦ†‹˜‡”•‹–›ǡ ™‡

ƒ†‡ ‘””‡Žƒ–‹‘•„‡–™‡‡‰‡‘‰”ƒ’Š‹ ƒŽ†‹•–ƒ ‡ƒ†‡–™‘”EǦ†‹˜‡”•‹–›ȋEȌǡEǦ†‹˜‡”•‹–›

†—‡ –‘ •’‡ ‹‡• –—”‘˜‡” ȋEȌ ƒ† EǦ†‹˜‡”•‹–› †—‡ –‘ ”‡™‹”‹‰ ȋEȌ —•‹‰ ’‡ƒ”ƒǯ• ‘””‡Žƒ–‹‘Ǥ





Figure 3.1ƒ”‡ˆƒ –‹‘ —”˜‡• ˆ‘” Š‘•– ƒ† ’ƒ”ƒ•‹–‘‹† •’‡ ‹‡• ƒ†  –Š‡‹”‹–‡”ƒ –‹‘•Ǥƒ•Š‡†Ž‹‡•‹†‹ ƒ–‡ͻͷΨ ‘ˆ‹†‡ ‡‹–‡”˜ƒŽ•Ǥ



 3.3.Results

‡‘„–ƒ‹‡†ͳͷͶͳ‡•–•ȋͶͻͷͶ ‡ŽŽ•Ȍˆ”‘ͶͳŠ‘•–•’‡ ‹‡•Ǥ‡˜‡–‡‡‘ˆ–Š‡•‡•’‡ ‹‡•™‡”‡ „‡‡•ȋ‡‰ƒ Š‹ŽŽ‹†ƒ‡Ȃͳͷ•’Ǥǡ‘ŽŽ‡–‹†ƒ‡Ȃʹ•’ǤȌƒ†ʹͶ™‡”‡™ƒ•’•ȋ”ƒ„”‘‹†ƒ‡Ȃͳʹ•’Ǥǡ ‘’‹Ž‹†ƒ‡Ȃͷ•’Ǥǡ‡•’‹†ƒ‡Ȃͷ•’Ǥǡ’Š‡ ‹†ƒ‡Ȃͳ•’Ǥǡ’—Ž‹ ‹†ƒ‡Ȃͳ•’ǤȌǤ™‡–›Ǧˆ‘—”

 83 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

Š‘•– •’‡ ‹‡• Šƒ†ǡ ƒ– Ž‡ƒ•–ǡ ‘‡ ‹†‹˜‹†—ƒŽ ’ƒ”ƒ•‹–‹œ‡†Ǥ ‡ ˆ‘—† ʹ͸ ’ƒ”ƒ•‹–‘‹† •’‡ ‹‡• ȋͳ͸ ™ƒ•’•ǡ ͷ ˆŽ‹‡•ǡ ͵ „‡‡–Ž‡•ǡ ͳ „‡‡ ƒ† ͳ ‹–‡Ȍ ƒ••‘ ‹ƒ–‡† –‘ –Š‡ ‡•–•ǡ ”‡•—Ž–‹‰ ‹ ͸ͷͶ ’ƒ”ƒ•‹–‹œ‡† ‡ŽŽ•Ǥˆ–Š‡•‡ʹ͸•’‡ ‹‡•ǡʹͲ™‡”‡’ƒ”ƒ•‹–‘‹†•ǡͶ™‡”‡ Ž‡’–‘’ƒ”ƒ•‹–‡•ǡͳ™ƒ•ƒ ’”‡†ƒ–‘”ƒ†ͳ™ƒ•ƒ• ƒ˜‡‰‡”Ǥ˜‡”ƒŽŽ’ƒ”ƒ•‹–‹•”ƒ–‡™ƒ•ͳ͵ǤʹΨǤ



3.3.1. Hostcommunity

ƒ”‡ˆƒ –‹‘ —”˜‡•‹†‹ ƒ–‡–Šƒ–‘—”Š‘•– ‘—‹–›™ƒ•”‡Žƒ–‹˜‡Ž›™‡ŽŽ•ƒ’Ž‡†ȋ ‹‰ǤͳȌǤ• ‡š’‡ –‡†ǡ Š‘•– ƒ„—†ƒ ‡ ƒ† ”‹ Š‡•• •Š‘™‡† ƒ ’‘•‹–‹˜‡ ”‡Žƒ–‹‘•Š‹’ ȋ’‡ƒ”ƒ ‘””‡Žƒ–‹‘ǣ ƒŽŽ Š‘•–•ǣ ”Š‘α ͲǤͷͻǡ ’δ ͲǤͲͲʹǢ ’ƒ”ƒ•‹–‹œ‡† Š‘•–•ǣ ”Š‘α ͲǤͷ͹ǡ ’δ ͲǤͲͲ͵Ǣ α ʹͷ ’Ž‘–•ȌǤ ‘‰ –Š‡ ͳͳ ‘•– ƒ„—†ƒ– Š‘•– •’‡ ‹‡• ȋεͳͲͲ ‡ŽŽ•Ȍǡ ˆ‹˜‡ •Š‘™‡† •’ƒ–‹ƒŽ ƒ—–‘ ‘””‡Žƒ–‹‘ǡ™Š‹Ž‡–Š‡”‡ƒ‹‹‰•‹š†‹†‘–Ǥ

‘•–EǦ†‹˜‡”•‹–›™ƒ•Š‹‰ŠȋEǦ αͲǤͺͻȌǡ‘•–Ž›†—‡–‘•’‡ ‹‡•–—”‘˜‡”ȋE Ǧ αͲǤͺͷȌǡ™‹–Šƒ

’‘‘” ‘–”‹„—–‹‘ ‘ˆ ‡•–‡†‡•• ȋEǦ α ͲǤͲͶȌ ȋƒ„Ž‡ ͳȌǤ ƒ”ƒ•‹–‹œ‡† Š‘•– EǦ†‹˜‡”•‹–› ™ƒ•

‡“—ƒŽŽ› Š‹‰Š ȋEǦ α ͲǤͻͳȌǡ ƒ‰ƒ‹ ‘•–Ž› †—‡ –‘ •’‡ ‹‡• –—”‘˜‡” ȋE Ǧ α ͲǤͺ͹Ȍǡ ™‹–Š ƒ

™‡‡ ‘–”‹„—–‹‘ ‘ˆ ‡•–‡†‡•• ȋEǦ αͲǤͲͶȌǤŠƒ–‹•ǡŽ‘ ƒŽŠ‘•– ‘—‹–‹‡•†‹ˆˆ‡”‹ •’‡ ‹‡• ‘’‘•‹–‹‘ǡ‘–†—‡–‘•’‡ ‹‡•Ž‘••ǡ„—–„‡ ƒ—•‡–Š‡”‡‹•ƒ•’‡ ‹‡•”‡’Žƒ ‡‡–ƒ† –Š—•–Š‡‹†‡–‹–›‘ˆ–Š‡•’‡ ‹‡•†‹ˆˆ‡”•ƒ‘‰•‹–‡•Ǥ



3.3.2. Parasitoidcommunity

ƒ”‡ˆƒ –‹‘ —”˜‡• ‹†‹ ƒ–‡ –Šƒ– ‘—” ’ƒ”ƒ•‹–‘‹† ‘—‹–› ™ƒ• ”‡Žƒ–‹˜‡Ž› ™‡ŽŽ •ƒ’Ž‡†ǡ ˆ‘ŽŽ‘™‹‰ ƒ ’ƒ––‡” ’ƒ”ƒŽŽ‡Ž –‘ –Š‡ Š‘•– ‘—‹–› ȋ ‹‰Ǥ ͳȌǤ •‡š’‡ –‡†ǡ’ƒ”ƒ•‹–‘‹† ƒ„—†ƒ ‡ƒ†”‹ Š‡•••Š‘™‡†ƒ’‘•‹–‹˜‡ ‘””‡Žƒ–‹‘ȋ”αͲǤͷ͹ǡ’δͲǤͲͲ͵ȌǤŠ‡’ƒ”ƒ•‹–‘‹† ‘—‹–›™ƒ•Š‹‰ŠŽ›†‡’‡†‡–‘–Š‡Š‘•– ‘—‹–›ǣ–Š‡”‡™ƒ•ƒ•‹‰‹ˆ‹ ƒ–’‘•‹–‹˜‡ ‘””‡Žƒ–‹‘ „‡–™‡‡ ’ƒ”ƒ•‹–‘‹† ƒ† Š‘•– ƒ„—†ƒ ‡ ȋƒŽŽ Š‘•–•ǣ ”Š‘α ͲǤ͸ʹǡ ’α ͲǤͲͲͳǡ αʹͷ ’Ž‘–•Ȍǡƒ†„‡–™‡‡’ƒ”ƒ•‹–‘‹†ƒ†Š‘•–”‹ Š‡••ȋƒŽŽŠ‘•–•ǣ”Š‘αͲǤ͹ͷǡ’δͲǤͲͲͲͳȌǤ

 84 









 Figure3.2‡‰‹‘ƒŽŠ‘•–Ǧ’ƒ”ƒ•‹–‘‹†•™‡„ȋαʹͷ’Ž‘–•ȌǤ’‡ ‹‡•ƒ‡• ƒ„‡ˆ‘—†‹’’‡†‹šǡƒ„Ž‡•ͳƒ†ʹǤ‹†–Š‘ˆ „Žƒ „ƒ”•†‡‘–‡•ƒ„—†ƒ ‡‘ˆ‹–‡”ƒ –‹‰•’‡ ‹‡•ȋ‘Ž›’ƒ”ƒ•‹–‹œ‡†Š‘•–‹†‹˜‹†—ƒŽ•‹ Ž—†‡†ȌǤ 

 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡• 

ƒ”ƒ•‹–‘‹† EǦ†‹˜‡”•‹–›™ƒ•ƒ‰ƒ‹ Š‹‰Š ȋEǦα ͲǤͻͳȌǡ ƒ† ‘•–Ž› ƒ––”‹„—–ƒ„Ž‡ –‘ •’‡ ‹‡•

–—”‘˜‡” ȋE Ǧα ͲǤͺͷȌǡ ”ƒ–Š‡” –Šƒ ‡•–‡†‡•• ȋEǦα ͲǤͲ͸Ȍ ȋƒ„Ž‡ ͳȌǤ EǦ†‹˜‡”•‹–› ‘ˆ

’ƒ”ƒ•‹–‘‹†• ȋEȌ ™ƒ• •‹‰‹ˆ‹ ƒ–Ž› ‘””‡Žƒ–‡† –‘ EǦ†‹˜‡”•‹–› ‘ˆ ƒŽŽ Š‘•–• ȋE Ȍȋ”Š‘αͲǤʹͳǡ ’α

ͲǤͲͲͲʹȌƒ†EǦ†‹˜‡”•‹–›‘ˆ’ƒ”ƒ•‹–‹œ‡†Š‘•–•ȋE Ȍȋ”Š‘αͲǤ͵͹ǡ’δͲǤͲͲͲͳȌǤ





Figure3.3‡Žƒ–‹‘•Š‹’„‡–™‡‡‡–™‘”EǦ†‹˜‡”•‹–›ƒ†ȋƒȌ’ƒ”ƒ•‹–‹œ‡†Š‘•–EǦ†‹˜‡”•‹–› ƒ†ȋ„Ȍ’ƒ”ƒ•‹–‘‹†EǦ†‹˜‡”•‹–›Ǥƒ”–‹ƒŽ”‡‰”‡••‹‘’Ž‘–•‘„–ƒ‹‡†ˆ”‘ƒŽ‹‡ƒ”‘†‡Ž™Š‡ –Š‡‡ˆˆ‡ –‘ˆ–Š‡‘–Š‡”‡š’Žƒƒ–‘”›˜ƒ”‹ƒ„Ž‡‹•”‡‘˜‡†Ǥ

  3.3.3. Interactions

ƒ”‡ˆƒ –‹‘ —”˜‡••Š‘™–Šƒ–ǡ ‘–”ƒ”›–‘Š‘•–ƒ†’ƒ”ƒ•‹–‘‹†•’‡ ‹‡•ǡ‹–‡”ƒ –‹‘•™‡”‡‘– •—ˆˆ‹ ‹‡–Ž› •ƒ’Ž‡† ȋ ‹‰Ǥ ͳȌǤ ƒ”ƒ•‹–‹• ”ƒ–‡ ™ƒ• ‘– ‡š’Žƒ‹‡†„›Š‘•–ƒ„—†ƒ ‡ ƒ– –Š‡ ’Ž‘– Ž‡˜‡Ž ȋ ǣ ’•‡—†‘ǦʹαͲǤͲͲͷǡ’αͲǤ͹ͷǡ α ʹͷ ’Ž‘–•ȌǤ ‘™‡˜‡”ǡ ƒ– •’‡ ‹‡• Ž‡˜‡Žǡ ’ƒ”ƒ•‹–‹• ”ƒ–‡ ‘ˆƒ‰‹˜‡Š‘•– •’‡ ‹‡• ™ƒ• ’ƒ”–‹ƒŽŽ› ‡š’Žƒ‹‡†„›‹–•ƒ„—†ƒ ‡ ȋ ǣ ’•‡—†‘ǦʹαͲǤͳ͵ǡ’αͲǤͲʹǡαʹͶ•’‡ ‹‡•ȌǤ

‡ ‘„–ƒ‹‡† ƒ ”‡‰‹‘ƒŽ ™‡„ ™‹–Š–Š‡†ƒ–ƒ ‘ˆ ƒŽŽ –Š‡ ’Ž‘–• •ƒ’Ž‡† ȋ ‹‰Ǥ ʹȌǤ ŽŽ ‡–™‘” ‡–”‹ • ƒŽ —Žƒ–‡††‡‘–‡ ƒ Š‹‰ŠŽ›•’‡ ‹ƒŽ‹œ‡†”‡‰‹‘ƒŽ™‡„ȋ‰‡‡”ƒŽ‹–›α ʹǤͻͻ͹ǡ’δͲǤͲͲͲͳǢ

˜—Ž‡”ƒ„‹Ž‹–›α ʹǤͺͳ͵ǡ ’δ ͲǤͲͲͲͳǢ ‹–‡”ƒ –‹‘ ‡˜‡‡••α ͲǤͶ͸ͺǡ ’δ ͲǤͲͲͲͳǢ ʹǯα ͲǤ͸ͷǡ ’δ ͲǤͲͲͲͳȌǤ

 86 Šƒ’–‡”͵

‡ˆ‘—†͹ʹ†‹ˆˆ‡”‡–•’‡ ‹ˆ‹ ‹–‡”ƒ –‹‘•Ǥ ‘™‡˜‡”ǡ‘•–‘ˆ–Š‡ȋ͸͵ǤͻΨȌ™‡”‡ˆ‘—†‹ Œ—•–‘‡’Ž‘–ǡƒ†ͶͲǤ͵Ψ‘ˆ–Š‡™‡”‡ˆ‘—†‘Ž›‘ ‡Ǥƒ‹”™‹•‡‹–‡”ƒ –‹‘†‹••‹‹Žƒ”‹–›

„‡–™‡‡ ’Ž‘–• ™ƒ• Š‹‰Š ȋEǡ ‡ƒrǣͲǤͺ͵άͲǤͳ͸ȌǤ‘•–‘ˆ–Š‹•†‹••‹‹Žƒ”‹–›™ƒ•†—‡–‘

•’‡ ‹‡•–—”‘˜‡”ȋEǡ‡ƒrǣͲǤ͸ͲάͲǤ͵ʹȌǡ™‹–Šƒ— ŠŽ‡••‡” ‘–”‹„—–‹‘‘ˆ‹–‡”ƒ –‹‘

”‡™‹”‹‰ȋEǡ‡ƒrǣͲǤʹ͵άͲǤ͵ʹȌǤEǯ˜ƒŽ—‡•™‡”‡Ž‘™ȋ„‡–™‡‡Ͳƒ†ͲǤ͵ͺȌǡ‹†‹ ƒ–‹‰ –Šƒ–•’‡ ‹‡•‹–‡”ƒ –•‹‹Žƒ”Ž›ƒ ”‘••Ž‘ ƒŽ‡–™‘”•Ǥ



3.3.4. RelationshipbetweenspeciesandinteractionEǦdiversity

ƒ”ƒ•‹–‹œ‡† Š‘•– EǦ†‹˜‡”•‹–› ȋE Ȍ ƒ† ’ƒ”ƒ•‹–‘‹† EǦ†‹˜‡”•‹–› ȋEȌ ™‡”‡ ‰‘‘† ’”‡†‹ –‘”• ‘ˆ

ʹ ‡–™‘”EǦ†‹˜‡”•‹–›ȋEȌȋ” αͲǤ͸͵ǡ’δͲǤͲͲͲͳȌȋ ‹‰Ǥ͵ȌǤ ‘™‡˜‡”ǡ™Š‹Ž‡’ƒ”ƒ•‹–‹œ‡†Š‘•–EǦ †‹˜‡”•‹–› ‡š’Žƒ‹‡† ͶͻǤͳΨ ‘ˆ –Š‡ ˜ƒ”‹ƒ ‡ǡ ’ƒ”ƒ•‹–‘‹† EǦ†‹˜‡”•‹–› ‘Ž› ‡š’Žƒ‹‡† ͳ͵ǤͻΨǡ ‹†‹ ƒ–‹‰–Šƒ–’ƒ”ƒ•‹–‹œ‡†Š‘•–EǦ†‹˜‡”•‹–›‹•ƒ„‡––‡”’”‡†‹ –‘”‘ˆ‡–™‘”EǦ†‹˜‡”•‹–›–Šƒ ’ƒ”ƒ•‹–‘‹†EǦ†‹˜‡”•‹–›Ǥ



Table 3.1  ‘•–ǡ ’ƒ”ƒ•‹–‹œ‡† Š‘•– ƒ† ’ƒ”ƒ•‹–‘‹† EǦ†‹˜‡”•‹–›  „ƒ•‡† ‘ ޔ‡•‡ †‹••‹‹Žƒ”‹–› ȋEȌǡ ’ƒ”–‹–‹‘‡† ‹–‘ –Š‡

‘’‘‡–‘ˆEǦ†‹˜‡”•‹–›†—‡–‘•’‡ ‹‡•–—”‘˜‡”ȋE Ȍƒ†  –Š‡ ‘’‘‡– ‘ˆ EǦ†‹˜‡”•‹–› †—‡ –‘ ‡•–‡†‡•• ȋEȌ ȋƒ•‡Ž‰ƒʹͲͳͲȌ 

 E E  E ‘•–• ͲǤͺͻ ͲǤͺͷ ͲǤͲͶ ƒ”ƒ•‹–‹œ‡†Š‘•–• ͲǤͻͳ ͲǤͺ͹ ͲǤͲͶ ƒ”ƒ•‹–‘‹†• ͲǤͻͳ ͲǤͺͷ ͲǤͲ͸ 

 3.3.5. DistancedecayofEǦdiversity

‘•–EǦ†‹˜‡”•‹–›ȋE ǣ”Š‘αͲǤʹͻǡ’δͲǤͲͲͲͳȌƒ†’ƒ”ƒ•‹–‹œ‡†Š‘•–EǦ†‹˜‡”•‹–›ȋE ǣ”Š‘αͲǤͳͻǡ ’α ͲǤͲͲͳȌ ™‡”‡ ’‘•‹–‹˜‡Ž› ‘””‡Žƒ–‡† –‘ ‰‡‘‰”ƒ’Š‹ ƒŽ †‹•–ƒ ‡Ǥ ‘˜‡”•‡Ž›ǡ ’ƒ”ƒ•‹–‘‹† EǦ

†‹˜‡”•‹–›ȋEǣ”Š‘αͲǤͲͺǡ’αͲǤͳͺȌȋ ‹‰ǤͶȌ™ƒ•‘”‡Žƒ–‡†–‘†‹•–ƒ ‡Ǥ‹‹Žƒ”Ž›ǡ‡–™‘”EǦ

†‹˜‡”•‹–› ȋEǣ ”Š‘αͲǤͲ͹ǡ ’α ͲǤʹͲȌ ƒ† ‹–• –™‘ ‘’‘‡–•ǡ EǦ†‹˜‡”•‹–›†—‡–‘•’‡ ‹‡•  87 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡• 

–—”‘˜‡”ȋEǣ”Š‘αͲǤͳͲǡ’αͲǤͲͺȌƒ†EǦ†‹˜‡”•‹–›†—‡–‘”‡™‹”‹‰ȋEǣ”Š‘αǦͲǤͲ͸ǡ’αͲǤʹͻȌ

™‡”‡‘–”‡Žƒ–‡†–‘‰‡‘‰”ƒ’Š‹ ƒŽ†‹•–ƒ ‡ȋ ‹‰ǤͶȌǤ





 Figure 3.4‹•–ƒ ‡†‡ ƒ›‘ˆŠ‘•– EǦ†‹˜‡”•‹–› ȋE Ȍǡ ’ƒ”ƒ•‹–‹œ‡† Š‘•– EǦ†‹˜‡”•‹–› ȋE Ȍǡ ’ƒ”ƒ•‹–‘‹† EǦ†‹˜‡”•‹–› ȋEȌǡ ‡–™‘” EǦ†‹˜‡”•‹–› ȋEȌǡ ‘’‘‡– ‘ˆ ‡–™‘” EǦ†‹˜‡”•‹–›  †—‡ –‘•’‡ ‹‡•–—”‘˜‡” ȋEȌǡ ƒ† ‘’‘‡– ‘ˆ EǦ†‹˜‡”•‹–› †—‡ –‘ ”‡™‹”‹‰ ȋEȌǤ ȋ–ƒ–‹•–‹ ƒŽ•‹‰‹ˆ‹ ƒ ‡Ǣȗȗȗǣ’ζͲǤͲͲͳǢȗȗȗȗǣ’ζͲǤͲͲͲͳȌǤ

  3.4. Discussion

ƒ”‡ˆƒ –‹‘ ”‡•—Ž–•‹†‹ ƒ–‡–Šƒ– •ƒ’Ž‹‰ ʹͷ ’Ž‘–•‘˜‡”ƒƒ”‡ƒ ‘ˆ ͵ʹ ʹ’”‘˜‹†‡† ƒ ”‡Žƒ–‹˜‡Ž›ƒ†‡“—ƒ–‡ Šƒ”ƒ –‡”‹œƒ–‹‘‘ˆ–Š‡Š‘•–ƒ†’ƒ”ƒ•‹–‘‹† ‘—‹–›‹‘—”•–—†›ƒ”‡ƒǤ  –Š‡‘–Š‡”Šƒ†ǡ –Š‡ ”ƒ”‡ˆƒ –‹‘ —”˜‡ ˆ‘”‹–‡”ƒ –‹‘• •—‰‰‡•–• –Šƒ– ’”‘’‡” Šƒ”ƒ –‡”‹œƒ–‹‘ ‘ˆ –Š‡ ‹–‡”ƒ –‹‘ ‘—‹–› ™‘—Ž† ”‡“—‹”‡ƒ— Š‰”‡ƒ–‡” •ƒ’Ž‹‰ ‡ˆˆ‘”–Ǥ‘‹•‘–‡–ƒŽǤȋʹͲͳʹȌ‘„–ƒ‹‡†•‹‹Žƒ””‡•—Ž–•™Š‡–Š‡›•ƒ’Ž‡†ƒƒŽǦ‡ –‘’ƒ”ƒ•‹–‡ ‹–‡”ƒ –‹‘•‹ͳͳ͵Ž‘ ƒŽ‹–‹‡•‘˜‡”‡–”ƒŽ—”‘’‡Ǥ ‘™‡˜‡”ǡ”ƒ”‡ˆƒ –‹‘ —”˜‡•ƒ”‡’”‘„ƒ„Ž› ‹ŽŽǦ•—‹–‡† –‘ ƒ••‡•••ƒ’Ž‹‰ ‘’Ž‡–‡‡•• ‘ˆ ‡ ‘Ž‘‰‹ ƒŽ ‹–‡”ƒ –‹‘•ǡ ƒ• –Š‡› ƒ••—‡ ƒŽŽ

 88 Šƒ’–‡”͵

•’‡ ‹‡• ƒ‹–‡”ƒ –™‹–ŠƒŽŽ•’‡ ‹‡•‹–Š‡‘’’‘•‹–‡–”‘’Š‹ Ž‡˜‡Žǡ–Š—•‘˜‡”Ž‘‘‹‰ˆ‘”„‹††‡ Ž‹•”‡•—Ž–‹‰ˆ”‘†‹ˆˆ‡”‡–‹†•‘ˆ‹–‡”ƒ –‹‘ ‘•–”ƒ‹–•ȋ ‘”†ƒ‘‡–ƒŽǤʹͲͲ͵ȌǤŠ‹•‹• ‡•’‡ ‹ƒŽŽ›–”—‡‹ƒ•›•–‡Ž‹‡‘—”•ǡ™‹–ŠŠ‹‰ŠŽ‡˜‡Ž•‘ˆ•’‡ ‹ƒŽ‹œƒ–‹‘ ƒ† Ž‘™ Ž‡˜‡Ž• ‘ˆ ”‡™‹”‹‰Ǥ

‘—” ‘—‹–›ǡ’ƒ”ƒ•‹–‹•”ƒ–‡™ƒ•‘–‹ˆŽ—‡ ‡†„›Š‘•–ƒ„—†ƒ ‡ƒ––Š‡ ‘—‹–› Ž‡˜‡ŽǤ–Š‡”•–—†‹‡•‘ ƒ˜‹–›Ǧ‡•–‹‰„‡‡•ƒ†™ƒ•’•Šƒ˜‡ˆ‘—†•‹‹Žƒ””‡•—Ž–•ȋ›Ž‹ƒƒ‹• ‡–ƒŽǤʹͲͲ͸ǡŽ„”‡ Š–‡–ƒŽǤʹͲͲ͹Ȍǡ›‡–‘–Š‡”•Šƒ˜‡ˆ‘—†ƒ’‘•‹–‹˜‡ ‘””‡Žƒ–‹‘„‡–™‡‡–Š‡ ȋ–‡ˆˆƒǦ‡™‡–‡” ʹͲͲ͵ȌǤ ‘™‡˜‡”ǡ ‹ ‘—” ‘—‹–›ǡ ‘”‡ ƒ„—†ƒ– Š‘•– •’‡ ‹‡• Šƒ† Š‹‰Š‡”’ƒ”ƒ•‹–‹•”ƒ–‡Ǥ™‘‡ Šƒ‹••ƒ›‡š’Žƒ‹–Š‹•”‡•—Ž–Ǥ ‹”•–ǡ’”‡›ƒ„—†ƒ ‡Šƒ• „‡‡ †‡‘•–”ƒ–‡† –‘ ‹ˆŽ—‡ ‡ ’”‡› Š‘‹ ‡ǡ ƒ• ’”‡†ƒ–‘”• ™‹–Š –Š‡ƒ„‹Ž‹–›–‘’”‡›‘ †‹ˆˆ‡”‡–Š‘•–•‘„–ƒ‹‰”‡ƒ–‡””‡–—”•„›Ž‡ƒ”‹‰Š‘™–‘Šƒ†Ž‡–Š‡‘•–ƒ„—†ƒ– ’”‡› ƒ† ˆ‘ —•‹‰ ‘ ‹– ȋ •Š‹‹ ƒ† Š‹ƒ†ƒ ʹͲͳʹȌǤ ‡ ‘†ǡ ˆ‘” •’‡ ‹ƒŽ‹•– ƒ–ƒ‰‘‹•–•ǡ ™‹–Š ƒ ”‡•–”‹ –‡†”ƒ‰‡‘ˆ’‘–‡–‹ƒŽ’”‡›•ȀŠ‘•–•ǡ‹–ƒ›„‡†‹ˆˆ‹ —Ž––‘„—‹Ž†ƒ•–ƒ„Ž‡’‘’—Žƒ–‹‘‘ƒ Ž‘ ƒŽŽ›”ƒ”‡Š‘•–Ǥ

Ž–Š‘—‰Š –Š‡ •–—†› ™ƒ• ‘†— –‡† ‹ ƒ ‘–‹—‘—• Šƒ„‹–ƒ–ǡ „‘–Š –Š‡Š‘•–ƒ†’ƒ”ƒ•‹–‘‹† ‘—‹–‹‡•ǡƒ•™‡ŽŽƒ•–Š‡‹”‹–‡”ƒ –‹‘•ǡ•Š‘™‡†•–”‘‰•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›ƒ–ƒŽ‘ ƒŽ• ƒŽ‡ ƒ• ‹†‹ ƒ–‡† „› –Š‡ Š‹‰Š EǦ†‹˜‡”•‹–›˜ƒŽ—‡•Ǥ ‹˜‡–Šƒ–„‡‡•ƒ†™ƒ•’•ƒ”‡Š‹‰ŠŽ›‘„‹Ž‡ ‘”‰ƒ‹••ǡ ‘‡ ‹‰Š– Šƒ˜‡ ‡š’‡ –‡† ƒ ‘”‡ —‹ˆ‘” Š‘•– †‹•–”‹„—–‹‘Ǥ Š‡ Ž‘ ƒŽ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ„‹Ž‹–› ˆ‘—† ‹†‹ ƒ–‡• ƒ •–”‘‰ ‡ˆˆ‡ – ‘ˆ Ž‘ ƒŽ ˆƒ –‘”•ǡ ƒ• ˆ‘—† ˆ‘” –Š‡ ‡–‹”‡ „‡‡ ‘—‹–›‹ƒ’”‡˜‹‘—••–—†›‹–Š‡•ƒ‡ƒ”‡ƒȋ‘”±Ǧ‘‰—‡”ƒ‡–ƒŽǤʹͲͳͶȌǤ

‘•– EǦ†‹˜‡”•‹–› ȋE Ȍ ƒ† ’ƒ”ƒ•‹–‹œ‡† Š‘•– EǦ†‹˜‡”•‹–› ȋE Ȍ ‹ ”‡ƒ•‡† ™‹–Š ‰‡‘‰”ƒ’Š‹ ƒŽ †‹•–ƒ ‡ǡ‹†‹ ƒ–‹‰–Šƒ–Š‘•–•’‡ ‹‡•–—”‘˜‡”‹ ”‡ƒ•‡•™‹–Š†‹•–ƒ ‡Ǥ–Š‡‘–Š‡”Šƒ†ǡ

’ƒ”ƒ•‹–‘‹†EǦ†‹˜‡”•‹–›ȋEȌƒ†‹–‡”ƒ –‹‘EǦ†‹˜‡”•‹–›ȋEȌǡƒ•™‡ŽŽƒ•‹–•–™‘ ‘’‘‡–•

ȋEǦ†‹˜‡”•‹–›†—‡–‘•’‡ ‹‡•–—”‘˜‡”ȋEȌƒ†EǦ†‹˜‡”•‹–›†—‡–‘”‡™‹”‹‰ȋEȌȌǡ†‹†‘–˜ƒ”› ™‹–Š‰‡‘‰”ƒ’Š‹ ƒŽ†‹•–ƒ ‡Ǥ˜‡–Š‘—‰Šˆ‡™•–—†‹‡•ƒ”‡ƒ˜ƒ‹Žƒ„Ž‡ˆ‘” ‘’ƒ”‹•‘ǡ†‹•–ƒ ‡ †‡ ƒ› ‘ˆ •’‡ ‹‡• ‘’‘•‹–‹‘ ‘ˆ –Š‡ Ž‘™‡” –”‘’Š‹  Ž‡˜‡Ž ƒ† ‘ˆ ‹–‡”ƒ –‹‘• •‡‡• –‘ „‡ ™‹†‡•’”‡ƒ†ȋ‘˜‘–›ʹͲͲͻǡƒ”•–‡•‡‡–ƒŽǤʹͲͳͶǡ”ÞŒ‡Ž•‰ƒƒ”†‡–ƒŽǤʹͲͳͷȌǤ‘’‘•‹–‹‘‘ˆ –Š‡Š‹‰Š‡”–”‘’Š‹ Ž‡˜‡Žǡ‘–Š‡‘–Š‡”Šƒ†ǡŠƒ•„‡‡ˆ‘—†–‘‡‹–Š‡”‹ ”‡ƒ•‡ȋ”ÞŒ‡Ž•‰ƒƒ”† ‡–ƒŽǤʹͲͳͷȌǡ†‡ ”‡ƒ•‡ȋ‘˜‘–›ʹͲͲͻȌ‘”‘–˜ƒ”›ȋƒ”•–‡•‡‡–ƒŽǤʹͲͳͶȌ™‹–Š‰‡‘‰”ƒ’Š‹ ƒŽ †‹•–ƒ ‡Ǥ‹• ”‡’ƒ ‹‡• ƒ‘‰–Š‡•‡ •–—†‹‡•ǡ‹ Ž—†‹‰‘—”•ǡƒ›„‡†—‡ –‘†‹ˆˆ‡”‡ ‡•‹ –Š‡ „‹‘Ž‘‰‹ ƒŽ •›•–‡• ”‡•‡ƒ” Š‡† ȋˆ”‘ ‘”‡ ‰‡‡”ƒŽ‹œ‡† ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘” ‹–‡”ƒ –‹‘• –‘

 89 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

‘”‡•’‡ ‹ƒŽ‹œ‡†Š‘•–Ǧ’ƒ”ƒ•‹–‡‹–‡”ƒ –‹‘•Ȍƒ†‹‰‡‘‰”ƒ’Š‹ ƒŽ †‹•–ƒ ‡ ȋˆ”‘ ͸ͲͲ  –‘ ͷͲͲȌǤ

—” ”‡•—Ž–• ƒ††‹–‹‘ƒŽŽ› •Š‘™ –Šƒ– ’ƒ”ƒ•‹–‘‹† ‘—‹–› ‹• Š‹‰ŠŽ›†‡’‡†‡–‘Š‘•– ‘—‹–›ǡ •‹ ‡ ƒ„—†ƒ ‡ǡ ”‹ Š‡•• ƒ† ȾǦ†‹˜‡”•‹–› ‘ˆ –Š‡ –™‘ ‘—‹–‹‡•ƒ”‡ ‘””‡Žƒ–‡†Ǥ ‘”‡” •–—†‹‡• ‘†— –‡† ‹ †‹ˆˆ‡”‡– Šƒ„‹–ƒ– –›’‡• ƒŽ•‘ •Š‘™‡† ƒ Š‹‰Š †‡’‡†‡ › ‘ˆ –Š‡ Š‹‰Š‡” –”‘’Š‹  Ž‡˜‡Ž ‘ –Š‡ Ž‘™‡” –”‘’Š‹  Ž‡˜‡Ž ȋŽ„”‡ Š– ‡– ƒŽǤ ʹͲͲ͹ǡ „‡Ž‹‰‡–ƒŽǤʹͲͳʹǡ‡‹‡”‡–ƒŽǤʹͲͳͶǡ•‘”‹‘‡–ƒŽǤʹͲͳͷȌǤ ‘•–Ǧ’ƒ”ƒ•‹–‘‹†”‡Žƒ–‹‘•Š‹’•ƒ”‡ǡ ‹ ‰‡‡”ƒŽǡ Ž‡•• ˆŽ‡š‹„Ž‡ ȋ’ƒ”–‡” ˆ‹†‡Ž‹–› ‹• ‘ˆ–‡ ‘„Ž‹‰ƒ–‘”›Ǣ ƒ™‹• ͳͻͻͶȌ –Šƒ ’Žƒ–Ǧ ’‘ŽŽ‹ƒ–‘””‡Žƒ–‹‘•Š‹’•ȋƒ•‡”‡–ƒŽǤͳͻͻ͸ȌǡŽ‡ƒ˜‹‰Ž‹––Ž‡”‘‘–‘‘’’‘”–—‹•–‹ ‹–‡”ƒ –‹‘•Ǥ  ‘”†‹‰Ž›ǡ ‘—” ”‡•—Ž–• ›‹‡Ž† ƒ Š‹‰ŠŽ› •’‡ ‹ƒŽ‹œ‡† ‡–™‘” ‘’ƒ”‡†–‘‘–Š‡”–›’‡•‘ˆ ‡–™‘”••— Šƒ•’‘ŽŽ‹ƒ–‹‘ǡ•‡‡††‹•’‡”•ƒŽ‘”ƒ–Ǧ‡ –ƒ”’Žƒ–‡–™‘”•ȋŽò–Š‰‡‡–ƒŽǤ ʹͲͲ͹ǡ ŠŽ‡—‹‰‡–ƒŽǤʹͲͳʹȌǤ—”˜ƒŽ—‡•ƒ”‡ƒƒŽ‘‰‘—•–‘–Š‘•‡ˆ‘—†ˆ‘”‘”‡•’‡ ‹ƒŽ‹œ‡† •›•–‡•ǡ•— Šƒ•Š‘•–Ǧ’ƒ”ƒ•‹–‘‹†ȋ‘””‹•‡–ƒŽǤʹͲͳͶȌƒ†ƒ–Ǧ›”‡ ‘’Š›–‡ȋŽò–Š‰‡‡–ƒŽǤ ʹͲͲ͹Ȍ‡–™‘”•Ǥ

‡•—Ž–••Š‘™–Šƒ–Š‘•–ƒ†’ƒ”ƒ•‹–‘‹† ‘—‹–‹‡•Šƒ†Š‹‰Š˜ƒŽ—‡•‘ˆEǦ†‹˜‡”•‹–›ǡ‘•–Ž› †—‡–‘•’‡ ‹‡•–—”‘˜‡”™‹–ŠƒŽ‘•–‹””‡Ž‡˜ƒ–•’‡ ‹‡•Ž‘••ǤŠƒ–‹•ǡŽ‘ ƒŽ ‘—‹–‹‡•†‹† ‘–‘Ž›†‹ˆˆ‡”‹•’‡ ‹‡•”‹ Š‡••ǡ„—––‡†‡†–‘Šƒ˜‡•’‡ ‹‡• ‘’‘•‹–‹‘•‹†‹‘•› ”ƒ–‹ –‘ ‡ƒ Š •‹–‡Ǥ ƒŽ›•‹• ‘ˆ EǦ†‹˜‡”•‹–› ‘ˆ Ž‘ ƒŽ ‡–™‘”• ‘ˆ‹”‡† –Š‡•‡ ”‡•—Ž–•Ǥ ƒ‹”™‹•‡

‘’ƒ”‹•‘• „‡–™‡‡ Ž‘ ƒŽ ‡–™‘”• ”‡˜‡ƒŽ‡† Š‹‰Š †‹••‹‹Žƒ”‹–› ‘ˆ ‹–‡”ƒ –‹‘• ȋȾǡ ‡ƒrǣ ͲǤͺ͵άͲǤͳ͸Ȍǡ ƒ† •’‡ ‹‡• –—”‘˜‡” ™ƒ• –Š‡ ƒ‹ ˆƒ –‘” ‘–”‹„—–‹‰ –‘ •— Š

†‹••‹‹Žƒ”‹–› ȋȾǡ ‡ƒrǣ ͲǤ͸ͲrͲǤ͵ʹȌǤ ‹˜‡ –Š‡ Š‹‰Š Ž‡˜‡Ž ‘ˆ •’‡ ‹ƒŽ‹œƒ–‹‘ ‹ ‘—” ‡–™‘”ǡ•’‡ ‹‡•–—”‘˜‡”•Š‘—Ž†„‡‡š’‡ –‡†–‘„‡–Š‡ƒŒ‘”†”‹˜‡”‘ˆ‹–‡”ƒ –‹‘†‹˜‡”•‹–› ȋŽ‡•‡ ‡– ƒŽǤ ʹͲͳͳȌǡ ƒ† –Š‡ ‡ˆˆ‡ – ‘ˆ ”‡™‹”‹‰ ȋ•Šƒ”‡† •’‡ ‹‡• ‹–‡”ƒ –‹‰ †‹ˆˆ‡”‡–Ž›Ȍ

•Š‘—Ž†„‡‡š’‡ –‡†–‘„‡— ŠŽ‘™‡”ȋȾǡ‡ƒrǣͲǤʹ͵rͲǤ͵ʹǡ‹–Š‹••–—†›ȌǤŠ‹•”‡•—Ž– ƒ‰”‡‡•™‹–Š’”‡˜‹‘—••–—†‹‡•ȋ‘˜‘–›ʹͲͲͻǡ‹ƒ‘‘ƒ†—”Ž‡ʹͲͳͶǡ”ÞŒ‡Ž•‰ƒƒ”†‡– ƒŽǤʹͲͳͷǡ„—–•‡‡ƒ”•–‡•‡‡–ƒŽǤʹͲͳͶˆ‘”ƒ•‹‹Žƒ” ‘–”‹„—–‹‘‘ˆ„‘–Š ‘’‘‡–•‘ˆ „‘–Š ‘’‘‡–•‘ˆ‹–‡”ƒ –‹‘†‹˜‡”•‹–›ȌǤŠ‡•‡•–—†‹‡•ǡ‹ Ž—†‹‰‘—”•ǡ ‘˜‡”ƒ˜ƒ”‹‡–›‘ˆ ‹–‡”ƒ –‹‘•›•–‡•ȋ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘”ǡ’Žƒ–Ǧ ƒ–‡”’‹ŽŽƒ”ƒ†Š‘•–Ǧ’ƒ”ƒ•‹–‘‹†Ȍƒ†ƒ”ƒ‰‡‘ˆ ‰‡‘‰”ƒ’Š‹ ƒŽ• ƒŽ‡•ȋˆ”‘ͶͲͲ–‘ͷͲͲȌǤŠ—•ǡ•’‡ ‹‡•–—”‘˜‡”ƒ’’‡ƒ”•–‘„‡–Š‡ƒ‹ †”‹˜‡”‘ˆ‡–™‘”EǦ†‹˜‡”•‹–›ƒ ”‘••„‹‘Ž‘‰‹ ƒŽ•›•–‡•ƒ†•’ƒ–‹ƒŽ• ƒŽ‡•‘ˆ‘„•‡”˜ƒ–‹‘Ǥ

 90 Šƒ’–‡”͵

–‡”ƒ –‹‘ •‹‹Žƒ”‹–› „‡–™‡‡ Ž‘ ƒŽ ‡–™‘”• ƒ† –Š‡‹” ‘—–‡”’ƒ”–•‹–Š‡”‡‰‹‘ƒŽ

‡–™‘”‹•Š‹‰Šǡƒ•”‡ˆŽ‡ –‡†„›–Š‡Ž‘™Eǯ ȋ‡ƒrα ͲǤʹάͲǤͳȌǤ  ‘–Š‡” ™‘”†•ǡ ‹–‡”ƒ –‹‘•‹Ž‘ ƒŽ‡–™‘”•ƒ”‡‡•–‡†‹–Š‡”‡‰‹‘ƒŽ‡–™‘”Ǥ –‡”ƒ –‹‘‡•–‡†‡••‹ ‘—”•›•–‡”‡•—Ž–•ˆ”‘ƒ ‘„‹ƒ–‹‘‘ˆƒȌ•’‡ ‹‡•‹–‡”ƒ –‹‰•‹‹Žƒ”Ž›ƒ ”‘••’Ž‘–•ǡƒ†„Ȍ Žƒ”‰‡ †‹ˆˆ‡”‡ ‡• „‡–™‡‡ ’Ž‘–• ‹ •’‡ ‹‡• ‘’‘•‹–‹‘Ǥ ‡ ƒ—•‡ Ž‘ ƒŽ ‘—‹–‹‡• ƒ”‡ Š‹‰ŠŽ›‹†‹‘•› ”ƒ–‹ ȋ͸͵ǤͻΨ‘ˆ‹–‡”ƒ –‹‘•ƒ”‡‘Ž›ˆ‘—†‹‘‡’Ž‘–Ȍǡ–Š‡ ‘–”‹„—–‹‘‘ˆ ‡ƒ Š’Ž‘––‘–Š‡”‡‰‹‘ƒŽ™‡„‹•Š‹‰ŠǤŠ‹•‹–‡”’”‡–ƒ–‹‘†‹ˆˆ‡”•ˆ”‘–Š‡‡š’Žƒƒ–‹‘‰‹˜‡ „› ‘‹•‘– ƒ† ‘ŽŽƒ„‘”ƒ–‘”• ‹ –Š‡‹” ƒƒŽ›•‹• ‘ˆ ƒ ƒƒŽǦ‡ –‘’ƒ”ƒ•‹–‡ ”‡‰‹‘ƒŽ ‡–™‘”

ȋ‘‹•‘– ‡– ƒŽǤ ʹͲͳʹȌǤ Š‡•‡ ƒ—–Š‘”• ‘ Ž—†‡ –Šƒ– Š‹‰Š ˜ƒŽ—‡• ‘ˆ Eǯ ‹†‹ ƒ–‡ –Šƒ– ƒ› ‹–‡”ƒ –‹‘•‘Ž›‘ —”‹ƒˆ‡™Ž‘ ƒŽ™‡„•ǡ’‘‹–‹‰–Šƒ–‹–‡”ƒ –‹‘•ƒ”‡Š‹‰ŠŽ›†‡–‡”‹‡† „›Ž‘ ƒŽ ‘†‹–‹‘•Ǥ ‘™‡˜‡”ǡ‹ˆ–Š‡”‡‹•ƒŽ‘™’”‘’‘”–‹‘‘ˆ•Šƒ”‡†•’‡ ‹‡•„‡–™‡‡Ž‘ ƒŽ

™‡„•ǡƒ†•Šƒ”‡†•’‡ ‹‡•‹–‡”ƒ –•‹‹Žƒ”Ž›ǡEǯ˜ƒŽ—‡•™‹ŽŽ„‡Ž‘™ƒ†‹–™‹ŽŽ•–‹ŽŽ„‡–”—‡ –Šƒ––Š‡”‡ƒ”‡ƒŽ‘–‘ˆ‹–‡”ƒ –‹‘•‘Ž›‘ —””‹‰‹ƒˆ‡™Ž‘ ƒŽ™‡„•Ǥ –Š‹• ƒ•‡ǡ™Š‹ Š‹• ‘—” ƒ•‡ǡ ‹–‡”ƒ –‹‘• ƒ”‡ •‹–‡Ǧ‹†‹‘•› ”ƒ–‹  ‘– „‡ ƒ—•‡ ‹–‡”ƒ –‹‘ ‘’‘•‹–‹‘ per se‹• †‡–‡”‹‡† „› Ž‘ ƒŽ ‘†‹–‹‘•ǡ „—– „‡ ƒ—•‡ •’‡ ‹‡• ‘’‘•‹–‹‘ ‹• †‡–‡”‹‡† „› Ž‘ ƒŽ ‘†‹–‹‘•Ǥ— Š”‡•—Ž–•ƒ‰”‡‡™‹–Š’”‡˜‹‘—•™‘”‹–Š‡•ƒ‡ƒ”‡ƒȋ‘”±Ǧ‘‰—‡”ƒ‡–ƒŽǤ ʹͲͳͶȌǡ™Š‹ Šˆ‘—†–Š‡‘˜‡”ƒŽŽ„‡‡ ‘—‹–›–‘„‡•–”‘‰Ž› ‘†‹–‹‘‡†„›Ž‘ ƒŽ‡ˆˆ‡ –•Ǥ —””‡•—Ž–•ƒ”‡ƒŽ•‘•—’’‘”–‡†„›ƒ•–—†›–Šƒ–ˆ‘—†Ž‘ ƒŽˆƒ –‘”• –‘ „‡ — Š ‘”‡ ‹’‘”–ƒ––ŠƒŽƒ†• ƒ’‡ˆƒ –‘”•ƒ•†‡–‡”‹ƒ–•‘ˆ ‘—‹–›ƒ†‡–™‘”•–”— –—”‡ƒ† ‘’‘•‹–‹‘‹ ƒ˜‹–›Ǧ‡•–‹‰„‡‡•ƒ†™ƒ•’•ȋ•‘”‹‘‡–ƒŽǤʹͲͳͷȌǤ

—””‡•—Ž–•ƒ††‹–‹‘ƒŽŽ›•Š‘™–Šƒ–EǦ†‹˜‡”•‹–›‘ˆ’ƒ”ƒ•‹–‹œ‡†Š‘•–•‹•ƒ„‡––‡”’”‡†‹ –‘”‘ˆEǦ †‹˜‡”•‹–›‘ˆ‹–‡”ƒ –‹‘•–ŠƒEǦ†‹˜‡”•‹–›‘ˆ’ƒ”ƒ•‹–‘‹†•Ǥ‡ƒ˜ƒŽ—‡‘ˆ’ƒ‹”™‹•‡ ‘—‹–›

EǦ†‹˜‡”•‹–›‹•Š‹‰Š‡”ˆ‘”’ƒ”ƒ•‹–‹œ‡†Š‘•–•ȋE ǣ‡ƒrαͲǤ͸ͺrͲǤʹͳȌ–Šƒˆ‘”’ƒ”ƒ•‹–‘‹†•

ȋEǣ‡ƒrαͲǤͷ͹rͲǤͳͺȌǤ ‘–Š‡”™‘”†•ǡ‘—”Š‘•– ‘—‹–›–—”‘˜‡”‹•Š‹‰Š‡”ƒ†–Š—• ‘–”‹„—–‡• ‘”‡ –Šƒ ’ƒ”ƒ•‹–‘‹† ‘—‹–› –—”‘˜‡” –‘ ‹–‡”ƒ –‹‘ –—”‘˜‡”Ǥ ”‡ƒ–‡” –—”‘˜‡”‘ˆ–Š‡Ž‘™‡”–”‘’Š‹ Ž‡˜‡Ž•‡‡•–‘„‡ƒˆ‡ƒ–—”‡ ‘‘–‘‘–Š‡”‹†•‘ˆ‡–™‘”• ȋ‘˜‘–›ʹͲͲͻǡƒ”•–‡•‡‡–ƒŽǤʹͲͳͶǡ‹ƒ‘‘ƒ†—”Ž‡ʹͲͳͶ„—–•‡‡”ÞŒ‡Ž•‰ƒƒ”†‡– ƒŽǤʹͲͳͷȌǤ

˜‡”ƒŽŽǡ –Š‹• •–—†› †‡‘•–”ƒ–‡• –Šƒ– ‘—‹–‹‡• ‘ˆ ‘„‹Ž‡ ‘”‰ƒ‹•• ƒ† –Š‡‹” ‹–‡”ƒ –‹‘•˜ƒ”›ƒ–ƒŽ‘ ƒŽ• ƒŽ‡ȋ̱ͷͲͲȌ‡˜‡‹–Š‡ƒ„•‡ ‡ ‘ˆ ‡ ‘Ž‘‰‹ ƒŽ „ƒ””‹‡”• ‘” ‡˜‹”‘‡–ƒŽ‰”ƒ†‹‡–•Ǥ ‘•–•’ƒ–‹ƒŽ–—”‘˜‡”‹•–Š‡ƒŒ‘”†”‹˜‡”‘ˆ–Š‡‘„•‡”˜‡†•’ƒ–‹ƒŽ Š‡–‡”‘‰‡‡‹–›ǡ•‹ ‡–Š‡’ƒ”ƒ•‹–‘‹† ‘—‹–›‹•Š‹‰ŠŽ›†‡’‡†‡–‘–Š‡Š‘•– ‘—‹–›ǡ  91 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

ƒ†‹–‡”ƒ –‹‘•†‡’‡†‘„‘–Š ‘—‹–‹‡•Ǥ‘‡–Š‡Ž‡••ǡ’ƒ”ƒ•‹–‘‹†•ƒ†‹–‡”ƒ –‹‘•ƒ”‡ ƒŽ•‘ •—„Œ‡ –‡† –‘ –Š‡‹” ‘™ ‹–”‹•‹  ˜ƒ”‹ƒ„‹Ž‹–›ǡ ”‡•—Ž–‹‰ ‹†‹ˆˆ‡”‡–”‡•’‘•‡•ƒ ”‘•• •’ƒ ‡ǤŠ—•ǡ‹–‹•‹’‘”–ƒ––‘–ƒ‡‹–‘ƒ ‘—–„‘–ŠŽ‡˜‡Ž•‘ˆ˜ƒ”‹ƒ„‹Ž‹–›ȋ ‘—‹–‹‡•ƒ† ‹–‡”ƒ –‹‘•Ȍ–‘ƒ†‡“—ƒ–‡Ž› Šƒ”ƒ –‡”‹œ‡‡ ‘Ž‘‰‹ ƒŽˆ— –‹‘Ǥ—””‡•—Ž–•ƒŽ•‘•Š‘™–Š‡‡‡† –‘ •ƒ’Ž‡ ƒ Žƒ”‰‡ —„‡” ‘ˆ ’Ž‘–• –‘ ƒ†‡“—ƒ–‡Ž› Šƒ”ƒ –‡”‹œ‡ ƒ ”‡‰‹‘ƒŽ ‡–™‘”ǡ •‹ ‡ ‡–™‘”•ƒ›˜ƒ”›•’ƒ–‹ƒŽŽ›ƒ–ƒ˜‡”›Ž‘ ƒŽ• ƒŽ‡‡˜‡‹ƒ ‘–‹—‘—•Šƒ„‹–ƒ–Ǥ‹•–‹‰—‹•Š‹‰ „‡–™‡‡ –Š‡ –™‘ ‘’‘‡–• ‘ˆ ‹–‡”ƒ –‹‘ EǦ†‹˜‡”•‹–› ‹• ‡••‡–‹ƒŽ –‘ —†‡”•–ƒ† –Š‡ †”‹˜‡”•‘ˆ•— Š†‹˜‡”•‹–›ǤŠ‡‡–™‘”EǦ†‹˜‡”•‹–›‹•‘•–Ž›†—‡–‘‹–‡”ƒ –‹‘”‡™‹”‹‰ǡ –Š‡ ”‡•—Ž–‹‰ ‡–™‘” ™‹ŽŽ ”‡ˆŽ‡ – –Š‡ ”‡‰‹‘ƒŽǡ „—– ‘– –Š‡ Ž‘ ƒŽ „‡Šƒ˜‹‘” ‘ˆ ‹–‡”ƒ –‹‰ •’‡ ‹‡•Ǥ  ‘–”ƒ•–ǡ ™Š‡ ‡–™‘” EǦ†‹˜‡”•‹–› ‹• ‘•–Ž› †—‡ –‘ •’‡ ‹‡• –—”‘˜‡”ǡ –Š‡ ”‡‰‹‘ƒŽ ‡–™‘” ™‹ŽŽ ”‡ˆŽ‡ – –Š‡ Ž‘ ƒŽ „‡Šƒ˜‹‘” ‘ˆ ‹–‡”ƒ –‹‰ •’‡ ‹‡•Ǥ ‹–Š Š‹‰Š Ž‡˜‡Ž• ‘ˆ •’‡ ‹ƒŽ‹œƒ–‹‘ƒ†Ž‘™Ž‡˜‡Ž•‘””‡™‹”‹‰ǡ‘—”•›•–‡ˆƒŽŽ•™‹–Š‹ –Š‡ Žƒ––‡” • ‡ƒ”‹‘Ǥ ‘™‡˜‡”ǡ‡˜‡–Š‡‹–‹•‡ ‡••ƒ”›–‘•ƒ’Ž‡ƒ•—ˆˆ‹ ‹‡–Ž›Žƒ”‰‡—„‡”‘ˆ’Ž‘–•„‡ ƒ—•‡ •’‡ ‹‡• ‘’‘•‹–‹‘‹•Š‹‰ŠŽ›†‡’‡†‡–‘Ž‘ ƒŽˆƒ –‘”•ƒ†•—„Œ‡ –‡†–‘˜ƒ”‹ƒ–‹‘ƒ–˜‡”› •ƒŽŽ• ƒŽ‡•Ǥ



Aknowledgements

‡ƒ”‡˜‡”›‰”ƒ–‡ˆ—Ž–‘ Ǥƒ””‹ŽǦ ”ƒ‡ŽŽ•ǡǤƒŽƒƒ”ƒǡǤ‘˜‡ŽŽƒǡ Ǥ‘„ƒ–‘ǡ Ǥƒ‘‡†ƒǡǤ • ‘ŽǡǤ‡˜‡”–±ǡǤג‡œƒ†Ǥ—ƒ†”ƒˆ‘”–Š‡‹”Š‡Ž’‹–Š‡ˆ‹‡Ž†ƒ†‹–Š‡Žƒ„‘”ƒ–‘”›Ǥ‡ ƒŽ•‘–ŠƒǤŽò–Š‰‡ȋ‡ Š‹ ƒŽ‹˜‡”•‹–›‘ˆƒ”•–ƒ†–Ȍˆ‘”Š‹•ƒƒŽ›–‹ ƒŽƒ†˜‹ ‡ƒ† Ǥ ‡†‡”‘•ȋ—•‡—‘ˆƒ–—”ƒŽ ‹‡ ‡•ǡƒ” ‡Ž‘ƒȌˆ‘”Š‹•Š‡Ž’™‹–Š‹’–‡”ƒ‹†‡–‹ˆ‹ ƒ–‹‘Ǥ ‡ƒ”‡ƒŽ•‘–Šƒˆ—Ž–‘‹’—–ƒ ‹×†‡ƒ” ‡Ž‘ƒˆ‘”’‡”‹••‹‘–‘™‘”‹–Š‡ƒ–—”ƒŽƒ”‘ˆ ‡Ž ƒ””ƒˆǤŠ‹••–—†›™ƒ•ˆ—†‡†„›–Š‡’ƒ‹•Š‹‹•–”›‘ˆ ‹‡ ‡ ƒ† ‘˜ƒ–‹‘ ȋ  Ȍǡ’”‘Œ‡ –• ʹͲͲͻǦͳʹ͸Ͷ͸ƒ†ʹͲͲͺǦͲͲͲͶͲǡƒ†ƒ  ˆ‡ŽŽ‘™•Š‹’ȋǦʹͲͳͲǦ ͲͶʹͷʹͲȌ–‘ǤǦǤ



References

Ž„”‡ Š–ǡ Ǥ ‡– ƒŽǤ ʹͲͲ͹Ǥ –‡”ƒ –‹‘ †‹˜‡”•‹–› ™‹–Š‹ “—ƒ–‹ˆ‹‡† ‹•‡ – ˆ‘‘† ™‡„• ‹ ”‡•–‘”‡† ƒ† ƒ†Œƒ ‡–‹–‡•‹˜‡Ž›ƒƒ‰‡†‡ƒ†‘™•ǤǦ Ǥ‹Ǥ ‘ŽǤ͹͸ǣͳͲͳͷȂͳͲʹͷǤ

 92 Šƒ’–‡”͵

ƒ•‡Ž‰ƒǡǤʹͲͳͲǤƒ”–‹–‹‘‹‰–Š‡–—”‘˜‡”ƒ†‡•–‡†‡•• ‘’‘‡–•‘ˆ„‡–ƒ†‹˜‡”•‹–›ǤǦ Ž‘„Ǥ ‘ŽǤ ‹‘‰‡‘‰”Ǥͳͻǣͳ͵ͶȂͳͶ͵Ǥ

ƒ•‡Ž‰ƒǡǤƒ†”‡ǡǤǤǤʹͲͳʹǤ„‡–ƒ’ƒ”–᩿ǣƒ’ƒ ƒ‰‡ˆ‘”–Š‡•–—†›‘ˆ„‡–ƒ†‹˜‡”•‹–›ǤǦ‡–Š‘†•  ‘ŽǤ˜‘ŽǤ͵ǣͺͲͺȂͺͳʹǤ

Žò–Š‰‡ǡǤ‡–ƒŽǤʹͲͲ͸Ǥ‡ƒ•—”‹‰•’‡ ‹ƒŽ‹œƒ–‹‘‹•’‡ ‹‡•‹–‡”ƒ –‹‘‡–™‘”•ǤǦ ‘ŽǤ͸ǣͻǤ

Žò–Š‰‡ǡǤ‡–ƒŽǤʹͲͲ͹Ǥ’‡ ‹ƒŽ‹œƒ–‹‘ǡ ‘•–”ƒ‹–•ǡƒ† ‘ˆŽ‹ –‹‰‹–‡”‡•–•‹—–—ƒŽ‹•–‹ ‡–™‘”•ǤǦ —””Ǥ‹‘ŽǤͳ͹ǣ͵ͶͳȂ͸Ǥ

—”Ž‡ǡǤǤƒ†Žƒ” אǡǤʹͲͳͳǤŠ‡ˆ—–—”‡‘ˆ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘”†‹˜‡”•‹–›ǣ—†‡”•–ƒ†‹‰‹–‡”ƒ –‹‘ ‡–™‘”•ƒ ”‘••–‹‡ǡ•’ƒ ‡ǡƒ†‰Ž‘„ƒŽ Šƒ‰‡ǤǦǤ Ǥ‘–ǤͻͺǣͷʹͺȂ͵ͺǤ

ƒ”•–‡•‡ǡǤǤ‡–ƒŽǤʹͲͳͶǤ‡–ƒ†‹˜‡”•‹–›‘ˆ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘”‡–™‘”•ƒ†–Š‡•’ƒ–‹ƒŽ–—”‘˜‡”‘ˆ ’ƒ‹”™‹•‡‹–‡”ƒ –‹‘•ǤǦ‘‡ͻǣ‡ͳͳʹͻͲ͵Ǥ

‘†‹–ǡǤ‡–ƒŽǤʹͲͲʹǤ‡–ƒǦ†‹˜‡”•‹–›‹–”‘’‹ ƒŽˆ‘”‡•––”‡‡•ǤǦ ‹‡ ‡ʹͻͷǣ͸͸͸Ȃ͸͸ͻǤ

ž––‹Ž‘ǡǤ‡–ƒŽǤʹͲͳ͵Ǥ’ƒ–‹ƒŽ•–”— –—”‡‘ˆƒ–Ǧ’Žƒ–—–—ƒŽ‹•–‹ ‡–™‘”•ǤǦ‹‘•ͳʹʹǣͳ͸Ͷ͵Ȃͳ͸ͶͺǤ

‘”ƒǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͺǤ –”‘†— ‹‰ –Š‡ „‹’ƒ”–‹–‡ ƒ ƒ‰‡᩿ǣ ƒŽ›•‹‰  ‘Ž‘‰‹ ƒŽ ‡–™‘”•Ǥ Ǧ  ‡™•ͺǣͺȂͳͳǤ

—’‘–ǡ Ǥ Ǥ‡– ƒŽǤ ʹͲͳͶǤ ’ƒ–‹ƒŽ •–”— –—”‡ ‘ˆ ƒ ‹†‹˜‹†—ƒŽǦ„ƒ•‡†’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘” ‡–™‘”Ǥ Ǧ ‹‘• ͳʹ͵ǣͳ͵ͲͳȂͳ͵ͳͲǤ

„‡Ž‹‰ǡ Ǥ ‡– ƒŽǤ ʹͲͳʹǤ —Ž–‹–”‘’Š‹  ‡ˆˆ‡ –• ‘ˆ ‡š’‡”‹‡–ƒŽ Šƒ‰‡• ‹ ’Žƒ– †‹˜‡”•‹–› ‘ ƒ˜‹–›Ǧ ‡•–‹‰„‡‡•ǡ™ƒ•’•ǡƒ†–Š‡‹”’ƒ”ƒ•‹–‘‹†•ǤǦ‡ ‘Ž‘‰‹ƒͳ͸ͻǣͶͷ͵ȂͶ͸ͷǤ

‹•Š‡”ǡǤǤ‡–ƒŽǤͳͻͶ͵ǤŠ‡”‡Žƒ–‹‘„‡–™‡‡–Š‡—„‡”‘ˆ•’‡ ‹‡•ƒ†–Š‡—„‡”‘ˆ‹†‹˜‹†—ƒŽ•‹ ƒ”ƒ†‘•ƒ’Ž‡‘ˆƒƒ‹ƒŽ’‘’—Žƒ–‹‘ǤǦ Ǥ‹Ǥ ‘ŽǤͳʹǣͶʹȂͷͺǤ

”ڏ’‹‰ǡ Ǥ ʹͲͲ͸Ǥ ‡Žƒ–‹˜‡ ‹’‘”–ƒ ‡ ˆ‘” Ž‹‡ƒ” ”‡‰”‡••‹‘ ‹ ǣ –Š‡ ’ƒ ƒ‰‡ ”‡Žƒ‹’‘Ǥ Ǧ Ǥ –ƒ–Ǥ ‘ˆ–™Ǥͳ͹ǣͳ͵ͻȂͳͶ͹Ǥ

ƒ™‹•ǡ Ǥ Ǥ ͳͻͻͶǤ ƒ––‡” ƒ† ’”‘ ‡•• ‹ Š‘•–Ǧ’ƒ”ƒ•‹–‘‹† ‹–‡”ƒ –‹‘•ǤǦƒ„”‹†‰‡‹˜‡”•‹–› ”‡••Ǥ

 93 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

•Š‹‹ǡ Ǥ ƒ† Š‹ƒ†ƒǡ Ǥ ʹͲͳʹǤ ‡ƒ”‹‰ ’”‡†ƒ–‘” ’”‘‘–‡• ‘‡š‹•–‡ ‡ ‘ˆ ’”‡› •’‡ ‹‡• ‹ Š‘•– Ȃ ’ƒ”ƒ•‹–‘‹†•›•–‡•ǤǦ”‘ Ǥƒ–ŽǤ ƒ†Ǥ ‹ǤǤǤǤͳͲͻǣͷͳͳͻǤ

‘”†ƒ‘ǡǤ‡–ƒŽǤʹͲͲ͵Ǥ ˜ƒ”‹ƒ–’”‘’‡”–‹‡•‹ ‘‡˜‘Ž—–‹‘ƒ”›‡–™‘”•‘ˆ’Žƒ–Ǧƒ‹ƒŽ‹–‡”ƒ –‹‘•ǤǦ  ‘ŽǤ‡––Ǥ͸ǣ͸ͻȂͺͳǤ

‘”ƒŽ‡•ǡ Ǥ Ǥ ƒ† žœ“—‡œǡ Ǥ Ǥ ʹͲͲͺǤ Š‡ ‡ˆˆ‡ – ‘ˆ •’ƒ ‡ ‹ ’Žƒ–Ǧƒ‹ƒŽ —–—ƒŽ‹•–‹  ‡–™‘”•ǣ ‹•‹‰Š–•ˆ”‘ƒ•‹—Žƒ–‹‘•–—†›ǤǦ‹‘•ͳͳ͹ǣͳ͵͸ʹȂͳ͵͹ͲǤ

‘””‹•ǡǤ Ǥ‡–ƒŽǤʹͲͳͶǤ–ƒ‰‘‹•–‹ ‹–‡”ƒ –‹‘‡–™‘”•ƒ”‡•–”— –—”‡†‹†‡’‡†‡–Ž›‘ˆŽƒ–‹–—†‡ ƒ†Š‘•–‰—‹Ž†ǤǦ ‘ŽǤ‡––Ǥͳ͹ǣ͵ͶͲȂͻǤ

‡‘Žƒǡ ǤǤƒ†Š‹–‡ǡǤǤͳͻͻͻǤŠ‡†‹•–ƒ ‡†‡ ƒ›‘ˆ•‹‹Žƒ”‹–›‹„‹‘‰‡‘‰”ƒ’Š›ƒ†‡ ‘Ž‘‰›ǤǦ Ǥ ‹‘‰‡‘‰”Ǥʹ͸ǣͺ͸͹Ȃͺ͹ͺǤ

‘”ˆ‘Žǡ Ǥ ‡– ƒŽǤ ʹͲͳͶǤ ‘–”ƒ•–‹‰ ’ƒ––‡”• ‘ˆ –—”‘˜‡” „‡–™‡‡’Žƒ–•ǡ’‘ŽŽ‹ƒ–‘”•ƒ†–Š‡‹” ‹–‡”ƒ –‹‘•ǤǦ‹˜‡”•Ǥ‹•–”‹„ǤǣͳȂͳͳǤ

‘˜‘–›ǡ Ǥ ʹͲͲͻǤ ‡–ƒ †‹˜‡”•‹–› ‘ˆ ’Žƒ–Ǧ‹•‡ – ˆ‘‘† ™‡„• ‹ –”‘’‹ ƒŽ ˆ‘”‡•–•ǣ  ‘ ‡’–—ƒŽ ˆ”ƒ‡™‘”ǤǦ •‡ –‘•‡”˜Ǥ‹˜‡”•ǤʹǣͷȂͻǤ

•ƒ‡ǡ Ǥ‡–ƒŽǤʹͲͳͷǤ‡‰ƒǣ‘—‹–› ‘Ž‘‰›ƒ ƒ‰‡Ǥ

Ž‡•‡ǡ Ǥ Ǥ ƒ† ‘”†ƒ‘ǡ Ǥ ʹͲͲʹǤ ‡‘‰”ƒ’Š‹  ’ƒ––‡”• ‹ ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘” —–—ƒŽ‹•–‹  ‡–™‘”•Ǥ Ǧ  ‘Ž‘‰›ͺ͵ǣʹͶͳ͸ȂʹͶʹͶǤ

Ž‡•‡ǡ ǤǤ‡–ƒŽǤʹͲͳͳǤ–”‘‰ǡŽ‘‰Ǧ–‡”–‡’‘”ƒŽ†›ƒ‹ •‘ˆƒ‡ ‘Ž‘‰‹ ƒŽ‡–™‘”ǤǦ‘‡͸ǣ ʹȂ͸Ǥ

•‘”‹‘ǡ Ǥ ‡– ƒŽǤ ʹͲͳͷǤ ‘ ƒŽ ƒ† Žƒ†• ƒ’‡ ‡ˆˆ‡ –• ‹ ƒ Š‘•–Ǧ’ƒ”ƒ•‹–‘‹† ‹–‡”ƒ –‹‘ ‡–™‘” ƒŽ‘‰ ƒ ˆ‘”‡•–Ǧ ”‘’Žƒ†‰”ƒ†‹‡–ǤǦ ‘ŽǤ’’ŽǤ‹’”‡••Ǥ

‘‹•‘–ǡǤ‡–ƒŽǤʹͲͳʹǤŠ‡†‹••‹‹Žƒ”‹–›‘ˆ•’‡ ‹‡•‹–‡”ƒ –‹‘‡–™‘”•ǤǦ ‘ŽǤ‡––Ǥͳͷǣͳ͵ͷ͵Ȃ͸ͳǤ

‘”‡‡ƒʹͲͳͶǤǣŽƒ‰—ƒ‰‡ƒ†‡˜‹”‘‡–ˆ‘”•–ƒ–‹•–‹ ƒŽ ‘’—–‹‰Ǥ

 ŠŽ‡—‹‰ǡ Ǥ ‡– ƒŽǤ ʹͲͳʹǤ ’‡ ‹ƒŽ‹œƒ–‹‘ ‘ˆ —–—ƒŽ‹•–‹  ‹–‡”ƒ –‹‘ ‡–™‘”• †‡ ”‡ƒ•‡• –‘™ƒ”† –”‘’‹ ƒŽŽƒ–‹–—†‡•ǤǦ—””Ǥ‹‘ŽǤʹʹǣͳͻʹͷȂ͵ͳǤ

 94 Šƒ’–‡”͵

‹ƒ‘‘ǡǤǤƒ†—”Ž‡ǡǤǤʹͲͳͶǤƒ”–‹–‹‘‹‰‹–‡”ƒ –‹‘–—”‘˜‡”ƒ‘‰ƒŽ’‹‡’‘ŽŽ‹ƒ–‹‘ ‡–™‘”•᩿ǣ•’ƒ–‹ƒŽǡ–‡’‘”ƒŽǡƒ†‡˜‹”‘‡–ƒŽ’ƒ––‡”•ǤǦ ‘•’Š‡”‡ͷǣͳȂͳ͹Ǥ

‘‹‹‡ǡ Ǥ‡–ƒŽǤʹͲͲ͹ǤŠ‡†‹•–ƒ ‡†‡ ƒ›‘ˆ•‹‹Žƒ”‹–›‹‡ ‘Ž‘‰‹ ƒŽ ‘—‹–‹‡•ǤǦ ‘‰”ƒ’Š›͵Ͳǣ͵Ȃ ͳʹǤ

–‡ˆˆƒǦ‡™‡–‡”ǡ ǤʹͲͲ͵Ǥ ’‘”–ƒ ‡‘ˆŠƒ„‹–ƒ–ƒ”‡ƒƒ†Žƒ†• ƒ’‡ ‘–‡š–ˆ‘”•’‡ ‹‡•”‹ Š‡••‘ˆ „‡‡•ƒ†™ƒ•’•‹ˆ”ƒ‰‡–‡†‘” Šƒ”†‡ƒ†‘™•ǤǦ‘•‡”˜Ǥ‹‘ŽǤͳ͹ǣͳͲ͵͸ȂͳͲͶͶǤ

‘”±Ǧ‘‰—‡”ƒǡ Ǥ ‡– ƒŽǤ ʹͲͳͶǤ ‡–‡”‹ƒ–• ‘ˆ •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‹ ƒ „‡‡ ‘—‹–›ǣ ‡•–‹‰ ”‡•‘—” ‡•ǡˆŽ‘™‡””‡•‘—” ‡•ǡƒ†„‘†›•‹œ‡ǤǦ‘‡ͻǣ‡ͻ͹ʹͷͷǤ

”ÞŒ‡Ž•‰ƒƒ”†ǡǤ‡–ƒŽǤʹͲͳͷǤ ‡‘‰”ƒ’Š‹ ƒŽ˜ƒ”‹ƒ–‹‘‹—–—ƒŽ‹•–‹ ‡–™‘”•ǣ•‹‹Žƒ”‹–›ǡ–—”‘˜‡”ƒ† ’ƒ”–‡”ˆ‹†‡Ž‹–›ǤǦ”‘ ǤǤ‘ Ǥ‹‘ŽǤ ‹ǤʹͺʹǣʹͲͳͶʹͻʹͷǤ

›Ž‹ƒƒ‹•ǡ ǤǤ‡–ƒŽǤʹͲͲ͸Ǥ‹˜‡”•‹–›ǡ‡ ‘•›•–‡ˆ— –‹‘ǡƒ†•–ƒ„‹Ž‹–›‘ˆ’ƒ”ƒ•‹–‘‹†ǦŠ‘•–‹–‡”ƒ –‹‘• ƒ ”‘••ƒ–”‘’‹ ƒŽŠƒ„‹–ƒ–‰”ƒ†‹‡–ǤǦ ‘Ž‘‰›ͺ͹ǣ͵ͲͶ͹Ȃ͵Ͳͷ͹Ǥ

›Ž‹ƒƒ‹•ǡ ǤǤ‡–ƒŽǤʹͲͲ͹Ǥ ƒ„‹–ƒ–‘†‹ˆ‹ ƒ–‹‘ƒŽ–‡”•–Š‡•–”— –—”‡‘ˆ–”‘’‹ ƒŽŠ‘•–Ǧ’ƒ”ƒ•‹–‘‹†ˆ‘‘† ™‡„•ǤǦƒ–—”‡ͶͶͷǣʹͲʹȂͷǤ

žœ“—‡œǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲ͹Ǥ ’‡ ‹‡• ƒ„—†ƒ ‡ ƒ† ƒ•›‡–”‹  ‹–‡”ƒ –‹‘ •–”‡‰–Š ‹ ‡ ‘Ž‘‰‹ ƒŽ ‡–™‘”•ǤǦ‹‘•ͳͳ͸ǣͳͳʹͲȂͳͳʹ͹Ǥ

žœ“—‡œǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͻƒǤ ˜ƒŽ—ƒ–‹‰ —Ž–‹’Ž‡ †‡–‡”‹ƒ–• ‘ˆ –Š‡ •–”— –—”‡ ‘ˆ ’Žƒ–Ǧƒ‹ƒŽ —–—ƒŽ‹•–‹ ‡–™‘”•ǤǦ ‘Ž‘‰›ͻͲǣʹͲ͵ͻȂʹͲͶ͸Ǥ

žœ“—‡œǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͲͻ„Ǥ ‹–‹‰ ’ƒ––‡” ƒ† ’”‘ ‡•• ‹ ’Žƒ–Ǧƒ‹ƒŽ —–—ƒŽ‹•–‹  ‡–™‘”•ǣ ƒ ”‡˜‹‡™ǤǦǤ‘–ǤͳͲ͵ǣͳͶͶͷȂͷ͹Ǥ

ƒ•‡”ǡǤǤ‡–ƒŽǤͳͻͻ͸Ǥ ‡‡”ƒŽ‹œƒ–‹‘‹’‘ŽŽ‹ƒ–‹‘•›•–‡•ǡƒ†™Š›‹–ƒ––‡”•ǤǦ ‘Ž‘‰›͹͹ǣͳͲͶ͵Ǥ

‡‹‡”ǡ Ǥ Ǥ ‡– ƒŽǤ ʹͲͳͶǤ ƒ†Ǧ—•‡ ‹’ƒ –• ‘ ’Žƒ–Ǧ’‘ŽŽ‹ƒ–‘”‡–™‘”•ǣ‹–‡”ƒ –‹‘•–”‡‰–Šƒ† •’‡ ‹ƒŽ‹œƒ–‹‘’”‡†‹ –’‘ŽŽ‹ƒ–‘”†‡ Ž‹‡•ǤǦ ‘Ž‘‰›ͻͷǣͶ͸͸Ȃ͹ͶǤ

Š‹––ƒ‡”ǡǤ Ǥͳͻ͸ͲǤ‡‰‡–ƒ–‹‘‘ˆ–Š‡‹•‹›‘—‘—–ƒ‹•ǡ”‡‰‘ƒ†ƒŽ‹ˆ‘”‹ƒǤǦ ‘ŽǤ‘‘‰”Ǥ ͵Ͳǣʹ͹ͻȂ͵͵ͺǤ



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Generalconclusions















  ‡‡”ƒŽ ‘ Ž—•‹‘•

Generalconclusions

ƒ Ž‘™‡”• ƒ† ‡•–‹‰ •—„•–”ƒ–‡• ƒ”‡ —‡˜‡Ž› †‹•–”‹„—–‡† ƒ ”‘•• ƒ ‘–‹—‘—• Šƒ„‹–ƒ–ǡ „—– ‘Ž›ˆŽ‘™‡”•ˆ‘ŽŽ‘™ƒ•’ƒ–‹ƒŽ’ƒ––‡”ƒ––Š‡• ƒŽ‡ ‘•‹†‡”‡†Ǥ‡•’‹–‡„‡‡•„‡‹‰Š‹‰ŠŽ›‘„‹Ž‡ ‘”‰ƒ‹••ǡ„‡‡•’‡ ‹‡• ‘’‘•‹–‹‘ƒŽ•‘ˆ‘ŽŽ‘™•ƒ•’ƒ–‹ƒŽ’ƒ––‡”ǡ †‡‘•–”ƒ–‹‰ –Šƒ– „‡‡ ‘—‹–‹‡•ƒ›†‹•’Žƒ›•–”‘‰Š‡–‡”‘‰‡‡‹–›ƒ–ƒŽ‘ ƒŽ• ƒŽ‡ȋͷͲͲǦͳͲͲͲȌ‹ƒ ‘–‹—‘—• Šƒ„‹–ƒ– ȋŠƒ’–‡” ͳȌǤ ƒ˜‹–›Ǧ‡•–‹‰ „‡‡ ƒ† ™ƒ•’ ‘—‹–› ȋŠ‘•–•Ȍ ƒ† –Š‡‹” ƒ••‘ ‹ƒ–‡† ˆƒ—ƒȋ’ƒ”ƒ•‹–‘‹†•Ȍǡƒ•™‡ŽŽƒ•–Š‡‹”‹–‡”ƒ –‹‘•ǡƒŽ•‘†‹•’Žƒ›Š‹‰Š•’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›ƒ–ƒŽ‘ ƒŽ • ƒŽ‡Ǥ ‘™‡˜‡”ǡ‘Ž›–Š‡Š‘•– ‘—‹–›•Š‘™•ƒ Ž‡ƒ”‰‡‘‰”ƒ’Š‹ ƒŽ’ƒ––‡”ȋŠƒ’–‡”͵ȌǤ

ƒ ‡•‘—” ‡ ƒ˜ƒ‹Žƒ„‹Ž‹–› Šƒ• —‡“—ƒŽ ‹ˆŽ—‡ ‡ ‹ •Šƒ’‹‰ –Š‡ •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‘ˆ –Š‡ „‡‡ ‘—‹–›Ǥ Ž‘”ƒŽ ”‡•‘—” ‡• ’ƒ”–‹ƒŽŽ› •Šƒ’‡ •’ƒ–‹ƒŽ †‹•–”‹„—–‹‘ ‘ˆ –Š‡ ‘—‹–›ǡ ™Š‹Ž‡ ‡•–‹‰”‡•‘—” ‡•ƒ”‡ƒŽ‘•–‹””‡Ž‡˜ƒ–ǤCistus•’’ǤˆŽ‘™‡”•ǡ‹’ƒ”–‹ —Žƒ”ǡ’Žƒ›ƒˆ‘”‡‘•–”‘Ž‡ ‹ •–”— –—”‹‰ –Š‡ „‡‡ ‘—‹–› ƒ– ƒ””ƒˆ ƒ–—”ƒŽ ƒ”ǡ „‡ ‘‹‰ ƒ ‹’‘”–ƒ– ’‘ŽŽ‡Ȁ‡ –ƒ”•‘—” ‡ƒ–ƒ–‹‡™Š‡‘–Š‡”ˆŽ‘™‡””‡•‘—” ‡•ƒ”‡• ƒ” ‡ȋŠƒ’–‡”•ͳƬʹȌǤ

ƒ ’‡ ‹‡• „‘†› •‹œ‡ ’Žƒ›• ƒ ‹’‘”–ƒ– ”‘Ž‡ †‡–‡”‹‹‰ –Š‡ •’ƒ–‹ƒŽ ’ƒ––‡”• ‘ˆ –Š‡ „‡‡ ‘—‹–›Ǥ ƒŽŽ •’‡ ‹‡• •Š‘™ •’ƒ–‹ƒŽ •–”— –—”‡ǡ ™Š‡”‡ƒ• Žƒ”‰‡ •’‡ ‹‡• –‡† –‘ •Š‘™ ƒ Š‘‘‰‡‡‘—• †‹•–”‹„—–‹‘Ǥ — Š ”‡Žƒ–‹‘•Š‹’ ‹• ’”‘„ƒ„Ž› †—‡ –‘ –Š‡ ‰”‡ƒ–‡•– ˆ‘”ƒ‰‹‰ †‹•–ƒ ‡•‘ˆŽƒ”‰‡•’‡ ‹‡•ǡ‹”‡Žƒ–‹‘–‘–Š‡‹”‰”‡ƒ–‡”ˆ‡‡†‹‰”‡“—‹”‡‡–•ȋŠƒ’–‡”ͳȌǤ

ƒ ƒ••‹˜‡ ‹–”‘†— –‹‘ ‘ˆ Š‘‡› „‡‡ ȋApis melliferaȌ ’‘’—Žƒ–‹‘• ”‡•—Ž–‹‰ ˆ”‘ „‡‡‡‡’‹‰ Šƒ••‹‰‹ˆ‹ ƒ–‡ˆˆ‡ –•‘’‘ŽŽ‡Ȁ‡ –ƒ”ƒ˜ƒ‹Žƒ„‹Ž‹–›Ǥ ‘‡›„‡‡•ƒ”‡–Š‡ƒ‹ ‘–”‹„—–‘”•–‘ ’‘ŽŽ‡Ȁ‡ –ƒ” ‘•—’–‹‘‘ˆ–Š‡–™‘ƒ‹ˆŽ‘™‡”‹‰•’‡ ‹‡•‹–Š‡ •–—†› ƒ”‡ƒǤ Š‡ ‘–”‹„—–‹‘‘ˆ„—„Ž‡„‡‡•ȋBombusterrestrisȌ‹•— ŠŽ‘™‡”ǡƒ†–Šƒ–‘ˆ‘–Š‡”™‹Ž†„‡‡•‹• ‹””‡Ž‡˜ƒ–ȋŠƒ’–‡”ʹȌǤ

ƒ Š‡™‹Ž†„‡‡ ‘—‹–›‹•‘†‹ˆ‹‡†‹ƒ”‡ƒ• Ž‘•‡–‘ƒ’‹ƒ”‹‡•Ǥ‹Ž†„‡‡„‹‘ƒ••†‡ ”‡ƒ•‡• ™‹–Š’”‘š‹‹–›–‘ƒ’‹ƒ”‹‡•†—‡–‘ƒŽ‘™‡”ƒ„—†ƒ ‡‘ˆŽƒ”‰‡„‡‡•ȋŠƒ’–‡”ʹȌǤƒ”‰‡„‡‡•ƒ”‡ ‡š’‡ –‡†–‘„‡–Š‡‘•–ƒˆˆ‡ –‡†„›’‘–‡–‹ƒŽ‡š’Ž‘‹–ƒ–‹˜‡ ‘’‡–‹–‹‘™‹–ŠŠ‘‡›„‡‡•†—‡ –‘–Š‡‹”Š‹‰Š‡”ˆ‡‡†‹‰”‡“—‹”‡‡–•ȋŠƒ’–‡”ʹȌǤ

ƒ ‡ ƒ—•‡ ˆŽ‘™‡” ƒ˜ƒ‹Žƒ„‹Ž‹–› ‹• Š‹‰ŠŽ› ˜ƒ”‹ƒ„Ž‡ ‹ •’ƒ ‡ ƒ† –‹‡ǡ‹–‹•†‹ˆˆ‹ —Ž––‘’”‘˜‹†‡ ‡•–‹ƒ–‡• ‘ˆ –Š‡ ƒ””›‹‰ ƒ’ƒ ‹–›ȋ•—•–ƒ‹ƒ„Ž‡Š‹˜‡†‡•‹–›Ȍ‘ˆ ƒ Šƒ„‹–ƒ–Ǥ ‘‡–Š‡Ž‡••ǡ †‡•‹–‹‡•‘˜‡”͵ǤͷŠ‹˜‡•’‡”ʹ•‡‡–‘ƒˆˆ‡ ––Š‡™‹Ž†„‡‡ ‘—‹–›‹‘—”•–—†›ƒ”‡ƒ ȋŠƒ’–‡”ʹȌǤ

 99 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

ƒ ƒ”ƒ•‹–‹•”ƒ–‡‘ˆ ƒ˜‹–›Ǧ‡•–‹‰„‡‡•ƒ†™ƒ•’•‹•‘–‡š’Žƒ‹‡†„›Š‘•–ƒ„—†ƒ ‡ƒ––Š‡ ‘—‹–›Ž‡˜‡ŽǤ ‘™‡˜‡”ǡƒ„—†ƒ–•’‡ ‹‡•–‡†–‘Šƒ˜‡Š‹‰Š‡”’ƒ”ƒ•‹–‹•”ƒ–‡Ǥ”‘„ƒ„Ž›ǡ ‰‡‡”ƒŽ‹•–’ƒ”ƒ•‹–‘‹†•ˆ‘ŽŽ‘™ƒˆ”‡“—‡ ›Ǧ†‡’‡†‡–†›ƒ‹ ƒ†Šƒ˜‡–Š‡ƒ„‹Ž‹–›–‘ˆ‘ —•‘ –Š‡‘•–ƒ„—†ƒ–•’‡ ‹‡•Ǥ–Š‡‘–Š‡”Šƒ†ǡ•’‡ ‹ƒŽ‹•–’ƒ”ƒ•‹–‘‹†•ƒ”‡’”‘„ƒ„Ž›•’‡ ‹ƒŽ‹œ‡† ‘ƒ„—†ƒ–•’‡ ‹‡•ǡ•‹ ‡‹–ƒ›„‡†‹ˆˆ‹ —Ž––‘‡•–ƒ„Ž‹•Šƒ•–ƒ„Ž‡’‘’—Žƒ–‹‘‘ƒŽ‘ ƒŽŽ›”ƒ”‡ Š‘•–ȋŠƒ’–‡”͵ȌǤ

ƒ Š‡ ‘—‹–›‘ˆ ƒ˜‹–›Ǧ‡•–‹‰„‡‡•ƒ†™ƒ•’•ȋŠ‘•–•Ȍƒ†–Šƒ– ‘ˆ ‹–• ƒ••‘ ‹ƒ–‡† ’ƒ”ƒ•‹–‘‹†•ǡ ƒ• ™‡ŽŽ ƒ• –Š‡‹” ‹–‡”ƒ –‹‘•ǡ •Š‘™ Š‹‰Š EǦ†‹˜‡”•‹–› ƒ– Ž‘ ƒŽ • ƒŽ‡Ǥ ‘•– ƒ† ’ƒ”ƒ•‹–‘‹†EǦ†‹˜‡”•‹–›’ƒ––‡”•ƒ”‡ƒ‹Ž›†—‡–‘•’‡ ‹‡•–—”‘˜‡””ƒ–Š‡”–Šƒ•’‡ ‹‡•Ž‘••ǡ ‡ƒ‹‰–Šƒ–™Š‹Ž‡•’‡ ‹‡•”‹ Š‡••†‘‡•‘–†‹ˆˆ‡”ƒ‘‰Ž‘ ƒŽ ‘—‹–‹‡•ǡ •’‡ ‹‡• ‘’‘•‹–‹‘ ‹• ‹†‹‘•› ”ƒ–‹  –‘ ‡ƒ Š •‹–‡Ǥ ‹‹Žƒ”Ž›ǡ EǦ†‹˜‡”•‹–› ‘ˆ •’‡ ‹‡• ‹–‡”ƒ –‹‘• ‹• ‘•–Ž›†—‡–‘•’‡ ‹‡•–—”‘˜‡””ƒ–Š‡”–ŠƒŽ‹”‡™‹”‹‰ȋ•Šƒ”‡† •’‡ ‹‡• ‹–‡”ƒ –‹‰ †‹ˆˆ‡”‡–Ž›ȌǤŠƒ–‹•ǡ‡˜‡‹ˆŽ‘ ƒŽ ‘—‹–‹‡•ƒ†‡–™‘”•†‹ˆˆ‡”ˆ”‘‡ƒ Š‘–Š‡”ǡ•’‡ ‹‡• ‹–‡”ƒ –•‹‹Žƒ”Ž›ƒ ”‘••–Š‡•–—†›ƒ”‡ƒȋŠƒ’–‡”͵ȌǤ

ƒ EǦ†‹˜‡”•‹–›‘ˆ–Š‡Š‘•– ‘—‹–›Šƒ•ƒƒ”‡†•’ƒ–‹ƒŽ•–”— –—”‡ǡ„‡ ‘‹‰‘”‡†‹••‹‹Žƒ” ™‹–Š‹ ”‡ƒ•‹‰‰‡‘‰”ƒ’Š‹ ƒŽ†‹•–ƒ ‡Ǥ ‘™‡˜‡”ǡEǦ†‹˜‡”•‹–›‘ˆ–Š‡’ƒ”ƒ•‹–‘‹† ‘—‹–›ƒ† EǦ†‹˜‡”•‹–›‘ˆ–Š‡‹–‡”ƒ –‹‘•†‘‘– Šƒ‰‡™‹–Š‰‡‘‰”ƒ’Š‹ ƒŽ†‹•–ƒ ‡ȋŠƒ’–‡”͵ȌǤ

ƒ EǦ†‹˜‡”•‹–›‘ˆŠ‘•–•ȋŽ‘™‡”–”‘’Š‹ Ž‡˜‡ŽȌ‹•ƒ„‡––‡”’”‡†‹ –‘”‘ˆEǦ†‹˜‡”•‹–›‘ˆ‹–‡”ƒ –‹‘• –Šƒ EǦ†‹˜‡”•‹–› ‘ˆ ’ƒ”ƒ•‹–‘‹†• ȋŠ‹‰Š‡” –”‘’Š‹  Ž‡˜‡ŽȌǤ ‹ ‡ Š‘•– ‘—‹–› –—”‘˜‡” ‹• ‰”‡ƒ–‡”–Šƒ’ƒ”ƒ•‹–‘‹† ‘—‹–›–—”‘˜‡”ǡ–Š‡ ‘–”‹„—–‹‘‘ˆŠ‘•––—”‘˜‡”–‘‹–‡”ƒ –‹‘ –—”‘˜‡”‹•‰”‡ƒ–‡”ȋŠƒ’–‡”͵ȌǤ

ƒ ’‡ ‹‡• ‘’‘•‹–‹‘ ‹• Š‹‰ŠŽ› †‡’‡†‡– ‘ Ž‘ ƒŽ ˆƒ –‘”• ȋŠƒ’–‡”• ͳ Ƭ ͵ȌǤ Š‹• Šƒ• ‹’Ž‹ ƒ–‹‘• ‘ –Š‡ •’ƒ–‹ƒŽ ˜ƒ”‹ƒ„‹Ž‹–› ‘ˆ ‹–‡”ƒ –‹‘•ǡ ™Š‹ Š ƒ ˜ƒ”› ƒ– ƒ ˜‡”› Ž‘ ƒŽ • ƒŽ‡ ȋͷͲͲ Ȃ ͳͲͲͲȌǡ Š‹‰ŠŽ‹‰Š–‹‰ –Š‡ ‡‡† –‘ •ƒ’Ž‡ ƒ Žƒ”‰‡ —„‡” ‘ˆ ’Ž‘–• –‘ ƒ†‡“—ƒ–‡Ž› Šƒ”ƒ –‡”‹œ‡–Š‡”‡‰‹‘ƒŽ‡–™‘”‹ƒ‰‹˜‡ƒ”‡ƒȋŠƒ’–‡”͵ȌǤ

ƒ ™‡–›Ǧˆ‹˜‡’Ž‘–•ƒ’’‡ƒ”–‘„‡•—ˆˆ‹ ‹‡––‘•ƒ’Ž‡ƒ ƒ˜‹–›Ǧ‡•–‹‰„‡‡ƒ†™ƒ•’ ‘—‹–› ƒ† ‹–• ‡•– ƒ••‘ ‹ƒ–‡• ‹ ƒ ͵ʹʹƒ”‡ƒ‘ˆ—‹–‡””—’–‡†Šƒ„‹–ƒ–ǡ„—–‘––‘•ƒ’Ž‡–Š‡‹” ‹–‡”ƒ –‹‘•Ǥ ‘™‡˜‡”ǡ”ƒ”‡ˆƒ –‹‘‡–Š‘†•†‘‘–•‡‡–‘„‡ƒ‰‘‘†ƒ’’”‘ƒ Š–‘‡˜ƒŽ—ƒ–‡ •ƒ’Ž‹‰“—ƒŽ‹–›‘ˆ‹–‡”ƒ –‹‘•„‡ ƒ—•‡–Š‡›‘˜‡”Ž‘‘ˆ‘”„‹††‡Ž‹•ȋŠƒ’–‡”͵ȌǤ





 100 

   

AppendixA

















  ’’‡†‹š

TableA1‡‡•’‡ ‹‡•ǡ–Š‡‹” ‘†‡—„‡”•ƒ†„‘†›•‹œ‡ ƒ–‡‰‘”›Ǥ

Family Species Codenumber Bodysize †”‡‹†ƒ‡ Andrenaangustior ͳ ƒŽŽ  Andrenacinerea ʹ ƒŽŽ  Andrenacombinata ͵ ƒŽŽ  Andrenadjelfensis Ͷ ƒŽŽ  Andrenaferrugineicrus ͷ ƒ”‰‡  Andrenafertoni ͸ ƒŽŽ  Andrenaflavipes ͹ ƒ”‰‡  Andrenahesperia ͺ ƒŽŽ  Andrenalabialis ͻ ƒ”‰‡  Andrenalagopus ͳͲ ƒŽŽ  Andrenalepida ͳͳ ƒŽŽ  Andrenalimata ͳʹ ƒ”‰‡  Andrenalivens ͳ͵ ƒŽŽ  Andrenanigroaenea ͳͶ ƒ”‰‡  Andrenanigroolivacea ͳͷ ƒ”‰‡  Andrenasenecionis ͳ͸ ƒŽŽ  Andrenasimilis ͳ͹ ƒŽŽ  Andrenasolenopalpa ͳͺ ƒŽŽ  Andrenasp. ͳͻ ƒŽŽ  Andrenatrimmerana ʹͲ ƒ”‰‡  Andrenaverticalis ʹͳ ƒŽŽ  Andrenavulpecula ʹʹ ƒŽŽ  Panurgusdentipes ʹ͵ ƒŽŽ ’‹†ƒ‡ Amegillaquadrifasciata ʹͶ ƒ”‰‡  Anthophoraacervorum ʹͷ ƒ”‰‡  Anthophoradispar ʹ͸ ƒ”‰‡  Apismellifera ʹ͹ ƒ”‰‡  Bombuspascuorum ʹͺ ƒ”‰‡  Bombuspratorum ʹͻ ƒ”‰‡  Bombusterrestris ͵Ͳ ƒ”‰‡  Ceratinacucurbitina ͵ͳ ƒŽŽ  Ceratinacyanea ͵ʹ ƒŽŽ  Euceraalternans ͵͵ ƒ”‰‡  Euceracaspica ͵Ͷ ƒ”‰‡  Eucerachrysopyga ͵ͷ ƒ”‰‡  Euceracollaris ͵͸ ƒ”‰‡  Euceraelongatula ͵͹ ƒ”‰‡

 103 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

 Euceranigrilabris ͵ͺ ƒ”‰‡  Eucerataurica ͵ͻ ƒ”‰‡  Melectaluctuosa ͶͲ ƒ”‰‡  Nomadaconnectens Ͷͳ ƒŽŽ  Nomadadicrepans Ͷʹ ƒŽŽ  Nomadadiscedens Ͷ͵ ƒŽŽ  Nomadaflavoguttata ͶͶ ƒŽŽ  Nomadahispanica Ͷͷ ƒŽŽ  Nomadaintegra Ͷ͸ ƒŽŽ  Nomadapanurgina Ͷ͹ ƒŽŽ  Nomadaserricornis Ͷͺ ƒŽŽ  Nomadasheppardana Ͷͻ ƒŽŽ  Xylocopaviolacea ͷͲ ƒ”‰‡ ‘ŽŽ‡–‹†ƒ‡ Hylaeusgarrulus ͷͳ ƒŽŽ  Hylaeusgibbus ͷʹ ƒŽŽ  Hylaeushyalinatus ͷ͵ ƒŽŽ  Hylaeustaeniolatus ͷͶ ƒŽŽ ƒŽ‹ –‹†ƒ‡ Halictusfulvipes ͷͷ ƒŽŽ  Halictusgemmeus ͷ͸ ƒŽŽ  Halictusquadricinctus ͷ͹ ƒ”‰‡  Halictusscabiosae ͷͺ ƒ”‰‡  Halictussimplex ͷͻ ƒŽŽ  Lasioglossumalbocinctum ͸Ͳ ƒŽŽ  Lasioglossumbimaculatum ͸ͳ ƒŽŽ  Lasioglossumgriseolum ͸ʹ ƒŽŽ  Lasioglossumibericum ͸͵ ƒŽŽ  Lasioglossuminterruptum ͸Ͷ ƒŽŽ  Lasioglossummalachurum ͸ͷ ƒŽŽ  Lasioglossummediterraneum ͸͸ ƒŽŽ  Lasioglossummorio ͸͹ ƒŽŽ  Lasioglossumsubhirtum ͸ͺ ƒŽŽ  Lasioglossumtransitorium ͸ͻ ƒŽŽ  Sphecodespseudofasciatus ͹Ͳ ƒŽŽ  Sphecodespuncticeps ͹ͳ ƒŽŽ  Sphecodesruficrus ͹ʹ ƒŽŽ ‡‰ƒ Š‹Ž‹†ƒ‡ Chelostomaflorisomne ͹͵ ƒŽŽ  Hoplitis(Anthocopa)sp. ͹Ͷ ƒŽŽ  Hoplitisadunca ͹ͷ ƒŽŽ  Hoplitisanthocopoides ͹͸ ƒŽŽ  Hoplitisbenoisti ͹͹ ƒŽŽ

 104   ’’‡†‹š

 Hoplosmialigurica ͹ͺ ƒŽŽ  Megachilebaetica ͹ͻ ƒ”‰‡  Megachileericetorum ͺͲ ƒ”‰‡  Megachilepyrenaica ͺͳ ƒ”‰‡  Osmiaaurulenta ͺʹ ƒŽŽ  Osmiagallarum ͺ͵ ƒŽŽ  Osmialatreillei ͺͶ ƒ”‰‡  Osmiamelanogaster/leaiana ͺͷ ƒŽŽ  Osmiamustelina ͺ͸ ƒ”‰‡  Osmianasoproducta ͺ͹ ƒ”‰‡  Osmianasuta ͺͺ ƒŽŽ  Osmianiveata ͺͻ ƒ”‰‡  Osmianiveocincta ͻͲ ƒ”‰‡  Osmiarufohirta ͻͳ ƒŽŽ  Osmiasubmicans ͻʹ ƒŽŽ  Osmiatricornis ͻ͵ ƒ”‰‡  Protosmia(Nanosmia)sp. ͻͶ ƒŽŽ  Protosmiacapitata ͻͷ ƒŽŽ  Protosmiaexenterata ͻ͸ ƒŽŽ  Rhodanthidiumseptemdentatum ͻ͹ ƒ”‰‡  Rhodanthidiumsticticum ͻͺ ƒ”‰‡

 105 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

TableA2‡ƒƒ† ‘‡ˆˆ‹ ‹‡–‘ˆ˜ƒ”‹ƒ–‹‘ȋαʹͳ’Ž‘–•Ȍ‘ˆˆŽ‘™‡”ƒ†‡•–‹‰”‡•‘—” ‡ ˜ƒ”‹ƒ„Ž‡•Ǥ

 Mean CV Flowers Ž‘™‡”ƒ„—†ƒ ‡ȋˆŽ‘™‡”•ȀʹȌ ͳͳͲǤͷͺ ͲǤ͸ͳ Ž‘™‡””‹ Š‡•• ͳͷǤͷ͹ ͲǤ͵Ͷ DorycniumpentaphyllumˆŽ‘™‡”•Ȁʹ ͺǤͺ͹ ͳǤͳͷ RosmarinusofficinalisˆŽ‘™‡”•Ȁʹ ͷʹǤͲͳ ͲǤ͹Ͷ ThymusvulgarisˆŽ‘™‡”•Ȁʹ ͶͻǤͳͳ ͳǤͲ͵ Cistus•’’ǤˆŽ‘™‡”•Ȁʹ ͲǤͷͻ ͳǤͷͶ Nestingresources Ψƒ”‡•‘‹Ž ͳ͵Ǥͳ ͲǤ͹ͳ Ψ‡ƒ†™‘‘† ͲǤͳ ʹǤͷͶ ΨQuercuscoccifera ‘˜‡” ͳʹǤͻ ͳǤͲͺ ΨAmpelodesmosmauritanica ‘˜‡” ͸Ǥͳ ͳǤͲͺ —„‡”‘ˆŠ‘Ž‡•‹”‘ •Ȁʹ ͲǤͷͷ ͳǤʹͺ —„‡”‘ˆ•ƒ‹Ž•Š‡ŽŽ•Ȁʹ ͲǤͳͻ ͳǤͶͲ 

 106   ’’‡†‹š

TableA3‹•–‘ˆˆŽ‘™‡”‹‰’Žƒ–•’‡ ‹‡•Ǥ

Flowerspecies Alliumroseum Alliumsphaerocephalon Anagallisarvensis Antirrhinumbarrelieri Aphyllanthesmonspeliensis Argyrolobiumzanonii Aristolochiapistolochia Asperulacynanchia Biscutellalaevigata Centaurealinifolia Centaureamontanassp.semidecurrens Centauriumerythraea Cistusalbidus Cistusmonspeliensis Cistussalviifolius Clematisflammula Convolvuluslanuginosus Corismonspeliensis Cytisophyllumsessilifolium Dorycniumhirsutum Dorycniumpentaphyllum Echiumvulgare Ericamultiflora Euphorbiacharacias Euphorbiaflavicoma Fumanaericifolia Fumanaericoides Fumanalaevipes Fumanalaevis Galiumaparine Galiumpalustre Genistascorpius Gladiolusillyricus Helianthemumoelandicumspp.italicum Helichrysumstoechas Hippocrepiscomosa Hypericumperforatum

 107 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

Lathyrus Leuzeaconifera Linumstrictum Lithospermumfruticosum Loniceraimplexa Muscarineglectum Narcissusassoanus Ononisminutissima Orobanchelatisquama Phlomislychnitis Polygalarupestris Potentillasp. Psoralerabituminosa Ranunculusgramineus Rosmarinusofficinalis Rubiaperegrina Scorpiurusmuricatus Sedumsediforme Sideritishirsuta Teucriumchamaedrys Thalictrumtuberosum Thesiumdivaricatum Thymusvulgaris Torilisarvensis Vicentoxicumhirundinaria Vicentoxicumnigrum Viciacracca 

 108 

TableA4‘†‡Ž•‡Ž‡ –‹‘„ƒ•‡†‘ƒ‹‡ǯ• ˆ‘”ƒ–‹‘”‹–‡”‹‘ȋ ȌǤ

‡’‡†‡–˜ƒ”‹ƒ„Ž‡ †‡’‡†‡–˜ƒ”‹ƒ„Ž‡•    ȟ   –‡” ‡’– ‘ˆˆ ˜—Ž ’‡ ‹• ™ • ‹” •ƒ‹Ž    ’‡ ‹‡•”‹ Š‡•• š š š š š š š š š ͻ ͳ͵͹ǤͺͶ ͳͲǤͶͷ  š š š š š š  š š ͺ ͳ͵͸ǤͳͶ ͺǤ͹ͷ  š š š  š š  š š ͹ ͳ͵ͶǤͶ͸ ͹ǤͲ͹  š š š  š   š š ͸ ͳ͵͵Ǥʹͻ ͷǤͻ  š  š  š   š š ͷ ͳ͵ʹǤͲ͹ ͶǤ͸ͺ  š  š  š   š  Ͷ ͳ͵ͲǤͷʹ ͵Ǥͳ͵  š    š   š  ͵ ͳʹͻǤͲ͹ ͳǤ͸ͺ  x x2 127.39 0.00  šͳͳʹͺǤͳͲ ͲǤ͹ͳ „—†ƒ ‡ š š š š š š š š š ͻ ͳͳǤ͵͹ ͳͲǤͻͳ  š š š š š  š š š ͺ ͻǤ͵ͺ ͺǤͻʹ  š  š š š  š š š ͹ ͹ǤͶͳ ͸Ǥͻͷ  š  š š š  š  š ͸ ͷǤͶ͸ ͷǤͲͲ  š  š š š    š ͷ ͵Ǥ͸Ͷ ͵Ǥͳͺ  ššššͶͳǤ͹͸ͳǤ͵Ͳ  xxx3 0.46 0.00  ššʹͲǤ͹͵ͲǤʹ͹  šͳͷǤͻͶͷǤͶͺ ƒŽ›•‡•‘ˆ–Š‡”‡Žƒ–‹‘•Š‹’‘ˆ„‡‡•’‡ ‹‡•”‹ Š‡••ƒ†ƒ„—†ƒ ‡ȋαʹͳ’Ž‘–•Ȍ™‹–ŠˆŽ‘™‡”ƒ†‡•–‹‰”‡•‘—” ‡˜ƒ”‹ƒ„Ž‡•Ǥƒ”‹ƒ„Ž‡•‹ Ž—†‡†‹‡ƒ Š‘†‡Žƒ”‡ƒ”‡†™‹–ŠƒšǤŠ‡•‡Ž‡ –‡† ‘†‡Ž‹•‹„‘Ž†ǡȋȌ‹•–Š‡—„‡”‘ˆ’ƒ”ƒ‡–‡”•‹–Š‡‘†‡Žƒ†ȋȟ Ȍ‹•–Š‡†‹ˆˆ‡”‡ ‡‹ „‡–™‡‡–Š‡•‡Ž‡ –‡†‘†‡Žƒ†–Š‡‰‹˜‡‘†‡ŽǤ †‡’‡†‡–˜ƒ”‹ƒ„Ž‡•ƒ”‡†‡•‹–›‘ˆR.officinalis ˆŽ‘™‡”•Ȃ‘ˆˆǦǢT.vulgarisˆŽ‘™‡”•–˜—ŽǦǡD.pentaphyllumˆŽ‘™‡”•–’‡ǦǡCistus•’’ǤˆŽ‘™‡”•Ȃ‹•Ǧǡ’”‡•‡ ‡‘ˆ†‡ƒ†™‘‘†Ȃ™ǦǡΨ‘ˆ„ƒ”‡•‘‹ŽȂ•Ǧǡ—„‡”‘ˆŠ‘Ž‡•‹”‘ •Ȃ ‹”Ǧƒ†—„‡”‘ˆ ˜ƒ ƒ–•ƒ‹Ž•Š‡ŽŽ•Ȃ•ƒ‹ŽǦǤ





   

AppendixB















  ’’‡†‹š

TableB1ƒ˜‹–›Ǧ‡•–‹‰„‡‡ƒ†™ƒ•’•’‡ ‹‡•ȋŠ‘•–•Ȍ™‹–Š–Š‡‹” ‘†‡•Ǥ

Code Species Bee/wasp Ͳͳ Ampulexruficollis ƒ•’ Ͳʹ Ancistroceruslongispinosus ƒ•’ Ͳ͵ Anthidiumflorentinum ‡‡ ͲͶ Anthidiumnigricolle ‡‡ Ͳͷ Auplopuscarbonarius ƒ•’ Ͳ͸ Chelostomaedentulum ‡‡ Ͳ͹ Diodontussp.1 ƒ•’ Ͳͺ Dipogonsp.1 ƒ•’ Ͳͻ Dipogonsp.2 ƒ•’ ͳͲ Dipogonsp.3 ƒ•’ ͳͳ Eumenidaesp.1 ƒ•’ ͳʹ Eumenidaesp.2 ƒ•’ ͳ͵ Eumenidaesp.3 ƒ•’ ͳͶ Euodynerussp.1 ƒ•’ ͳͷ Heriadescrenulatus ‡‡ ͳ͸ Hoplitisadunca ‡‡ ͳ͹ Hylaeushyalinatus ‡‡ ͳͺ Hylaeustaeniolatus ‡‡ ͳͻ Isodontiamexicana ƒ•’ ʹͲ Megachileapicalis ‡‡ ʹͳ Megachileericetorum ‡‡ ʹʹ Megachilerotundata ‡‡ ʹ͵ Nitelafallax ƒ•’ ʹͶ Nitelasp.1 ƒ•’ ʹͷ Nitelatruncata ƒ•’ ʹ͸ Osmiabicornis ‡‡ ʹ͹ Osmiacaerulescens ‡‡ ʹͺ Osmialatreillei ‡‡ ʹͻ Osmiamelanogaster ‡‡ ͵Ͳ Osmianasoproducta ‡‡ ͵ͳ Osmiasubmicans ‡‡ ͵ʹ Osmiatricornis ‡‡ ͵͵ Passaloecusgracilis ƒ•’ ͵Ͷ Passaloecuspictus ƒ•’ ͵ͷ Pisonatrum ƒ•’ ͵͸ ‘’‹Ž‹†ƒ‡Gen.sp.1 ƒ•’ ͵͹ Psenulusfuscipennis ƒ•’ ͵ͺ Solierellacompedita ƒ•’  113 ’ƒ–‹ƒŽ˜ƒ”‹ƒ„‹Ž‹–›‘ˆ„‡‡ ‘—‹–‹‡•  

͵ͻ Solierellasp.1 ƒ•’ ͶͲ Trypoxylonsp.1 ƒ•’ Ͷͳ Trypoxylonsp.2 ƒ•’  

 114   ’’‡†‹š

TableB2ƒ”ƒ•‹–‡ǡ’ƒ”ƒ•‹–‘‹†ǡ Ž‡’–‘’ƒ”ƒ•‹–‡ƒ†• ƒ˜‡‰‡”•’‡ ‹‡•ƒ••‘ ‹ƒ–‡†–‘–”ƒ’Ǧ‡•–‹‰Š‘•–•ǡ –Š‡‹” ‘†‡•ƒ†–›’‡‘ˆ‹–‡”ƒ –‹‘Ǥ

Code Species Order Interactiontype Ͳͳ Anthraxanthrax ‹’–‡”ƒ ƒ”ƒ•‹–‘‹† Ͳʹ Anthraxsp.2 ‹’–‡”ƒ ƒ”ƒ•‹–‘‹† Ͳ͵ Calliphoridaesp.1 ‹’–‡”ƒ ƒ”ƒ•‹–‘‹† ͲͶ Chaetodactylusosmiae ƒ” ‘’–‹ˆ‘”‡ Ž‡’–‘’ƒ”ƒ•‹–‡ Ͳͷ Chalcissp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† Ͳ͸ Chrysisignita ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† Ͳ͹ Chrysurasp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† Ͳͺ Coelioxysechinata ›‡‘’–‡”ƒ Ž‡’–‘’ƒ”ƒ•‹–‡ Ͳͻ Cystomutillasp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ͳͲ Gasteruptionsp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ͳͳ Hedycridiumsp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ͳʹ Hybomischossp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ͳ͵ Leucospisdorsigera ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ͳͶ Melittobiaacasta ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ͳͷ Zonitisimmactulata ‘Ž‡‘’–‡”ƒ Ž‡’–‘’ƒ”ƒ•‹–‡ ͳ͸ Miltogrammaspp. ‹’–‡”ƒ ƒ”ƒ•‹–‘‹† ͳ͹ —–‹ŽŽ‹†ƒ‡Gen.sp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ͳͺ Omalussp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ͳͻ Omalussp.2 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ʹͲ –‡”‘ƒŽ‹†ƒ‡Gen.sp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ʹͳ Sapygaquinquepunctata ›‡‘’–‡”ƒ Ž‡’–‘’ƒ”ƒ•‹–‡ ʹʹ Toxophorafasciculata ‹’–‡”ƒ ƒ”ƒ•‹–‘‹† ʹ͵ Trichodesleucopsideus ‘Ž‡‘’–‡”ƒ ”‡†ƒ–‘” ʹͶ Trichrysiscyanea ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹† ʹͷ Trogodermasp.1 ‘Ž‡‘’–‡”ƒ  ƒ˜‡‰‡” ʹ͸ Xoridessp.1 ›‡‘’–‡”ƒ ƒ”ƒ•‹–‘‹†  

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