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Individual Variation of Behaviour in Firebugs (Pyrrhocoris Apterus ): Causes and Consequences

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Individual Variation of Behaviour in Firebugs (Pyrrhocoris Apterus ): Causes and Consequences DE TTK 1949 Individual variation of behaviour in firebugs (Pyrrhocoris apterus ): causes and consequences A viselkedés egyedi változatossága a ver őkölt ő bodobácsnál (Pyrrhocoris apterus ): okok és következmények Egyetemi doktori (PhD) értekezés Gyuris Enik ő Témavezet ő: Prof. Barta Zoltán DEBRECENI EGYETEM Természettudományi Doktori Tanács Juhász-Nagy Pál Doktori Iskola Debrecen, 2013. Ezen értekezést a Debreceni Egyetem Természettudományi Doktori Tanács Juhász-Nagy Pál Doktori Iskola Biodiverzitás programja keretében készítettem a Debreceni Egyetem természettudományi doktori (PhD) fokozatának elnyerése céljából. Debrecen, 2013 . ……………………………. Gyuris Enik ő Tanúsítom, hogy Gyuris Enik ő doktorjelölt 2009-2012 között a fent megnevezett Doktori Iskola Biodiverzitás programjának keretében irányításommal végezte munkáját. Az értekezésben foglalt eredményekhez a jelölt önálló alkotó tevékenységével meghatározóan hozzájárult. Az értekezés elfogadását javasolom. Debrecen, 2013 . ……………………………… Prof. Barta Zoltán ii Individual variation of behaviour in firebugs (Pyrrhocoris apterus ): causes and consequences A viselkedés egyedi változatossága a ver őkölt ő bodobácsnál (Pyrrhocoris apterus ): okok és következmények Értekezés a doktori (Ph.D.) fokozat megszerzése érdekében a biológia tudományágban Írta: Gyuris Enik ő okleveles biológus Készült a Debreceni Egyetem Juhász-Nagy Pál doktori iskolája (Biodiverzitás programja) keretében Témavezet ő: Prof. Barta Zoltán A doktori szigorlati bizottság: elnök: Dr. Bánfalvi Gáspár........................ .............................................. tagok: Dr. Pénzes Zsolt.............................. .............................................. Dr. Végvári Zsolt............................ .............................................. A doktori szigorlat id őpontja: 2012.október 16. Az értekezés bírálói: Dr. .................................................. .............................................. Dr. .................................................. .............................................. Dr. .................................................. .............................................. A bírálóbizottság: elnök: Dr. .................................................. .............................................. tagok: Dr. .................................................. .............................................. Dr. .................................................. .............................................. Dr. .................................................. .............................................. Dr. .................................................. .............................................. Az értekezés védésének id őpontja: 201… . ……………… … . iii Tartalomjegyzék - Contents CHAPTER I. 1 A VISELKEDÉS EGYEDI VÁLTOZATOSSÁGA A VER ŐKÖLT Ő BODOBÁCSNÁL (PYRRHOCORIS APTERUS ): OKOK ÉS KÖVETKEZMÉNYEK I. 1. Bevezetés 2 I. 2. Célkit űzések 9 I. 2. 1. Konzisztens egyedi különbségek a viselkedésben 15 I. 2. 2. Személyiségjegyek stabilitása és változásai az egyedfejl ődés során 16 I. 2. 3. A Hemipteroseius adleri atka megjelenése hazánkban 17 I. 2. 4. A ver őkölt ő bodobácsok ( Pyrrhocoris apterus ) személyiségjegyeinek és a Hemipteroseius adleri atkának kapcsolata 17 I. 3. Az értekezés új tudományos eredményei 18 I. 3. 1. Konzisztens egyedi különbségek a viselkedésben 18 I. 3. 2. Személyiségjegyek stabilitása és változásai az egyedfejl ődés során 19 I. 3. 3. A Hemipteroseius adleri atka megjelenése hazánkban 15 I. 3. 4. A ver őkölt ő bodobácsok ( Pyrrhocoris apterus ) személyiségjegyeinek és a Hemipteroseius adleri atkának kapcsolata 15 I. 4. Összefoglalás és jöv őbeni tervek 16 I. 5. Köszönetnyilvánítás 18 I. 6. Irodalomjegyzék 19 CHAPTER II. 25 INDIVIDUAL VARIATION OF BEHAVIOUR IN FIREBUGS ( PYRRHOCORIS APTERUS ): CAUSES AND CONSEQUENCES II. 1. Introduction 26 II. 2. Aims 37 II. 2. 1. Consistent individual differences in behaviour 39 II. 2. 2. Stability and changes of personality traits across ontogeny 39 II. 2. 3. Occurrence of Hemipteroseius adleri ectoparasitic mites in Hungary 35 II. 2. 4. Ectoparasitic mites ( Hemipteroseius adleri ) and firebugs’ ( Pyrrhocoris apterus ) personality traits 36 II. 3. New scientific results of the dissertation 37 II. 3. 1. Consistent individual differences in behaviour 37 II. 3. 2. Stability and changes of personality traits across ontogeny 37 II. 3. 3. Occurrence of ectoparasitic mites ( Hemipteroseius adleri ) in Hungary 38 II. 3. 4. Ectoparasitic mites ( Hemipteroseius adleri ) and firebugs’ ( Pyrrhocoris apterus ) personality traits 38 II. 4. Conclusion and future prospects 39 II. 5. Acknowledgements 41 II. 6. References 42 iv CHAPTER III. 48 STUDY 1: INDIVIDUAL BEHAVIOUR IN FIREBUGS ( PYRRHOCORIS APTERUS ) CHAPTER IV. 55 STUDY 2: PERSONALITY TRAITS ACROSS ONTOGENY IN FIREBUGS, PYRRHOCORIS APTERUS CHAPTER V. 63 STUDY 3: HEMIPTEROSEIUS ADLERI COSTA, 1968 COLLECTED ON RED FIREBUG: THE FIRST RECORD OF THE FAMILY OTOPHEIDOMENIDAE TREAT, 1955 (ACARI: MESOSTIGMATA) IN HUNGARY CHAPTER VI. 67 STUDY 4: ECTOPARASITIC MITES (HEMIPTEROSEIUS ADLERI ) AND FIREBUGS’ (PYRRHOCORIS APTERUS ) PERSONALITY TRAITS v Chapter I. A viselkedés egyedi változatossága a ver őkölt ő bodobácsnál (Pyrrhocoris apterus ): okok és következmények 1 I. 1. Bevezetés Az állati viselkedés tanulmányozása az utóbbi évtizedben egy jelent ős változáson ment keresztül. A viselkedésben fellép ő egyedi különbségek jelensége, melyet régebben „zajnak”, az egyedfejl ődés során fellép ő „hibák” következményének tekintettek, ma már a viselkedésökológiai kutatások egyik központi kérdése lett (Réale és mtsai 2010). Ezeket a viselkedésben megjelen ő egyedei különbségeket, melyek id őn és szituációkon keresztül állandóak, és gyakran egymással összefüggnek szokták viselkedési szindrómának, vérmérsékletnek (temperamentumnak), viselkedési típusnak illetve, állati személyiségnek is definiálni (Sih és mtsai 2004, Réale és mtsai 2007, Sih és Bell 2008). Egy nemrég megjelent tanulmány szerint azonban ezeket az eddig egymás szinonimájaként használt fogalmakat az alábbiak szerint pontosíthatjuk (Herczeg és Garamszegi 2012): Vérmérséklet : Egy egyedre jellemz ő hipotetikus jelleg, mely különböz ő élethelyzetben jelenik meg különböz ő viselkedésekként (pl. kockázat vállaló viselkedés szintje predációs környezetben). Állati személyiség : Egyedi viselkedési különbségek, mely kontextusokon és id őn keresztül konzisztensek. Leggyakrabban vizsgált személyiség jellegek a bátorság, felfedez őképesség, aktivitás, szociabilitás és agresszivitás (Réale és mtsai 2007). Viselkedési típus : Egy egyedre jellemz ő személyiség jelleg. Viselkedési szindróma : Csoport szinten (pl. populáció) értelmezhet ő legalább két személyiség jelleg között megjelen ő rang-sorrenden alapuló korreláció (Bell 2007). Az, hogy az emberek rendelkeznek stabil egyedi viselkedésbeli különbséggel már egy régóta köztudott tény (Eysenck 1953), azonban viselkedésökológusok nemrég felfigyeltek arra a jelenségre, hogy ezek a különbségek leegyszer űsödve, de nagyon hasonlóképpen megjelenhetnek állatfajok számunkra teljesen azonosnak t űnő egyedeinél is. Riechert és Hedrik (1990) a személyiség kutatás kezdeti szakaszán végzett vizsgálata arra a kérdésre kereste a választ, hogy a környezeti hatások hogyan hatnak a korrelált viselkedési minták evolúciójára. Az észak-amerikai sivatagi „füves” pók ( Agelenopsis aperta ) el őfordul mind száraz, füves él őhelyeken, mind pedig páradús, folyó menti habitaton is. A száraz él őhelyen kevesebb a préda és kisebb a predációs kockázat, ugyanakkor nagyobb a kompetíció a területért. Ezzel szemben a folyó melletti habitaton a pókok több predátorral 2 találkozhatnak, de magasabb a táplálék ellátottság is és így kisebb a kompetíció. Megfigyelték, hogy a vízi él őhelyen el őforduló pókok félénkebbek, sok id őt töltenek el a ragadozó el őli bujkálással, ugyanis az ebb ől származó el őny nagyobb, mint az adott esetben elszalasztott táplálékból származó hátrány, hiszen nagy a táplálék ellátottság. Ezzel szemben a füves él őhelyeken él ő pókoknak a territórium fenntartásáért el őnyös agresszívabbként viselkedniük. Ezen természetben megfigyelt két személyiség jelleget (bátorság, agresszivitás) kísérleti úton is tesztelték, és azt tapasztalták, hogy a pókok viselkedése a helyi környezethez való adaptációnak tekinthet ő, azaz a természetes szelekció a vizes él őhelyen a félénkebb, kevésbé agresszívebb egyedeket, míg a száraz él őhelyeken a bátrabb, agresszívebb egyedeket részesíti el őnyben (Riechert és Hall 2000). A viselkedésben megjelen ő egyedi különbségek evolúciós alapjainak megértéséhez egy másik érdekes jelenség a viselkedési „túlfolyás ( spillover )”, mely azon eseteket írja le, amikor egy jellegre ható pozitív szelekció egy adott helyzetben maladaptív viselkedést eredményez egy másik helyzetben a jellegek közötti kapcsoltság miatt. (Sih és mtsai 2004). Egy észak-amerikai halászpók fajnál ( Dolomedes triton ) azt találták, hogy egyes n őstény pókok gyakran a párzás el őtt felfalják a nekik udvarló hímeket (Johnson és Sih 2005). Ez a „prekopulációs kannibalizmus” valószín űleg annak az eredménye, hogy a természetes szelekció azokat a juvenilis n őstényeket részesíti el őnyben, akik a táplálékhiány miatt agresszíven viselkedtek. Megfigyelték, hogy az agresszivitás feln őttkorban is stabilan megmarad és korrelál a fekunditással, ugyanakkor egyes esetekben a n őstények a túlzott agressziónak súlyos árát fizetik, azaz nem szaporodnak. A viselkedési és fiziológiai jellegek sokszor nem függetlenek
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