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FILOGENETSKA SISTEMATIKA in EVOLUCIJA MREŽ PRI PAJKIH MREŽARJIH SKUPIN Zygiella SENSU LATO in Caerostris

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FILOGENETSKA SISTEMATIKA in EVOLUCIJA MREŽ PRI PAJKIH MREŽARJIH SKUPIN Zygiella SENSU LATO in Caerostris UNIVERZA V LJUBLJANI BIOTEHNIŠKA FAKULTETA Matjaž GREGORIČ FILOGENETSKA SISTEMATIKA IN EVOLUCIJA MREŽ PRI PAJKIH MREŽARJIH SKUPIN Zygiella SENSU LATO IN Caerostris DOKTORSKA DISERTACIJA Ljubljana, 2013 UNIVERZA V LJUBLJANI BIOTEHNIŠKA FAKULTETA Matjaž GREGORIČ FILOGENETSKA SISTEMATIKA IN EVOLUCIJA MREŽ PRI PAJKIH MREŽARJIH SKUPIN Zygiella SENSU LATO IN Caerostris DOKTORSKA DISERTACIJA PHYLOGENETIC SYSTEMATICS AND WEB EVOLUTION IN THE ORB WEAVING Zygiella SENSU LATO AND Caerostris DOCTORAL DISSERTATION Ljubljana, 2013 Gregorič M., Filogenetska sistematika in evolucija mrež pri pajkih mrežarjih...Zygiella...Caerostris. Dokt. disertacija. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, 2013 II Doktorska disertacija je bila opravljena na Biološkem inštitutu Jovana Hadžija, ZRC SAZU, Slovenija. Dodatno laboratorijsko delo je bilo opravljeno na Department of Biology, University of Akron, Ohio, ZDA in na Department of Biology, University of Puerto Rico, San Juan, Portoriko, ZDA. Terensko delo je bilo opravljeno v Sloveniji, Indoneziji, Singapurju, Maleziji, Portoriku, na Kitajskem in na Madagaskarju. Komisija za doktorski študij Univerze v Ljubljani je za mentorja doktorske disertacije imenovala viš. znan. sod. doc. dr. Matjaža Kuntnerja. Komisija za oceno in zagovor: Predsednik: prof. dr. Rok KOSTANJŠEK Univerza v Ljubljani, Biotehniška fakulteta, Odelek za biologijo Član: doc. dr. Cene FIŠER Univerza v Ljubljani, Biotehniška fakulteta, Odelek za biologijo Član in mentor: višji znan. sod. doc. dr. Matjaž KUNTNER Biološki Inštitut Jovana Hadžija ZRC SAZU Datum zagovora: 27. 9. 2013 Doktorsko delo je rezultat lastnega raziskovalnega dela. Podpisani se strinjam z objavo svoje naloge v polnem tekstu na spletni strani Digitalne knjižnice biotehniške fakultete. Izjavljam, da je naloga, ki sem jo oddal v elektronski obliki, identična tiskani verziji. Matjaž Gregorič Gregorič M., Filogenetska sistematika in evolucija mrež pri pajkih mrežarjih...Zygiella...Caerostris. Dokt. disertacija. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, 2013 III KLJUČNA DOKUMENTACIJSKA INFORMACIJA (KDI) ŠD Dd DK UDK 565.4:575.85(043.3) KG Zygiellidae/Caerostris/evolucija mrež/filogenetska sistematika/vedenjska plastičnost AV GREGORIČ, Matjaž, univ. dipl. biol. SA KUNTNER, Matjaž (mentor) KZ SI-1000 Ljubljana, Večna pot 111 ZA Univerza v Ljubljani, Biotehnična fakulteta, Oddelek za biologijo LI 2013 IN FILOGENETSKA SISTEMATIKA IN EVOLUCIJA MREŽ PRI PAJKIH MREŽARJIH SKUPIN Zygiella SENSU LATO IN Caerostris TD Doktorska disertacija OP XIII, 164 str., 7 pregl., 27 sl., 3 pril., 164 vir. IJ sl JI sl / en AI Pajki mrežarji iz naddružine Araneoidea so pomembni in pogosto celo modelni organizmi v mnogih evolucijskih študijah, npr. v raziskovanju adaptivne evolucije, evolucijske ekologije in evolucije vedenja v najširšem smislu. V doktorski disertaciji smo tako raziskovali ali različne lastnosti pajčjih mrež, ki vplivajo na vrstno pestrost pajkov, med seboj koevoluirajo, in ali ontogenetske spremembe v vedenju resnično sledijo biogenetskemu pravilu in kako individualni pajki glede na svojo velikost spreminjajo energijski vložek v mreže. Večino študij smo izvajali na rodovih Zygiella in Caerostris, oba iz družine križevcev (Araneidae), ki sta filogenetsko problematična in sta zanimiva tudi po biologiji mrež. Ta rodova smo v disertaciji filogenetsko umestili in raziskali evolucijo njunih mrež. Pokazali smo, da ločene filogenetske linije obeh rodov kažejo značilne vedenjske posebnosti. Rezultati raziskav v doktorski disertaciji nakazujejo, da prvič, arhitektura mrež in mehanske lastnosti dejansko koevoluirajo z ekološkimi lastnostmi mrež in vedenjem ob njihovi gradnji. Drugič, pajčje mreže med ontogenetskim razvojem pajkov ne sledijo splošnemu biogenetskemu pravilu, ampak se spreminjajo tako, da osebku omogočajo čim učikovitejše bivanje v svojem mikrookolju, v našem primeru učinkovito plenjenje. Tretjič, energijski vložek v plenjenje je verjetno pod močnim selekcijskim pritiskom, pajki pa ga spreminjajo s prirejanjem arhitekture mrež in spreminjanjem svilenih niti in lepila, zato je za natančno oceno materialnega vložka v plenjenje potrebna kvantifikacija vsega naštetega. Četrtič, skupino Zygiellidae lahko obravnavamo kot veljaven taksonomski pojem in je verjetno sestrska skupina ostalim predstavnikom družine Araneidae. Petič, moji rezultati rod Caerostris filogenetsko uvrščajo med evolucijsko izvorne predstavnike družine Araneidae, rod je verjetno precej raznovrstnejši, opisali pa smo tudi pet novih vrst. Gregorič M., Filogenetska sistematika in evolucija mrež pri pajkih mrežarjih...Zygiella...Caerostris. Dokt. disertacija. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, 2013 IV KEYWORDS DOCUMENTATION (KWD) DN Dd DC UDK 565.4:575.85(043.3) CX Zygiellidae/Caerostris/web evolution/phylogenetic systematics/behavioral plasticity AU GREGORIČ, Matjaž, univ. dipl. biol. AA KUNTNER, Matjaž (mentor) PP SI-1000 Ljubljana, Večna pot 111 PB Univerza v Ljubljani, Biotehnična fakulteta, Oddelek za biologijo PY 2013 TI PHYLOGENETIC SYSTEMATICS AND WEB EVOLUTION IN THE ORB WEAVING Zygiella SENSU LATO AND Caerostris DT Doctoral dissertation NO XIII, 164 p., 7 tabl., 27 fig., 3 ann., 164 ref. LA sl AL sl / en AB The orb weaving araneoid spiders are considered important or even model organisms in several topics of evolutionary research, like adaptive evolution, evolutionary ecology and evolution of behavior in the broadest sense. In my doctoral thesis, I investigated whether spider webs coevolve with other traits that influence spider diversification, whether ontogenetic changes in spider webs really follow the biogenetic law, and how individual spiders alter their foraging investment in relation to body size. I conducted most studies on the genera Zygiella s.l. and Caerostris from the family Araneidae. Both genera are taxonomically controversial and exhibit interesting web biology. Thus, I phylogenetically placed both genera and investigated the evolution of their webs. The results in the doctoral dissertation show that first, the results continue to build evidence for the coevolution of behavioral (web building), ecological (web microhabitat) and biomaterial (silk biomechanics) traits. Second, biogenetic law cannot explain spider orb web allometry, but our results instead support optimization of foraging area in response to spider size. Third, in relation to body size, spiders alter their webs through i) web architecture, ii) investment in silk, and iii) investment in glue, and that quantifying all three parameters is necessary for meaningful estimates of total foraging investment. Fourth, Zygiellidae can be regarded a valid taxonomic concept and is probably sister to other araneids. Fifth, my results recover Caerostris as basal araneid, the genus probably contains several more species, and we also describe five new species. Gregorič M., Filogenetska sistematika in evolucija mrež pri pajkih mrežarjih...Zygiella...Caerostris. Dokt. disertacija. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, 2013 V KAZALO VSEBINE KLJUČNA DOKUMENTACIJSKA INFORMACIJA .....................................................III KEY WORDS DOCUMENTATION ............................................................................... IV KAZALO VSEBINE ......................................................................................................... V KAZALO ZNANSTVENIH DEL ................................................................................ VIII KAZALO PREGLEDNIC ............................................................................................. IX KAZALO SLIK ................................................................................................................. X KAZALO PRILOG ....................................................................................................... XII OKRAJŠAVE IN SIMBOLI ........................................................................................ XIII 1 PREDSTAVITEV PROBLEMATIKE IN HIPOTEZE .........................................1 1.1 PAJKI KOT POMEMBNI RAZISKOVALNI OBJEKTI ....................................... 2 1.2 MREŽE PAJKOV KOT POMEMBNI RAZISKOVALNI OBJEKTI .................... 2 1.2.1 Vloga v raziskavah biomaterialov ................................................................ 3 1.3 ZGRADBA KOLESASTE MREŽE ........................................................................ 4 1.4 EVOLUCIJA KOLESASTE MREŽE ..................................................................... 7 1.5 DELOVANJE KOLESASTE MREŽE .................................................................. 10 1.5.1 Individualna variabilnost v mrežah ............................................................. 11 1.6 CILJI DOKTORSKE NALOGE............................................................................ 12 1.6.1 Koevolucija vedenja, ekologije in biomateriala .......................................... 12 1.6.2 Ontogenetske spremembe v arhitekturi mrež ............................................. 13 Gregorič M., Filogenetska sistematika in evolucija mrež pri pajkih mrežarjih...Zygiella...Caerostris. Dokt. disertacija. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, 2013 VI 1.6.3 Energijski vložek v plenjenje in vedenjska plastičnost ............................... 13 1.6.4 Filogenetska sistematika rodov Zygiella s.l. in Caerostris ......................... 14 2 ZNANSTVENA DELA ............................................................................................16 2.1 OBJAVLJENA ZNANSTVENA DELA ..............................................................
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