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Interactions Between Liometopum Microcephalum (Formicidae) and Other Dominant Ant Species of Sympatric Occurrence

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Interactions Between Liometopum Microcephalum (Formicidae) and Other Dominant Ant Species of Sympatric Occurrence MASARYK UNIVERSITY FACULTY OF SCIENCE DEPARTMENT OF BOTANY AND ZOOLOGY Populations of the ant Liometopum microcephalum (Panzer, 1798) at different spatial scales Ph.D. Dissertation Lenka Petráková Supervisor: Doc. Jiří Schlaghamerský, Ph.D. Brno 2016 Bibliographic Entry Author Mgr. Lenka Petráková Faculty of Science, Masaryk University Department of Botany and Zoology Title of Thesis: Populations of Liometopum microcephalum (Panzer, 1798) at different spatial scales Degree programme: Biology Field of Study: Zoology Supervisor: Doc. Jiří Schlaghamerský, Ph.D. Academic Year: 2015/2016 Number of Pages: 166 Keywords: Liometopum microcephalum, ant colonies, competitors, territories, worker size polymorphism, trophobiosis, dispersal, genetic diversity, South Moravian population, distribution range, lineage divergence, Pleistocene refugia Bibliografický záznam Autor: Mgr. Lenka Petráková Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Populace mravence Liometopum microcephalum (Panzer, 1798) na různých prostorových škálách Studijní program: Biologie Studijní obor: Zoologie Vedoucí práce: Doc. Jiří Schlaghamerský, Ph.D. Akademický rok: 2015/2016 Počet stran: 166 Klíčová slova: Liometopum microcephalum, mravenčí kolonie, konkurenti, teritoria, velikostní polymorfismus dělnic, trofobioza, šíření, genetická diverzita, jihomoravská populace, areál druhu, rozdílnost linií, pleistocenní refugia Abstract The ant Liometopum microcephalum is scattered from Italy in the west to the Lower Volga River in Russia and western Iran in the east, from South Moravia in the Czech Republic in the north to northern Israel in the south. The studied species is the only representative of the genus Liometopum occurring in Europe, albeit 14 fossil species of this genus have been found here. Despite the fact that this arboricolous species is locally ecologically important and its colonies consist of hundreds of thousands of workers, several questions regarding its biology remain unknown. The present thesis includes studies on L. microcephalum conducted at three spatial scales: local situations of one or several neighbouring colonies, the South Moravian population and the entire distribution area. Individual colonies within the South Moravian population were studied in regard to territory size and shape, interactions with competitors, trophobiosis with aphids, and morphological and genetic variability. The morphological and genetic variability was compared among colonies from different sites in South Moravia. The relatedness among colonies and populations was further assessed across the species’ distribution range to explore the phylogeography of the species. The thesis includes three scientific papers, which were published in ISI-indexed journals, and one submitted manuscript, which has already undergone a first review. The first paper deals with the position of the species in the hierarchy of ant communities and with its interactions with two other territorial ants, Lasius fuliginosus and Formica rufa, occurring in the same habitats. Liometopum microcephalum was able to withstand the competitive pressure of these competitors. Aggression was observed particularly in the beginning of the vegetation season, when the ants fought about foraging trees. Later in the season, conflicts occurred only when workers of different species were experimentally transferred into a competitor’s territory. Otherwise the competitors avoided each other. When defending its territories, L. microcephalum profited from the cooperation of its workers, which were biting the body of their rivals with their strong mandibles. The aim of the second study was to clarify the extent of morphological variability within and among L. microcephalum colonies. In all fifteen studied colonies a broad range of worker sizes was recorded. Variability was continuous and there were no distinct morphs. Several degrees of polymorphism were found in the studied colonies. These degrees did not correlate with the measured variables, but often varied in the course of time. Workers sampled in spring were significantly bigger than the workers collected in summer. There was a positive relationship between worker sizes and the colonies’ territory areas. In the third study the importance of honeydew intake in the studied species was evaluated by measuring the amount of total and reducing sugars in ant gasters of workers collected in different situations. The highest amount of sugars occurred in workers descending from foraging trees. This confirmed that the ants visit those trees to tend aphids. The amount of sugars in the gasters increased from April to June, but decreased rapidly in July. That is apparently related to the aphids’ life cycles and to the increased need of protein-rich food. In the fourth study the relatedness of colonies sampled at 40 sites across the distribution range was determined. In the studied colonies, 36 mitochondrial haplotypes were found. The most recent common ancestor of all studied colonies lived almost four million years ago. The Levantine population differed distinctly from the rest of the samples, having evolved separately since the Pliocene. The species probably dispersed from Anatolia in north-western direction. The origin of the European clades is related to the Pleistocene climate oscillations. The species survived glaciations in traditional Mediterranean refugia (the Balkan and Apennine Peninsulas) but also within the Carpathian Arc, at the Black Sea coast and at the south-eastern margin of the Alps. Abstrakt Mravenec lužní (Liometopum microcephalum) je ostrůvkovitě rozšířený od Itálie na západě po dolní tok Volhy v Rusku a západní Írán na východě, od Jižní Moravy na severu po severní Izrael na jihu. Studovaný druh je jediným zástupcem svého rodu v celé Evropě, ačkoli zde bylo nalezeno 14 fosilních druhů tohoto rodu. Přestože je tento arborikolní druh lokálně ekologicky významný a jeho kolonie čítají až stovky tisíc dělnic, různé aspekty jeho biologie jsou stále neznámé. Práce zahrnuje studie druhu L. microcephalum provedené na třech prostorových měřítkách: lokální situace u jedné nebo několika sousedících kolonií, v jihomoravské populaci a v rámci celého areálu druhu. U jednotlivých kolonií uvnitř jihomoravské populace byla studována velikost a tvar teritorií, vztahy s konkurenty, trofobioza se mšicemi, morfologická a genetická variabilita. Morfologická a genetická variabilita byla porovnávána mezi koloniemi z různých stanovišť na jižní Moravě. Příbuznost kolonií a populací pak byla sledována napříč areálem druhu s cílem objasnit fylogeografii druhu. Součástí práce jsou tři vědecké články publikované v časopisech s impakt faktorem (WoS) a jeden rukopis zaslaný do redakce, který již prošel první recenzí. První studie se zabývá pozicí druhu v hierarchii mravenčích společenstev a jeho vztahy ke dvěma dalším teritoriálním druhům, Lasius fuliginosus a Formica rufa, které obývají stejný habitat. Liometopum microcephalum dokázal obstát v konkurenci těchto soupeřů. Agresivní chování bylo pozorováno zejména na počátku vegetační sezóny, kdy docházelo k bojům o potravní stromy. Později ke střetům docházelo pouze tehdy, pokud byly jedinci jiného druhu experimentálně přeneseni do konkurentova teritoria. Jinak se všechny studované druhy svým konkurentům vyhýbaly. Při obraně svých teritorií L. microcephalum těžil ze spolupráce mezi dělnicemi, které vetřelce napadaly svými silných kusadly. Druhá studie měla za cíl objasnit míru morfologické variability uvnitř a mezi koloniemi mravence lužního. U všech patnácti studovaných kolonií byla zjištěna široká škála velikostí dělnic. Proměnlivost byla plynulá, bez jasně vymezených morf. U jednotlivých kolonií byly zjištěny různé stupně polymorfismu. Tyto stupně nesouvisely s žádnou z měřených proměnných, měnily se však v čase. Dělnice vzorkované na jaře byly významně větší než dělnice sbírané v létě. Byla potvrzena pozitivní korelace mezi velikostí dělnic a rozlohou teritoria kolonie. Ve třetí studii byl hodnocen význam příjmu medovice u mravence lužního pomocí množství celkových a redukujících cukrů v gasterech dělnic sbíraných v různých situacích. Nejvyšší obsah cukrů byl naměřen u dělnic sestupujících z potravních stromů. Tím se potvrdilo, že mravenci tyto stromy navštěvují za účelem dojení mšic. Obsah zjištěných cukrů narůstal od dubna do června, v červenci však rapidně poklesl. To patrně souvisí s generačními cykly mšic a zvýšenou potřebou proteinů v potravě. Ve čtvrté studii byla zjišťována příbuznost kolonií ze 40 lokalit napříč areálem druhu. U studovaných kolonií bylo nalezeno celkem 36 různých mitochondriálních haplotypů. Nejbližší společný předek všech studovaných kolonií žil před téměř čtyřmi miliony lety. Populace z Levanty se zřetelně odlišovala od ostatních vzorků, vyvíjela se samostatně od Pliocénu. Druh se pravděpodobně šířil z Anatolie severozápadním směrem. Původ evropských kládů souvisí s pleistocenními oscilacemi klimatu. Druh přežíval glaciály v tradičních mediteránních refugiích (Balkánský a Apeninský poloostrov), ale i v Karpatském oblouku, na pobřeží Černého moře a pod jihovýchodním okrajem Alp. © Lenka Petráková, Masaryk University, 2016 Acknowledgements I would like to thank all those people, who have trusted me and believed that my work had sense. My special thanks go to my supervisor Jiří Schlaghamerský for his ideas, advice, comments, and especially for his patience with corrections of my texts written in English. I´d like to thank him also for the opportunity to study this remarkable
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