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Maksims Zolovs UNIVERSITY OF DAUGAVPILS INSTITUTE OF LIFE SCIENCES AND TECHNOLOGY DEPARTMENT OF BIOSYSTEMATICS MAKSIMS ZOLOVS IMPACT OF ENVIRONMENT FACTORS ON ECTOPARASITES DISTRIBUTION WITHIN FISH GILL APPARATUS: PREFERENCES, CO-OCCURRENCE AND INTERACTION APKĀRTĒJĀS VIDES FAKTORU IETEKME UZ EKTOPARAZĪTU IZPLATĪBU ZIVJU ŽAUNU APARĀTĀ: PREFERENCES, LĪDZĀSPASTĀVĒŠANA UN MIJIEDARBĪBA Doctoral Thesis in Biology for Obtaining the Doctoral Degree (branch: zoology) Supervisor: Dr. biol., Senior Researcher Voldemārs Spuņģis Daugavpils 2018 Department of Biosystematics, Institute of Life Sciences and technology, University of Daugavpils, Latvia Type of work: doctoral thesis (a set of publications) in biology, the branch of zoology. The thesis was performed at University of Daugavpils in 2013 – 2018 and was partly supported by the European Social Fund within the project Nr.2013/0016/1DP/1.1.1.2.0/APIA/VIAA/055. Supervisor: Dr. biol., Senior Researcher Voldemārs Spuņģis (University of Latvia, Latvia) Scientific adviser: PhD, Prof. Boris Krasnov (Ben-Gurion University of the Negev, Israel) Opponents: Dr. med. vet., Asoc. Prof. Dace Keidāne (Latvia University of Agriculture, Latvia) PhD, Assist. Prof. Grzegorz Zaleśny (Wrocław University of Environmental and Life Sciences, Poland) Dr. biol., Senior Researcher Maksims Balalaikins (Daugavpils University, Latvia) The head of the Promotion Council: Dr. biol., Prof. Arvīds Barševskis Commencement: Room 130, Parādes street 1A, University of Daugavpils, Daugavpils, on June 1, 2018, at 12:00. The Doctoral Thesis and it’s summary are available at the library of University of Daugavpils, Parādes street 1, Daugavpils and: http://du.lv/lv/zinatne/promocija/aizstavetie_promocijas_darbi/2018_gads. Comments are welcome. Send them to the secretary of the Promotion Council, Parādes street 1A, Daugavpils, LV-5401; mob. +37126002593; e-mail: [email protected] The secretary of the Promotion Council: Dr. biol. Jana Paidere, researcher of University of Daugavpils. © Maksims Zolovs, 2018 CONTENTS LIST OF ORIGINAL PAPERS........................................................................................ 4 1. INTRODUCTION..................................................................................................... 5 1.1 Actuality and novelty of the study.............................................................................. 5 1.2. Parasite preferences and interactions.......................................................................... 7 1.3. Morphological plasticity of monogeneans.................................................................. 9 1.4. Biotic and abiotic factors affecting parasite occurrence............................................. 10 2. MATERIAL AND METHODS............................................................................... 12 2.1. Preliminary investigation............................................................................................ 12 2.2. Examination of fish and collection of parasties.......................................................... 12 2.3. Morphometric analysis of monogeneans.................................................................... 14 2.4. Collecting of water samples and quality analysis....................................................... 15 2.5. Basic statistics............................................................................................................. 15 2.6. Calculation of niche breadth and overlap................................................................... 15 2.7. Statistical data analysis of parasites microhabitat preference and co-occurrence...... 16 2.8. Ethical note................................................................................................................. 17 3. RESULTS................................................................................................................... 18 3.1 Morphometric analysis of the hard parts of monogeneans......................................... 18 3.2. The distribution of digenean metacercariae within bream (Abramis brama) gill apparatus: preferences, co-occurrence and interactions of parasties.......................... 22 3.3. The effect of chemical water quality parameters on the spatial distribution of perch (Perca fluviatilis) ectoparasites................................................................................... 26 4. DISCUSSION............................................................................................................. 30 SUMMARY.......................................................................................................................... 35 ACKNOWLEDGEMENTS................................................................................................ 36 DISSERTATION IN LATVIAN........................................................................................ 37 REFERENCES.................................................................................................................... 70 ORIGINAL PAPERS.......................................................................................................... 84 3 LIST OF ORIGINAL PAPERS / PUBLIKĀCIJU SARAKSTS This thesis is based on the following papers, which are referred to in the text by their Roman numerals. Original papers are reproduced with permissions from the publishers. Promocijas darbs ir balstīts uz publikācijām, kuras disertācijas tekstā ir norādītas ar romiešu cipariem. Oriģinālie raksti ir publicēti ar izdevēju atļaujām. I Zolovs M., Priekule M., Gasperovich O., Kolesnikova J., Osipovs S., Spuņģis V. 2018. The spatial distribution of perch (Perca fluviatilis) ectoparasites and the effect of chemical water quality parameters on the ectoparasites’ spatial niche size. Proceedings of the Latvian Academy of Sciences (in press). II Zolovs M., Kanto J., Jakubāne I. 2017. The distribution of digenean metacercariae within bream (Abramis brama) gill apparatus: preferences, co-occurrence and interactions of parasites. Journal of Helminthology: 1 – 11. DOI: 10.1017/S0022149X1700044X III Zolovs M., Deksne G., Daukšte J., Aizups J., Kirjušina M. 2016. Morphometric analysis of the hard parts of Pseudodactylogyrus anguillae and Pseudodactylogyrus bini (Monogenea: Dactylogyridae) on the gill apparatus of the European eels (Anguilla anguilla) from the freshwaters of Latvia. Journal of Parasitology 102: 388 – 394. DOI: 10.1645/15-789 The author’s contribution (%) to the papers / Autora ieguldījums (%) pētījumos: I II III Original idea / Pētījuma ideja 60 80 80 Study design / Pētījuma dizains 90 90 90 Data collection / Datu ievākšana 50 65 65 Data analysis / Datu analīze 90 90 100 Manuscript preparation / Manuskripta sagatavošana 90 90 90 The order of the authors’ names reflects their involvement in the paper. Autora pieminēšanas kārtība publikācijā norāda viņa ieguldījumu pētījuma veikšanā. 4 1. INTRODUCTION 1.1. Actuality and novelty of the study Host–parasites relationships and relationships among parasites are the focus of many parasitological studies. Thanks to extensive research in this field, our understanding of those relationships is markedly progressed. Some topics are well studied from various sides. For example, parasites negative impact on host organism. However, several themes have only fragmental data or even gaps. The main goals of this thesis were to evaluate host-parasite relationships at the morphological level, to analyze the interaction among distantly related parasites species (e.g., species belonging to different major taxa) and to investigate the effect of chemical water quality parameters on ectoparasites spatial niche breadth. The theory of phenotypic plasticity suggests that a single genotype is capable to exhibit variable phenotype in different environments (Fusco & Minelli 2010). Parasites live in two environments. The first environment is a host that provides a place for parasites to live, forage and/or mate. The second environment is a habitat, in which a host organism lives (Dogel 1962; Kennedy 1975). Various studies found the relationship between water environment (the second environment) and morphology of attachment apparatus of parasites. However, the effect of microhabitats within a host (the first environment) on parasite attachment apparatus was not studied. The first aim of the study was to test whether the occurrence of parasites in different gill patches is associated with the size of parasite attachment apparatus (III). The length of gill filaments gradually increases along the dorsal third of gill arch and then gradually decreases to the ventral end. Moreover, the gill filament lamellae differ in shape and number both within the gill arch and filament (Hoar & Randall 1984). In light of this, we hypothesized that microenvironmental condition of patches within gill apparatus would affect the size of parasite attachment apparatus. Interactions among parasites have mainly been investigated in closely related species, as they exploit the same resources (El Hafidi et al. 1998; Šimková et al. 2000; Buchmann & Lindenstrøm 2002; Koskivaara et al. 2009). If distantly related parasites (e.g., species belonging to different major taxa) exploit similar resources they also can interact. However, such interactions have rarely been studied due to parasites aggregation among hosts where most hosts harbour a few parasite species and only some hosts species harbour large infracommunities (Shaw & Dobson 1995). The second aim of the study was to test whether digenean metacercariae a) prefer to encyst at specific patches
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