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Masarykova Univerzita Dizertačná Práca MASARYKOVA UNIVERZITA Přírodovědecká fakulta Ústav botaniky a zoologie DIZERTAČNÁ PRÁCA Štúdium biosystematiky a fylogenézy stredoeurópskych druhov čeľade Ceratopogonidae (Diptera) s využitím morfologických i molekulárno-biologických znakov Brno, 2006 Andrea Tóthová MASARYKOVA UNIVERZITA Přírodovědecká fakulta Andrea TÓTHOVÁ Štúdium biosystematiky a fylogenézy stredoeurópskych druhov čeľade Ceratopogonidae (Diptera) s využitím morfologických i molekulárno-biologických znakov Dizertačná práca Školitel: Prof. RNDr. Jaromír Vaňhara, CSc. Brno, 2006 Bibliografická identifikácia Meno a priezvisko autora: RNDr. Andrea Tóthová Názov dizertačnej práce: Štúdium biosystematiky a fylogenézy stredoeurópskych druhov čeľade Ceratopogonidae (Diptera) s využitím morfologických i molekulárno-biologických znakov Názov dizertačnej práce anglicky: Biosystematics and phylogeny of the family Ceratopogonidae (Diptera) of central Europe based on morphological and molecular –biological characters Študijný program: Biologie Študijný obor (smer), kombinácia oborov: 1501V - Zoologie Školiteľ: Prof. RNDr. Jaromír Vaňhara, CSc. Rok obhajoby: 2006 Kľúčové slová: Ceratopogonidae, Diptera, systematika, fylogenéza, 16S, génové markery, taxonómia Kľúčové slová v angličtine: Ceratopogonidae, Diptera, systematics, phylogeny, 16S, gene markers, taxonomy © Andrea Tóthová, Masarykova univerzita, 2006 Poďakovanie Dizertačná práca bola vypracovaná v rámci interného doktorského štúdia na Ústave botaniky a zoologie Přírodovědecké fakulty Masarykovy Univerzity v Brne. Práca bola finančne podporená grantmi FRVŠ 1237/2005 „Využití molekulárně-biologických metod pro studium systematiky a fylogenetických vztahů pakomárců (Diptera, Ceratopogonidae)“ (riešiteľ A. Tóthová), GAČR 524/05/H536 (hlavný riešiteľ M. Chytrý) a výskumnými zámermi „Časoprostorová dynamika biodiverzity v ekosystémech střední Evropy“ (MSM 143100010, hlavný riešiteľ J. Vaňhara) a „Diverzita biotických společenstev a populací: kauzální analýza variability v prostoru a čase“ (MSM 0021622416, hlavný riešiteľ M. Chytrý). Na tomto mieste by som rada poďakovala svojmu školiteľovi prof. Jaromírovi Vaňharovi za jeho skutočne všestrannú pomoc a podporu pri riešení ako problematiky tak i problémov. Za skvelú podporu a prínos poznatkov o morfológii a systematike pakomárikov patrí moja vďaka konzultantovi prof. Janovi Knozovi. Ďalej by som chcela poďakovať Pepovi Bryjovi za zasvätenie do tajov laboratórnej práce, spoluprácu pri spracovávaní vzoriek a revíziu rukopisov, ako i osadenstvu laboratória bývalej Katedry botaniky Jakubovi Šmerdovi a Lucke Horovej za nenahraditeľné a cenné rady „z praxe molekulárnika“. Moja vďaka patrí samozrejme všetkým, ktorí prispeli k úspešnosti mojej práce, či už radami, alebo tvrdou prácou (Honza Mendel, osadenstvo laboratória v Studenci, Andrea Šimková, Peťo Mikulíček...) V neposledom rade patrí obrovská vďaka mojim rodičom a priateľovi Radimovi za ich skvelú a všestrannú podporu. 4 Obsah Abstrakt 6 Abstract 7 Úvod 8 1.1. Morfológia čeľade 9 1.2. Determinácia 10 2. Materiál a metodika 2.1. Metódy zberu 10 2.2. Aplikácia génových analýz v štúdiu dvojkrídleho hmyzu 11 Ciele dizertačnej práce 13 3. Súhrn výsledkov 14 4. Diskusia a záver 21 Literatúra 23 Príloha 1 (vložené publikácie) 28 Článok A 29 Článok B 36 Článok C 42 Článok D 50 Článok E 79 Článok F 107 Článok G 118 Príloha 2 (Curriculum vitae) 138 Príloha 3 (zoznam publikácií) 140 5 Abstrakt Abstrakt ŠTÚDIUM BIOSYSTEMATIKY A FYLOGENÉZY STREDOEURÓPSKYCH DRUHOV ČEĽADE CERATOPOGONIDAE (DIPTERA) S VYUŽITÍM MORFOLOGICKÝCH I MOLEKULÁRNO- BIOLOGICKÝCH ZNAKOV Andrea Tóthová Čeľaď Ceratopogonidae zahŕňa dvojkrídly hmyz, ktorý je náročné správne determinovať do úrovne druhu. Predložená práca je prierezom štúdií založených na klasických morfologických, ale i molekulárno-biologických metódach, snažiacich sa objasniť vybrané taxonomické problémy a zhodnotiť fylogenetické vzťahy v rámci čeľade. Zástupcovia čeľade sa všeobecne zbierajú pomocou pascí svetelných, prípadne pascí založených na CO2 či vnadených nulliparných samičkách. Predložené práce zaoberajúce sa druhovým zložením sú založené na zbere materiálu nesvetelnými pascami. Zber materiálu pakomárikov prebiehal v NP Podyjí v rámci komplexného monitoring čeľadí dvojkrídlovcov. Nový zoznam druhov pakomárikov Českej a Slovenskej republiky bol od roku 1997 obohatený o 33 nových druhov. Počas monitoringu fauny pakomárikov bolo zistených niekoľko taxonomických nejasností v podrode Meloehelea rodu Atrichopogon. Podľa v súčasnosti jediného determinačného kľúča bolo niekoľko jedincov identifikovaných ako severoamerický druh Atrichopogon epicautae. Pre porovnanie bol zapožičaný typový materiál tohto druhu z USA a ďalšie jedince Kanady. Po dôkladnom zhodnotení všetkých morfologických znakov boli zistené zaujímavé skutočnosti (rozdiely medzi kanadským a americkými jedincami, existujúce samce patria do podrodu Atrichopogon, atd.). Pre objasnenie taxonomie v podrode Meloehelea i v problematickom rode Dasyhelea bola využitá analýza sekvencií mitochodriálneho génu 16S DNA. U jedincov podrodu A. (M.) aff. epicautae bola preukázaná totožnosť s európskym druhom A. (M.) lucorum, čo potvrdilo variabilitu kľúčových determinačných znakov a ich nevhodnosť pre rozlíšenie týchto dvoch druhov. Analýza sekvencií génu 16S potvrdila validitu 3 sporných druhov rodu Dasyhelea s obrovskou variabilitou determinačných znakov (sfarbenie nôh a tela, tvar postgenitálnej plôšky samíc).Bola potvrdená monofýlia celej čeľade a intergenerické vzťahy plne podporujúce súčasnú systematiku založenú na morfologických znakoch. Vzťah rodov a tribov odvodený z mitochondriálneho génu 16S je veľmi podobný tomu, ktorý bol odvodený inými autormi z génu cox2. Obe tieto analýzy sa zhodujú v rozdielnom pohľade na monofyletický pôvod podčeľade Ceratopogoninae odvodený z kladistických štúdií rôznych autorov. Ich závery neboli potvrdené žiadnou z molekulárnych analýz. 6 Abstract Abstract BIOSYSTEMATICS AND PHYLOGENY OF THE FAMILY CERATOPOGONIDAE (DIPTERA) OF CENTRAL EUROPE BASED ON MORPHOLOGICAL AND MOLECULAR –BIOLOGICAL CHARACTERS The Biting Midges (Diptera: Ceratopogonidae) are small flies which are often difficult to identify, especially at the species level. The current study reviews a cross-section of faunistic and taxonomic studies based on classic morphological characters as well as more modern, molecular-based methods in order to solve specific taxonomic problems and reconstruct phylogenetic relationships within the family. Biting midges are usually collected with various types of light traps, CO2 traps or traps with nulliparous females. Some studies focused on monitoring of species were made with non-light traps, such as Malaise traps. In the present study, collections were made in the Podyjí NP, Czech Republic in order to obtain biomonitoring data of these flies. A revised checklist of Ceratopogonidae of the Czech Republic and Slovakia is provided, including 33 species which are recorded in this area for the first time. During this faunistic study, some taxonomic problems were discovered within the subgenus Meloehelea of Atrichopogon. Using the available morphogically based keys; several specimens were identified within the North American Atrichopogon epicautae complex. We borrowed the type-series of A. epicautae as well as examples of several additional species from the USA and Canada and discovered some differences between the American and Canadian specimens. In an attempt to clarify this taxonomic situation in Atrichopogon (Meloehelea) and the genus Dasyhelea, an analysis of the 16S region of mitchondrial DNA (mtDNA) was performed on selected species of these two groups. Our results indicated that putative specimens of the European A. (M.) epicautae, were in fact, the Palaearctic species, A. (M.) lucorum. Therefore, the extreme variability of previously defined morphological characters used to distinguish species of A. (Meloehelea) was supported by our mtDNA results. However, our 16S analysis supported the validity of three species of Dasyhelea that we originally considered to be conspecific, the monophyly of the family and the intergeneric relationships previously based on morphology. The relation of the genera and tribes is very similar to that inferred from the cox2 gene; however, both of these analyses are consistent in their differences of the subfamilies based on cladistic analyses - the monophyly of the subfamily Ceratopogoninae was not supported by any of the molecular analyses. 7 Úvod ÚVOD Čeľaď pakomárikovitých (Ceratopogonidae) patrí do radu dvojkrídlovcov (Diptera) a zahŕňa 5155 druhov známych zo sveta (BORKENT & WIRTH 1997), 2970 druhov z palearktického regiónu (REMM 1988) a 588 druhov z Európy (SADZIEWSKI & BORKENT 2005). Zo systematického hľadiska sa čeľaď rozdeľuje na 5 podčeľadí (Leptoconopinae, Austroconopinae, Ceratopogoninae, Dasyheleinae a Forcipomyiinae), hoci ZILÁHI-SEBESS (1960) i KRIVOSHEINA (1962) sa snažili povýšiť podčeľaď Leptoconopinae na čeľaď. Čeľad je v úzkom príbuzenstve s čeľaďou Chironomidae, od ktorej sa ale pakomáriky odlišujú prítomnosťou bodavo-cicavého ústneho ústroja , výrazne kratšími prednými nohami a krídelnou žilnatinou. Larvy pakomárikovitých sa vyvíjajú v najrôznejších typoch habitatov od vody a vlhkých brehov stojatých vôd, cez rozkladajúcu sa opadanku, machy a hríby až po vtáčie hniezda. Spoločným ukazovateľom pre ne je istý stupeň vhlkosti, pri jej nedostatku larvy hynú. Dospelé jedince pakomárikov je možné nájsť v blízkosti ich liahnisk. Vo väčšine prípadov sa jedná o druhy so súmračnou
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