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MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE DISERTAČNÍ PRÁCE Brno 2018 Martina Hrušková-Martišová 1 MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE EKOLOGIE A ETOLOGIE SOLIFUG (ARACHNIDA: SOLIFUGAE) Disertační práce Martina Hrušková-Martišová Vedoucí práce: prof. Mgr. Stanislav Pekár, PhD. Brno 2018 2 Bibliografický záznam Autor: Mgr. Martina Hrušková-Martišová Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Ekologie a etologie solifug (Arachnida: Solifugae) Studijní program: Biologie Studijní obor: Ekologie Vedoucí práce: prof. Mgr. Stanislav Pekár, PhD. Akademický rok: 2017/2018 Počet stran: 75+52 Klíčová slova: Pavoukovci; Habitat; Aktivita; Kořist; Reprodukce; Námluvy; Násilná kopulace; Sexuální kanibalismus; Pohlavní dimorfismus; Defenzivní chování; Obranné strategie; Stridulační orgán 3 Bibliographic Entry Author: Mgr. Martina Hrušková-Martišová Faculty of Science, Masaryk University Department of Botany and Zoology Title of Thesis: Ecology and behaviour of solifuges (Arachnida: Solifugae) Degree programme: Biology Field of Study: Ecology Supervisor: prof. Mgr. Stanislav Pekár, PhD. Academic Year: 2017/2018 Number of Pages: 75+52 Keywords: Arachnida; Camel-spiders; Habitat; Activity; Prey; Reproduction; Courtship; Forced copulation; Sexual cannibalism; Sexual dimorphism; Defensive behaviour; Defensive strategies; Stridulatory organ 4 Abstrakt Solifugy (Arachnida: Solifugae) představují jednu z nejdůležitějších složek fauny většiny aridních a semiaridních oblastí, přesto existuje jen velmi málo studií zabývajících se různými aspekty biologie těchto pozoruhodných pavoukovců. Cílem mé práce bylo detailně prozkoumat biologii dvou druhů solifug, středoasijské solifugy Galeodes caspius subfuscus Birula 1937 (Galeodidae) a iberijské Gluvia dorsalis (Latreille 1817) (Daesidae), a vypracovat aktualizovaný přehled poznatků o solifugách zaměřený především na ekologii a etologii, založený na výsledcích vlastních pozorování a shrnutí dostupných informací. Zjistila jsem, že oba druhy G. caspius a G. dorsalis se nejčastěji vyskytují na otevřených písčitých plochách s řídkým nebo nízkým vegetačním pokryvem. Jsou noční, sezónní aktivita je u obou druhů ovlivněna teplotou a množstvím srážek. Výsledky mé studie ukázaly, že G. dorsalis je biennální druh, se 7-9 juvenilními stádii a délkou života minimálně 2 roky. Oba druhy jsou vysoce žraví generalističní predátoři, lovící především různé druhy členovců. Juvenilní jedinci a dospělci mohou využívat odlišné predační strategie. Dále jsem ve své práci zkoumala stridulaci solifugy G. caspius. Popsala jsem stridulační orgán a nahrála a analyzovala produkovaný zvuk. Provedla jsem komparativní analýzu stridulačního orgánu a stridulace během ontogeneze. Pomocí akustické analýzy jsem zjistila primárně defenzivní funkci stridulace. Detailně jsem studovala reprodukční chování a zjistila jsem, že páření G. caspius a G. dorsalis se výrazně liší. Nicméně u obou druhů několik zásadních behaviorálních charakteristik jasně potvrdilo násilný charakter kopulace. U druhu G. caspius jsem navíc odhalila rysy typické pro námluvy a dokázala jsem tak, že tyto dvě navzájem se vylučující strategie se mohou vyskytovat současně v rámci jedné kopulace. Zatímco násilná kopulace se vyvinula za účelem snížení rizika sexuálního kanibalismu, námluvy mohou reprezentovat komplementární strategii samců s cílem zvýšit reprodukční úspěch a šanci na paternitu. 5 Abstract Solifuges (Arachnida: Solifugae) belong to the most important arthropods especially in arid and semi-arid environments. Unfortunately, only little data are available on biology of these fascinating arachnids. In my thesis I aimed to study two different species: Central Asian Galeodes caspiius subfuscus Birula 1937 (Galeodidae) and Iberian Gluvia dorsalis (Latreille 1817) (Daesidae), and provide an updated overview of solifuge biology, particularly their ecology and behaviour, based on my observations and available published information. I found that both G. caspius and G. dorsalis prefer open sandy areas with low or sparce vegetation cover and are strictly nocturnal in their activity patterns. The seasonal activity is strongly influenced by temperature and humidity. My results showed that the G. dorsalis is a biennal species, with 7-9 juvenile instars and longevity of about 2 years. Both species can be characterised as generalist voracious predators, feeding on various arthropods. Foraging strategies may differ between juveniles and adults. Further, the stridulation of G. caspius was examined. I described the stridulatory organ and recorded and analyzed produced sound. Then I performed comparative analysis of both stridulatory organ and stridulation during the onthogenesis. Acoustical analysis suggested that the stridulation has a primarily defensive role. I studied mating behaviour of G. caspius and G. dorsalis in detail. I found that mating differed markedly between the two species, however, several behavioural features demontrated coercive copulation in both of them. In the coercively mating G. caspius I revealed presence of luring traits too thus I proved that these two mutually exlusive strategies may coexist during one copulation. Coercive mating may have evolved to reduce risk of sexual cannibalism, luring behaviour may represent complementary strategy to increase mating and paternity succes of the male. 6 © Martina Hrušková-Martišová, Masarykova Univerzita, 2018 7 Poděkování Tato dizertační práce by nikdy nevznikla, nebýt mnoha lidí, kterým bych proto ráda touto cestou poděkovala. Na prvním místě děkuji především mému školiteli, prof. Mgr. Stano Pekárovi, PhD., který se mě i mého nápadu studovat solifugy ujal, a otevřel mi tak cestu za poznáním těchto úžasných a tajemných tvorů. Po celou dobu mého studia mi byl trpělivým průvodcem, pomocníkem, rádcem i podnětným kritikem. Děkuji mu také za finanční podporu, kterou mi poskytoval, a která mi umožnila podniknout několik cest do oblastí, kde se tito tvorové vyskytují. Dále bych ráda poděkovala panu prof. RNDr. Jaroslavu Smržovi, CSc., který ve mě během svých přednášek probudil fascinaci těmito tvory a jako první mě tak přivedl k myšlence zkoumat solifugy. Děkuji Dr. Alexandru V. Gromovovi za vřelé přijetí i za zázemí, které nám poskytl při terénním výzkumu v Kazachstánu. Bez jeho pomoci, znalostí a kontaktů by výzkum solifug v této zemi nebyl tak úspěšný, a pravděpodobně by ani nebyl možný. Poděkování patří i doc. RNDr. Jiřímu Královi, CSc. a Dr. Pedru Cardoso, PhD., díky kterým byla práce v terénnu ještě zajímavější. V neposlední řadě děkuji členům mé rodiny za podporu a pomoc především v závěru mého studia a mým dětem Tereze, Matoušovi a Jáchymovi za pochopení a trpělivost při sepisování této práce. Výzkum byl podporován projektem NATO-Science project č. LST.CLG.980430: Biology of solifuges of central Asia, granty č. 1P05ME777, 1P06ME851 a 0021622416 MŠMT ČR, projektem č. 20061431C008 MU a projektem č. SFRH/BD/1195/200 FCT. 8 Prohlášení Prohlašuji, že jsem svoji disertační práci vypracovala samostatně s využitím informačních zdrojů, které jsou v práci citovány. Brno 7. března 2018 ……………………………… Martina Hrušková-Martišová 9 OBSAH I. PŘEDMLUVA … 12 II. ÚVOD … 14 1. SOLIFUGY - OBECNÁ CHARAKTERISTIKA … 14 2. SPECIFICKÉ MORFOLOGICKÉ RYSY SOLIFUG … 15 3. POPIS STUDOVANÝCH DRUHŮ … 20 4. EKOLOGIE SOLIFUG … 21 4.1 Geografické rozšíření … 21 4.2 Biotop … 21 4.3 Potravní vztahy … 23 4.4 Cirkadiální aktivita … 25 4.5 Sezónní výskyt … 26 4.6 Životní cyklus … 27 5. ETOLOGIE SOLIFUG … 30 5.1 Predační chování … 30 5.2 Defenzivní chování a obranné strategie … 32 5.3 Reprodukční chování … 37 5.4 Ovipozice a péče o potomstvo … 45 III. CÍLE PRÁCE … 46 IV. SHRNUTÍ JEDNOTLIVÝCH RUKOPISŮ … 47 1. Rukopis A … 47 2. Rukopis B … 48 3. Rukopis C … 49 4. Rukopis D … 51 V. ZÁVĚR … 52 VI. SEZNAM POUŽITÉ LITERATURY … 55 VII. PŘÍLOHY … 76 10 „Žije zde podivný tvor, nestvůrné zjevení. Jak zvláštního vzhledu, jak divných proporcí, jak zle vyhlížející a odpudivé svým ohyzdným tvarem a pokryvem dlouhých, zježených chlupů! Možná je to jakási předpotopní forma života, jež byla zachována výhradně pro toto nepřívětivé místo. Zdá se, jako by bylo přímo stvořeno pro tuto neúrodnou zemi, vyprahlou a nevlídnou pustinu. Jen zde můžeme nalézt něco tak neobvyklého a nepřirozeného; zdá se, že pro obyčejný život není v této vyschlé písečné divočině místo.“ (Hingston, R. W. G. 1925. Nature at the Desert's Edge.) 11 I. PŘEDMLUVA Solifugy byly a dosud jsou předmětem mnoha rozličných studií. Navzdory tomu představují jednu z nejméně probádaných skupin pavoukovců. Většina informací, které máme, se totiž týká jen relativně málo druhů především z oblastí Severní Ameriky a Afriky, a často jen dospělých vývojových stádií. Zatímco morfologie a fyziologie byla v rámci této skupiny zkoumána velmi podrobně, o ekologických a behaviorálních aspektech jejich života víme velmi málo. Předkládaná dizertační práce shrnuje výsledky výzkumu vybraných ekologických a etologických charakteristik dvou druhů solifug, iberské solifugy Gluvia dorsalis (Daesidae) a středoasijské solifugy Galeodes caspius subfuscus (Galeodidae). Dizertační práce se skládá z literární rešerše a čtyř vědeckých publikací, které jsou založeny na výsledcích tří nezávislých studií. Dvě práce (A, C) se podrobně zabývají biologií těchto dvou druhů solifug, jedna práce (B) se věnuje stridulaci u druhu Galeodes caspius a třetí práce (D) popisuje reprodukční chování solifug a násilnou kopulaci. Všechny rukopisy jsem vypracovala
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  • Changes in the Numbers of Chromosomes and Sex Determination System in Bushcrickets of the Genus Odontura (Orthoptera: Tettigoniidae: Phaneropterinae)

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  • T.C. Harran Üniversitesi Fen Bilimleri Enstitüsü

    T.C. Harran Üniversitesi Fen Bilimleri Enstitüsü

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