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Gastrotricha of Sweden – Biodiversity and Phylogeny

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Gastrotricha of Sweden – Biodiversity and Phylogeny Gastrotricha of Sweden – Biodiversity and Phylogeny Tobias Kånneby Department of Zoology Stockholm University 2011 Gastrotricha of Sweden – Biodiversity and Phylogeny Doctoral dissertation 2011 Tobias Kånneby Department of Zoology Stockholm University SE-106 91 Stockholm Sweden Department of Invertebrate Zoology Swedish Museum of Natural History PO Box 50007 SE-104 05 Stockholm Sweden [email protected] [email protected] ©Tobias Kånneby, Stockholm 2011 ISBN 978-91-7447-397-1 Cover Illustration: Therése Pettersson Printed in Sweden by US-AB, Stockholm 2011 Distributor: Department of Zoology, Stockholm University Till Mamma och Pappa ABSTRACT Gastrotricha are small aquatic invertebrates with approximately 770 known species. The group has a cosmopolitan distribution and is currently classified into two orders, Chaetonotida and Macrodasyida. The gastrotrich fauna of Sweden is poorly known: a couple of years ago only 29 species had been reported. In Paper I, III, and IV, 5 freshwater species new to science are described. In total 56 species have been recorded for the first time in Sweden during the course of this thesis. Common species with a cosmopolitan distribution, e. g. Chaetonotus hystrix and Lepidodermella squamata, as well as rarer species, e. g. Haltidytes crassus, Ichthydium diacanthum and Stylochaeta scirtetica, are reported. In Paper II molecular data is used to infer phylogenetic relationships within the morphologically very diverse marine family Thaumastodermatidae (Macrodasyida). Results give high support for monophyly of Thaumastodermatidae and also the subfamilies Diplodasyinae and Thaumastoder- matinae. In Paper III the hypothesis of cryptic speciation is tested in widely distributed freshwater gastrotrichs. Heterolepidoderma ocellatum f. sphagnophilum is raised to species under the name H. acidophilum n. sp. The results indicate that L. squamata may be a complex of at least two species. In Paper III and V the phylogeny of Chaetonotidae (Chaetonotida), the largest family within Gastrotricha, is inferred. The group suffers from a troubled taxonomy and is hypothesized to be non- monophyletic. Results show that members of Dasydytidae are nested within the group. Since only 3 of 17 sampled genera are monophyletic, it is hypothesized that the cuticular structures used in current classification do not reflect phylogenetic relationships. The phylogenetic hypothesis generated in Paper V indicates a marine origin of the predominantly limnic Chaetonotidae with a subsequent secondary invasion to marine environments of some taxa. Keywords: taxonomy, systematics, phylogeny, new species, cryptic species, Chaetonotidae, Dasydytidae, Thaumastodermatidae. Swedish Gastrotricha Tobias Kånneby CONTENTS AVHANDLINGEN I KORTHET .........................................................................................................2 LIST OF PAPERS .................................................................................................................................4 1. INTRODUCTION .............................................................................................................................5 1.1 Historical overview......................................................................................................................5 1.2 Morphology..................................................................................................................................8 1.2.1 Size and shape.......................................................................................................................8 1.2.2 Cuticle and body wall ...........................................................................................................9 1.2.3 Alimentary canal.................................................................................................................10 1.2.4 Nervous system and sense organs.......................................................................................10 1.2.5 Excretory organs .................................................................................................................11 1.2.6 Respiration and circulation .................................................................................................11 1.2.7 Reproduction and life cycle ................................................................................................11 1.3 Biogeography.............................................................................................................................13 1.3.1 Dispersal .............................................................................................................................13 1.4 Aims...........................................................................................................................................14 2. METHODS ......................................................................................................................................14 2.1 Collection and documentation ...................................................................................................14 2.2 Further treatment........................................................................................................................15 2.3 Molecular studies.......................................................................................................................15 3. SWEDISH FAUNA.........................................................................................................................16 3.1 Freshwater fauna........................................................................................................................16 3.2 Marine fauna ..............................................................................................................................17 4. GASTROTRICH RELATIONSHIPS..............................................................................................17 4.1 Gastrotrich interrelationships.....................................................................................................18 4.2 Cryptic species...........................................................................................................................19 5. FUTURE PERSPECTIVES.............................................................................................................20 6. SUMMARY OF PAPERS ...............................................................................................................21 7. LITERATURE CITED ....................................................................................................................23 8. ACKNOWLEDGEMENTS.............................................................................................................28 APPENDIX I – GASTROTRICHA REPORTED FROM SWEDEN.................................................30 APPENDIX II – REFERENCES FIGURE 1 AND FIGURE 2 ..........................................................34 Swedish Gastrotricha Tobias Kånneby AVHANDLINGEN I KORTHET Bukhårsdjur (Gastrotricha) är mycket små maskformiga ryggradslösa djur som lever i både sötvatten och marina miljöer. Gruppen utgör en del av det man brukar kalla meiofauna, ett samhälle av små djur som ofta lever mellan sandkorn och andra partiklar i akvatiska miljöer. I sötvatten kan bukhårsdjuren också leva på växter, ovanpå bottnen eller till och med i den fria vattenmassan. De lever av små organiska partiklar, bakterier eller alger och äts i sin tur av lite större ryggradslösa djur. Bukhårsdjur kännetecknas av att de har små hår, så kallade cilier, på undersidan som de använder för att ta sig fram. De har också häftrör med vilka de kan klistra fast sig mot underlaget om något skulle störa dem. Djuren är relativt vanliga men kan oftast inte ses med blotta ögat, de minsta bukhårsdjuren är bara en tiondels millimeter långa. Därför behöver man mikroskop för att dokumentera och artbestämma dem. Hittills har man hittat nästan 770 arter i hela världen men det finns fortfarande många delar av jordklotet som inte är undersökta. Bukhårsdjur delas in i två grupper. Den ena gruppen (Macrodasyida) innehåller nästan uteslutande marina avlånga maskar med häftrör på bakkroppen, på sidorna och kanske även på ryggen och i framändan. De är hermafroditer, samma individ är samtidigt både hane och hona eller så alternerar individen mellan hona och hane. Den andra gruppen (Chaetonotida) innehåller både marina och sötvattenslevande maskar som med få undantag har en kägelformad kropp. Hos denna grupp finns häftrören oftast bara på bakkroppen och sitter på en gaffelliknande struktur som kallas furka. Många arter inom gruppen förökar sig genom jungfrufödsel, vilket innebär att avkomman utvecklas ur obefruktade ägg, men hermafroditer förekommer också hos marina arter. I Sverige är bukhårsdjur en dåligt studerad grupp och 2007 kände man bara till 22 marina och 7 sötvattenslevande arter. Det enklaste sättet att samla in bukhårsdjur är att ta upp bottenmaterial, växter eller sand med en mycket finmaskig håv eller för hand. Proverna letas därefter igenom under en stereolupp och djuren kan plockas ut. Oftast måste man söva djuren med magnesiumklorid, som sätter nervsystemet ur spel, och hindrar dem från att häfta fast vid olika ytor. För att identifiera djuret måste det studeras levande under ett speciellt mikroskop, ett så kallat differential interferens kontrast mikroskop (DIC) som ökar kontrasten på de viktiga strukturerna hos det mer eller mindre transparenta djuret. När djuret är fotograferat och väl dokumenterat kan
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