PARASITIC ARTHROPODS: A COMPARATIVE ZOOLOGICAL-PALAEONTOLOGICAL STUDY Dissertation der Fakultät für Biologie der Ludwig-Maximilians-Universität München zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) Vorgelegt von M. Sc. Christina Nagler München, 2017 Für meine Eltern Diese Dissertation wurde angefertigt unter der Leitung von Prof. Dr. Matthias J. Starck & Dr. Joachim T. Haug im Bereich Zoologie an der Ludwig-Maximilians-Universität München Erstgutachter: Prof. Dr. Matthias J. Starck Zweitgutachter: Prof. Dr. Roland R. Melzer Tag der Abgabe: 22.05.2017 Tag der mündlichen Prüfung: 09.08.2017 ii EIDESSTATTLICHE ERKLÄRUNG Ich versichere hiermit an Eides statt, dass die vorgelegte Dissertation von mir selbstständig und ohne unerlaubte Hilfe angefertigt ist. Ort, Datum Christina Nagler ERKLÄRUNG Hiermit erkläre ich, dass diese Dissertation nicht einer anderen Prüfungskommission vorgelegt worden ist und dass ich mich nicht anderweitig einer Doktorprüfung ohne Erfolg unterzogen habe. Ort, Datum Christina Nagler iii LIST OF PUBLICATIONS INCORPORATED IN THIS DISSERTATION I. Nagler C, Haug C, Resch U, Kriwet J, Haug JT. 2016. 150 million years old isopods on fishes: a possible case of palaeo-parasitism. Bulletin of Geosciences, 91: 1–12. DOI: 10.3140/bull.geosci.1586. II. Nagler C, Haug JT. 2016. Functional morphology of parasitic isopods: understanding morphological adaptations of attachment and feeding structures in Nerocila as a pre-requisite for reconstructing the evolution of Cymothoidae. PeerJ, 4: e2188. DOI: 10.7717/peerj.2188 III. Serrano-Sánchez ML, Nagler C, Haug C, Haug JT, Centeno-García E, Vega FJ. 2016. The first fossil record of larval stages of parasitic isopods: cryptoniscus larvae preserved in Miocene amber. Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen, 279: 97–106. DOI: 10.1127/njgpa/2016/0543. IV. Nagler C, Hyžný M., Haug JT. 2017a. 168 million years old ―marine mallophagan‖ and the evolution of parasitism within isopods. BMC Evolutionary Biology, 17: 76. DOI: 10.1186/s12862-017-0915-1. V. Nagler C, Haug JT, Glenner H, Buckeridge J. 2017b. Litholepas klausreschi gen. et sp. nov., a new neolepadine barnacle (Cirripedia, Thoracica) on a sponge from the Upper Jurassic lithographic limestone‘s of southern Germany. Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen, 284: 29–42. DOI: 10.1127/njgpa/2017/0648 VI. Nagler C, Høeg J, Haug C, Haug JT. 2017c. A possible 150 million years old cirripede crustacean nauplius and the phenomenon of giant larvae. Contributions to Zoology, accepted. VII. Nagler C, Hörnig MK, Haug JT, Noever C, Glenner H. 2017d. The bigger the better? Volume measurements of parasites and hosts: parasitic barnacles (Cirripedia, Rhizocephala) and their decapod hosts. PlosOne, 12: e0179958. DOI: 10.1371/journal.pone.0179958. VIII. Nagler C, Wagner P, Olesen J, Kerp H, Waloszek D, Haug JT, Haug C. in prep a. The 400-million-year old eucrustacean Ebullitiocaris oviformis re-evaluated: a thecostracan with parasite-type morphology in the Rhynie chert. Palaeontology. IX. Nagler C, Wagner P, Olesen J, Haug JT, Haug C. in prep b. Re-evaluation of nauplius larvae from Rhynie chert supports a phylogenetic affinity to Thecostraca. Palaeontology. iv LIST OF ADDITIONAL PUBLICATIONS I. Matzke-Karasz R, Nagler C, Hofmann S. 2014. The ostracod springtail – camera recordings of a previously undescribed high-speed escape jump in the genus Tanycypris (Cypridoidea, Ostracoda). Crustaceana, 87: 1072–1094. DOI: 10.1163/15685403-00003343. II. Nagler C, Matzke-Karasz R, Geist J. 2014. A revision of the genus Tanycypris (Ostracoda, Cypricercinae) with the description of a new species and a dichotomous key of the genus. Zootaxa. 38214: 401–424. DOI:10.11646/zootaxa.3821.4.1 III. Nagler C, Haug JT. 2015. From fossil parasitoids to vectors: insects as parasites and hosts. In: DeBaets K, Littlewood T (eds.), Advances in Parasitology, 90: 137– 200. DOI: 10.1016/bs.apar.2015.09.003. IV. Haug JT, Nagler C, Haug C, Hörnig MK. 2017. A group of assassin fly pupae preserved in a single piece of amber. Bulletin of Geosciences, in revision. v STATEMENT OF AUTHOR’S CONTRIBUTION In this thesis, I present the results from my doctoral research conducted from 2014 until 2017, carried out under the supervision of Prof. Dr. Matthias J. Starck and Dr. Joachim T. Haug at the Ludwig-Maximilians-Universität München. The author contribution is written according to the CRediT taxonomy. CONTRIBUTION TO PAPER I: NAGLER ET AL. 2016 Conceptualization, CN, CH, UR and JTH; Methodology, CN, CH and JTH; Formal Analysis, CN and JTH; Investigation, CN; Resources, UR and JK; Data curation, CN; Writing – Original Draft, CN; Writing – Review & Editing, CH, JK and JTH; Visualization, CN; Supervision, CH and JTH; Funding Acquisition, CN, CH, JK and JTH. CONTRIBUTION TO PAPER II: NAGLER & HAUG 2016 Conceptualization, CN and JTH; Methodology, CN and JTH, Formal Analysis, CN, Investigation, CN; Resources, CN and JTH, Data curation, CN; Writing – Original Draft, CN; Writing – Review & Editing, JTH; Visualization, CN; Supervision, JTH; Funding Acquisition, CN and JTH. CONTRIBUTION TO PAPER III: SERRANO-SÁNCHEZ ET AL. 2016 Conceptualization, MLSS, CCE and FJV; Methodology, CN, CH and JTH; Formal Analysis, CN and JTH; Investigation, MLSS, CN, CH, JTH, FJV; Resources, CCE and FJV; Data curation, MLSS and FJV; Writing – Original Draft, MLSS and CN; Writing – Review & Editing, CH, JTH, CCE and FJV; Visualization, CN and JTH; Supervision, JTH and FJV; Funding Acquisition, CN, CH, JTH, CCE, FJV. CONTRIBUTION TO PAPER IV: NAGLER ET AL. 2017a: Conceptualization, CN and JTH; Methodology, CN and JTH; Formal Analysis, CN and JTH.; Investigation, CN, MH and JTH; Resources, BSPG (MN and AN), ZSM (RM, ES and SF), MH; Data curation, CN; Writing – Original Draft, CN; Writing – Review & Editing, MH and JTH; Visualization, CN; Supervision, MH and JTH; Funding acquisition, CN, MH and JTH. CONTRIBUTION TO PAPER V: NAGLER ET AL. 2017b: Conceptualization, CN and HG; Methodology, CN; Formal Analysis, CN; Investigation, CN, JTH, HG, JB; Resources, CN, JB, SMNS Stuttgart, ZMUC Copenhagen; Data curation, CN; Writing – Original Draft, CN; Writing – Review & Editing, JTH, HG and JB; Visualization, CN; Supervision, HG and JB; Funding Acquisition, CN, JTH, HG and JB. CONTRIBUTION TO PAPER VI: NAGLER ET AL. 2017c: Conceptualization, CN, CH and JTH; Methodology, CN, CH and JTH; Formal Analysis, CN, CH and JTH.; Investigation, CN; Resources, SMNS Stuttgart, MNHN Paris, ZMUC Copenhagen, JTHo and JTH; Data curation, CN; Writing – Original Draft, CN; Writing – Review & Editing, JTHo, CH and JTH; Visualization, CN; Supervision, CH and JTH; Funding acquisition, CN, JTHo, CH and JTH. vi CONTRIBUTION TO PAPER VII: NAGLER ET AL. 2017d: Conceptualization, CN and HG; Methodology, CN, MKH and CNo; Validation, CN; Formal Analysis, CN and HG; Investigation: CN; Resources, CN and CNo; Data curation, CN and CNo; Writing – Original Draft, CN; Writing – Review & Editing, MKH, JTH, CNo, JTH and HG; Visualization, CN and MKH; Supervision, JTH, JH and HG; Project Administration, CN and HG; Funding acquisition, CN, MKH, JTH, JH and HG. CONTRIBUTION TO PAPER VIII: NAGLER ET AL. in prep a: Conceptualization, CN, PW, JTH and CH; Methodology, CN, PW, JO and CH; Formal Analysis, CN and PW; Investigation, CN, PW, JTH and CH; Resources, Paleobotanical Collection University Münster, CN and JO; Data curation, MorphDBase, CN; Writing – Original Draft, CN and PW; Writing – Review & Editing, JO, WD, HK, JTH and CH; Visualization, CN and PW; Supervision, JTH and CH; Funding Acquisition, CN, JO, JTH and CH. CONTRIBUTION TO PAPER IX: NAGLER ET AL. in prep b: Conceptualization, CN, PW, JTH and CH; Methodology, CN, PW, JO and CH; Formal Analysis, CN and PW; Investigation, CN, PW, JTH and CH; Resources, Science Collections of the National Museum of Scotland, CN and JO; Data curation, MorphDBase and CN; Writing – Original Draft, CN and PW; Writing – Review & Editing, JO, JTH and CH; Visualization, CN and PW; Supervision, JTH and CH; Funding Acquisition, CN, JO, JTH and CH. Ort, Datum Christina Nagler Ort, Datum Dr. Joachim T. Haug Prof. Dr. Matthias J. Starck vii „Ingen flat vei førrer på toppen!” (Unknown author, Norway) CONTENTS EIDESSTATTLICHE ERKLÄRUNG ................................................................................ iii ERKLÄRUNG ..................................................................................................................... iii LIST OF PUBLICATIONS INCORPORATED IN THIS DISSERTATION ..................... iv LIST OF ADDITIONAL PUBLICATIONS ......................................................................... v STATEMENT OF AUTHOR‘S CONTRIBUTION ............................................................ vi CONTENTS ......................................................................................................................... ix SUMMARY ........................................................................................................................... 0 1 INTRODUCTION .......................................................................................................... 1 1.1 PARASITISM ........................................................................................................ 1 1.2 ARTHROPODS ..................................................................................................... 4 1.2.1 CYMOTHOIDA .............................................................................................. 5 1.2.2 THECOSTRACA
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