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(COLEOPTERA: CUCUJIFORMIA) Ph.D. Thesis

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(COLEOPTERA: CUCUJIFORMIA) Ph.D. Thesis Palacký University in Olomouc Faculty of Science Department of Zoology and Anthropology Zuzana Levkani čová MOLECULAR PHYLOGENY OF THE SUPERFAMILY TENEBRIONOIDEA (COLEOPTERA: CUCUJIFORMIA) Ph.D. Thesis P1501 Zoology Supervisor: Prof. Ing. Ladislav Bocák, Ph.D. Olomouc 2009 I undersigned Zuzana Levkani čová declare to have written this Ph.D. thesis alone during 2004-2009 in the Department of Zoology and Anthropology under the supervision of Prof. Ing. Ladislav Bocák, Ph.D. and used the references enclosed in the Ph.D. thesis. Olomouc 2009 Supervisor: Prof. Ing. Ladislav Bocák, Ph.D. Ph.D.candidate: Mgr. Zuzana Levkani čová BIBLIOGRAPHICAL IDENTIFICATION Author’s first name and surname: Zuzana Levkani čová Title: Phylogeny of the superfamily Tenebrionoidea (Coleoptera: Cucujiformia) Type of thesis: Ph.D. thesis Department: Department of Zoology and Anthropology Supervisor: Prof. Ing. Ladislav Bocák, Ph.D. The year of presentation: 2009 Abstract: The phylogenetic relationships of the superfamily Tenebrionoidea are investigated here for the first time. The Tenebrionoidea (darkling beetles) is superfamily of speciose rich and complex series Cucujiformia, that is considered as the most derived among the Coloeptera. The Tenebrionoidea itself is very diverse group and contain approximately 30.000 species classified in 30 families. It has been recognized as a relative to the Cucujoidea superfamily, however the position within the Cucujiformia has not been stabilized yet. The intrarelationships of the Tenebrionoidea are also poorly known, since only the studies on either generic or subfamilial level have been published. Here, two nuclear genes SSU and LSU rDNA and two mitochondrial genes rrnL rDNA and cox1 mtDNA of total length approximately 3700 bp were used to reveal the phylogeny of this puzzling group. There were sampled 154 taxa representing 20 families of the superfamily. Both, static and dynamic multiple alignments of combined dataset were performed, followed by the analyses of maximum parsimony and maximum likelihood and bayesian analysis. They confirm the monophyly of the group, proposing its closer relationship to the Lymexyloidea than it has been recognized before. Within the superfamily, four clades of families have been established- tenebrionid, melandryinid, ripiphorid-mordellid-meloid, and scraptiid- pyrochroid. The monophyly of most of families has been confirmed as well, except the families Salpingidae, Pyrochroidae, Anthicidae that have been found paraphyletic and the families Tetratomidae, Melandryidae and Zopheridae found polyphyletic. The paraphyletic status would be changed to monophyletic if certain taxa as Ischaliinae and Eurygeniinae (Anthicidae), Agnathinae (Pyrochroidae), Othniinae (Salpingidae) are excluded out of the family. The polyphyletic families should be revised considering their division in smaller units. There are the high degree of homoplasy and the complexity of the group found as reasons of unsatisfyingly resolved phylogeny of the group. More comprehensive and extensive studies, that would involve both molecular and morphological characters, inclusion of all families as well as of members of the Cucujiformia series and more extensive analyses, will be needed to recognize true relationships within the Tenebrionoidea. Keywords: Alignment, phylogeny, rDNA, mtDNA, systematics, taxonomy, Tenebrionoidea. Number of pages: 98 Number of appendices: 2 Language: English Contents 1. Introduction -1- 2. Aims of the Ph.D. thesis -5- 3. Literature review -6- 3.1 The superfamily Tenebrionoidea -6- 3.2 The families of the superfamily Tenebrionoidea -7- 3.3 Molecular markers -41- 4. Material and methods -44- 4.1 Sampling of taxons -44- 4.2 Laboratory methods -44- 4.3 Phylogenetic analyses -46- 4.3.1 Sequences analyses -46- 4.3.2 Multiple alignment -46- 4.3.3 Tree search methods -48- 4.3.3.1 Maximum Parsimony -48- 4.3.3.2 Bayesian analysis -48- 4.3.3.3 Maximum Likelihood -49- 4.3.4 Taxonomic Retention Index -49- 5. Results -51- 5.1 Sequences and alignment -51- 5.2 Phylogenetic analyses -55- 5.2.1 Maximum Parsimony analyses -55- 5.2.2 Bayesian analyses -62- 5.2.3 Maximum Likelihood analyses -65- 5.2.4 Taxonomic Retention Index -67- 6. Discussion -69- 6.1 Monophyly of the Tenebrionoidea -69- 6.2 Internal relationships within the Tenebrionoidea -69- 6.3 The evolution of hypermetamorphosis -81- 7. Conclusion -83- 8. Souhrn -84- 9. Acknowledgements -85- 10. Abbreviations -86- 11. List of figures -88- 12. List of tables -89- 13. References -90- Supplementary material A. Classification B. Sampling list Author’s publications 1 1. Introduction. The Tenebrionoidea, formerly known as Heteromera, is a speciose, morphologically and ecologically heterogenous superfamily of polyphagan beetles. It is placed within the Cucujiformia series. Tenebrionoidea contain approximately 30 000 species classified in 30 families and 71 subfamilies (Lawrence & Newton, 1995). Generally known large families are Tenebrionidae (darkling beetles) and Meloidae (blister beetles). Other species rich families are Anthicidae, Mordellidae, Oedemeridae, Zopheridae and Aderidae, while other families include only one or a few genera. Traditionally, Tenebrionoidea have been accepted as a lineage within Cucujiformia. The suborder Polyphaga, where they are placed, may have originated ca 270 Mya, the Cucujiformia ca 236 Mya, and the Tenebrionoidea in the Late Triassic according to Hunt et al. (2007). The origin of the Meloidae has been determined by the fossil record to an Early Cretaceous period (125–135 Mya), the period of flowering plant radiation (Bologna et al. , 2008). The Heteromera, as a separated section, were for the first time distinguished in the beetle system by P. A. Latreille (1803), the first entomologist, who divided the Coleoptera in supergeneric taxa, based on the tarsal segmentation. Since, Heteromera have been recognized in every classification, though in different positions. Lameere placed it in 1900 in suborder Cantharidiformia; Kolbe, in 1901, found Heteromera in suborder Heterophaga; in 1903 Ganglbauer similarly put it in suborder Polyphaga. All these authors left families Mycetophagidae, Ciidae, Colydiidae either in Clavicornia or Diversicornia section. This trend continued in the beginning of the 20 th century. All classifications, including those by Sharp and Muir’s (1912) based on the male genital tube, Forbes’s (1926) based on the wing venation and wing folding patterns and Poll’s (1932) based on the structure of the Malpighian tubules, found a separated superfamily Heteromera. However Sharp and Muir (1912) admitted only few families allied to Tenebrionidae to be a part of the Tenebrionoidea and they placed all remaining families in the Cucujoidea. Böving and Craighead’ study of larval types, (Böving & Craighead, 1931), united the Heteromera and Clavicornia in a single superfamily Cucujoidea and they elevated the family Mordellidae to the separated superfamily Mordelloidea and families Meloidae and Rhipiphoridae to the superfamily Meloidea at a coordinated taxon with Cucujoidea. The Peyerimhoff’s classification (Peyerimhoff, 1933) merged Heteromera with Cucujoidea, but the cryptonephridial groups were placed in the end 2 of the system, as the most derived ones. Later, Jeannel and Paulian (1944) published the classification based on structure of the aedeagus and other abdominal features and they established tenebrionoids as the division Heteromeroidea of suborder Heterogastra independently of the division Cucujoidea. They discriminated four sections of Heteromeroidea: Lyttaria, Tenebrionaria, Mordellaria and Oedemeraria. Crowson’s (1955) detailed morphological study of both larvae and adults kept the superfamily Cucujoidea with two recognized sections, Clavicornia and Heteromera. He did not find the differences between them enough substantial to define both of them as superfamilies. According to Crowson, Heteromera arose from primitive Clavicorn types and were the most difficult section to divide into well-characterised families. In this study, he established several new families- Merycidae, Pterogeniidae, elevated several other to family rank: Boridae, Elacatidae, Mycteridae, Inopeplidae and Tetratomidae, the families like Mycetophagidae, Colydiidae, Inopeplidae and Hemipeplidae were transferred from Clavicornia to Heteromera. Although Crowson (1960) mentioned a possibility to establish two or more superfamilies, corresponding with Clavicornia and Heteromera sections, he finally decided to retain a single superfamily because of the unresolved complexity of relationships between families. Crowson (1960) also suggested, that the families Byturidae and Bihpyllidae could be transferred from Clavicornia to Heteromera. Later, Crowson (1966) recognized family Synchroidae and discussed presumable phylogeny of the group. He tentatively proposed a common ancestor of Heteromera, that resembles the family Tetratomidae, both in larval and adult features. As direct descendants were proposed the families Tetratomidae and Mycetophagidae and perhaps Pterogeniidae-Ciidae. The second possible ancestor arose from a tetratomid-like ancestor and might have larval characters like the Zopheridae and adult characters like Synchroa and Stenotrachelus . From this ancestor might be derived (1) the aderid-anthicid-meloid line, (2) a line leading via Pythidae and Pyrochroidae to Salpingidae, Mycteridae, Boridae and Inopeplidae, (3) a line leading via Synchroid and Zopherid-like forms to Merycidae and Monommidae and Colydiidae and perhaps to true Zopheridae
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