The first section of this thesis mainly contains alpha-taxonomical contributions to the knowledge of fossil odonates from several outcrops.  

* Triassic odonates: the odonate fauna of the Upper Triassic of Bergamo is revised. A new genus and species Italomyrmeleon bergomensis is described as first Protomyrmeleontidae from the Triassic of Europe. A tiny fossil odonate which belongs to a new genus and species is described but not named because it is a poorly preserved specimen. Italophlebia gervasuttii is redescribed and a new species of the same genus, Italophlebia paganoniae, is described. The phylogenetic position of Italophlebia is discussed and the genus is shown to be one of the oldest known stemgroup representatives of Anisoptera, and is therefore transferred from Zygoptera - Hemiphlebioidea to "Anisozygoptera" - Isophlebioptera. The taxonomy of all fossil odonates from other Triassic localities is discussed, and several new positions and new synonymies are suggested. Reisia nana is described as new species from France. Reisia (= Triadotypus) and Triassologus are both transferred from "Protodonata" to Triadophlebioptera, so that there are no Triassic protodonates known any longer. Thuringopteryx gimmi is transferred from "Protodonata" to Palaeodictyoptera, as the first known Triassic representative of this group. 
  
* Anisoptera from the Upper Jurassic of Germany: The dragonfly genus Urogomphus from the Upper Jurassic of Germany is revised and its position in Aeschnidiidae is confirmed. Urogomphus giganteus and U. eximius are redescribed, and a lectotype for U. eximius is designated. Lithoaeschnidium viohli is considered as a synonym of U.  eximius. A new species Urogomphus nusplingensis is described from the Upper Jurassic limestones of Nusplingen, while the 20 other known specimens of this genus have been found in the Solnhofen Lithographic Limestone. Urogomphus abscissus is considered as conspecific with Bergeriaeschnidia inexpectata, and the holotype of the latter species is designated as neotype of U. abscissus, so that its valid name is now Bergeriaeschnidia abscissa. The phylogenetic position of Urogomphus and Aeschnidiidae is discussed, and an explanation for the extinction of Aeschnidiidae, Mesozoic anisozygopteres, and Palaeozoic protodonates is proposed. 
  
Protolindenia wittei and the genus Aeschnogomphus from the Upper Jurassic of Germany, are revised. Aeschnogomphus buchi is recognized as valid name for Aeschnogomphus charpentieri. Protolindenia, formerly considered to be a gomphid, is transferred to Petalurida, as most basal member of the stemgroup of Petaluridae. Aeschnogomphus, formerly considered to be a Cordulegastridae or Gomphidae, is transferred to Aktassiidae in Petalurida. 
  
The type species of the genus Mesuropetala is redescribed, and Mesuropetala muensteri is considered as its valid name instead of Mesuropetala koehleri. The genus Aeschnopsis is transferred to Mesuropetalidae, and two new species, Aeschnopsis perkinsi and A. tischlingeri are described from the Solnhofen Lithographic Limestone. Cymatophlebia longialata is redescribed. Curious (autapomorphic) structures on the male abdomen of Cymatophlebia spp. are described in detail and their function is discussed. Four new species of the genus Cymatophlebia are described: Cymatophlebia kuempeli and Cymatophlebia pumilio from the Solnhofen Lithographic Limestone, and Cymatophlebia suevica and Cymatophlebia herrlenae from the Malm beta of the Swabian Alb. The two holotype specimens of Cymatophlebia suevica and Cymatophlebia herrlenae represent the oldest known crowngroup representatives of Anisoptera, and the first fossil insect remains known from the Malm beta of the Swabian Alb. With an estimated wing span of more than 22 cm, Cymatophlebia suevica seems to represent the biggest Anisoptera and even the biggest crowngroup Odonata known at all. A new genus Eumorbaeschna with the type species "Cymatophlebia" jurassica (Carpenter, 1932) is proposed as "replacement" name for the aeshnid described by Needham (1907) under the incorrect name "Morbaeschna muensteri" because of a misidentified type species. The genus Morbaeschna is synonymized with the genus Mesuropetala. 
  
The adult fossil dragonflies from the Lower Cretaceous of Mongolia that were previously attributed to the species Sona nectes (Sonidae) are classified as new taxon Proterogomphus krauseorum in a new family Proterogomphidae within the monophylum Gomphides. A new species Proterogomphus renateae is described from the Solnhofen Lithographic Limestone. The status of the alleged hemeroscopid larvae is discussed and they are preliminarily transferred as new (unnamed) species to Sonidae, which is restricted to the referring fossil larvae that have to be regarded as larval Aeschnidiidae. 
  
Nannogomphus bavaricus from the Solnhofen Lithographic Limestone is redescribed. The phylogenetic affinities of Nannogomphus within Anisoptera are discussed. The alleged gomphid affinities are dismissed as based on symplesiomorphies and convergences. Nannogomphus is classified in a new family Nannogomphidae, which is regarded as most basal representative of Brachystigmata. This new family probably also includes a new genus with two new species, Prohemeroscopus jurassicus and P. kuehnapfeli, that are described from the same layers. 
  
A new genus and species Juracordulia schiemenzi is described from the Solnhofen Lithographic Limestone. The analysis of the wing venation reveals that this new dragonfly species belongs to the stemgroup of Eurypalpida (Libelluloidea auct.). It is the first certain record of Eurypalpida from this locality and the first Jurassic and thus oldest record of this taxon at all.

* New Anisoptera from the Lower Cretaceous of China: a new genus and species Progobiaeshna liaoningensis is described from the Lower Cretaceous of China in a new family Progobiaeshnidae, which is regarded as sistergroup of Aeshnida within the new higher taxa Aeshnomorpha - Panaeshnida. Gobiaeshna occulta is preliminarily attributed to this new family, too. 
  
* Odonata from the Lower Cretaceous of Brazil: an overview of the fossil odonate fauna of the Crato Formation from the Lower Cretaceous of Brazil is given. Currently 384 specimens (264 adults and 120 larvae) in 12 families and 34 species are known to science. More than half of the adult and larval fossil odonates belong to the clade Gomphides, especially to the Cordulagomphinae which supports the hypothesis of an allochthonous origin of the aquatic insects. The new genus and species Araripeliupanshania annesuseae is described in Mesuropetaloidea - Liupanshaniidae, and represents the first New World record of this group. Two new genera and three new species, Paramorbaeschna araripensis, Progomphaeschnaoides ursulae, and Progomphaeschnaoides staniczeki, are described within Neoaeshnida - Gomphaeschnidae in a new subfamily Gomphaeschnaoidinae. In the same group three further new species, Gomphaeschnaoides magnus, Gomphaeschnaoides petersi, and Gomphaeschnaoides betoreti are described, together with a redescription of the type species Gomphaeschnaoides obliquus, including its previously unknown forewings and body. Six further new taxa are described: Araripegomphus andreneli (Araripegomphidae), Cordulagomphus (Procordulagomphus) senckenbergi (Proterogomphidae - Cordulagomphinae), Araripephlebia mirabilis (Araripephlebiidae), Cratocordulia borschukewitzi (Araripelibellulidae), Cretarchistigma essweini (Zygoptera incertae sedis), and Parahemiphlebia mickoleiti (Hemiphlebiidae). With a wing length of only 9 mm the latter new species represents one of the smallest odonates of all times. Araripephlebia mirabilis is classified in a new family Araripephlebiidae that probably represents the sistergroup of Valdicorduliidae. A still unnamed new genus and species represents the first fossil record and the first New World record for Chlorogomphida. Four further new species are illustrated and discussed, but not yet named. The phylogenetic relationship of several known species is discussed, and some diagnoses are amended or corrected. Giant dragonfly larvae of up to 70 mm length are described, regarded as older stages of Nothomacromia sensibilis, and considered as larval Aeschnidiidae. The position of Araripegomphidae in the stemgroup of Gomphides rather than Eurypalpida (= Libelluloidea auct.) is advocated (contra Lohmann, 1996). The former genus Procordulagomphus is down-ranked to a subgenus of Cordulagomphus. "Cordulagomphus" santanensis is recognized as earwig and thus transferred from Odonata - Cordulagomphinae to Dermaptera incertae sedis. The taphonomy of the Crato limestones is discussed and an origin in a brackish lagoon without autochthonous insects is advocated. A comparison with the odonate fauna of the Upper Jurassic Lithographic Limestones of Solnhofen reveals several remarkable differences. Because of the absence of typical Mesozoic odonate groups, such as anisozygopteres, Archizygoptera and Steleopteridae, as well as the presence of extant families of Zygoptera, the odonate fauna of the Crato Formation appears to be already significantly more advanced. 
  
* Odonata from amber and fossil Epallagidae: Litheuphaea ludwigi is described as new species and first representative of Epallagidae from Baltic amber, and the holotype of Litheuphaea carpenteri is redescribed.  
The authorship of Selys (1853) for the family-group name Euphaeidae is rejected, since the "légion Euphaea" proposed by Selys is neither a noun in the nominative plural, nor ending in a latinized suffix. Consequently, the correct family name must be Epallagidae Needham, 1903, since Euphaeidae were first established by Jacobson & Bianchi (1905) and thus have to be considered as a junior subjective synonym. Similarly, all the other "légions" proposed by Selys are rejected as available family-group taxa, so that the next available family-group name has to be used, e.g. Heliocharitidae Tillyard & Fraser, 1939 instead of Dicteriadidae Montgomery, 1959 (nec Selys, 1853). A unique fossil odonate is briefly described, which represents a damselfly in Baltic amber that is just emerging from the exuvia (probably Platystictidae or Megapodagrionidae). An annotated new catalogue of all known odonates in amber is provided, including 70 specimens from Lebanon, Dominican, Paris, Baltic and Saxonian amber, of which 6 specimens are adult Anisoptera and 5 specimens are exuviae. A lectotype for Platycnemis antiqua is designated and illustrated. 
  
The second section of the thesis contains phylogenetic systematic analyses and conclusions about the evolution of odonates.  
  
Some theoretical problems of modern phylogenetic research are discussed, with special consideration of the issue of character weighting and its implications for the principle of parsimony. The cladistic paradigm of parsimony is rejected and a broader concept of phylogenetic parsimony is advocated. Pattern cladism is shown to be fundamentally different from Hennigian phylogenetic systematics, and more similar to numerical taxonomy (phenetics). 
  
A set of basic principles and rules for phylogenetic taxonomy and classification is proposed, including a new notation for synonymic lists that also considers taxa that are synonymous in content but no synonyms according to the current rules of zoological nomenclature. 
  
Based on a very comprehensive "manual" phylogenetic analysis, a reclassification of all higher taxa of fossil and extant odonates above the genus-group level is introduced. For all proposed monophyla a complete list of all known autapomorphies (including wing venational characters, as well as all other available evidence, e.g. body anatomy, larval morphology, molecular data, behaviour, etc.), a recommended usage, and a complete synonymy is provided. Numerous new higher taxa are described (see list below). The most important results include the confirmation of the sistergroup relationship of Ephemeroptera and Odonatoptera (monophyly of Palaeoptera), the demonstration of monophyly of Zygoptera (contra Fraser, 1954, 1957; Hennig, 1969, 1981; Pfau, 1986, 1991; Zessin, 1991; Trueman, 1991, 1996; Nel et al., 1993), the identification of Tarsophlebiidae as fossil sistergroup of crowngroup Odonata, the identification of the basal splitting events in Anisoptera with Petalurida being recognized as most basal clade, as well as a revised position of Epiophlebiidae within the "anisozygopteroid" grade. Only the ingroup relationships within extant Coenagrionidae, Gomphidae and Libellulidae could not be completely resolved yet. The main differences of the proposed system to other recently proposed phylogenetic systems of Odonata (Carle, 1995, 1996; Trueman, 1996; Lohmann, 1996) are discussed. 
 
At the example of Aeshnoptera, it is demonstrated that many derived wing venational characters have a very incongruent pattern, which implies either multiple convergent gains, or reductions, of the referring characters. The problem of this high level of homoplasy for the phylogenetic reconstructions is discussed. 
  
A numerical cladistic parsimony analysis of the major clades of Anisoptera confirmed the results of the "manual" phylogenetic analysis. Only the phylogenetic position of Gomphides (incl. Proterogomphidae) could not be correctly resolved, because of the lack of appropriate wing venational apomorphies. A "manual" phylogenetic analysis, based on the morphology of extant dragonflies, identifies Petalurida as sistergroup of all remaining crowngroup Anisoptera. Several important plesiomorphic features of Petalurida are discussed, and the alleged plesiomorphy of semi-terrestrial larvae is rejected. A numerical cladistic analysis and "manual" phylogenetic analysis of all fossil and extant petalurid genera is provided. Two different scenarios of the historical biogeography of Petalurida are discussed. 

Taxonomy:  

This thesis includes the description of 83 new higher taxa (above the family-group level), 26 new families, 18 new subfamilies, 10 new tribes, 5 new subtribes, 15 new genera, 1 new subgenus, and 30 new species. Furthermore, there are 80 taxa stat. nov. (nomina translata), 10 taxa stat. rest., 95 taxa sens. nov., 73 taxa pos. nov., 27 new synonymies, 3 type designations, and 5 corrected spellings (justified emendations and mandatory changes). Please note that all listed taxonomical acts are only becoming available and valid with the referring publications from this PhD thesis (see list below), which itself has to be regarded as unpublished according to Art. 8 and 9 IRZN. 
 
This thesis is published in several parts with the following publications:   
  
Bechly, G. (1996): Morphologische Untersuchungen am Flügelgeäder der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata), unter besonderer Berücksichtigung der Phylogenetischen Systematik und des Grundplanes der *Odonata. - Petalura, spec. vol. 2: 1-402; Böblingen.  

Bechly, G. (1997): New fossil odonates from the Upper Triassic of Italy, with a redescription of Italophlebia gervasuttii, and a reclassification of Triassic dragonflies (Insecta: Odonata). - Rivista del Museo Civico di Scienze Naturali "Enrico Caffi", 19: 31-70; Bergamo.  

Bechly, G. (1998a): New fossil dragonflies from the Lower Cretaceous Crato Formation of north-east Brazil (Insecta: Odonata). - Stuttgarter Beiträge zur Naturkunde (B), 264: 1-66; Stuttgart.  

Bechly, G. (1998b): Phylogenetic Systematics of Odonata. - [Website on the Internet, new URL: http://members.tripod.de/GBechly/index.htm]. [will be electronically published on CD-ROM: Schorr, M. & Lindeboom, M. (eds) (2000): Fundamentals of odonatological research. Tübingen]  

Bechly, G. (1998c): Juracordulia schiemenzi gen. et. sp. nov., eine neue Libelle aus den Solnhofener Plattenkalken (Insecta: Odonata: Anisoptera). - Archaeopteryx, 16: 29-36; Munich.  

Bechly, G. (1998d): New fossil damselflies from Baltic amber, with description of a new species, a redescription of Litheuphaea carpenteri Fraser, and a discussion on the phylogeny of Epallagidae (Zygoptera: Caloptera). - International Journal of Odonatology (Pantala), 1(1): 33-63; Leiden.  

Bechly, G. (1998e): A revision of the fossil dragonfly genus Urogomphus, with description of a new species (Insecta: Odonata: Pananisoptera: Aeschnidiidae). - Stuttgarter Beiträge zur Naturkunde (B), 270: 1-47; Stuttgart.  

Bechly, G., Brauckmann, C. & Zessin, W. (in prep.): Neue Erkenntnisse zur Morphologie der ältesten Libellen (Insecta: Odonatoptera) aus dem Namurium von Hagen-Vorhalle (Deutschland).  

Bechly, G., Nel A. & Martínez-Delclòs, X. (1996): Redescription of Nannogomphus bavaricus Handlirsch, 1906-1908, from the Upper Jurassic of Germany, with an analysis of its phylogenetic position (Odonata: Anisoptera: Gomphidae or Libelluloidea). - Archaeopteryx, 14: 51-66; Munich.  

Bechly, G.,Nel, A., Martínez-Delclòs, X. & Fleck, G. (1998): Four new dragonfly species from the Upper Jurassic of Germany and the Lower Cretaceous of Mongolia (Anisoptera: Hemeroscopidae, Sonidae, and Proterogomphidae fam. nov.). - Odonatologica, 27(2): 149-187; Bilthoven.  
  
Bechly, G., Nel, A., Martínez-Delclòs, X., Jarzembowski, E.A., Coram, R., Martill, D.,Fleck, G., Escuillié, F., Wisshak, M.M. & Maisch,M. (in press): A revision and phylogenetic study of Mesozoic Aeshnoptera, with description of several new families, genera and species (Insecta: Odonata: Anisoptera). Neue Paläontologische Abhandlungen, 4, Dresden. 
  
Nel, A., Bechly, G., Jarzembowski, E.A. & Martínez-Delclòs, X. (1998): A revision of the fossil petalurid dragonflies (Insecta: Odonata: Anisoptera: Petalurida). Paleontologia Lombarda (N.S.), 10: 1-68, Milano. 

Rust, J. 1999.- Biologie der Insekten aus dem ältesten Tertiär Nordeuropas [Biology of insects from the Lower Tertiary of Northern Europe]. Habilitationsschrift, Biologische Fakultät der Georg-August-Universität Göttingen, 482 S., 131 Abb., 34 Taf., Germany. 
Email: jrust@gwdg.de 

The following section is a summary of my postdoctoral thesis about the investigation of fossil insects from the Paleogene Fur- and Ølst-Formations ("Mo clay") in Denmark. The publication of the complete work in English is in progress.

Abstract: Until recently little was known about the biology of Palaeocene and lowermost Eocene insects due to the rarity of insect-bearing strata from those periods. Owing to particularly favourite climatic conditions, the far-reaching occurrence of rain forests and the rich diversification of angiosperms the Lower Tertiary was of special importance for the evolution of insects without a doubt. In the present work an as-complete-as possible scenario of insect biocenoses of those times including the prevailing conditions which insects had to face is reconstructed. 

The present investigations are based on the systematic examination of all insect taxa (about 20.000 specimens) from the Paleogene marine deposits of the Ølst - and Fur-formations of Denmark which are about 55 million years old. The findings are characterised by a brilliant preservation of even the finest details. An inventory of 182 species was identified, 107 of which were new to science. The most common insects belong to the Diptera, Heteroptera and Auchenorrhyncha. These are particularly diverse, and especially the Fulgoromorpha are represented by higher numbers of different forms than anywhere else. Compared with all other known fossil-bearing sites the Lepidoptera are represented by numbers high above average (more than 1700 specimen out of 6 species). For one fossil Lepidopteran species evidence of mass migrations as known for recent species was found for the first time. Saltatoria, Neuroptera and Hymenoptera are also rather rich in species. For the Saltatoria the oldest known fossil evidence of Tettigoniidae, Pteroplistinae, Oecanthinae and Acrididae in Danish Paleogene sediments is presented. For the most abundant tettigoniid (Pseudotettigonia) an exact reconstruction of the original singing and hearing ability is presented. Other insect taxa (Odonata, Mantodea, Blattoidea, Dermaptera, Aphidoidea, Coleoptera, Mecoptera and Trichoptera) are likewise represented in outstanding findings, some of which allow detailed statements on the habits of these species. 

The insects were transported from southern and south-western Scandinavia to the former North Sea by active flight, passive winddrift or with plant remains. Flightless forms were not found. A plausible model for the special condition under which the deposition of insects took place in a highly productive upwelling area is postulated. The original flight ability is ascertained for all taxa by comparing them to recent representatives of these taxa. The investigations are used for reconstructing the distance from deposition site to the mainland. 
The original former flight ability as well as the taxonomic composition of the different strata show that the sediments of the Ølst-formation were deposited closer to the shore than those of the Fur Formation. In addition the faunal composition shows that insects always flew during summers with high temperatures and prevailing northerly winds. The climate of the North Sea area was influenced by season and the insect fauna gives hints of subtropical or paratropical conditions during the Lower Tertiary. Examples from the Danish Paleogene are known which show that climatic adaptations of some of the recent insects to moderate temperatures may have developed as late as during the course of the Tertiary and Quaternary. 

The high diversity of plant-sucking insects gives evidence of lush and highly diverse vegetation on the Scandinavian mainland. A broad spectrum of freshwater insects suggests the existence of large and diverse lakes, ponds and rivers. In addition there are a lot of fossil forms whose extant relatives are inhabitants of swamplands. Especially diverse are those insects that give evidence of the former existence of forests and open areas with shrub vegetation. 

Very close paleobiogeographic relations are found for the congruent occurrence of seven species between the Ølst- and Fur-formations and the Lower Tertiary of Havighorst near Hamburg. No close relationship was found with other Paleogene insect assemblies which were found in sediments (France, England, Greenland, Svalbard, North America) or amber (Sibiria). However, general faunistic correspondence is found between the entomofaunas discussed herein and those from North America. Close relations to faunas which have their main distribution in South and Southeast Asia or America are found when considering recent species. In other cases the representatives of today show a Gondwana-like distribution or are cosmopolites. 

The in part dramatic events at the Cretaceous-Tertiary-boundary (volcanism, meteorite) obviously had no major impact on the evolution of insects. The insects of the Ølst- and Fur-formations were highly developed and closely resemble recent species. 34 species may be attributed to 25 genera which still exist today. This value (about 50 % of all identified genera) is very high even when compared to more recent entomofaunas. It thus may be concluded that the systematic and ecological basis of recent insects developed during the Mesozoic and it is likely that these events are closely connected with the rich development of the angiosperms during the Middle and Upper Cretaceous.

 
Abstract: The present work is an analysis of the historical development of insect faunas and the comparison of these data with the results of the studies of mammals for understanding the processes of the regional development of faunas in general. These aims are achieved by the following: 

  1. Description of Pleistocene and Holocene insect faunas from the central and northern parts of West Siberia within the taiga zone. 
  2. Ascertaining the time of appearance of modern insect communities. 
  3. Comparison of these data with the results of the studies of small mammals from the Quaternary deposits known from the literature. 
  4. Identifying the stages of faunal complex formation in the West Siberian taiga zone. 

Fossil insect material was collected from 37 localities, these faunas dating from the Early, Middle, and Late Pleistocene and Holocene. Pleistocene and Holocene insect faunas were divided into four types: tundra-like, forest-tundra, forest and intrazonal; this subdivision was based on the alignment of arctic, hemiarctic, boreal and polyzonal species of insects. 

Most Pleistocene insect faunas can be characterised as tundra-like differing from modern tundra insect communities by the predominance of arctic species (ground beetles Pterostichus costatus Men., P. sp.cf. pinguedineus Esch., P. sp. cf. ventricosus Esch., Curtonotus alpinus Pk.; rare beetle Tachinus arcticus Maekl.; weevil Lepyrus nordenskjoldi Fst. et al.) with the presence of species not found in modern tundras (ground beetles Epaphius rivularis Gyll., Pterostichus mannerheimi Dej., Synuchus vivalis Pk., Poecilus sp. cf. ravus Lutschn. et al.). Early Holocene insect faunas are referred to tundra-like, forest and intrazonal types; Late Holocene faunas to forest and intrazonal types. 

 The formation of modern insect faunas, like mammals, coincided with a period at the end of the Last Glaciation and the beginning of the Holocene. This process corresponded to a subdivision of tundra-like and periglacial faunas which was accompanied by extinction of some large mammals (mammoth, woolly rhinoceros), changes in the configuration of areas of tundra and distribution of xerophilous species (insects and mammals), and the conservation of some boreal and polyzonal species. This data confirms the allochthonous character of West Siberian faunas.