Current Position: Postdoctoral Fellow, Steinmann Institute, Division Paleontology, University of Bonn
Education: BSc, Geology, University of Clausthal, 1999 PhD, Paleoentomology, University of Clausthal, 2003 Habilitation, Paleontology, University of Bonn, 2010
Abstract: Altitudinal gradients provide an excellent opportunity to examine the effects of climate change. Ants (Hymenoptera) are ideally suited for studying Ã-diversity along altitudinal gradients because not only are they one of the most ecologically significant components of an ecosystem, many species are restricted to their preferred (micro-) habitats. Here we show the possible effects of global warming on the diversity of ants in the Wurmberg Mountain (Harz Mountains), Germany. We identified three subfamilies, ten genera, and 41 species, collecting 971 ants on Wurmberg Mountain. Higher diversity indices, numbers of individuals and species of ants were observed outside the forest than inside the pine forest, as well as with the decrease of the altitudinal gradient. We observed 18 species of Myrmicinae and 12 of Formicinae (total 30 species) at lower altitudes against seven and five (total 12 species), respectively, at higher altitudes. We found that the ants of the Harz Mountains react as expected to changes in altitude. That said, some species, specifically low-altitude thermophilic ones, show signs of expanding into higher altitudes, a possible reaction to climate warming. The distributions of host-specific herbivorous insects along altitudinal gradients, particularly within montane environments, provide useful analogs for predicted future changes that are likely to occur over time at any one location, given a gradually changing thermal environment.
Abstract: The enigmatic Svalbard Palaeocene fossil taxon Hymenopterites deperditus is revised, and is neither a wasp nor a plant seed, but turns out to be a bug hemelytra corresponding to the oldest described reduviid bug. It can be attributable to the âemesine-saicine cladeâ. The presence in the Svalbard Palaeocene of this insectivorous bug, showing possible mimicry of a wasp model, confirms the presence of diverse entomofauna and of rather warm palaeoclimatic conditions.
Abstract: The first fossil bumble bee (Apinae: Bombini) from the Miocene Randeck Maar of southwestern Germany is described and illustrated. The specimen is subjected to a geometric morphometric analysis along with a diversity of other bumble bee species representing most major extant lineages, and particularly the subgenus Bombus s. str. The morphometric analysis supports the placement of the Randeck Maar species within Bombus s. str., as a species distinct from all others in the subgenus. It shows that extant subgenera of bumblebees were already derived in Early/Middle Miocene. The Randeck Maar fossil is formally described as Bombus (Bombus) randeckensis sp. n..
Abstract: Although plantâarthropod relationships underpin the dramatic rise in diversity and ecological dominance of flowering plants and their associated arthropods, direct observations of such interactions in the fossil record are rare, as these ephemeral moments are difficult to preserve. Three-dimensionally preserved charred remains of Chloranthistemon flowers from the Late Albian to Early Cenomanian of Germany preserve scales of mosquitoes and an oribatid mite with mouthparts inserted into the pollen sac. Mosquitoes, which today are frequent nectar feeders, and the mite were feeding on pollen at the time wildfire consumed the flowers. These findings document directly arthropod feeding strategies and their role in decomposition.
Abstract: Terrestrial plants and insects account for the majority of the Earth's biodiversity today, and herbivorous interactions are dated back more than 400 million yr. However, investigation of their associations remains in its infancy in Europe. The Miocene is characterized by palaeogeographic re-organization due to the collision of the African with the Eurasian plates. Antarctica's enormous impact on global climatic conditions, and thus on European palaeoenvironment, resulted from a series of episodes of minor glaciations in the Early Miocene after the initial cooling and ice sheet formed during the Oligocene.
More than 3500 plant remains showing various kinds of feeding damage were available for the present study. These trace fossils are classified according to their external morphology into damage types (DT) and grouped to functional feeding-groups. The Neogene plant record in Europe is rich and diverse, offering a profound large-scale understanding of the floristic and vegetational development. A database of fossil traces from the Most Basin was compiled and analyzed by various statistical methods in terms of the diversity and intensity of palaeoherbivory. The primary objective is to present results on the development of insect herbivory through the section of the BÃlina Mine in North Bohemia, with the aim of understanding the principal factors that caused the observed phenomena. The research was focused on two horizonsâDelta Sandy Horizon (DSH) and Lake Clayey Horizon (LCH)âboth sufficiently represented to compare their palaeoecological and palaeoclimatological signals on the basis of the presence of damages caused by insects and other herbivorous arthropods. A total sample of 60 different damage types, attributed to eight main functional feeding groups, was examined. Results from analyses of the frequency and diversity of the selected categories of plant arthropod associations within both examined horizons significantly support different environmental conditions. The LCH seems to be affected by the relatively colder and drier climatic conditions as indicated by a four times greater frequency of leaves with galls and lower taxonomic diversity and species equability, whereas DSH indicates warmer and more humid conditions reflecting the higher diversity of the plant species and damage types.
Abstract: Early Tertiary High Arctic forest ecosystems are unique in that they have no equivalent among modern forests. Today, no forest ecosystem exists at such high latitudes. To assess the potential role of herbivory during the early Tertiary warm period at high latitudes, we have surveyed 1567 fossil angiosperm leaves from Svalbard for the presence or absence of 35 insect damage types (DTs). Our investigation for the first time uncovered a wealth of insect trace fossils from the early Tertiary northern high latitudes. These include galls, mines, and feeding traces on fossil leaves. Most of the folivory includes unspecific external foliage feeding that cannot be ascribed to a particular group of insects. Exceptions are the mining damage types that are most similar to those made by leaf-mining moths (Lepidoptera: Gracillaroidea). Nevertheless, the abundance of folivory indicates that herbivorous insects were an important component of the forests thriving in the Arctic realm. The observed change in insect herbivory from the Middle Paleocene to the Late Eocene/Early Oligocene in Spitsbergen may be attributed to climatic variables because they influence insect life-cycle timing, population density, and geographic range.
Abstract: The Miocene Randeck Maar (southwestern Germany) is one of the only sites with abundant material of fossil honey bees. The fauna has been the focus of much scrutiny by early authors who recognized multiple species or subspecies within the fauna. The history of work on the Randeck Maar is briefly reviewed and these fossils placed into context with other Tertiary and living species of the genus Apis Linnaeus (Apinae: Apini). Previously unrecorded specimens from Randeck Maar were compared with earlier series in an attempt to evaluate the observed variation. A morphometric analysis of forewing venation angles across representative Recent and Tertiary species of Apis as well as various non-Apini controls was undertaken to evaluate the distribution of variation in fossil honey bees. The resulting dendrogram shows considerable variation concerning the wing venation of Miocene Apini, but intergradation of other morphological characters reveals no clear pattern of separate species. This suggests that a single, highly variable species was present in Europe during the Miocene. The pattern also supports the notion that the multiple species and subspecies proposed by earlier authors for the Randeck Maar honey bee fauna are not valid, and all are accordingly recognized as Apis armbrusteri Zeuner.
Abstract: The purpose of this application is to provide stability to the name Apis armbrusteri Zeuner, 1931 for a species of fossil honey bee occurring in the Miocene fauna of southwestern Germany. The type series designated by Zeuner (1931) is the hollow impression of a bee from the Early Miocene Böttingen Marmor and, aside from attributing it to the tribe Apini, no details regarding its specific identity can be gleaned from this specimen. Nonetheless, this name has been universally applied to the Early Miocene honey bees from Böttingen Marmor and the related contemporaneous site from the same crater series, Randeck Maar, since Zeuner & Manning (1976). To recognize A. armbrusteri as a nomen dubium and resurrect the unused specific epithet Apis scheuthlei (Armbruster, 1938) for these bees would be counter to current usage and destabilize a voluminous literature on honey bee evolution and ecology. It would also render the subgeneric name Cascapis Engel, 1999, invalid as A. armbrusteri sensu Zeuner & Manning (1976), i.e., based on the Randeck Maar material, is its type species by original designation. Accordingly, it is proposed that the original, unidentifiable type be set aside and one of the more exquisitely preserved and easily diagnosable specimens from this same fauna be designated as neotype, thereby stabilizing the honey bee taxonomy and bringing the application of the name A. armbrusteri into line with universal current usage.
Abstract: Parasites commonly manipulate host behaviour and among the most dramatic examples are diverse fungi that cause insects to die attached to leaves. This death- grip behaviour functions to place insects in an ideal location for spore dispersal from a dead body following host death. Fossil leaves record many aspects of insect behavior (feeding, galls, leaf mining) but to date there are no known examples of behavioral manipulation. Here, we document the first example of the stereotypical death grip from 48-million-year-old leaves of Messel, Germany, indicating the antiquity of this behavior. As well as being the first example of behavioural manipulation in the fossil record, these data support a biogeographical parallelism between mid Eocene northern Europe and recent Southeast Asia.
Abstract: The first find of fossil Heteroptera (Aradidae, Aneurinae) from New Zealand is figured and described. The ventrally preserved specimen from early Miocene lake sediments at Foulden Maar, Otago, is assigned to Aneurus, a flat-bug genus that is present with six species in the modern New Zealand fauna. Aneurus sp. from Foulden Maar represents the first fossil Southern Hemisphere record of the family Aradidae. Its occurrence in the early Miocene of Otago indicates that the genus has been present in the Australasian region at least since the early Miocene and confirms previous suggestions of a long evolutionary history of New Zealand aradids closely linked to rainforest habitats.
Abstract: About thirty fossil bark-gnawing beetles (Trogossitidae) have been described from several Mesozoic and Cenozoic fossil sites. The fossil record is best for the Eocene because of many excellent localities. Fossils from the Mesozoic in contrast are taxonomically problematic and comparatively rare. We have examined fossil specimens of bark-gnawing beetles in excellent condi- tion preserved in limnic sediments and Baltic amber regarding their phylogeny and biogeography. Morphological parameters of the beetles were recorded by using light microscopy as well as by using a micro-CT scanner (μ-CT).
Abstract: Laurophyllum lanigeroides (Engelhardt) Wilde, one of the most common laurel leaves of the Messel flora, is well-known from leaf and cuticle characters but there is no sufficient evidence for confirming affinities to an extant genus of the Lauraceae. In the present study we describe two of the most conspicuous damage types on L. lanigeroides in detail and compare them with known insect damage from extant Lauraceae. They appear to resemble galls induced by extant gall midges known from Asia and the eastern Palaearctic regions. These distinctive damage types lead to a more taxonomically precise identification of the fossil host lineage. Furthermore, the discovery of advanced forms of cecidomyiid insects and their activities in the Tertiary suggests the diversification and establishment of cecidomyiid leaf galls on the nearest living relatives of the Tertiary flora.
Abstract: Insects form an important part of modern terrestrial ecosystems, but while their body remains are rare in the fossil record, their trace fossils, such as feeding damage, are more common. Studies of insect herbivory on fossil leaves can provide key information for an ecological understanding of disturbance and biotic response in deep time, such as the response of insect-damage frequency and diversity to changing vegetation and climate. In this contribution, I provide the first, high-resolution study documenting insect damage of fossil leaves that indicate the reaction of insect herbivores to changing regional climates and vegetation during the latest Paleogene in Europe. Insect damage censuses were conducted at six stratigraphic levels ranging in age from 27 to 23 Ma in the Siebengebirge area near Bonn, Germany. A total of 3122 fossil angiosperm leaves pertaining to 135 species were examined for the presence or absence of insect damage types (DTs); 59 damage types were recorded. These results provide the first data on the representation of insect herbivory in the Tertiary of Germany that describe temporal trends in herbivory, and correlate these trends to external variables such as floral diversity and temperature. The most parsimonious explanation for the trends observed in this study is that the fossil insect damage represents a regional response to global environmental changes.
Abstract: The Palaeontinidae (Insecta: Cicadomorpha) from the Upper Jurassic Solnhofen Limestone of Bavaria are revised. The diagnostic characters for three monotypic genera Eocicada Oppenheim, 1888, Prolystra Oppenheim, 1888 and Archipsyche Handlirsch, 1906 are reassessed based on newly discovered material. Beloptesis gigantea (Weyenbergh, 1874), B. oppenheimi Handlirsch, 1906, Limacodites mesozoicus Handlirsch, 1906, and Protopsyche braueri Handlirsch, 1906 are considered to be junior synonyms of Prolystra lithographica Oppenheim, 1888. Eocicada lameerei Handlirsch, 1908 is a junior synonym of E. microcephala Oppenheim, 1888. A key to the species of Solnhofen Palaeontinidae is presented. Solnhofen Palaeontinidae and most Cretaceous Palaeontinidae most probably form a monophyletic group based on the following characters: costal area narrow, vein RA branching from stem R basally, vein ScP not fused with vein RA, clavus much reduced and hindwing smaller. Furthermore, Solnhofen Palaeontinidae are probably basal to Cretaceous Palaeontinidae by the mesonotum lacking distinct longitudinal carinae. A fast succession from early to more derived Palaeontinidae took place during Late Jurassic times. Early Palaeontinidae declined sharply in the Late Jurassic, probably owing to the rise of newly evolved insectivorous animals like early birds and mammals. Late Palaeontinidae with better flight ability survived and became a dominant insect group during latest Jurassic times.
Abstract: Fossilized leaf mines and other traces of phytophagous insects provide a unique window into ecological and evolutionary associations of the past. Leaf-mining flies (Diptera: Agromyzidae) are an important component of the recent leaf-mining fauna, but their fossil record is sparse compared to other mining insect lineages; many putative agromyzid body fossils and traces are dubiously assigned. Agromyzid leaf mines often can be distinguished from those of other insects by the presence of an intermittent, fluidized frass trail that may alternate between the sides of the mine. Here, we describe two new Paleogene leaf mine fossils, Phytomyzites biliapchaensis Winkler, Labandeira and Wilf n. sp. from the early Paleocene of southeastern Montana, USA, occurring in leaves of Platanus raynoldsi (Platanaceae); and Phytomyzites schaarschmidti Wappler n. sp., from the middle Eocene of Messel, Germany, occurring in leaves of Toddalia ovata (Rutaceae). These fossils both exhibit frass trails indicative of an agromyzid origin, and P. biliapchaensis also exhibits associated stereotypical marks identical to damage caused by feeding punctures of extant adult female Agromyzidae prior to oviposition. Phytomyzites biliapchaensis represents the earliest confirmed record of Agromyzidae, and one of the earliest records for the large dipteran clade Schizophora. Plant hosts of both species belong to families that are no longer hosts of leaf-mining Agromyzidae, suggesting a complex and dynamic history of early host-plant associations and, for the early Paleocene example, an evolutionary, possibly opportunistic colonization in the midst of the ecological chaos following the end-Cretaceous event in North America.
Abstract: A new genus and species of Trogossitidae (Lophocaterinae: Lophocaterini), Promanodes serafini, is described from Baltic amber. The fossil is considered related to three species of the extant Promanus Sharp, 1877 from New Zealand. It differs from the latter especially in larger postcoxal projection and coniform maxillary palpi. A close morphological similarity between recent and fossil representatives of the both genera allows to suppose predaceous way of life of the new Eocene species. All extant species of the genus Promanus are reviewed, their types illustrated and a differential diagnosis given. Biogeography of Promanus is discussed.
Abstract: Terrestrial plants and insects currently account for the majority of the Earth's biodiversity, and approximately half of insect species are herbivores. Thus, insects and plants share ancient associations date back for more than 400 million years. However, investigation of their past interactions are at the preliminary stages in Western Europe. Herein we present first results of study various feeding damages based on dataset counting of nearly 3500 examined plant specimens from Lower Miocene of the Lagerstätte BÃlina Mine in Most Basin, Czech Republic. This site provides unique view into the Neogene freshwater ecosystems. It was longtime studied by scientists working in different branches of sedimentology, palaeobotany and palaeozoology. The fossils are preserved in three characteristic horizons overlaying the coal-seam (Clayey Superseam Horizon, Delta Sandy Horizon, Lake Clayey Horizon) reflecting palaeoenvironmental changes in short time period of development. The trace fossils are classified as functional feeding-groups or "guilds" without searching for direct causer or recent analogue host relation. About 23% of specimens of dicotyledonous plant leaves were found damaged and associated with some leaf âmorphotypesâ. Deciduous plantâhost taxa and a those with chartaceous texture typical of riparian habitats were frequently damaged, such as Populus recorded with two species P. zaddachii and P. populina show 57.9% and 31.0% levels for herbivory, respectively, followed by the Acer, Alnus, and Carya averaging almost 30% of damaged leaves/leaflets. Evidence of 60 damage types representing all functional feeding groups recorded at the BÃlina Mine including 12 types of leaf mines and 16 gall type DTs. In total it exhibit a high level of external foliage feeding types (23.7%), and a low level of more specialized damage types, such as galls (4.3%) and leaf mines (<1%). Broader comparison based on damage types of the main sedimentary environments shows significancy supporting different biomes by frequency of damage levels and DT diversities.
Abstract: To mark the occasion of the 175th anniversary of the Rheinische Naturforschende Gesellschaft in 2009 and of the centennial of the Mainz Natural History Museum in 2010 we present a short account of our present knowledge of the Eckfeld Maar after 20 years of continuous research. This paper does not attempt to include all of the detailed results on the geology of the Eckfeld site or its biota. To date, nearly 250 papers and books have been published since the start of our project. An up-to-date list of these publications can be found at www.eckfelder-maar.de.
Abstract: Aside from pollen and nectar, bees of the subfamily Megachilinae are closely associated with plants as a source of materials for nest construction. Megachilines use resins, masticated leaves, trichomes and other plant materials sometimes along with mud to construct nests in cavities or in soil. Among these, the leafcutter bees (Megachile s.l.) are the most famous for their behaviour to line their brood cells with discs cut from various plants. We report on fossil records of one body fossil of a new nonleafcutting megachiline and of 12 leafcuttings from three European sitesâEckfeld and Messel, both in Germany (Eocene), and Menat, France (Paleocene). The excisions include the currently earliest record of probable Megachile activity and suggest the presence of such bees in the Paleocene European fauna. Comparison with extant leafcuttings permits the interpretation of a minimal number of species that produced these excisions. The wide range of size for the leafcuttings indirectly might suggest at least two species of Megachile for the fauna of Messel in addition to the other megachiline bee described here. The presence of several cuttings on most leaves from Eckfeld implies that the preferential foraging behaviour of extant Megachile arose early in megachiline evolution. These results demonstrate that combined investigation of body and trace fossils complement each other in understanding past biodiversity, the latter permitting the detection of taxa not otherwise directly sampled and inferences on behavioural evolution.
Abstract: A new fossil bark-gnawing beetle, Ancyrona eocenica Schmied, Wappler, KolibáÄ, new species (Coleoptera: Cleroidea: Trogossitidae), is described and figured from a specimen preserved in Eocene limnic sediments of the Eckfeld Maar, Germany. Ancyrona eocenica sp. nov. (Peltinae: Ancyronini) is the first trogossitid described from the Eckfeld Maar as well as the first fossil ever of this genus. The species is distinguished from recent congeners which are distributed in Ethiopian and Oriental regions by the following: minute body (2.6 mm), anterior margin of pronotum weakly concave, body broadly oval. A taxonomic catalogue of previously described fossil bark-gnawing beetles has been complied and is given here.
Abstract: The fossil history of the family Geinitziidae is reviewed. New taxa are described in the family: Shurabia hissarica, sp. nov. (Lower Jurassic of Tajikistan), Sh. shartegica, sp. nov. (Upper Jurassic of Mongolia), Sh. serrata, sp. nov. and Ginitzia sagulensis, sp. nov. (both Lower Jurassic of Kyrgyzstan). The subfamily Stegopterinae Sharov, 1961 is synonymized under Geinitziinae Handlirsch, 1906. The genera Minesedes Fujiyama, 1973 and Ominea Fujiyama, 1973 (Carnian of Japan) are synonymized under Shurabia Martynov, 1937 and Geinitzia Handlirsch, 1906, respectively. Geinitzia varia Bode, 1953 (Lower Jurassic of Germany) and Fletchizia picturata Riek, 1976 (Upper Triassic of South Africa) are redescribed from their holotypes. F. kapokraalensis Wappler, 2001 and F. aleda Wappler, 2001 (both Upper Triassic of South Africa) are transferred to the genus Shurabia.
Abstract: Insect herbivores are considered vulnerable to extinctions of their plant hosts. Previous studies of insect-damaged fossil leaves in the US Western Interior showed major plant and insect herbivore extinction at the Cretaceous â Palaeogene (K â T) boundary. Further, the regional plant â insect system remained depressed or ecologically unbalanced throughout the Palaeocene. Whereas Cretaceous ï¬oras had high plant and insect-feeding diversity, all Palaeocene assemblages to date had low richness of plants, insect feeding or both. Here, we use leaf fossils from the middle Palaeocene Menat site, France, which has the oldest well-preserved leaf assemblage from the Palaeocene of Europe, to test the generality of the observed Palaeocene US pattern. Surprisingly, Menat combines high ï¬oral diversity with high insect activity, making it the ï¬rst obser vation of a âhealthyâ Palaeocene plant â insect system. Furthermore, rich and abundant leaf mines across plant species indicate well-developed host specialization. The diversity and complexity of plant â insect interactions at Menat suggest that the net effects of the K â T extinction were less at this greater distance from the Chicxulub, Mexico, impact site. Along with the available data from other regions, our results show that the end-Cretaceous event did not cause a uniform, long-lasting depression of global terrestrial ecosystems. Rather, it gave rise to varying regional patterns of ecological collapse and recover y that appear to have been strongly inï¬uenced by distance from the Chicxulub structure.
Abstract: Polyrhachis annosus n. sp., based on part and counterpart of a unique ant worker specimen from Late Miocene deposits of the island of Crete (Greece), is figured and described here. Features of P. annosus unique for the large recent genus Polyrhachis include a large first gastral segment, a scapus attached at some distance from the clypeus, and an alitrunk (mesosoma) and petiole each with a pair of teeth or spines. The new taxon represents the first occurrence of the genus in the fossil record. The origin and rise of one of the worldâs largest and most distinct ant genera are still a puzzle, and closely related genera are quite diverse in Tertiary amber deposits. Interestingly, the abundance of Polyrhachis in Indonesian copal confirms the statement that their absence in Baltic amber is not casual.
Abstract: To mark the occasion of the 175th anniversary of the Rheinische Naturforschende Gesellschaft in 2009, and of the centennial of the Naturhistorisches Museum Mainz in 2010, we present a preliminary account of our present know-ledge of the Eckfeld Maar after 20 years of continuous research. To date, nearly 250 papers and books have been published since the start of our excavations. This paper does not attempt to include all of the detailed results on the geology of the Eckfeld site or its biota, but an up-to-date list of publications can be found under the address www.eckfelder-maar.de.
Abstract: Of the 30 extant orders of insects, scale insects (Hemiptera: Coccoidea) have one of the best fossil records among insects that probably extends to the early Mesozoic. Most fossil Coccoidea records are of adult males which have been trapped in amber, whereas adult females are less common, probably because of their sedentary occurrence on plants. Descriptions, photos, and SEM micrographs are presented of over 250 wellâpreserved scale covers of adult females and nymphal stages of armoured scale insects (Hemiptera: Coccoidea: Diaspididae) which were found on dicotyledonous and monocotyledonous fossil leaves in Middle Eocene deposits from Germany. The structure of these fossil scale covers closely agrees with that of extant Diaspididae. These armoured scale insects belong to the subfamily Aspidiotinae. The age of this insectâplant associationâ47 to 44 millionâyearâoldâindicates that the Diaspididae have a long history in the Palaearctic region.
Abstract: Two new species of weaver ant are described from the Eocene of Germany. Males and gynes of Oecophylla longiceps
Dlussky sp. nov. are found in the middle Eocene (about 47 Ma) of Grube Messel, Germany. Males, gynes and two workers of O. eckfeldiana Dlussky sp. nov. are recorded from the middle Eocene (about 43 Ma) of Eckfeld maar, Germany. The two new species are among the oldest records of the extant genus Oecophylla. First adaptations for moving in the forest canopy are present in the workers of O. eckfeldiana. Even more specialized adaptations for arboreal life are found in the workers of O. brischkei from Baltic amber. The coexistence of two species in Baltic amber and in the Bembridge marls suggests that in these cases different ecological niches were realised by sympatric species. Comparisons of the queens from different fossil and extant species indicate morphological trends, partly connected with increasing fertility. Most likely Oecophylla originated in the early Paleogene in the Palaearctic realm, radiating strongly during the climatic changes of the EoceneâOligocene transition.
Abstract: Terrestrial plants and insects today account for the majority of the Earth's biodiversity, and almost half of all insect species are herbivores. Therefore, the ecology of plant-insect associations is a significant aspect of modern ecological research today. The high species diversity has been attributed to several factors, including the small size of insects resulting in many more exploitable niches. Also, insects have a high genetic heterogeneity or elasticity which may promote speciation in the face of environmental change. Increasingly it is argued that the interactions with other organisms such as plants is also itself a major reason for its great diversity.
The field of insect-plant interactions is overwhelmingly interesting because of the enormous diversity of both groups, as well as their apparently intimate relationships. Its complexity is already evident from the sheer numbers of species. The high degree of food specialization of most herbivorous insects, whereby even so-called generalists show distinct and refined food preferences, indicating the geologically long-term persistence of some of these associations.
Fossil leaves are extremely sensitive indicators of past climates, evolution of plant-insect interactions, development of biodiversity, and the effects of major environmental disturbances. In ongoing studies granted by the German Science Foundation from the abundant, diverse, and well-preserved insect-feeding damage record found on fossil leaves from different Paleogene European localities provide for the first time deep insights in these patterns from a time period immediately before the Paleocene Eocene Thermal Maximum (PETM) and after the middle Eocene Climatic Optimum towards the Late Oligocene. The Paleogene is characterized by cycles of cooling and warming superimposed on the broad global trend of progressive cooling, and thus provides a record of deep time climate change. The inventory of insect damage occurring on plant hosts indicates that an impressive variety of plant-insect associations were present during Paleogene times. The analysis of the European floras show a greater diversity of insect feeding than in contemporaneous, comparable floras from North America, revealing another important, previously unknown component of high diversity in the European Paleogene. The results further indicate a high diversity of damage types in particular for highly specialized feeding types like mining and galling. This may be tied to the fact that host specificity characterized most known lineages of gallers and miners to a high degree.
Abstract: Three fossil taxa of megapodagrionid damselflies are described and figured from the Paleogene localities in Europe on the basis of isolated wings. Eckfeldia superstes (Wappler, 2003) gen. nov. is described from the laminated mudstones of middle Eocene age from Eckfeld Maar, Germany. Furagrion jutlandicus (Henriksen, 1922) gen. nov. is recorded from the laminated claystones of lowermost Eocene age from the Ãlst and Fur-Formation, Denmark, and an undetermined megapodagrionid damselfly is recognized from middle Eocene strata. Taphonomy and color preservation in the fossils are briefly considered. Characters used for phylogenetic analyses in extant and fossil Megapodagrionidae are discussed. The biogeographic and paleoecological implications of the new European fossils are briefly discussed.
Abstract: Koevolutive Beziehungen zwischen Pflanzen und Insekten werden seit Jahren als wichtigster âMotorâ für den enormen Artenreichtum heutiger terrestrischer Ãkosysteme angesehen. Dabei steht die Erforschung ihrer Wechselbeziehungen aber noch weitgehend am Anfang. Untersuchungen zur Dynamik von Insekten-Pflanzen Interaktionen in erdgeschichtlichen Zeiträumen anhand fossilen Pflanzenmaterials wurden bislang nur für einige nordamerikanische Fundstellen durchgeführt. Hier bieten die eozänen Fossillagerstätten Messel und Eckfeld wegen ihres Artenreichtums und ihrer hervorragenden Erhaltungsqualität von Pflanzenfossilien und FraÃspuren günstigste Voraussetzungen. Nahezu 40.000 Pflanzenreste stehen für die Untersuchungen zur Verfügung, von denen nach einer ersten Durchsicht 15-20% Schäden wie BlattfraÃ, Minen oder Gallen aufweisen.
Die Vielfalt dieser Beschädigungen, die als sogenannte âGildenâ bzw. funktionale Nahrungsgruppen klassifiziert werden, liefert ein hervorragendes Beispiel für die Komplexität, die Struktur und den Wandel von terrestrischen Ãkosystemen im Bereich des eozänen Klimaoptimums.
Für manche Insektengruppen sind solche indirekten Nachweise besonders bedeutsam, da ihre fragilen, dünngepanzerten oder auch extrem kleinen Larven- und auch Adultstadien fossil nur schlecht oder gar nicht erhalten sind.
In diesem Zusammenhang sind vor allem Vertreter der âMikrolepidopteraâ zu nennen, die sich fossil bislang in beiden Fundstellen nicht nachweisen lieÃen. Aufgrund der durchaus diversen Minentypen, die von gewundenen Minen, flächigen Platzminen sowie ober- und unterseitigen Minen reichen, zeigt sich, dass diese Tiergruppe in Messel mit mindestens 4-5 unterschiedlichen Familien verbreitet gewesen ist. Für das Eckfelder Maar hingegen ist die Anzahl an Minentypen im Vergleich zu Messel relativ gering, wohingegen die Diversität an Pflanzengallen prozentual erhöht zu sein scheint. Inwieweit dies eine allmähliche lokale oder gar überregionale Klimaveränderung widerspiegelt, müssen weitere Untersuchungen zeigen.
Die Ergebnisse sollen im Vergleich mit Fundstellen Nordamerikas und im Hinblick auf moderne Theorien zur Entwicklung der Biodiversität ausgewertet werden.
Dieses von der DFG geförderte Projekt ist die erste umfangreiche Untersuchung von Pflanzen-Insekten Interaktionen für zwei der bedeutendsten terrestrischen Fossillagerstätten Europas.
Abstract: We describe Priscalestes germanica PetruleviÄius & Wappler, a new genus and species of Lestinoidea Calvert (1901) (sensu Bechly 1996) from the Eocene of Germany. The new genus represents a new family, Priscalestidae PetruleviÄius & Wappler fam. nov., with close relationship to Megalestidae, Lestidae and the genus Promegalestes PetruleviÄius & Nel 2004 from the late Paleocene of Argentina. The new family seems to be in a basal position with respect to the Lestidae because of the lack of their synapomorphies, i. e. MA strongly zigzagged and the area between IR2 and RP3/4 distally strongly widened with three rows of cells between these two veins instead of only one. The new family differs also from Lestidae and Megalestidae in the presence of two autapomorphies, i.e. (1) all secondary longitudinal veins (except IR1 and IR2) suppressed, resulting in presence of unicellular rows between IR1 and RP2, RP2 and IR2, RP3 and MA; and (2) the midfork closer to the subnodus than to the arculus. The sharing of last character with Promegalestes let us thinking in a close relationship of these two genera.
Abstract: Heteroptera is an insect group that has been successful since a long time. First fossils are known from the Permian period. Actually, true bugs comprise seven subgroups. A short review of the fossil history of these subgroups is given. Additionally, fossil bug faunas of several important European Fossillager-stätten are presented. The lower Eocene Moler (Denmark) is characterized by a diverse Gerromor-phan fauna, while fossils of other bug taxa are very abundant, but not as diverse. In the middle Eo-cene fossil site Messel pit the Heteroptera represent a quite large portion of the insects, while true bugs are significantly rarer in the middle Eocene Eckfeld maar and in Enspel (both Germany).
Abstract: A new genus of cantacaderid lace bugs, Lutetiacader gen. et sp. nov., is described from the lower Middle Eocene of Messel Quarry, Germany. It clearly belongs to the cantacaderid lineage of Tingidae, and is the second known fossil genus of the tribe Cantacaderini (after Paleocader Froeschner). It appears to be most similar to extant species of the genera Nectocader Drake (Brazil) and Teratocader Drake (Malay Peninsula). Lutetiacader occupies a central position with respect to the geographical gap between the ranges of these two extant genera. Its biogeographical significance is discussed.
Abstract: Three new genera of extinct Dictyopharidae, Lophopidae and Eurybrachyidae respectively are described. Wedelphus gen. nov. with Wedelphus dichopteroides sp. nov. placed in Dictyopharidae, Baninus gen. nov. with Baninus thuringiorum sp. nov. of the family Lophopidae and Amalaberga gen. nov. with Amalaberga ostrogothiorum sp. nov. placed in Eurybrachidae (first fossil record of the family) are described, all from deposits of the Grube Messel Lagerstätte in Germany. A representative of an unrecognised family, possibly related to Henriksenopterix PetruleviÄius, 2005 is reported. The oil shales of the Messel maar in Hessen are well known for their extremely rich fossil flora and fauna. They are of Lower Middle Eocene age (about 48 million years) and contain a highly diverse insect fauna.
Abstract: The oil shales of the famous Messel maar in Hessen, Germany are well known for their extremely rich fossil flora and fauna. They are of Lower Middle Eocene age (about 48 million years) and contain also a highly diverse insect fauna. In addition to previous publications on some taxa belonging to the Heteropteran infraorders Pentatomomorpha, Cimicomorpha and Gerromorpha, three species of the Pentatomomorphan family Aradidae (flat bugs) are described and figured : Neuroctenus kotejai sp. n., Neuroctenus messelensis sp. n. and Mezira eocenica sp. n.
Abstract: Neuroctenus enspelensis n. sp., a new fossil flat bug species of the family Aradidae, subfamily Mezirinae from the Upper Oligocene lacustrine shales of Enspel, Germany is described and figured.
Abstract: In addition to the description of the first three fossil species of Aradidae from limnic sediments from the Messel maar in Germany (WAPPLER & HEISS in press), three more species from the Eckfeld maar: Mezira eckfeldensis n. sp., Mezira crassifemur n. sp. and Neuroctenus incompletus n. sp. are described and figured. This fossil deposit is dated as Middle Eocene (about 44 million years) and thus somewhat younger than that of Messel with an age of about 47 million years. The taxa are congeneric but differences in habitus, structure of head, antennae, pronotum and legs indicates that they belong to other undescribed species.
Abstract: A new species of coleseed sawfly (Tenthredinidae: Athaliini) is described and figured from the Eocene-Oligocene âRebbergâ quarry within the Fossilliferous Zone, a member of the Middle Salt Formation. Athalia vetuecclesiae n. sp. is the first representative of the Athalia group from the geological record. The new species is most similar to members of the A. vollenhoveni-group represented by only 6 species from Africa but can be distinguished by details of antennal structure and length of spurs of the hind tibia. The phylogenetic position of the fossil within the Tenthredinidae*, its paleoenvironmental implication, and the geological setting of the quarry is briefly discussed.
Abstract: The middle Eocene (Lutetian Stage) earwig fauna of the Eckfeld maar, Germany is reviewed and three species are reported â two are undeterminable as to family, the third is representative of a new species of Forficulidae. Forficula kotejai sp. n., is referred to this group of European earwigs on the basis of comparisons with extant and extinct specimens. A brief summary of the fossil history of Dermaptera is provided and a catalog of fossil earwigs appended.
Abstract: Only very few specimens of dictyopharid plant-hoppers are known from the fossil record. For an additional specimen which was recovered from the fossil site Grube Messel (Lower Middle Eocene) near Darmstadt a detailed description and figures are given. Judging from characters of the wing venation and head capsule it is described under open nomenclature as âDictyopharinae gen. et sp. indet.â. Due to the lack of sufficient information it is not possible to determine it on tribus or genus level.
Abstract: Water striders (Hemiptera, Gerromorpha) are a very distinct ecological group specially adapted for life on the water surface. The present paper reports on four species of Gerromorpha from the Middle Eocene fossil sites of Eckfeld and Messel describing Lutetiabates eckfeldensis n.gen. et n.sp., Cylindrobates messelensis n.gen. et n.sp. (both Gerridae), and two nymphs most probably of the genus Gerris. The record of two new members of Gerridae from the Paleogene as weil as the hitherto known Gerromorpha from fossiliferous resins document a distinctly higher diversity of water striders within die European Paleogene than today. Lastly, comments are made on the fossil history as weil as on the palaeobiological and palaeobiogeographical significance of the faunas.
Abstract: Out of the 30 extant orders of insects, all but one, the parasitic lice (Insecta: Phthiraptera), have a confirmed fossil record. Here, we report the discovery of what appears to be the first bird louse fossil: an exceptionally well-preserved specimen collected from the crater of the Eckfeld maar near Manderscheid, Germany. The 44-million-year-old specimen shows close phylogenetic affinities with modern feather louse ectoparasites of aquatic birds. Preservation of feather remnants in the specimen's foregut confirms its association as a bird ectoparasite. Based on a phylogenetic analysis of the specimen and palaeoecological data, we suggest that this louse was the parasite of a large ancestor to modern Anseriformes (swans, geese and ducks) or Charadriiformes (shorebirds). The crown group position of this fossil in the phylogeny of lice confirms the group's long coevolutionary history with birds and points to an early origin for lice, perhaps inherited from early-feathered theropod dinosaurs.
Abstract: The rich insect taphocoenosis from the Middle Eocene lacustrine deposits of the Eckfelder Maar is compreÂhensively presented for the first time. Comparison between this fossil record and the insect taphocoenoses from Messel and the Geiseltal permit detailed insights into Middle Eocene paleoecosystems, the last epoch with a worldwide greenhouse climate. Furthermore, this study permits a rather extensive reconstruction of this now exÂtinct ecosystem. The results are in congruence with other data, for example, from sedimentological and paleobotanical researches.
In total, the Eckfelder Maar insect taphocoenosis contains 4617 fossil specimens. Most of them are in good condition with a potential for exceptional details of preservation. This fossil record documents a highly diverse terrestrial fauna and flora, while aquatic life is rather poor in species. The insect taphocoenosis is predominantly composed of Coleoptera (84 %). They consist of 21 families among which the Curculionoidea (Curculionidae and Brentidae) is the dominant group, comprising 40 % of all specimens. In second place are Chrysomelidae which are quite common. Elateridae, Scarabaeidae, Buprestidae, and Tenebrionidae are also frequently found. Odonata are extremely rare, represented only by isolated wings but in one instance preserved with the original wing pattern. Dermaptera, Blattaria, Isoptera, Diptera, Plecoptera, and Phthiraptera are represented by only a few fossils. Much more common are Auchenorryncha and Heteroptera. Trichoptera are recorded from the Eckfelder Maar by four different types of portable tubecases, body fossils are absent. Main research has focused on Hymenoptera which show a great diversity in Apoidea (bees) and Formicidae (ants), the latter exclusively repreÂsented by winged reproductives (male and female). The new material from Eckfeld is in such excellent condition that bees could completely be revised.
Analysis of the giant ants from the extinct subfamily Formiciinae, known only from Eckfeld and Messel by complete specimens, shows that the intraspecific variability during the Geiseltalian time period (4-5 Ma) is negligible. Specimens from both localities are probably conspecific. Similar results could be proposed for represenÂtatives of Elateridae, Buprestidae, and Chrysomelidae. In these groups, as well as in some others, the original structural colouring is preserved, comparable with taxa from Messel and Geiseltal.
Altogether there are numerous xylophagous and parasitic insect taxa present. They indicate densely wooded surroundings, with a highly complex paleoecosystem. The disposal of heteropteran taxa over the digging profile clearly shows that some kind of temporary xerothermic habitats (e.g., small, restricted temporary areas without vegetation) were also present close to the maar.
Abstract: The rich insect taphocoenosis from the Middle Eocene lacustrine deposits of the Eckfelder Maar is compreÂhensively presented for the first time. Comparison between this fossil record and the insect taphocoenoses from Messel and the Geiseltal permit detailed insights into Middle Eocene paleoecosystems, the last epoch with a worldwide greenhouse climate. Furthermore, this study permits a rather extensive reconstruction of this now exÂtinct ecosystem. The results are in congruence with other data, for example, from sedimentological and paleobotanical researches.
In total, the Eckfelder Maar insect taphocoenosis contains 4617 fossil specimens. Most of them are in good condition with a potential for exceptional details of preservation. This fossil record documents a highly diverse terrestrial fauna and flora, while aquatic life is rather poor in species. The insect taphocoenosis is predominantly composed of Coleoptera (84 %). They consist of 21 families among which the Curculionoidea (Curculionidae and Brentidae) is the dominant group, comprising 40 % of all specimens. In second place are Chrysomelidae which are quite common. Elateridae, Scarabaeidae, Buprestidae, and Tenebrionidae are also frequently found. Odonata are extremely rare, represented only by isolated wings but in one instance preserved with the original wing pattern. Dermaptera, Blattaria, Isoptera, Diptera, Plecoptera, and Phthiraptera are represented by only a few fossils. Much more common are Auchenorryncha and Heteroptera. Trichoptera are recorded from the Eckfelder Maar by four different types of portable tubecases, body fossils are absent. Main research has focused on Hymenoptera which show a great diversity in Apoidea (bees) and Formicidae (ants), the latter exclusively repreÂsented by winged reproductives (male and female). The new material from Eckfeld is in such excellent condition that bees could completely be revised.
Analysis of the giant ants from the extinct subfamily Formiciinae, known only from Eckfeld and Messel by complete specimens, shows that the intraspecific variability during the Geiseltalian time period (4-5 Ma) is negligible. Specimens from both localities are probably conspecific. Similar results could be proposed for represenÂtatives of Elateridae, Buprestidae, and Chrysomelidae. In these groups, as well as in some others, the original structural colouring is preserved, comparable with taxa from Messel and Geiseltal.
Altogether there are numerous xylophagous and parasitic insect taxa present. They indicate densely wooded surroundings, with a highly complex paleoecosystem. The disposal of heteropteran taxa over the digging profile clearly shows that some kind of temporary xerothermic habitats (e.g., small, restricted temporary areas without vegetation) were also present close to the maar.
Abstract: The Middle Eocene (Lutetian) bee faunas of Eckfeld and Messel, Germany are revised. In addition to the previously known Electrapis electrapoides (Lutz), five additional species are recognized. Four new species of the Electrapini (Apidae: Apinae) are described: Electrapis micheneri Wappler and Engel, E. prolata Engel and Wappler, Protobombus pristinus Wappler and Engel, and P. messelensis Engel and Wappler. In addition, the new genus Pygomelissa Engel and Wappler is proposed for Pygomelissa lutetia Engel and Wappler new species, which cannot presently be classified into any tribe of the Apidae. The tribe Megachilini (Megachilidae: Megachilinae) is also recorded from Eckfeld but in the absence of any body fossil. Megachilines include the leaf-cutter bees (Megachile) and from the occurrence of the distinctive semicircular damage they produce in leaves, we conclude that such bees were also present in the fauna. The bee fauna is compared with that of the contemporaneous Baltic amber. As with Baltic amber, the majority of bee specimens are from the advanced eusocial lineages of the corbiculate Apinae. Lastly, comments are made on the phylogenetic and paleobiological significance of the faunas.
Abstract: The rich insect taphocoenosis from the Middle Eocene lacustrine deposits of the Eckfelder Maar is compreÂhensively presented for the first time. Comparison between this fossil record and the insect taphocoenoses from Messel and the Geiseltal permit detailed insights into Middle Eocene paleoecosystems, the last epoch with a worldwide greenhouse climate. Furthermore, this study permits a rather extensive reconstruction of this now exÂtinct ecosystem. The results are in congruence with other data, for example, from sedimentological and paleobotanical researches.
In total, the Eckfelder Maar insect taphocoenosis contains 4617 fossil specimens. Most of them are in good condition with a potential for exceptional details of preservation. This fossil record documents a highly diverse terrestrial fauna and flora, while aquatic life is rather poor in species. The insect taphocoenosis is predominantly composed of Coleoptera (84 %). They consist of 21 families among which the Curculionoidea (Curculionidae and Brentidae) is the dominant group, comprising 40 % of all specimens. In second place are Chrysomelidae which are quite common. Elateridae, Scarabaeidae, Buprestidae, and Tenebrionidae are also frequently found. Odonata are extremely rare, represented only by isolated wings but in one instance preserved with the original wing pattern. Dermaptera, Blattaria, Isoptera, Diptera, Plecoptera, and Phthiraptera are represented by only a few fossils. Much more common are Auchenorryncha and Heteroptera. Trichoptera are recorded from the Eckfelder Maar by four different types of portable tubecases, body fossils are absent. Main research has focused on Hymenoptera which show a great diversity in Apoidea (bees) and Formicidae (ants), the latter exclusively repreÂsented by winged reproductives (male and female). The new material from Eckfeld is in such excellent condition that bees could completely be revised.
Analysis of the giant ants from the extinct subfamily Formiciinae, known only from Eckfeld and Messel by complete specimens, shows that the intraspecific variability during the Geiseltalian time period (4-5 Ma) is negligible. Specimens from both localities are probably conspecific. Similar results could be proposed for represenÂtatives of Elateridae, Buprestidae, and Chrysomelidae. In these groups, as well as in some others, the original structural colouring is preserved, comparable with taxa from Messel and Geiseltal.
Altogether there are numerous xylophagous and parasitic insect taxa present. They indicate densely wooded surroundings, with a highly complex paleoecosystem. The disposal of heteropteran taxa over the digging profile clearly shows that some kind of temporary xerothermic habitats (e.g., small, restricted temporary areas without vegetation) were also present close to the maar.
The systematic revision of the Eckfelder taphocoenosis reveals that most of the taxa are generally allied with lineages that are now inhabiting tropical or subtropical climates. Taxa with a Palearctic or Holarctic distribution are less common. The main paleobiogeographic pattern shows strong relationships to faunas from South America and SE-Asia, i.e. continents previously allocated to Gondwana.
The Apinae show a strong relationship to groups currently known from the roughly contemporaneous and geographically near Baltic amber. This also applies to some other insect taxa (e.g., Carabidae: Paussinae and Gerromorpha).
Taphonomic aspects are investigated over the whole section of the digging site. Besides the dominant process of selectivity during the post-mortem drifting on the water surface, effects of water density (e.g., changes of the electrolyte concentration in the monimolimnion), and water level changes determine the composition of the asÂsemblages of fossil insects. These investigations are supported by carbon and oxygen isotopes of authigen-formed siderite which are useful proxies for paleolimnologic models. The main trend in the corresponding profile segments clearly show that the curve for the siderite content correlates positively with the relative abundance of Coleoptera.
The Eckfelder Maar insect taphocoenosis shows no striking evidence for a cooling at the end of the Middle Eocene. By contrast, the fossils indicate a paratropical rainforest community, similar to recent ones in SE-Asia, and referring to the generally warm climate conditions prevailing in central Europe during the early Paleogene (PETM = Paleocene-Eocene Thermal Maximum).
