Two new species of European Microgaster Latreille, 1804 (Hymenoptera: Braconidae, Microgastrinae), with host data on some further species

1) A survey of cytological, genetical, and other evidence for phylogenetic differentiation of higher plant groups in Mediterranean and Central European areas is presented.2) Methods and possible results are demonstrated, using as an example the closely related Rubiaceous genera Cruciata and Valantia, where polyploidy, descending dysploidy, increase of heterochromatin, change from allogamy to autogamy, and from perennials to annuals suggest salient features of geographical and ecological differentiation.3) The phylogeny of more than 70 other Europe groups of Ferns and Angiosperms from the Mediterranean and Central is briefly discussed in chapters on woody plants, perennials of forest, alpine, and other open habitats, and annuals.4) The sequence from woody plants to annuals is on the average paralleled by increasing intensity of cytogenetical differentiation (polyploidy, dysploidy, etc.) and evolutionary versatility (instead of evolutionary conservatism), and by decreasing age of groups. Furtheron, there are more Holarctic and Eurasian elements in the former, but more European, Mediterranean and Oriental elements in the later groups.5) In woody groups our examples suggest ancient affinities between relic Macaronesian resp. Colchic laurel forest elements and even more primitive Southeast‐Asian relatives (Prunus subg. Laurocerasus), origin of Mediterranean sclerophylls form laurel forest progenitors related to species in Macaronesia and East Asia (Viburnum sect. Tinus), and close affinities between Mediterranean evergreens and European summergreens (Rubus sect. Moriferi); these phenomena probably date back from the early to the later Tertiary.6) Examples from perennial groups stress the importance of Macaronesian, Mediterranean, and Oriental areas as centers for old and primary geographical differentiation (West/East etc.) as well as for ecological radiation from laurel, sclerophyll and temperate forest elements to those of open, warm and/or cold (alpine‐arctic) habitats, probably from Mid‐Tertiary onward (Polystichum setiferum‐P. lonchitis. Lotus corniculatus agg., Cyclamen purpurascens, Cruciata‐Valantia, Galium mollugo agg., Campanula subsect. Heterophylla, Brachypodium, Dactylis glomerata agg., etc.).7) The repeated origin of annual from perennial groups (Anagallis subg. Anagallis. Cruciata‐Valantia, Carthamus, Calendula, Crepis, Brachypodium, etc.) and their further differentiation was centered in Oriental, southeastern and southwestern Mediterranean areas, and seems to have been furthered by succsssive xerothermic periods since the later Tertiary.8) During the Pleistocene and its extreme climatic changes European floras became depauperate : Mediterranean areas then served as most important centers of conservation for many primitive progenitors of younger Central European elements. This has been demonstrated for numerous groups from forest (e. g. Polypodium vulgare agg., Paeonia, Rubus sect. Morveri. Lamiastrum, etc.), open (Pulsatilla vulgaris agg., Pinguicula sect. Pinguicula, Veronica hederifolia agg., Galium aparine agg., Anthoxanthum odoraturn agg., Dactylis glomerata agg., etc.), and even alpine habitats (Erysimum sylvestre agg., Poa badensis agg., etc.).9) During and after the glaciations of the Pleistocene and during the Holocene intensive secondary phylogenetic differentiation and speciation took place: In the Central Europeanvegetation and flora many of the depleted niches and lateron those newly created by man were filled predominantly from the South. Proof for this comes from dysploid series leading from Mediterranean to Central European representatives (Myosotis sylwatica, Leucojum, Carex sempervirens group, Crepis sect. Phytodesia, etc.), but more typically from numerous polyploid (end partly apomictic) complexes. Many of these polyploid Central European groups and species have originated through hybrid combinations from geographically and/or ecologically diverse diploids : from various (sub)mediterranean( ‐montane) progenitors (Rubus sect. Morgeri, Aphanes, Sorbus sect. Aria and sect. Torminaria, Paeonia, Dentaria, Galium pusillum agg., Cr uciata glabra, Knautia drymeia agg. and K. arvensis agg., Carthamus, Poaceae‐Honermeae, etc.), from further addition of genomes from eastern‐continental (Pulsatilla halleri agg., Lamiastrum, Veronica ser. Austriacae and V. hederifolia agg., etc.), boreal (Polypodium vulgare agg., Polystichum aculeatum agg., etc.), or montanealpine elements (Cerastium arvense, Arenaria serpyllifolia agg., Ceum subg. Geum, Lotus corniculatus agg., Tanacetum alpinum agg., Anthoxanthum odoratum agg., Poa annua agg., etc.), or even from several of such types (Potentilla werna agg., Thlaspi, Campanula sect. Heterophylla, Galium mollugo agg., Iris pumila, ete.).

A molecular phylogeny of the Sylvia cantillans complex: Cryptic species within the Mediterranean basin

Large parts of northern and central Europe were covered by ice sheets and permafrost due to climate changes in Europe during the last ice age (2.7 million to 11.7 kya). Plant and animal species had to adapt to lower temperatures, retreated to warmer areas in the south or went extinct. Once, after the Last Glacial Maximum (LGM, 26.5 kya to 19 kya) higher temperatures induced ice free habitats and these new habitats could be recolonized from different refugia. Collembola are one of the most abundant soil living decomposer animals and play a major role in aboveground - belowground interactions. Surprisingly, little is known about genetic and phylogeographic patterns, dispersal routes and anthropogenic influences of sexual and parthenogenetic reproducing European Collembola species, neglecting one important part of the global biodiversity, the belowground system. This thesis focuses on genetic patterns of four Collembola species with different reproductive modes and overlapping ecology across Europe. Collembola existed millions of years in stable habitats, as Eocene fossils show only little variation to extant taxa. In contrast, Cenozoic and Quaternary climatic changes reduced diversity and changed genetic structure of above living animals and plants. 
\nIn Chapter 2 I investigated the phylogeographic patterns of three common species of Collembola (Ceratophysella denticulata, Folsomia quadrioculata and Isotomiella minor) at a pan-European scale to identify glacial refuges and post-glacial colonization patterns with three genetic markers to cover different time scales. Results suggested density dependent processes for the establishment of new populations, as genetic diversity was high between but low within populations. This founder-takes-it-all principle is common in animal and plant species and suggests that only few early colonizing individuals founded the populations which grew and expanded rapidly. Arrival and invasion of other alleles into these populations was prevented by competition. Surprisingly and in contrast to the post-glacial recolonization patterns of aboveground organisms, the last ice age little affected the genetic composition of the studied Collembola species, indicating that soil provided habitat and resources for survival. The results show that divergence of populations took place during the Miocene (20-5 mya), when climatic conditions were favorable (warm and humid) for little sclerotized arthropods, susceptible to desiccation. Thus, the Miocene facilitated large scale expansion of European Collembola species. The results suggest that evolutionary processes of soil-living species are slowed-down, compared to above the ground living species, resulting in stable populations for millions of years.
\nIn chapter 3 I investigated differences in phylogeographic patterns due to different reproductive modes. As no partner is needed for reproduction, parthenogenesis provides a colonization advantage. To investigate the significance of reproductive modes for colonization, I compared the genetic structure of one sexual (Folsomia quadrioculata) and one parthenogenetic (Isotomiella minor) Collembola species with similar ecology across Europe, using one mitochondrial and two nuclear markers. Molecular variance was similar in both species and genetic differences were high between populations, indicating old diversifications. Northern and central Europe populations of I. minor were genetically homogenous suggesting that few lineages of this parthenogenetic species colonized these regions after LGM. Compared to I. minor the genetic structure of F. quadrioculata was more complex with more synonymous substitutions in protein coding genes. The results suggest that in addition to founder-effects and old diversifications, different forces affected sexual and parthenogenetic species, resulting in different phylogeographic patterns. In addition, mitonuclear compatibility among mating partners likely contributed to the more complex genetic structure in F. quadrioculata, whereas gene-environment interactions were of greater importance in I. minor. Overall, results indicate that the widespread view of central and northern European species being shaped by postglacial colonization patterns does not hold for both parthenogenetic and sexual soil-living species.
\nIn chapter 4 I investigated colonization patterns including cryptic diversity and the anthropogenic influence of the ubiquitous Collembola species Parisotoma notabilis in Europe. P. notabilis is the most widespread and abundant Collembola species in Europe colonizing many anthropogenic and disturbed habitats. Three molecular markers were used to investigate how anthropogenic factors and climate affected the present-day genetic structure of P. notabilis in Europe. The results showed that P. notabilis forms one morphologically coherent species comprising of several discrete genetic lineages. Molecular divergence estimates suggest that these lineages diverged in the Miocene during wet and warm climate, and a biome change in central and Eastern Europe from forest to grassland. The results further suggest that human activities favored the dispersal of P. notabilis as genetic lineages reflect human trading and migration routes such as the Channel and Mediterranean areas. These lineages are rather young and genetically uniform as compared to other soil-living arthropods.
\nOverall, the results suggest that phylogeographic patterns of soil-living European Collembola follow the ‘southern-richness and northern-purity’ scenario, but in contrast to this scenario, the pattern is not due to climate driven extinction of populations in central Europe during the Quaternary and recolonization thereafter. Rather, the pattern originated during the Miocene (20-5 mya) as indicated by divergence times of most clades. This indicates that the soil forms a stable habitat buffering climatic variability. Moreover, the results suggest that the dispersal of Collembola in Europe was affected by human activities. Overall, colonization in the Miocene and human activities in the Holocene resulted in a more complex genetic structure in soil-living species as compared to those living above the ground.

One. Introduction.- 1. On invading species and invaded ecosystems: the interplay of historical chance and biological necessity.- Two. Plant invasions.- 2. Plant invasions in Central Europe: historical and ecological aspects.- 3. History of the impact of man on the distribution of plant species.- 4. Recent plant invasions in the Circum-Mediterranean region.- 5. The invading weeds within the Western Mediterranean Basin.- 6. Widespread adventive plants in Catalonia.- 7. History and patterns of plant invasion in Northern Africa.- 8. Invasions of adventive plants in Israel.- 9. Man and vegetation in the Mediterranean area during the last 20,000 years.- 10. Plant invasions in Southern Europe from the Paleoecological point of view.- 11. Mediterranean weeds: exchanges of invasive plants between the five Mediterranean regions of the world.- Three. Animal invasions.- 12. The invasion of Northern Europe during the Pleistocene by Mediterranean species of Coleoptera.- 13. Migratory Phenomena in European animal species.- 14. The bean beetle (Acanthoscelides obtectus) and its host, the French bean (Phaseolus vulgaris): a two-way colonization story.- 15. Some recent bird invasions in Europe and the Mediterranean Basin.- 16. Of mice and men.- 17. Invasions by parasites in continental Europe.- 18. Human activities and modification of ichtyofauna of the Mediterranean sea: effect on parasitosis.- 19. Influence of environmental factors on the invasion of molluscs by parasites: with special reference to Europe.- Four. Mechanisms of invasions.- 20. In search of the characteristics of plant invaders.- 21. Biogeographical and physiological aspects of the invasion by Dittrichia (ex Inula) viscosa W. Greuter, a ruderal species in the Mediterranean Basin.- 22. Invaders and disequilibrium.- 23. Species-specific pollination: a help or a limitation to range extension?.- 24. Genetic differentiation in beech (Fagus sylvatica L.) during periods of invasion and regeneration.- 25. Invasion of natural pastures by a cultivated grass (Dactylis glomerata L.) in Galicia, Spain: process and consequence on plant-cattle interactions.- 26. Introduced and cultivated fleshy-fruited plants: consequences of a mutualistic Mediterranean plant-bird system.- 27. Fire as an agent of plant invasion? A case study in the French Mediterranean vegetation.- List of contributors.- Index of Genera and Species.- General index.

The phylogeography of Eurasian Fraxinus species reveals ancient transcontinental reticulation

The family Hexathelidae ranks among the smaller mygalomorph spider families. Most species are endemic to the Australasian region and the family was traditionally considered an example of a Gondwanan lineage. However, recent studies have cast some doubt on the monophyly of the family. Macrothele is the only genus with an out-of-Gondwana distribution. The bulk of the Macrothele diversity is found in South-east Asia, few species are known from central Africa and two species inhabit Europe: Macrothele calpeiana (Walckenaer, 1805) from the Iberian Peninsula and Macrothele cretica Kulczynski, 1903 endemic to Crete. Here we investigate the origins of the European Macrothele species by means of a multi-locus phylogenetic approach and by inferring the time frame of the diversification of the genus using Bayesian relaxed clock methods. We also provide further insights into the phylogenetic status of the family Hexathelidae. Our results indicate that the diversification of Macrothele traces back to the period of the Gondwana break-up and its present-day distribution most likely reflects the subsequent tectonic plate movements. The two European species were not recovered as sister taxa, suggesting that Macrothele colonised the Mediterranean region twice independently. The polyphyly of the family Hexathelidae is further confirmed and the subfamily Atracinae is identified as the conflicting lineage.

The Lonicera clade Caprifolium contains approximately 25 species distributed around the Northern Hemisphere, including in the Mediterranean climates of California and Europe. We sequenced the second intron of LFY to help resolve relationships within the clade where the internal transcribed spacer and chloroplast markers had previously failed to do so. Divergence time estimation and biogeographic analyses over the posterior distribution of dated trees suggest that a widespread ancestor was distributed across the Northern Hemisphere some 7-17 million years ago. Asian species form a sister group to a clade in which the European species are sister to the North American species. We use climatic niche modeling and divergence time estimates to explore the evolution of climate variables in the group. Principal component analyses help to identify instances of convergence, especially between distantly related species in the Mediterranean basin and in the chaparral of California. We document several cases of significant divergence between sister species in eastern North America and western North America. Climatic models were projected from one continent into the others (e.g., North American species projected into Asia and Europe) to examine whether species living in these areas occupy similar climates. This study demonstrates the utility of combining niche modeling with historical biogeographic analyses and documents significant climatic niche evolution within a group of species distributed throughout the Northern Hemisphere. These results suggest a possible model for the origin of the Madrean-Tethyan disjunction pattern.

Aim To infer the phylogenetic relationships and biogeography of Hydromantes, with special emphasis on the European taxa. In particular, we aimed to test: (1) the monophyly of the European species and current views on their interrelationships; and (2) previously proposed timings of the separation of European and American Hydromantes, and of biogeographically important events within Europe.Location California and the Western Mediterranean Basin, specifically south‐east France, Italy, and the island of Sardinia.Methods Partial sequences of mitochondrial genes (cytochrome b and 12S rRNA) were obtained from 45 specimens of Hydromantes, including all European extant species and subspecies, and two species from California. In addition, a fragment of the mitochondrial 16S rRNA gene was amplified for 16 specimens. Data sets were aligned using ClustalX, and well‐supported phylogenetic trees were produced using maximum‐likelihood, Bayesian and maximum‐parsimony methods. Estimates of divergence times were obtained with the program r8s, the molecular clock being calibrated using the opening of the Strait of Gibraltar, the final event in the Messinian Salinity Crisis of 5.3 Ma.Results Separation between the American and European clades occurred approximately 13.5 Ma, most probably before or after westward dispersal across the Bering Land Bridge. In Europe, divergence started in the late Miocene, when Hydromantes (A.) genei separated from other members of the genus 9 Ma and colonized south‐west Sardinia. Movement between the European mainland and Sardinia, by a member of the subgenus Speleomantes, occurred in the Messinian Salinity Crisis, after the Mediterranean Basin desiccated almost completely 5.96 Ma. Subsequent widespread aridification fragmented the geographical ranges of Hydromantes, which live in cool and humid conditions, resulting in the origin of the six species in the subgenus Speleomantes. In contrast, a second period of diversification, in continental Europe 2–1.3 Ma, was probably caused by very cold interludes during the climatic oscillations that characterized the Pleistocene.Main conclusions The molecular clock used here indicates that the separation of Californian and European Hydromantes occurred more recently than previously believed, and the same is true of some subsequent phylogenetic divergences within Europe. Estimated dates for these divergence events are consistent with known geophysical and climatic events that could have caused or facilitated them.

Spider conservation in Europe: a review

BackgroundThe glacial-interglacial oscillations caused severe range modifications of biota. Thermophilic species became extinct in the North and survived in southern retreats, e.g. the Mediterranean Basin. These repeated extinction and (re)colonisation events led to long-term isolation and intermixing of populations and thus resulted in strong genetic imprints in many European species therefore being composed of several genetic lineages. To better understand these cycles of repeated expansion and retraction, we selected the Marbled White butterfly Melanargia galathea. Fourty-one populations scattered over Europe and the Maghreb and one population of the sibling taxon M. lachesis were analysed using allozyme electrophoresis.ResultsWe obtained seven distinct lineages applying neighbour joining and STRUCTURE analyses: (i) Morocco, (ii) Tunisia, (iii) Sicily, (iv) Italy and southern France, (v) eastern Balkans extending to Central Europe, (vi) western Balkans with western Carpathian Basin as well as (vii) south-western Alps. The hierarchy of these splits is well matching the chronology of glacial and interglacial cycles since the Günz ice age starting with an initial split between the galathea group in North Africa and the lachesis group in Iberia. These genetic structures were compared with past distribution patterns during the last glacial stage calculated with distribution models.ConclusionsBoth methods suggest climatically suitable areas in the Maghreb and the southern European peninsulas with distinct refugia during the last glacial period and underpin strong range expansions to the North during the Postglacial. However, the allozyme patterns reveal biogeographical structures not detected by distribution modelling as two distinct refugia in the Maghreb, two or more distinct refugia at the Balkans and a close link between the eastern Maghreb and Sicily. Furthermore, the genetically highly diverse western Maghreb might have acted as source or speciation centre of this taxon, while the eastern, genetically impoverished Maghreb population might result from a relatively recent recolonisation from Europe via Sicily.

Comb cells and puparial silk in the oriental hornet nest: Structure and function.

Bechstein's bat (Myotis bechsteinii) is a European species, restricted to woodlands with preference for mature deciduous forests in lowlands. It is considered rare throughout its range, although it may be common in optimal habitats. Roosts play crucial roles in the ecology of bats, and survival is partially dependant on the extent to which roosts protect bats from environmental extremes and predators. Roost selection is especially important for reproductive females, due to the higher energetic demands imposed by reproduction so understanding roost selection by maternity colonies is important for conservation. We investigated maternity roost selection by M. bechsteinii in southwestern Spain as a hierarchical process that proceeds from broad landscape scales to fine-scale local habitat characteristics. Radio-tracking of 28 lactating females allowed location and census of 13 maternity roost sites. Roosts were characterised at four detail scales (cavity, tree, stand, and landscape). All the roosts occurred in Quercus pyrenaica trees, within stands of the same species of very diverse structure. Ten of the roosts were former woodpecker holes, among which seven had their entrance modified by nuthatch. Roosts were located inside the forest and close (< 620 m) to permanent water sources. Roost trees were characterised by a higher proportion of dead branches. Other explored variables such as tree height, orientation, foliar cover, and elevation did not explain distribution of roosts at any scale. The species' breeding roost selection is described for the first time in a Mediterranean area.

The Mediterranean Basin is expected to be more strongly affected by ongoing global climate change than most other regions of the earth. Given the magnitude of forecasted trends, there are great concerns for the particularly rich biodiversity found in the region. Studies of the consequences of past climate shifts on biodiversity represent one of the best sources of data to validate models of the ecological and evolutionary consequences of future changes. Here we review recent findings from palaeoecology, phylogeography and climate change research to (1) explore possible antecedents of the predicted climate warming in the younger geological history of the Mediterranean Basin, (2) assess how tree populations have reacted to them, and (3) evaluate the significance of the evolutionary heritage that is at stake. A major question of our retrospective approach is whether Quaternary tree extinctions took place primarily during glacial or during interglacial episodes. Available data are scanty and somewhat conflicting. In contrast, abundant phylogeographic evidence clearly indicates that the bulk of genetic diversity in European temperate tree species is almost invariably located in the southernmost part of their range. Long‐term persistence of isolated populations have been common phenomena in the Mediterranean, to the point that the current genetic structure in this area probably often reflects population divergence that pre‐dates the onset of the Mediterranean climate in the Pliocene. In particular, Tertiary migrations into the Mediterranean of tree taxa originating from Asia seem to have left their footprints in the current genetic structure in these slowly evolving organisms. Moreover, phylogeographic studies point to heterogeneous rates of molecular evolution across lineages that are inversely related with their stability. We conclude that relict tree populations in the Mediterranean Basin represent an evolutionary heritage of disproportionate significance for the conservation of European plant biodiversity.

Many recent studies of intraspecific geographic variation in maternally inherited chloroplast DNA (cpDNA) in European trees have revealed haplotype distributions that can be interpreted in terms of scenarios of postglacial migration and range expansion. However, there is still a lack of comparable information from widespread herb species. In the present study, we investigated the geographic distribution of cpDNA variation in 124 populations, covering a large part of the range of the widespread, dioecious, European herb, Silene dioica. PCR-RFLP analysis revealed 24 different cpDNA haplotypes. As in the majority of European tree species, the large-scale geographic distributions of the most common S. dioica haplotypes suggest that the species colonized Europe from more than one geographic source. Material from 16 populations of S. latifolia and five hybrid populations was also included in the study for comparative purposes. Five out of seven haplotypes detected in S. latifolia were shared with S. dioica. The similarity of the geographic distributions of the shared haplotypes in both species is consistent with a history of past and/or recent interspecific hybridization and introgression between these closely related plants. The two haplotypes detected only in S. latifolia were present in populations in the Mediterranean region – on the southern margin of the species’ area of sympatry, or outside the range of S. dioica.

ABSTRACTAll ecosystems are affected by climate change, but differences in the pace of change will render some areas more exposed than others. Such spatial patterns of risk are important when assessing the continued functionality of protected area (PA) networks or planning for their expansion. Europe is undertaking an expansion of the PA network to cover 30% of its land and sea surface by 2030, but this must account for climate risk. Here, we estimate four metrics of future climate risk across Europe: local velocity, analog velocity, magnitude, and residence time, and assess the level of climate exposure of European PAs vs. nonprotected control sites. We also evaluate the intensity of climate risks on > 1000 European species of conservation concern associated with Natura 2000 sites. Our results show large spatial differences in climate change exposure across Europe, with a faster pace and farther shifts in the Boreal, Steppic, and Pannonian regions but slower changes in the Mediterranean, Alpine, Arctic, and Macaronesia regions. The magnitude of climate change was higher for the Arctic, Alpine, and Mediterranean regions, implying large local differences between present and future climate. These spatial risk patterns were largely consistent across scenarios, but with up to three times higher risk under the most pessimistic vs. the most optimistic scenario. Large variation in climate exposure for species of conservation concern was revealed, including 11 species that are highly dependent on Natura 2000 sites and predicted to experience rapid climate change. Our results provide guidance for managing European PAs and expanding their coverage by pinpointing areas offering more stable climates. We emphasize the need for connectivity across the network to support species adaptation via range shifting. This is especially the case in areas facing high climate change magnitude but low velocity, implying that climate conditions similar to current ones will be found nearby.