Extra-Mediterranean refugia: The rule and not the exception?

ABSTRACT: Some decades ago, biogeographers distinguished three major faunal types of high importance for Europe: (i) Mediterranean elements with exclusive glacial survival in the Mediterranean refugia, (ii) Siberian elements with glacial refugia in the eastern Palearctic and only postglacial expansion to Europe and (iii) arctic and/or alpine elements with large zonal distributions in the periglacial areas and postglacial retreat to the North and/or into the high mountain systems. Genetic analyses have unravelled numerous additional refugia both of continental and Mediterranean species, thus strongly modifying the biogeographical view of Europe. This modified notion is particularly true for the so-called Siberian species, which in many cases have not immigrated into Europe during the postglacial period, but most likely have survived the last, or even several glacial phases, in extra-Mediterranean refugia in some climatically favourable but geographically limited areas of southern Central and Eastern Europe. Recently, genetic analyses revealed that typical Mediterranean species have also survived the Last Glacial Maximum in cryptic northern refugia (e.g. in the Carpathians or even north of the Alps) in addition to their Mediterranean refuge areas.     

贺兰山天牛科昆虫区系组成及垂直分布

2013-2015年对贺兰山不同海拔和垂直植被带的天牛科昆虫进行系统调查,结果表明:贺兰山共有天牛科昆虫6亚科31属45种,其中,沟胫天牛亚科属、种数最多,有14属21种,其种数占总种数46.67%,天牛亚科有9属12种,幽天牛亚科有4属5种,花天牛亚科有2属4种,锯天牛亚科2属2种,膜花天牛亚科仅有1属1种。区系组成上,古北界有17种,占总种数的37.78%,其中中亚成分占35.29%,广布型28种,占总种数的62.22%。在我国昆虫地理区划中的分布共有13个分布类型,全国广布型最多,占总种数的20%。以蒙新区种类为主体,并与东北区和华北区关系密切联系。在垂直分布上,以山前阔叶林带采集到的天牛种类最多,有22种,占总数的48.89%;从低海拔的山前荒漠半荒漠带到高海拔的山地针叶林带,天牛物种数量逐渐增加;除山地草原带外,随海拔增高,古北界种类逐渐增多。通过聚类分析显示,山地针叶林带和针阔混交林带天牛种类组成相似性较高;山地灌丛带、山地疏林带和山前阔叶林带相似性较高;山前荒漠半荒漠带、山地草地带与其他植被带天牛组成差异较大。     

Climatic preferences and frequent co-occurrence of boreal and temperate plants in Hokkaido Island,northern Japan

Effects of climatic factors on the forest plant distribution were examined in Hokkaido Island, northern Japan, where boreal and temperate plants frequently co-occur, and the relationship of floral make-up with phytogeography and life form was analysed. From the climatic preference of tree species consisting of mixed forest, the co-occurrence of boreal and temperate plants was found over a wider thermal range in regions of little rainfall. However, co-occurrence in a certain forest site was more notable in the understory than in the crown. A particular case of this involves southern evergreen shrubs and herbs in a boreal coniferous forest. Another is the presence of northern summergreen herbs in a temperate hardwood forest. It is speculated that the co-occurrence dates back at least to the late Quaternary, when a decrease in temperature associated with the glacial period forced understory plants to adapt their life form or leaf habit to snowcover and the light conditions of interior forests. Recent geographical evidence suggests that the island has been isolated from the southern island for at least 70000 years. Thus, the postglacial re-expansion of southern species does not appear to be due to immigration but rather due to their expansion from refuges on the island such as Oshima Peninsula for snow-tolerant species and Hidaka District for snow-intolerant species.     

Phylogeographical patterns and genetic diversity in three species of Eurasian boreal forest beetles

Many species that occur in formerly glaciated areas of Fennoscandia have reached their current ranges from glacial refugial areas in Eurasia. Little is known of the refugia and postglacial colonization routes of insect species that are confined to boreal forests. Here, we investigate the phylogeography of three species of saproxylic beetles distributed across Eurasia: two rare boreal forest specialists, Pytho kolwensis and Pytho abieticola , and a common, less specialized species, Pytho depressus . In all species, there were two well-defined haplotype clades based on 645 bp of cytochrome oxidase subunit I gene sequence. In each species one clade was found only in China. The other clade occurred from China to north-western Europe in both P. kolwensis and P. depressus , but was apparently absent from China in P. abieticola. In spite of common phylogeographical patterns, the distribution of genetic variation differed markedly between the three species. In P. kolwensis , a highly-threatened species in old-growth forests in Fennoscandia, there was an extremely low level of genetic variation throughout Eurasia. One common haplotype, represented by 86% of the samples, dominated in all sampling localities. Levels of genetic variation were higher in both P. abieticola and P. depressus , with 31% and 58%, respectively, of the samples representing a unique haplotype. In each species, relationships between haplotypes were not well resolved, and haplotypes from one sampling locality were generally not clustered in either Neighbour-joining trees or statistical parsimony networks. These patterns in the distribution of genetic variation can be attributed to differences in the species' population sizes, ecologies, glacial refugial areas, and postglacial colonization dynamics.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 267–279.     

The boreal forests of north-eastern Eurasia

The distribution, diversity and structure of boreal forests of monsoon Asia are reviewed. The boreal zone is circumscribed by the isotherms of 15°C and 45°C of Kira's Warmth Index (WI) and includes the territory of the Amur basin, the seashores of the Sea of Okhotsk, Kamchatka, Sakhalin and the Kuril islands. Boreal forests also occur on the mountains of Hokkaido, Honshu, Korea and NE China. The oceanic sectors of the boreal zone are composed mainly of birch forests, the marine sectors of spruce-fir forests and the continental sectors of larch and pine forests. The upper limit of the boreal forests ascends from the lowest level in the North to ca. 1500 m a.s.l. in the South of the zone, and reaches above 2500 m in the mountains of the temperate zone. In the boreal zone the secondary forests are common. They form after fires and cutting. Volcanism is an essential factor in the forest dynamics in the oceanic part of the region. It is suggested that as a result of global warming one may expect a strong alteration in vegetation pattern, especially in the northern and mountain areas.Paper presented at the International Symposium on Global Change Impacts on Terrestrial Ecosystems in Monsoon Asia (TEMA) Waseda University, Tokyo, 4–6 September 1993.     

The Niobrara River Valley,a postglacial migration corridor and refugium of forest plants and animals in the grasslands of central North America

The Niobrara River Valley of northern Nebraska contains numerous bryophyte, vascular plant, and animal species more typical of forests far to the east, north, and west than to other forests in the grasslands that surround the Valley. Some species are probably relicts of cooler glacial and early postglacial times, when much of what is now grassland was covered by boreal and cool-temperate forests. Others entered the Valley from the east in postglacial times, and some entered from the west as the climate became semi-arid. There is a steep decline in total number of vascular plant species from the mouth of the Missouri River up through the Niobrara Valley, suggesting an environmental gradient and differential migration and extirpation of species at various times since the Pleistocene.     

Spatial patterns of genetic diversity across European subspecies of the mountain hare, Lepus timidus L

Fossil evidence shows that populations of species that currently inhabit arctic and boreal regions were not isolated in refugia during glacial periods, but instead maintained populations across large areas of central Europe. These species commonly display little reduction in genetic diversity in northern areas of their range, in contrast to many temperate species. The mountain hare currently inhabits both temperate and arctic-boreal regions. We used nuclear microsatellite and mtDNA sequence data to examine population structure and alternate phylogeographic hypotheses for the mountain hare, that is, temperate type (lower genetic diversity in northern areas) and arctic-boreal type (high northern genetic diversity). Both data sets revealed concordant patterns. Highest allelic richness, expected heterozygosity and mtDNA haplotype diversity were identified in the most northerly subspecies, indicating that this species more closely maps to phylogeographic patterns observed in arctic-boreal rather than temperate species. With regard to population structure, the Alpine and Fennoscandian subspecies were most genetically similar (F(ST) approximately 0.1). These subspecies also clustered together on the mtDNA tree and were assigned with highest likelihood to a common Bayesian cluster. This is consistent with fossil evidence for intermediate populations in the central European plain, persisting well into the postglacial period. In contrast, the geographically close Scottish and Irish populations occupied separate Bayesian clusters, distinct clades on the mtDNA maximum likelihood tree and were genetically divergent from each other (F(ST) > 0.4) indicating the influence of genetic drift, long isolation (possibly dating from the late glacial era) and/or separate postglacial colonisation routes.     

Horizontal and elevational phylogeographic patterns of Himalayan and Southeast Asian forest passerines (Aves: Passeriformes)

Aim Zoogeographic patterns in the Himalayas and their neighbouring Southeast Asian mountain ranges include elevational parapatry and ecological segregation, particularly among passerine bird species. We estimate timings of lineage splits among close relatives from the north Palaearctic, the Sino‐Himalayan mountain forests and from adjacent Southeast Asia. We also compare phylogeographic affinities and timing of radiation among members of avian communities from different elevational belts. Location East Asia. Methods We reconstructed molecular phylogenies based on a mitochondrial marker (cytochrome b) and multilocus data sets for seven passerine groups: Aegithalidae, Certhiidae (Certhia), Fringillidae (Pyrrhula), Paridae (Periparus), Phylloscopidae, Regulidae and Timaliidae (Garrulax sensu lato). Molecular dating was carried out using a Bayesian approach applying a relaxed clock in beast . Time estimates were inferred from three independent calibrations based on either a fixed mean substitution rate or fixed node ages. The biogeographic history of each group was reconstructed using a parsimony‐based approach. Results Passerine radiation in Southeast Asia can be divided into roughly three major phases of separation events. We infer that an initial Miocene radiation within the Southeast Asian region included invasions of (sub)tropical faunal elements from the Indo‐Burmese region to the Himalayan foothills and further successive invasions to Central Asia and Taiwan towards the early Pliocene. During two further Pliocene/Pleistocene phases, the subalpine mountain belt of the Sino‐Himalayas was initially invaded by boreal species with clear phylogenetic affinities to the north Palaearctic taiga belt. Most terminal splits between boreal Himalayan/Chinese sister taxa were dated to the Pleistocene. Main conclusions Extant patterns of elevational parapatry and faunal transition in the Sino‐Himalayas originated from successive invasions from different climatic regions. The initiation of Southeast Asian passerine diversification and colonization of the Himalayan foothills in the mid‐Miocene coincides with the postulated onset of Asian monsoon climate and the resulting floral and faunal turnovers. Patterns of elevational parapatry were established by southward invasions of boreal avifaunal elements to the subalpine Sino‐Himalayan forest belt that were strongly connected to climate cooling towards the end of the Pliocene. Current patterns of allopatry and parapatry in boreal species (groups) were shaped through Pleistocene forest fragmentation in East Asia.     

Vegetation surveys in the circumboreal coniferous forests: A review

As an introduction to the special issue “Vegetation of Circumboreal Coniferous Forests”, a brief history is provided of the development and present state of floristic vegetation surveys of the boreal and mountain coniferous forests of Europe, northern Asia and North America. The focus is on forests assigned to the phytosociological classVaccinio-Piceetea and closely related vegetation types.     

Vegetation history since the last glacial period in the Mikata lowland,the Sea of Japan area,western Japan

Local and regional vegetation since the last glacial period was reconstructed on the basis of a palynological study of sediment at Iwaya, in the Sea of Japan area, western Japan. During the interstade (before about 30 000 years BP), forests were composed predominantly ofCryptomeria japonica withTsuga sieboldii and cool-temperate deciduous broad-leaved trees. In the pre-full-glacial, the full-glacial and the early late-glacial (30 000-12 000 years BP), forests were dominated by temperate (montane) and boreal (subalpine) Pinaceae andBetula. During the early full-glacial, the pinaceous forests were mixed with cool-temperate trees such asFagus crenata. In the late full-glacial (18 000-16 000 years BP), the maximum development of pinaceous conifer forests was recognized. Cool-temperate broad-leaved forests composed mainly ofF. crenata andQuercus (Lepidobalanus) replaced the pinaceous forests at about 12 000 years BP and were maintained to the early postglacial.Cryptomeria japonica was distributed around the Mikata lowland during the last glacial.Cryptomeria japonica, which began to increase at 16 000 years BP, increased abruptly in the early postglacial and spread throughout the postglacial in the lowlands. After 6300 years BP, lucidophyllous forests composed mainly ofQuercus (Cyclobalanopsis) andCastanopsis were established in the Mikata district; this was later than in the inland and the Pacific Ocean areas in the Kinki region, western Japan. In historic times (afterca 2000 years BP), secondary forest ofPinus densiflora, which can grow as a pioneer in disturbed habitats, spread.     

On the estimated age of recent fauna in different latitudinal zones (by the example of brachiopods)

Maximum period of existence of endemic genera in the temperately warm-water (subtropical) and temperately cold-water (low boreal and subantarctic) faunal belts is revealed on the base of latest data on the composition of brachiopod fauna of recent seas and oceans and most full data on the paleontological finds of recent brachiopod genera. Most of the subtropical endemics inhabit southern hemisphere; one of them is Bouchardia, which is known beginning with the Late Cretaceous (Maastrichtian), that brings together southern subtropical faunal belt with the tropical zone and suggests that subtropical waters could serve as a refuge for representatives of ancient warm-water faunas during large geological and hydrological reconstructions at the Mesozoic-Cenozoic boundary. Most of the younger endemics of low boreal and subantarctic faunal belts appeared at the Paleogene-Neogene boundary, during the period of formation of modern contrast climate on the Earth that was caused by the formation of Antarctic Circumpolar Current. The appearance of recent species and genera in the Holocene was influenced by last considerable postglacial reconstruction of climate.     

Ecological factors explaining the location of the boundary between the mixedwood and coniferous bioclimatic zones in the boreal biome of eastern North America

Aim Climate is often regarded as the primary control determining the location of an ecotone between two vegetation zones. However, other ecological factors may also be important, especially when the northern limit of the dominant species of a vegetation zone extends further than the limit of the zone itself. This study aimed to identify the ecological variables explaining the transition between two zones within the boreal biome in Quebec (eastern Canada): the southern mixedwood forests dominated by balsam fir (Abies balsamea) and white birch (Betula papyrifera), and the northern coniferous forests dominated by black spruce (Picea mariana). Location Quebec (eastern Canada). Methods Data from 5023 sampling plots from the ecological inventory of the Québec Ministry of Natural Resources distributed throughout the two bioclimatic zones were used in logistic regressions to determine the relationships between the presence or absence of balsam fir stands and different abiotic and biotic variables, at both stand and landscape scales. Results The presence of balsam fir stands was negatively related to the thick organic horizons, coarse xeric deposits and low positions on the slope, whereas stands were favoured by high elevations, steep slopes and moderate drainage. These results defined the suitable conditions for the development of balsam fir stands. In the coniferous zone these suitable conditions were less abundant. Furthermore, the saturation level of suitable sites was lower, as well as the incidence of balsam fir stands in unsuitable sites (overflow). Balsam fir stands were mostly located near lakes and rivers. All significant variables at both the stand and landscape scales explained between 34 and 42% of the location of the potential northern distribution limit of the mixedwood zone. Main conclusions Our results suggest the important role of historical factors related to post‐glacial vegetation and past disturbances in determining the relative abundance of balsam fir in both zones of the boreal biome.     

Glacial refugia and postglacial expansion of the alpine–prealpine plant species Polygala chamaebuxus

The shrubby milkwort (Polygala chamaebuxus L.) is widely distributed in the Alps, but occurs also in the lower mountain ranges of Central Europe such as the Franconian Jura or the Bohemian uplands. Populations in these regions may either originate from glacial survival or from postglacial recolonization. In this study, we analyzed 30 populations of P. chamaebuxus from the whole distribution range using AFLP (Amplified Fragment Length Polymorphism) analysis to identify glacial refugia and to illuminate the origin of P. chamaebuxus in the lower mountain ranges of Central Europe. Genetic variation and the number of rare fragments within populations were highest in populations from the central part of the distribution range, especially in the Southern Alps (from the Tessin Alps and the Prealps of Lugano to the Triglav Massiv) and in the middle part of the northern Alps. These regions may have served, in accordance with previous studies, as long‐term refugia for the glacial survival of the species. The geographic pattern of genetic variation, as revealed by analysis of molecular variance, Bayesian cluster analysis and a PopGraph genetic network was, however, only weak. Instead of postglacial recolonization from only few long‐term refugia, which would have resulted in deeper genetic splits within the data set, broad waves of postglacial expansion from several short‐term isolated populations in the center to the actual periphery of the distribution range seem to be the scenario explaining the observed pattern of genetic variation most likely. The populations from the lower mountain ranges in Central Europe were more closely related to the populations from the southwestern and northern than from the nearby eastern Alps. Although glacial survival in the Bohemian uplands cannot fully be excluded, P. chamaebuxus seems to have immigrated postglacially from the southwestern or central‐northern parts of the Alps into these regions during the expansion of the pine forests in the early Holocene.     

Ecological patterns strongly impact the biogeography of western Palaearctic longhorn beetles (Coleoptera: Cerambycoidea)

We aim to unravel the biogeographic structuring of western Palaearctic longhorn beetles with focus on the location of different refugia, barriers to dispersal and postglacial range expansions with their particular filters. The interaction of different ecological features with these structures is analysed. The western Palaearctic was divided into 95 geographic entities. We produced presence-only matrices for all 955 Cerambycoidea species autochthonous to this area and derived species richness distributions and extracted faunal regions and faunal elements by cluster analyses and principal component analyses. Similar analyses were performed for sub-families and ecological groups. Longhorn beetles show a strong biogeographic structuring in the western Palaearctic. Species numbers strongly decrease to the north and west. Less mobile species and root feeders mostly contribute to the fauna of the Mediterranean region, whilst mobile species are more widespread. Feeders on broad-leaved trees dominate in western Europe, whilst feeders on coniferous trees are most important in northern Europe. Our results support multiple refugia in the Mediterranean region and underline the importance of Provence, Crimea and Crete as such refugia. Crete even might be an area of old endemism. The Atlanto- and the Ponto-Mediterranean regions are more strongly structured than assumed in classical biogeography. Mediterranean assemblages are mostly composed of non-flying species, root feeders and species with small distributions not found outside their glacial refugia. Tree feeders left their glacial retreats with their host plants. These range dynamics result in biogeographic structures with several dispersal barriers and filters composed of mountains, sea straits and climatic conditions.     

Genetic differentiation of the Western Capercaillie highlights the importance of South-eastern Europe for understanding the species phylogeography

The Western Capercaillie (Tetrao urogallus L.) is a grouse species of open boreal or high altitude forests of Eurasia. It is endangered throughout most mountain range habitat areas in Europe. Two major genetically identifiable lineages of Western Capercaillie have been described to date: the southern lineage at the species' southernmost range of distribution in Europe, and the boreal lineage. We address the question of genetic differentiation of capercaillie populations from the Rhodope and Rila Mountains in Bulgaria, across the Dinaric Mountains to the Slovenian Alps. The two lineages' contact zone and resulting conservation strategies in this so-far understudied area of distribution have not been previously determined. The results of analysis of mitochondrial DNA control region sequences of 319 samples from the studied populations show that Alpine populations were composed exclusively of boreal lineage; Dinaric populations of both, but predominantly (96%) of boreal lineage; and Rhodope-Rila populations predominantly (>90%) of southern lineage individuals. The Bulgarian mountains were identified as the core area of the southern lineage, and the Dinaric Mountains as the western contact zone between both lineages in the Balkans. Bulgarian populations appeared genetically distinct from Alpine and Dinaric populations and exhibited characteristics of a long-term stationary population, suggesting that they should be considered as a glacial relict and probably a distinct subspecies. Although all of the studied populations suffered a decline in the past, the significantly lower level of genetic diversity when compared with the neighbouring Alpine and Bulgarian populations suggests that the isolated Dinaric capercaillie is particularly vulnerable to continuing population decline. The results are discussed in the context of conservation of the species in the Balkans, its principal threats and legal protection status. Potential conservation strategies should consider the existence of the two lineages and their vulnerable Dinaric contact zone and support the specificities of the populations.     

Genetic differentiation of an endangered capercaillie (<Emphasis Type="Italic">Tetrao urogallus</Emphasis>) population at the Southern edge of the species range

The low-latitude limits of species ranges are thought to be particularly important as long-term stores of genetic diversity and hot spots for speciation. The Iberian Peninsula, one of the main glacial refugia in Europe, houses the southern distribution limits of a number of boreal species. The capercaillie is one such species with a range extending northwards to cover most of Europe from Iberia to Scandinavia and East to Siberia. The Cantabrian Range, in North Spain, constitutes the contemporary south-western distribution limit of the species. In contrast to all other populations, which live in pure or mixed coniferous forests, the Cantabrian population is unique in inhabiting pure deciduous forests. We have assessed the existence of genetic differentiation between this and other European populations using microsatellite and mitochondrial DNA (mtDNA) extracted from capercaillie feathers. Samples were collected between 2001 and 2004 across most of the current distribution of the Cantabrian population. Mitochondrial DNA analysis showed that the Cantabrian birds form a distinct clade with respect to all the other European populations analysed, including the Alps, Black Forest, Scandinavia and Russia, which are all members of a discrete clade. Microsatellite DNA from Cantabrian birds reveals the lowest genetic variation within the species in Europe. The existence of birds from both mtDNA clades in the Pyrenees and evidence from microsatellite frequencies for two different groups, points to the existence of a Pyrenean contact zone between European and Cantabrian type birds. The ecological and genetic differences of the Cantabrian capercaillies qualify them as an Evolutionarily Significant Unit and support the idea of the importance of the rear edge for speciation. Implications for capercaillie taxonomy and conservation are discussed.     

Classification of ultracontinental boreal forests in central Yakutia

Using the Braun-Blanquet approach, five associations of boreal forests were distinguished in central Yakutia, the most continental part of eastern Siberia. Ecological features of the syntaxa were explained with the use of the DCA ordination of 50 relevés. All available data from eastern Siberia were involved in the study for syntaxonomic analysis. Central Yakutian boreal forests were classified into two classes:Rhytidio-Laricetea sibiricae Korotkov etErmakov 1999 — ultracontinental light coniferous hemiboreal forests, andVaccinio-Piceetea Br.-Bl. inBr.-Bl. et al. 1939 — typical coniferous taiga forests of northern Eurasia. A new concept of higher syntaxonomic units of the classVaccinio-Piceetea in eastern Siberia has been developed. Three orders represent the diversity of taiga forests: (1)Cladonio-Vaccinietalia Kielland-Lund 1967 (with alliancesHieracio umbellati-Pinion sylvestris Anenkhonov etChytry 1998 andSaxifrago bronchialis-Pinion sylvestris all. nov.) — light coniferous boreal forests occurring in dry and moderately dry oligotrophic sites in various climatic sectors of Northern Eurasia; (2)Lathyro humilis-Laricetalia cajanderi ord. nov. (with alliancesAulacomnio acuminati-Laricion cajanderi all. nov. andRhododendro daurici-Laricion gmelinii all. nov.) — zonal boreal forests with xeric elements, which are typical of regions of northern Asia with cold, dry ultracontinental climate; (3)Ledo-Laricetalia cajanderi ord. prov. (with allianceLedo-Laricion cajanderi prov.) — North Eurasian boreal forests occurring in cold sites with excessive soil moisture, sometimes water-logged. Phytogeography and ecology of these orders are discussed in comparison with other regions of northern Asia.     

Seasonal variability in habitat structure may have shaped acoustic signals and repertoires in the black-capped and boreal chickadees

Many songbird species have evolved multiple vocalizations, or repertoires, that function to communicate various biological signals. More diverse repertoires may have evolved in response to the effects of seasonal variation in habitat structure on signal transmission. Such changes in habitat necessarily occur for migrating species, but they also occur for resident species that occupy deciduous forests. The black-capped chickadee (Poecile atricapillus) possesses a chick-a-dee call and a fee-bee song, but the closely related boreal chickadee (P. borealis) lacks a song. Consistent with the habitat variability hypothesis, the black-capped chickadee possesses a larger repertoire and primarily occupies deciduous forests, whereas the songless boreal chickadee occurs more often in coniferous forests. We explored the ecological basis of this hypothesis by recording audio playbacks of two species in two habitat types during two seasons. Specifically, we played both songs and calls of the black-capped chickadee and calls of the boreal chickadee in deciduous and coniferous habitats, prior to and after leaf-out. We measured attenuation and degradation in re-recorded vocalizations. For black-capped chickadees, the song was less degraded than the call in post-leaf, deciduous forests. The boreal chickadee call attenuated more quickly in all treatments, but maintained its acoustic structure better than both black-capped chickadee vocalizations in coniferous forests. Our results support the hypothesis that variable habitats provided a seasonal transmission benefit for both song and call in the black-capped chickadee, but that the transmission benefit of song is lost in the less variant coniferous forests, which may underlie the absence of a song in the boreal chickadee.     

Einwanderungsrouten nach Mitteleuropa

Butterflies – who is coming, who is going? Immigration routes to Central Europe The majority of butterflies and burnet moths distributed in Central Europe today results from postglacial colonisation of this region mostly from expansions out of the well known Mediterranean refuge areas. The study region was mostly colonised west or east of the Alps, which represent a strong dispersal obstacle. Species being widespread under glacial conditions in Central Europe have retreated under the warm conditions of the Postglacial to the high mountain systems or the Arctic. Therefore, they are mostly restricted to the high altitudes of the Alps in Western Central Europe. Nevertheless, some species have even survived in lowland areas of Central Europe during glacial and inter‐/postglacial conditions, but these species are either better adapted to glacial or to inter‐/postglacial conditions.     

Are habitat islands islands? Woodliving beetles (Coleoptera) in deciduous forest fragments in boreal forest

Modern forestry has changed the fire dynamics in the boreal forest and as a result the size and number of deciduous forest patches have been reduced as well as the number of deciduous trees within the coniferous forest This has exaggerated the insularity of deciduous forest patches within the boreal coniferous forest zone In this paper I examine whether the diversity of beetles living in dead stems of deciduous trees follows the relationship with patch area to be predicted from island biogeographic theory, and to what extent the species assemblages differ between large and small patches and single deciduous trees within managed coniferous forests Three larger patches of deciduous forest (>120 ha) arisen as successions after forest fires in the late 1880's are compared with 6 small (<20 ha) patches of similar origin No difference in diversity could be detected between large and small patches A statistically significant difference between assembly composition was detected using matrix regression between a matrix of observed assembly similarities and a hypothetical similarity matrix based on the type of area in which the plot was situated (large- small or matrix) Finally I examined the distribution of the 56 most common beetle species over the habitat types investigated Thirty four species did not show any Significant habitat preference, 12 were found more often in clear-cut areas than expected 5 were over-represented in small patches, and 4 in large areas My explanation to this apparent lack of insularity effects is the relative recent commencement of intensive forestry in these areas and the fact that the matrix is of rather high quality that is inhabitable for numerous beetle species Thus the patches may be viewed as incipient islands still exchanging biota with their surroundings