Institut für Ökologie, Evolution und Diversität
Refine
Year of publication
Document Type
- Article (344)
- Preprint (31)
- Doctoral Thesis (20)
- Part of a Book (2)
- Conference Proceeding (2)
Language
- English (399) (remove)
Has Fulltext
- yes (399)
Is part of the Bibliography
- no (399)
Keywords
- Invasive species (11)
- Biogeography (10)
- Biodiversity (9)
- taxonomy (8)
- Thailand (5)
- phylogeny (5)
- Community ecology (4)
- Crustacea (4)
- Ecological modelling (4)
- Ecology (4)
Institute
- Institut für Ökologie, Evolution und Diversität (399)
- Senckenbergische Naturforschende Gesellschaft (213)
- Biodiversität und Klima Forschungszentrum (BiK-F) (130)
- Biowissenschaften (94)
- Medizin (14)
- Zentrum für Interdisziplinäre Afrikaforschung (ZIAF) (9)
- Geowissenschaften (6)
- LOEWE-Schwerpunkt für Integrative Pilzforschung (5)
- Institut für sozial-ökologische Forschung (ISOE) (3)
- Exzellenzcluster Makromolekulare Komplexe (2)
- Geographie (2)
- Informatik (2)
- Biochemie und Chemie (1)
- Sonderforschungsbereiche / Forschungskollegs (1)
- Universität des 3. Lebensalters e.V. (1)
Sacoglossan sea slugs are unique in the animal kingdom in that they sequester and maintain active plastids that they acquire from the siphonaceous algae upon which they feed, making the animals photosynthetic. While most sacoglossan species digest their freshly ingested plastids within hours, four species from the family Plakobranchidae retain their stolen plastids (kleptoplasts) in a photosynthetically active state on time scales of weeks to months. The molecular basis of plastid maintenance within the cytosol of digestive gland cells in these photosynthetic metazoans is yet unknown, but is widely thought to involve gene transfer from the algal food source to the slugs based upon previous investigations of single genes. Indeed, normal plastid development requires hundreds of nuclear-encoded proteins, with protein turnover in photosystem II in particular known to be rapid under various conditions. Moreover, only algal plastids, not the algal nuclei, are sequestered by the animals during feeding. If algal nuclear genes are transferred to the animal either during feeding or in the germ line, and if they are expressed, then they should be readily detectable with deep-sequencing methods. We have sequenced expressed mRNAs from actively photosynthesizing, starved individuals of two photosynthetic sea slug species, Plakobranchus ocellatus Van Hasselt, 1824 and Elysia timida Risso, 1818. We find that nuclear-encoded, algal-derived genes specific to photosynthetic function are expressed neither in P. ocellatus nor in E. timida. Despite their dramatic plastid longevity, these photosynthetic sacoglossan slugs do not express genes acquired from algal nuclei in order to maintain plastid function.
Responding to inadequate awareness of the outstanding importance of biodiversity, the BioFrankfurt network was founded in 2004 in the State of Hesse, Germany. It is presented here as a case study and may serve as a model for other parts of the world, such as the Middle East. In 2007, only about 26% of the German population were familiar with the term “Biodiversity”, and most of them only had a vague idea about its meaning. The BioFrankfurt network of institutions addressed this problem, raising public awareness and supporting research, education and conservation. A regional biodiversity education program has been developed and delivered to more than 500 schools. Since 2007, an innovative public relations campaign combines raising awareness on regional biodiversity issues with activities to improve the public image of the Frankfurt area. Because of its geographical focus, the network’s activities gained the attention of local and regional politicians and other decision makers, culminating in the joint establishment of a new Biodiversity and Climate Research Centre by BioFrankfurt member institutions. The success of current activities attracts interesting partners, resulting in challenging cooperation initiatives. The authors are convinced that the network’s concepts and activities have a great potential to profoundly enhance the notion and acceptance of biodiversity issues elsewhere. Keywords: BioFrankfurt, biodiversity network, education, public awareness, scientifi c communication
The huntsman spider genera Sparianthina Banks, 1929 and Anaptomecus Simon, 1903 are reviewed. The type species of Sparianthina, Sparianthina selenopoides Banks, 1929, is redescribed, illustrated, and recorded from Costa Rica for the first time; a lectotype and paralectotype are designated. Three species are transferred to the genus: Sparianthina pumilla (Keyserling, 1880) comb. n. from Heteropoda Latreille, 1804 (lectotype and paralectotype are designated), Sparianthina rufescens (Mello-Leitão, 1940) comb. n. from Anaptomecus and Sparianthina milleri (Caporiacco, 1955) comb. n. from Macrinus Simon, 1887. The ♂ of S. rufescens (Mello-Leitão, 1940) comb. n. and the ♀ of S. milleri comb. n. are described for the first time. Three new species are described: Sparianthina adisi sp. n., S. deltshevi sp. n., and S. saaristoi sp. n. The male and female of Anaptomecus longiventris Simon, 1903 are described for the first time and the species is recorded from Panama for the first time. Two new species are described: Anaptomecus temii sp. n. and A. levyi sp. n. Keywords: Taxonomy, re-description, transfer, review
A new species of the basal araneomorph spider genus Ectatosticta (Araneae, Hypochilidae) from China
(2009)
The hypochilid spider Ectatosticta davidi (Simon) is redescribed on the basis of adults from Mt. Taibaishan in Shaanxi Province, China; the specimens from Qinghai Province previously identified as E. davidi by most modern authors belong to a new species described as E. deltshevi. Keywords: Araneae, Araneomorphae, Hypochilidae, Ectatosticta, China
Despite prevailing arid conditions, the diversity of terrestrial and freshwater biota in the Middle East is amazingly high and marine biodiversity is among the highest on Earth. Th roughout the Region, threats to the environment are moderate to severe. Despite the outstanding economic and ecological importance of biological diversity, the capacity in biodiversity-related research and academic education is inadequate. The "Middle Eastern Biodiversity Network" (MEBN), founded in 2006 by six universities and research institutes in Iran, Jordan, Germany, Lebanon and Yemen was designed to fi ll this gap. An integrated approach is taken to upgrade biodiversity research and education in order to improve regional ecosystem conservation and management capacities. A wide range of activities are carried out in the framework of the Network, including capacity building in biological collection management and professional natural history curatorship, developing university curricula in biodiversity, conducting scientifi c research, organising workshops and conferences on Middle Eastern biodiversity, and translating the results of biodiversity research into conservation and sustainable development. Keywords: Middle Eastern biodiversity, nature museums, biodiversity research, biodiversity education, biodiversity conservation, biodiversity networks
Background: Local adaptation to divergent environmental conditions can promote population genetic differentiation even in the absence of geographic barriers and hence, lead to speciation. Perturbations by catastrophic events, however, can distort such parapatric ecological speciation processes. Here, we asked whether an exceptionally strong flood led to homogenization of gene pools among locally adapted populations of the Atlantic molly (Poecilia mexicana, Poeciliidae) in the Cueva del Azufre system in southern Mexico, where two strong environmental selection factors (darkness within caves and/or presence of toxic H2S in sulfidic springs) drive the diversification of P. mexicana. Nine nuclear microsatellites as well as heritable female life history traits (both as a proxy for quantitative genetics and for trait divergence) were used as markers to compare genetic differentiation, genetic diversity, and especially population mixing (immigration and emigration) before and after the flood. Results: Habitat type (i.e., non-sulfidic surface, sulfidic surface, or sulfidic cave), but not geographic distance was the major predictor of genetic differentiation. Before and after the flood, each habitat type harbored a genetically distinct population. Only a weak signal of individual dislocation among ecologically divergent habitat types was uncovered (with the exception of slightly increased dislocation from the Cueva del Azufre into the sulfidic creek, El Azufre). By contrast, several lines of evidence are indicative of increased flood-induced dislocation within the same habitat type, e.g., between different cave chambers of the Cueva del Azufre. Conclusions: The virtual absence of individual dislocation among ecologically different habitat types indicates strong natural selection against migrants. Thus, our current study exemplifies that ecological speciation in this and other systems, in which extreme environmental factors drive speciation, may be little affected by temporary perturbations, as adaptations to physico-chemical stressors may directly affect the survival probability in divergent habitat types.
Background: Pythium ultimum (P. ultimum) is a ubiquitous oomycete plant pathogen responsible for a variety of diseases on a broad range of crop and ornamental species. Results: The P. ultimum genome (42.8 Mb) encodes 15,290 genes and has extensive sequence similarity and synteny with related Phytophthora species, including the potato blight pathogen Phytophthora infestans. Whole transcriptome sequencing revealed expression of 86% of genes, with detectable differential expression of suites of genes under abiotic stress and in the presence of a host. The predicted proteome includes a large repertoire of proteins involved in plant pathogen interactions although surprisingly, the P. ultimum genome does not encode any classical RXLR effectors and relatively few Crinkler genes in comparison to related phytopathogenic oomycetes. A lower number of enzymes involved in carbohydrate metabolism were present compared to Phytophthora species, with the notable absence of cutinases, suggesting a significant difference in virulence mechanisms between P. ultimum and more host specific oomycete species. Although we observed a high degree of orthology with Phytophthora genomes, there were novel features of the P. ultimum proteome including an expansion of genes involved in proteolysis and genes unique to Pythium. We identified a small gene family of cadherins, proteins involved in cell adhesion, the first report in a genome outside the metazoans. Conclusions: Access to the P. ultimum genome has revealed not only core pathogenic mechanisms within the oomycetes but also lineage specific genes associated with the alternative virulence and lifestyles found within the pythiaceous lineages compared to the Peronosporaceae.
In over 100 genera of tropical angiosperms, one or more species possess specialised structures for housing ants. The longevity and intimacy of these associations has often facilitated an increasing specialisation of both the ants and the plants, leading to a number of highly specific and obligate symbioses. Early literature contained only few anecdotal reports of the ant genus Cladomyrma WHEELER inhabiting (unidentified) plants. This work presents the new findings on Cladomyrma and its host plants that accumulated over the last two decades. My studies of Cladomyrma reveal that there is a largely overlooked community of south-east Asian plant-ants and their associated plants. Currently the genus consists of at least 12 species. Cladomyrma has been thought to be restricted to the ever-wet part of the West Malesian floristic region, comprising the Malay Peninsula, Borneo, and Sumatra, but recent collections from Thailand and Vietnam indicate that species of the genus penetrate the seasonal tropical forests of Continental Asia. Cladomyrma inhabits 24 plant species belonging to a surprisingly extensive range of plant taxa: Callerya, Saraca, Spatholobus (Fabaceae), Crypteronia (Crypteroniaceae), Drypetes (Putranjivaceae), Ryparosa (Achariaceae), Strychnos (Loganiaceae), Neonauclea (Rubiaceae), Luvunga (Rutaceae) and Sphenodesme (Verbenaceae). In terms of taxonomic diversity on the genus and family level the range of hosts utilised by Cladomyrma is one of the broadest ever recorded for any live stem-nesting plant-ant lineage worldwide. This work provides a species-level overview of all Cladomyrma host plants known from Borneo, the Malay Peninsula and Sumatra, including descriptions of ant-housing structures (domatia), ant inhabitant identity, onset of colonisation during plant ontogeny, nest structure, occupancy rate, and considerations of results obtained from herbarium specimens. Both the regularity of ant association and the degree of morphological specialisation toward myrmecophytism are assessed. The behavioural traits of Cladomyrma are compatible with traits exhibited by other protective plant-ants. This work demonstrates that all species of Cladomyrma investigated (dianeae, maschwitzi, yongi, petalae) confer antiherbivore protection to young leaves of its host. The ants also attack and repell or kill herbivorous insect larvae encountered on young foliage. Cleaning behaviour appears to be a trait shared by all members of the genus, and the two species tested (maschwitzi, petalae) successfully removed termite eggs experimentally placed onto young leaves. Another trait common to all known species of the genus is that the ants preferentially patrol young shoots and leaves ('neophily'). These behavioural traits of Cladomyrma likely reduce stem damage and pathogenic infection of their host. The ants prune encroaching vegetation (tested in dianeae maschwitzi, petalae, yongi, observed in crypteroniae) and attack paper tape used to mark host plants (observed in andrei, dianeae, hobbyi, nudidorsalis, maschwitzi, yongi, petalae). If these traits combined translate into a better reproductive success of the hosts has yet to be verified. Evidence for lifetime fitness benefits is particularly difficult to quantify for the long-lived woody host plants of Cladomyrma. The predominant food source of Cladomyrma appears to be the honeydew of scale insects (Coccidae and Pseudococcidae) which the ants tend inside their nest cavities. Observations on scale insect acquisition by Cladomyrma foundress queens show that hemipteran trophobionts are not transported by the queens on their nuptial flight but they nevertheless arrive on the host plant independently of the ants. Entry into nest chambers is facilitated by small holes kept open by the foundress queen. Most Cladomyrma species have been recorded from only one or two (three) host plant species (andrei, crypteroniae, hobbyi, maschwitzi, nudidorsalis, scopulosa, yongi), but two species, Cladomyrma petalae and C. dianeae, are more catholic in their host usage; the first being a 'generalist' plant-ant colonising hosts across a broad taxonomic range, the second inhabiting several members of the genus Neonauclea. First results of host-choice experiments with C. petalae are presented and the potential mechanisms promoting host specificity are discussed. My studies of the Cladomyrma/plant associations indicate that codiversification and host shifts or host expansions, rather than cospeciation, shape the pattern of species interactions in this system. Finally, I propose a scenario in which three key traits of Cladomyrma –access to live stems, utilisation of indirect food rewards via trophobionts and 'neophily'– are hypothesised to favour niche differentiation and the acquisition of new hosts over evolutionary time.
Background: Mitochondrial DNA sequencing increasingly results in the recognition of genetically divergent, but morphologically cryptic lineages. Species delimitation approaches that rely on multiple lines of evidence in areas of co-occurrence are particularly powerful to infer their specific status. We investigated the species boundaries of two cryptic lineages of the land snail genus Trochulus in a contact zone, using mitochondrial and nuclear DNA marker as well as shell morphometrics.
Results: Both mitochondrial lineages have a distinct geographical distribution with a small zone of co-occurrence. In the same area, we detected two nuclear genotype clusters, each being highly significantly associated to one mitochondrial lineage. This association however had exceptions: a small number of individuals in the contact zone showed intermediate genotypes (4%) or cytonuclear disequilibrium (12%). Both mitochondrial lineage and nuclear cluster were statistically significant predictors for the shell shape indicating morphological divergence. Nevertheless, the lineage morphospaces largely overlapped (low posterior classification success rate of 69% and 78%, respectively): the two lineages are truly cryptic.
Conclusions: The integrative approach using multiple lines of evidence supported the hypothesis that the investigated Trochulus lineages are reproductively isolated species. In the small contact area, however, the lineages hybridise to a limited extent. This detection of a hybrid zone adds an instance to the rare reported cases of hybridisation in land snails.
Background, aim, and scope Food consumption is an important route of human exposure to endocrine-disrupting chemicals. So far, this has been demonstrated by exposure modeling or analytical identification of single substances in foodstuff (e.g., phthalates) and human body fluids (e.g., urine and blood). Since the research in this field is focused on few chemicals (and thus missing mixture effects), the overall contamination of edibles with xenohormones is largely unknown. The aim of this study was to assess the integrated estrogenic burden of bottled mineral water as model foodstuff and to characterize the potential sources of the estrogenic contamination. Materials, methods, and results In the present study, we analyzed commercially available mineral water in an in vitro system with the human estrogen receptor alpha and detected estrogenic contamination in 60% of all samples with a maximum activity equivalent to 75.2 ng/l of the natural sex hormone 17beta-estradiol. Furthermore, breeding of the molluskan model Potamopyrgus antipodarum in water bottles made of glass and plastic [polyethylene terephthalate (PET)] resulted in an increased reproductive output of snails cultured in PET bottles. This provides first evidence that substances leaching from plastic food packaging materials act as functional estrogens in vivo. Discussion and conclusions Our results demonstrate a widespread contamination of mineral water with xenoestrogens that partly originates from compounds leaching from the plastic packaging material. These substances possess potent estrogenic activity in vivo in a molluskan sentinel. Overall, the results indicate that a broader range of foodstuff may be contaminated with endocrine disruptors when packed in plastics. Keywords Endocrine disrupting chemicals - Estradiol equivalents - Human exposure - In vitro effects - In vivo effects - Mineral water - Plastic bottles - Plastic packaging - Polyethylene terephthalate - Potamopyrgus antipodarum - Yeast estrogen screen - Xenoestrogens
Impact of protected areas and land use on regeneration of Acacia woodland’s in Eastern Burkina Faso
(2008)
Regeneration success, persistence strategies (seedlings vs. coppicing), and population trend of Acacia spp. were tested under two land-use regimes in eastern Burkina Faso: (i) protected areas shielded to livestock grazing pressure, to logging, and using early annual fire as a management system; (ii) areas with high human impact (heavily and extensive livestock grazing, harvesting for wood and for medicinal plants). Generally, a good regeneration rate of Acacia species was observed in protected areas and a poor regeneration rate in areas with high human impact. Nevertheless, some species affiliated to the subgenus Aculeiferum as A. dudgeoni and A. polyacantha showed a good regeneration under both land use regimes. Juvenile plants less than 25 cm height of A. dudgeoni and A. gourmaensis increased by 116 to 50 % in areas with human impact as compared to their populations in protected areas. With SCD slopes varying from –0.40 to -0.70, the protected Acacia woodland displayed a stable population structure due to abundance of recruitment, and coppicing persistence (more common in the subgenus Aculeiferum) favoured by early annual fire. Consequently, the protected areas are favourable for Acacia woodland regeneration. Conversely, SCD slopes are positive or close to zero in areas of anthropogenic regime and showed a declining population, especially more marked with the subgenus Acacia due to permanent seed and seedling removal by livestock grazing. Nevertheless, the number of seedlings of some species was higher in areas under human pressure than in protected areas, especially for the subgenus Aculeiferum, improving the genetic variability and thus the long-term maintenance of the population. Key words: Acacia, early fire, land use, sudano-sahelien zone
We welcome critical appraisals that help to provide balance; however, Dietrich et al. gave an unjustified reproach. We feel that Dietrich’s position is severely compromised because he serves as an expert for the bisphenol A (BPA) Industry Group (Brussels, Belgium). We would like to respond to the issues raised by Dietrich et al., as well as to their oversights and inappropriate interpretations of our findings...
There is a growing body of evidence that indicates common environmental pollutants are capable of disrupting reproductive and developmental processes by interfering with the actions of endogenous hormones. Many reports of endocrine disruption describe changes in the normal development of organs and tissues that are consistent with genetic damage, and recent studies confirm that many chemicals classified to have hormone-modulating effects also possess carcinogenic and mutagenic potential. To date, however, there have been no conclusive examples linking genetic damage with perturbation of endocrine function and adverse effects in vivo. Here, we provide the first evidence of DNA damage associated with the development of imposex (the masculinization of female gastropods considered to be the result of alterations to endocrine-mediated pathways) in the dog-whelk Nucella lapillus. Animals (n = 257) that displayed various stages of tributyltin (TBT)-induced imposex were collected from sites in southwest England, and their imposex status was determined by physical examination. Linear regression analysis revealed a very strong relationship (correlation coefficient of 0.935, p < 0.0001) between the degree of imposex and the extent of DNA damage (micronucleus formation) in hemocytes. Moreover, histological examination of a larger number of dog-whelks collected from sites throughout Europe confirmed the presence of hyperplastic growths, primarily on the vas deferens and penis in both TBT-exposed male snails and in females that exhibited imposex. A strong association was found between TBT body burden and the prevalence of abnormal growths, thereby providing compelling evidence to support the hypothesis that environmental chemicals that affect reproductive processes do so partly through DNA damage pathways.
Previous investigations have shown that bisphenol A (BPA) induces a superfeminization syndrome in the freshwater snail Marisa cornuarietis at concentrations as low as 1 μg/L. Superfemales are characterized by the formation of additional female organs, enlarged accessory sex glands, gross malformations of the pallial oviduct, and a stimulation of egg and clutch production, resulting in increased female mortality. However, these studies were challenged on the basis of incomplete experimentation. Therefore, the objective of the current approach was to bridge several gaps in knowledge by conducting additional experiments. In an initial series of experiments, study results from the reproductive phase of the snails were evaluated in the sub-micrograms per liter range. Before and after the spawning season, superfemale responses were observed [NOEC (no observed effect concentration) 7.9 ng/L, EC10 (effective concentration at 10%) 13.9 ng/L], which were absent during the spawning season. A further experiment investigated the temperature dependence of BPA responses by exposing snails at two temperatures in parallel. The adverse effect of BPA was at least partially masked at 27°C (EC10 998 ng/L) when compared with 20°C (EC10 14.8 ng/L). In M. cornuarietis, BPA acts as an estrogen receptor (ER) agonist, because effects were completely antagonized by a co-exposure to tamoxifen and Faslodex. Antiandrogenic effects of BPA, such as a significant decrease in penis length at 20°C, were also observed. Competitive receptor displacement experiments indicate the presence of androgen- and estrogen-specific binding sites. The affinity for BPA of the estrogen binding sites in M. cornuarietis is higher than that of the ER in aquatic vertebrates. The results emphasize that prosobranchs are affected by BPA at lower concentrations than are other wildlife groups, and the findings also highlight the importance of exposure conditions.
In an effort to link quantitative morphometric information with molecular data on the population level, we have analysed 19 populations of the conchologically variable land snail Candidula unifasciata from across the species range for variation in quantitative shell traits and at the mitochondrial 16S ribosomal (r)DNA locus. In genetic analysis, including 21 additional populations, we observed two fundamental haplotype clades with an average pairwise sequence divergence of 0.209 ± 0.009 between clades compared to 0.017 ± 0.012 within clades, suggesting the presence of two different evolutionary lineages. Integrating additional shell material from the Senckenberg Malacological Collection, a highly significant discriminant analysis on the morphological shell traits with fundamental haplotype clades as grouping variable suggested that the less frequent haplotype corresponds to the described subspecies C. u. rugosiuscula, which we propose to regard as a distinct species. Both taxa were highly subdivided genetically (FST = 0.648 and 0.777 P < 0.001). This was contrasted by the partition of morphological variance, where only 29.6% and 21.9% of the variance were distributed among populations, respectively. In C. unifasciata, no significant association between population pairwise FST estimates and corresponding morphological fixation indices could be detected, indicating independent evolution of the two character sets. Partial least square analysis of environmental factors against shell trait variables in C. u. unifasciata revealed significant correlations between environmental factors and certain quantitative shell traits, whose potential adaptational values are discussed.
Population structure was estimated in a continuous population of a small land snail (Trochoidea geyeri). Mark-recapture experiments and randomly amplified polymorphic DNA analyses indicate that the population structure can be described by the isolation by distance model of Wright (1946). Estimates of density and dispersal suggest a neighbourhood size of 70-208 individuals on an area of 13-21 m². A principal component analysis of the randomly amplified polymorphic DNA data reveals clinal variation of genetic composition across the population, as predicted by the neighbourhood concept. An analysis of molecular variance indicates substantial genetic structuring. Comparisons of the genetic distances, expressed as euclidean distances among individuals, versus the geographic distance between sampling sites yield a highly significant positive correlation (Mantel test: r = 0.567, p<0.0001). The revealed pattern of populational subdivision on a microgeographic scale seems to be one of the principal processes generating and maintaining genetic diversity within populations of small land gastropods.
In the present study the population genetic structure of the terrestrial snail Pomatias elegans was related to habitat structure on a microspatial scale. The genetic variability of 1607 individuals from 51 sampling sites in five different populations in Provence, France, was studied with an allozyme marker using population genetic methods, Mantel tests and spatial autocorrelation techniques were applied to different connectivity networks accounting for the structural features of the landscape. It is suggested that the population structure is, to a large extent, a function of the habitat quality, quantified as population density, and of the spatial arrangement of the habitat in the landscape and not of the geographical distance per se. In fragmented habitats, random genetic drift was the prevailing force for sampling sites separated by a few hundred meters.
The study of organisms with restricted dispersal abilities and presence in the fossil record is particularly adequate to understand the impact of climate changes on the distribution and genetic structure of species. Trochoidea geyeri (Soós 1926) is a land snail restricted to a patchy, insular distribution in Germany and France. Fossil evidence suggests that current populations of T. geyeri are relicts of a much more widespread distribution during more favourable climatic periods in the Pleistocene. Results: Phylogeographic analysis of the mitochondrial 16S rDNA and nuclear ITS-1 sequence variation was used to infer the history of the remnant populations of T. geyeri. Nested clade analysis for both loci suggested that the origin of the species is in the Provence from where it expanded its range first to Southwest France and subsequently from there to Germany. Estimated divergence times predating the last glacial maximum between 25–17 ka implied that the colonization of the northern part of the current species range occurred during the Pleistocene. Conclusion: We conclude that T. geyeri could quite successfully persist in cryptic refugia during major climatic changes in the past, despite of a restricted capacity of individuals to actively avoid unfavourable conditions.