Habitat loss and fragmentation are among the biggest threats to biodiversity. Anthropogenic habitat fragmentation leads to small and isolated remnant plant and animal populations. The combination of increased random genetic drift, inbreeding, and reduced gene flow may substantially reduce genetic variation of remnant populations. However, the magnitude of these responses may depend on several poorly understood factors including organism group, habitat type of both the fragment and the surrounding matrix, life-history traits, and time since fragmentation. We compiled data for 83 plant and 52 animal species and conducted a meta-analysis following best practices to evaluate how these factors mediate the effects of anthropogenic habitat fragmentation. We calculated 206 effect sizes as correlations between one of four measures of population-level genetic diversity and fragment area. All analyses were repeated using models of increasing complexity (traditional random-effects models, multilevel models accounting for non-independent data, and multilevel models additionally correcting for phylogenetic relatedness). We confirmed that anthropogenic habitat fragmentation has overall negative effects on genetic diversity of organisms. Our meta-analysis shows, however, that plant species responded in general stronger to fragmentation than animal species and that the largest negative impacts of fragmentation occurred in tropical and temperate forest fragments, surrounded by a non-forest matrix. In contrast, we found only weak responses in non-forest fragments. Genetic diversity measured as mean number of alleles (A) showed the strongest response to fragmentation. Expected heterozygosity (He) and percentage of polymorphic loci (PLP) showed similar but weaker responses. In contrast, our meta-analysis indicated that inbreeding (Fis) was not measurably affected by anthropogenic habitat fragmentation. Additionally, our models revealed that effects on genetic diversity became stronger with age of fragments: We found significant negative responses for fragments older than 50 yr but not for those more recently isolated. Our meta-analyses also showed that currently animals are underrepresented in the literature on genetic effects of anthropogenic fragmentation, as are certain geographical regions and habitat types. We expect that future field studies using state-of-the-art approaches will provide further evidence of negative genetic effects, which may reinforce the here reported patterns, even for groups not yet studied.
Summary 1. Field observations in the Swiss Jura mountains showed that males and females of the bivoltine Adonis Blue butterfly Lysandra bellargus Rott. differed significantly in their flower visitation patterns. 2. In both generations, females visited a broader range of available nectar plants than did males. The specific flower visitation patterns of males and females were not affected by the general availability and abundance of potential nectar plant species during both flight periods, indicating high selectivity for nectar plants by both males and females. 3. In addition, the sexes differed in their nectar foraging behaviours: distances between successively visited flowers were significantly longer in males than in females, indicating that male and female butterflies have different foraging strategies. 4. Investigations of nectar characteristics showed that the sexes preferred flowers with different nectar compositions. Males of both generations preferred flowers with high proportions of sucrose and high amounts of total sugar, whereas females preferred flowers with high portions of glucose in their nectar, and, in the spring generation, flowers rich in amino acids. 5. Flowers visited exclusively by males or females in spring differed significantly in their amino acid composition. 6. This clear‐cut pattern did not hold for the autumn generation, most probably due to the limited availability of flowers. 7. The observed nectar foraging patterns underline the importance of adult feeding for longevity and reproduction in butterflies. The findings are particularly relevant for conservation, because L. bellargus is an increasingly threatened species in many European countries.
The fragmentation of natural habitats is generally considered to be a major threat to biodiversity. We investigated short-term responses of vascular plants (grasses and forbs) and four groups of invertebrates (ants, butterflies, grasshoppers and gastropods) to experimental fragmentation of calcareous grassland in the north-western Jura mountains, Switzerland. Three years after the initiation of fragmentation - which was created and maintained by mowing the area between the fragments - we compared species richness, diversity and composition of the different groups and the abundance of single species in fragments of different size (area: 20.25 m, 2.25 m and 0.25 m) with those in corresponding control plots. The abundances of 19 (29%) of the 65 common species examined were affected by fragmentation. However, the experimental fragmentation affected different taxonomic groups and single species to a different extent. Butterflies, the most mobile animals among the invertebrates studied, reacted most sensitively: species richness and foraging abundances of single butterfly species were lower in fragments than in control plots. Of the few other taxonomic groups or single species that were affected by the experimental fragmentation, most had a higher species richness or abundance in fragments than in control plots. This is probably because the type of fragmentation used is beneficial to some plants via decreased competition intensity along the fragment edges, and because some animals may use fragments as retreats between foraging bouts into the mown isolation area.
Exploring the effects of allelopathic plant chemicals on the growth of native vegetation is essential to understand their ecological roles and importance in exotic plant invasion. Naphthoquinones have been identified as potential growth inhibitors produced by Impatiens glandulifera, an exotic annual plant that recently invaded temperate forests in Europe. However, naphthoquinone release and inhibitory potential have not been examined. We quantified the naphthoquinone content in cotyledons, leaves, stems, and roots from plants of different ages of both the invasive I. glandulifera and native Impatiens noli-tangere as well as in soil extracts and rainwater rinsed from leaves of either plant species by using ultra-high pressure liquid chromatography-mass spectrometry (UHPLC-MS). We identified the compound 2-methoxy-1,4-naphthoquinone (2-MNQ) exclusively in plant organs of I. glandulifera, in resin bags buried into the soil of patches invaded by I. glandulifera, and in rainwater rinsed from its leaves. This indicates that 2-MNQ is released from the roots of I. glandulifera and leached from its leaves by rain. Specific bioassays using aqueous shoot and root extracts revealed a strong inhibitory effect on the germination of two native forest herbs and on the mycelium growth of three ectomycorrhiza fungi. These findings suggest that the release of 2-MNQ may contribute to the invasion success of I. glandulifera and support the novel weapons hypothesis.
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