Population genetic structure of parthenogenetic flatworm populations with occasional sex

1. Sexual populations are expected to perform better in fluctuating environments than asexuals because recombination provides the potential to adapt to changing environments due to increased genetic variation. Nevertheless, some asexual species show comparably high levels of genotypic diversity. Such diversity might be achieved through gene flow between coexisting sexual and asexual populations or through sexual events within asexual populations. 2. Evidence for occasional sex in the flatworm Schmidtea polychroa was previously found at one specific site that is inhabited by parthenogenetic forms. There, varying rates of sex between subpopulations, reaching up to 12%, were observed. Past recurrent sexual processes left a significant genetic signature in the population genetic structure of this population. In the present study, we examined the population genetic structure of six independent metapopulations (lakes) of the freshwater planarian flatworm S. polychroa, to confirm the presence of occasional sex and that its population genetic consequences can be generalised. 3. Using microsatellites, we found varying rates of occasional sex among subpopulations. Metapopulations showed medium to high levels of genotypic diversity that correlated with the rate of sex. 4. We conclude that occasional sex has considerable consequences for population genetic structure of parthenogenetic species and promotes diversity that might allow response to the particular type of selection that is usually predicted to favour sexual reproduction. This reproductive strategy provides genetic characteristics required for selection to act on, and might, therefore, explain the success of this parthenogenetic species.     

Evaluation of approaches for measuring taxonomic completeness of lake profundal macroinvertebrate assemblages

1. Benthic macroinvertebrates (MI) are commonly used to assess freshwater ecosystems with the reference condition approach. Such assessments necessitate control for natural community variation, either by categorical typologies or by predictive models that have been widely and successfully developed for running water biota but not previously for lake profundal invertebrates. 2. We evaluated four modelling techniques [multivariate regression tree (MRT), limiting environmental differences, nonparametric multiplicative regression (NPMR) and River Invertebrate Prediction And Classification System (RIVPACS) and the operative Finnish lake typology for assessing taxonomic completeness (observed‐to‐expected number of taxa, O/E) of profundal MI assemblages. We used data from 74 and 33 minimally disturbed reference lake basins for calibration and validation of the approaches, respectively, and 72 test basins subject to various anthropogenic pressures to evaluate sensitivity to detect impact. Either all predicted taxa (threshold probability of capture P_t = 0+) or only those predicted to be captured with ≥0.25 probability were used to calculate O/E. 3. With P_t = 0.25, all four modelling approaches were accurate (mean O/E = 0.966–1.053) but imprecise (SD of O/E = 0.279–0.304) in predicting the fauna actually observed in validation sites. All models were subtly more precise than a null model (mean 1.038, SD 0.343) or the typology (1.046, 0.327). The taxon‐specific NPMR model was slightly more precise than the other three models based on site groupings. 4. The O/E values correlated relatively weakly (r = 0.55–0.86) among the approaches, which thus produced contrasting lake‐specific assessments, despite their seemingly comparable performances. Indeed, typology, suggesting that MI assemblages were impaired in 56% of test sites, was more sensitive than the other approaches (26–46%) as an indicator of human‐induced deterioration. However, this greater ostensible sensitivity seemed to be biased, as lake morphometry, a main driver of natural community variation, remained uncontrolled by the typology. 5. Generally, our exercise illustrates the inconclusiveness of the common validation criteria for the assessment methods. The apparent poor predictability of the profundal fauna, irrespective of the method, may partly stem from large observation error, which could be alleviated by more intensive sampling. However, instead of an O/E‐taxa index, some other metric encompassing quantitative aspects might be preferable for assessing these species‐poor communities.     

Aminopeptidase and phosphatase activities in basins of Lake Hiidenvesi dominated by cyanobacteria and in laboratory grown Anabaena

1. Extracellular enzyme activities were examined in freshwater basins representing a transition from hypertrophy to mesotrophy and in axenic cyanobacterial cultures to evaluate the ecological role of extracellular enzyme activities of cyanobacteria.
2. Aminopeptidase activity was related to the trophic status of the lake basins. The activity was highest in the most eutrophic basin and decreased in the less nutrient-rich basins. Cyanobacteria were the most important autotrophic organisms and aminopeptidase activity was positively associated with cyanobacterial biomass.
3. In an axenic Anabaena batch culture, nitrogenase activity was several orders of magnitude higher than leucine aminopeptidase activity. Nitrate did not have an effect on aminopeptidase activity or growth, but significantly reduced the rate of nitrogen fixation. A high phosphorus concentration at the beginning of the Anabaena batch-culture experiment resulted in reduced phosphatase activity.
4. In Lake Hiidenvesi, aminopeptidase activity probably originated mostly from attached bacteria and less so from cyanobacteria.     

Spatial patterns of litter decomposition in the littoral zone of boreal lakes

1. We studied the patterns of litter decomposition in lake littoral habitats and investigated whether decay rates, as an integrating proxy for environmental conditions in the sediment, would co‐vary with net carbon dioxide (CO₂) exchange and methane (CH₄) efflux. These gas fluxes are known to be sensitive to environmental conditions. Losses in the mass of cellulose, root, rhizome and moss litter were measured during 2 years in boreal littoral wetlands in Finland and compared with published data on concurrently measured gas fluxes. Four study sites covered a range of sediment types and hydrological conditions. 2. Decomposition was not linearly related to the duration of flooding but depended on sediment type. Readily decomposable litter fractions, such as cellulose and rhizome litter, lost mass at a faster rate in marshes with a longer period of flooding but wide water level fluctuations that hinder establishment of a Sphagnum cover, than in peat‐forming fens. In marshes, the mean first‐year mass losses were 83–99% and 19–62% for cellulose and rhizomes, respectively. In fens, the respective losses were 40–53% and 33%. In the first year, the loss in the mass of the more recalcitrant root litter did not differ between sites (mean 19–30%) and moss litter lost no mass. 3. The estimated first‐year carbon loss from belowground litter was about 0.1–0.3 times ecosystem respiration and roughly similar to net carbon gas (CO₂, CH₄) efflux, suggesting that vascular plants and recent plant residues contribute substantially to ecosystem release of carbon gases. On the other hand, at least 40% of the mass of the belowground litter remained on a littoral site after the first 2 years of decomposition. Slow decomposition may indicate the accumulation of organic‐rich sediments. The accumulated carbon could explain the excess CO₂ release found in most littoral sites. In continuously inundated sites decomposition rates were similar to those in periodically flooded sites, but ecosystem‐atmosphere CO₂ exchange fell to close to zero. This discrepancy implies that the released CO₂ is dissolved in water and may be exported into the pelagic zone of the lake.     

The seasonal dynamics and distribution of Chaoborus flavicans larvae in adjacent lake basins of different morphometry and degree of eutrophication

1.  Seasonal dynamics, spatial distribution and population size of the phantom midge Chaoborus flavicans in different parts of the eutrophic Lake Hiidenvesi (30.3 km²) were studied.
2.  Density of larvae was low in the shallow, most eutrophic parts of the lake, while the deep Kiihkelyksenselkä basin was inhabited by a dense population. In the deepest part of Kiihkelyksenselkä (33 m) density was 13 989 ± 3542 m^–2 in May, declined to 1102 ± 274 m^–2 in July and recovered to 7225 ± 1314 m^–2 by October. In spring and autumn the majority of larvae were benthic while, during high summer, few larvae were found in the sediment.
3.  Horizontal distribution fluctuated seasonally. On 3 June < 5% of the population inhabited areas shallower than 10 m. On 6 July the limnetic fraction was still restricted to regions deeper than 10 m, but 43% of benthic larvae were found between 6 and 10 m depths. In October both limnetic and benthic larvae were concentrated in areas deeper than 20 m.
4.  Within the lake, distribution was mainly regulated by stratification characteristics, degree of eutrophy being less important. The seasonal horizontal movements were probably induced by food shortage. Larvae could not meet their energetic demands in stratified areas and dispersed to shallower water, reducing predation risk by use of the benthic habitat.     

Age‐dependent responses to alarm calls depend on foraging activity in Willow Tits Poecile montanus

Survivorship in animals depends on both foraging activities and avoidance of predation, and thus behavioural decisions often reflect a trade‐off between predation risk and foraging efficiency. In this experimental study, we compared behavioural responses of free‐living adult and juvenile Willow Tits Poecile montanus to a conspecific alarm call in two treatments. The alarm call was played back when a focal bird was either not feeding, or feeding on a sunflower seed on the middle part of a spruce branch. When feeding at the time of the alarm call, juveniles more often stayed motionless or moved shorter distances than adults. Our results suggest that in hierarchical groups, juveniles are forced to take greater risks to maintain access to food or lack experience to optimize between food and safety.     

Effects of short- and long-term water-level drawdown on the populations and activity of aerobic decomposers in a boreal peatland

We analysed the response of microbial communities, characterized by phospholipid fatty acids (PLFAs), to changing hydrological conditions at sites with different nutrient levels in a southern boreal peatland. Although PLFAs of Gram‐negative bacteria were characteristic of the peatland complex, microbial communities differed among sites (ombrotrophic bog, oligotrophic fen, mesotrophic fen) and sampling depths (0–5, 5–10, 10–20, 20–30 cm). The microbial communities in each site changed significantly following water‐level drawdown. The patterns of change varied among sites and sampling depths. The relative proportion of Gram‐negative bacteria decreased in the upper 10 cm but increased in deeper layers of the fen sites. Fungi benefited from water‐level drawdown in the upper 5 cm of the mesotrophic fen, but suffered in the drier surfaces of the ombrotrophic bog, especially in the 5–10 cm layer. In contrast, actinobacteria suffered from water‐level drawdown in the mesotrophic fen, but benefited in the drier surfaces of the ombrotrophic bog. Basal respiration rate correlated positively with pH and fungal PLFA, and negatively with depth. We suggest that the changes in microbial community structure after persistent water‐level drawdown follow not only the hydrological conditions but also the patterns of vegetation change. Our results imply that changes in structure and activity of the microbial community in response to climate change will be strongly dependent on the type of peatland.     

Genetic variation in sexual and clonal lineages of a freshwater snail

Sexual reproduction within natural populations of most plants and animals continues to remain an enigma in evolutionary biology. That the enigma persists is not for lack of testable hypotheses but rather because of the lack of suitable study systems in which sexual and asexual females coexist. Here we review our studies on one such organism, the freshwater snail Potamopyrgus antipodarum (Gray). We also present new data that bear on hypotheses for the maintenance of sex and its relationship to clonal diversity. We have found that sexual populations of the snail are composed of diploid females and males, while clonal populations are composed of a high diversity of triploid apomictic females. Sexual and asexual individuals coexist in stable frequencies in many ‘mixed’ populations; genetic data indicate that clones from these mixed populations originated from the local population of sexual individuals without interspecific hybridization. Field data show that clonal and sexual snails have completely overlapping life histories, but individual clonal genotypes are less variable than individuals from the sympatric sexual population. Field data also show segregation of clones among depth‐specific habitat zones within a lake, but clonal diversity remains high even within habitats. A new laboratory experiment revealed extensive clonal variation in reproductive rate, a result which suggests that clonal diversity would be low in nature without some form of frequency‐dependent selection. New results from a long‐term field study of a natural, asexual population reveal that clonal diversity remained nearly constant over a 10‐year period. Nonetheless, clonal turnover occurs, and it occurs in a manner that is consistent with parasite‐mediated, frequency‐dependent selection. Reciprocal cross‐infection experiments have further shown that parasites are more infective to sympatric host snails than to allopatric snails, and that they are also more infective to common clones than rare clones within asexual host populations. Hence we suggest that sexual reproduction in these snails may be maintained, at least in part, by locally adapted parasites. Parasite‐mediated selection possibly also contributes to the maintenance of local clonal diversity within habitats, while clonal selection may be responsible for the distribution of clones among habitats. © 2003 The Linnean Society of London. Biological Journal of the Linnean Society 2003, 79 , 165–181.     

Land cover change 2002–2005 in Borneo and the role of fire derived from MODIS imagery

Borneo has experienced heavy deforestation and forest degradation during the past two decades. In this study the Moderate Resolution Imaging Spectroradiometer was used to monitor land cover change in Borneo between 2002 and 2005 in order to assess the current extent of the forest cover, the deforestation rate and the role of fire. Using Landsat and ground observation for validation it was possible to discriminate 11 land cover classes. In 2002 57% of the land surface of Borneo was covered with forest of which 74% was dipterocarp and more than 23% peat swamp forest. The average deforestation rate between 2002 and 2005 was 1.7% yr^{− 1}. The carbon-rich ecosystem of peat swamp forests showed a deforestation rate of 2.2%. Almost 98% of all deforestation occurred within a range of 5 km to the forest edge. Fire is highly correlated with land cover changes. Most fires were detected in degraded forests. Ninety-eight per cent of all forest fires were detected in the 5 km buffer zone, underlining that fire is the major driver for forest degradation and deforestation.     

Spawning coloration and sperm quality in a large lake population of Arctic charr (Salmonidae: Salvelinus alpinus L.)

The modern theories of sexual selection predict that male sexual ornaments may have evolved as reliable signals of male fertilization efficiency. However, among the studies of fishes with external fertilization, the results have yielded ambiguous evidence. In the present study, we present data on the phenotypic relationships between red spawning coloration and ejaculate quality (spermatocrit, sperm motility) from Arctic charr, Salvelinus alpinus. We studied two generations (F₁ and F₂) of males from a large lake population, reared in a standardized hatchery environment, to determine whether differential hatchery history, or duration of hatchery selection, affected the variation in ejaculate characteristics or abdominal coloration. After controlling for body length, there was no difference between the hatchery generations in these traits. However, the degree of redness increased with fish size. We found a positive correlation between sperm velocity and sperm longevity, indicating a functional integration between these sperm features across generations. Sperm velocity was also positively correlated with male redness. Therefore, the finding obtained in the present study suggests that the carotenoid‐based ornamentation in Arctic charr may provide information about differences between males in their fertilization potential. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 794–802.