Ane T. Laugen

Latitudinal countergradient variation in the common frog (Rana temporaria) development rates – evidence for local adaptation

Adaptive genetic differentiation along a climatic gradient as a response to natural selection is not necessarily expressed at phenotypic level if environmental effects on population mean phenotypes oppose the genotypic effects. This form of cryptic evolution – called countergradient variation – has seldom been explicitly demonstrated for terrestrial vertebrates. We investigated the patterns of phenotypic and genotypic differentiation in developmental rates of common frogs (Rana temporaria) along a ca. 1600 km latitudinal gradient across Scandinavia. Developmental rates in the field were not latitudinally ordered, but displayed large variation even among different ponds within a given latitudinal area. In contrast, development rates assessed in the laboratory increased strongly and linearly with increasing latitude, suggesting a genetic capacity for faster development in the northern than the southern larvae. Experiments further revealed that environmental effects (temperature and food) could easily override the genetic effects on developmental rates, providing a possible mechanistic explanation as to why the genetic differentiation was not seen in the samples collected from the wild. Our results suggest that the higher developmental rates of the northern larvae are likely to be related to selection stemming from seasonal time constrains, rather than from selection dictated by low ambient temperatures per se. All in all, the results provide a demonstration of environmental effects concealing substantial latitudinally ordered genetic differentiation understandable in terms of adaptation to clinal variation in time constrains.

Latitudinal divergence of common frog (Rana temporaria) life history traits by natural selection: evidence from a comparison of molecular and quantitative genetic data

The relative roles of natural selection and direct environmental induction, as well as of natural selection and genetic drift, in creating clinal latitudinal variation in quantitative traits have seldom been assessed in vertebrates. To address these issues, we compared molecular and quantitative genetic differentiation between six common frog (Rana temporaria) populations along an approximately 1600 km long latitudinal gradient across Scandinavia. The degree of population differentiation (QST ≈ 0.81) in three heritable quantitative traits (age and size at metamorphosis, growth rate) exceeded that in eight (neutral) microsatellite loci (FST = 0.24). Isolation by distance was clear for both neutral markers and quantitative traits, but considerably stronger for one of the three quantitative traits than for neutral markers. QST estimates obtained using animals subjected to different rearing conditions (temperature and food treatments) revealed some environmental dependency in patterns of population divergence in quantitative traits, but in general, these effects were weak in comparison to overall patterns. Pairwise comparisons of FST and QST estimates across populations and treatments revealed that the degree of quantitative trait differentiation was not generally predictable from knowledge of that in molecular markers. In fact, both positive and negative correlations were observed depending on conditions where the quantitative genetic variability had been measured. All in all, the results suggest a very high degree of genetic subdivision both in neutral marker genes and genes coding quantitative traits across a relatively recently (< 9000 years) colonized environmental gradient. In particular, they give evidence for natural selection being the primary agent behind the observed latitudinal differentiation in quantitative traits.

ANTIPREDATOR DEFENSES ALONG A LATITUDINAL GRADIENT IN RANA TEMPORARIA

Antipredator defenses are expected to decrease toward higher latitudes because predation rates are predicted to decrease with latitude. However, latitudinal variation in predator avoidance and defense mechanisms has seldom been studied. We studied tadpole antipredator defenses in seven Rana temporaria populations collected along a 1500-km latitudinal gradient across Sweden, along which previous studies have found increasing tadpole growth and development rates. In a laboratory common garden experiment, we measured behavioral and morphological defenses by raising tadpoles in the presence and absence of a predator (Aeshna dragonfly larva) in two temperature treatments. We also estimated tadpole survival in the presence of free-ranging predators and compared predator densities between R. temporaria breeding ponds situated at low and high latitudes. Activity and foraging were generally positively correlated with latitude in the common garden experiment. While all populations responded to predator presence by decreasing activity and foraging, high-latitude populations maintained higher activity levels in the presence of the predator. All populations exhibited defensive morphology in body and tail shape. However, whereas tail depth tended to increase with latitude in the presence of predator, it did not change with latitude in the absence of the predator. Predator presence generally increased larval period and decreased growth rate. In the southern populations, predator presence tended to have a negative effect on metamorphic size, whereas in the northern populations predators had little or a positive effect on size. Latitude of origin had a strong effect on survival in the presence of a free-ranging predator, with high-latitude tadpoles experiencing higher mortality than those from the low latitudes. In the wild, predator densities were significantly lower in high-latitude than in mid-latitude breeding ponds. Although the higher activity level in the northern populations seems to confer a significant survival disadvantage under predation risk, it is probably needed to maintain the high growth and development rates. However, the occurrence of R. temporaria at high latitudes may be facilitated by the lower predator densities in the north.

Evolutionary impact assessment: accounting for evolutionary consequences of fishing in an ecosystem approach to fisheries management

Managing fisheries resources to maintain healthy ecosystems is one of the main goals of the ecosystem approach to fisheries (EAF). While a number of international treaties call for the implementation of EAF, there are still gaps in the underlying methodology. One aspect that has received substantial scientific attention recently is fisheries-induced evolution (FIE). Increasing evidence indicates that intensive fishing has the potential to exert strong directional selection on life-history traits, behaviour, physiology, and morphology of exploited fish. Of particular concern is that reversing evolutionary responses to fishing can be much more difficult than reversing demographic or phenotypically plastic responses. Furthermore, like climate change, multiple agents cause FIE, with effects accumulating over time. Consequently, FIE may alter the utility derived from fish stocks, which in turn can modify the monetary value living aquatic resources provide to society. Quantifying and predicting the evolutionary effects of fishing is therefore important for both ecological and economic reasons. An important reason this is not happening is the lack of an appropriate assessment framework. We therefore describe the evolutionary impact assessment (EvoIA) as a structured approach for assessing the evolutionary consequences of fishing and evaluating the predicted evolutionary outcomes of alternative management options. EvoIA can contribute to EAF by clarifying how evolution may alter stock properties and ecological relations, support the precautionary approach to fisheries management by addressing a previously overlooked source of uncertainty and risk, and thus contribute to sustainable fisheries.

Adaptive phenotypic plasticity in timing of metamorphosis in the common frog Rana temporaria

Abstract Unpredictable environments are expected to select for adaptive plasticity in traits enabling adjustment of phenotype to prevailing environmental conditions. Common frogs (Rana temporaria) breed frequently in ponds, which dry up before the aquatic larvae have metamorphosed, and consequently, plasticity in timing of metamorphosis in response to pond drying could be adaptive. We investigated the responses of half- and full-sib R. temporaria larvae to simulated pond drying in a factorial experiment to test whether there is adaptive phenotypic plasticity in timing of metamorphosis, and whether this plasticity is genetically determined. As expected under the adaptive hypothesis, we found that larvae exposed to the decreasing water treatment metamorphosed significantly faster than their sibs in the constant water treatments. Furthermore, age and size at metamorphosis were positively correlated in the constant water treatments, but negatively correlated in the decreasing water treatment. However, larvae from decreasing water treatment metamorphosed, on average, at a smaller size as compared to larvae from the constant water treatments, even after controlling for variation in developmental time. Since smaller size at metamorphosis is likely to be related to reduced fitness, this indicates that faster development may trade off with other components of fitness. The results further show that the amount of food received during the four first days of development influenced age and size at metamorphosis, reinforcing the view that early life nutrition may have a significant impact on later life fitness. Although there was a genetic component to developmental rates, we found no evidence for genetic variation in plasticity. In accordance with evidence from other studies, our results suggest existence of adaptive phenotypic plasticity in amphibian development.

Latitudinal and temperature-dependent variation in embryonic development and growth in Rana temporaria

Variation in seasonal time constraints and temperature along latitudinal gradients are expected to select for life history trait differentiation, but information about the relative importance of these factors in shaping patterns of divergence in embryonic traits remains sparse. We studied embryonic survival, growth and development rates in the common frog (Rana temporaria) along a 1,400-km latitudinal gradient across Sweden by raising embryos from four populations in the laboratory at seven temperatures (9°C, 12°C, 15°C, 18°C, 21°C, 24°C, 27°C). We found significant differences in mean values of all traits between the populations and temperature treatments, but this variation was not latitudinally ordered. In general, embryonic survival decreased at the two highest temperatures in all populations, but less so in the southernmost as compared to the other populations. The northernmost population developed slowest at the lowest temperature, while the two mid-latitude populations were slowest at the other temperatures. Hatchling size increased with increasing temperature especially in the two northern populations, whereas the two southern populations showed peak hatchling size at 15°C. Analyses of within-population genetic variation with a half-sib design revealed that there was significant additive genetic variation in all traits, and egg size-related maternal effects were important in the case of hatchling size. Overall, our results indicate that unlike larval growth and development, variation in embryonic development and growth in R. temporaria cannot be explained in terms of a latitudinal gradient in season length. While adaptation to a latitudinal variation in temperature might have contributed to the observed differentiation in embryonic performance, the effects of other, perhaps more local environmental factors, seem to have overridden them in importance.

Quantitative genetics of larval life-history traits in Rana temporaria in different environmental conditions

The degree to which genetic variation in a given trait varies among different populations of the same species and across different environments has seldom been quantified in wild vertebrate species. We investigated the expression of genetic variability and maternal effects in three larval life-history traits of the amphibian Rana temporaria. In a factorial laboratory experiment, five widely separated populations (max. 1600 km) were subjected to two different environmental treatments. Animal model analyses revealed that all traits were heritable (h(2) approximately 0.20) in all populations and under most treatment combinations. Although the cross-food treatment genetic correlations were close to unity, heritabilities under a restricted food regime tended to be lower than those under an ad libitum food regime. Likewise, maternal effects (m(2) approximately 0.05) were detected in most traits, and they tended to be most pronounced under restricted food conditions. We detected several cross-temperature genetic and maternal effects correlations that were lower than unity, suggesting that genotype-environment interactions and maternal effect-environment interactions are a significant source of phenotypic variation. The results reinforce the perspective that although the expression of genetic and maternal effects may be relatively homogeneous across different populations of the same species, local variation in environmental conditions can lead to significant variation in phenotypic expression of quantitative traits through genotype-environment and maternal effect-environment interactions.

Allen’s rule revisited: quantitative genetics of extremity length in the common frog along a latitudinal gradient

Ecogeographical rules linking climate to morphology have gained renewed interest because of climate change. Yet few studies have evaluated to what extent geographical trends ascribed to these rules have a genetic, rather than environmentally determined, basis. This applies especially to Allen’s rule, which states that the relative extremity length decreases with increasing latitude. We studied leg length in the common frog (Rana temporaria) along a 1500 km latitudinal gradient utilizing wild and common garden data. In the wild, the body size–corrected femur and tibia lengths did not conform to Allen’s rule but peaked at mid‐latitudes. However, the ratio of femur to tibia length increased in the north, and the common garden data revealed a genetic cline consistent with Allen’s rule in some trait and treatment combinations. While selection may have shortened the leg length in the north, the genetic trend seems to be partially masked by environmental effects.

Endophilic Anopheles mosquitoes in Guinea Bissau, West Africa in relation to human housing conditions

Abstract Environmental risk factors associated with increased malaria mosquito (Anopheles) abundance indoors were studied in a suburban area, Antula, of Guinea Bissau, during the rainy seasons of 1993–1995. All bedrooms in 30 houses were searched for resting mosquitoes three times each year. The most abundant mosquito was An. gambiae s.s. Significantly greater numbers of resting mosquitoes were present in rooms with open eaves and in houses with a well on the compound. Pigs were the most common domestic animals in Antula. Presence of pigs in a house was associated with increased mosquito abundance in the bedrooms of the same house. The abundance of female mosquitoes also increased with increasing human biomass per square meter of bedroom area.

Prognostic value of using bee and hive debris samples for the detection of American foulbrood disease in honey bee colonies

The objective of this study was to compare how well the detection of Paenibacillus larvae in samples of live bees or in accumulated winter debris collected from the bottom of beehives relates to symptoms of American foulbrood in honey bee colonies. Fifty-eight colonies in one commercial beekeeping operation were inspected for disease symptoms and assayed for P. larvae using culture-based techniques and PCR. The results show that culture-based techniques are more accurate than recently published PCR methods for detecting the bacterium in clinically diseased colonies, and that the prognostic value of bacterial colony counts from bee samples is superior to colony counts from debris. However, if the objective is to monitor the prevalence of the bacterium irrespective of disease symptoms, the preferable method is PCR analysis of accumulated winter hive debris.