Panu Välimäki

Latitudinal insect body size clines revisited: a critical evaluation of the saw‐tooth model

1. Insect body size is predicted to increase with decreasing latitude because time available for growth increases. In insects with changing voltinism (i.e. number of generations per season), sharp decreases in development time and body size are expected at season lengths where new generations are added to the phenology of a species, giving rise to saw-tooth clines in these traits across latitudes. Growth rate variation may affect the magnitude of variation in body size or even reverse the saw-tooth cline. 2. In this study, we analyse latitudinal body size clines in four geometrid moths with changing voltinism in a common laboratory environment. In addition to body size, we measured larval development time and growth rate and genetic correlations among the three traits. 3. The patterns of clinal variation in body size were diverse, and the theory was not supported even when saw-tooth body size clines were found. Larval development time increased and growth rate decreased consistently with increasing season length, the clines in these traits being uniform. 4. The consistencies of development time and growth rate clines suggest a common mechanism underlying the observations. Such a mechanism is discussed in relation to the complex interdependencies among the traits.

SEASONALITY MAINTAINS ALTERNATIVE LIFE-HISTORY PHENOTYPES

Many organisms express discrete alternative phenotypes (polyphenisms) in relation to predictable environmental variation. However, the evolution of alternative life‐history phenotypes remains poorly understood. Here, we analyze the evolution of alternative life histories in seasonal environments by using temperate insects as a model system. Temperate insects express alternative developmental pathways of diapause and direct development, the induction of a certain pathway affecting fitness through its life‐history correlates. We develop a methodologically novel and holistic simulation model and optimize development time, growth rate, body size, reproductive effort, and adult life span simultaneously in both developmental pathways. The model predicts that direct development should be associated with shorter development time (duration of growth) and adult life span, higher growth rate and reproductive effort, smaller body size as well as lower fecundity compared to the diapause pathway, because the two generations divide the available time unequally. These predictions are consistent with many empirical data. Our analysis shows that seasonality alone can explain the evolution of alternative life histories.

Divergent timing of egg‐laying may maintain life history polymorphism in potentially multivoltine insects in seasonal environments

The length of the favourable season determines voltinism in insect populations. In some insects, there is variation in fecundity and timing of reproduction among females. If the length of the favourable season does not allow all offspring to develop into adults without diapause, the benefits of high early fecundity may outweigh the associated cost of low lifetime fecundity. We tested this by exploring mating frequencies of Pieris napi females along a latitudinal gradient in different generations. Pieris napi is a bivoltine butterfly, and genetically polyandrous females enjoy higher lifetime fecundity than monandrous ones. Polyandry is, however, coupled with a relatively low early fecundity. We found that monandrous females are more likely to produce an additional generation than polyandrous ones under conditions that allow production of only a partial summer generation. Monandrous females were also the first to emerge and slightly over‐represented in the summer generation under conditions that allow the development of a complete summer generation. Further, a stochastic model shows that variation in the timing of reproduction between strategies is sufficient to explain the observed patterns. Thus, seasonality may counter‐select against polyandry, or more generally against low early reproductive rate, and promote maintenance of polymorphism in life history strategies.

Latitudinal clines in alternative life histories in a geometrid moth

The relative roles of genetic differentiation and developmental plasticity in generating latitudinal gradients in life histories remain insufficiently understood. In particular, this applies to determination of voltinism (annual number of generations) in short‐lived ectotherms, and the associated trait values. We studied different components of variation in development of Chiasmia clathrata (Lepidoptera: Geometridae) larvae that originated from populations expressing univoltine, partially bivoltine or bivoltine phenology along a latitudinal gradient of season length. Indicative of population‐level genetic differentiation, larval period became longer while growth rate decreased with increasing season length within a particular phenology, but saw‐tooth clines emerged across the phenologies. Indicative of phenotypic plasticity, individuals that developed directly into reproductive adults had shorter development times and higher growth rates than those entering diapause. The most marked differences between the alternative developmental pathways were found in the bivoltine region suggesting that the adaptive correlates of the direct development evolve if exposed to selection. Pupal mass followed a complex cline without clear reference to the shift in voltinism or developmental pathway probably due to varying interplay between the responses in development time and growth rate. The results highlight the multidimensionality of evolutionary trajectories of life‐history traits, which either facilitate or constrain the evolution of integrated traits in alternative phenotypes.

Geographical variation in host use of a blood-feeding ectoparasitic fly: implications for population invasiveness

Invasive generalist ectoparasites provide a tool to study factors affecting expansion rates. An increase in the number of host species may facilitate geographic range expansion by increasing the number of suitable habitats and by affecting local extinction and colonization rates. A geographic perspective on parasite host specificity and its implications on range expansion are, however, insufficiently understood. We conducted a field study to explore if divergent host specificity could explain the observed variation in expansion rates between Fennoscandian populations of the deer ked (Lipoptena cervi), which is a blood-feeding ectoparasitic fly of cervids. We found that the rapidly expanding eastern population in Finland appears to specialize on moose, whereas the slowly expanding western population in Norway breeds successfully on both moose and roe deer. The eastern population was also found to utilize the wild forest reindeer as an auxiliary host, but this species is apparently of low value for L. cervi in terms of adult maintenance, reproductive output and offspring quality. Abundant numbers of roe deer and white-tailed deer were observed to be apparently uninfected in Finland, suggesting that host use is not a plastic response to host availability, but rather a consequence of population-level evolutionary changes. Locally compatible hosts were found to be the ones sharing a long history with the deer ked in the area. Cervids that sustained adult deer keds also allowed successful reproduction. Thus, host use is probably determined by the ability of the adult to exploit particular host species. We conclude that a wide host range alone does not account for the high expansion rate or wide geographic distribution of the deer ked, although loose ecological requirements would increase habitat availability.

Genetic and phenotypic variation in juvenile development in relation to temperature and developmental pathway in a geometrid moth

Life histories show genetic population‐level variation due to spatial variation in selection pressures. Phenotypic plasticity in life histories is also common, facilitating fine‐tuning of the phenotype in relation to the prevailing selection regime. In multivoltine (≥ 2 generations per year) insects, individuals following alternative developmental pathways (diapause/direct development) experience different selection regimes. We studied the genetic and phenotypic components of juvenile development in Cabera exanthemata (Lepidoptera: Geometridae) in a factorial split‐brood experiment. F2 offspring of individuals originating from populations in northern and central Finland were divided among manipulations defined by temperature (14 °C/20 °C) and day length (24 h/15 h). Short day length invariably induced diapause, whereas continuous light almost invariably induced direct development in both regions, although northern populations are strictly univoltine in the wild. Individuals from northern Finland had higher growth rates, shorter development times and higher pupal masses than individuals from central Finland across the conditions, indicating genetic differences between regions. Individuals that developed directly into adults tended to have higher growth rates, shorter development times and higher pupal masses than those entering diapause, indicating phenotypic plasticity. Temperature‐induced plasticity was substantial; growth rate was much higher, development time much shorter and pupal mass higher at 20 °C than at 14 °C. The degree of plasticity in relation to developmental pathway was pronounced at 20 °C in growth rate and development time and at 14 °C in pupal mass, emphasizing multidimensionality of reaction norms. The observed genetic variation and developmental plasticity seem adaptive in relation to time‐stress due to seasonality.

Seasonal Clines of Evolutionarily Stable Reproductive Effort in Insects

For ectotherms, growth and reproduction are possible only during a limited period of the year in seasonal environments. In insects, fitness is generally maximized by producing as many generations as possible within a season, and in many species, the number of generations produced (voltinism) increases with increasing season length. In this study, we analyzed variation in adult life histories in insects along a climatic gradient. The analyzed trait is reproductive effort (resource allocation to reproduction). We begin by formalizing the trade‐off between current reproduction and subsequent survival generated by reproductive effort. It appeared that reproductive effort is correlated positively with early fecundity and negatively with lifetime fecundity and life span. Then, deriving from that trade‐off, we analyze the evolutionary stability of different schedules of age‐specific fecundity that are generated by divergent reproductive effort. The analysis was carried out in season lengths that promote either univoltine or bivoltine phenology. The evolutionarily stable reproductive effort decreases with increasing season length in both phenologies, with a sudden increase when a change from univoltine to partially bivoltine phenology takes place. Reproductive effort responds strongly to changing phenology when density‐dependent mortality occurs during diapause and weakly when juvenile mortality is density dependent.

The value of open power line habitat in conservation of ground beetles (Coleoptera: Carabidae) associated with mires

The draining of mires for silvicultural purposes has caused one of the most dramatic changes in the landscape during the last century in Finland. To study the effects of mire drainage, carabid beetle assemblages were sampled using pitfall trapping in three different mire habitat types. Carabids were sampled from mires in their natural state, drained mires and drained mires with an open power line to see whether the cleared power line can serve as an alternative habitat for mire dependent carabids. The draining of mires greatly increases the species richness of the carabid assemblages. Yet, the conservation value of the environment has dropped following the draining, since only common and abundant forest carabids have benefited from human impact. The role of the open power line as an alternative habitat for mire specialists remains questionable. A few carabid species have, however, benefited from the open habitat of the power line. The vegetation structure had a significant effect in determining the compositions of the carabid assemblages on the studied habitat types. It seems that mire dwelling carabids cannot survive on the drained mires, unless at least some characteristics, other than the mere openness of the cleared power lines, of natural mires remain.

Larval melanism in a geometrid moth: promoted neither by a thermal nor seasonal adaptation but desiccating environments

Spatiotemporal variation in the degree of melanism is often considered in the context of thermal adaptation, melanism being advantageous under suboptimal thermal conditions. Yet, other mutually nonexclusive explanations exist. Analysis of geographical patterns combined with laboratory experiments on the mechanisms of morph induction helps to unveil the adaptive value of particular cases of polyphenism. In the context of the thermal melanism hypothesis and seasonal adaptations, we explored an array of environmental factors that may affect the expression and performance of nonmelanic vs. melanic larval morphs in different latitudinal populations of the facultatively bivoltine moth Chiasmia clathrata (Lepidoptera: Geometridae). Geographical variation in larval coloration was independent of average temperatures experienced by the populations in the wild. The melanic morph was, however, more abundant in dry than in mesic habitats. In the laboratory, the melanic morph was induced especially under a high level of incident radiation but also at relatively high temperatures, but independently of photoperiod. Melanic larvae had higher growth rates and shorter development times than the nonmelanic ones when both temperature and the level of incident radiation were high. Our results that melanism is induced and advantageous in warm desiccating conditions contradict the thermal melanism hypothesis for this species. Neither has melanism evolved to compensate time constraints due to forthcoming autumn. Instead, larvae solve seasonal variation in the time available for growth by an elevated growth rate and a shortened larval period in the face of autumnal photoperiods. The phenotypic response to the level of incident radiation and a lack of adaptive adjustment of larval growth trajectories in univoltine populations underpin the role of deterministic environmental variation in the evolution of irreversible adaptive plasticity and seasonal polyphenism.

Evolution of alternative insect life histories in stochastic seasonal environments

Abstract Deterministic seasonality can explain the evolution of alternative life history phenotypes (i.e., life history polyphenism) expressed in different generations emerging within the same year. However, the influence of stochastic variation on the expression of such life history polyphenisms in seasonal environments is insufficiently understood. Here, we use insects as a model and explore (1) the effects of stochastic variation in seasonality and (2) the life cycle on the degree of life history differentiation among the alternative developmental pathways of direct development and diapause (overwintering), and (3) the evolution of phenology. With numerical simulation, we determine the values of development (growth) time, growth rate, body size, reproductive effort, adult life span, and fecundity in both the overwintering and directly developing generations that maximize geometric mean fitness. The results suggest that natural selection favors the expression of alternative life histories in the alternative developmental pathways even when there is stochastic variation in seasonality, but that trait differentiation is affected by the developmental stage that overwinters. Increasing environmental unpredictability induced a switch to a bet‐hedging type of life history strategy, which is consistent with general life history theory. Bet‐hedging appeared in our study system as reduced expression of the direct development phenotype, with associated changes in life history phenotypes, because the fitness value of direct development is highly variable in uncertain environments. Our main result is that seasonality itself is a key factor promoting the evolution of seasonally polyphenic life histories but that environmental stochasticity may modulate the expression of life history phenotypes.