Alternative life histories in the water strider Gerris lacustris: time constraint on wing morph and voltinism

The wing dimorphic water strider Gerris lacustris L. (Heteroptera: Gerridae) switches to a bivoltine life cycle under favorable climatic conditions. The switch in voltinism is accompanied by a reduction of wing development in the directly reproducing midsummer generation, while the diapausing generation has a high fraction of long‐winged individuals. We investigated whether the thermal energy (degree‐days) available in natural habitats constrains the combination of developmental pathway and wing morph. Offspring of G. lacustris were reared under quasi‐natural conditions at two temperature regimes to determine the thermal constant k required to complete adult development in either wing morph. The thermal constant for egg‐to‐adult development of the short‐winged morph was about 20% lower than of the long‐winged morph. Based on the results from the outdoor laboratory experiment, we calculated the total degree‐days necessary to complete the possible combinations of wing morph pattern and voltinism. Comparison of these estimates with the thermal energy actually available during the reproductive season of 2004 for various natural habitats (sun‐exposed field ponds and shaded forest ponds) suggests that voltinism as well as wing morph pattern is strongly limited by the number of degree‐days available in these habitats. On forest ponds, only univoltine life cycles were possible, whereas on field ponds temperature allowed bivoltine life cycles. However, only the eggs laid at the very beginning of the season had the potential to accumulate enough degree‐days to complete a bivoltine life cycle with both generations long‐winged. We conclude that thermal energy is the main environmental constraint limiting voltinism of populations in the two habitat types. Furthermore, the available thermal energy also seems to influence the determination of the seasonal wing pattern in G. lacustris.     

Variability in the life history of the water strider Gerris lacustris (Heteroptera: Gerridae) across small spatial scales

Abstract.  1. Variability in life-history traits of the common pond skater Gerris lacustris (L.) (Heteroptera: Gerridae) was investigated on a small geographical scale in Bavaria, Germany (≈ 49 °N). In an area of approximately 3.5 × 3.0 km, voltinism and wing length data of G. lacustris were recorded from 65 ponds located either in an open field or surrounded by forest.
2. Life history differed significantly between populations in the two habitat types. Gerris lacustris showed a bivoltine life-cycle in most of the field ponds with a high proportion of long-winged individuals in the diapause generation and a large increase of the short-winged morph in the midsummer generation. In contrast, almost all populations inhabiting forest ponds were univoltine. Only a few forest populations produced any second generation larvae. The long-winged morph dominated during the whole season in forest ponds.
3. The appearance of first-instar larvae was delayed by 3–4 weeks in forest ponds compared with field ponds. Presumably, low temperatures led to this reduction in the developmental rate of eggs and larvae. Consequently, 'field larvae' of the first generation developed under increasing daylight conditions (triggering direct reproduction and brachyptery) but 'forest larvae' developed under a shortening photoperiod (triggering diapause and macroptery).
4. Offspring raised from 'forest parents' in the laboratory showed the same variability in life history when reared in a cool (16.9 ± 2.4 °C) or warm (23.1 ± 1.8 °C) temperature regime.
5. Given the small spatial scale of the study, it is concluded that the variability in the life history of G. lacustris has a strong phenotypic component.     

Seasonal phenotypic plasticity of wing melanisation in the cabbage white butterfly, Pieris rapae L. (Lepidoptera: Pieridae)

Abstract.  1. Effective thermoregulation is crucial for the fitness of small flying insects. Phenotypic plasticity of the ventral hindwing of pierid butterflies is widely recognised as adaptive for effective thermoregulation. Butterflies eclosing in cooler environments have more heavily melanised wings that absorb solar radiation, thus allowing flight under these cool conditions.
2. Many pierids also exhibit phenotypic plasticity of dorsal forewing melanisation but in this case, cooler environments reduce melanisation. It has been hypothesised that this plasticity is also adaptive because it increases solar reflection from the wing surfaces onto the body in certain basking postures.
3. The degree of seasonal variation in ventral hindwing and dorsal forewing melanisation of wild-caught Pieris rapae was quantified to determine if it shows patterns of plasticity similar to that documented for other Pieris species.
4. Male wing melanisation on both wing surfaces shows the characteristic seasonal, adaptive plasticity. However, only some dorsal forewing pattern elements of females conformed to the predictions of the hypothesis of adaptive dorsal forewing melanisation. Sexual dimorphism of wing pattern plasticity may result from, and/or affect, sexual dimorphism of behaviour and physiology of these butterflies.     

Reduction and variability of trunk spines in the acanthocephalan Corynosoma cetaceum: the role of physical constraints on attachment

Abstract. In this study, we investigated a functional trade-off between trunk attachment and trunk-spine development in the acanthocephalan Corynosoma cetaceum . The worms live attached to the stomach and upper intestine of their cetacean definitive hosts, using the proboscis and spiny foretrunk as the main holdfast; the spiny hindtrunk can also attach by bending ventrally. When the hindtrunk bends, ventral compression generates an anterior fold (AF) and a posterior fold (PF). A morphological analysis based on 7,823 individuals collected from 10 franciscana dolphins, Pontoporia blainvillei , revealed that spines were smaller and more variable in size and occurrence in the folds than on neighboring areas; the growth of fold spines seemed to be inhibited to various degrees. Spines were more reduced in the AF than in the PF, and spines of both folds were more reduced in females than in males. Patterns of reduction appeared to be directly related to the intensity of fold compression associated with hindtrunk bending. Fold compression could induce plastic inhibition of spine growth, and/or could make fold spines maladaptive, spines being reduced by natural selection. Apparently, fold spines neither contact the substrate, nor are they exposed to the environment when the hindtrunk attaches. Therefore, fold spines could have reduced, or lost, their primary function, at least in the definitive host. The reduction and variability of spines in C. cetaceum seem to be unique among Corynosoma species.     

Effect of photoperiod on development and growth in a pentatomid bug, Dolycoris baccarum

The effect of photoperiod on nymphal development, growth and adult size was examined in a pentatomid bug, Dolycoris baccarum , collected in Osaka (a warm temperate region) and Hokkaido (a subfrigid region), Japan. When insects were reared from eggs at 25°C, the developmental period was long and adult size was large under photoperiods close to the critical photoperiod for the induction of adult diapause. Adults of the Hokkaido population were larger than those of the Osaka population. There was no significant correlation between developmental period and adult size. Insects also showed variation in their growth rate: growth rate was low under photoperiods a little longer than the critical photoperiod for the induction of diapause. The ecological significance of variation in development and growth is discussed.     

The effect of temperature and wing morphology on quantitative genetic variation in the cricket Gryllus firmus, with an appendix examining the statistical properties of the Jackknife-MANOVA method of matrix comparison

We investigated the effect of temperature and wing morphology on the quantitative genetic variances and covariances of five size-related traits in the sand cricket, Gryllus firmus. Micropterous and macropterous crickets were reared in the laboratory at 24, 28 and 32 degrees C. Quantitative genetic parameters were estimated using a nested full-sib family design, and (co)variance matrices were compared using the T method, Flury hierarchy and Jackknife-manova method. The results revealed that the mean phenotypic value of each trait varied significantly among temperatures and wing morphs, but temperature reaction norms were not similar across all traits. Micropterous individuals were always smaller than macropterous individuals while expressing more phenotypic variation, a finding discussed in terms of canalization and life-history trade-offs. We observed little variation between the matrices of among-family (co)variation corresponding to each combination of temperature and wing morphology, with only one matrix of six differing in structure from the others. The implications of this result are discussed with respect to the prediction of evolutionary trajectories.     

Effects of food availability on predator-induced morphological defence in the ciliate Euplotes octocarinatus (Protista)

The effect of a prey's food resources on predator-induced morphological transformation was studied in two ciliate species: Euplotes octocarinatus (prey) and Stylonychia mytilus (predator). The ability to produce defence in Euplotes was much reduced by prolonged starvation. The extent of prey morphological transformation and therefore the effectiveness of the defence was inversely related to the length of time without food. The results suggest that the defence involves an energetic cost which may be a significant part of the whole cell budget, at least when food is in short supply. Not only the morphological transformation but also the maintenance of the anti-predator phenotype were energetically costly for Euplotes, the cost being proportional to the defence level. The morphological transformation was affected by the kind of food (algae) provided to the prey. Ciliates fed Chlorogonium elongatum attained their maximum width sooner than those fed Chlamydomonas. Thus, both the quantity and the quality of the food available may significantly affect the magnitude of response and therefore the potential success of an induced defence.     

Seasonal butterfly design: morphological plasticity among three developmental pathways relative to sex,flight and thermoregulation

The temperate‐zone butterfly Pararge aegeria can use three developmental pathways corresponding to different seasonal cohorts: (1) development with a pupal winter diapause resulting in early spring adults; (2) development with a larval winter diapause resulting in late‐spring adults and (3) direct development resulting in summer or second generation adults. In order to test adaptive predictions, we compared variation in flight‐ and thermoregulation‐related morphology among adult males and females from the three pathways using both field data (i.e. wild‐caught butterflies) and experimental breeding data (i.e. reared under different photoperiod regimes). Morphological patterns among the pathways were largely similar in the field and rearing data. Seasonal patterns differed between the sexes for most traits, including (relative) size measures and wing colour. Our results suggest sex‐related, adaptive seasonal plasticity for morphological traits related to flight behaviour in a multivoltine insect.     

The change in phototactic behaviour of a Daphnia magna clone in the presence of fish kairomones: the effect of exposure time

Using a clone that responds to the presence of fishkairomones by a pronounced change in phototacticbehaviour, we determined how fast a change to morenegatively phototactic behaviour occurs in Daphnia magnaadults that are exposed to a highconcentration of fish kairomones. Kairomone exposedanimals showed an approximately linear decrease in thevalue of the phototactic index with time. Though theresponse was almost immediate, it took two hoursbefore the difference between fish-induced and controlanimals was significant. Extrapolation of the observedresponse indicates that a maximal change inphototactic behaviour, equivalent to animals that havebeen cultured in the presence of fish kairomones sincebirth, occurs after about 13 hours exposure. Weconclude that the predator-induced change in dielvertical migration of zooplankton is fast, and isfully developed in less than a day. The response timeto fish kairomones of Daphnia is shorter forphototactic behaviour than for life history traits,which may have important consequences with respect tothe evolution of trait-dependence in induced defenceresponses.     

Pleiotropic effects of flowering time genes in the annual crucifer Arabidopsis thaliana (Brassicaceae)

Variation in flowering time of Arabidopsis thaliana was studied in an experiment with mutant lines. The pleiotropic effects of flowering time genes on morphology and reproductive yield were assessed under three levels of nutrient supply. At all nutrient levels flowering time and number of rosette leaves at flowering varied among mutant lines. The relationship between these two traits depended strongly on nutrient supply. A lower nutrient supply first led to an extension of the vegetative phase, while the mean number of leaves at flowering was hardly affected. A further reduction resulted in no further extension of the vegetative phase and, on average, plants started flowering with a lower leaf number. At low nutrients, early flowering affected the timing of production of siliques rather than the total output, whereas late flowering was favorable at high nutrients. This may explain the fact that many plant species flower at a relatively small size under poor conditions. Flowering time genes had pleiotropic effects on the leaf length, number of rosette and cauline leaves, and number of axillary flowering shoots of the main inflorescence. Silique production was positively correlated with the number of axillary shoots of the main inflorescence; the number of axillary primordia appeared to have a large impact on reproductive yield.     

Antagonistic and reinforcing pleiotropy: a study of differences in development time in wing dimorphic insects

Morphological dimorphisms are found in many different taxa. Wing dimorphism in insects, in which some individuals possess wings and associated flight muscles and are thus volant while others lack a functional flight apparatus and are thus flightless, is a typical example of such types of dimorphisms. It has been extensively studied and such studies have demonstrated that the volant form, although possessing the advantage of flight capability, suffers a fitness cost in a delay in the onset of reproduction after emergence into the adult form and a reduced fecundity. Previous comparative analyses have suggested that there is no consistent trend for development time (hatching to adult) to differ between the two morphs. The present study analyses the phenotypic and genetic correlations between development time and wing morph in the cricket Gryllus firmus. It is shown that the macropterous (volant) morph develops faster than the micropterous (flightless morph). This trade-off is manifested at both thephenotypic and genetic level. Further, a comparative analysis shows that the same phenotypic trade-off is generally found in other Orthopteran species so far studied, but in other orders the micropterous morph develops faster. Provided that the phenotypic trade-off is genetically based, in the Orthoptera the fitness advantage of the earlier onset of reproduction in micropterous females is offset by the extended development time (antagonistic pleiotropy). However, in other orders there is reinforcing pleiotropy in that the micropterous females develop faster and reproduce sooner than the macropterous morph. These results highlight the complexity of fitness interactions and the need to study a phenomenon across several taxa.     

Late‐moulting Black‐necked Grebes Podiceps nigricollis show greater body mass in the face of failing food supply

From August to December, thousands of Black‐necked Grebes Podiceps nigricollis concentrate during the flightless moult period in salt ponds in the Odiel Marshes, southern Spain, where they feed on the brine shrimp Artemia parthenogenetica. We predicted that because Black‐necked Grebes moulted in a food‐rich, predator‐free environment, there would be no net loss of body mass caused by the use of fat stored to meet energy needs during remigial feather replacement (as is the case for some other diving waterbirds). However, because the food resource disappears in winter, we predicted that grebes moulting later in the season would put on more body mass prior to moult because of the increasing risk of an Artemia population crash before the moult period is completed. Body mass determinations of thousands of birds captured during 2000–2010 showed that grebes in active wing‐moult showed greater mass with date of capture. Early‐moulting grebes were significantly lighter at all stages than late‐moulting birds. Grebes captured with new feathers post‐moult were significantly lighter than those in moult. This is the first study to support the hypothesis that individual waterbirds adopt different strategies in body mass accumulation according to timing of moult: early‐season grebes were able to acquire an excess of energy over expenditure and accumulate fat stores while moulting. Delayed moulters acquired greater fat stores in advance of moult to contribute to energy expenditure for feather replacement and retained extra stores later, most likely as a bet hedge against the increasing probability of failing food supply and higher thermoregulatory demands late in the season. An alternative hypothesis, that mass change is affected by a trophically transmitted cestode using brine shrimps as an intermediate host and Black‐necked Grebes as final host, was not supported by the data.     

Sexual size dimorphism decreases with temperature in a blowfly,Chrysomya megacephala (Fabricius) (Diptera: Calliphoridae)

1. There is wide intra‐specific variation in sexual size dimorphism (SSD). Much of this variation is probably as a result of sexual differences in the selective pressure on body size. However, environmental variables could affect males and females differently, causing variation in SSD. 2. We examined the effects of two temperatures (20 and 30 °C) on SSD in six populations of the blowfly, Chrysomya megacephala. 3. We found that body size increased with temperature in all the populations studied, and the sexes differed in phenotypic plasticity of body size in response to rearing temperature. This created substantial temperature‐induced variation in SSD (i.e. sex × temperature interaction). Males were often smaller than females, but the degree of dimorphism was smaller at the higher temperature (30 °C) and larger at the lower temperature (20 °C). This change in SSD was not because of a gender difference in the effect of temperature on development time. Further studies should address whether this variation can be produced by adaptive canalisation of one sex against variation in temperature, or whether it may be a consequence of non‐adaptive developmental differences between the sexes. 4. Although most studies assume that the magnitude of SSD is fixed within a species, the present study demonstrates that rearing temperature can generate considerable intra‐specific variation in the degree of SSD.     

The effect of prey availability on offspring survival depends on maternal food resources

Maternal nutrition can strongly influence embryo development and offspring fitness. The environmental matching hypothesis posits that developmental conditions affect offspring in ways that enable them to appropriately deal with similar post‐developmental conditions, although mismatches between developmental and post‐developmental environments will reduce fitness. To test this hypothesis, reproductive lizards (Anolis sagrei) were reared in environments with high versus low prey availability. The resultant offspring were then reared reciprocally under the same two prey conditions that their mothers experienced. High levels of prey available to mothers increased egg production, although the survival of eggs was low compared to those produced by mothers in the low‐prey treatment. Low prey availability to offspring reduced growth, regardless of the amount of prey available to their mothers. Low prey availability also compromised offspring survival, although this negative effect was only present when mothers experienced high‐prey conditions, whereas matching of low‐prey conditions in maternal and offspring stages resulted in high survival. However, because the mismatch of low maternal and high offspring prey availability resulted in similar offspring survival to the matched treatments, our results do not fully support the environmental matching hypothesis. Nevertheless, the present study highlights the interactive role of maternal and post‐hatching environments in generating variation in offspring fitness. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 437–447.     

Brain plasticity over the metamorphic boundary: carry-over effect of larval environment on froglet brain development

Brain development shows high plasticity in response to environmental heterogeneity. However, it is unknown how environmental variation during development may affect brain architecture across life history switch points in species with complex life cycles. Previously, we showed that predation and competition affect brain development in common frog (Rana temporaria) tadpoles. Here, we studied whether larval environment had carry-over effects in brains of metamorphs. Tadpoles grown at high density had large optic tecta at metamorphosis, whereas tadpoles grown under predation risk had small diencephala. We found that larval density had a carry-over effect on froglet optic tectum size, whereas the effect of larval predation risk had vanished by metamorphosis. We discuss the possibility that the observed changes may be adaptive, reflecting the needs of an organism in given environmental and developmental contexts.     

Rainfall-induced changes in food availability modify the spring departure programme of a migratory bird

Climatic warming has intensified selection for earlier reproduction in many organisms, but potential constraints imposed by climate change outside the breeding period have received little attention. Migratory birds provide an ideal model for exploring such constraints because they face warming temperatures on temperate breeding grounds and declining rainfall on many tropical non-breeding areas. Here, we use longitudinal data on spring departure dates of American redstarts (Setophaga ruticilla) to show that annual variation in tropical rainfall and food resources are associated with marked change in the timing of spring departure of the same individuals among years. This finding challenges the idea that photoperiod alone regulates the onset of migration, providing evidence that intensifying drought in the tropical winter could hinder adaptive responses to climatic warming in the temperate zone.     

Variation in wing dimorphism of oriental mole cricket Gryllotalpa orientalis: Comparative study in Okinawa and Hyogo populations

The oriental mole cricket Gryllotalpa orientalis exhibits variation in wing dimorphism. In an Okinawa population, no short‐winged individuals were observed, and wing dimorphism has not been detected. Flight behavior of G. orientalis was observed from April to October in Okinawa. In contrast, a Hyogo population exhibited seasonal wing dimorphism and long‐winged individuals appear from June to September. The flight period of the long‐winged morph coincided with this period. Short‐winged individuals appeared from September to the following June and they never fly. Both populations showed univoltine life cycles. Considering the possible flight period, wing pattern and life cycle of mole crickets in these two areas, it is presumed that flightlessness is expected to arise when adults can not experience suitable temperatures for flight activity.     

Interclonal differences, plasticity and trade-offs of life history traits of Cyperus esculentus in relation to water availability

Cyperus esculentus is an exotic clonal (or pseudoannual) weed in Japan, and its range is steadily increasing. To investigate its interclonal variation and phenotypic plasticity in response to water availability, five clones of C. esculentus , collected from different sites in Japan, were grown singly in pots placed outdoors under dry and wet conditions. All the traits examined showed considerable variation among the five clones. However, two clones from Tochigi were similar to each other; thus, they might have originated from the same founder population. The clone from Ibaraki was quite different from the others. Therefore, it is suggested that the Japanese populations of C. esculentus might have resulted from multiple introductions of genotypes from geographically separated and, hence, genetically differentiated, source populations. All the clones also showed considerable plasticity in response to water availability. Clones with a larger ramet number had a greater plasticity, whereas tuber size was invariant across water treatments. Highly plastic traits had generally low interclonal variation in plasticity. All the clones had high productivity and produced more ramets and tubers under wet conditions than under dry conditions. Moreover, water availability could partially regulate the mode of its reproduction; wet conditions favored tuber production (vegetative propagation) while dry conditions favored sexual reproduction. A number of trade-offs occurred between the traits of clonal growth, storage and sexual reproduction, indicating that allocation among the competing functions/organs is mutually exclusive in plants. The results obtained here suggest that C. esculentus is more likely to invade wet habitats than dry habitats.