Thermal physiological ecology of Colias butterflies in flight

Summary As a comparison to the many studies of larger flying insects, we carried out an initial study of heat balance and thermal dependence of flight of a small butterfly (Colias) in a wind tunnel and in the wild.Unlike many larger, or facultatively endothermic insects, Colias do not regulate heat loss by altering hemolymph circulation between thorax and abdomen as a function of body temperature. During flight, thermal excess of the abdomen above ambient temperature is weakly but consistently coupled to that of the thorax. Total heat loss is best expressed as the sum of heat loss from the head and thorex combined plus heat loss from the abdomen because the whole body is not isothermal. Convective cooling is a simple linear function of the square root of air speed from 0.2 to 2.0 m/s in the wind tunnel. Solar heat flux is the main source of heat gain in flight, just as it is the exclusive source for warmup at rest. The balance of heat gain from sunlight versus heat loss from convection and radiation does not appear to change by more than a few percent between the wings-closed basking posture and the variable opening of wings in flight, although several aspects require further study. Heat generation by action of the flight muscles is small (on the order of 100 m W/g tissue) compared to values reported for other strongly flying insects. Colias appears to have only very limited capacity to modulate flight performance. Wing beat frequency varies from 12–19 Hz depending on body mass, air speed, and thoracic temperature. At suboptimal flight temperatures, wing beat frequency increases significantly with thoracic temperature and body mass but is independent of air speed. Within the reported thermal optimum of 35–39°C, wing beat frequency is negatively dependent on air speed at values above 1.5 m/s, but independent of mass and body temperature. Flight preference of butterflies in the wind tunnel is for air speeds of 0.5–1.5 m/s, and no flight occurs at or above 2.5 m/s. Voluntary flight initiation in the wild occurs only at air speeds 1.4 m/s.In the field, Colias fly just above the vegetation at body temperatures of 1–2°C greater than when basking at the top of the vegetation. These measurements are consistent with our findings on low heat gain from muscular activity during flight. Basking temperatures of butterflies sheltered from the wind within the vegetation were 1–2°C greater than flight temperatures at vegetation height.     

Compensating for climate change–induced cue‐environment mismatches: evidence for contemporary evolution of a photoperiodic reaction norm in Colias butterflies

Anthropogenic climate change alters seasonal conditions without altering photoperiod and can thus create a cue‐environment mismatch for organisms that use photoperiod as a cue for seasonal plasticity. We investigated whether evolution of the photoperiodic reaction norm has compensated for this mismatch in Colias eurytheme. This butterfly’s wing melanization has a thermoregulatory function and changes seasonally. In 1971, Hoffmann quantified how larval photoperiod determines adult wing melanization. We recreated his experiment 47 years later using a contemporary population. Comparing our results to his, we found decreased melanization at short photoperiods but no change in melanization at long photoperiods, which is consistent with the greater increase in spring than summer temperatures recorded for this region. Our study shows that evolution can help correct cue‐environment mismatches but not in the same way under all conditions. Studies of contemporary evolution may miss important changes if they focus on only a limited range of conditions.     

Evolutionary genetics of dorsal wing colour in Colias butterflies

The evolution of butterfly wing colouration is strongly affected by its multiple functions and by the correlated evolution of wing colour elements. Both factors may prevent local adaptation to ecological conditions. We investigated one aspect of wing colouration, the degree of dorsal wing melanization, in the butterfly Colias philodice eriphyle across an elevational gradient and its correlation with another aspect of wing colouration, ventral wing melanization. Dorsal wing melanization increased with elevation and these differences persisted in a common environment. Full-sibling analysis revealed high heritability for males but only intermediate heritability for females. The correlation between ventral and dorsal melanization showed significant elevational and sex-specific differences. In males the two traits were highly correlated, whereas in females the strength of the correlation decreased with increasing elevation. We conclude that uncoupling of ventral and dorsal melanization has evolved in females but not in males and discuss possible mechanisms underlying uncoupling.     

Thermoregulation and the determinants of heat transfer in Colias butterflies

Summary As a means of exploring behavioral and morphological adaptations for thermoregulation in Colias butterflies, convective heat transfer coefficients of real and model butterflies were measured in a wind tunnel as a function of wind speed and body orientation (yaw angle). Results are reported in terms of a dimensionless heat transfer coefficient (Nusselt number, Nu) and a dimensionless wind speed (Reynolds number, Re), for a wind speed range typical of that experienced by basking Colias in the field. The resultant Nusselt-Reynolds (Nu-Re) plots thus indicate the rates of heat transfer by forced convection as a function of wind speed for particular model geometries.For Reynolds numbers throughout the measured range, Nusselt numbers for C. eurytheme butterflies are consistently lower than those for long cylinders, and are independent of yaw angle. There is significant variation among individual butterflies in heat transfer coefficients throughout the Re range. Model butterflies without artificial fur have Nu-Re relations similar to those for cylinders. Heat transfer in these models depends upon yaw angle, with higher heat transfer at intermediate yaw angles (30–60°); these yaw effects increase with increasing Reynolds number. Models with artificial fur, like real Colias, have Nusselt numbers which are consistently lower than those for models without fur at given Reynolds numbers throughout the Re range. Unlike real Colias, however, the models with fur do show yaw angle effects similar to those for models without fur.The independence of heat loss from yaw angle for real Colias is consistent with field observations indicating no behavioral orientation to wind direction. The presence of fur on the models reduces heat loss but does not affect yaw dependence. The large individual variation in heat transfer coefficients among butterflies is probably due to differences in fur characteristics rather than to differences in wing morphology.Finally, a physical model of a butterfly was constructed which accurately simulates the body temperatures of basking Colias in the field for a variety of radiation and wind velocity conditions. The success of the butterfly simulator in mimicking Colias thermal characteristics confirms our preliminary understanding of the physical bases for and heat transfer mechanisms underlying thermoregulatory adaptations in these butterflies.     

The role of chemotactile stimuli in the oviposition preferences of Colias butterflies

Summary The legume foodplants of Colias butterflies possess specific chemical components which stimulate oviposition in females of the genus. Different legumes provide different degrees of stimulation, as demonstrated by close correlation between field observations and laboratory experiments with a new behavioral assay. Several generalizations emerge from this study. 1) Chemical preferences for various foodplants are under genetic control in C. meadii, and are at least partly independent of an individual's previous exposure to different legumes. 2) In some cases, chemical cues alone do not allow females to discriminate between species. Long-range search cues may be used to enhance discrimination in such instances. 3) Lupinus, a legume which is not usually utilized by Colias, stimulates oviposition in the laboratory, indicating its chemical affinity with other Leguminosae. The behavioral assay described should prove useful for the future identification of specific oviposition stimulants and deterrents. It is stressed, however, that chemotactile cues are involved only in the final step of oviposition, and that understanding foodplant choice in nature will require in-depth investigation into the mechanics of individual search processes.     

Sources of intraspecific variation in the hostplant seeking behavior of Colias butterflies

Summary Egg-laying females of the legume-feeding butterfly Colias philodice eriphyle were observed within a high-altitude study site during 1977 and 1978. Average oviposition selectivity showed two patterns of seasonal variation. First, second-brood females laid eggs more frequently after alighting upon hostplants than did first-brood females in both years. Second, a consistent decline in post-alighting oviposition probability near the end of the second brood corresponded with a dramatic decrease in the water content of hostplant foliage near the end of the dry summer season. In addition to seasonal variation in oviposition behavior, individual females landed and oviposited upon widely varying sets of legume hostplant species. By sampling hostplant abundances along the flightpaths of observed females, we show that individuals varied in their tendency to land upon different hostplants. Females tended to specialize upon one or two species, at least in the short term, and the pattern of oligophagy for the population as a whole is partly generated by variation in the host-seeking behavior of individual females.     

Functional ecological implications of intraspecific differences in wing melanization in Colias butterflies

Variation in the degree of insect wing melanin affects thermoregulation, and is expected to be adapted to local environmental conditions, for example over an elevational gradient. The effects of melanization on flight activity and egg maturation rate were assessed in the closely related butterflies Colias philodice eriphyle and C. eurytheme using experimental manipulation of wing darkness and transplant experiments between high and low elevation sites. Experimental manipulation of wing darkness in C. p. eriphyle demonstrated that light males had reduced flight activity at high elevations, and darkened males had reduced flight activity at low elevations. In contrast, the transplant experiments revealed asymmetrical adaptation for male C. p. eriphyle . At high elevations darker, high-elevation males had higher flight activity than lighter, low-elevation males, but there was no difference between the two groups at low elevation. For females, melanization had no effect on flight activity. However, an increase in female C. eurytheme wing darkness led to a significantly higher egg maturation rate at cold ambient temperatures, which may increase female reproductive output under natural conditions. Therefore, dispersers moving down in elevation may be more successful than those moving up.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 79–87.     

Growth,developmental and stress responses of larvae of the clouded sulphur butterfly Colias eriphyle to repeated exposure to high,sub‐lethal temperatures

The optimal temperature at which an organism grows and develops is commonly correlated with latitude and elevation; however, the maximum temperature for physiological performance often is not. This makes performance at temperatures between the optimum and the maximum of particular interest. Temperature can influence long‐term performance (growth and development), as well as short‐term performance (heat shock protein) responses differentially. In the present study, two populations of the clouded sulphur butterfly Colias eriphyle Edwards that differ in elevation, thermal regime and optimal and maximum temperatures are studied to quantify their responses to repeated, sub‐lethal heat treatments early in development (second instar). Heat treatments accelerate development during the second to fourth instars in both populations initially, although this effect disappears by pupation. Heat treatment decreases pupal mass in the lower elevation population, suggesting that repeated exposure to high temperatures early in development may reduce final size and fecundity in this population. Heat shock protein gene (hsp70) expression levels in the lower elevation (1633 m a.s.l.) population are highest 24 h after the start of the heat treatment and then the fall to pre‐exposure levels by 36–72 h, suggesting a rapid response to stressful temperatures. By contrast, heat treatment has no significant effect on pupal mass in the higher elevation (2347 m a.s.l.) population. This population has higher levels of hsp70 expression overall but constant expression levels, suggesting that the temperature treatments used are insufficient to elicit a heat stress response. Overall, the effects of repeated exposure to sub‐lethal high temperatures early in development on growth, final size and gene expression differ between populations that differ in thermal sensitivity.     

Climate variability slows evolutionary responses of Colias butterflies to recent climate change

How does recent climate warming and climate variability alter fitness, phenotypic selection and evolution in natural populations? We combine biophysical, demographic and evolutionary models with recent climate data to address this question for the subalpine and alpine butterfly, Colias meadii, in the southern Rocky Mountains. We focus on predicting patterns of selection and evolution for a key thermoregulatory trait, melanin (solar absorptivity) on the posterior ventral hindwings, which affects patterns of body temperature, flight activity, adult and egg survival, and reproductive success in Colias. Both mean annual summer temperatures and thermal variability within summers have increased during the past 60 years at subalpine and alpine sites. At the subalpine site, predicted directional selection on wing absorptivity has shifted from generally positive (favouring increased wing melanin) to generally negative during the past 60 years, but there is substantial variation among years in the predicted magnitude and direction of selection and the optimal absorptivity. The predicted magnitude of directional selection at the alpine site declined during the past 60 years and varies substantially among years, but selection has generally been positive at this site. Predicted evolutionary responses to mean climate warming at the subalpine site since 1980 is small, because of the variability in selection and asymmetry of the fitness function. At both sites, the predicted effects of adaptive evolution on mean population fitness are much smaller than the fluctuations in mean fitness due to climate variability among years. Our analyses suggest that variation in climate within and among years may strongly limit evolutionary responses of ectotherms to mean climate warming in these habitats.     

Drivers of local densities of endangered Lycaena helle butterflies in a fragmented landscape

Due to their specialised habitat requirements, butterflies are particularly vulnerable to habitat loss and fragmentation. Understanding the drivers of local abundances of species is essential for their effective conservation in fragmented landscapes. We investigated factors affecting population densities of an endangered European butterfly, the Violet Copper (Lycaena helle), occurring in a small metapopulation near the city of Kraków, southern Poland. The environmental parameters tested as predictors of the local densities of the species included both the variables associated with spatial structure of habitats such as patch sizes, their isolation and fragmentation as well as those potentially reflecting habitat quality. Patch area and vegetation height turned out to be the only factors significantly influencing L. helle densities, both having a positive effect. The positive impact of patch area is a bit surprising, since its relationship with population densities is typically negative in butterflies. In our study system it is likely to derive from source-sink dynamics as the smaller habitat patches are apparently too small to sustain viable local populations. In turn, the positive influence of vegetation height implies that the ongoing succession does not deteriorate the quality of the recently abandoned meadows yet, whereas higher turf may provide better sheltering places. The loss of almost half of L. helle habitat patches in the study area in recent years is alarming. However, its inclusion into the Natura 2000 system should help to conserve the species as long as this act is followed by proper management of its habitats.     

Population structure of pierid butterflies IV. Genetic and physiological investment in offspring by male Colias

Summary Population structure encompasses all the rules by which a population's gametes come together, including genetic and physiological investment in offspring. We document female use of nutrients donated by males at mating, and complete sperm precedence, in Colias eurytheme Boisduval. The effect of these phenomena on the population structure of this species is discussed.     

Natural hybridization in heliconiine butterflies: the species boundary as a continuum

Background  

To understand speciation and the maintenance of taxa as separate entities, we need information about natural hybridization and gene flow among species.     

Parallel evolution in sympatric, hybridizing species: performance of Colias butterflies on their introduced host plants

Shared ancestry and introgression can contribute to genetic similarity between hybridizing species, and it is generally difficult to disentangle these causes. However, shared ancestry plays a more limited role in traits that have recently undergone parallel directional selection in the two species, permitting the role of introgression to be better understood. The butterflies Colias eurytheme (Boisduval) and Colias philodice (Godart) (Lepidoptera, Pieridae) are native to North America and have shifted their host ranges in parallel onto several introduced weedy and agricultural legumes. These butterflies hybridize at moderate rates throughout their range, and there is a strong possibility that they could be sharing host‐associated adaptations. We split families of each species among nine introduced, prospective hosts and measured survivorship, larval duration, pupal weight, and a new variable, effective daily growth rate (DGR), analogous to a compound daily interest rate in economics. We found strong effects of host, sex, and family (species), but negligible effects of the host*species interaction that would indicate species‐specific differences in performance on different hosts. We found species‐specific life‐history differences: C. eurytheme matured significantly later and reached a significantly larger body size than C. philodice while growing at the same DGR. Protandry was strong, and males, in addition to pupating sooner than females, grew significantly faster than females as measured by DGR. We measured broad‐sense heritabilities and genetic correlations for host‐associated performance variables. Most pairwise comparisons of performance among hosts and most pairwise comparisons between performance variables showed positive genetic correlations, except survivorship where little heritability was found. Nevertheless, a factorial multivariate analysis of variance of G‐matrices showed highly significant species, host, and host*species interactions, suggesting differentially evolving genetic architectures underlying host adaptation in these two species, despite the small differences in overall performance. At least some of the genes affecting host performance in Colias are likely to be in the small, species‐diagnostic regions and not shared via introgression between these hybridizing species. For biologists interested in the evolutionary ecology of their host associations, including applied biologists managing their agricultural pest potential, C. eurytheme and C. philodice are most usefully studied as if they were a single polymorphic species wherever they co‐exist. In studying species that hybridize readily with a sympatric congener, it may often be necessary to include the second species in the experimental design.     

The evolution of wing color in Colias butterflies: heritability,sex linkage,and population divergence

We investigated the genetic background of intraspecific variation in wing color across an elevational gradient in the butterfly Colias philodice eriphyle. The degree of wing melanization was an accelerating function of elevation, and differences in wing melanization persisted in a common environment. Full-sibling analysis and parent-offspring regression yielded consistent, moderate to high heritabilities for the degree of wing melanization. The breeding experiments also demonstrated that wing melanization is strongly sex linked. Because traits that differentiate sister species also tend to be sex linked, our results suggest that the genetic mechanisms underlying intraspecific differences in wing melanization are not fundamentally different from those that have been shown to differentiate sister species.     

The evolution of wing color: male mate choice opposes adaptive wing color divergence in Colias butterflies

Abstract Correlated evolution of mate signals and mate preference may be constrained if selection pressures acting on mate preference differ from those acting on mate signals. In particular, opposing selection pressures may act on mate preference and signals when traits have sexual as well as nonsexual functions. In the butterfly Colias philodice eriphyle , divergent selection on wing color across an elevational gradient in response to the thermal environment has led to increasing wing melanization at higher elevations. Wing color is also a long-range signal used by males in mate searching. We conducted experiments to test whether sexual selection on wing melanization via male mate choice acts in the same direction as natural selection on mate signals due to the thermal environment. We performed controlled mate choice experiments in the field over an elevational range of 1500 meters using decoy butterflies with different melanization levels. Also, we obtained a more direct estimate of the relation between wing color and sexual selection by measuring mating success in wild-caught females. Both our experiments showed that wing melanization is an important determinant of female mating success in C. p. eriphyle . However, a lack of elevational variation in male mate preference prevents coevolution of mate signals and mate preference, as males at all elevations prefer less-melanized females. We suggest that this apparently maladaptive mate choice may be maintained by differences in detectability between the morphs or by preservation of species recognition.     

Environmental control of fine root dynamics in a northern hardwood forest

Understanding how exogenous and endogenous factors control the distribution, production and mortality of fine roots is fundamental to assessing the implications of global change, yet our knowledge of control over fine root dynamics remains rudimentary. To improve understanding of these processes, the present study developed regression relationships between environmental variables and fine root dynamics within a northern hardwood forest in New Hampshire, USA, which was experimentally manipulated with a snow removal treatment. Fine roots (< 1 mm diameter) were observed using minirhizotrons for 2 years in sugar maple and yellow birch stands and analyzed in relation to temperature, water and nutrient availability. Fine root dynamics at this site fluctuated seasonally, with growth and mortality peaking during warmer months. Monthly fine root production was strongly associated with mean monthly air temperature and neither soil moisture nor nutrient availability added additional predictive power to this relationship. This relationship exhibited a seasonal temperature hysteresis, which was altered by snow removal treatment. These results suggest that both exogenous and endogenous cues may be important in controlling fine root growth in this system. Proportional fine root mortality was directly associated with mean monthly soil temperature, and proportional fine root mortality during the over‐winter interval was strongly related to whether the soil froze. The strong relationship between fine root production and air temperature reported herein contrasts with findings from some hardwood forest sites and indicates that controls on fine root dynamics vary geographically. Future research must more clearly distinguish between endogenous and exogenous control over fine root dynamics in various ecosystems.     

The role of local ecology during hybridization at the initial stages of ecological speciation in a marine snail

Hybrid zones of ecologically divergent populations are ideal systems to study the interaction between natural selection and gene flow during the initial stages of speciation. Here, we perform an amplified fragment length polymorphism (AFLP) genome scan in parallel hybrid zones between divergent ecotypes of the marine snail Littorina saxatilis, which is considered a model case for the study of ecological speciation. Ridged‐Banded (RB) and Smooth‐Unbanded (SU) ecotypes are adapted to different shore levels and microhabitats, although they present a sympatric distribution at the mid‐shore where they meet and mate (partially assortatively). We used shell morphology, outlier and nonoutlier AFLP loci from RB, SU and hybrid specimens captured in sympatry to determine the level of phenotypic and genetic introgression. We found different levels of introgression at parallel hybrid zones and nonoutlier loci showed more gene flow with greater phenotypic introgression. These results were independent from the phylogeography of the studied populations, but not from the local ecological conditions. Genetic variation at outlier loci was highly correlated with phenotypic variation. In addition, we used the relationship between genetic and phenotypic variation to estimate the heritability of morphological traits and to identify potential Quantitative Trait Loci to be confirmed in future crosses. These results suggest that ecology (exogenous selection) plays an important role in this hybrid zone. Thus, ecologically based divergent natural selection is responsible, simultaneously, for both ecotype divergence and hybridization. On the other hand, genetic introgression occurs only at neutral loci (nonoutliers). In the future, genome‐wide studies and controlled crosses would give more valuable information about this process of speciation in the face of gene flow.     

Are legume‐feeding herbivores buffered against direct effects of elevated carbon dioxide on host plants? A test with the sulfur butterfly,Colias philodice

When grown under elevated atmospheric carbon dioxide (CO₂), leaf nitrogen content decreases less for legumes than for nonlegume C₃ plants. Given that elevated CO₂ adversely affects insect herbivores primarily through dilution of plant nitrogen, it is reasonable to expect that legume-feeding herbivores will be relatively buffered against CO₂-induced reduction in performance. However, despite their ecological and economic importance, very few studies have addressed the effects of elevated CO₂ on legume-feeding herbivores. Unlike the responses of the vast majority of nonlegume C₃ plants, when the legumes Trifolium pratense and Melilotus alba were grown under elevated (742 ppm) CO₂, leaf nitrogen and carbon contents and C : N ratios did not change. For Colias philodice larvae fed T. pratense , elevated CO₂ had little or no effect on consumption, digestion, or conversion of whole food or nitrogen and, consequently, no effect on growth rate, instar duration, or pupal weight. For larvae fed M. alba , elevated CO₂ had little or no effect on consumption of whole food or nitrogen, increased digestion but decreased conversion of both and, consequently, had no effect on growth rate, instar duration or pupal weight. These results suggest that, relative to herbivores of nonlegume C₃ plants, legume-feeding herbivores will be less affected as atmospheric CO₂ continues to rise.     

植物激素对体细胞胚胎发生的诱导与调节

以作者自己的工作为背景,结合国内外近几年的有关报道,综述了几种外源和内源激素对植物体细胞胚胎发生的诱导与调节作用。外源生长素和细胞分裂素是诱导离体培养细胞分化与增殖所必需的,2,4-D是诱导胚性愈伤组织的重要激素。在体细胞胚胎发生中内源激素含量和代谢的平衡起着关键的作用,而且外源和内源激素对诱导体细胞胚胎发生起相互调节作用。ABA在提高体细胞胚胎发生频率和质量上具有重要作用,同时,外源与内源ABA对体细胞胚胎发生起相互促进作用。本文还较为深入地讨论了这些激素诱导体细胞胚胎发生的可能作用机制。 Abstract:The paper summarizes the induced and regulatory effects of a few exogenous and endogenous hormones in plant somatic embryogenesis by our studies and related international reports.The exogenous auxin and cytokinin are necessary to induced differentiation and proliferation of cells of culture in vitro.2,4-D is an important hormone of induced embryogenic calluses.The contents and the metabolic balances of endogenous hormones have key effects for somatic embryogenesis.In addition,the exogenous and endogenous hormones have mutual regulatory effects for somatic embryogenesis.ABA has an important effect to improving the frequency and quality of somatic embryogenesis.Meanwhile,the exogenous and endogenous ABA have mutual promoted effects for somatic embryogenesis.The paper discusses possible mechanism of hormones-induced somatic embryogenesis in a deep-going way.