Consumption rates and the evolution of diet-induced plasticity in the head morphology of Melanoplus femurrubrum (Orthoptera: Acrididae)

Summary Phenotypic plasticity may be an ecologically important evolutionary response to natural selection in multiple environments. I have determined the effect of diet-induced developmental plasticity in the head size of grasshoppers (Melanoplus femurrubrum) onfeeding performance on two types of plants. Full-sib families were divided and raised on either red clover, Trifolium repens, or rye grass, Lolium perenne. In three different stages of ontogeny, grasshoppers raised on rye grass had significantly larger heads, relative to body size, than full-sibs raised on clover. A principal components analysis indicated that two to five relative head size characters covaried as a block in their plastic response to the feeding environment. Regressions of adjusted consumption rates (mg/sec) against relative head size revealed that larger head sizes, induced by the rye grass diet, enhanced consumption rates of rye grass, but not clover. Unexpectedly, a similar positive association was observed between head size and consumption rate for grasshoppers raised on clover when they were feeding on clover. These results support the inference that grasshoppers exhibit adaptive phenotypic plasticity. However, the unexpected influence of head size on consumption rates of clover indicates that the functional relationship between head morphology and feeding performance is complex and that variation in this relationship among plant environments is not sufficient to explain the evolution of diet-induced phenotypic plasticity.     

Visual behaviors of the migratory grasshopper,Melanoplus sanguinipes F. (Orthoptera: Acrididae)

The visual behavior of adult Melanoplus sanguinipesF. (Orthoptera: Acrididae) was investigated by placing individuals in the center of an arena and recording their orientation responses to visual targets at the perimeter of the arena. Targets that reflected more 540- to 570-nm light were approached more frequently; however, when reflectance in the 540 -to 570-nm region was combined with reflectance in the 400- to 520-nm region, orientation responses were reduced significantly. This suggests that spectral discrimination in M. sanguinipesinvolves at least two classes of photoreceptors, which respond to different regions of the wavelength spectrum. In addition, grasshoppers oriented to vertical, but not horizontal, contrasting stripes. However, when vertical stripes were added to targets reflecting 520- to 650-nm light, responses to verticals on these targets were not enhanced relative to verticals presented against a target background of 400- to 520-nm + 520- to 650-nm light. Thus, spectral discrimination and vertical stripe fixation appear to be two distinct visual behaviors, controlled by outputs from two classes of photoreceptors and a single class of photoreceptors, respectively, and may be used in different physiological or ecological contexts.     

The evolution of the phenotypic covariance matrix: evidence for selection and drift in Melanoplus

Phenotypic variation in trait means is a common observation for geographically separated populations. Such variation is typically retained under common garden conditions, indicating that there has been evolutionary change in the populations, as a result of selection and/or drift. Much less frequently studied is variation in the phenotypic covariance matrix (hereafter, P matrix), although this is an important component of evolutionary change. In this paper, we examine variation in the phenotypic means and P matrices in two species of grasshopper, Melanoplus sanguinipes and M. devastator. Using the P matrices estimated for 14 populations of M. sanguinipes and three populations of M. devastator we find that (1) significant differences between the sexes can be attributed to scaling effects; (2) there is no significant difference between the two species; (3) there are highly significant differences among populations that cannot be accounted for by scaling effects; (4) these differences are a consequence of statistically significant patterns of covariation with geographic and environmental factors, phenotypic variances and covariances increasing with increased temperature but decreasing with increased latitude and altitude. This covariation suggests that selection has been important in the evolution of the P matrix in these populations Finally, we find a significant positive correlation between the average difference between matrices and the genetic distance between the populations, indicating that drift has caused some of the variation in the P matrices.     

The influence of temperature on the duration of egg development in West European grasshoppers (Orthoptera: Acrididae)

Summary The relationship between embryonic postdiapause development (PDD) and temperature in ten Acridid species was studied by culturing eggpods from diapause onwards at constant temperatures (ranging from 15.0° C to 37.5° C). In six species PDD was also studied at fluctuating temperatures (12 h at 27.5° C and 12 h at 13.0° C). The PDD duration was strongly temperature dependent in all species studied (Q₁₀ values between 3.0 and 4.1) and differed greatly between species. To assess whether these differences affect habitat determination, PDD duration was related to habitat microclimate. We found that species with a relatively long PDD are restricted to relatively warm grasslands, while species with a short PDD are also found in colder grasslands. Two species have a more limited occurrence than predicted from PDD duration. It is argued that the length of PDD (and consequently hatching date) is crucial to the completion of the annual cycle. The results are applicable to conservation of grasshoppers in grasslands subject to eutrophication. Here increase of vegetation density and height will lead to lower maximum temperatures in the egg environment (i.e. near the soil surface) and consequently to a delay in hatching and loss of thermophilous species. Grassland management rules are proposed that should aid conservation of these species.     

Genetic and environmental components of growth in nestling blue tits (Parus caeruleus)

We investigated the effect of brood‐size mediated food availability on the genetic and environmental components of nestling growth in the blue tit (Parus caeruleus), using a cross‐fostering technique. We found genetic variation for body size at most nestling ages, and for duration of mass increase, but not of tarsus growth. Hence, nestling growth in our study population seems to have the potential to evolve further. Furthermore, significant genotype–environment interactions indicated heritable variation in reaction norms of growth rates and growth periods, i.e. that our study population had a heritable plasticity in the growth response to environmental conditions. The decreasing phenotypic variance with nestling age indicated compensatory growth in all body traits. Furthermore, the period of weight increase was longer for nestlings growing up in enlarged broods, while there was no difference to reduced broods in the period of tarsus growth. At fledging, birds in enlarged broods had shorter tarsi and lower weights than birds in reduced broods, but there was no difference in wing length or body condition between the two experimental groups. The observed flexibility in nestling growth suggests that growing nestlings are able to respond adaptively to food constraint by protecting the growth of ecologically important traits.     

Selection in a fluctuating environment leads to decreased genetic variation and facilitates the evolution of phenotypic plasticity

Changes in the environment are expected to induce changes in the quantitative genetic variation, which influences the ability of a population to adapt to environmental change. Furthermore, environmental changes are not constant in time, but fluctuate. Here, we investigate the effect of rapid, continuous and/or fluctuating temperature changes in the seed beetle Callosobruchus maculatus, using an evolution experiment followed by a split-brood experiment. In line with expectations, individuals responded in a plastic way and had an overall higher potential to respond to selection after a rapid change in the environment. After selection in an environment with increasing temperature, plasticity remained unchanged (or decreased) and environmental variation decreased, especially when fluctuations were added; these results were unexpected. As expected, the genetic variation decreased after fluctuating selection. Our results suggest that fluctuations in the environment have major impact on the response of a population to environmental change; in a highly variable environment with low predictability, a plastic response might not be beneficial and the response is genetically and environmentally canalized resulting in a low potential to respond to selection and low environmental sensitivity. Interestingly, we found greater variation for phenotypic plasticity after selection, suggesting that the potential for plasticity to evolve is facilitated after exposure to environmental fluctuations. Our study highlights that environmental fluctuations should be considered when investigating the response of a population to environmental change.     

Quantitative trait loci mapping of phenotypic plasticity and genotype-environment interactions in plant and insect performance

Community genetic studies generally ignore the plasticity of the functional traits through which the effect is passed from individuals to the associated community. However, the ability of organisms to be phenotypically plastic allows them to rapidly adapt to changing environments and plasticity is commonly observed across all taxa. Owing to the fitness benefits of phenotypic plasticity, evolutionary biologists are interested in its genetic basis, which could explain how phenotypic plasticity is involved in the evolution of species interactions. Two current ideas exist: (i) phenotypic plasticity is caused by environmentally sensitive loci associated with a phenotype; (ii) phenotypic plasticity is caused by regulatory genes that simply influence the plasticity of a phenotype. Here, we designed a quantitative trait loci (QTL) mapping experiment to locate QTL on the barley genome associated with barley performance when the environment varies in the presence of aphids, and the composition of the rhizosphere. We simultaneously mapped aphid performance across variable rhizosphere environments. We mapped main effects, QTL × environment interaction (QTL×E), and phenotypic plasticity (measured as the difference in mean trait values) for barley and aphid performance onto the barley genome using an interval mapping procedure. We found that QTL associated with phenotypic plasticity were co-located with main effect QTL and QTL×E. We also located phenotypic plasticity QTL that were located separately from main effect QTL. These results support both of the current ideas of how phenotypic plasticity is genetically based and provide an initial insight into the functional genetic basis of how phenotypically plastic traits may still be important sources of community genetic effects.     

Male and female genitalia in some Libyan species of Acrididae (Orthoptera: Acridoidea)

A comparative study of male and female genitalia was carried out in thirty‐seven Libyan species representing twenty genera of the family Acrididae. An attempt has been made to describe and illustrate the different structures, namely, epiphallus, aedeagus, subgenital plate, supra‐anal plate and cerci of the male, and spermatheca, ovipositor, subgenital plate, supra‐anal plate and cerci of the female, in Acridids, with an aim to discover their significance in order to make the identification of genera and species, together with other generic characters, more perfect and convenient. Distinct family characters are shield or bridge‐shaped condition of epiphallus; presence or absence of dorso‐lateral appendices, oval sclerites and lophi on epiphallus; divided, undivided or flexured condition of aedeagus; presence or absence of gonopore process on aedeagus; condition of apical and pre‐apical diverticula of spermatheca; presence or absence of glandular pouches of Cornstock and Kellog on female subgenital plate; and rudimentary or well developed condition of egg‐guide. Stable characters for separating the subfamilies are taken to be presence or absence of ancorae on epiphallus, long or short condition of aedeagal sclerites; elongate, slender or short and broad condition of ovipositor valves: presence or absence of Jannone's organs and setae on posterior margin of female subgenital plate; and shape of diverticula of spermatheca. Useful generic characters are shape of male subgenital plate, supra‐anal plate and cerci, broad or narrow condition of bridge, presence or absence of branch of bridge connecting lophi with bridge of epiphallus; mono‐, bi‐ or tri‐lobate condition of lophi of epiphallus, length and upcurved or downcurved condition of apical valve of aedeagus, shape of posterior margin of female subgenital plate, presence of setae on the whole posterior margin or confined to lateral margins only; and toothed, tuberculate or smooth condition of ovipositor valves, length of the lateral apodeme in relation to the dorsal valves. Specific characters are shape of egg‐guide of female subgenital plate, shape of ovipositor valves and apical tips, shape of male supra‐anal plate and cerci, size of anterior and posterior lobes of lophi of epiphallus, size and shape of ancorae, shape of apical valves of aedeagus; and size of apical and pre‐apical diverticula and presence of protuberance on pre‐apical diverticulum.     

Response of grasshoppers (Orthoptera: Acrididae) to livestock grazing in southeastern Arizona: differences between seasons and subfamilies

Summary Grasshopper densities were compared between grazed and ungrazed semidesert grassland sites in southeastern Arizona. Bouteloua-dominated perennial grass cover was about 1.5 times greater on the livestock exclosure. Grasshoppers were 3.7 times more abundant on the protected area in the summers of 1983 and 1984, when dominant species were grass-feeding members of the subfamily Gomphocerinae. In fall 1984, grasshoppers were 3.8 times more common on the grazed site, when dominants were mainly herb-feeders in the subfamily Melanoplinae. These results indicate important seasonal and taxonomic differences in the responses of grasshoppers to the activities of vertebrate grazers.     

Genetics of behavioural and morphological differences between parapatric subspecies of Chorthippus parallelus (Orthoptera: Acrididae)

The grasshoppers Chorthippus parallelus parallelus and C. p. erythropus form a narrow hybrid zone in the Pyrenees. They differ in several characters of the pattern and structure of male stridulation and in the morphology of the stridulatory file. These characters are considered to be involved in the species' mate recognition system.
Crosses have been made between a Pyrenean C. p. erythropus population and two C. p. parallelus populations, one in the Pyrenees and one in the Massif Central. Reciprocal Fls and backcrosses have been examined for a set of stridulation and associated morphological characters. The crosses confirm that the subspecific differences have a genetic basis and suggest that they are polygenically determined. However the mode of inheritance is not simple. There is evidence for dominance and epistatic effects and for sex-linkage or maternal effects. Genetic correlations exist between some pairs of characters in the backcrosses.
These results are discussed in the context of the hybrid zone and in relation to the general problem of the evolutionary divergence of mate recognition systems.     

Perceptual,developmental, experiential,and physiological parameters of swimming in Melanopline grasshoppers (Orthoptera: Acrididae)

The roles of perception, development, experience, and physiology on the swimming behavior of grasshoppers in an arena were investigated usingMelanoplus bivittatus, M. differentialis, andM. sanguinipes. The parameters of swimming included capacity (the ability to reach the edge of a 45-cm-diameter water basin within 3 min), net displacement time (time required to reach the edge of the arena), and orientation (ability to make initial contact with a target pattern).Melanoplus sanguinipes adults were most attracted to a vertical rectangle set at 90° to the water surface (the standard target). Orientation ability decreased significantly as the target became square or it declined to 60°, and there was virtually no orientation to horizontal patterns or those at 45°. All nymphal instars ofM. sanguinipes exhibited the capacity to swim and orient to a standard target. Although the net displacement time was lowest from the third instar through the adult stages, the earlier instars (first through fourth) had the greatest relative net displacement rates. As adults aged, their net displacement time and orientation ability declined significantly, particularly in males. BothM. differentialis andM. sanguinipes adults showed a decline in net displacement time over a 10-day trial period. The loss of a hind leg did not significantly alter swimming performance of eitherM. bivittatus orM. sanguinipes adults, but this alteration significantly reduced the net displacement time ofM. differentialis. DecerebratedM. differentialis adults were capable of swimming for several minutes.     

Genotype x environment interaction for male attractiveness in an acoustic moth: evidence for plasticity and canalization

The lek paradox arises when choosy females deplete the genetic variance for male display traits from a population, yet substantial additive genetic variation (V_A) in male traits persists. Thus, the lek paradox can be more generally stated as one of the most fundamental evolutionary questions: What maintains genetic variation in natural populations? One solution to this problem may be found in the condition‐dependent nature of many sexually selected traits. Genotype × environment (G × E) interactions can maintain V_A under conditions of environmental heterogeneity provided certain restrictions are met, although antagonistic pleiotropy has also been proposed as a mechanism. Here, we provide evidence for G × E interactions and against the role of antagonistic pleiotropy in the maintenance of V_A for sexually selected traits. Using inbred lines of the lesser waxmoth Achroia grisella, we measured V_A for song attractiveness, condition and development rate under different competitive environments and found that genotypes differed in their plasticity. We argue that variation persists in natural populations because G × E interactions prevent any one variant from producing the optimal phenotype across all environments.     

Stage-Specific Behavioral Responses of Ageneotettix deorum (Orthoptera: Acrididae) in the Presence of Lycosid Spider Predators

Grasshoppers must gather food while avoiding size-selective predation from other arthropods, especially spiders, potentially leading to a trade-off between foraging and defensive behaviors. This trade-off becomes less intense as prey grow larger and are less susceptible to arthropod predation. Activity budgets were constructed for three nymphal (third- to fifth- instar) and adult life cycle stages of Ageneotettix deorum, a common rangeland grasshopper, for three conditions of predation risk by lycosid spiders (spider absence, spider presence, and presence of a nonlethal, chelicerae-modified spider). In third and fourth instars, exposure to predators resulted in reduced feeding activity, increased time spent in antipredator and defensive behaviors, and reduced general activity compared to individuals not exposed to spiders. No significant shifts in behaviors were observed for fifth-instar nymphs and adult A. deorum in response to spider presence. Activity levels in functional spiders and chelicerae-modified spiders were statistically indistinguishable.     

Genetic variation and phenotypic plasticity in a trophically polymorphic population of pumpkinseed sunfish (Lepomis gibbosus)

Summary Adaptive variation can exist at a variety of scales in biological systems, including among species, among local populations of a single species and among individuals within a single population. Trophic or resource polymorphisms in fishes are a good example of the lowest level of this hierarchy. In lakes without bluegill sunfish (Lepomis macrochirus), pumpkinseed sunfish (Lepomis gibbosus) can be trophically polymorphic, including a planktivorous limnetic form found in the pelagic habitat, in addition to the usual benthic form found in the littoral zone. In this paper we examine the degree to which morphological differences between the two forms are caused by genetic differences versus phenotypic plasticity. Adults from pelagic and littoral sites in Paradox Lake, NY, were bred separately and their progeny were raised in cages both in the open water and shallow water habitats of an artificial pond. The experimental design permitted two tests of genetic differences between the breeding stocks (in open and shallow water cages, respectively) and two tests of phenotypic plasticity (in the limnetic and benthic offspring, respectively). Limnetic progeny were more fusiform than benthic progeny raised in the same habitat. In addition, progeny of both stocks displayed limnetic-type characteristics when raised in the open water and benthic-type characteristics in the shallow water. Thus, genetic differences and phenotypic plasticity both contributed to the trophic polymorphism. Phenotypic plasticity and genetic differentiation accounted for 53 and 14%, respectively, of the variation in morphology. This study addresses the nature of subtle phenotypic differences among individuals from a single population that is embedded within a complex community, a condition that is likely to be the norm for most natural populations, as opposed to very large differences that have evolved in relatively few populations that reside in species-poor environments.     

Genetic and environmental components of phenotypic variation in body shape among populations of Atlantic cod (Gadus morhua L.)

A common-garden experiment was conducted on larvae to test for genetic differences in body shape among populations of Atlantic cod ( Gadus morhua ). Offspring from four north-west Atlantic regions were reared from hatching to postmetamorphosis at two temperatures (7 ± 1 °C and 11 ± 1 °C) and two food levels (1500 and 4500 prey L⁻¹). Body shape differed between populations and treatments. Population differences were greatest between south-west Scotian Shelf cod and those further north; the former were characterized by a deeper body, larger head, and longer caudal peduncle than cod from the other populations. Significant differences were also observed between two putative populations on the south-west Scotian Shelf, suggesting genetic divergence between spawning aggregations at small spatial scales (< 100 km). Temperature and food supply also influenced body shape, with the effect of the former being more pronounced. Individuals reared at the higher temperature or food level had a deeper body and a larger head than those reared at the lower temperature or food supply. Phenotypic responses to changes in the rearing environment also differed among populations, indicating genetic differences in phenotypic plasticity. Differences between populations in morphology and in phenotypic plasticity suggest genetic divergence at both large (> 1000 km) and small (< 100 km) spatial scales. The genetic differences at large spatial scales counteracted the expected effects of temperature differences in the wild, suggesting countergradient variation in morphology among these populations.  © 2006 Her Majesty the Queen in Right of Canada. Journal compilation © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 351–365.     

Assessing the sensitivity of Melanoplus frigidus (Orthoptera: Acrididae) to different weather conditions: A modeling approach focussing on development times

The temperature and soil moisture conditions as well as vegetation patterns were studied to describe the habitat and to model the life cycle of Melanoplus frigidus, a true alpine grasshopper of the Scandes. In the low alpine belt of the Norwegian Scandes the species colonizes only the warmest microhabitats with maximum soil surface temperatures of 31°C. Vegetation of these habitats consists of shrub‐rich heath dominated by Vaccinium myrtillus and Calluna vulgaris. Using continuously measured temperature data, the development times for four different seasons were modeled and related to field observations. The maximum delay of adult molt was estimated to amount to 3 weeks, the delay being determined by the variation in spring temperature conditions between different years. The possibilities of using M. frigidus as an indicator organism of climate change effects on alpine zoo‐coenoses of the Scandes are discussed.     

Transmission and perception of acoustic signals in the desert clicker,Ligurotettix coquilletti (Orthoptera: Acrididae)

Males of the desert clicker, Ligurotettix coquilletti (Acrididae: Orthoptera) defend a femalerequired resource, the creosote bush Larrea tridentata, in desert habitats of the southwestern United States. Males signal acoustically to each other as well as to searching females. The call is produced by tegminal/femoral stridulation where one or both legs are used for sound production. Sound pressure levels, measured laterally, are influenced by the intervening tegmen between the stridulating leg and the microphone. Differences in measured sound pressure levels between sides can vary up to 7 dB. When clicks are produced multiply,these multiple clicks may be 4 dB louder than single clicks. We examine the structure of the call and the effective broadcast area of single males by monitoring acoustic ascending neurons of the ventral nerve cord in the neck. By taking the neurophysiological preparation into the field, we were able to map the broadcast area of isolated males and also of males calling within aggregations. The distance over which the signal of isolated males could be detected was 8–14 m, whereas neural representation of the calls of males within aggregation were detectable within 4–6 m. The sound spectrum of the song, although having a major lower-frequency component around 10 kHz, has extensive power in the ultrasonic range. The tuning characteristics of the ascending auditory neuron matched the overall structure of the male call. The importance of the acoustic cue, as compared to visual cues, is discussed in relation to female attraction.     

Genetic and maternal influences on life history plasticity in milkweed bugs (Oncopeltus): response to temperature

This study was designed to examine life history flexibility arising from phenotypic plasticity in response to temperature and from maternal effects in response to reproductive diapause in a temperate zone population of the milkweek bug (Oncopeltus fasciatus). We employed a split-family, first-cousin, full-sib design with siblings reared at different temperatures in order to quantify phenotypic plasticity, maternal effects, and variation for each. The following traits were analyzed: development time, age at first reproduction, longevity, early-life fecundity, and wing length. We found both life history plasticity and maternal effects on life history traits which tend to enhance the colonizing ability of offspring born to mothers that have undergone reproductive diapause. We were unable to demonstrate additive genetic variation for plasticity for any of the traits, while for development time and wing length we found variation due to non-additive genetic or common-environmental sources. We were also unable to demonstrate additive genetic variation for maternal effects, although variation may exist at low levels that are difficult to detect using cousin-families. The apparent lack of variation in this population would constrain evolution of life history flexibility even though considerable flexibility exists in the phenotype.