Green‐brown polymorphism in alpine grasshoppers affects body temperature

1. Ectothermic animals depend on external heat sources for pursuing their daily activities. However, reaching sufficiently high temperature can be limiting at high altitudes, where nights are cold and seasons short. We focus on the role of a green‐brown color polymorphism in grasshoppers from alpine habitats. The green‐brown polymorphism is phylogenetically and spatially widespread among Orthopterans and the eco‐evolutionary processes that contribute to its maintenance have not yet been identified.
2. We here test whether green and brown individuals heat up to different temperatures under field conditions. If they do, this would suggest that thermoregulatory capacity might contribute to the maintenance of the green‐brown polymorphism.
3. We recorded thorax temperatures of individuals sampled and measured under field conditions. Overall, thorax temperatures ranged 1.7–42.1°C. Heat up during morning hours was particularly rapid, and temperatures stabilized between 31 and 36°C during the warm parts of the day. Female body temperatures were significantly higher than body temperatures of males by an average of 2.4°C. We also found that brown morphs were warmer by 1.5°C on average, a pattern that was particularly supported in the polymorphic club‐legged grasshopper Gomphocerus sibiricus and the meadow grasshopper Pseudochorthippus parallelus.
4. The difference in body temperature between morphs might lead to fitness differences that can contribute to the maintenance of the color polymorphism in combination with other components, such as crypsis, that functionally trade‐off with the ability to heat up. The data may be of more general relevance to the maintenance of a high prevalence polymorphism in Orthopteran insects.

     

Density‐dependent polyphenism and geographic variation in size among two populations of lubber grasshoppers (Romalea microptera)

1. Density‐dependent phase polyphenism occurs when changes in density during the juvenile stages result in a developmental shift from one phenotype to another. Density‐dependent phase polyphenism is common among locusts (Orthoptera: Acrididae). 2. Previously, we demonstrated a longitudinal geographic cline in adult body size (western populations = small adults; eastern populations = large adults) in the eastern lubber grasshopper (Romalea microptera) in south Florida. As lubbers are confamilial with locusts, we hypothesised that the longitudinal size cline was partly due to density‐dependent phase polyphenism. 3. We tested the effect of density, population, and density×population interaction on life‐history traits (pronotum length, mass, cumulative development time, growth rate) of, and proportion surviving to, each of the five instars and the adult stage in a 2 × 3 factorial laboratory experiment with two lubber populations, each reared from hatchling to adult at three different densities. 4. The effect of density on life history and survival was independent of the effects of population on life history and survival. Higher densities led to larger adult sizes (pronotum, mass) and lower survivorship. The western population had smaller adult masses, fewer cumulative days to the adult stage, and higher survivorship than the eastern population. 5. Our data suggest that lubber grasshoppers exhibit density‐dependent phase polyphenism initiated by the physical presence of conspecifics. However, the plastic response of adult size to density observed in the laboratory is not consistent with the relationship between phenotypes and adult density in the field. Genetic differences between populations observed in the laboratory could contribute to size and life‐history differences among lubber populations in the field.     

Dynamics of two Montana grasshopper populations: relationships among weather,food abundance and intraspecific competition

The population dynamics of two grasshoppers (Melanoplus femurrubrum and M. sanguinipes) were studied using experimental microcosms over 8 years at a Palouse prairie site in Montana. Grasshopper density, survival and reproduction in the experimental populations responded in a density-dependent fashion to natural and experimental changes in food availability for all grasshopper developmental stages, both within and between all years. We observed that field populations of the grasshoppers at the site exhibited density, survival and reproductive responses similar to the experimental populations over the period of the study. Because we could not identify any differences between the field and microcosm environments or the grasshopper individuals in them, we contend that field populations were ultimately limited by food within and between years. Density-dependent food limitation occurred for all age categories over the entire summer, because food abundance declined relative to grasshopper food requirements over the summer. Food limitation occurred between years, because in years with the lowest food abundance, the populations produced more hatchlings for the next year than could be supported by the highest observed food abundance. Finally, the observed annual changes in food abundance were correlated with the annual variation in weather (rainfall and temperature), which indicated that the long established relationship between grasshopper densities and weather conditions does not imply population limitation by density-independent processes.     

Feeding preferences of Melanoplus femurrubrum grasshoppers on native and exotic grasses: behavioral and molecular approaches

Generalist insect herbivores, such as grasshoppers, may either avoid feeding on exotic plants, potentially enabling these plants to become invasive in the introduced range, or insects may incorporate exotic plants into their diet, contributing to the biotic resistance of native communities and potentially preventing plant invasions. Accurate determination of insect diet preferences with regard to native and exotic plants can be challenging, but this information is critical for understanding the interaction between native herbivores and exotic plants, and ultimately the mechanisms underlying plant invasions. To address this, we combined behavioral and molecular approaches to accurately compare food consumption of the polyphagous red‐legged grasshopper, Melanoplus femurrubrum (De Geer) (Orthoptera: Acrididae), on native [Andropogon gerardii Vitman and Bouteloua curtipendula (Michx.) Torr.] and exotic, potentially invasive grasses [Miscanthus sinensis Andersson and Bothriochloa ischaemum (L.) Keng] (all Poaceae). We found that M. femurrubrum grasshoppers demonstrated strong feeding preferences toward exotic grasses in experiments with intact plants under both field and greenhouse conditions, but they showed no preference in experiments with clipped leaves. Additionally, we sampled the gut contents of M. femurrubrum collected in the field and identified the ingested plant species based on DNA sequences for the non‐coding region of the chloroplast trnL (UAA) gene. We found that exotic plants were prevalent in the gut contents of grasshoppers collected at study sites in Ohio and Maryland, USA. These results suggest that the generalist herbivore M. femurrubrum does not avoid feeding on exotic grasses with which they do not share coevolutionary history. In addition, by demonstrating greater food consumption of exotic plants, these grasshoppers potentially provide biotic resistance should these grasses escape cultivation and become invasive in the introduced range.     

Evidence that reducing mammalian predators is beneficial for threatened and declining New Zealand grasshoppers

We evaluate evidence that reducing mammalian predators benefits threatened and declining grasshoppers in the Mackenzie Basin, New Zealand. Long-term population trends of Sigaus minutus are investigated under three control regimes: high intensity, indirect control through prey reduction and no control. We then test whether predator control benefits conservation management of Brachaspis robustus by translocating wild-caught individuals to areas of moderate versus no predator control. A significant positive trend in S. minutus counts occurred under high intensity and indirect control, suggesting that mammalian predator control is beneficial. Differences in the decline of translocated B. robustus were observed between moderate and no predator control release sites but could not be unequivocally attributed to predator densities. We recommend replicated predator control studies be undertaken to develop a predator management strategy which will enable grasshopper recovery, and investigate the potential for meso-predator release and prey-switching under regimes that target specific mammals.     

A molecular biogeographic analysis of the relationship between North American melanoploid grasshoppers and their Eurasian and South American relatives

The Melanoplinae constitute one of the two largest subfamilies of Acrididae. Distributed mainly throughout the New World and parts of Eurasia, this group of grasshoppers includes over 100 genera and 800 species. Over the past five decades there has been considerable speculation on the origins of North and South American taxa. The most favored hypothesis proposes an ancient division of Laurasian taxa accompanying the separation of North America and Eurasia, with subsequent radiations within those continents, followed by a recent incursion of Nearctic melanoplines into the southern hemisphere with the establishment of the Isthmus of Panama. This research tests that scenario by phylogenetic analysis using as characters portions of five mitochondrial gene sequences, totaling 2285 bp. Three tree-building methods, maximum-parsimony, neighbor-joining, and maximum-likelihood, strongly support the different view that melanopline grasshoppers originated somewhere in the Americas and spread to the Old World. The feasibility of these findings is discussed within a geological context.     

Condition‐dependence and sexual ornamentation: Effects of immune challenges on a highly sexually dimorphic grasshopper

Sexual ornaments contribute substantially to phenotypic diversity and it is particularly relevant to understand their evolution. Ornaments can assume the function of signals‐of‐quality that the choosy sex uses to evaluate potential mating partners. Often there are no obvious direct benefits and investment into mate choice is primarily rewarded by beneficial alleles that are inherited to the offspring. Inter‐sexual communication via sexual ornaments requires honesty of the sexual signal, yet the question of what maintains honesty remains only partially solved. One solution is that honesty is maintained by trait expression being dependent on individual condition, since condition‐dependent trait expression offers an effectively inexhaustible source of genetic variability. Here we test in the highly sexually dimorphic club‐legged grasshopper Gomphocerus sibiricus if putative sexual ornaments, in particular the striking front‐leg clubs, are more strongly affected by a lipopolysaccharide (LPS) immune challenge than putatively not sexually selected traits. Our results show overall little condition‐dependent expression of morphological and song traits, with sexually selected traits exhibiting effects comparable to nonsexually selected traits (with the possible exception of stridulatory file length and syllable‐to‐pause ratio in advertisement songs). Interestingly, field observations of individuals of lethally parasitized individuals suggest that a very strong environmental challenge can specifically affect the expression of the front‐leg clubs. The presence of 1% of males in natural populations with missing or heavily deformed clubs plus 5% with minor club deformations furthermore indicate that there are risks associated with club development during final ecdysis and this might act as a filter against deleterious alleles.     

Seeking salt: herbivorous prairie insects can be co‐limited by macronutrients and sodium

The canonical factors typically thought to determine herbivore community structure often explain only a small fraction of the variation in herbivore abundance and diversity. We tested how macronutrients and relatively understudied micronutrients interacted to influence the structure of insect herbivore (orthopteran) communities. We conducted a factorial fertilisation experiment manipulating macronutrients (N and P, added together) and micronutrients (Ca, Na and K) in large plots (30 × 30 m²) in a Texas coastal prairie. Although no single or combination of micronutrients affected herbivore communities in the absence of additional macronutrients, macronutrients and sodium added together increased herbivore abundance by 60%, richness by 15% and diversity by 20%. These results represent the first large‐scale manipulation of single micronutrients and macronutrients in concert, and revealed an herbivore community co‐limited by macronutrients and Na. Our work supports an emerging paradigm that Na may be important in limiting herbivore communities.     

Dietary selection and nutritional regulation in a common mixed‐feeding insect herbivore

The geometric framework provides a way for understanding the multi‐dimensional nutritional relationships between consumers and their food. We use this approach to further our understanding of the feeding and nutritional ecology of a ubiquitous mixed‐feeding insect herbivore that consumes a variety of host plants spanning a wide range of nutritional composition. Our overall objective was to examine feeding decisions, resulting performance, and post‐ingestive consequences in a common mixed‐feeding insect herbivore, Melanoplus bivittatus (Say) (Orthoptera: Acrididae), when presented with paired diets differing in protein:carbohydrate (p:c) ratio. Intake p:c of M. bivittatus differed among all but two treatments and in many cases was farther than expected from the previously identified p:c intake target for this species. Despite this variability in intake of protein and carbohydrate, we found few effects of the diet treatments on performance or post‐ingestive processing. However, our results suggest that when feeding on high‐quality diets, nutrients consumed in excess may be stored rather than excreted.     

Coevolution of color pattern and thermoregulatory behavior in polymorphic pygmy grasshoppers Tetrix undulata

Ectothermic organisms, such as insects and reptiles, rely on external heat sources to control body temperature and possess physiological and behavioral traits that are temperature dependent. It has therefore been hypothesised that differences in body temperature resulting from phenotypic properties, such as color pattern, may translate into selection against thermally inferior phenotypes. We tested for costs and benefits of pale versus dark coloration by comparing the behaviors (i.e., basking duration and bouts) of pygmy grasshopper (Tetrix undulata) individuals exposed to experimental situations imposing a trade-off between temperature regulation and feeding. We used pairs consisting of two full-siblings of the same sex that represented different (genetically coded) color morphs but had shared identical conditions from the time of fertilization. Our results revealed significant differences in behavioral thermoregulation between dark and pale individuals in females, but not in males. Pale females spent more time feeding than dark females, regardless of whether feeding was associated with a risk of either hypothermia or overheating. In contrast, only minor differences in behavior (if any) were evident between individuals that belonged to the same color morph but had been painted black or gray to increase and decrease their heating rates. This suggests that the behavioral differences between individuals belonging to different color morphs are genetically determined, rather than simply reflecting a response to different heating rates. To test for effects of acclimation on behaviors, we used pairs of individuals that had been reared from hatchlings to adults under controlled conditions in either low or high temperature. The thermal regime experienced during rearing had little effect on behaviors during the experiments reported above, but significantly influenced the body temperatures selected in a laboratory thermal gradient. In females (but not in males) preferred body temperature also varied among individuals born to mothers belonging to different color morphs, suggesting that a genetic correlation exists between color pattern and temperature preferences. Collectively, these findings, at least in females, are consistent with the hypothesis of multiple-trait coevolution and suggest that the different color morphs represent alternative evolutionary strategies.     

Effects of temperature on nutrient self‐selection in the silverfish Lepisma saccharina

Nutrient self‐selection represents an important behaviour that has been measured across many taxa. Despite the amount of research on this phenomenon, few studies report the evaluation of the effects of environmental variables such as temperature on nutrient selection by animals. In the present study, the nutrient selections of the silverfish Lepisma saccharina L. are measured across a range of temperatures (10, 15, 20, 25, 30 and 35 °C) using feeding arenas with three nutrient choices: carbohydrate (sucrose), protein (casein) and fat (lard). An overall preference for carbohydrates is shown across the range of temperatures, followed by protein, and then fat. However, the proportional consumption of each dietary component changes with temperature; the proportional carbohydrate consumption decreases dramatically with increasing temperature (>94% of the diet at 15 °C but <58% at 30 °C), whereas the proportional protein and lipid consumption increases with increasing temperature up to 30 °C. Changes in nutrient selection with temperature may be related to the dietary requirements of the insect at different temperatures.     

Tests of pleistocene speciation in montane grasshoppers (genus Melanoplus) from the sky islands of western North America

There has a been a resurgence of debate on whether the Pleistocene glaciations inhibited speciation. This study tests a model of Pleistocene speciation, estimating the phylogenetic relationships and divergence times of 10 species of montane grasshoppers, genus Melanoplus, using 1300 bp of the mitochondrial gene cytochrome oxidase I (COI). Based on average pairwise distances (corrected for multiple substitutions using Kimura's two-parameter model), all species appear to have originated within the Pleistocene. Sequence divergences between species are less than 4%, corresponding to divergence times less than 1.7 million years ago. Branching patterns among the species suggest that speciation was associated with more than one glacial-interglacial cycle. A likelihood-ratio test rejected a model of simultaneous species origins, the predicted branching pattern if species arose from the fragmentation of a widespread ancestor. These grasshoppers live in an area that was previously glaciated and, as inhabitants of the northern Rocky Mountain sky islands, underwent latitudinal and probably altitudinal shifts in distribution in response to climatic fluctuations. Given the repeated distributional shifts and range overlap of the taxa, there most likely has been ample opportunity for population mixing. However, despite periodic glacial cycles, with more than 10 major glaciations over the past million years and climatic fluctuations over as short a time scale as 10(3) to 10(4) years, the dynamic history of the Pleistocene did not preclude speciation. Although relationships among some taxa remain unresolved, these grasshopper species, even with their recent origins, exhibit genetic coherence and monophyletic or paraphyletic gene trees. The frequency of glacial cycles suggests that the speciation process must have been extremely rapid. These species of grasshoppers are morphologically very similar, differing primarily in the shape of the male genitalia. These characters are posited to be under sexual selection, may play an important role in reproductive isolation, and are known to diverge rapidly. This suggests the rapidity of evolution of reproductive isolation may determine whether species divergences occurred during the Pleistocene glaciations.     

Varied tastes: home range implications of foraging‐patch selection

Despite evidence of home range behaviour across many taxa, the mechanisms underlying the development of home ranges are still unknown. Recently, models have been developed to explore these mechanisms for both territorial and non‐ territorial species. One such model for a generic forager suggests animal memory and optimal foraging theory as underlying mechanisms driving forager movement and the development of stable home ranges. Although this is a promising model for ungulate home range development, assumptions of the model have yet to be evaluated. Using GPS relocation data from two populations of elk, we explored how foraging patch selection might influence the structure and development of home ranges in elk Cervus elaphus. During the summer growing season, we identified and sampled foraging patches used by elk. Points along elk paths not used for foraging were sampled identically for comparison. We contrasted ‘patch’ and ‘nonpatch’ data points, to identify foraging selection differences across herd, sex and season using a combination of directly sampled and remotely sensed covariates. In general, elk selected patches with higher biomass, cover, slope and lower traffic on the nearest road. These patch‐selection results speak directly to differences between foraging areas and other areas used by elk and demonstrate that both physiographic and anthropocentric features influence foraging patch selection. Our results offer insight as to what defines a valuable foraging patch for elk and how these patches might influence the development and structure of home ranges in a free‐ranging ungulate.     

Escape behaviour in blue‐winged grasshoppers,Oedipoda caerulescens

Blue‐winged grasshoppers Oedipoda caerulescens (Linnaeus, 1758) are commonly found in flat, open, unprotected areas. In the event of immediate danger, they leave their camouflaged position and jump away at the last moment. The present study conducted in a flight arena shows that, despite jumping at short notice from a crouching position, the grasshoppers achieve the correct timing for an optimal leap. If both compound eyes are blinded and the animals are stimulated by touch to execute an unprepared jump, the take‐off of the flightless nymphs is delayed, and adults are delayed in raising their wings; the animals tumble backward during the leap (in the case of adults, if they do not open their wings). This is a result of the unprepared take‐off position; because the entire length of the hind legs cannot be used for acceleration, the body is rotated backward. However, the escape path is not ultimately affected because, in the air, physical processes compensate for the unfavourable starting conditions. In addition, no disadvantage is evident upon landing. In each case, a hook landing was completed safely (i.e. the grasshopper landed and swung round to face the direction it had come from). The impact force is reduced and the grasshopper stabilizes itself by rotating from a forward to a backward position, immediately after the first contact with the ground. The hook landing also serves to confuse the potential attacker, and the disappearance of the bright blue hind wings of the adult makes it difficult for predators to shift quickly enough to a different kind of search to relocate their prey. In conclusion, the present study shows that the escape behaviour of blue‐winged grasshoppers is adapted to extremely short escape distances.     

Allozyme variation in sympatric populations of British grasshoppers,—evidence of natural selection

Seven species of British grasshopper were analysed for variation at 15 enzyme loci. Where possible, sympatric populations were collected so that direct comparisons could be made. Allozyme frequencies in Chorthippus brunneus and C. albomarginatus were compared using correlation analysis. A positive correlation was noted for a phosphoglucomutase allozyme and a negative correlation for an isocitrate dehydrogenase allozyme. It was suggested that natural selection may affect allozyme frequencies in the wild.
Using cluster analysis and principal component analysis, evolutionary relationships of grasshoppers have been examined and compared with interpretations based on morphology.     

Individual‐ and population‐level personalities in a floriphilic katydid

Behaviour, including personality, informs us about the response of animals towards their changing environment. Despite the widespread occurrence of florivorous insects and the important but often underrated ecological roles that they play, the study of florivore behaviour is neglected relative to that of pollinators and other herbivores. Specifically, we do not know how different personality types can develop among florivores and enable them to persist in habitats with an ephemeral and dynamic availability of food resources. To address this knowledge gap, we investigated the following questions: whether the (a) inter‐individual differences of exploration and boldness are consistent; (b) inter‐population differences of exploration and boldness are consistent; (c) exploration and boldness are correlated. We collected individuals of the polyphagous floriphilic katydid, Phaneroptera brevis from four populations from wasteland sites in Singapore and performed a personality assay conducted in an insectary to investigate the exploratory and boldness levels of the individuals and populations. The major novel finding was that the floriphilic P. brevis katydids exhibit population‐level personality types for boldness, but not for exploration. Some katydid individuals were consistently more exploratory and bolder than other individuals. However, contrary to our predictions, we did not find any evidence of behavioural syndromes in the katydid individuals, as the boldness level for individuals was not significantly correlated with exploration for individuals. This suggests that an individual which is more exploratory may not be equally keen to take risks and consume novel food that it encounters. Our findings also suggest that boldness and exploration are linked to ecologically important behaviours, but more studies are needed to better understand population‐level personality and how and why natural selection may favour the evolution of personality in certain populations.     

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.     

Characterizing phase‐related differences in behaviour of Schistocerca gregaria with spatial distribution analysis

The behaviour of locusts has been studied extensively using two approaches: (1) analysing a single individual's response to a group stimulus or (2) using group conditions to look at aggregation patterns. The second approach has, in contrast with the first one, not been improved in terms of statistical analyses since the 1960s. In the present study, we propose a spatial statistics approach of point‐pattern analysis to improve the group‐based assessment of behavioural phase characterization. This diagnostic tool was developed and tested in the laboratory with comparative analysis of solitarious (isolation‐reared) and gregarious (crowd‐reared) desert locusts, Schistocerca gregaria (Forskål) (Orthoptera: Acrididae). The spatial distribution patterns of 10 either solitarious or gregarious third‐instar hoppers were characterized with nearest neighbour distance measurements in a circular arena. The temporal sequence of spatial disposition of locusts was recorded with a digital camera taking photographs at regular intervals. The approach of point‐pattern analysis focused on the spatial distribution of observed events and allowed us to make inferences about the underlying process that generated them. The results confirmed that our diagnostic tool could identify that crowd‐reared hoppers tended to aggregate more to conspecifics than isolation‐reared ones. We could also verify that isolation‐reared hoppers were less active than crowd‐reared ones, but this was only true at the beginning of the experiments. The spatial statistics approach proposed in the present study could help with observations of phase‐related differences in the behaviour of locusts.