I examined the relative importance of colour pattern, body size and shape for thermoregulation by... more I examined the relative importance of colour pattern, body size and shape for thermoregulation by experimentally subjecting adult female Tetrix subulata belonging to four different colour morphs (black, striped, brown, and white) to augmented irradiation levels. The results revealed significant variation in temperature excess (i.e., the difference between ambient and body temperature) among colour morphs, with black individuals having a mean temperature excess 49% greater than that of white individuals. Temperature excess was not significantly influenced by overall body size (measured as principal component 1 (PC1)) but increased with increasing values of PC2, suggesting that stout bodied individuals attained higher body temperatures than did individuals with low body mass relative to surface area. These results show that coloration may play an important role in determining body temperatures of grasshoppers exposed to direct irradiation, and suggest that reproductive females may be able to maintain higher body temperatures than non-reproductive females and males.
Because body temperature influences the physiology, behaviour and performance of ectothermic orga... more Because body temperature influences the physiology, behaviour and performance of ectothermic organisms, thermal conditions might influence the frequency and outcome of predator-prey interactions. Furthermore, differences in colour pattern among prey individuals may indirectly influence predation risk, through effects of coloration on body temperature. We performed staged encounters between lizards (Psammodromus algirus) and grasshoppers (Tetrix subulata). We painted the dorsal surface of grasshoppers either black or grey to manipulate their body temperature and behaviour, and exposed them to predation under two different thermal conditions (sun and shade). Grasshoppers were less active in the presence of a lizard, suggesting that they modified their behaviour to avoid detection. Lizards were more active and attacked the grasshoppers more frequently in the sun than in the shade. However, the proportion of unsuccessful attacks did not differ between sun and shade, suggesting that thermal environment did not influence the relative performance of predators and prey. In the sun, black grasshoppers were more active and tended to be attacked more frequently than grey ones, albeit not significantly so. However, mortality did not differ between black and grey individuals. This suggests that an elevated activity need not translate into increased predation if accompanied by enhanced escape performance. Conversely, individuals may compensate for a poor escape performance associated with low body temperatures by reducing activity. Our findings suggest that selection imposed by predators may favour certain combinations of prey coloration and behaviour, and that colour polymorphism in ectotherms may be influenced also by indirect effects of coloration on predation risk. Counter to intuition, the existence of such indirect effects raises the possibility that the evolution of prey coloration may be governed by selection imposed also by colour-blind or even totally blind predators.
Random left minus right deviations from symmetry in otherwise bilaterally symmetric traits may ar... more Random left minus right deviations from symmetry in otherwise bilaterally symmetric traits may arise due to developmental instability in response to environmental stress. Here we test for variation in developmental instability, measured as asymmetry of (femur) size, among individuals belonging to four different genetically encoded colour morphs of the pygmy grasshopper Tetrix undulata (Sow.) (Orthoptera: Tetrigidae). Such a difference is expected under the hypotheses that perturbed growth and development and concomitant morphological asymmetry may result from exposure to unfavourable temperatures or costs associated with melanization, or reflect a by-product of past disease in individuals with poor immune responsiveness. Results from mixed model two-way ANOVAs uncovered no statistically significant directional asymmetry in femur size, whereas non-directional asymmetry was significant in each of the four different colour morphs. The degree of intraindividual, interlateral variance in femur size varied significantly among individuals belonging to different colour morphs, being considerably higher in the black morph, which also suffers the greatest risk of overheating, compared to the brown, striped and grey morphs. Asymmetry in femur size was not associated with one measure of immune responsiveness, the encapsulation response of individuals experimentally implanted with a novel antigen consisting of a nylon mono-filament. These results are consistent with the notion that individual pigmentation pattern may indirectly influence developmental instability and morphological asymmetry, via the effects of coloration on body temperature.
Ecological and evolutionary consequences of host-parasite interactions have attracted considerabl... more Ecological and evolutionary consequences of host-parasite interactions have attracted considerable attention from evolutionary biologists. Previous studies have suggested that immune responsiveness may be genetically or developmentally linked with colour pattern, such that the evolution of animal colour patterns may be influenced by correlated responses to selection for parasite resistance. We studied interactions between the endoparasitic fly Leiophora innoxia (Meigen) (Diptera: Tachinidae) and its colour polymorphic pygmy grasshopper host Tetrix undulata (Sow.) (Orthoptera: Tetrigidae) to test for morph-specific differences in parasitization and immune defence, and hostinduced variation in parasite phenotypes. Our results revealed that c. 2 and 30% of adult grasshoppers collected from the same natural population in two subsequent years, respectively were parasitized. Parasite prevalence was independent of host sex and colour morph. Pupae were larger if the parasite had developed in a female than in a male host, possibly reflecting host resource value or a physical constraint on larval growth imposed by host body size. Pupal size was also associated with host colour morph, with individuals that had developed in dark morphs being smaller at pupation compared to those that developed in paler morphs. However, immune defence, measured as the encapsulation response to a novel antigen, did not differ among individuals belonging to alternative colour morphs or sexes. Darker morphs warm up more quickly and prefer higher body temperatures than paler ones. Encapsulation was not influenced by maintenance temperature (15 vs. 30 ∞ C), however, suggesting that indirect effects of coloration on parasite resistance mediated via differential body temperature are unlikely. The dependence of parasite body size on host colour morph may thus reflect plasticity of growth and development of the larvae in response to differential host body temperature, rather than variable host immune defence.
Ectothermic organisms, such as insects and reptiles, rely on external heat sources to control bod... more 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.
Recent studies have interpreted intraspecific divergence in relative head sizes in snakes as evid... more Recent studies have interpreted intraspecific divergence in relative head sizes in snakes as evidence for adaptation of the trophic apparatus in gape-limited predators to local prey size. However, such variation might also arise from non-adaptive processes (such as allometry, correlated response, genetic drift, or non-adaptive phenotypic plasticity). We test predictions from these alternative hypotheses using data on the allometric relationship between head size and body size in two wide-ranging snake species: eight populations of adders (Vipera berus) and 30 populations of common gartersnakes (Thamnophis sirtalis). Our data enable strong rejection of the alternative (non-adaptive) hypotheses, because the relationship between head and body size differed significantly among populations, the geographic distance separating pairs of populations explained less than 1.5% of their divergence in allometric coefficients, and the within-population allometric coefficients were higher than the among-population coefficients in each species. In addition, the geographical variability of allometric coefficients in females did not parallel that in males, suggesting that allometric coefficients have evolved independently in the two sexes. Phenotypic plasticity also cannot explain the data, because laboratory studies show that the allometric relationship between head size and body size is relatively insensitive to differing growth rates. We conclude that the intraspecific head size divergence in these snakes is better explained by spatially heterogeneous selection to optimize prey handling ability, than by non-adaptive processes.
Females of Lampruphulis delicata are dimorphic for colour pattern, the difference between morphs ... more Females of Lampruphulis delicata are dimorphic for colour pattern, the difference between morphs being the presence or absence of a distinct white mid-lateral stripe. A less distinct striped morph occurs also in males. We evaluated alternative hypotheses for the maintenance of this polymorphism by examining temporal and spatial variation in morph frequency, testing for differential selection among morphs using data on body size and reproductive traits from preserved specimens, and experimentally manipulating colour pattern in freeranging lizards of both sexes, to assess the influence of the lateral stripe on survival rates. We found that the relative frequency of striped individuals varied among populations and decreased from north to south in both sexes, coincident with an increasing incidence of regenerated tails. Morph frequencies did not change through time within a population. Striped gravid females appeared to survive better and produced larger clutches than did non-striped females. In our experimental study, the relationship between survival and colour morph differed between the two sexes; males painted with a white lateral stripe had lower survival than control (brown stripe) males, but survival did not differ between striped and control females. The different response in the two sexes may be due partly to differences in temperature and microhabitat selection. We propose that the white lateral stripe decreases susceptibility to predators in gravid females but increases risk of predation in males, especially in combination with low temperatures. The polymorphism might be maintained by: ( I ) opposing fitness consequences of the stripe in males and females; ('2) sex-specific habitat selection; and (3) gene flow in combination with spatial variation in relative fitness of the two morphs.
Habitat selection behavior is affected by complex interplays between competing requirements. Here... more Habitat selection behavior is affected by complex interplays between competing requirements. Here we combine field observations with laboratory experiments to examine how thermal benefits and predator avoidance influences habitat selection by different color morphs of the pygmy grasshopper Tetrix undulata. The composition of substrate types and surface temperatures in areas selected by free-ranging individuals did not reflect relative availability, and varied among morphs and sexes. Surface temperatures of selected habitats deviated less from the range of preferred body temperatures than would result from a random utilization of surface temperatures, suggesting that grasshoppers selected habitats with thermal properties which were suitable for maintaining preferred body temperatures. The thermal property of habitats occupied by different color morphs suggests that darker morphs (which absorb more solar radiation) selected cooler habitats to avoid overheating. Dissimilarities in substrate use among color morphs in the field and laboratory emphasize a role also of predator avoidance by background matching for habitat choice. The degree of habitat selectivity was lowest in the striped morph, supporting the notion that a disruptive color pattern may constitute a solution to the trade-off between relative crypsis in different visual backgrounds. Finally, individuals modified their habitat use when subjected to elevated risk of predation, showing that habitat choice is governed by conflicting priorities. Collectively, our findings suggest that, as a result of direct and indirect effects of coloration on performance, alternative color morphs use different solutions to the trade-off between competing requirements. Our results also lend support to the notion that relative fitness of alternative color morphs and sexes may be dependent on microhabitat selection, as predicted by the theory of multiple niche polymorphisms.
I examined the relative importance of colour pattern, body size and shape for thermoregulation by... more I examined the relative importance of colour pattern, body size and shape for thermoregulation by experimentally subjecting adult female Tetrix subulata belonging to four different colour morphs (black, striped, brown, and white) to augmented irradiation levels. The results revealed significant variation in temperature excess (i.e., the difference between ambient and body temperature) among colour morphs, with black individuals having a mean temperature excess 49% greater than that of white individuals. Temperature excess was not significantly influenced by overall body size (measured as principal component 1 (PC1)) but increased with increasing values of PC2, suggesting that stout bodied individuals attained higher body temperatures than did individuals with low body mass relative to surface area. These results show that coloration may play an important role in determining body temperatures of grasshoppers exposed to direct irradiation, and suggest that reproductive females may be able to maintain higher body temperatures than non-reproductive females and males.
Because body temperature influences the physiology, behaviour and performance of ectothermic orga... more Because body temperature influences the physiology, behaviour and performance of ectothermic organisms, thermal conditions might influence the frequency and outcome of predator-prey interactions. Furthermore, differences in colour pattern among prey individuals may indirectly influence predation risk, through effects of coloration on body temperature. We performed staged encounters between lizards (Psammodromus algirus) and grasshoppers (Tetrix subulata). We painted the dorsal surface of grasshoppers either black or grey to manipulate their body temperature and behaviour, and exposed them to predation under two different thermal conditions (sun and shade). Grasshoppers were less active in the presence of a lizard, suggesting that they modified their behaviour to avoid detection. Lizards were more active and attacked the grasshoppers more frequently in the sun than in the shade. However, the proportion of unsuccessful attacks did not differ between sun and shade, suggesting that thermal environment did not influence the relative performance of predators and prey. In the sun, black grasshoppers were more active and tended to be attacked more frequently than grey ones, albeit not significantly so. However, mortality did not differ between black and grey individuals. This suggests that an elevated activity need not translate into increased predation if accompanied by enhanced escape performance. Conversely, individuals may compensate for a poor escape performance associated with low body temperatures by reducing activity. Our findings suggest that selection imposed by predators may favour certain combinations of prey coloration and behaviour, and that colour polymorphism in ectotherms may be influenced also by indirect effects of coloration on predation risk. Counter to intuition, the existence of such indirect effects raises the possibility that the evolution of prey coloration may be governed by selection imposed also by colour-blind or even totally blind predators.
Random left minus right deviations from symmetry in otherwise bilaterally symmetric traits may ar... more Random left minus right deviations from symmetry in otherwise bilaterally symmetric traits may arise due to developmental instability in response to environmental stress. Here we test for variation in developmental instability, measured as asymmetry of (femur) size, among individuals belonging to four different genetically encoded colour morphs of the pygmy grasshopper Tetrix undulata (Sow.) (Orthoptera: Tetrigidae). Such a difference is expected under the hypotheses that perturbed growth and development and concomitant morphological asymmetry may result from exposure to unfavourable temperatures or costs associated with melanization, or reflect a by-product of past disease in individuals with poor immune responsiveness. Results from mixed model two-way ANOVAs uncovered no statistically significant directional asymmetry in femur size, whereas non-directional asymmetry was significant in each of the four different colour morphs. The degree of intraindividual, interlateral variance in femur size varied significantly among individuals belonging to different colour morphs, being considerably higher in the black morph, which also suffers the greatest risk of overheating, compared to the brown, striped and grey morphs. Asymmetry in femur size was not associated with one measure of immune responsiveness, the encapsulation response of individuals experimentally implanted with a novel antigen consisting of a nylon mono-filament. These results are consistent with the notion that individual pigmentation pattern may indirectly influence developmental instability and morphological asymmetry, via the effects of coloration on body temperature.
Ecological and evolutionary consequences of host-parasite interactions have attracted considerabl... more Ecological and evolutionary consequences of host-parasite interactions have attracted considerable attention from evolutionary biologists. Previous studies have suggested that immune responsiveness may be genetically or developmentally linked with colour pattern, such that the evolution of animal colour patterns may be influenced by correlated responses to selection for parasite resistance. We studied interactions between the endoparasitic fly Leiophora innoxia (Meigen) (Diptera: Tachinidae) and its colour polymorphic pygmy grasshopper host Tetrix undulata (Sow.) (Orthoptera: Tetrigidae) to test for morph-specific differences in parasitization and immune defence, and hostinduced variation in parasite phenotypes. Our results revealed that c. 2 and 30% of adult grasshoppers collected from the same natural population in two subsequent years, respectively were parasitized. Parasite prevalence was independent of host sex and colour morph. Pupae were larger if the parasite had developed in a female than in a male host, possibly reflecting host resource value or a physical constraint on larval growth imposed by host body size. Pupal size was also associated with host colour morph, with individuals that had developed in dark morphs being smaller at pupation compared to those that developed in paler morphs. However, immune defence, measured as the encapsulation response to a novel antigen, did not differ among individuals belonging to alternative colour morphs or sexes. Darker morphs warm up more quickly and prefer higher body temperatures than paler ones. Encapsulation was not influenced by maintenance temperature (15 vs. 30 ∞ C), however, suggesting that indirect effects of coloration on parasite resistance mediated via differential body temperature are unlikely. The dependence of parasite body size on host colour morph may thus reflect plasticity of growth and development of the larvae in response to differential host body temperature, rather than variable host immune defence.
Ectothermic organisms, such as insects and reptiles, rely on external heat sources to control bod... more 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.
Recent studies have interpreted intraspecific divergence in relative head sizes in snakes as evid... more Recent studies have interpreted intraspecific divergence in relative head sizes in snakes as evidence for adaptation of the trophic apparatus in gape-limited predators to local prey size. However, such variation might also arise from non-adaptive processes (such as allometry, correlated response, genetic drift, or non-adaptive phenotypic plasticity). We test predictions from these alternative hypotheses using data on the allometric relationship between head size and body size in two wide-ranging snake species: eight populations of adders (Vipera berus) and 30 populations of common gartersnakes (Thamnophis sirtalis). Our data enable strong rejection of the alternative (non-adaptive) hypotheses, because the relationship between head and body size differed significantly among populations, the geographic distance separating pairs of populations explained less than 1.5% of their divergence in allometric coefficients, and the within-population allometric coefficients were higher than the among-population coefficients in each species. In addition, the geographical variability of allometric coefficients in females did not parallel that in males, suggesting that allometric coefficients have evolved independently in the two sexes. Phenotypic plasticity also cannot explain the data, because laboratory studies show that the allometric relationship between head size and body size is relatively insensitive to differing growth rates. We conclude that the intraspecific head size divergence in these snakes is better explained by spatially heterogeneous selection to optimize prey handling ability, than by non-adaptive processes.
Females of Lampruphulis delicata are dimorphic for colour pattern, the difference between morphs ... more Females of Lampruphulis delicata are dimorphic for colour pattern, the difference between morphs being the presence or absence of a distinct white mid-lateral stripe. A less distinct striped morph occurs also in males. We evaluated alternative hypotheses for the maintenance of this polymorphism by examining temporal and spatial variation in morph frequency, testing for differential selection among morphs using data on body size and reproductive traits from preserved specimens, and experimentally manipulating colour pattern in freeranging lizards of both sexes, to assess the influence of the lateral stripe on survival rates. We found that the relative frequency of striped individuals varied among populations and decreased from north to south in both sexes, coincident with an increasing incidence of regenerated tails. Morph frequencies did not change through time within a population. Striped gravid females appeared to survive better and produced larger clutches than did non-striped females. In our experimental study, the relationship between survival and colour morph differed between the two sexes; males painted with a white lateral stripe had lower survival than control (brown stripe) males, but survival did not differ between striped and control females. The different response in the two sexes may be due partly to differences in temperature and microhabitat selection. We propose that the white lateral stripe decreases susceptibility to predators in gravid females but increases risk of predation in males, especially in combination with low temperatures. The polymorphism might be maintained by: ( I ) opposing fitness consequences of the stripe in males and females; ('2) sex-specific habitat selection; and (3) gene flow in combination with spatial variation in relative fitness of the two morphs.
Habitat selection behavior is affected by complex interplays between competing requirements. Here... more Habitat selection behavior is affected by complex interplays between competing requirements. Here we combine field observations with laboratory experiments to examine how thermal benefits and predator avoidance influences habitat selection by different color morphs of the pygmy grasshopper Tetrix undulata. The composition of substrate types and surface temperatures in areas selected by free-ranging individuals did not reflect relative availability, and varied among morphs and sexes. Surface temperatures of selected habitats deviated less from the range of preferred body temperatures than would result from a random utilization of surface temperatures, suggesting that grasshoppers selected habitats with thermal properties which were suitable for maintaining preferred body temperatures. The thermal property of habitats occupied by different color morphs suggests that darker morphs (which absorb more solar radiation) selected cooler habitats to avoid overheating. Dissimilarities in substrate use among color morphs in the field and laboratory emphasize a role also of predator avoidance by background matching for habitat choice. The degree of habitat selectivity was lowest in the striped morph, supporting the notion that a disruptive color pattern may constitute a solution to the trade-off between relative crypsis in different visual backgrounds. Finally, individuals modified their habitat use when subjected to elevated risk of predation, showing that habitat choice is governed by conflicting priorities. Collectively, our findings suggest that, as a result of direct and indirect effects of coloration on performance, alternative color morphs use different solutions to the trade-off between competing requirements. Our results also lend support to the notion that relative fitness of alternative color morphs and sexes may be dependent on microhabitat selection, as predicted by the theory of multiple niche polymorphisms.
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Papers by Anders Forsman