Colour morph related performance in the meadow grasshopper Chorthippus parallelus (Orthoptera,Acrididae)

Abstract 1. Polymorphism has been described for a number of herbivorous insects, but little is known about whether differences in body colour cause fitness differences. In Chorthippus parallelus, three main colour morphs occur, namely brown, green, and dorsally striped. 2. The present study examined colour morph abundances and morph‐related differences in body size, oviposition rate, and offspring numbers in females of C. parallelus collected in 15 montane grasslands. The study also examined the effect of plant species richness, composition, community productivity, and solar radiation on colour morph frequency and fitness. 3. The relative frequencies of the three colour morphs was 31.7% (brown), 33.1% (green), and 35.2% (dorsally striped), but the morphs were not evenly distributed across the 15 sites. 4. There was no effect of the habitat variables on the distribution of the green and the striped morph in the study sites, however 80% of the variation in the abundances of the brown morph was explained by plant species richness and composition. 5. Grasshopper size was equal among the morphs. Brown females laid significantly more egg pods than the green and dorsally striped morphs. There were no significant differences in offspring numbers among the colour morphs. 6. Body colour in C. parallelus seems to be a fitness‐relevant trait, raising the question of the evolutionary maintenance of polymorphism.     

Differential visual predation on morphs of Timema cristinae (Phasmatodeae:Timemidae) and its consequences for host range

Timema cristinae is a herbivorous insect that exhibits polymorphism for body coloration (green, red and grey morphs) and for pattern (striped, expressed only in the green morph, and unstriped). The striped green morph is associated with ceanothus ( Ceanothus spinosus ) and the unstriped green morph is associated with chamise ( Adenostoma fasciculatum ). This study examines the relative vulnerabilities to predation of the different pattern and colour morphs on their natural backgrounds. The vulnerabilities of the striped and unstriped morphs on their two food plants were tested using uncaged wild birds (Scrub Jays) and captive western fence lizards. Strong differential predation was observed suggesting that each morph is most cryptic on the food plant on which it is most common. Furthermore, in a mark-recapture experiment in a patch of ceanothus the unstriped and red morphs were recaptured in higher proportion than the other morphs. The vulnerabilities of the grey and green morphs on the ground and foliage were tested using lizards. The grey morph was more vulnerable on the plants than the green morph, but the inverse was observed on the ground (where they drop after a disturbance). This may be why the grey morph is not associated with specific food plants. The striped and colour polymorphisms in T. cristinae appear to be an evolutionary consequence of differential predation on different backgrounds. The implications of differential predation to food-plant utilization are discussed.     

Predation Determines Different Selective Pressure on Pea Aphid Host Races in a Complex Agricultural Mosaic

Field assessments were conducted to examine the interplay between host plant and predation in complex agricultural mosaic on pea aphid clover and alfalfa races. In one experiment, we examined the relative fitness on clover race (CR) and alfalfa race (AR) pea aphids on broad bean, red clover and alfalfa alone. But because clover is typically grown in a more complex agricultural mosaic with alfalfa and broad bean, a second experiment was conducted to assess the fitness consequences under predation in a more complex agricultural field setting that also included potential apparent competition with AR pea aphids. In a third experiment we tested for the effect of differential host race density on the fitness of the other host race mediated by a predator effect. CR pea aphids always had fitness losses when on broad bean (had lower fitness on broad bean relative to red clover) and fitness benefits when on red clover (higher fitness on red clover relative to broad bean), whether or not in apparent competition with alfalfa race aphids on bean and alfalfa. AR suffered fitness loss on both alfalfa and bean in apparent competition with CR on clover. Therefore we can conclude that the predation rate between host races was highly asymmetrical. The complexity of the agricultural mosaic thus can influence prey selection by predators on different host plants. These may have evolutionary consequences through context dependent fitness benefits on particular host plants.     

<Emphasis Type="Italic">Aphidius ervi</Emphasis> Preferentially Attacks the Green Morph of the Pea Aphid, <Emphasis Type="Italic">Acyrthosiphon pisum</Emphasis>

The pea aphid, Acyrthosiphon pisum Harris (Hemiptera: Aphididae) is found in red and green color morphs. Previous work has suggested that the aphidiine parasitoid Aphidius ervi Haliday preferentially attacks green pea aphids in the field. It is not clear whether these results reflect a real preference, or some unknown clonal difference, such as in immunity, between the aphids used in the previous studies. We used three susceptibility-matched pairs of red and green morph pea aphid clones to test for preferences. In a no-choice situation, the parasitoids attacked equal proportions of each color morph. When provided with a choice, A. ervi was significantly more likely to oviposit into colonies formed from green morphs when the neighboring colony was formed from red morph aphids. In contrast, red morphs were less likely to be attacked when their neighboring colony was of the green morph. By preferentially attacking green colonies, A. ervi may reduce the likelihood of intraguild predation, as it is suggested that visually foraging predators preferentially attack red aphid colonies. Furthermore, if this host choice behavior is replicated in the field, we speculate that color morphs of the pea aphid may interact indirectly through their shared natural enemies, leading to intraspecific apparent competition.     

Interaction between female mating preferences and predation may explain the maintenance of rare males in the pentamorphic fish Poecilia parae

Variation in mating preferences coupled with selective predation may allow for the maintenance of alternative mating strategies. Males of the South American live‐bearing fish Poecilia parae fall in one of five discrete morphs: red, yellow, blue, stripe‐coloured tail (parae) and female mimic (immaculata). Field surveys indicate that the red and yellow morphs are the rarest and that their rarity is consistent across years. We explored the role of variable female mating preference and selective predation by visual predators in explaining the rarity of red and yellow males, and more generally, the maintenance of this extreme colour polymorphism. We presented wild‐caught P. parae females and Aequidens tetramerus, the most common cichlid predator, with the five male colour morphs in separate trials to determine mating and prey preferences, respectively. We found that a large proportion of females shared a strong preference for the rare carotenoid‐based red and yellow males, but a distinct group also preferred the blue and parae morphs. The cichlid predator strongly preferred red and yellow males as prey. Together, these results suggest that the interaction between premating sexual selection favouring and predation acting against the red and yellow morphs may explain their rarity in the wild. The trade‐off between sexual and natural selection, accompanied by variation in female mating preferences, may therefore facilitate the maintenance of the striking colour polymorphism in P. parae.     

The influence of light environment on host colour preference in a parasitoid wasp

1. Visual chromatic cues and contrast effects are widely used by insects in behaviours involving host/prey/mate‐finding and recognition. However, naturally changing light conditions may challenge the visual perception of cues for these organisms. 2. We used the host/parasitoid system Acyrthosiphon pisum/Aphidius ervi to determine if apparent visual preference of the wasp for green over pink host aphids was a visually based choice or a post‐attack mechanism based on host susceptibility depending on anti‐parasitoid symbiotic bacteria. 3. The study tested the ability of the wasp to recognise and attack pea aphid clones expressing variation based on colour and/or symbionts under a broad range of LED‐controlled light environments mimicking natural variations. 4. Results showed that the amount of reflected light of pink morphs was about half that of the green morphs in the cyan‐green components. Both host colours were recognised and attacked under all tested light conditions, even red light (660 nm). The previously reported preference of A. ervi for green pea aphids, was clear only for naive females given a choice between two aphid colours under all light environments, but quickly disappeared. 5. Wasps showed no tendency of avoiding oviposition in clones with defensive symbionts. 6. These findings suggest that variable rates of pea aphid parasitism by A. ervi in fields do not depend on host colour discrimination, but rather on susceptibility variation among aphid clones in allowing larval development after egg‐laying. Further studies should consider deeper investigation of the impact of red lights used in modified light environments in greenhouses and the proportion of host colour morph available.     

Body colour and genetic variation in winged morph production in the pea aphid

Aphids (Homoptera: Aphidoidea) produce a number of different phenotypes in their life-cycle, among which are winged (alate) and wingless (apterous) morphs. Lowe & Taylor (1964) and Sutherland (1969a, b) were the first to suggest that aphid clones differ in their propensity to produce the winged morph and that in the pea aphid (Acyrthosiphon pisum Harris), this propensity is linked to the colour of the phenotype. We tested for the occurrence of genetic variation in winged morph production by rearing individuals from red and green clones of pea aphid under wing-inducing (crowding) and control conditions, and scored the phenotypes of their offspring. Clones differed significantly in alate production and red clones produced on average a higher proportion of winged morphs than green clones. Importantly, however, there was considerable variation between clones of the same colour. Broad-sense heritabilities of winged morph production were 0.69 (crowding treatment) and 0.63 (control). Clones also differed in the number of offspring they produced. When exposed to the crowding stimulus, aphids deferred offspring production, resulting in a higher number of offspring produced in the crowding treatment than in the control.     

不同大豆品种上红绿两种色型豌豆蚜的种群参数

【目的】探讨不同大豆品种对红绿两种色型豌豆蚜Acyrthosiphon pisum种群参数的影响,为大豆品种的抗蚜性评价及豌豆蚜的预测预报与综合防治提供试验依据。【方法】在光周期10L∶14D、温度23±1℃,相对湿度60%±10%,光照强度212μmol/m~2·s的人工气候箱中,观察和分析4个大豆品种(汾豆牧绿2号、南夏豆25、南黑豆20和南豆5号)叶片上红绿两种色型豌豆蚜的成虫寿命、繁殖和种群生命表参数。【结果】在大豆品种汾豆牧绿2号上,红绿两种色型豌豆蚜成虫寿命均最短,繁殖力均最弱,且红色型豌豆蚜成虫寿命比绿色型长1.07 d,内禀增长率是绿色型豌豆蚜的29.93倍;在南夏豆25上,红色型豌豆蚜成虫寿命比绿色型长2.46 d,内禀增长率是绿色型豌豆蚜的5.86倍;在南黑豆20上,红色型豌豆蚜成虫寿命比绿色型长2.47 d,内禀增长率是绿色型豌豆蚜的1.54倍;在南豆5号上,红色型豌豆蚜成虫寿命比绿色型短0.02 d,内禀增长率是绿色型豌豆蚜的1.41倍。【结论】不同大豆品种对红色和绿色型豌豆蚜种群参数的影响不同,且两种色型豌豆蚜对不同大豆品种适应性反应也不相同。     

Biased predation could promote convergence yet maintain diversity within Müllerian mimicry rings of Oreina leaf beetles

Müllerian mimicry is a classic example of adaptation, yet Müller's original theory does not account for the diversity often observed in mimicry rings. Here, we aimed to assess how well classical Müllerian mimicry can account for the colour polymorphism found in chemically defended Oreina leaf beetles by using field data and laboratory assays of predator behaviour. We also evaluated the hypothesis that thermoregulation can explain diversity between Oreina mimicry rings. We found that frequencies of each colour morph were positively correlated among species, a critical prediction of Müllerian mimicry. Predators learned to associate colour with chemical defences. Learned avoidance of the green morph of one species protected green morphs of another species. Avoidance of blue morphs was completely generalized to green morphs, but surprisingly, avoidance of green morphs was less generalized to blue morphs. This asymmetrical generalization should favour green morphs: indeed, green morphs persist in blue communities, whereas blue morphs are entirely excluded from green communities. We did not find a correlation between elevation and coloration, rejecting thermoregulation as an explanation for diversity between mimicry rings. Biased predation could explain within‐community diversity in warning coloration, providing a solution to a long‐standing puzzle. We propose testable hypotheses for why asymmetric generalization occurs, and how predators maintain the predominance of blue morphs in a community, despite asymmetric generalization.     

Prey Preference of Aphidoletes Aphidimyza on Acyrthosiphon Pisum: Effect of Prey Color and Size

Color polymorphism in insects as well as factors contributing to its occurrence and maintenance have been of a great interest. Pea aphid (A. pisum) exhibits a noteworthy color polymorphism which occurs as red and green. The preference of the predatory gall midge Aphidoletes aphidimyza (Rondani) for the two color morphs of pea aphid at two life stages (adult and second instar) was investigated. Red adults, red nymphs, green adults, and green nymphs were offered to the larvae of the predator in different sets. The larvae attacked red aphids more than green ones of a same size. But whenever adults were offered along with nymphs, the nymphs experienced significantly more attacks. Although visual cues could result in more predation on red aphids, our findings showed that the size of aphids was the more important factor. The results showed that previous findings, suggesting more predation on the red morph, are valid when the same size of aphids is used. The ovipositing females exhibited no differences in oviposition choice between stalks infested with either of two colors.     

Differential predation on colour morphs of the Midas cichlid, Cichlasoma citrinellum

Predation has often been invoked as a selective agent affecting colour patterns in a wide range of organisms, but data relating susceptibility to predation and objective measures of conspicuousness are rare. To investigate relative crypticity in free-swimming fishes, two colour morphs of the Midas cichlid were exposed to predation by largemouth bass, Micropterus salmoides. The two colour morphs, normal and amelanic golds, were viewed by predators in ponds against a uniform, natural background. Swimming pools, 4 m in diameter, were stocked with equal numbers of gold and normal juvenile Midas cichlids. Bass took a significantly higher proportion of normal-coloured fishes (69·2% of fish) than gold morphs overall, and in a significantly higher proportion of trials (68·8% of experiments). No differences were found between colour morphs in their behaviour or in the willingness of bass to attack either morph. The significance of these results are discussed in relation to the visual system of the predator and the relative conspicuousness of the prey colour patterns.     

The use of visual cues in host evaluation by aphidiid wasps I. Comparison between threeAphidius parasitoids of the pea aphid

Host evaluation behaviour was examined in three species of aphid parasitoids,Aphidius ervi haliday,A. pisivorus Smith, andA. smithi Sharma & Subba Rao (Hymenoptera: Aphidiidae). Parasitoids were provided under laboratory conditions with three kinds of hosts representing two aphid species: (green) pea aphid,Acyrthosiphon pisum (Harris), and green and pink colour morphs of the alfalfa aphid,Macrosiphum creelii Davis. Females of all threeAphidius species distinguished between aphids on the basis of colour, movement, and host species. Patterns of host acceptance by parasitoids were species-specific. InA. ervi, host preference was the same in light and dark conditions: pea aphid>green alfalfa aphid≫pink alfalfa aphid. In contrast,A. pisivorus attacked and accepted pea aphid and green alfalfa aphid equally in the light and preferred both of these over pink alfalfa aphid; however, it made no distinction between pea aphid and pink alfalfa aphid in the dark. Females ofA. smithi attacked all three kinds of hosts (pea aphid>green alfalfa aphid≫pink alfalfa aphid) but apparently laid eggs only in pea aphid. The frequencies of attack and oviposition by all wasps were higher on ‘normal’ pea aphids than on those anaesthetized with CO₂. Host recognition is confirmed by chemical cues in the aphid cuticle that are detected during antennation, and host acceptance is dependent on an assessment of host quality during ovipositor probing.     

Prey selection by wild birds can allow novel and conspicuous colour morphs to spread in prey populations

Conspicuous warning coloration helps to protect prey because it signals to potential predators that the prey is unprofitable. However, such signals only work once predators have come to associate the conspicuous colour with the unprofitability of the prey. The evolution of warning coloration is generally considered to be paradoxical, because it has traditionally been assumed that the first brightly coloured individuals would be at an immediate selective disadvantage because of their greater conspicuousness to predators that are naïve to the meaning of the signal. As a result, it has been difficult to understand how a novel conspicuous colour morph could ever avoid rapid extinction, and instead survive and spread in the population until predators have become educated about the signal. In the present study, we experimentally simulated the appearance of a single novel coloured mutant in small populations (20 individuals) of palatable artificial pastry "prey". The colour morph frequencies in each "generation" of prey (presented on successive days of a trial) were determined by the relative survival of the previous generation under predation by free-living birds. We found that the novel colour morphs regularly persisted and increased from a starting frequency of 1/20 to reach fixation (100%), despite being fully palatable, even when the novel morph was much more conspicuous against the background than the familiar morph. This was true for both green (not normally considered a warning colour) and red (a classic warning colour) novel morphs. Novel colours reached fixation significantly faster than could be accounted for by random drift, indicating differential predation in relation to prey colour by the birds. Our experiments show that the immediate demise of a fully palatable new prey morph is not an inevitable outcome of predator behaviour, because even very conspicuous prey can gain protection from conservative foragers, simply by being novel.     

Morph‐dependent fitness and directional change of morph frequencies over time in a Dutch population of Common buzzards Buteo buteo

How genetic polymorphisms are maintained in a population is a key question in evolutionary ecology. Previous work on a plumage colour polymorphism in the common buzzard Buteo buteo suggested heterozygote advantage as the mechanism maintaining the co‐existence of three morphs (light, intermediate and dark). We took advantage of 20 years of life‐history data collected in a Dutch population to replicate earlier studies on the relationship between colour morph and fitness in this species. We examined differences between morphs in adult apparent survival, breeding success, annual number of fledglings produced and cumulative reproductive success. We found that cumulative reproductive success differed among morphs, with the intermediate morph having highest fitness. We also found assortative mating for colour morph, whereby assortative pairs were more likely to produce offspring and had longer‐lasting pair bonds than disassortative pairs. Over the 20‐year study period, the proportion of individuals with an intermediate morph increased. This apparent evolutionary change did not just arise from selection on individual phenotypes, but also from fitness benefits of assortative mating. The increased frequency of intermediates might also be due to immigration or drift. We hypothesize that genetic variation is maintained through spatial variation in selection pressures. Further studies should investigate morph‐dependent dispersal behaviour and habitat choice.     

Spatial variability in seed predation in Primula farinosa: local population legacy versus patch selection

Spatio-temporal variation in seed predation may strongly influence both plant population dynamics and selection on plant traits. The intensity of seed predation may depend on a number of factors, but the relative importance of previous predator abundance (“local legacy”), spatial distribution of the host plant, environmental factors and plant characteristics has been explored in few species. We monitored seed predation in the perennial herb Primula farinosa, which is dimorphic for scape length, during 5 consecutive years, in a 10-km × 4-km area comprising 79 P. farinosa populations. A transplant experiment showed that the seed predator, the oligophagous tortricid moth Falseuncaria ruficiliana, was not dispersal limited at the spatial scale corresponding to typical distances between P. farinosa populations. Correlations between population characteristics and incidence and intensity of seed predation varied among years. The incidence of the seed predator was positively correlated with host population size and mean number of flowers, while intensity of seed predation in occupied patches was positively related to the frequency of the long-scaped morph in 2 years and negatively related to host population size in 1 year. In both scape morphs, predation tended to increase with increasing frequency of the long morph. There was no evidence of a local legacy; incidence and intensity of seed predation were not related to the abundance of the seed predator in the population in the previous year. Taken together, the results indicate that among-population variation in seed predation intensity is determined largely by patch selection and that the seed predator’s preference for tall and many-flowered inflorescences may not only affect selection on plant traits within host plant populations, but also the overall intensity of seed predation.     

Differential predation on sexes affects colour polymorphism of the isopod Idotea baltica (Pallas)

Variable selection, including spatio-temporal variation, frequency-dependent selection and differential selection due to habitat choice, may maintain polymorphism in heterogeneous environments. We studied predation as a selective agent on colour polymorphism of the aquatic isopod I baltica. Variable predation on this species can arise from at least three sources. First, apostatic selection was studied by testing the formation of preferences on colour morphs in the perch, a common predator of I baltica. Such acquired preferences should induce apostatic selection. While our results indicate some acquired preferences, there was significant heterogeneity in the behaviour of predator individuals. Second, temporal variation in selection can arise due to habitat shift from the green algae juvenile habitat to the bladderwrack adult habitat, and the consequent change in the crypsis of the morphs. Different crypsis between sexes probably promoted high predation mortality among females in the juvenile habitat. The high rate of male mortality during the breeding period, on the other hand, was presumably due to their high mate-searching activity. Third, the sex-dependent habitat choice of I baltica leads to sexual differences in the susceptibility of morphs to predation. Predators preferred the white-spotted morph over the uniform one in males but not in females, supporting the 'dimorphic niche' hypothesis as an explanation of sexual differences in morph frequencies. Finally, no evidence was found that the colouration patterns were under sexual selection. We therefore conclude diat variable predation is the most promising explanation for the maintenance of polymorphism in I. baltica.     

Direct and indirect selection on floral pigmentation by pollinators and seed predators in a color polymorphic South African shrub

The coexistence of different color morphs is often attributed to variable selection pressures across space, time, morph frequencies, or selection agents, but the routes by which each morph is favored are rarely identified. In this study we investigated factors that influence floral color polymorphisms on a local scale in Protea, within which approximately 40% of species are polymorphic. Previous work shows that seed predators and reproductive differences likely contribute to maintaining polymorphism in four Protea species. We explored whether selection acts directly or indirectly on floral color in two populations of Protea aurea, using path analysis of pollinator behavior, nectar production, seed predation, color, morphology, and maternal fecundity fitness components. We found that avian pollinators spent more time on white morphs, likely due to nectar differences, but that this had no apparent consequences for fecundity. Instead, the number of flowers per inflorescence underpinned many of the reproductively important differences between color morphs. White morphs had more flowers per inflorescence, which itself was positively correlated with nectar production, seed predator occurrence, and total long-term seed production. The number of seeds per plant to survive predation, in contrast, was not directly associated with color or any other floral trait. Thus, although color differences may be associated with conflicting selection pressures, the selection appears to be associated with the number of flowers per inflorescence and its unmeasured correlates, rather than with inflorescence color itself.     

The role of vision and color in the close proximity foraging behavior of four coccinellid species

The role of vision and color in close-proximity foraging behavior was investigated for four species of lady beetles: Coccinella septempunctata, Hippodamia convergens, Harmonia axyridis, and Coleomegilla maculata. The effect of light level and color cues on consumption rates varied among the four predator species. The consumption rates of these predators on the pea aphid Acyrthosiphon pisum (Harris) was measured under light and dark conditions. C. septempunctata,H. convergens, and Ha. axyridis consumed significantly more aphids in the light than in the dark, while the consumption rate of Col. maculata was not affected by light level. Foraging ability was also measured on red and green color morphs of the pea aphid on red, green, and white backgrounds. C. septempunctata consumed significantly more of the aphid morph that contrasted with the background color, and showed no difference between morphs on the white background. H. axyridis consumed significantly more red morph aphids regardless of background. The remaining two species showed no difference in consumption rates on the two color morphs. The variation in the use of visual cues demonstrates how different species of predators can exhibit different foraging behaviors when searching for the same prey. Received: 4 August 1997 / Accepted: 3 February 1998     

Learning by the parasitoid wasp, Aphidius ervi (Hymenoptera: Braconidae), alters individual fixed preferences for pea aphid color morphs

Learning, defined as changes in behavior that occur due to past experience, has been well documented for nearly 20 species of hymenopterous parasitoids. Few studies, however, have explored the influence of learning on population-level patterns of host use by parasitoids in field populations. Our study explores learning in the parasitoid Aphidius ervi Haliday that attacks pea aphids, Acyrthosiphon pisum (Harris). We used a sequence of laboratory experiments to investigate whether there is a learned component in the selection of red or green aphid color morphs. We then used the results of these experiments to parameterize a model that examines whether learned behaviors can explain the changes in the rates of parasitism observed in field populations in South-central Wisconsin, USA. In the first of two experiments, we measured the sequence of host choice by A. ervi on pea aphid color morphs and analyzed this sequence for patterns in biased host selection. Parasitoids exhibited an inherent preference for green aphid morphs, but this preference was malleable; initial encounters with red aphids led to a greater chance of subsequent orientation towards red aphids than predicted by chance. In a second experiment, we found no evidence that parasitoids specialize on red or green morphs; for the same parasitoids tested in trials separated by 2 h, color preference in the first trial did not predict color preference in the second, as would be expected if they differed in fixed preferences or exhibited long-term (> 2 h) learning. Using data from the two experiments, we parameterized a population dynamics model and found that learning of the magnitude observed in our experiments leads to biased parasitism towards the most common color morph. This bias is sufficient to explain changes in the ratio of aphid color morphs observed in field sites over multiple years. Our study suggests that for even relatively simple organisms, learned behaviors may be important for explaining the population dynamics of their hosts.     

Reproductive success and risk of predation in normal and melanistic colour morphs of the adder, Vipera berus

In a population of adders (Vipera berus) in Southwest Sweden, melanistic males were heavier than normal coloured males of the same length. Victory in male-male sexual combats was positively related to size. Higher risk of predation in the black morph was inferred from experiments showing a high predator attack rate on models of the black morph. Even the bright colour in newly moulted basking males of the normal morph gives cryptic protection. In females, melanism probably also affects body size and risk of predation by visually searching predators. The thermoregulatory influence of black colour, the reproductive success and the maintenance of two colour morphs in the population are discussed.