An unusual phylogeography in the bushcricket Ephippiger ephippiger from Southern France

Pleistocene glaciations have played a major role in species divergence. The bushcricket Ephippiger ephippiger shows unusual patterns of intraspecific variation in multiple traits across Southern Europe. This is centred in Southern France, and evidence implies that it results from secondary contact after differentiation in Pleistocene refugia. However, the possible time scales involved, locations of the refugia and patterns of expansion remain obscure. This study sequenced the COII (507 BP) and cyt b (428 BP) mitochondrial genes to examine the intraspecific phylogeography of Western European samples of E. ephippiger. A minimum evolution tree revealed little resolution between described subspecies of E. ephippiger. Strikingly, populations from the Pyrenees and Mediterranean coastal region contained a complex genetic structure corresponding to major river valleys, independent of the traditional taxonomy. Samples of the subspecies E. e. vitium formed a distinct clade, perhaps supporting their taxonomic status. However, other forms (cruciger and cunii) were not genetically distinct, which is surprising given differences in their morphology and behaviour. The extent of the genetic divergence between Pyreneen valleys is unexpectedly deep, with average Tamura-Nei distances of around 14% (net distances of 11%) separating the main clades of coding COII sequences. Cyt b showed a similar pattern, but was confounded by some non-coding probable pseudogenes. If a conventional insect molecular clock is applied, these cryptic clades must pre-date the Pleistocene, and hypotheses for their history are discussed. However, mtDNA divergence in Ephippiger is not evolving in a clock-like manner, because a likelihood ratio test rejects clock assumptions for the COII sequences.     

Effects of ambient colour on colour preference and growth of juvenile rainbow trout Oncorhynchus mykiss (Walbaum)

Colour preference of individual juvenile rainbow trout Oncorhynchus mykiss was tested at 1 and 12° C, and also at 12° C after a 42 day growth experiment under white, blue, green, yellow or red ambient colour. All experiments were carried out under controlled laboratory conditions and the preference was assessed by the location of the fish in a preference tank with four chambers. Rainbow trout showed a preference for blue and green at 1° C and for green at 12° C. After the growth experiment the fish reared in blue tanks preferred blue and green but green was the most preferred colour for the fish reared in green, yellow and red tanks. Yellow and especially red chambers were avoided, irrespective of the ambient colour during the growth trial. The final mass of fish reared in the red aquaria was significantly smaller than that of the fish in green tanks. In addition, when the data of the preference tests were correlated with the data of the growth experiment using mean values of the four tested colours, a very good linear relationship was observed between the preference ( i.e. visit frequency in coloured compartments) and growth rate as well as food intake. When considering the results both from the preference and growth trials it is suggested that green is the best environmental colour for rearing juvenile rainbow trout while rearing in a red environment cannot be recommended.     

Observations on skin colour changes in juvenile lumpsuckers

This study reports on an experimental investigation of colour change in young lumpsuckers Cyclopterus lumpus , employing Munsell colour charts to quantify skin hue, colour value and chroma. Juvenile lumpsuckers can live pelagically close to the sea surface, or can attach to floating seaweed and debris with their suckers. When swimming they adopt a cryptic default light green coloration, with the dorsum being darker than the belly (countershading). This default colour is also seen in complete darkness. When attached to objects they take a colour stimulus from below and to the sides and change colour. They can achieve good colour matches (noticeable in 3 min; complete within 15 min) with substrata that are green-yellow in colour, especially the brown alga Ascophyllum nodosum . Colour matching is largely dependent upon changes in colour value (i.e. lightness/darkness) produced by melanin dispersal or concentration in the melanophores. Thus, on artificial backgrounds outside the range of achievable hue matching, they still achieve good matchings of colour value. Slight hue changes were recorded (probably reflecting interaction of basic skin colour with melanin dispersal state), but chroma (brightness) variations were negligible. Lumpsucker colour depends upon light intensity, allowing sustained colour matching as weed illumination changes.     

The major carotenoid pigments of the grain aphid, Sitobion avenae (F.) (Hemiptera: Aphididae)

The economically important grain aphid, Sitobion avenae (F.) shows colour polymorphism, with brown and green forms predominating. Colour is determined both genetically and in response to environmental factors, including nutrition. The biological significance of the colour polymorphism is unknown, although seasonal changes occur in the frequency of colour morphs in the field, whilst the brown morph may have adaptive significance in terms of hymenopterous endoparasitism. The ground colour of aphids is produced by haemolymph pigments, aphins (glucosides) and carotenoids. The latter may be under the synthetic control of intracellular endosymbiotic bacteria. In this study, the major carotenoid pigments of a brown and a green clone of S. avenae were examined using thin layer chromatography (TLC) and high-performance liquid chromatography (HPLC), and their absorbance spectra recorded. Using TLC, the brown clone produced five bands of different R_f, ranging from yellow, to orange-pink to pink in colour. In contrast, the green clone gave only a single yellow band of higher R_f than any of the bands of brown aphids. Following separation of carotenoids by HPLC, brown aphids gave seven peaks and green aphids five. Comparison of absorbance maxima with known published values for carotenoids provides strong evidence for the identification of four of the carotenoid pigments from brown aphids (RB-4, 3,4-didehydrolycopene; RB-5, torulene; RB-6; lycopene; RB-7, γ-carotene) and one from green aphids (RG-2, α-carotene). The other carotenoids remain unidentified. The biosynthesis and possible biological relevance of the various pigments of S. avenae are briefly discussed.     

Effects of hatchling body colour and rearing density on body colouration in last-stadium nymphs of the desert locust, Schistocerca gregaria

Abstract.  The influences of hatchling character and rearing density on body colour at the last-nymphal stadium are investigated for the desert locust, Schistocerca gregaria . Hatchlings are divided into five groups based on the darkness of the body colour and reared either under isolated or crowded conditions. Two types of body colour variation at the last-nymphal stadium are separately analysed (i.e. the background colour and black patterns). Under isolated conditions, the background body colour is either greenish or brownish. Most individuals are greenish and the highest percentage of brownish insects is obtained from hatchlings with the darkest body colour. Under crowded conditions, the background colour is yellow or orange and the percentage of yellowish nymphs tends to decrease when they are darker at hatching. The intensity of black patterns differs depending on the body colour at hatching and subsequent rearing density. Most isolated-reared nymphs exhibit few or no black patterns but nymphs with some black patterns also appear, particularly among those that had been dark at hatching. Under crowded conditions, the black patterns become more intense when they are darker at hatching. Therefore, last-stadium nymphs with typical solitarious or gregarious body colouration appear when they have the phase-specific body colouration at hatching as well. The present results demonstrate that both body colour at hatching and rearing density during nymphal development influence body colouration at the last-nymphal stadium.     

Colour constancy in insects

Colour constancy is the perceptual phenomenon that the colour of an object appears largely unchanged, even if the spectral composition of the illuminating light changes. Colour constancy has been found in all insect species so far tested. Especially the pollinating insects offer a remarkable opportunity to study the ecological significance of colour constancy since they spend much of their adult lives identifying and choosing between colour targets (flowers) under continuously changing ambient lighting conditions. In bees, whose colour vision is best studied among the insects, the compensation provided by colour constancy is only partial and its efficiency depends on the area of colour space. There is no evidence for complete ‘discounting’ of the illuminant in bees, and the spectral composition of the light can itself be used as adaptive information. In patchy illumination, bees adjust their spatial foraging to minimise transitions between variously illuminated zones. Modelling allows the quantification of the adaptive benefits of various colour constancy mechanisms in the economy of nature. We also discuss the neural mechanisms and cognitive operations that might underpin colour constancy in insects.     

颜色对梨小食心虫产卵选择性的影响

梨小食心虫(Grapholitha molesta Busck)是一种重要的果树害虫,早春产卵喜在桃嫩梢叶上,为探明寄主颜色在其产卵选择中的作用,利用彩色卡纸模拟寄主颜色,室内比较了红、粉红、浅粉、橙黄、深黄、浅黄、青绿、深绿、浅绿、蓝、紫、褐色等12种不同颜色基质对其成虫产卵选择性的影响。结果发现:基质颜色对梨小食心虫的产卵选择性有显著影响,其产卵偏嗜浅黄和浅绿色,白色和黑色参比时其产卵选择率分别依次为68.9%、63.8%和64.1%、65.5%,蓝和浅粉色则表现一定的拒避作用,白色和黑色参比时其产卵选择率分别依次为47.7%、40.4%和47.2%、42.7%,且参比色对其产卵选择性影响差异显著。基质颜色对其产卵量亦有显著性影响,无论黑或白色参比,黄、绿颜色上的产卵量均较多,尤其是深黄、深绿和青绿色。基质颜色对1—7日龄梨小食心虫成虫的产卵选择性均有显著影响。白色参比时,2、3日龄蛾对浅绿和浅黄色的产卵选择率显著高于其他颜色;黑色参比时,2日龄时明显偏嗜浅绿色(79.7%),6日龄时明显偏嗜浅黄色(74.8%)。表明寄主颜色在梨小食心虫产卵场所选择中具有重要作用,为其卵期监测和防控中颜色应用乃至进一步揭示其产卵寄主选择机理提供了依据和参考。     

Receptor based models for spontaneous colour choices in flies and butterflies

Colour is one of several stimuli used by herbivorous insects in host choice. Insects have between 2 and 5 different types of photoreceptors to catch quanta of different wavelengths of the spectrum. Many insects have been shown to possess opponent neural interactions between the receptors that enable them to see colour. I present simple models to describe colour choices as functions of the receptor quantum catches and linear interactions of the receptor types. Models are applied to data sets obtained from own experiments and from the literature, on Pieris brassicae and P. rapae (Lepidoptera, Pieridae), Papilio aegeus (Papilionidae), Dacus oleae (Diptera, Tephritidae) and Eristalis tenax (Syrphidae). In fruit flies, detection of green fruit is based on an inhibitory interaction between a green-sensitive receptor type and a blue-sensitive receptor type. This might explain the preference many herbivorous insects have for yellow over green stimuli. Pollen feeding in hoverflies might have evolved from yellow pollen being a super-normal stimulus for herbivorous insects. In butterflies, an additional red-receptor is involved in the colour choice for an oviposition substratum and leads to them choosing green and not yellow. The models introduced in this study open new perspective for a physiological understanding of the design of visual stimuli for monitoring and trapping pest insects.     

A test of the relationship between cuticular melanism and immune function in wild‐caught Mormon crickets

Cuticular melanism and innate immune parameters can share common physiological pathways in insects, and this functional connection may contribute to the maintenance of insect colour polymorphisms. However, evidence linking colouration and immune function has been equivocal, particularly when tested in wild populations. The present study investigates phenotypic links between colouration and immune function in migratory Mormon crickets (Anabrus simplex, Haldeman), in which juveniles occur in conspicuous colour variants but mature to become uniformly melanic adults. Wild‐caught insects are used to evaluate the relationship between juvenile colouration and three immune parameters: encapsulation ability, lysozyme‐like activity and phenoloxidase activity. As nymphs, brown crickets are better able to encapsulate an inert implant introduced into the haemocoel than green crickets, although the difference is slight and ceases after they all become darkly‐coloured adults. By contrast, adults that develop from brown nymphs have a higher basal phenoloxidase activity than those that develop from green nymphs, regardless of the fact that all adults are brown. Intrinsic factors other than colouration exert larger effects on immunity: males show stronger encapsulation responses but lower phenoloxidase activity than females, suggesting a sex‐specific trade‐off between these two immune parameters, and adults exhibit higher immune function than nymphs. In summary, modest support is found for a correlation between cuticular melanism and increased immune function in wild Mormon crickets. Additional intrinsic factors such as developmental stage and sex appear to interact with colouration and have a more substantial connection to immune function in the wild.     

The influence of density on frequency-dependent selection by wild birds feeding on artificial prey

Previous work has demonstrated frequency-dependent selection by wild garden birds when feeding on green and brown pastry ''baits''. When the density of baits is low, the common colour is eaten disproportionately more than the rare colour (apostatic selection), and when the density is very high, the rare colour is eaten disproportionately more than the common (anti-apostatic selection). We explored the relationship between frequency-dependent predation and density in an experiment at 16 separate sites, using four levels of density and two frequencies of green and brown. Analysis of estimates of log-relative risk ratios showed little evidence for frequency-independent selection, but frequency-dependent selection changed gradually from apostatic at low density to anti-apostatic at high density. The validity of these conclusions in terms of individual bird behaviour was confirmed by Monte-Carlo simulations. We thus conclude that selection by wild birds feeding on green and brown artificial prey is frequency dependent, and that the strength and direction of this selection changes with prey density in a gradual and predictable manner.     

A seed parasitoid wasp prevents berries from changing their colour,reducing their attractiveness to frugivorous birds

1. Frugivorous and seed‐feeding insects may alter the traits of fruits, such as shape and size, which may influence fruit attractiveness to frugivorous birds. Consequently, trait‐mediated interactions may occur in systems where plants, seed‐dispersing frugivorous vertebrates, and frugivorous or seed‐feeding insects interact. We investigated colour manipulation in Ilex integra Thunb. berries caused by the seed parasitoid wasp Macrodasyceras hirsutum Kamijo and how that manipulation relates to fruit attractiveness for frugivorous birds. 2. In winter, the colour of I. integra berries varied from green to red, but most berries were greenish, indicating that the berries were immature. Berry dissection indicated that the number of live parasitoid larvae present within each berry was closely related to berry colour – the greater the number of live larvae, more intense is the green colour of the berry. However, the wasp larvae did not modify the shape or size of the berries. More than 98% of berries that were protected from the insects by gauze bags ripened and turned red. In the present study, berries with unfertilised seeds alone turned red. Field‐feeding preference tests showed that the brown‐eared bulbul Hypsipetes amaurotis Temminck preferred red berries to green berries. 3. We demonstrated that the seed parasitoid wasp manipulates the berry colour, but not its shape or size, in a density‐dependent manner. Because green berries suffered less from bird foraging, we believe that this colour manipulation helps the wasps to avoid being killed by the birds. The present study indicates that manipulation by wasps may reduce the level of mutualism between the tree and seed‐dispersing birds.     

Selection by passerine birds is anti-apostatic at high prey density

Most of the results from past experiments with wild birds and green and brown pastry 'baits' have suggested that disproportionately more of the rare forms are eaten when bait density is high (i.e. selection is anti-apostatic). In two separate series of experiments we presented birds with dishes containing 270 baits of one colour and 30 of another. In series I, five different pairs of colours were presented simultaneously to wild birds at two sites. One colour of each pair was common at one site and the same colour was rare at the other site. After 35 days the ratios of the colours were reversed and the dishes were presented for another 35 days. There was a statistically significant tendency for the colours to be at a higher risk when rare. In series II, three caged blackbirds were offered green and brown baits in two dishes simultaneously; in one dish green was rare and in the other brown was rare. Selection over 6 days was anti-apostatic for all three birds combined but the data proved heterogeneous both between and within individuals. At any one time, each bird tended to concentrate on one colour, irrespective of whether that colour was rare or common. We believe that this behaviour leads to anti-apostatic selection, as has been observed in these and other experiments with pastry prey.     

Variation in plumage colour of the Great tit Parus major in relation to habitat, season and food

The plumage colour of nestling and adult Great tits Parus major major L. was studied at Trondheim (63 °N, 10 °E), Norway. Nestlings reared in deciduous woodland were yellower than those reared in coniferous woodland, those reared in 1983 were yellower than those in 1982, and the later in the breeding season they were reared the more yellow they were. Furthermore, adults were yellower than nestlings, and for both groups the males were yellower than the females. Experimental interchange of eggs between nests showed that the variation in plumage colour observed between broods was related primarily to the food and not to genetic differences. Sampling, by the neck-collar method, of the food brought by the parent birds showed that the yellower nestlings received a greater proportion of lepidopteran larvae, in particular greenish-coloured ones, than did the nestlings with a paler plumage. A speculation is whether or not the variation in plumage colour has some adaptive value (viz. background matching).
In avian systematics, plumage coloration has been used to distinguish and describe subspecies. The present study shows that such a classification should only be made with certain reservations, until possible variation in the food quality of the subspecies is known to have no influence on plumage coloration.     

The colour of fitness: plumage coloration and lifetime reproductive success in the tawny owl

We studied variation in plumage colour and life history in a population of tawny owls (Strix aluco) in southern Finland, using 26 years of data on individually marked male and female owls. Colour was scored on a semi-continuous scale from pale grey to reddish brown. Colour scoring was repeatable and showed a bimodal distribution (grey and brown morph) in both sexes. During the study period, colour composition was stable in the study population in both sexes. The sexes did not mate assortatively with respect to their colour. Colour was a highly heritable trait and was under selection. Grey-coloured male and female owls had a higher lifetime production of fledglings, and grey-coloured male (but not female) owls produced more recruits during their lifetime than brown individuals. Selection on colour was mediated through viability selection and not through fecundity selection. Our results reveal remarkably strong selection on a genetically determined phenotypic trait.     

The pale brown allele in Cepaea nemoralis

The pale brown colour morph in Cepaea nemoralis appears to be determined by an allele at the C (colour) locus ( C ^{P B}). Pale brown is dominant to yellow, codominant with pink and recessive to dark brown. It is linked to the B locus (which controls the presence or absence of banding on the shell), but not to the U locus, which determines whether there is one band or five. In segregations of pale brown and yellow there is a significant deficiency of pale brown, suggesting that there are differences in viability between the morphs.     

Colour, shape and defence in aphidophagous syrphid larvae (Diptera)

Colour patterns, shape and behaviour are analysed in 22 species of aphidophagous syrphid larvae in relation to their possible role in defence against visually hunting predators. Crypsis is the most important method of primary defence. There is much variation in colour pattern. Some species are translucent, others green, brown or otherwise coloured. Most have disruptive stripes, bars or other markings that break up the body outline at short range. There is one example of a bird dropping resemblance. Behavioural requirements for effective crypsis include diurnal immobility, resting in sites that provide maximum concealment, and slow interrupted locomotion. Secondary defences are less variable and involve catalepsis, attack with saliva, and rolling and dropping from the plant.     

The environmental control of colouration in a bushcricket, Mygalopsis marki Bailey (Orthoptera: Tettigoniidae)

The environmental factors proximately influencing variable colouration in the bushcricket, Mygalopsis marki , were studied in natural populations and under controlled laboratory conditions. Two major components of variable colouration were identified; a green/brown dimorphism in ground colour hue (dominant wave length) and continuous variation in the darkness of brown insects, representing a contrast between ground colour value (brightness) and the intensity of overlying black pattern. Ground colour hue differed between natural populations in different seasons and at different localities. The proportion of brown individuals was greater in populations developing over late summer than over spring at the same locality, and in populations from drier localities, as indicated by vegetation type, in the same season. Experimental studies showed that colour change occurred at moults, with green colouration promoted by increased water intake in developing nymphs. Ground colour value and pattern differed between natural populations at different localities, but not in different seasons. Ground colour value was less, and pattern was greater, in populations from localities with a dense cover of forbs and grasses. Experimental studies showed that ground colour value and pattern were influenced by the intensity of incident radiation received by developing nymphs, with decreased intensity promoting lower value and greater pattern.     

Visible variation in the dog-whelk, Nucella lapillus

Shell shape and colour variation is common in littoral molluscs. As far as shape is concerned, exposure to wave action acts as a strong agent of natural selection, but only speculation seems to have been directed towards elucidating the determinants of colour variation. In the dog-whelk ( Nucella lapillus (L.)) diet may play some part in producing the pigment for colour. However white and banded forms are often found in otherwise coloured colonies. This study describes the classification for colour and banding of 83,524 whelks from 444 areas around the British Isles and west Atlantic coast. Colour and banding were significantly correlated r =0.45±0.05). Because of the possible involvement of diet in shell colour, attention has been confined to banded forms. No obvious pattern in distribution could be discerned. Possible factors affecting the occurrence and incidence of banding are: climate; geology; exposure to wave action; predation; intra- or interspecific competition; cryptic genetical factors; or intraspecific recognition. The only one which was definitely associated with banding frequency was exposure ( r = 0.33±0.05), but there were so many exceptions to the correlation (e.g. highly exposed populations which were entirely unhanded), that this is unlikely to be causal. The most probable explanation seems to be the local spread of mutant forms (probably linked to fitness traits), which have not attained equilibrium frequencies–even though N. lapillus has apparently occurred in large colonies round much of the British coast since the end of the Pleistocene.     

Pupal colour dimorphism in a desert swallowtail (Lepidoptera: Papilionidae) is driven by changes in food availability,not photoperiod

1. The swallowtail butterfly Battus polydamas archidamas Boisduval, 1936, exhibits polyphenism for pupal coloration (green and brown). It is distributed across arid regions with winter rains and is monophagous on Aristolochia plants, which emerge after the winter rains and dry out the during summer. Thus, day length does not covary positively with host plant productivity. It was hypothesised that pupal colour was driven by food availability, not photoperiod. The benefits of pupal coloration matching the colour of pupation sites in terms of field survival were also investigated to evaluate the adaptive value of pupa colour. 2. Larvae were reared under a factorial array of two photoperiods (LD 10:14 h and LD 14:10 h) and two food availability regimes (leaves ad libitum and available every other day) to assess the frequency of green and brown pupae. Field survival of green and brown pupae was quantified in three commonly used habitats that differ in background coloration (cacti, rocks and shrubs). 3. Food availability determined pupal colour. Larvae in the ad libitum regime resulted mostly in green pupae, while those with restricted food were mostly brown. In contrast, photoperiod did not influence pupal colour. Survival probability of pupae placed on cacti was higher than those placed on rocks and shrubs, and the lowest predation risk across habitats was for green pupae on cacti. 4. Food availability plays a major role in the seasonal polyphenism for pupal colour of specialist butterflies inhabiting arid environments with winter rains.     

Use of ITS rDNA for discrimination of European green- and brown-banded sporocysts within the genus Leucochloridium Carus, 1835 (Digenea: Leucochloriidae)

Transmission of Leucochloridium species to their definitive avian hosts may be facilitated by the rhythmic movement of coloured sporocyst broodsacs in the ocular tentacles of infected snails. These broodsacs resemble caterpillars and by enticement increase the probability of predation by birds. Broodsac banding pattern and colour (green, yellow to red/brown) have traditionally formed part of the taxonomic criteria for the genus. In this study, sequence divergence of the 5.8S rDNA gene and associated internal transcribed spacers (ITS1 and ITS2) in two of the most frequently observed Leucochloridium taxa from Europe is related to broodsac type based on colour and banding pattern. The present green-banded broodsac (L. paradoxum Carus, 1835) and brown-banded broodsac forms (L. variae McIntosh, 1932) differ in ITS sequence by 6.8%, confirming their distinctness. No intraspecific differences were noted within each colour morph in specimens collected from Poland, Denmark or Norway, indicating that a single taxon of each type occurs in Europe. The significance of these findings to our understanding of metapopulation dynamics and evolutionary ecology of Leucochloridium is discussed.