Insect colour preference compared to flower colours in the Australian Alps

An apparent predominance of plant taxa with pale flowers in the alpine floras of Australia and New Zealand may be due to the prevalence of insects, such as flies, that prefer pale colours and the absence of other types of potential pollinators that are attracted to bright colours such as social bees and birds. In this study, the diversity of flower colours, and the preference of insects for different colours were examined for the largest contiguous alpine area in Australia, around Mt Kosciuszko. Out of an alpine flora of 204 taxa, 127 species were found to have large showy flowers. The most common flower colour among these taxa was white (53.5%), then yellow (21.3%), followed by pink (6.3%), and cream (6.3%). Only a handful of taxa had red, blue, brown, green, orange or purple flowers. When the colour preference of insects was tested using five different coloured traps (white, yellow, orange, red and purple), the most successful traps were white then yellow, with these two colours accounting for 66% of all individual insects collected. Diptera were the most common insects caught (576 insects greater than 4 mm in length, 31 morphotaxa) showing an apparent preference for white and yellow coloured traps over others. Therefore, the results add some support to the proposition that the 'white' flora of the Australian Alps may be associated with the colour preference of flies, which have previously been found to be the most common type of pollinators in the Kosciuszko alpine zone.     

Flower colours along an alpine altitude gradient, seen through the eyes of fly and bee pollinators

Alpine flowers face multiple challenges in terms of abiotic and biotic factors, some of which may result in selection for certain colours at increasing altitude, in particular the changing pollinator species composition, which tends to move from bee-dominated at lower elevations to fly-dominated in high-alpine regions. To evaluate whether growing at altitude—and the associated change in the dominant pollinator groups present—has an effect on the colour of flowers, we analysed data collected from the Dovrefjell National Park in Norway. Unlike previous studies, however, we considered the flower colours according to ecologically relevant models of bee and fly colour vision and also their physical spectral properties independently of any colour vision system, rather than merely looking at human colour categories. The shift from bee to fly pollination with elevation might, according to the pollination syndrome hypothesis, lead to the prediction that flower colours should shift from more bee-blue and UV-blue flowers (blue/violet to humans, i.e. colours traditionally associated with large bee pollinators) at low elevations to more bee-blue-green and green (yellow and white to humans—colours often linked to fly pollination) flowers at higher altitude. However, although there was a slight increase in bee-blue-green flowers and a decrease in bee-blue flowers with increasing elevation, there were no statistically significant effects of altitude on flower colour as seen either by bees or by flies. Although flower colour is known to be constrained by evolutionary history, in this sample we also did not find evidence that phylogeny and elevation interact to determine flower colours in alpine areas. Handling editor: Neal Williams     

Red oilseed rape? The potential for manipulation of petal colour in control strategies for the pollen beetle (Meligethes aeneus)

The pollen beetle (Meligethes aeneus) is a major pest of oilseed rape (Brassica napus) at the inflorescence stage and is well known to prefer colours called yellow by human observers over many other colours. While commercial cultivars of oilseed rape have yellow flowers, little is known about the potential to manipulate host plant location and reduce subsequent infestation by this pest through variation in flower colour. We investigated the responses of pollen beetles to flowers of a white-petalled oilseed rape variety that had been dyed different colours in semi-field arena and field experiments. Flowers dyed blue or red were less heavily infested than those dyed yellow or the white flowers, indicating that blue and red flowers were less attractive than yellow and white ones. This response was most likely due to differences in petal colour because olfactometer studies showed that beetle responses to the odours of the coloured treatments did not differ. The comparatively high infestation of untreated white flowers is interpreted as a consequence of their high UV reflectance; the presence of a UV receptor in M. aeneus is suggested, and its role in visually guided insect–plant interactions in this species described. The potential for manipulation of petal colour in control strategies for the pollen beetle is discussed.     

Frequency-dependent selection by pollinators: mechanisms and consequences with regard to behaviour of bumblebees Bombus terrestris (L.) (Hymenoptera: Apidae)

Bombus terrestris , a typical pollinating insect species, was offered artificial flowers of two different corolla colours with the same sucrose solution reward in an array. Common colours were significantly preferred, and the strength of the frequency-dependent response increased as a result of learning. There were also frequency-independent biases towards blue flowers, probably because blue flowers appeared more conspicuous to bumblebees than yellow flowers, and the degree of preference for blue was greater when flowers had low nectar rewards. Flower-to-flower movements by individual bumblebees between flowers were non-random, were biased to movements within the same flower colour, and were also dependent on morph frequency. The mechanisms governing flower selection in bumblebees are discussed. Pollinators foraging similarly in a natural situation would induce positive frequency-dependent selection, assortative mating, and directional selection on different corolla colour morphs of the plant population being visited, resulting in stabilizing selection for a single flower colour.     

The Genus Lechenaultia

Summary.  Lechenaultia is a genus of about 30 species in the largely Australian family Goodeniaceae. Most are small shrubs but some are annuals. They have short, fine leaves and terminal sprays of hand-like flowers with usually winged lobes. The range of flower colour is remarkable and includes shades of red, blue, yellow, white and green.     

Spring versus autumn leaf colours: Evidence for different selective agents and evolution in various species and floras

Red colouration is common in young and old leaves of broadleaf woody species. Assuming that leaf colours are adaptive, we examined, by comparing the colouration in young versus old leaves, the possibility that different selection agents may have operated on spring versus autumn leaf colouration. We observed spring versus autumn colouration in three very different woody floras (Finland, Japan and Israel) in order to allow for a broad ecological and evolutionary spectrum. The null hypothesis was that if the same selective agents operated in spring and autumn, it is expected that when spring leaves are red, they should always be red in autumn, and when spring leaves are green, they should be green or yellow in autumn. We found that green spring leaves are almost exclusively associated with yellow leaf colour at senescence in autumn. Species with red autumn leaves almost always have at least some red colouration in their spring leaves. However, about half of the species with red spring leaves have yellow autumn leaves. Brown autumn leaves were not common in the species we studied. As about half of the species with red spring leaves have yellow autumn leaves but not vice versa, we conclude that there are many cases in which the selecting agents for spring versus autumn leaf colour were not the same.     

Reproductive Traits and Phenology of Plants in Tropical Dry Evergreen Forest on the Coromandel Coast of India

Qualitative reproductive traits of 84 plant species belonging to 41 families were studied in tropical dry evergreen forest on the Coromandel coast of India. Majority of species had rotate type, white-coloured, scented flowers, rewarding nectar and pollen and pollinated chiefly by bees. An association between floral traits and pollination spectrum is evident. Bee pollination was prevalent in pollination systems. Among the fruit types, drupe and berry were common in black and red colour respectively, and dispersed by zoochorous mode. Seeds of brown- and green-coloured dry fruits, without any reward were disseminated by wind and explosion. The reproductive phenophase of trees and lianas occurred mostly during the dry period from January to June, which receives rainfall of less than 50 mm a month. However, shrubs showed a peak in flowering and fruiting in wet period. Detailed phenological observations of 22 woody species revealed a seasonal and unimodal pattern in flowering. Although some species were in flower round the year, flowering activity was skewed towards the dry season. The fruiting activity showed a bimodal pattern, one peak in dry season and another in wet season. Many species displayed a temporal aggregation in flowering and fruiting. The significant relation was obtained between reproductive traits and phenology of plants in the tropical dry evergreen forest.     

What colour of flowers do Lepidoptera prefer for foraging?

Food plant preferences of some Lepidoptera species associated with particular colour of the flowers were investigated. Based on 1,329 field observations of 43 Lepidoptera and 66 plant species, Lepidoptera showed a high tendency (G-test, G _adj = 698.6, df = 6, P < 0.001) to use the yellow (29%) and pink (28%) coloured flowers for foraging. Compared to the other colours it was evident that plants with red flowers (2%) were not preferred. Moreover, the plants with red (H = 0.435) and yellow-white (H = 0.543) flowers were not visited by diverse Lepidoptera species. Although yellow and pink flowers were most frequently visited, the highest degrees of the Lepidoptera diversity values were associated with the plants having blue (H = 0.647) and purple (H = 0.634) flowers. Species of Nymphalidae were most numerous (14 spp.) in the study area and the members of this family were observed 430 times on 39 different plant species, but never on plants with red flowers. Pieris rapae was the most abundant species that occurred 136 times on a total of 21 different plant species of which eight had yellow flowers. But, this species has never been seen while feeding on red flowers.     

Biological significance of distinguishing between similar colours in spectrally variable illumination: bumblebees (<Emphasis Type="Italic">Bombus terrestris</Emphasis>) as a case study

Individual bumblebees were trained to choose between rewarded target flowers and non-rewarded distractor flowers in a controlled illumination laboratory. Bees learnt to discriminate similar colours, but with smaller colour distances the frequency of errors increased. This indicates that pollen transfer might occur between flowers with similar colours, even if these colours are distinguishable. The effect of similar colours on reducing foraging accuracy of bees is evident for colour distances high above discrimination threshold, which explains previous field observations showing that bees do not exhibit complete flower constancy unless flower colour between species is distinct. Bees tested in spectrally different illumination conditions experienced a significant decrease in their ability to discriminate between similar colours. The extent to which this happens differs in different areas of colour space, which is consistent with a von Kries-type model of colour constancy. We find that it would be beneficial for plant species to have highly distinctive colour signals to overcome limitations on the bees performance in reliably judging differences between similar colours. An exception to this finding was flowers that varied in shape, in which case bees used this cue to compensate for inaccuracies of colour vision.     

Colour and size of flowers in relation to pollination of Erica species

Summary Data on flower colour polymorphism were recorded for 341 of some 426 species of Erica occurring in the south-western Cape, South Africa. Thirty-eight per cent of these Erica species are colour polymorphic, the incidence of polymorphism being greater than expected for ornithophilous species and lower than expected for anemophilous species. Both altitude and season of flowering are correlated with the incidence of colour polymorphism, with most polymorphs occurring in species which have relatively large altitudinal ranges and extended flowering periods. The mean corolla length for each of pink, purple and white flowers is significantly shorter than that for each of red, orange, yellow and green flowers, suggesting that these two sets of colours correspond with entomophily and ornithophily, respectively. There are no Erica species with blue flowers. We suggest that the patterns of colour polymorphism, because of their relationships with the behaviour of pollinators, may reflect patterns of speciation in the genus.     

Responses to olfactory and visual cues by over-wintered and summer generations of the pollen beetle, Meligethes aeneus

Abstract.  Behavioural responses to odours of oilseed rape in bud and flower stage and to green and yellow colours are studied for the two main phenological stages of pollen beetles, Meligethes aeneus , a major pest of oilseed rape, Brassica napus . Over-wintered individuals oviposit in buds of oilseed rape. Adults of the new generation (i.e. the summer generation) feed on flowers of different plant species before over wintering. In olfactometer experiments, the over-wintered beetles display a higher preference for odour of oilseed rape at the bud stage compared with the summer generation, both with and without colour stimuli. Flower odours are preferred in combination with yellow colour. Colour stimuli presented alone do not affect the behaviour. The summer generation beetles respond to both bud and flower odour. Adding colour stimuli changes the summer generations preference towards yellow and flower odour.     

The role of beetle marks and flower colour on visitation by monkey beetles (hopliini) in the greater cape floral region, South Africa

BACKGROUND AND AIMS: A deviation from the classical beetle pollination syndrome of dull-coloured flowers with an unpleasant scent is found in the Greater Cape Floral Region of South Africa. Here, monkey beetles (Scarabaeidae) visit brightly coloured, odourless flowers with conspicuous dark spots and centres (beetle marks). The role of flower colour and markings in attracting monkey beetles is still poorly understood. METHODS: Artificial model flowers with different marking patterns were used to test the effect of beetle marks on visitation by monkey beetles. To test whether monkey beetles are conditioned to the colour of the local matrix species, model flowers of different colours were placed in populations of three differently coloured species of Iridaceae. KEY RESULTS: Among all three matrix species the presence of dark markings of some kind (either centres or spots) increased visitation rates but the different matrix species differed in whether the effect was due to a dark centre or to dark spots. Monkey beetles were not conditioned for the colour of the matrix species: model colour was not significant in the Hesperantha vaginata and in the Romulea monadelpha matrices, whereas yellow model flowers were preferred over orange ones in the orange-flowered Sparaxis elegans matrix. CONCLUSIONS: This study is the first to demonstrate that beetle marks attract pollinating monkey beetles in the Greater Cape Floral Region. In contrast to plants with the classical beetle pollination syndrome that use floral scent as the most important attractant of pollinating beetles, plants with the monkey beetle pollination syndrome rely on visual signals, and, in some areas at least, monkey beetles favour flowers with dark beetle markings over unmarked flowers.     

滇西北野生观赏花卉调查

依据野外调查结果和有关文献资料,运用统计和比较的方法,对滇西北地区野生花卉的多样性进行了研究。结果表明,滇西北野生花卉有83科324属2206种。其中草本花卉1,163种,木本花卉。743种,滇西北特有野生花卉751种,珍稀濒危花卉35种。本文对滇西北地区野生花卉植物的种类、分布、观赏类型、花色、花期、受威胁状况以及特有现象进行了详细地统计分析。发现滇西北地区花卉植物的丰富度依次为丽江、中甸、贡山、德钦、维西、鹤庆、福贡、洱源、大理、兰坪。从垂直分布上看,海拔2400～3000m以及海拔3500m以上的地段花卉植物较为丰富,且特有和珍稀濒危野生花卉植物也较多地集中在这两个地段。按花色紫蓝、橙黄、红、白四大类型分类,则以紫蓝花种类最为丰富(约400种)、橙黄花次之(约230种)、红花较少(约170种)、白花最少(约140种)。     

Parallel evolution of angiosperm colour signals: common evolutionary pressures linked to hymenopteran vision

Flowering plants in Australia have been geographically isolated for more than 34 million years. In the Northern Hemisphere, previous work has revealed a close fit between the optimal discrimination capabilities of hymenopteran pollinators and the flower colours that have most frequently evolved. We collected spectral data from 111 Australian native flowers and tested signal appearance considering the colour discrimination capabilities of potentially important pollinators. The highest frequency of flower reflectance curves is consistent with data reported for the Northern Hemisphere. The subsequent mapping of Australian flower reflectances into a bee colour space reveals a very similar distribution of flower colour evolution to the Northern Hemisphere. Thus, flowering plants in Australia are likely to have independently evolved spectral signals that maximize colour discrimination by hymenoptera. Moreover, we found that the degree of variability in flower coloration for particular angiosperm species matched the range of reflectance colours that can only be discriminated by bees that have experienced differential conditioning. This observation suggests a requirement for plasticity in the nervous systems of pollinators to allow generalization of flowers of the same species while overcoming the possible presence of non-rewarding flower mimics.     

湖北星斗山地形变化对不同生活型植物叶功能性状的影响

探究地形变化对不同生活型植物叶功能性状的影响有助于深入理解森林群落物种组成的维持特征。该研究以湖北星斗山常绿落叶阔叶混交林为研究对象, 测量了50个样地中224种木本植物的叶面积、叶厚度、叶干质量、叶干物质含量和比叶面积, 运用单因素方差分析揭示了乔木、灌木和木质藤本的叶功能性状变异特征, 并采用偏曼特尔检验分别从群落水平和物种水平分析了地形变化对不同生活型木本植物叶功能性状的影响。研究发现: 不同生活型植物叶性状变异系数分布范围为23.42%-110.45%; 不同生活型之间的植物叶功能性状差异明显。群落水平上, 海拔与乔木叶干质量、灌木叶面积和木质藤本叶厚度显著正相关, 坡度仅对灌木和木质藤本比叶面积具有显著影响, 坡向与灌木叶厚度、叶干质量和比叶面积显著正相关。物种水平上, 海拔比坡度和坡向对植物叶功能性状影响更为显著, 且不同物种对地形变化的敏感度不一致; 在控制空间结构影响后, 地形因子对植物叶功能性状的影响降低。该研究结果表明, 不同生活型植物的叶功能性状对地形变化的响应格局不同, 这可能是星斗山常绿落叶阔叶混交林植物多样性的主要维持机制。     

Honeybee (Apis mellifera ligustica) Response to Differences in Handling Time, Rewards and Flower Colours

Free flying honeybees were tested outdoors on blue–white and blue–yellow dimorphic artificial flower patches to examine the influence of reward difference, flower handling‐time difference and flower colour choice on foraging decisions. We employed different flower‐well depths to vary handling times (costs), and differences in sucrose molarity to vary reward quality. Tests were performed with 2 and 6 μl rewards to vary quantity. We show that when handling time is correlated with flower‐colour morphs on a pedicellate artificial flower patch, a honeybee's foraging behaviour is dependent on the flower colours used in the choice tests. This supports a honeybee foraging model where constraints are a significant factor in decision making. Bees visiting blue–yellow flower patches exhibited flower constancy to colour, where they restricted most visits to a single flower colour, some bees to blue and others to yellow, irrespective of handing time differences. When offered a choice of equally rewarding blue or white flowers, bees were not constrained by flower colour and chose to visit flowers with a lower handling time. When reward molarity varied with well depth between blue and white flowers, foragers chose shallow‐well flowers (short‐handling time) with a smaller net harvest rate over deep‐well flowers (long‐handling time) with a greater net harvest rate. Results using the blue–white dimorphic flower patch suggest that when foraging options simultaneously involve reward and handling‐time choices, honeybee forager behaviour is inconsistent with an absolute method of evaluating profit.     

Thrips see red – flower colour and the host relationships of a polyphagous anthophilic thrips

Abstract 1. The common blossom thrips, Frankliniella schultzei, is a polyphagous anthophilic species that colonises a wide range of host‐plant species across different plant taxa. The environmental cues used by these polyphagous insects to recognise and locate host plants are not known. We therefore determined if colour is an important environmental signal used by F. schultzei to recognise flowers of eight of its more significant host‐plant species. 2. The effect of flower colour on the colonisation of different host plant species by F. schultzei was investigated by collecting and analysing the following: (a) numbers of thrips from different heights and aspects of the primary host plant Malvaviscus arboreus, (b) thrips distribution within flowers of Hibiscus rosasinensis, (c) colour reflectance from flowers of eight different host‐plant species, and (d) reflectance from different coloured sticky traps and the number of thrips trapped on them at different times of the day and on different dates. 3. The results indicate that: (a) the thrips (both sexes) concentrate towards the top of the primary host plant M. arboreus and are not distributed differentially according to sunny or shady aspect of the plant, (b) the number of female thrips on H. rosasinensis was higher in anthers compared to petals (corolla) and the basal parts of the flower, and males were as numerous on the petals as were females, and (c) there is a common floral reflectance pattern (but with different intensities) across the eight host plant species, mainly in the red part of the spectrum (600–700 nm). 4. Results of colour sticky trapping show that red attracts more female thrips compared to any other colour and that most were caught between 09.00 and 11.00 hours. By contrast, more male thrips were trapped between 07.00 and 09.00 hours. Males were more evenly distributed across the different colours but the highest numbers were associated with the yellow traps. 5. The higher densities of thrips at the top of their host plant may be related to the early morning (07.00–11.00 hours) activity of the thrips, when the top portions of the plant are more exposed to sunlight. The sex‐related distributions of F. schultzei thrips across time, coloured sticky traps, and various parts of the flowers seem to be related to mating swarm formation by the males, on the one hand, and the relative frequency and intensity of the use of M. arboreus by the females, on the other, as a feeding and oviposition site. Frankliniella schultzei females respond more strongly to red than to any other colours, so it is predicted that the spectral properties of colour recognition by this species will correlate with the predominant red reflectance of its primary host, M. arboreus, and that there may well be a sex‐related difference in colour recognition within this species.     

The significance of flower colour change in eight co-occurring shrub species

Flower colour changes from white or yellow to various shades of red at or near the sites of harvestable pollen in Calytrix glutinosa, Grevillea pilulifera, Isopogon dubius and Petrophile biloba , and over most of the flower in Hypocalymma angustifolium, Verticordia chrysantha and V. huegelii and over the pseudanthium in Darwinia citriodora. All bee, wasp, beetle, fly, butterfly and moth visitors select flowers in the white/yellow phase rather than the red or intermediate phase.
Nectar is produced by five species, harvestable pollen by four species and detectable perfume by three species, all of which features are usually absent from the red phase. The timing of the colour change in all species also corresponds to loss of stigma receptivity, completion of pollination and onset of ovule seed) swelling. Six species also undergo minor morphometric changes which discourage visitation. In all species, colour change is non-inducible by pollinators, taking 2–30 days to complete. In three protandrous species, all available pollen may be removed in the first visit, requiring transport of non-self pollen to rewardless flowers during the 10 h period of the yellow phase.
These species are highly floriferous and occur in dense patches. Since only a small proportion of flowers may be receptive at any one time, it is concluded that retention of flower parts essentially serves to enhance long-distance attraction, while colour change maximizes pollination and foraging efficiency.     

Flower colour adaptation in a mimetic orchid

Although the tremendous variability in floral colour among angiosperms is often attributed to divergent selection by pollinators, it is usually difficult to preclude the possibility that floral colour shifts were driven by non-pollinator processes. Here, we examine the adaptive significance of flower colour in Disa ferruginea, a non-rewarding orchid that is thought to attract its butterfly pollinator by mimicking the flowers of sympatric nectar-producing species. Disa ferruginea has red flowers in the western part of its range and orange flowers in the eastern part--a colour shift that we hypothesized to be the outcome of selection for resemblance to different local nectar-producing plants. Using reciprocal translocations of red and orange phenotypes as well as arrays of artificial flowers, we found that the butterfly Aeropetes tulbaghia, the only pollinator of the orchid, preferred both the red phenotype and red artificial flowers in the west where its main nectar plant also has red flowers, and both the orange phenotype and orange artificial flowers in the east, where its main nectar plant has orange flowers. This phenotype by environment interaction demonstrates that the flower colour shift in D. ferruginea is adaptive and driven by local colour preference in its pollinator.     

A Matter of Contrast: Yellow Flower Colour Constrains Style Length in Crocus species

Most flowers display distinct colour patterns comprising two different areas. The peripheral large-area component of floral colour patterns attracts flower visitors from some distance and the central small-area component guides flower visitors towards landing sites. Whereas the peripheral colour is largely variable among species, the central colour, produced mostly by anthers and pollen or pollen mimicking floral guides, is predominantly yellow and UV-absorbing. This holds also for yellow flowers that regularly display a UV bull’s eye pattern. Here we show that yellow-flowering Crocus species are a noticeable exception, since yellow-flowering Crocus species–being entirely UV-absorbing–exhibit low colour contrast between yellow reproductive organs and yellow tepals. The elongated yellow or orange-yellow style of Crocus flowers is a stamen-mimicking structure promoting cross-pollination by facilitating flower visitors’ contact with the apical stigma before the flower visitors are touching the anthers. Since Crocus species possess either yellow, violet or white tepals, the colour contrast between the stamen-mimicking style and the tepals varies among species. In this study comprising 106 Crocus species, it was tested whether the style length of Crocus flowers is dependent on the corolla colour. The results show that members of the genus Crocus with yellow tepals have evolved independently up to twelve times in the genus Crocus and that yellow-flowering Crocus species possess shorter styles as compared to violet- and white-flowering ones. The manipulation of flower visitors by anther-mimicking elongated styles in Crocus flowers is discussed.