Colour preferences influences odour learning in the hawkmoth, Macroglossum stellatarum

The hummingbird hawkmoth, Macroglossum stellatarum, learns colour fast and reliably. It has earlier been shown to spontaneously feed from odourless artificial flowers. Now, we have studied odour learning. The moths were trained to discriminate feeders of the same colour but marked with different odours. They did not learn to discriminate two natural flower odours when they were presented with the innately preferred colour blue, but they did learn this discrimination combined with yellow or green colours that are less attractive to the moth. The yellow colour could be trained to become as attractive as the innately preferred blue colour and the blue colour could be trained to become less attractive. This is the first proof of odour learning in a diurnal moth. The results show that M. stellatarum can use more than one modality in their foraging behaviour and that the system is plastic. By manipulating the preferences for the different colours, their influence on odour learning could be changed.     

Retinal regionalization and heterogeneity of butterfly eyes

The regional characteristics of the eyes of butterflies from different families have been surveyed using epi-illumination microscopy, utilizing the eyeshine visible due to the tapetum situated proximally to the rhabdom. All butterflies studied have a high spatial acuity in the frontal region. The facet diameter varies slightly across the eye, and the interommatidial angle and the eye parameter p are especially large dorsally. Whereas the ommatidial lattice is generally highly regular, the eyeshine colours distinctly depend on the species. Sometimes the eyeshine is locally uniform, but often it is heterogeneous. It is hypothesized that the regional characteristics as well as the local heterogeneity are adaptations that optimize spectral discrimination.     

Hydrocarbon synthesis by enzymatically dissociated oenocytes of the abdominal integument of the German Cockroach,<Emphasis Type="Italic"> Blattella germanica</Emphasis>

In insects, hydrocarbons waterproof the cuticle, protect the insect from the external environment, and serve as semiochemicals or their metabolic precursors. In the German cockroach, Blattella germanica, hydrocarbons are synthesized by the abdominal integument, but the precise site of biosynthesis is not known. We developed a method for separation of oenocytes from other cells in the abdominal integument using enzymatic dissociation followed by Percoll gradient centrifugation. Radiolabeled propionate was then used to monitor de novo synthesis of hydrocarbons by dissociated cells. Oenocyte-enriched cell suspensions of abdominal sternites synthesized hydrocarbons, whereas suspensions enriched with epidermal cells did not. Our results show conclusively that hydrocarbons are produced by oenocytes not only in insects whose oenocytes are localized within the hemocoel, but also in those insects whose oenocytes are within the abdominal integument. Furthermore, these data support a hemolymph pathway for transport and delivery of hydrocarbons to both external and internal tissues, including the epicuticle, fat body, and ovaries.     

A comparative analysis of colour preferences in temperate and tropical social bees

The spontaneous occurrence of colour preferences without learning has been demonstrated in several insect species; however, the underlying mechanisms are still not understood. Here, we use a comparative approach to investigate spontaneous and learned colour preferences in foraging bees of two tropical and one temperate species. We hypothesised that tropical bees utilise different sets of plants and therefore might differ in their spontaneous colour preferences. We tested colour-naive bees and foragers from colonies that had been enclosed in large flight cages for a long time. Bees were shortly trained with triplets of neutral, UV-grey stimuli placed randomly at eight locations on a black training disk to induce foraging motivation. During unrewarded tests, the bees’ responses to eight colours were video-recorded. Bees explored all colours and displayed an overall preference for colours dominated by long or short wavelengths, rather than a single colour stimulus. Naive Apis cerana and Bombus terrestris showed similar choices. Both inspected long-wavelength stimuli more than short-wavelength stimuli, whilst responses of the tropical stingless bee Tetragonula iridipennis differed, suggesting that resource partitioning could be a determinant of spontaneous colour preferences. Reward on an unsaturated yellow colour shifted the bees’ preference curves as predicted, which is in line with previous findings that brief colour experience overrides the expression of spontaneous preferences. We conclude that rather than determining foraging behaviour in inflexible ways, spontaneous colour preferences vary depending on experimental settings and reflect potential biases in mechanisms of learning and decision-making in pollinating insects.     

Fine colour discrimination requires differential conditioning in bumblebees

Accurate recognition requires that visual systems must be able to discriminate between target and distractor stimuli. Flowers are learned and recognised by bees using visual cues including colour and shape. We investigated whether bees were able to learn to discriminate between colours differently depending upon absolute or differential conditioning. For absolute conditioning bees were rewarded with sucrose solution for visits to target flowers. When distractor stimuli were subsequently presented, a high level of discrimination was observed if there was a perceptually large colour distance separating distractors and targets, but for a perceptually small colour distance the bees generalised and did not discriminate between stimuli. When provided with differential conditioning where both target and distractors were present, the bees learnt to discriminate stimuli separated by a perceptually small colour distance. This shows that for bees to learn fine colour discrimination tasks it is important to use differential conditioning. The findings are discussed within the context of the necessity for plants to produce distinctive flower colours.     

Colour mimicry and sexual deception by Tongue orchids (Cryptostylis)

Typically, floral colour attracts pollinators by advertising rewards such as nectar, but how does colour function when pollinators are deceived, unrewarded, and may even suffer fitness costs? Sexually deceptive orchids are pollinated only by male insects fooled into mating with orchid flowers and inadvertently transferring orchid pollinia. Over long distances, sexually deceptive orchids lure pollinators with counterfeit insect sex pheromones, but close-range deception with colour mimicry is a tantalising possibility. Here, for the first time, we analyse the colours of four sexually deceptive Cryptostylis orchid species and the female wasp they mimic (Lissopimpla excelsa, Ichneumonidae), from the perspective of the orchids’ single, shared pollinator, male Lissopimpla excelsa. Despite appearing different to humans, the colours of the orchids and female wasps were effectively identical when mapped into a hymenopteran hexagonal colour space. The orchids and wasps reflected predominantly red-orange wavelengths, but UV was also reflected by raised bumps on two orchid species and by female wasp wings. The orchids’ bright yellow pollinia contrasted significantly with their overall red colour. Orchid deception may therefore involve accurate and species-specific mimicry of wavelengths reflected by female wasps, and potentially, exploitation of insects’ innate attraction to UV and yellow wavelengths. In general, mimicry may be facilitated by exploiting visual vulnerabilities and evolve more readily at the peripheries of sensory perception. Many sexually deceptive orchids are predominantly red, green or white: colours that are all potentially difficult for hymenoptera to detect or distinguish from the background.     

Does contrast between eggshell ground and spot coloration affect egg rejection?

Obligate avian brood parasitic species impose the costs of incubating foreign eggs and raising young upon their unrelated hosts. The most common host defence is the rejection of parasitic eggs from the nest. Both egg colours and spot patterns influence egg rejection decisions in many host species, yet no studies have explicitly examined the role of variation in spot coloration. We studied the American robin Turdus migratorius, a blue-green unspotted egg-laying host of the brown-headed cowbird Molothrus ater, a brood parasite that lays non-mimetic spotted eggs. We examined host responses to model eggs with variable spot coloration against a constant robin-mimetic ground colour to identify patterns of rejection associated with perceived contrast between spot and ground colours. By using avian visual modelling, we found that robins were more likely to reject eggs whose spots had greater chromatic (hue) but not achromatic (brightness) contrast. Therefore, egg rejection decision rules in the American robin may depend on the colour contrast between parasite eggshell spot and host ground coloration. Our study also suggests that egg recognition in relation to spot coloration, like ground colour recognition, is tuned to the natural variation of avian eggshell spot colours but not to unnatural spot colours.     

A colony-level response to disease control in a leaf-cutting ant

Parasites and pathogens often impose significant costs on their hosts. This is particularly true for social organisms, where the genetic structure of groups and the accumulation of contaminated waste facilitate disease transmission. In response, hosts have evolved many mechanisms of defence against parasites. Here we present evidence that Atta colombica, a leaf-cutting ant, may combat Escovopsis, a dangerous parasite of Atta's garden fungus, through a colony-level behavioural response. In A. colombica, garden waste is removed from within the colony and transported to the midden - an external waste dump - where it is processed by a group of midden workers. We found that colonies infected with Escovopsis have higher numbers of workers on the midden, where Escovopsis is deposited. Further, midden workers are highly effective in dispersing newly deposited waste away from the dumping site. Thus, the colony-level task allocation strategies of the Atta superorganism may change in response to the threat of disease to a third, essential party.     

Ancient specimens and DNA contamination: a case study from the 12S rRNA gene sequence of the "Linh Duong" bovid (Pseudonovibos spiralis)

In 1993, several unusual horn sheaths, collected in the South of Vietnam, were regarded as evidence for a new mammal species, Pseudonovibos spiralis. However, the taxonomic status of P. spiralis remains a highly controversial subject: firstly, it has been related to three different groups of the family Bovidae: Antilopini (gazelles), Bovini (oxen, bisons, and buffaloes), and Caprini sensu lato (goats, sheep and allies). Secondly, certain horns have been shown to be faked. Most recently, it has been suggested, on the basis of 12S rRNA gene sequences, that P. spiralis was a new species of buffalo. I demonstrate here that this conclusion was inaccurate, and base this on the grounds that: (1) the putative sequence of P. spiralis is shown to be a chimera obtained from three different species: Bos taurus, Bubalus bubalis and Saiga tatarica, and (2) several factors indicate that the specimen used was not authentic. This new study confirms earlier suggestions that the horns used to formally describe P. spiralis were bogus and were in fact derived from the horns of domesticated cattle (Bos taurus). The data suggest that P. spiralis never existed.     

Floral colour change in <Emphasis Type="Italic">Byrsonima variabilis</Emphasis> (Malpighiaceae) as a visual cue for pollen but not oil foraging by oil-collecting bees

Pollinators search for multiple flora resources throughout their life cycle. Most studies, however, only assess how bees discriminate floral cues in the context of nectar foraging. In the present study, we sought to elucidate whether oil-collecting bees discriminate flowers of Byrsonima variabilis (Malpighiaceae) with petals of different colours when foraging for pollen or oil. As the colour of the standard petal changes during anthesis, we characterised the spectral reflectance patterns of flowers throughout anthesis and modelled chromatic perceptual space to determine how these colour patterns are perceived by bees. Through the quantification of flower pollen in the different phases, we found that the colour of the standard petal is an honest cue of the presence of pollen. Centridine bees preferentially visited flowers with a yellow (bee’s green) colour when searching for pollen, but indiscriminately visited flowers with different petal colours when searching for floral oil. We suggest that standard petals, in the species studied and others of the genus, like nectar guides, act as pollen guides, which oil-collecting females use to detect pollen-rich flowers. Moreover, they use different floral clues during foraging for different resources in the same host plant.     

Molecular phylogeny of cycads inferred from rbcL sequences

The chloroplast gene rbcL was sequenced to elucidate the evolution of the gymnosperm plant order Cycadales. In accordance with traditional systematics, the order Cycadales and the corresponding genera cluster as monophyletic clades. Among them, the genus Cycas forms a basal group. The genetic distances within the genus Encephalartos and between the sister groups Encephalartos, Lepidozamia and Macrozamia, are unexpectedly small, suggesting that the extant species are the result of Miocene and Pliocene speciation. Their distribution in Africa or Australia, respectively, may therefore rather be due to long-distance dispersal than to Cretaceous continental drift, as had previously been assumed. The rbcL sequences also indicate that the colonisation of Madagascar by Cycas thouarsii occurred only recently as the sequences of C. thouarsii and Cycas rumphii from Indonesia are identical. In contrast, the divergence of the Cycadaceae and Zamiaceae apparently occurred in the Mesozoic.     

Assortative mating by carotenoid-based plumage colour: a quality indicator in American goldfinches,<Emphasis Type="Italic"> Carduelis tristis</Emphasis>

We investigated the mating patterns in 22 breeding pairs of wild American goldfinches (Carduelis tristis) with respect to their body size, condition, and carotenoid-based, yellow plumage colour. Using reflectance spectrometry, we objectively quantified plumage colours across the bird-visible wavelengths, revealing a unexpected UV peak in the reflectance spectrum from yellow feathers. We summarized our colour measurements using a principal components analysis to create a single variable, carotenoid PC1, that represents the intensity of this carotenoid-based yellow colour, a measure of phenotypic quality in this species. We found no evidence of assortative mating with respect to measures of body size or condition but there was positive assortative mating by carotenoid PC1, such that the yellow plumage colours of males and females were significantly correlated within pairs. We argue that the yellow carotenoid coloration of goldfinches may be important in mutual mate choice and, thus, that sexual selection in this species may act upon female ornamentation, as well as the more obvious plumage signals of males. Because assortative mating results in an increase in genetic variance, we suggest that this might be a mechanism that maintains variance in ornamental traits in spite of the variance-eroding effects of sexual selection.     

Split-second escape decisions in blue tits (Parus caeruleus)

Bird mortality is heavily affected by birds of prey. Under attack, take-off is crucial for survival and even minor mistakes in initial escape response can have devastating consequences. Birds may respond differently depending on the character of the predator's attack and these split-second decisions were studied using a model merlin (Falco columbarius) that attacked feeding blue tits (Parus caeruleus) from two different attack angles in two different speeds. When attacked from a low attack angle they took off more steeply than when attacked from a high angle. This is the first study to show that escape behaviour also depends on predator attack speed. The blue tits responded to a high-speed attack by dodging sideways more often than when attacked at a low speed. Escape speed was not significantly affected by the different treatments. Although they have only a split-second before escaping an attack, blue tits do adjust their escape strategy to the prevailing attack conditions.     

Plumage coloration and nutritional condition in the great tit<Emphasis Type="Italic"> Parus major</Emphasis>: the roles of carotenoids and melanins differ

The size and coloration of some body characters seem to influence mate choice in many species. Most animal colours are either structural or based on melanin or carotenoid pigments. It has recently been suggested that carotenoid-based or structural coloration may be a condition-dependent trait, whereas melanin-based coloration is not; a difference that may be highly relevant when studying the evolution of multiple mating preferences. We tested this hypothesis in the great tit (Parus major). The size of the melanin breast band was not correlated to nutritional condition as estimated by the rate of tail growth (ptilochronology), controlling for locality, age, sex, year and season effects. However, the correlation was significant for the hue of yellow breast (carotenoid-based coloration), and the slopes of the regressions of the two pigments to growth bars differed significantly. These results suggest that the expression of the two traits may be regulated by different mechanisms.     

A non-policing honey bee colony (Apis mellifera capensis)

In the Cape honey bee Apis mellifera capensis, workers lay female eggs without mating by thelytokous parthenogenesis. As a result, workers are as related to worker-laid eggs as they are to queen-laid eggs and therefore worker policing is expected to be lower, or even absent. This was tested by transferring worker- and queen-laid eggs into three queenright A. m. capensis discriminator colonies and monitoring their removal. Our results show that worker policing is variable in A. m. capensis and that in one colony worker-laid eggs were not removed. This is the first report of a non-policing queenright honey bee colony. DNA microsatellite and morphometric analysis suggests that the racial composition of the three discriminator colonies was different. The variation in policing rates could be explained by differences in degrees of hybridisation between A. m. capensis and A. m. scutellata, although a larger survey is needed to confirm this.     

Cryptic mammalian species: a new species of whiskered bat (Myotis alcathoe n. sp.) in Europe

The analysis of morphological, behavioural and genetic characters of whiskered bats revealed a new European bat species within the family Vespertilionidae. We describe the morphology, karyology, genetic similarity, ecology and distribution of Myotis alcathoe n. sp. It closely resembles Myotis mystacinus, Myotis brandtii and Myotis ikonnikovi in morphology, but all four species show clear genetic differences in two mitochondrial genes (ND1 and 12S rRNA). Myotis alcathoe n. sp. is the smallest species among the European whiskered bats and uses the highest-frequency echolocation calls of all the European Myotis species. It prefers to hunt in small valleys with deciduous trees and flowing water, which is an endangered habitat. Records from Greece and Hungary indicate a distribution range in south-eastern Europe.     

Alcoholism in cockchafers: orientation of male Melolontha melolontha towards green leaf alcohols

Chemical orientation of the European cockchafer, Melolontha melolontha L., a serious pest in agriculture and horticulture, was investigated by field tests and electrophysiological experiments using plant volatiles. In total, 16 typical plant volatiles were shown to elicit electrophysiological responses in male cockchafers. Funnel trap field bioassays revealed that green leaf alcohols (i.e. (Z)-3-hexen-1-ol, (E)-2-hexen-1-ol and 1-hexanol) attracted males, whereas the corresponding aldehydes and acetates were behaviourally inactive. Furthermore, male cockchafers were attracted by volatiles from mechanically damaged leaves of Fagus sylvatica L., Quercus robur L. and Carpinus betulus L. However, volatiles emitted by damaged leaves of F. sylvatica attracted significantly more males than those from the other host plants. Odour from intact F. sylvatica leaves was not attractive to M. melolontha males. Females were not attracted by any of the tested volatile sources. The results suggest that plant volatiles play a similar role as a sexual kairomone in mate finding of M. melolontha, as has been shown for the forest cockchafer, Melolontha hippocastani F. Nevertheless, both species show remarkable differences in their reaction to green leaf alcohols.     

Feminization of genetic males by a symbiotic bacterium in a butterfly,Eurema hecabe (Lepidoptera: Pieridae)

Wolbachia are symbiotic bacteria found in many arthropods and filarian nematodes. They often manipulate the reproduction of host arthropods. In the present study, female-biased sex-ratio distortion in the butterfly Eurema hecabe was investigated. Breeding experiments showed that this distorted sex ratio is maternally inherited. When treated with tetracycline, adult females of the thelygenic line produced male progeny only. After PCR using Wolbachia-specific primers for the ftsZ gene a positive result was seen in the thelygenic females, but not in male progeny from tetracycline-treated females, or individuals from a Tokyo population with normal sex ratio and reproduction. Cytological observations showed that thelygenic females lack the sex chromatin body (W chromosome). The results strongly suggest that the sex-ratio distortion in E. hecabe is due to feminization of genetic males by Wolbachia.