Crypsis in a heterogeneous environment: relationships between changeable polymorphic colour patterns and behaviour in a galaxiid fish

1. The ability to achieve optimal camouflage varies between microhabitats in heterogeneous environments, potentially restricting individuals to a single habitat or imposing a compromise on crypsis to match several habitats. However, animals may exhibit morphological and behavioural attributes that enhance crypsis in different habitats. 2. We used an undescribed fish species, Galaxias‘nebula’, to investigate two objectives. First, we examined two potential methods of enhancing crypsis: change in colour pattern and selection of a suitable background. Second, we characterised the colour pattern of this unstudied fish and assessed its capacity for crypsis. 3. No background selection was apparent but the area of dark pigment expressed varied between backgrounds, which may negate the requirement to be choosy about habitats. The capacity to change colour and selection of a background that maximises crypsis are most likely separate, non‐mutually exclusive strategies. 4. Galaxias‘nebula’ exhibits polymorphic, non‐interchangeable colour patterns that have elements of both background pattern matching and disruptive colouration. This, coupled with habitat characteristics, suggests a combination of generalist and specialist strategies of habitat use. The fish’s camouflage strategy and air‐breathing ability may be key to survival under increasing pressure from habitat degradation and invasive predators.     

Extra-pair paternity and sexual dimorphism in birds

Differences in the strength of sexual selection between males and females can lead to sexual dimorphism. Extra-pair paternity (EPP) can increase the variance in male reproductive success and hence the opportunity for sexual selection. Previous research on birds suggests that EPP drives the evolution of dimorphism in plumage colour and in body size. Because EPP increases the intensity of sexual selection in males, it should lead to increased dimorphism in species with larger or more colourful males, but decreased dimorphism in species with larger or more colourful females. We explored the covariation between EPP and sexual dimorphism in wing length and plumage colouration in 401 bird species, while controlling for other, potentially confounding variables. Wing length dimorphism was associated positively with the frequency of EPP, but also with social polygamy, sex bias in parental behaviour and body size and negatively with migration distance. The frequency of EPP was the only predictor of plumage colour dimorphism. In support of our prediction, high EPP levels were associated with sexual dichromatism, positively in species in which males are more colourful and negatively in those in which females are more colourful. Contrary to our prediction, high EPP rates were associated with increased wing length dimorphism in species with both male- and female-biased dimorphism. The results support a role for EPP in the evolution of both size and plumage colour dimorphism. The two forms of dimorphism were weakly correlated and predicted by different reproductive, social and life-history traits, suggesting an independent evolution.     

Geographic variation for colour in the sandburrowing beetle Chaerodes trachyscelides White (Goleoptera: Tenebrionidae) on New Zealand beaches analysed using CIELAB L * values

Chaerodes trachyscelides White is a highly specialized, flightless burrowing beetle confined to the narrow strip of sand at and just above high water level on sandy marine beaches in New Zealand. Although the ventral surface of the beetle is always pale, the dorsal surface varies from pale to almost black. Large samples of this beetle were taken, together with the sand, from 11 beaches on New Zealand's three main islands. The colour of the dorsal surface of each individual beetle and that of the sand samples was measured using reflectance spectroscopy and expressed as CIE L*,a*,b* (CIELAB) values. The L* values, which are objective, quantitative measures of the degree of lightness of the beetles, were subjected to statistical and frequency analysis. Although the species was very variable in colour and the variation appeared to be continuous, a highly significant correlation was obtained between the mean of the L* values for the samples on each beach and that of the sand, the correlation coefficient being 0.961. This close association between the lightness of the beetles and that of the sand suggests the variable melanism functions as cryptic colouration. On most beaches, the distribution oflightness among the beetles sampled conformed to a normal curve. For beetles from sites where the sand was relatively uniform, such as the black Taranaki beaches, the L* frequency distribution curves were narrow and the coefficient of variation of mean beetle colour was relatively small indicating low colour variability. In contrast, the greatest within-site variability occurred on the two Stewart Island beaches sampled, where in each case there was less uniformity in the colour of the sand. At one of these sites, Maori Beach, darker sand present below the high water level is often deposited on the zone occupied by the beetles after storms. On Lonneker's Beach, the distribution of L* values among the beetles sampled was actually bimodal. On this small beach, there was an area of intensely black sand in the zone occupied by the beetles, but most of the rest was covered with light golden sand. These results are interpreted as evidence that the variability of colour of Chaerodes beetles has the effect of populations being able to match the colour of the sand of their home beaches, presumably as a consequence of the differential survival of individuals.     

Innate colour preferences of flower visitors

Freshly emerged flower visitors exhibit colour preferences prior to individual experience with flowers. The understanding of innate colour preferences in flower visitors requires a detailed analysis, as, on the one hand, colour is a multiple-signal stimulus, and, on the other hand, flower visits include a sequence of behavioural reactions each of which can be driven by a preferential behaviour. Behavioural reactions, such as the distant approach, the close-range orientation, the landing, and the extension of mouthparts can be triggered by colour stimuli. The physiological limitations of spectral sensitivity, the neuro-sensory filters, and the animals' different abilities to make use of visual information such as brightness perception, wavelength-specific behaviour and colour vision shape colour preferences. Besides these receiverbased factors, there are restrictions of flower colouration due to sender-based factors such as the absorption properties of floral pigments and the dual function of flower colours triggering both innate and learned behaviour. Recordings of the spectral reflection of coloured objects, which trigger innate colour preferences, provide an objective measure of the colour stimuli. Weighting the spectral reflection of coloured objects by the spectral composition of the ambient light and the spectral sensitivity of the flower visitors' photoreceptors allows the calculation of the effective stimuli. Perceptual dimensions are known for only a few taxa of flower visitors.     

Stabilizing survival selection on presenescent expression of a sexual ornament followed by a terminal decline

Senescence is a decrease in functional capacity, increasing mortality rate with age. Sexual signals indicate functional capacity, because costs of ornamentation ensure signal honesty, and are therefore expected to senesce, tracking physiological deterioration and mortality. For sexual traits, mixed associations with age and positive associations with life expectancy have been reported. However, whether these associations are caused by selective disappearance and/or within‐individual senescence of sexual signals, respectively, is not known. We previously reported that zebra finches with redder bills had greater life expectancy, based on a single bill colour measurement per individual. We here extend this analysis using longitudinal data and show that this finding is attributable to terminal declines in bill redness in the year before death, with no detectable change in presenescent redness. Additionally, there was a quadratic relationship between presenescent bill colouration and survival: individuals with intermediate bill redness have maximum survival prospects. This may reflect that redder individuals overinvest in colouration and/or associated physiological changes, while below‐average bill redness probably reflects poorer phenotypic quality. Together, this pattern suggests that bill colouration is defended against physiological deterioration, because of mate attraction benefits, or that physiological deterioration is not a gradual process, but accelerates sharply prior to death. We discuss these possibilities in the context of the reliability theory of ageing and sexual selection.     

Do stabilimenta in orb webs attract prey or defend spiders?

Orb-weaving spiders are ideal organisms for the study of conflictbetween behavioral investments in foraging and defense becausetheir webs provide physical manifestations of those investments.We examined the impact of including stabilimenta, designs ofbright-white noncapture silk, at the center of orb webs forforaging and defense in Argiope aurantia. Our findings suggestthat stabilimentum building is a defensive behavior, supportingthe "web advertisement" hypothesis that the high visibilityof stabilimenta can prevent birds from flying through webs.Yet, spiders often do not include stabilimenta in their webs,indicating that a serious cost is associated with them. We alsoshow, through comparison of paired webs with and without stabilimenta,that stabilimenta reduce the prey capture success of spidersby almost 30%. This demonstrates the potential impact that defensivebehaviors of spiders can have on their foraging success andsuggests that much of the variation in stabilimenta may be accountedfor by a cost—benefit trade-off made when including stabilimentain webs.     

Global biogeography of warning coloration in the butterfly Danaus chrysippus

Warning coloration provides a textbook example of natural selection, but the frequent observation of polymorphism in aposematic species presents an evolutionary puzzle. We investigated biogeography and polymorphism of warning patterns in the widespread butterfly Danaus chrysippus using records from citizen science (n = 5467), museums (n = 8864) and fieldwork (n = 2586). We find that polymorphism in three traits controlled by known mendelian loci is extensive. Broad allele frequency clines, hundreds of kilometres wide, suggest a balance between long-range dispersal and predation of unfamiliar morphs. Mismatched clines for the white hindwing and forewing tip in East Africa are consistent with a previous finding that the black wingtip allele has spread recently in the region through hitchhiking with a heritable endosymbiont. Light/dark background coloration shows more extensive polymorphism. The darker genotype is more common in cooler regions, possibly reflecting a trade-off between thermoregulation and predator warning. Overall, our findings show how studying local adaptation at the global scale provides a more complete picture of the evolutionary forces involved.     

Potential aposematism in an insular tree species: are signals dishonest early in ontogeny?

Recent investigations have suggested that some plants are aposematic. Our understanding of how aposematism varies through plant ontogeny, however, is incomplete. Furthermore, the potential for lower leaf surfaces to signal to vertebrate herbivores that are viewing leaves from below has not been investigated. Here, we investigate ontogenetic changes in leaf colour in Pseudopanax crassifolius (Araliaceae), a tree species that is endemic to New Zealand. We demonstrate that P. crassifolius produces lateral leaf spines that peak in size during the sapling stage of development. Spots of brightly coloured tissues on the upper leaf surfaces may be warning signals. The intensity of these signals, however, peaked at the seedling stage, providing a dishonest signal of defence. Conversely, signals on lower leaf surfaces peaked in the sapling stage, providing an honest defensive signal later in ontogeny. Lateral leaf spines and all potential warning colours were absent in adults, after they grow above the reach of the largest known native megaherbivores (moa – Aves: Dinornithiformes). Overall, these results suggest that aposematism may vary predictably through plant ontogeny in response to the changing perspective of herbivores as plants grow vertically.     

Discriminant analysis of Lepidopteran prey characteristics and their effects on the outcome of bird-feeding trials

Discriminant analysis was used to analyse the results of 348 bird-feeding trials conducted from 1982 to 1985 in Leverett, Massachusetts. Four size classes, seven appearance categories, and five larval host types, based on 163 species of moths and butterflies used as prey in two or more trials, were selected as predictor variables to discriminate between prey taken and not taken by birds. Discriminant analysis of individual feeding trials correctly classified 97.5 percent of prey taken or not-taken and ranked the predictor variables according to their relative importance in determining prey acceptability. Characteristics most acceptable to birds were: (1) large size, (2) bark-like appearance, (3) warning colouration, (4) woody generalist, (5) dead-leaf-like appearance, (6) woody specialist, and (7) medium size. Characteristics least acceptable to birds were: (1) small size, (2) mimetic appearance, (3) butterfly appearance, (4) herbaceous specialist food type, (5) black-and-white appearance, (6) extra large size, and (7) overall generalist feeder. A summary of the analyses includes a discriminant function based on lepidopteran characteristics that can be used to predict the prey acceptability of species not used in this study. A multiple regression analysis of prey taken revealed that size alone and larval host type combined with other prey characteristics were the most important variables in determining the selection of prey regardless of their abundance in the trials.