The responses of wild jacamars (Galbula ruficauda,Galbulidae) to aposematic,aposematic and cryptic,and cryptic butterflies in central Brazil

1. This article reports the responses of wild, adult jacamars to butterflies with distinct coloration types in central Brazil. Fully aposematic species, i.e. those exhibiting bright and/or contrasting colours on both wing surfaces (= A/A), were predominantly sight‐rejected by birds and, with one exception, the few butterflies attacked and captured were taste‐rejected afterwards. 2. Aposematic and cryptic butterflies, i.e. those exhibiting bright and/or contrasting colours on the upper and cryptic colours on the underwings (= A/C) were sight‐rejected while flying, when they show their conspicuous colours to predators. This suggests that birds associate butterfly colours with their difficulty of capture, as in the case of Morpho and several Coliadinae species. These butterflies, however, were heavily attacked at rest, when they are cryptic. 3, Fully cryptic butterflies, i.e. those exhibiting cryptic colours on both wing surfaces (= C/C) did not elicit sight rejections by birds. Comparisons involving the number of attacks and the capture success of flying and resting individuals showed no significant differences in species more frequently observed like some cracker butterflies (Hamadryas feronia and H. februa) and Taygetis laches. Compared with the A/C Coliadinae, these butterflies showed a lesser, although not significantly different, ability to escape while flying, but a greater and significantly different ability to escape while at rest. 4, A hunting tactic of jacamars, which consists of following flying A/C and C/C butterflies on sight, and waiting until they perch to locate and attack them, is described for the first time.     

Testosterone is correlated with courtship but not aggression in the tropical buff-banded rail,Gallirallus philippensis

Past studies into the roles of testosterone in birds have focused on species that occur in temperate regions. In such species, plasma testosterone levels are high during the establishment of territories in the spring and are associated with increased aggression. In contrast to most temperate species, tropical birds frequently defend territories year-round, during which time territoriality often occurs in a nonsexual context. The few studies that have been carried out on tropical birds show lower levels of circulating testosterone than occur in their temperate counterparts. In some year-round territorial tropical species, testosterone and aggression are dissociated, while in other species testosterone still plays a role in regulating aggression. This study examined the relationship between aggression and plasma testosterone levels in a year-round territorial, subtropical population of the buff-banded rail with characteristics typical of tropical species. Peak testosterone levels were substantially lower than those found in temperate species. Males displayed a seasonal peak in plasma testosterone level when their partners were most likely to be fertile. At other times, testosterone levels were mostly undetectable, despite year-round territoriality. We found that T levels increased with courtship behavior but showed no relationship with aggression, supporting the hypothesis that dissociation between testosterone and territoriality may be widespread among tropical avian taxa.     

Sexual and seasonal differences in the frequency of beak marks on the wings of two Lethe butterflies

To assess bird predation pressure on butterflies, I investigated beak marks on the wings of two Lethe butterflies for 3 years in secondary temperate forests. If bird predation had significant effects on average longevity of butterflies, and if the number of specimens preyed upon was proportionate to the number of beak-marked specimens, the beak mark frequency would be negatively correlated with average longevity of a butterfly. Bird predation pressure is generally thought to influence average longevity of butterflies. Therefore, if there is a negative correlation between beak mark frequency and average longevity, bird predation pressure would be reflected in beak mark frequency. Beak mark frequency was negatively correlated with longevity in Lethe diana (Butler), the more abundant of the two species; thus, the beak mark frequency was considered to be a suitable index of bird predation pressure on the butterflies investigated in this study. In both Lethe species, beak mark frequency was higher in females than in males. Because female butterflies have a relatively smaller thorax and flight muscles and a larger abdomen that contains eggs, they are presumably weaker or less agile fliers than males, and are probably attacked more easily by birds. In autumn, butterflies were heavily attacked by birds irrespective of sex and species. Because the numbers of lepidopteran larvae, which are the preferred prey of many birds, decreased in autumn, birds were thought to shift their diets to alternative prey such as adult butterflies.     

Predation,thermoregulation, and wing color in pierid butterflies

Summary This paper explores two hypotheses about the relationships among predation, thermoregulation, and wing color in butterflies: First, that butterflies are susceptible to predation during thermally marginal periods (e.g., cool weather) when effective thermoregulation and flight are not possible; second, that Pieris butterflies are relatively unpalatable to visual predators, supporting the idea that the white wing pigment of Pieris represents aposematic coloration. Field experiments with Pieris and Colias in 1984 and 1985 demonstrate that substantial predation may occur during the morning period before butterflies are able to actively fly. Circumstantial evidence is presented to suggest that at least some of the predation is by small, cursorial mammals. Feeding experiments in the field using Grey Jays as predators indicate that Pieris napi and P. occidentalis are less palatable than other sympatric butterflies, including confamial Colias alexandra. These and previous results suggest that Pieris are edible but less preferred as prey by birds, and that the degree of palatibility may vary among Pieris species. The relatively low palatability of these Pieris is consistent with the hypothesis that their white pigmentation represents aposematic coloration; however, the cues by which potential bird predators might discriminate against Pieris have not been established.     

Revealing the colourful side of birds: spatial distribution of conspicuous plumage colours on the body of Australian birds

In many species of birds, different body parts often display very different colours. This spatial distribution of coloured plumage patches may be determined, among other factors, by the balance between being cryptic to predators, and conspicuous to intended receivers. If this is the case, ventral and anterior body parts in birds – which are less visible to predators but more prominent to conspecifics – should present more conspicuous and sexually dichromatic plumage colours. Here, I test these predictions using reflectance spectrometric measurements of standardised plumage patches across males and females for nearly an entire avifauna (Australian landbirds, n = 538 species). My data show that, as predicted, conspicuous and sexually dichromatic colours are mainly located near the head, while the plumage of the back is the most cryptic. One clear exception to this pattern is the conspicuous rump coloration. In many species, this patch can be concealed by wings, and therefore exposed only when necessary. In addition, conspicuous rump coloration could deflect or confuse predators in case of attack. However, there is considerable variation across species, and this makes position on the body a very poor predictor of plumage elaboration (R² < 0.02). Future studies should try to determine whether differences between species in the distribution of colours across the plumage are due to variation in ecological factors (predation risk, habitat, etc.).     

Avoidance of an aposematically coloured butterfly by wild birds in a tropical forest

1. Birds are considered to be the primary selective agents for warning colouration in butterflies, and select for aposematic mimicry by learning to avoid brightly coloured prey after unpleasant experiences. It has long been thought that bright colouration plays an important role in promoting the avoidance of distasteful prey by birds. 2. The hypothesis that warning colouration facilitates memorability and promotes predator avoidance was tested by means of a field experiment using distasteful model butterflies. Artificial butterflies with a Heliconius colour pattern unknown to local birds were generated using bird vision models, either coloured or achromatic, and hung in tree branches in a tropical forest. Two sequential trials were conducted at each site to test avoidance by naïve and experienced predators. 3. There was a significant reduction in predation in the second trial. Also, coloured models were attacked less than achromatic models. Specifically, coloured butterflies were attacked significantly less in the second trial, but there was no significant decrease in predation on achromatic models. 4. The present results imply an important role for colour in enhancing aversion of aposematic butterflies. It has also been demonstrated that previous experience of distasteful prey can lead to enhanced avoidance in subsequent trials, supporting mimicry theory.     

The benefit of being a social butterfly: communal roosting deters predation

Aposematic passion-vine butterflies from the genus Heliconius form communal roosts on a nightly basis. This behaviour has been hypothesized to be beneficial in terms of information sharing and/or anti-predator defence. To better understand the adaptive value of communal roosting, we tested these two hypotheses in field studies. The information-sharing hypothesis was addressed by examining following behaviour of butterflies departing from natural roosts. We found no evidence of roost mates following one another to resources, thus providing no support for this hypothesis. The anti-predator defence hypothesis was tested using avian-indiscriminable Heliconius erato models placed singly and in aggregations at field sites. A significantly higher number of predation attempts were observed on solitary models versus aggregations of models. This relationship between aggregation size and attack rate suggests that communally roosting butterflies enjoy the benefits of both overall decreased attack frequency as well as a prey dilution effect. Communal roosts probably deter predators through collective aposematism in which aggregations of conspicuous, unpalatable prey communicate a more effective repel signal to predators. On the basis of our results, we propose that predation by birds is a key selective pressure maintaining Heliconius communal roosting behaviour.     

Do bright colors at nests incur a cost due to predation?

Summary Birds show much interspecific variation in the coloration and brightness of their plumage. I examine the hypothesis that selection due to predation on incubating birds and their nest contents can explain part of this diversity. First, I argue that rather than using absolute rates of nest predation to make predictions about the costs of conspicuous colours, we should measure experimentally whether increases in plumage conspicuousness elevate rates of nest predation. Second, I present experimental data investigating the cost of red and brown colour at ground and tree nests. These data provide the first evidence that bright colours do attract predators to nests and that, in addition, this cost varies according to the nesting site. Natural selection seems to most strongly oppose the evolution of conspicuous colours in ground-nesting birds.     

Can redistribution of breeding colonies on a landscape mitigate changing predation danger?

The reproductive success of colonially breeding species depends in part upon a trade‐off between the benefit of a dilution effect against nestling predation within larger colonies and colony conspicuousness. However, there may be no net survivorship benefit of dilution if smaller colonies are sufficiently inconspicuous. This raises the question about how the size distribution of breeding colonies on a landscape might change as the predation danger for nestlings changes. In southwest British Columbia, Canada, bald eagle Haliaeetus leucocephalus populations have increased exponentially at ～5% per year in recent decades and prey upon nestlings of colonial breeding great blue herons Ardea herodias faninni. Motivated by field data on reproductive success in relation to colony size, modeling is used to ask under which circumstances trading off a dilution benefit against colony conspicuousness can improve population reproductive success. That is, which colonial nesting distribution, dispersed and cryptic versus clumped and conspicuous, best mitigates predation danger on nestlings? When predators are territorial, the modeling predicts a dispersed nesting strategy as attack rate increases, but not as predator numbers increase. When predators are non‐territorial, the modeling predicts a dispersed nesting strategy as predator numbers and/or attack rates increase. When predators are both territorial and non‐territorial, colonial nesting within a predator's territory improves reproductive success when attack rates are low. This suggests nesting in association with territorial predators may offer decreased levels of predation when compared with nesting amongst non‐territorial predators. Thus a change in the colony size distribution of colonially breeding species might be anticipated on a landscape experiencing a change in predation danger.     

Nest predation and the evolution of nestling begging calls

Begging by nestling birds can be conspicuous and loud. Such displays are thought to function in signalling nestling condition and securing parental care, but they also may inadvertently attract the attention of predators. We compared the structure of nestling begging calls to the risk of predation among 24 species of birds breeding in a forest community in central Arizona. After controlling for body size and phylogeny, we found that species subject to greater nest predation had calls with higher frequency (pitch) and lower amplitude (loudness) than species subject to lower rates of nest predation. As these acoustic features make it difficult for potential predators to pinpoint the source of a sound, our results suggest that an increased risk of predation has led to the evolution of begging calls that minimize locatability. The relationship between call structure and the risk of predation also supports the hypothesis that attracting predators is a direct cost of begging and that such costs can constrain any evolutionary escalation in the intensity of nestling begging.     

Predator mixes and the conspicuousness of aposematic signals

Conspicuous warning signals of unprofitable prey are a defense against visually hunting predators. They work because predators learn to associate unprofitability with bright coloration and because strong signals are detectable and memorable. However, many species that can be considered defended are not very conspicuous; they have weak warning signals. This phenomenon has previously been ignored in models and experiments. In addition, there is significant within- and among-species variation among predators in their search behavior, in their visual, cognitive, and learning abilities, and in their resistance to defenses. In this article we explore the effects of variable predators on models that combine positive frequency-dependent, frequency-independent, and negative frequency-dependent predation and show that weak signaling of aposematic species can evolve if predators vary in their tendency to attack defended prey.     

Beak marks on the wings of butterflies and predation pressure in the field

Beak marks on butterfly wings have been used as an indicator of predation pressure. The relationship between butterflies and their predators in the field was examined to evaluate the beak mark rate as an indicator of predation pressure. Transect censuses were conducted to measure the beak mark rate on butterflies from May to November, 2010, in Shiga Prefecture, Japan. A total of 1216 butterflies of 42 species from five families were observed during the study period. The beak mark rate in the Papilionidae was significantly higher than those of all other families. Analysis of the monthly fluctuations in the beak mark rate and relative abundance of predators revealed that the higher beak mark rates observed for two butterfly families (Papilionidae and Nymphalidae) were significantly and positively related to predation pressure, while no significant relationship was found for other families (Lycaenidae, Pieridae and Hesperiidae). Beak marks in larger butterflies (Papilionidae and Nymphalidae) can be used as an indicator to evaluate the relative intensity of predation pressure in the field.     

Are Blackcaps (Sylvia atricapilla) defending their nests also calling for help from their neighbours?

One hypothesized function of conspicuous mobbing of intruders by bird nest owners is to attract neighbouring birds (“calling for help” hypothesis) or third-party predators (“attract the mightier” hypothesis). These may help the nest owners by distracting and/or attacking the mobbed intruder. To date, these hypotheses have been studied solely during the mobbing of predators. Here, for the first time, I have studied mobbing attraction in the context of brood parasitism. I experimentally tested the Blackcap (Sylvia atricapilla), a small passerine with a highly aggressive and conspicuous nest defence behaviour. I elicited the aggressive responses of Blackcaps by presenting stuffed dummies of the brood parasitic Common Cuckoo (Cuculus canorus) and controls near their nests. At 32% of the nests (n = 75), the responses of the Blackcaps to dummies attracted up to 15 birds per trial from 21 passerine species. Most of the attracted birds were heterospecifics and rarely participated in mobbing; thus the “calling for help” hypothesis was not supported. No potential predators of the Cuckoo were attracted despite them living in the study area and despite prolonged mobbing by Blackcaps; thus rejecting the “attract the mightier” hypothesis. I argue that this hypothesis is unlikely to apply to typical avian predators during nest predation acts because these only last for several seconds. The number of attracted birds was a positive function of the owner’s intensity of nest defence as measured by the rates of alarm calling but not visual cues (rates of attacks). Suitable and unsuitable Cuckoo hosts did not differ in their behaviour in the vicinity of defended nests. The observed pattern of the positive correlation between the intensity of nest defence and the number of attracted birds is most likely a proximate by-product of the conspicuous nest defence by Blackcaps (but may well be adaptive for recruiting neighbours themselves). Thus, the mobbing behaviour of the Blackcap is directed towards the brood parasite and not towards other audience predators or potential recruits to the mob.     

Why some rails have white tails: the evolution of white undertail plumage and anti-predator signaling

Conspicuous plumage patches have evolved in birds as conspecific signals for mate attraction and assessment, intersexual competition or to signal alarm. Signals may alternatively be directed at potential predators to discourage pursuit. Rails (Family Rallidae) are ground-dwelling birds, many of which inhabit wetlands, while others occur in forests and grasslands. They are renown for their secretive nature and the tendency to flick their tails when observed. This behavior is more conspicuous in species with white undertail coverts that contrast sharply with darker body plumage. Using species comparisons and controlling for phylogeny, we investigated four hypotheses for the evolution of white undertail coverts in rails. We found little support for the hypothesis that white tails are sexually selected: white tails were not more common in species with polygamous as opposed to monogamous mating systems, species with sexual dimorphism, nor species that display their tails in courtship. Nor did our results support the hypothesis that white tail plumage evolved for intersexual competition during territorial interactions. Instead, we found that species that flock for at least part of the year and species found in open as opposed to concealing habitats were significantly more likely to have white undertail coverts. Rail species inhabiting concealing habitats are less commonly gregarious and more likely selected for crypsis. Using phylogenetically-controlled statistical inference we found that adaptation to open wetland habitats significantly precedes the evolution of white undertails, whereas gregariousness likely evolved later in some lineages. The inferred order of trait evolution suggests that this plumage characteristic could have been selected primarily for enhancement of an anti-predator signal rather than a social signal for conspecifics.     

Cryptic differences in colour among Müllerian mimics: how can the visual capacities of predators and prey shape the evolution of wing colours?

Antagonistic interactions between predators and prey often lead to co‐evolution. In the case of toxic prey, aposematic colours act as warning signals for predators and play a protective role. Evolutionary convergence in colour patterns among toxic prey evolves due to positive density‐dependent selection and the benefits of mutual resemblance in spreading the mortality cost of educating predators over a larger prey assemblage. Comimetic species evolve highly similar colour patterns, but such convergence may interfere with intraspecific signalling and recognition in the prey community, especially for species involved in polymorphic mimicry. Using spectrophotometry measures, we investigated the variation in wing coloration among comimetic butterflies from distantly related lineages. We focused on seven morphs of the polymorphic species Heliconius numata and the seven corresponding comimetic species from the genus Melinaea. Significant differences in the yellow, orange and black patches of the wing were detected between genera. Perceptions of these cryptic differences by bird and butterfly observers were then estimated using models of animal vision based on physiological data. Our results showed that the most strikingly perceived differences were obtained for the contrast of yellow against a black background. The capacity to discriminate between comimetic genera based on this colour contrast was also evaluated to be higher for butterflies than for birds, suggesting that this variation in colour, likely undetectable to birds, might be used by butterflies for distinguishing mating partners without losing the benefits of mimicry. The evolution of wing colour in mimetic butterflies might thus be shaped by the opposite selective pressures exerted by predation and species recognition.     

A comparative study of territoriality in intertidal and subtidal blennioids (Teleostei, Blennioidei)

This paper presents a comparative analysis of territoriality in three intertidal (Lipophrys pholis, Coryphoblennius galerita, Salaria pavo) and two subtidal (Tripterygion delaisi, Parablennius pilicornis) blennioid fishes. Focal-animal observations of males guarding eggs showed that: (i) intertidal species had smaller territories that were less frequently patrolled; (ii) in subtidal species feeding was limited to the territory, while in intertidal fishes a substantial proportion of the feeding acts occurred outside the defended area; (iii) intertidal species spent less time out of the nest and showed a lower level of locomotory activity; (iv) subtidal species were subjected to a higher number of territorial intrusions with more species intruding their territory, some of them potential egg predators; (v) subtidal species performed several water column displays that played a significant role in courtship, while in intertidal fishes these displays are almost absent. Signalling without the presence of a female was almost absent in intertidal species. It is argued that intertidal fishes minimize the time spent out of the nest and the loss of contact with the substrate, features that may be of high survival value in conditions of strong turbulence. Although intertidal species have a reduced time available for feeding, this may be compensated by lower levels of locomotory activity, territorial defense and risk of egg predation. This revised version was published online in August 2006 with corrections to the Cover Date.     

The conceptual and practical implications of interpreting diet breadth mechanistically in generalist predatory insects

Seasonal polyphenism in animal colour patterns indicates that temporal variation in selection pressures maintains phenotypic plasticity. Spring generation of the polyphenic European map butterfly Araschnia levana has an orange–black fritillary‐like pattern whilst individuals of the summer generation are black with white bands across the wings. What selects for the colour difference is unknown. Because predation is a major selection pressure for insect coloration, we first tested whether map butterfly coloration could have a warning function (i.e. whether the butterflies are unpalatable to birds). In a following field experiment with butterfly dummies we tested whether the spring form is better protected than the summer form from predators in the spring, and vice versa in the summer. The butterflies were palatable to birds (blue tits Cyanistes caeruleus) and in the field the spring and summer form dummies were attacked equally irrespective of season. Therefore, we found no evidence that the map butterfly is warning‐coloured or that seasonal polyphenism is an adaptation to avian predation. Because insect coloration has multiple functions and map butterfly coloration is linked to morphology, life history and development it is likely that the interplay of several selection pressures explains the evolution of colour polyphenism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Agonistic display or courtship behavior? A review of contests over mating opportunity in butterflies

Male butterflies compete over mating opportunities. Two types of contest behavior are reported. Males of various butterfly species compete over a mating territory via aerial interactions until one of the two contestants retreats. Males of other butterfly species fly around larval food plants to find receptive females. Males of some species among the latter type can find a conspecific pupa, and they gather around it without expelling their rivals. Scramble competition over mating occurs when a female emerges from the pupa. Many studies have been performed on territorial species, and their contest resolution has often been understood from the point of view of contest models based on game theory. However, these models cannot explain why these butterflies perform contest displays despite the fact that they do not have the ability to attack their opponent. A recent study based on Lloyd Morgan’s Canon showed that territorial contests of male butterflies are better understood as erroneous courtship between sexually active males. In this paper, I review research on contests over mating opportunity in butterflies, and show that the erroneous courtship framework can explain not only territorial contests of butterflies but also why males do not determine the owner of a conspecific pupa.     

Response to Predation Risk in Urban and Rural House Sparrows

Habitat urbanization may change the density of predators, and it is often assumed that such changes lead to altered predation risk for urban populations of their prey. Although it is difficult to study predation hazard directly, behavior responses of prey species may be informative in inferring such habitat differences. In this study, we compared the risk‐taking behavior of urban and rural house sparrows (Passer domesticus) after simulated attacks by two of their important predators (sparrowhawk Accipiter nisus and domestic cat Felis catus). The birds were startled by moving dummies of these predators and respective control objects, and their risk taking was estimated as their latency to feed after the startle. We found that sparrows responded more strongly (had longer post‐startle feeding latencies) to sparrowhawk attacks than to the control object, and their responses differed between the habitats. First, risk taking of urban birds strongly decreased with age (older birds had longer latencies than young birds), while there was no such age difference in rural birds. Second, young urban birds responded less strongly, while older urban birds responded more strongly to the sparrowhawk than the same age groups of rural birds, respectively. We did not succeed in evoking antipredatory response by simulated cat attacks, because birds responded similarly to the dummy and the control object. Our results support that predation risk, posed at least by avian predators, is different in urban and rural habitats of house sparrows. The increased wariness of older, hence presumably more experienced, urban birds implies that sparrows may be more exposed to predation in cities.     

Manipulations of the AVT system shift social status and related courtship and aggressive behavior in the bluehead wrasse

Arginine vasotocin (AVT) and its mammalian homologoue arginine vasopressin (AVP) influence male sexual and aggressive behaviors in many species. We tested the effects of AVT and an AVP-V(1a) receptor antagonist on the display of alternative male tactics in a tropical coral reef fish, the bluehead wrasse Thalassoma bifasciatum. We gave AVT injections to territorial and nonterritorial males of the large and colorful phenotype (terminal phase) and an AVP-V(1a) receptor antagonist, Manning compound, to territorial males in the field. AVT increased courtship independent of status, while its effects on territoriality and aggression were dependent upon male status. In territorial males, AVT increased courtship and tended to decrease the number of chases toward initial phase individuals. In nonterritorial males, AVT increased courtship, chases toward initial phase individuals, and territorial behavior while decreasing feeding. These are all behaviors rarely seen in nonterritorial males, so AVT made these males act like territorial TP males. The AVP-V(1a) receptor antagonist had opposite effects. It decreased courtship and territorial defense, making these males act more like nonterritorial males. Manipulations of the AVT system shifted males within a single phenotype from the nonterritorial social status to the territorial social status and vice versa. Since the entire suite of behaviors related to territoriality was affected by AVT system manipulations, our results suggest that the AVT system may play a key role in motivation of behaviors related to mating.