Cryptically Patterned Moths Perceive Bark Structure When Choosing Body Orientations That Match Wing Color Pattern to the Bark Pattern

Many moths have wing patterns that resemble bark of trees on which they rest. The wing patterns help moths to become camouflaged and to avoid predation because the moths are able to assume specific body orientations that produce a very good match between the pattern on the bark and the pattern on the wings. Furthermore, after landing on a bark moths are able to perceive stimuli that correlate with their crypticity and are able to re-position their bodies to new more cryptic locations and body orientations. However, the proximate mechanisms, i.e. how a moth finds an appropriate resting position and orientation, are poorly studied. Here, we used a geometrid moth Jankowskia fuscaria to examine i) whether a choice of resting orientation by moths depends on the properties of natural background, and ii) what sensory cues moths use. We studied moths’ behavior on natural (a tree log) and artificial backgrounds, each of which was designed to mimic one of the hypothetical cues that moths may perceive on a tree trunk (visual pattern, directional furrow structure, and curvature). We found that moths mainly used structural cues from the background when choosing their resting position and orientation. Our findings highlight the possibility that moths use information from one type of sensory modality (structure of furrows is probably detected through tactile channel) to achieve crypticity in another sensory modality (visual). This study extends our knowledge of how behavior, sensory systems and morphology of animals interact to produce crypsis.     

Threatened tadpoles of Bokermannohyla alvarengai (Anura: Hylidae) choose backgrounds that enhance crypsis potential

Crypsis results from a complex interaction among prey coloration, background matching, behaviour and predator visual perception. Tadpoles are known to have varied adaptations to escape predation, but the use of crypsis is little explored, although it is likely for certain species. We investigated potential escape mechanisms related to active escape (fleeing) and crypsis improvement in Bokermannohyla alvarengai tadpoles, proposing a new method to measure cryptic potential. We studied the range of distances covered by threatened and fleeing tadpoles and the proportion of tadpoles that seek shelter or remain exposed after fleeing. We hypothesized that tadpoles that remain exposed may use alternative strategies to avoid detection, such as reaching deeper microhabitats or positioning themselves on substrates that confer greater crypsis than the ones they were on before disturbance. A significantly greater proportion of tadpoles remained exposed after disturbance and positioned themselves on backgrounds that offered greater cryptic potential, but did not differ in depth. Tadpoles may respond to a trade‐off between sheltering and being cryptic. On the one hand, they may remain close to retreat sites or they may escape to microhabitats that provide appropriate background matching as a means to achieve crypsis. On the other hand, the absence of matching backgrounds in the tadpoles' vicinities may induce them to seek shelter. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 437–446.     

Behaviourally mediated crypsis in two nocturnal moths with contrasting appearance

The natural resting orientations of several species of nocturnal moth on tree trunks were recorded over a three-month period in eastern Ontario, Canada. Moths from certain genera exhibited resting orientation distributions that differed significantly from random, whereas others did not. In particular, Catocala spp. collectively tended to orient vertically, whereas subfamily Larentiinae representatives showed a variety of orientations that did not differ significantly from random. To understand why different moth species adopted different orientations, we presented human subjects with a computer-based detection task of finding and ‘attacking’ Catocala cerogama and Euphyia intermediata target images at different orientations when superimposed on images of sugar maple (Acer saccharum) trees. For both C. cerogama and E. intermediata, orientation had a significant effect on survivorship, although the effect was more pronounced in C. cerogama. When the tree background images were flipped horizontally the optimal orientation changed accordingly, indicating that the detection rates were dependent on the interaction between certain directional appearance features of the moth and its background. Collectively, our results suggest that the contrasting wing patterns of the moths are involved in background matching, and that the moths are able to improve their crypsis through appropriate behavioural orientation.     

Moths use multimodal sensory information to adopt adaptive resting orientations

Camouflage conceals animals from predators and depends on the interplay between the morphology and behaviour of animals. Behavioural elements of animals, such as the choice of a resting spot or posture, are important for effective camouflage, as well as the animals’ cryptic appearance. To date, the type of sensory input that mediates resting site choice remains poorly understood. Previously, we showed that bark‐like moths perceive and rely on bark structure to seek out cryptic resting positions and body orientations on tree trunks. In the present study, we investigated the sensory organs through which moths perceive the structure of bark when positioning their bodies in adaptive resting orientations. We amputated (or blocked) each one of the hypothetical sensory organs in moths (antennae, forelegs, wings, and eyes) and tested whether they were still able to perceive bark structure properly and adopt adaptive resting orientations. We found that visual information or stimulation is crucial for adaptively orienting their bodies when resting and tactile information from wings may play an additional role. The present study reveals multimodal information use by moths to achieve visual camouflage and highlights the sensory mechanism that is responsible for the adaptive behaviour of cryptic insects. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 900–904.     

Resting orientation enhances prey survival on strongly structured background

Prey can use various camouflage types as defense against predators. One of the most common and important types is background matching, which occurs if an animal matches the background in color, brightness, and pattern. Although background matching has been studied intensively, the effects of the resting orientation of prey on the effectiveness of camouflage through background matching are not well known in natural conditions. Several past experimental studies have been conducted on resting orientation in the lab often using the visual system of humans. Their results revealed that the detection rates of predators hinge on the combination of the resting orientation of artificial moths and their background. Here, we studied whether survival rates of artificial moth-like models depend on their resting orientation in the wild where the visual conditions and detection distances vary. We used a 2 × 2 design of two resting positions of a horizontally and a vertically striped morph on tree bark. Our results show that the survival probability of moths depended mainly on the orientation of stripes relative to the vertical structure of tree bark. Thus, resting orientation relative to background affected survival. After reanalyzing Endler’s (Biol J Linn Soc 22:187–231, 1984) data on resting habitats of 317 species of North American moths, we found that horizontally striped moths occurred frequently on small herbs and tree bark. We suggest that it would be beneficial for striped moths to orient non-randomly on strongly structured background, like furrows of tree bark. We further suggest that background matching was more important than coincident disruptive coloration in determining the survival rates of our artificial moths.     

Progressive background in moths, and a quantitative measure of crypsis

A method is presented for quantitative estimation of the degree of crypsis of species seen by visual predators against known backgrounds. It is based upon a comparison between transects taken across animal and background colour patterns. The method was applied to day-resting moths in deciduous forest in New Jersey. Each species is found for two to four weeks at characteristic dates, and there is a constant turnover of species. In both moths and backgrounds there is a regular change in the colour pattern parameters from winter through spring to early summer. Moths are on average more cryptic at their normal dates than they would be if present earlier or later in the year. Species with known resting sites are on average more cryptic on their resting sites than other background habitats. Species that rest on more than one background habitat are less cryptic on their preferred habitats than are specialists. Species that rest under leaves and are not visible from above are not very cryptic. Specific v. general resemblance, disruptive coloration, and factors affecting 'aspect diversity' are discussed. The new method of estimating crypsis is useful for studies of crypsis as well as in sexual selection. It is necessary to know much about the resting sites and behaviour of moths, as well as other functions of colour patterns, to understand colour pattern evolution.     

Active background choice facilitates crypsis in a tropical crab

Animals can evade predators in multiple ways, one of the most effective of which is to avoid detection in the first place. We know much about the evolution of color patterns that match the visual background to avoid detection (i.e., crypsis), yet we know surprisingly less about the specific behaviors that have co‐evolved with these morphological traits to enhance or maintain crypsis. We here explore whether the match between body color and background in a seemingly well‐camouflaged tropical shore crab is a result of active background choice. Taking advantage of a coastal area in the Solomon Islands with variable sand color and a population of the pallid ghost crab Ocypode pallidula with varying carapace color, we experimentally tested whether individuals actively choose specific substrate that best matches their color patterns. We found that individuals taken from extreme sand colors chose substrate that maintained crypsis, with relatively darker crabs typically choosing dark sand and lighter crabs choosing light sand. Crabs of intermediate color pattern, in contrast, showed no clear preference for dark or light sand. Our results suggest that potential prey can actively choose specific backgrounds to enhance and maintain crypsis, providing insights into how behavior interacts with morphological traits to avoid predator detection.     

Background matching ability and the maintenance of a colour polymorphism in the red devil cichlid

The evolution and maintenance of colour polymorphisms remains a topic of considerable research interest. One key mechanism thought to contribute to the coexistence of different colour morphs is a bias in how conspicuous they are to visual predators. Although individuals of many species camouflage themselves against their background to avoid predation, differently coloured individuals within a species may vary in their capacity to do so. However, to date, very few studies have explicitly investigated the ability of different colour morphs to plastically adjust their colouration to match their background. The red devil (Amphilophus labiatus) is a Neotropical cichlid fish with a stable colour polymorphism, with the gold morph being genetically dominant and having a myriad of documented advantages over the dark morph. However, gold individuals are much rarer, which may be related to their heightened conspicuousness to would‐be predators. Here, we tested the ability of differently coloured individuals to phenotypically adjust the shade of their body colour and patterns to match their background. In particular, we filmed dark, gold and mottled (a transitioning phase from dark to gold) individuals under an identical set‐up on light vs. dark‐coloured substrates. We found that, in contrast to individuals of the dark morph, gold and mottled individuals were less capable of matching their body colouration to their background. As a result, gold individuals appeared to be more conspicuous. These results suggest that a difference in background matching ability could play an important role in the maintenance of colour polymorphisms.     

Spots and stripes: ecology and colour pattern evolution in butterflyfishes

The incredible diversity of colour patterns in coral reef fishes has intrigued biologists for centuries. Yet, despite the many proposed explanations for this diversity in coloration, definitive tests of the role of ecological factors in shaping the evolution of particular colour pattern traits are absent. Patterns such as spots and eyespots (spots surrounded by concentric rings of contrasting colour) have often been assumed to function for predator defence by mimicking predators'' enemies'' eyes, deflecting attacks or intimidating predators, but the evolutionary processes underlying these functions have never been addressed. Striped body patterns have been suggested to serve for both social communication and predator defence, but the impact of ecological constraints remains unclear. We conducted the first comparative analysis of colour pattern diversity in butterflyfishes (Family: Chaetodontidae), fishes with conspicuous spots, eyespots and wide variation in coloration. Using a dated molecular phylogeny of 95 species (approx. 75% of the family), we tested whether spots and eyespots have evolved characteristics that are consistent with their proposed defensive function and whether the presence of spots and body stripes is linked with species'' body length, dietary complexity, habitat diversity or social behaviour. Contrary to our expectations, spots and eyespots appeared relatively recently in butterflyfish evolution and are highly evolutionarily labile, suggesting that they are unlikely to have played an important part in the evolutionary history of the group. Striped body patterns showed correlated evolution with a number of ecological factors including habitat type, sociality and dietary complexity. Our findings question the prevailing view that eyespots are an evolutionary response to predation pressure, providing a valuable counter example to the role of these markings as revealed in other taxa.     

Fine nanostructural variation in the wing pattern of a moth <Emphasis Type="Italic">Chiasmia eleonora</Emphasis> Cramer (1780)

Butterflies and moths possess diverse patterns on their wings. Butterflies employ miscellaneous colour in the wings whereas moths use a combination of dull colours like white, grey, brown and black for the patterning of their wings. The exception is some of the toxic diurnal moths which possess bright wing colouration. Moths possess an obscure pattern in the dorsal part of the wings which may be a line, zigzag or swirl. Such patterns help in camouflage during resting period. Thus, the dorsal wing pattern of the moth is used for both intra- as well as inter-specific signal communication. Chiasmia eleonora is a nocturnal moth of greyish black colouration. The dorsal hindwing possesses yellow and black colour patches. A white-coloured oblique line crosses both left and right fore- and hindwings to form a V-shaped pattern across the dorsal wing. This V-shaped pattern possesses a UV signal. Closer to the body, the colour appears darker, which fades towards the margin. The fine nanostructural variation is observed throughout the wings. This study elucidates the wing pattern of the geometrid moth C. eleonora using high-resolution microscopy techniques that has not been described in previous studies.     

Threshold determination for olfactory receptors of the spiny lobster

The shallow water caridean shrimps Heptacarpus pictus and H. paludicola are polymorphic in colour pattern. Populations of these species collected over colour‐variable substrates showed the greatest degree of coloration in terms of the proportion of individuals displaying a distinct colour pattern. The frequencies of H. Pictus colour morphs varied significantly between most sampling periods. Apostatic selection by fish predators is suggested as one hypothesis which could explain these changes in morph frequency.

Experiments with Heptacarpus pictus on rapid colour change showed that, although some pigment migration did occur in the chromatosomes studied, the macroscopic appearance of the colour patterns was not altered when shrimps were shifted from black to white backgrounds or vice versa. The results of background choice experiments with H. pictus suggest that these shrimp do not seek out colour backgrounds that would seem to be a matching background in a concealing coloration. This behaviour is consistent with the morphology of the colour patterns which appears to be a disruptive coloration rather than a concealing coloration that closely matches a particular substrate.     

A new species of spiny driftwood catfish Spinipterus (Siluriformes: Auchenipteridae) from the Amazon basin

An expedition to the middle Rio Purus basin uncovered a remarkable new species of the genus Spinipterus. The new species has a very distinct and conspicuous colour pattern resembling a jaguar and it is almost four times larger than Spinipterus acsi, a small specimen (32 mm L_S) from Caño Santa Rita, a right bank tributary of Río Nanay in Peru and a second specimen was reported from Rio Juruá, Amazonas State, Brazil. Although the new species is more similar in size and colour pattern to Liosomadoras, it shares the synapomorphies for Spinipterus. The new species differs from the congener by the following characters: (a) colour pattern with large black rosette-like spots over a light yellow to brown background (v. brown background with small dark blotches over the body); (b) adult body size reaching 104.5 mm L_S (v. maximum known size 37.1 mm L_S); (c) posterior process of cleithrum short, never reaching vertical through the dorsal-fin origin (v. posterior process long, surpassing vertical through the dorsal-fin origin); (d) seven soft pectoral-fin rays (v. six); (e) caudal fin truncated (v. caudal fin rounded).     

Coral reef flounders,Bothus lunatus,choose substrates on which they can achieve camouflage with their limited body pattern repertoire

Camouflage is a common tactic to avoid detection or recognition by predators and prey. Flounders have adaptive camouflage but a limited body pattern repertoire. We tested whether peacock flounders actively select or avoid certain substrates to more effectively use their limited camouflaging ability. We acquired and analyzed ten 30‐min videos of individual flounders on a coral reef in Bonaire, Dutch Caribbean. Using Manly's beta resource selection indices, we were able to confirm that peacock flounders at this location preferred to settle on neutral‐coloured substrates, such as sand and dead coral. Moreover, they avoided live coral, cyanobacteria, and sponges, which are often brightly coloured (e.g. yellow, orange, and purple). Quantitative analyses of photographs of settled flounders indicate that they use uniform and mottled camouflage patterns, and that the small‐to‐moderate spatial scale of their physiologically controlled light and dark skin components limits their camouflage capabilities to substrates with similar colour and spatial frequencies. These fishes changed their body pattern very fast. We did not observe disruptive body patterns, which are generally characterized by large‐scale skin components and higher contrast. The results suggest that flounders are using visual information to actively choose substrates on which they can achieve general background resemblance. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 629–638.     

Adaptive body patterning,three‐dimensional skin morphology and camouflage measures of the slender filefish Monacanthus tuckeri on a Caribbean coral reef

The slender filefish is a master of adaptive camouflage and can change its appearance within 1–3 s. Videos and photographs of this animal's cryptic body patterning and behavior were collected in situ under natural light on a Caribbean coral reef. We present an ethogram of body patterning components that includes large‐ and small‐scale spots, stripes and bars that confer a variety of cryptic patterns amidst a range of complex backgrounds. Field images were analyzed to investigate two aspects of camouflage effectiveness: (1) the degree of colour resemblance between animals and their nearby visual stimuli; and (2) the visibility of each fish's actual body outline vs. its illusory outline. Most animals more closely matched the colour of nearby visual stimuli than that of the surrounding background. Three‐dimensional dermal flaps complement the melanophore skin patterns by enhancing the complexity of the fish's physical skin texture to disguise its actual body shape, and the morphology of these structures was studied. The results suggest that the body patterns, skin texture, postures and swimming orientations putatively hinder both the detection and recognition of the fish by potential visual predators. Overall, the rapid speed of change of multiple patterns, colour blending with nearby backgrounds, and the physically complicated edge produced by dermal flaps effectively camouflage this animal among soft corals and macroalgae in the Caribbean Sea.     

Variable colour patterns indicate multidimensional,intraspecific trait variation and ecological generalization in moths

Animal colour patterns long have provided information about key processes that drive the ecological and evolutionary dynamics of biological diversity. Theory and empirical evidence indicate that variation in colour patterns and other traits among individuals generally improves the performance of populations and species, for example by reducing predation risk, increasing establishment success, improving resilience to environmental change, and decreasing risk of extinction. However, little is known about whether and how variation in colour pattern among species is associated with variation in other phenotypic dimensions. To address this issue, we analysed associations of colour pattern with morphological, behavioural and life-history traits on the basis of data for nearly 400 species of noctuid moths. We found that moths with more variable colour patterns had longer flight activity periods, more diverse habitats and a greater number of host plant species than species with less variable colour patterns. Variable coloration in adult noctuid moths therefore can be considered as indicative of broader niches and generalist diets. Colour pattern variability was not significantly associated with overwintering stage or body size (wing span), and it was independent of whether the colour pattern of the larvae was non-variable, variable or highly variable. Colour pattern variation during the larval stage tended to increase as the duration of the flight activity period increased, but was independent of the length of the larval period, diet breadth and habitat use. The realization that information on colour pattern variation in adult moths, and possibly other organisms, offers a proxy for niche breadth and dietary generalization can inform management and conservation biology.     

Bergmann's rule does not apply to geometrid moths along an elevational gradient in an Andean montane rain forest

Aim Bergmann's rule generally predicts larger animal body sizes with colder climates. We tested whether Bergmann's rule at the interspecific level applies to moths (Lepidoptera: Geometridae) along an extended elevational gradient in the Ecuadorian Andes. Location Moths were sampled at 22 sites in the province Zamora‐Chinchipe in southern Ecuador in forest habitats ranging from 1040 m to 2677 m above sea level. Methods Wingspans of 2282 male geometrid moths representing 953 species were measured and analysed at the level of the family Geometridae, as well as for the subfamily Ennominae with the tribes Boarmiini and Ourapterygini, and the subfamily Larentiinae with the genera Eois, Eupithecia and Psaliodes. Results Bergmann's rule was not supported since the average wingspan of geometrid moths was negatively correlated with altitude (r = ?0.59, P < 0.005). The relationship between body size and altitude in Geometridae appears to be spurious because species of the subfamily Larentiinae are significantly smaller than species of the subfamily Ennominae and simultaneously increase in their proportion along the gradient. A significant decrease of wingspan was also found in the ennomine tribe Ourapterygini, but no consistent body size patterns were found in the other six taxa studied. In most taxa, body size variation increases with altitude, suggesting that factors acting to constrain body size might be weaker at high elevations. Main conclusions The results are in accordance with previous studies that could not detect consistent body size patterns in insects at the interspecific level along climatic gradients.     

Geographical and morphological variation within and between colour phases in Coris julis (L. 1758), a protogynous marine fish

The possible differences between sexes in patterns of morphological variation in geographical space have been explored only in gonochorist freshwater species. We explored patterns of body shape variation in geographical space in a marine sequential hermaphrodite species, Coris julis (L. 1758), analyzing variation both within and between colour phases, through the use of geometric morphometrics and spatially‐explicit statistical analyses. We also tested for the association of body shape with two environmental variables: temperature and chlorophyll a concentration, as obtained from time‐series of satellite‐derived data. Both colour phases showed a significant morphological variation in geographical space and patterns of variation divergent between phases. Although the morphological variation was qualitatively similar, individuals in the initial colour phase showed a more marked variation than individuals in the terminal phase. Body shape showed a weak but significant correlation with environmental variables, which was more pronounced in primary specimens. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 148–162.     

Mutually honest? Physiological ‘qualities’ signalled by colour ornaments in monomorphic king penguins

Mate choice is expected to be important for the fitness of both sexes for species in which successful reproduction relies strongly on shared and substantial parental investment by males and females. Reciprocal selection may then favour the evolution of morphological signals providing mutual information on the condition/quality of tentative partners. However, because males and females often have differing physiological constraints, it is unclear which proximate physiological pathways guarantee the honesty of male and female signals in similarly ornamented species. We used the monomorphic king penguin (Aptenodytes patagonicus) as a model to investigate the physiological qualities signalled by colour and morphological ornaments known to be under sexual selection (coloration of the beak spots and size of auricular feather patches). In both sexes of this slow‐breeding seabird, we investigated the links between ornaments and multiple indices of individual quality; including body condition, immunity, stress and energy status. In both sexes, individual innate immunity, resting metabolic rate, and the ability to mount a stress response in answer to an acute disturbance (capture) were similarly signalled by various aspects of beak coloration or auricular patch size. However, we also reveal interesting and contrasting relationships between males and females in how ornaments may signal individual quality. Body condition and oxidative stress status were signalled by beak coloration, although in opposite directions for the sexes. Over an exhaustive set of physiological variables, several suggestive patterns indicated the conveyance of honest information about mate quality in this monomorphic species. However, sex‐specific patterns suggested that monomorphic ornaments may signal different information concerning body mass and oxidative balance of males and females, at least in king penguins.     

PHYLOGENY AND BIOGEOGRAPHY OF THE ATLANTIC AND EASTERN PACIFIC HYPSELODORIS STIMPSON, 1855 (NUDIBRANCHIA, CHROMODORIDIDAE) WITH THE DESCRIPTION OF A NEW SPECIES FROM THE CARIBBEAN SEA

A new species, based on two specimens from Guana Island, BritishVirgin Islands, is described. The new species agrees with thegenus Hypselodoris in having a high body profile, a large vestibulargland and mantle glands. It differs from other members of thegenus in the Atlantic Ocean by having a reddish background bodycolour. In addition, dorsal colour patterns such as a broadcentral white line with lateral extensions and the lack of yellowlines or spots further differentiate this species. The radularformula of 52x41.0.41 and a smaller seminal receptacle are alsodistinctive. The phylogenetic relationships of 34 species andsubspecies of Hypselodoris from the eastern Pacific and Atlanticare examined using morphological characters. With the exceptionof the new species, these are characterized by a dark blue backgroundbody colour. The phylogenetic analysis of the data matrix resultedin eight most-parsimonious trees. The resulting consensus treeshows that eastern Pacific and Atlantic species of Hypselodorisconstitute a monophyletic group that is basally split into twosister clades. One clade contains the eastern Pacific speciesand most of the Caribbean species, whereas the other clade containsthe eastern Atlantic species. The new species is the sisterto the rest of the Caribbean species, which also form a monophyleticgroup. This phylogenetic hypothesis suggests that two consecutivevicariant events have affected the biogeography of Hypselodoris:(1) the closure of communication between the tropical Indo-Pacificregion and the Atlantic and eastern Pacific, completed withthe formation of the East Pacific Barrier; and (2) the riseof the Panama isthmus. (Received 19 December 2003; accepted 12 October 2005)