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Movement of fishes in the aquatic realm is fundamental to their ecology and survival. Movement can be driven by a variety of biological, physiological, and environmental factors occurring across all spatial and temporal scales. The intrinsic capacity of movement to impact fish individually (e.g., foraging) with potential knock-on effects throughout the ecosystem (e.g., food web dynamics) has garnered considerable interest in the field of movement ecology. The advancement of technology in recent decades, in combination with ever-growing threats to freshwater and marine systems, have further spurred empirical research and theoretical considerations. Given the rapid expansion within the field of movement ecology and its significant role in informing management and conservation efforts, a contemporary and multidisciplinary review about the various components influencing movement is outstanding. Using an established conceptual framework for movement ecology as a guide (i.e., Nathan et al., 2008 PNAS. 105:19052), we synthesize the environmental and individual factors that affect the movement of fishes. Specifically, internal (e.g., energy acquisition, endocrinology, and homeostasis) and external (biotic and abiotic) environmental elements are discussed, as well as the different processes that influence individual-level (or population) decisions, such as navigation cues, motion capacity, propagation characteristics, and group behaviours. In addition to environmental drivers and individual movement factors, we also explore how associated strategies help survival by optimizing physiological and other biological states. Next, we identify how movement ecology is increasingly being incorporated into management and conservation by highlighting the inherent benefits that spatio-temporal fish behaviour imbues into policy, regulatory, and remediation planning. Finally, we consider the future of movement ecology by evaluating ongoing technological innovations and both the challenges and opportunities that these advancements create for scientists and managers. As aquatic ecosystems continue to face alarming climate (and other human-driven) issues that impact animal movements, the comprehensive and multidisciplinary assessment of movement ecology will be instrumental in developing plans to guide research and promote sustainability measures for aquatic resources.

Visual perception is, in part, a function of the ambient illumination spectrum. In aquatic environments, illumination depends upon the water's optical properties and depth, both of which can change due to anthropogenic impacts: turbidity is increasing in many aquatic habitats, and many species have shifted deeper in response to warming surface waters (known as bathymetric shifts). Although increasing turbidity and bathymetric shifts can result in similarly large changes to a species' optical environment, no studies have yet examined the impact of the latter on visually mediated interactions. Here, we examine a potential link between climate change and visual perception, with a focus on colour. We discuss (i) what is known about bathymetric shifts; (ii) how the impacts of bathymetric shifts on visual interactions may be distributed across species; (iii) which interactions might be affected; and (iv) the ways that animals have to respond to these changes. As warming continues and temperature fluctuations grow more extreme, many species may move into even deeper waters. There is thus a need for studies that examine how such shifts can affect an organism's visual world, interfere with behaviour, and impact fitness, population dynamics, and community structure.

Summary Transparent zooplankton and nekton are often nearly invisible when viewed under ambient light in the pelagic zone [1–3]. However, in this environment, where the light field is directional (and thus likely to cause reflections), and under the bioluminescent searchlights of potential predators, animals may be revealed by reflections from their body surface [4–7]. We investigated the cuticle surfaces of seven species of hyperiids (Crustacea; Amphipoda) using scanning electron microscopy and found two undocumented features that may reduce reflectance. We found that the legs of Cystisoma spp. (n = 5) are covered with an ordered array of nanoprotuberances 200 ± 20 nm SD in height that function optically as a gradient refractive index material [6, 8, 9]. Additionally, we observed that Cystisoma and six other species of hyperiids are covered with a monolayer of homogenous spheres (diameters ranging from 52 ± 7 nm SD on Cystisoma spp. to 320 ± 15 nm SD on Phronima spp.). Optical modeling using effective medium theory and transfer matrix methods demonstrated that both the nanoprotuberances and the monolayers reduce reflectance by as much as 100-fold, depending on the wavelength and angle of the incident light and the thickness of the gradient layer. Even though we only consider surface reflectance and not internal light scattering, our study demonstrates that these nanoprotuberances and spheres can improve crypsis in a featureless habitat where the smallest reflection can render an animal vulnerable to visual predation.

The increased availability of genome sequences has provided remarkable advances in our understanding of the evolutionary history of non-model species. One important consideration in evolutionary studies is the role of demographic history in shaping contemporary levels and distribution of genetic variation. In green sea turtles (Chelonia mydas), a draft genome sequence has recently been made available, yet little is known regarding how past demographic events have shaped genomic variation in populations of this species. In this study, single nucleotide polymorphisms were identified in the green sea turtle’s genome and used to reconstruct past demographic events. It was found that this green sea turtle population, from the South China Sea, experienced a marked expansion ca. 0.8 million years ago near the mid-Pleistocene transition (MPT). Simulations revealed that the past demographic history can at least partially be explained by changes in population structure and gene flow; possibly associated with the climatic and geomagnetic events occurring since the MPT. The results demonstrate the importance in considering the effects of gene flow when reconstructing historical changes in population size and provide an extensive set of genomic resources for future evolutionary studies of green sea turtles.

The iconic red hourglass of the black widow spiders (genus Latrodectus) is traditionally considered an aposematic signal, yet experimental evidence is lacking. Here, we present data that suggest that black widow coloration may have evolved to be an aposematic signal that is more conspicuous to their vertebrate predators than to their insect prey. In choice experiments with wild birds, we found that the red-and-black coloration deters potential predators: Wild birds were ~3 times less likely to attack a black widow model with an hourglass than one without. Using visual-system appropriate models, we also found that a black widow’s red-and-black color combo is more apparent to a typical bird than a typical insect. Additionally, an ancestral reconstruction reveals that red dorsal coloration is ancestral in black widows and that at some point some North American widows lost their red dorsal coloration. Behaviorally, differences in red dorsal coloration between 2 North American species are accompanied by differences in microhabitat that affects how often a bird will view a black widow’s dorsal region. All observations are consistent with a cost–benefit trade-off of being more conspicuous to predators than to prey. We suggest that limiting detection by prey may help explain why red and black aposematic signals occur frequently in nature.

A major goal of sensory ecology is to identify factors that underlie sensory-trait variation. One open question centers on why fishes show the greatest diversity among vertebrates in their capacity to detect color (i.e., spectral sensitivity). Over the past several decades, λmax values (photoreceptor class peak sensitivity) and chromacy (photoreceptor class number) have been cataloged for hundreds of fish species, yet the ecological basis of this diversity and the functional significance of high chromacy levels (e.g., tetra- and pentachromacy) remain unclear. In this study, we examined phylogenetic, physiological, and ecological patterns of spectral sensitivity of ray-finned fishes (Actinoptergyii) via a meta-analysis of data compiled from 213 species. Across the fishes sampled, our results indicate that trichromacy is most common, ultraviolet λmax values are not found in monochromatic or dichromatic species, and increasing chromacy, including from tetra- to pentachromacy, significantly increases spectral sensitivity range. In an ecological analysis, multivariate phylogenetic latent liability modeling was performed to analyze correlations between chromacy and five hypothesized predictors (depth, habitat, diet, body coloration, and body size). In a model not accounting for phylogenetic relatedness, each predictor with the exception of habitat significantly correlated with chromacy: a positive relationship in body color and negative relationships with body size, diet, and depth. However, after phylogenetic correction, the only remaining correlated predictor was depth. The findings of this study indicate that phyletic heritage and depth are important factors in fish spectral sensitivity and impart caution about excluding phylogenetic comparative methods in studies of sensory trait variation.

Camouflage is exceptionally challenging in pelagic environments because of their featureless nature. Thus, it is perhaps no surprise that pelagic species have evolved highly sophisticated cryptic strategies, three of which—transparency, mirrors, and counterillumination—are rare or absent in other habitats. Pelagic visual systems are equally complex, and several visual capabilities, including UV and polarization sensitivity and intraocular filters, are thought to facilitate detection of camouflaged animals. This article reviews the optical nature of the pelagic realm and both the camouflage and camouflage-breaking strategies of its inhabitants, focusing primarily on underlying principles and what remains to be discovered. A theme throughout is that far more is known about the structures of the optical and visual systems involved than about their function, an imbalance that is due primarily to the rarity of observations of undisturbed behavior.

Fluorescence is a physico-chemical energy exchange where shorter-wavelength photons are absorbed by a molecule and are re-emitted as longer-wavelength photons. It has been suggested a means of communication in several taxa including flowers, pitcher plants, corals, algae, worms, squid, spiders, stomatopods, fish, reptiles, parrots and humans. The surface or object that the pigment molecule is part of appears to glow due to its setting rather than an actual production of light, and this may enhance both signals and, in some cases, camouflage. This review examines some known uses of fluorescence, mainly in the context of visual communication in animals, the challenge being to distinguish when fluorescence is a functional feature of biological coloration or when it is a by-product of a pigment or other molecule. In general, we conclude that most observations of fluorescence lack enough evidence to suggest they are used in visually driven behaviours. This article is part of the themed issue ‘Animal coloration: production, perception, function and application’.

Visual acuity (the ability to resolve spatial detail) is highly variable across fishes. However, little is known about the evolutionary pressures underlying this variation. We reviewed published literature to create an acuity database for 159 species of ray-finned fishes (Actinopterygii). Within a subset of those species for which we had phylogenetic information and anatomically-measured acuity data (n=81), we examined relationships between acuity and both morphological (eye size and body size) and ecological (light level, water turbidity, habitat spatial complexity, and diet) variables. Acuity was significantly correlated with eye size (p

Although the function of polarization vision, the ability to discern the polarization characteristics of light, is well established in many terrestrial and benthic species, its purpose in pelagic species (squid and certain fish and crustaceans) is poorly understood [1]. A long-held hypothesis is that polarization vision in open water is used to break the mirror camouflage of silvery fish, as biological mirrors can change the polarization of reflected light [2,3]. Although, the addition of polarization information may increase the conspicuousness of silvery fish at close range, direct evidence that silvery fish - or indeed any pelagic animal - are visible at longer distances using polarization vision rather than using radiance (i.e. brightness) vision is lacking. Here we show, using in situ polarization imagery and a new visual detection model, that polarization vision does not in fact appear to allow viewers to see silvery fish at greater distances.

Summary Animals in the lower mesopelagic zone (600–1,000 m depth) of the oceans have converged on two major strategies for camouflage: transparency and red or black pigmentation [1]. Transparency conveys excellent camouflage under ambient light conditions, greatly reducing the conspicuousness of the animal's silhouette [1, 2]. Transparent tissues are seldom perfectly so, resulting in unavoidable internal light scattering [2]. Under directed light, such as that emitted from photophores thought to function as searchlights [3–8], the scattered light returning to a viewer will be brighter than the background, rendering the animal conspicuous [2, 4]. At depths where bioluminescence becomes the dominant source of light, most animals are pigmented red or black, thereby reflecting little light at wavelengths generally associated with photophore emissions and visual sensitivities [3, 9–14]. However, pigmented animals are susceptible to being detected via their silhouettes [5, 9–11]. Here we show evidence for rapid switching between transparency and pigmentation under changing optical conditions in two mesopelagic cephalopods, Japetella heathi and Onychoteuthis banksii . Reflectance measurements of Japetella show that transparent tissue reflects twice as much light as pigmented tissue under direct light. This is consistent with a dynamic strategy to optimize camouflage under ambient and searchlight conditions.

Because light in the pelagic environment is partially polarized, it has been suggested that the polarization sensitivity found in certain pelagic species may serve to enhance the contrast of their transparent zooplankton prey. We examined its potential during cruises in the Gulf of Mexico and Atlantic Ocean and at a field station on the Great Barrier Reef. First, we collected various species of transparent zooplankton and micronekton and photographed them between crossed polarizers. Many groups, particularly the cephalopods, pelagic snails, salps and ctenophores, were found to have ciliary, muscular or connective tissues with striking birefringence.In situpolarization imagery of the same species showed that, while the degree of underwater polarization was fairly high (approx. 30% in horizontal lines of sight), tissue birefringence played little to no role in increasing visibility. This is most likely due to the low radiance of the horizontal background light when compared with the downwelling irradiance. In fact, the dominant radiance and polarization contrasts are due to unpolarized downwelling light that has been scattered from the animal viewed against the darker and polarized horizontal background light. We show that relatively simple algorithms can use this negative polarization contrast to increase visibility substantially.

SUMMARY Transparency is an effective form of camouflage, but it must be present throughout the entire volume of an animal to succeed. Certain environmental stressors may cause physiological responses that increase internal light scattering, making tissue less transparent and more conspicuous to predators. We tested this in the transparent grass shrimp, Palaemonetes pugio, which is found in shallow estuaries where both salinity and temperature change rapidly because of tidal cycles, evaporation and runoff. Animals originally kept at a salinity of 15 p.p.t. and a temperature of 20°C were placed into solutions with salinities of 0, 15, 25 or 30 p.p.t. and temperatures of 13, 20 or 27°C for 12 h (N=26 for each of 12 treatments). Under the control conditions of 15 p.p.t. at 20°C, the transparency of grass shrimp tails was 54±3% (mean ± s.e.). At higher salinities and at both higher and lower temperatures, transparency dropped significantly (P

Many arthropod species have evolved to thrive only on the leaves of a particular species of plant, which they must be capable of finding in order to survive accidental displacement, developmental transitions or the changing of the seasons. A number of studies have tested whether such species select leaves to land or oviposit on based on their color, shape or size. Unfortunately, many studies did not control for correlates of these characters, such as the brightness of different colors, the areas of different shapes, and the level of ambient illumination in the vicinity of different sizes of leaves. In the present study, we tested for leaf color, shape and size preferences in a leaf-dwelling jumping spider ( Lyssomanes viridis ) with known summer and winter host plants, while controlling for these correlates. First, color preferences were tested outdoors under the natural illumination of their forest habitat. Lyssomanes viridis did not prefer to perch on a green substrate compared with various shades of gray, but did prefer the second darkest shade of gray we presented them with. Of the green and gray substrates, this shade of gray9s integrated photon flux (350–700 nm), viewed from below, i.e. the spider9s perspective in the arena, was the most similar to that of real leaves. This relationship also held when we weighted the transmitted photon flux by the jumping spiders9 green photopigment spectral sensitivity. Spiders did not prefer the star-like leaf shape of their summer host plant, Liquidambar styraciflua , to a green circle of the same area. When given a choice between a L. styraciflua leaf-shaped stimulus that was half the area of an otherwise identical alternative, spiders preferred the larger stimulus. However, placing a neutral density filter over the side of the experimental arena with the smaller stimulus abolished this preference, with spiders then being more likely to choose the side of the arena with the smaller stimulus. In conclusion, L. viridis appears to use ambient illumination and possibly perceived leaf brightness but not leaf shape or color to locate its microhabitat. This calls for a careful re-examination of which visual cues a variety of arthropods are actually attending to when they search for their preferred host species or microhabitat.

The Hamilton–Zuk hypothesis proposes that females prefer male secondary sexual traits because they are honest indicators of parasite resistance. Despite the attention that this hypothesis has received, its role in sexual selection remains equivocal. This study presents the first field test in guppies of two key predictions of the Hamilton–Zuk hypothesis: (1) that within populations, the most highly ornamented males have the fewest parasites and (2) that among populations, males in high parasite populations have the most conspicuous ornaments. Five hundred male guppies from 19 distinct populations in the Northern Range of Trinidad were inspected for Gyrodactylus parasites and photographed. Eight measures of orange spot ornamentation were used to test the predictions: hue, saturation, lightness, relative area, number, and area-weighted hue, saturation, and lightness. Parasite load had no significant effect on any of these measures. There was also no relationship between orange spot ornamentation and parasite abundance among populations. Guppies from high-predation environments had significantly more parasites, and their orange coloration was lighter and less saturated than that in guppies from low-predation environments. Despite previous lab results, this study found no relationship between parasite load and male orange spot ornamentation.

Ultraviolet (UV) photoreceptors have been reported in a wide variety of freshwater and marine organisms, suggesting that UV vision is prominent in aquatic ecosystems. However, its adaptive significance remains speculative. The present study tested whether the foraging of juvenile bluegill (Lepomis macrochirus) is enhanced in the presence of UV radiation (UVR). Laboratory feeding trials were conducted in a laminar flow tank in which L. macrochirus juveniles between 2.3 and 3.5 cm standard length were fed the cladoceran Daphnia magna. Sighting and striking distances, as well as capture success, were measured in the presence and absence of UVR. Mean sighting and striking distances and capture success did not differ significantly between the two light treatments. There were also no significant differences in the frequency distributions of sighting and striking distance. These results suggest that UV vision may not be used to enhance foraging in L. macrochirus on Daphnia within the size class tested.

The colors of deep-sea species are generally assumed to be cryptic, but it is not known how cryptic they are and under what conditions. This study measured the color of approximately 70 deep-sea species, both pelagic and benthic, and compared the results with two sets of predictions: 1) optimal crypsis under ambient light, 2) optimal crypsis when viewed by bioluminescent "searchlights." The reflectances of the pelagic species at the blue-green wavelengths important for deep-sea vision were far lower than the predicted reflectances for crypsis under ambient light and closer to the zero reflectance prediction for crypsis under searchlights. This suggests that bioluminescence is more important than ambient light for the visual detection of pelagic species at mesopelagic depths. The reflectances of the benthic species were highly variable and a relatively poor match to the substrates on which they were found. However, estimates of the contrast sensitivity of deep-sea visual systems suggest that even approximate matches may be sufficient for crypsis in visually complex benthic habitats. Body coloration was generally uniform, but many crabs had striking patterns that may serve to disrupt the outlines of their bodies.

Mirrored and colored surfaces are common adaptations for crypsis in pelagic habitats. Although highly successful when optimized for a particular situation, either may become less successful if it is then viewed in a different situation. In this study we examine the relative robustness of these two strategies by determining how visible an organism becomes when viewed under optical conditions different from those under which the camouflage is optimal. Underwater radiance distributions were calculated using inherent optical properties measured in coastal waters 80 km off the coast of New Hampshire. These radiance distributions were then used to calculate optimally cryptic diffuse and specular reflectance spectra as a function of depth, solar elevation, viewing angle, and azimuth. Then the visibilities of organisms cryptic in one situation viewed in a different situation were calculated, using the Atlantic Cod, Gadus morhua, as the viewer. In contrast to benthic organisms, pelagic organisms cryptic under one set of optical conditions were quite visible under a different set, particularly when viewed from a different azimuth. The crypsis afforded by mirrored surfaces was generally more robust than that resulting from colored surfaces. However, because mirrored surfaces could never be perfectly cryptic when viewed in the azimuth of the sun, neither strategy clearly outperformed the other. In general, crypsis by colored or mirrored surfaces was not robust in nearsurface water, which may help explain both the predominance of transparent species in near-surface pelagic habitats and the vertical migration of many colored and mirrored species. The results also show that three common foraging strategies—circling, crepuscular activity, and driving prey toward the surface—all increase the visibility of cryptically colored or mirrored prey.

After emerging from underground nests, sea turtle hatchlings migrate through the surf zone and out to the open ocean. During this migration, both waves and water currents can disrupt hatchling orientation by unpredictably rotating the turtles away from their migratory headings. In addition, waves cause turtles to roll and pitch, temporarily impeding forward swimming by forcing the hatchlings into steeply inclined positions. To maintain seaward orientation and remain upright in the water column, hatchlings must continuously compensate for such displacements. As a first step toward determining how this is achieved, we studied the responses of loggerhead (Caretta caretta L.) sea turtle hatchlings to rotational displacements involving yaw, roll, and pitch. Hatchlings responded to rotations in the horizontal plane (yaw) by extending the rear flipper on the side opposite the direction of rotation. Thus, the flipper presumably acts as a rudder to help turn the turtle back toward its original heading. Turtles responded to rotations in the roll plane with stereotypic movements of the front flippers that act to right the hatchlings with respect to gravity. Finally, hatchlings responded to rotations in the pitch plane with movements of the hind flippers that appear likely to curtail or counteract the pitching motion. Thus, the results of these experiments imply that young sea turtles emerge from their nests possessing a suite of stereotypic behavioral responses that function to counteract rotational displacements, enable the animals to maintain equilibrium, and facilitate efficient movement toward the open sea.

Cuttlefish are cephalopods capable of rapid camouflage responses to visual stimuli. However, it is not always clear to what these animals are responding. Previous studies have found cuttlefish to be more responsive to lateral stimuli rather than substrate. However, in previous works, the cuttlefish were allowed to settle next to the lateral stimuli. In this study, we examine whether juvenile cuttlefish (Sepia officinalis) respond more strongly to visual stimuli seen on the sides versus the bottom of an experimental aquarium, specifically when the animals are not allowed to be adjacent to the tank walls. We used the Sub Sea Holodeck, a novel aquarium that employs plasma display screens to create a variety of artificial visual environments without disturbing the animals. Once the cuttlefish were acclimated, we compared the variability of camouflage patterns that were elicited from displaying various stimuli on the bottom versus the sides of the Holodeck. To characterize the camouflage patterns, we classified them in terms of uniform, disruptive, and mottled patterning. The elicited camouflage patterns from different bottom stimuli were more variable than those elicited by different side stimuli, suggesting that S. officinalis responds more strongly to the patterns displayed on the bottom than the sides of the tank. We argue that the cuttlefish pay more attention to the bottom of the Holodeck because it is closer and thus more relevant for camouflage., United States. Office of Naval Research (Grant N00014-09-1-1053)

One classic explanation for the remarkable diversity of flower colors across angiosperms involves evolutionary shifts among different types of pollinators with different color preferences. However, the pollinator shift model fails to account for the many examples of color variation within clades that share the same pollination system. An alternate explanation is the competition model, which suggests that color divergence evolves in response to interspecific competition for pollinators, as a means to decrease interspecific pollinator movements. This model predicts color overdispersion within communities relative to null assemblages. Here, we combine morphometric analyses, field surveys, and models of pollinator vision with a species-level phylogeny to test the competition model in the primarily hummingbird-pollinated clade Iochrominae (Solanaceae). Results show that flower color as perceived by pollinators is significantly overdispersed within sites. This pattern is not simply due to phylogenetic history: phylogenetic community structure does not deviate from random expectations, and flower color lacks phylogenetic signal. Moreover, taxa that occur in sympatry occupy a significantly larger volume of color space than those in allopatry, supporting the hypothesis that competition in sympatry drove the evolution of novel colors. We suggest that competition among close relatives may commonly underlie floral divergence, especially in species-rich habitats where congeners frequently co-occur.

Despite the prevalence and importance of transparency in organisms, particularly pelagic species, it is a poorly understood characteristic. This article reviews the current state of knowledge on the distribution, ecology, and physical basis of biological transparency. Particular attention is paid to the distribution of transparent species relative to their optical environment, the relationship between transparency and visual predation, the physics of transparency, and what is known about the anatomical and ultrastructural modifications required to achieve this condition. Transparency is shown to be primarily a pelagic trait, uncommon in other aquatic habitats and extremely rare on land. Experimental and theoretical studies in terrestrial, freshwater, and marine ecosystems have shown that transparency is a successful form of camouflage, and that several visual adaptations seem to counter it. The physical basis of transparency is still poorly understood, but anatomical observations and mathematical models show that there are various routes to transparency. Future avenues for research include examination of the ultrastructure and optical properties of transparent tissue, exploring the link between transparent species and special visual modifications in the species they interact with, and analysis of the evolution of transparency using comparative methods.

The use of transparency as camouflage in the epipelagic realm is complicated by the presence of ultraviolet radiation, because the presence of UV-protective pigments decreases UV transparency and may reveal transparent zooplankton to predators and prey with UV vision. During July 1999, September 1999, and June 2000, transparency measurements (from 280 to 500 nm) were made on living specimens of 15 epipelagic (collection depth: 0–20 m, average: 11 ± 1 m) and 19 mesopelagic (collection depth: 150–790 m, average: 370 ± 40 m) species of transparent zooplankton from Oceanographer Canyon and Wilkinson Basin in the Northwest Atlantic Ocean. In addition, measurements of downwelling irradiance (from 330 to 500 nm) versus depth were made. The tissues from epipelagic zooplankton had lower UV transparency than those from mesopelagic zooplankton, while the average visible transparency (at 480 nm) of the two groups was not significantly different. Percent transparency was positively correlated with wavelength over most of the measured range, with a rapid decrease below a certain cutoff wavelength. In mesopelagic tissues, the cutoff wavelength was generally 300 nm. In epipelagic tissues, the cutoff wavelength was between 300 and 400 nm. Twelve out of 19 epipelagic tissues had transparencies at 320 nm that were half or less than their 480 nm transparency values, versus only 4 out of 21 mesopelagic tissues. The effects of UV absorption on UV visibility and minimum attainable depth were modeled using contrast theory and the physics of light attenuation. Because UV absorption was generally significantly greater in the UVB than in the UVA spectrum (where UV vision occurs), and because the highest UV absorption was often found in less transparent individuals, its modeled effects on visibility were slight compared to its effects on minimum attainable depth.

As the open ocean environment lacks points of refuge from visual predators, it has favored the evolution of extraordinary adaptations for optical concealment, such as vertical migration, trans- parency and counterillumination. Bioluminescent plankton, which respond to a mechanical disturb- ance with a flash of light, are ubiquitous in the ocean and potentially a threat to any organism seeking darkness as a means to evade detection. Estimating encounter probabilities for organisms maneu- vering through luminescent 'minefields' requires characterization of the three-dimensional distri- bution patterns of the potential light emitters. In order to measure nearest neighbor distances (NNDs) of bioluminescent biota, we have developed a spatial plankton analysis technique (SPLAT) for 3D reconstruction and statistical analysis of the spatial point patterns of identified bioluminescent displays. Analysis of aggregations of bioluminescent biota in the Gulf of Maine (Wilkinson Basin) revealed that encounter probabilities were highest in the temperature minimum zone (temperature

To learn how organisms apportion space in the open ocean, biological oceanographers have sought to improve temporal and spatial resolution of ocean sampling systems. Their objectives are to simultaneously measure physical, chemical and biological structure in the water column in order to find significant correlations that may reveal underlying processes. Here we report one such correlation between intense peaks of bioluminescence and density discontinuities in the water column. Intensified video recordings made in these bioluminescent “hot spots” were analyzed with a computer image-recognition program that identifies organisms based on the temporal and spatial characteristics of their luminescent displays. Based on this analysis, the source of the “hot spots” was found to be very thin layers (0.5 m) of the bioluminescent copepod Metridia lucens present at from 5 to 100 times average background concentrations. Given the recent discovery that the vertical distribution of marine snow is also strongly correlated with density discontinuities in the water column, we suggest that this finding may provide a possible explanation for the disparity between estimated energy requirements of marine copepods and measurements of average in situ food concentrations. The energy costs associated with locating food-rich micro-patches is greatly reduced if those patches are spread out into very thin layers, because the search strategy can be reduced from three dimensions to one.

Transparency measurements (at 400 to 700 nm) were made on living specimens of 29 common spe- cies of gelatinous zooplankton from the Northwestern At- lantic Ocean and Gulf of Mexico. Percent transparency ranged from 91% for the hydromedusa Sibogota typa to 0.5 1% for the pteropod Clione limacina. Percent transpar- ency was linearly and positively correlated with wave- length, with slopes of the regression lines (normalized to the percent transparency at 480 nm) ranging from O.O27%/nm for Sibogota typa to 0.5 1 %/nm for the cteno- phore Mnemiopsis mactydi (average 0.17 2 O.O19%/nm). There was no significant correlation between the percent transparency of an animal and its daytime depth distribu- tion. The relationship between percent transparency and sighting distance when viewed from below was modeled and showed that, due to the increase of the minimum contrast threshold for object detection at lower light lev- els, the usefulness of transparency as camouflage in- creases dramatically with depth. A preliminary account of these results was presented by the authors at the four- teenth meeting of the Ocean Optics Society in November

Many morphological, chemical, and behavioural characteristics of echinoderms have been implicated as defences against ultraviolet light, though no studies have investigated whether adult echinoderms are damaged by this form of radiation. This study tests whether the brittlestar Ophioderma brevispinum (Ophiuroidea: Echinodermata) is damaged by solar ultraviolet radiation. Specimens of O. brevispinum were exposed to sunlight at a field station on the North Carolina coast. After 4 d of exposure, 12 out of 13 animals were dead and the remaining animal was moderately damaged. The animals in the control treatment, protected by a UV-opaque filter, suffered almost no damage.

Aggregation is a common life-history trait in open-water taxa. Qualitative understanding of how aggregation by prey influences their encounter rates with predators is critical for understanding pelagic predator-prey interactions and trophic webs.We extend a recently developed theory on underwater visibility to predict the consequences of grouping in open-water species in terms of increased visual detection of groups by predators. Our model suggests that enhanced visibility will be relatively modest, with maximum detection distance typically only doubling for a 100-fold increase in the number of prey in a group. This result suggests that although larger groups are more easily detected, this cost to aggregation will in many cases be dominated by benefits, especially through risk dilution in situations where predators cannot consume all members of a discovered group. This, in turn, helps to explain the ubiquity of grouping across a great variety of open-water taxa. Postprint

Weaponry and color badges are commonly theorized to function as visual signals of aggressiveness or fighting ability. However, few studies have supported a signaling function of weaponry, and the role of color in invertebrate competitive interactions remains virtually unexplored. Jumping spiders (Salticidae) make excellent invertebrate models for studying weaponry and color because males of many species are colorful and possess exaggerated chelicerae, which are used as weapons in escalated contests. To determine whether color or weaponry might function as visual signals in male-male competitions, we investigated relationships between contest success, cheliceral length, and red coloration in Lyssomanes viridis. Males having longer chelicerae than their opponents were significantly more likely to win (p=0.0008). Males who won, despite being smaller than their opponents, had significantly less red chelicerae than their opponents (p=0.01). Male and female cheliceral length, as well as foreleg length, correlated tightly with body size. Cheliceral and foreleg length showed significantly stronger positive allometry in males than in females. We conclude that male chelicerae and forelegs are under strong positive selection for their use in physical fights and/or as visual signals of fighting ability.

While transparency, cryptic coloration, and counterillumination are all highly successful cryptic strategies for pelagic species, they become less effective when confronted with varying optical conditions. Transparent species are susceptible to detection by reflections from their body surface, particularly at shallow depths. Colored and mirrored species are vulnerable to detection when viewed from certain angles, or at certain times of day. Counterilluminating species must cope with the changes in the angular distribution and spectra of downwelling light at different depths. In all cases the vulnerabilities are more pronounced at shallow depths and essentially negligible at depths greater than 200 m. The results suggest interesting adaptations both for crypsis (e.g., anti-reflection coatings, variable coloration, variable filters for photophores) and for visual detection (e.g., circling, crepuscular predation), all of which are potentially fruitful topics for future research.

Oceanography has seen the recent development of many new tools and techniques. The subfield of pelagic visual ecology in particular has benefited from the development of more reliable, portable, and economic tools and techniques that can be taken to sea including spectrometers, microspectrophotometery, electroretinography, and ultraviolet and polarization imaging systems. These advances have led to a relative wealth of data on the visual physiology of pelagic species and on the optical properties of these species and their environment. These data, particularly in combination with computational methods, have tested long-standing hypotheses in pelagic ecology and led to new hypotheses and research directions. While the ability to study pelagic species still lags far behind what is possible in terrestrial and coastal environments, a renaissance may be developing in the study of the integrative biology of pelagic species.

Polarization sensitivity is documented in a range of marine animals. The variety of tasks for which animals can use this sensitivity, and the range over which they do so, are confined by the visual systems of these animals and by the propagation of the polarization information in the aquatic environment. We examine the environmental physical constraints in an attempt to reveal the depth, range and other limitations to the use of polarization sensitivity by marine animals. In clear oceanic waters, navigation that is based on the polarization pattern of the sky appears to be limited to shallow waters, while solar-based navigation is possible down to 200–400 m. When combined with intensity difference, polarization sensitivity allows an increase in target detection range by 70–80% with an upper limit of 15 m for large-eyed animals. This distance will be significantly smaller for small animals, such as plankton, and in turbid waters. Polarization-contrast detection, which is relevant to object detection and communication, is strongly affected by water conditions and in clear waters its range limit may reach 15 m as well. We show that polarization sensitivity may also serve for target distance estimation, when examining point source bioluminescent objects in the photic mesopelagic depth range.

SUMMARY Visual displays often play a large role in animal communication,particularly in sexual interactions. The blue crab Callinectes sapidus is both colorful and highly visually responsive, yet almost all studies of their courtship have focused on chemical cues. In the blue crab's underwater environment, however, visual cues may function more rapidly and over a longer distance than chemical cues. Given that blue crabs are aggressive and cannibalistic, visual cues may therefore allow blue crabs to quickly evaluate potential mates from safer distances. In the present study we show that courtship and mate choice behavior in C. sapidus can be stimulated by visual cues alone. Further, we show that males have a preference for females with red claw dactyls. In binary choice experiments, males displayed more often to photographs of females with red claws than to those with white claws or to those with black claws that were isoluminant to the red ones. This strongly suggests that male blue crabs made their choices based on the hue of the red claws, further suggesting that blue crabs are capable of color vision and use color in mate choice.

Biological receptors used to detect underwater light vary in structure, from simple clusters of cells that only detect light intensity to complex organs that form detailed images. While most aquatic organisms perceive some portion of the visible spectrum, UV photoreceptors have recently been found in a wide diverse of species, from bacteria to fish. These UV photoreceptors are thought to aid in communication, foraging, and avoidance of damaging sunlight. However, they are not without costs as seeing in the UV can cause damage to the eye and image degradation. Given that the spectral composition of underwater light can directly affect an organism's physiology and behavior, environmental stressors that threaten to alter the color and clarity of fresh waters are a concern. These include nutrient and sediment loading associated with changes in climatic patterns and anthropogenic activities. This chapter discusses: (1) the underwater UV environment, (2) the structure and function of photoreceptors and the methods used to measure their spectral sensitivity, (3) the distribution of UV photoreceptors among freshwater organisms, (4) the adaptive significance of UV vision, and (5) potential effects of environmental stressors on UV vision.

Cephalopods, and in particular the cuttlefishSepia officinalis, are common models for studies of camouflage and predator avoidance behaviour. Preventing detection by predators is especially important to this group of animals, most of which are soft-bodied, lack physical defences, and are subject to both visually and non-visually mediated detection. Here, we report a novel cryptic mechanism inS. officinalisin which bioelectric cues are reduced via a behavioural freeze response to a predator stimulus. The reduction of bioelectric fields created by the freeze-simulating stimulus resulted in a possible decrease in shark predation risk by reducing detectability. The freeze response may also facilitate other non-visual cryptic mechanisms to lower predation risk from a wide range of predator types.

We measured downwelling spectral vector irradiance (from 350 to 800 nm) during evening civil and nautical twilight (solar elevation down to -12°). Nine sets of measurements were taken to cover the first half of the lunar cycle (from the new to full moon) and were also used to calculate chromaticity (CIE 1976 u'v'). The lunar phase had no consistent effect on downwelling irradiance until solar elevation was less than -8°. For lower solar elevations, the effect of the moon increased with the fraction of the illuminated lunar disk until the fraction was approximately 50%. For fractions greater than 50%, the brightness and chromaticity of the downwelling irradiance were approximately independent of the fraction illuminated, likely because the greater brightness of a fuller moon was offset by its lower elevation during twilight. Given the importance of crepuscular periods to animal activity, including predation, reproductive cycles, and color vision in dim light, these results may have significant implications for animal ecology.

Visual performance and visual interactions in pelagic animals are notoriously hard to investigate because of our restricted access to the habitat. The pelagic visual world is also dramatically different from benthic or terrestrial habitats, and our intuition is less helpful in understanding vision in unfamiliar environments. Here, we develop a computational approach to investigate visual ecology in the pelagic realm. Using information on eye size, key retinal properties, optical properties of the water and radiance, we develop expressions for calculating the visual range for detection of important types of pelagic targets. We also briefly apply the computations to a number of central questions in pelagic visual ecology, such as the relationship between eye size and visual performance, the maximum depth at which daylight is useful for vision, visual range relations between prey and predators, counter-illumination and the importance of various aspects of retinal physiology. We also argue that our present addition to computational visual ecology can be developed further, and that a computational approach offers plenty of unused potential for investigations of visual ecology in both aquatic and terrestrial habitats.

This optical feature of some iridescent wings catches a suitor's eye in the deep forest. Iridescent butterfly scales are visually stunning structures that reflect highly saturated colour. They also create an array of non-chromatic optical phenomena, such as polarization, polarization mixing and highly directional flashes1,2, but the ecological purpose of these effects is unclear3,4. Here we show that polarized light is used in mate recognition by Heliconius butterflies, a genus that is known to rely on visual cues in sexual selection and speciation5. This terrestrial example of exploitation of polarized light may have adaptive value in dense forest, where illumination varies greatly in spectrum and intensity.

We introduce a broadened framework to study aspects of coevolution based on the NK class of statistical models of rugged fitness landscapes. In these models the fitness contribution of each of N genes in a genotype depends epistatically on K other genes. Increasing epistatic interactions increases the rugged multipeaked character of the fitness landscape. Coevolution is thought of, at the lowest level, as a coupling of landscapes such that adaptive moves by one player deform the landscapes of its immediate partners. In these models we are able to tune the ruggedness of landscapes, how richly intercoupled any two landscapes are, and how many other players interact with each player. All these properties profoundly alter the character of the coevolutionary dynamics. In particular, these parameters govern how readily coevolving ecosystems achieve Nash equilibria, how stable to perturbations such equilibria are, and the sustained mean fitness of coevolving partners. In turn, this raises the possibility that an evolutionary metadynamics due to natural selection may sculpt landscapes and their couplings to achieve coevolutionary systems able to coadapt well. The results suggest that sustained fitness is optimized when landscape ruggedness relative to couplings between landscapes is tuned such that Nash equilibria just tenuously form across the ecosystem. In this poised state, coevolutionary avalanches appear to propagate on all length scales in a power law distribution. Such avalanches may be related to the distribution of small and large extinction events in the record.

Summary Almost all animals, no matter how humble, possess eyes. Only those that live in total darkness, such as in a pitch-dark cave, may lack eyes entirely. Even at tremendous depths in the ocean — where the only lights that are ever seen are rare and fitful sparks of bioluminescence — most animals have eyes, and often surprisingly well-developed eyes. And despite their diversity (there are currently ten generally recognised optical types) all eyes have evolved in response to the remarkably varied light environments that are present in the habitats where animals live. Variations in the intensity of light, as well as in its direction, colour and dominant planes of polarisation, have all had dramatic effects on visual evolution. In the terrestrial habitats where we ourselves have most recently evolved, the light environment can vary quite markedly from day to night and from one location to another. In aquatic habitats, this variation can be orders of magnitude greater. Even though the ecologies and life histories of animals have played a major role in visual evolution, it is arguably the physical limitations imposed on photodetection by a given habitat and its light environment that have defined the basic selective pressures that have driven the evolution of eyes.