Evolution of compound eye morphology underlies differences in vision between closely related Drosophila species.

BACKGROUND: Insects have evolved complex visual systems and display an astonishing range of adaptations for diverse ecological niches. Species of Drosophila melanogaster subgroup exhibit extensive intra- and interspecific differences in compound eye size. These differences provide an excellent opportunity to better understand variation in insect eye structure and the impact on vision. Here we further explored the difference in eye size between D. mauritiana and its sibling species D. simulans. RESULTS: We confirmed that D. mauritiana have rapidly evolved larger eyes as a result of more and wider ommatidia than D. simulans since they recently diverged approximately 240,000 years ago. The functional impact of eye size, and specifically ommatidia size, is often only estimated based on the rigid surface morphology of the compound eye. Therefore, we used 3D synchrotron radiation tomography to measure optical parameters in 3D, predict optical capacity, and compare the modelled vision to in vivo optomotor responses. Our optical models predicted higher contrast sensitivity for D. mauritiana, which we verified by presenting sinusoidal gratings to tethered flies in a flight arena. Similarly, we confirmed the higher spatial acuity predicted for Drosophila simulans with smaller ommatidia and found evidence for higher temporal resolution. CONCLUSIONS: Our study demonstrates that even subtle differences in ommatidia size between closely related Drosophila species can impact the vision of these insects. Therefore, further comparative studies of intra- and interspecific variation in eye morphology and the consequences for vision among other Drosophila species, other dipterans and other insects are needed to better understand compound eye structure-function and how the diversification of eye size, shape, and function has helped insects to adapt to the vast range of ecological niches.

Increased eye size is favoured in Trinidadian killifish experimentally transplanted into low light, high competition environments.

Intraspecific variation in vertebrate eye size is well known. Ecological factors such as light availability are often correlated with shifts in relative eye size. However, experimental tests of selection on eye size are lacking. Trinidadian killifish (Anablepsoides hartii) are found in sites that differ in predation intensity. Sites that lack predators are characterized by lower light, high killifish densities, low resource availability, and intense competition for food. We previously found that killifish in sites that lack predators have evolved a larger "relative" eye size (eye size corrected for body size) than fish from sites with predators. Here, we used transplant experiments to test how selection operates on eye size when fish that are adapted to sites with predators are translocated into sites where predators are absent. We observed a significant "population × relative eye size" interaction; the relationship between relative eye size and a proxy for fitness (rates of individual growth) was positive in the transplanted fish. The trend was the opposite for resident fish. Such results provide experimental support that larger eyes enhance fitness and are favoured in environments characterized by low light and high competition.

Turbidity drives plasticity in the eyes and brains of an African cichlid.

Natural variation in environmental turbidity correlates with variation in the visual sensory system of many fishes, suggesting that turbidity may act as a strong selective agent on visual systems. Since many aquatic systems experience increased turbidity due to anthropogenic perturbations, it is important to understand the degree to which fish can respond to rapid shifts in their visual environment, and whether such responses can occur within the lifetime of an individual. We examined whether developmental exposure to turbidity (clear, <5 NTU; turbid, ∼9 NTU) influenced the size of morphological structures associated with vision in the African blue-lip cichlid Pseudocrenilabrus multicolor. Parental fish were collected from two sites (clear swamp, turbid river) in western Uganda. F1 broods from each population were split and reared under clear and turbid rearing treatments until maturity. We measured morphological traits associated with the visual sensory system (eye diameter, pupil diameter, axial length, brain mass, optic tectum volume) over the course of development. Age was significant in explaining variation in visual traits even when standardized for body size, suggesting an ontogenetic shift in the relative size of eyes and brains. When age groups were analyzed separately, young fish reared in turbid water grew larger eyes than fish reared in clear conditions. Population was important in the older age category, with swamp-origin fish having relatively larger eyes and optic lobes relative to river-origin fish. Plastic responses during development may be important for coping with a more variable visual environment associated with anthropogenically induced turbidity.

Phenotypic signatures of urbanization? Resident, but not migratory, songbird eye size varies with urban-associated light pollution levels.

Urbanization now exposes large portions of the earth to sources of anthropogenic disturbance, driving rapid environmental change and producing novel environments. Changes in selective pressures as a result of urbanization are often associated with phenotypic divergence; however, the generality of phenotypic change remains unclear. In this study, we examined whether morphological phenotypes in two residential species (Carolina Wren [Thryothorus ludovicianus] and Northern Cardinal [Cardinalis cardinalis]) and two migratory species (Painted Bunting [Passerina ciris], and White-eyed Vireo [Vireo griseus]), differed between urban core and edge habitats in San Antonio, Texas, USA. More specifically, we examined whether urbanization, associated sensory pollution (light and noise) and brightness (open, bright areas cause by anthropogenic land use) influenced measures of avian body (mass and frame size) and lateral eye size. We found no differences in body size between urban core and edge habitats for all species except the Painted Bunting, in which core-urban individuals were smaller. Rather than a direct effect of urbanization, this was due to differences in age structure between habitats, with urban-core areas consisting of higher proportions of younger buntings which are, on average, smaller than older birds. Residential birds inhabiting urban-core areas had smaller eyes compared to their urban-edge counterparts, resulting from a negative association between eye size and light pollution and brightness across study sites; notably, we found no such association in the two migratory species. Our findings demonstrate how urbanization may indirectly influence phenotypes by altering population demographics and highlight the importance of accounting for age when assessing factors driving phenotypic change. We also provide some of the first evidence that birds may adapt to urban environments through changes in their eye morphology, demonstrating the need for future research into relationships among eye size, ambient light microenvironment use, and disassembly of avian communities as a result of urbanization.

Experimental test of the influence of light availability on the evolution of eye size and behaviour in Daphnia.

There exists extensive variation in eye size. Much work has provided a connection between light availability and differences in eye size across taxa. Experimental tests of the role of the light environment on the evolution of eye size are lacking. Here, we performed a selection experiment that examined the influence of light availability on shifts in eye size and the connection between eye size and phototactic (anti-predator) behaviour in Daphnia. We set-up replicate experimental populations of Daphnia, repeatedly evaluated phenotypic shifts in eye size during the ~50-day experiment, and performed a common garden experiment at the end of the experiment to test for evolutionary shifts in eye size and behaviour. Our phenotypic analyses showed that eye size rapidly diverged between the light treatments; relative eye size was consistently larger in the low versus high light treatments. Selection on eye size was also modified by variation in density as increases in Daphnia density favoured a larger eye. However, we did not observe differences in eye size between the light treatments following two generations of common garden rearing at the end of the experiment. We instead observed strong shifts in anti-predator behaviour. Daphnia from the low light treatment exhibited decreased phototactic responses to light. Our results show that decreased light relaxes selection on anti-predator behaviour. Such trends provide new insights into selection on eye size and behaviour.

Host and brood parasite coevolutionary interactions covary with comparative patterns of the avian visual system.

In coevolutionary arms-races, reciprocal ecological interactions and their fitness impacts shape the course of phenotypic evolution. The classic example of avian host-brood parasite interactions selects for host recognition and rejection of increasingly mimetic foreign eggs. An essential component of perceptual mimicry is that parasitic eggs escape detection by host sensory systems, yet there is no direct evidence that the avian visual system covaries with parasitic egg recognition or mimicry. Here, we used eye size measurements collected from preserved museum specimens as a metric of the avian visual system for species involved in host-brood parasite interactions. We discovered that (i) hosts had smaller eyes compared with non-hosts, (ii) parasites had larger eyes compared with hosts before but not after phylogenetic corrections, perhaps owing to the limited number of independent evolutionary origins of obligate brood parasitism, (iii) egg rejection in hosts with non-mimetic parasitic eggs positively correlated with eye size, and (iv) eye size was positively associated with increased avian-perceived host-parasite eggshell similarity. These results imply that both host-use by parasites and anti-parasitic responses by hosts covary with a metric of the visual system across relevant bird species, providing comparative evidence for coevolutionary patterns of host and brood parasite sensory systems.

Investigating the Predictive Power of Three Potential Sublethal Endpoints for the Fathead Minnow Fish Embryo Toxicity Test: Snout-Vent Length, Eye Size, and Pericardial Edema.

The fish embryo acute toxicity (FET) test is known to be less sensitive than the fish acute test for some chemicals, including neurotoxicants. Thus, there is an interest in identifying additional endpoints that can improve FET test performance. The goal of this project was to advance alternative toxicity testing methods by determining whether select developmental abnormalities-snout-vent length, eye size, and pericardial area-are linked to adverse alterations in ecologically-relevant behaviors and delayed mortality. Fathead minnow (Pimephales promelas) FET tests were conducted with 3,4-dicholoroaniline, cadmium, and perfluorooctanesulfonic acid (PFOS) and developmental abnormalities were quantified. Surviving eleutheroembryos were reared in clean water to 14 days post fertilization (dpf), during which time behaviors and mortality were evaluated. None of the abnormalities evaluated were predictive of behavioral alterations; however, embryos with ≥14% reductions in length or ≥3.54-fold increases in pericardial area had an 80% chance of mortality by 14 dpf. When these abnormalities were used as markers of mortality, the LC50s for cadmium and PFOS were less than those calculated using only standardized FET test endpoints and similar to those obtained via larval fish tests, indicating that the snout-vent length and pericardial area warrant consideration as standard FET test endpoints.

The effect of ecological factors on eye morphology in the western rainbowfish, Melanotaenia australis.

Ecological factors such as spatial habitat complexity and diet can explain variation in visual morphology, but few studies have sought to determine whether visual specialisation can occur among populations of the same species. We used a small Australian freshwater fish (the western rainbowfish, Melanotaenia australis) to determine whether populations showed variation in eye size and eye position, and whether this variation could be explained by environmental (light availability, turbidity) and ecological (predation risk, habitat complexity, invertebrate abundance) variables. We investigated three aspects of eye morphology - (1) eye size relative to body size, (2) pupil size relative to eye size and (3) eye position in the head - for fish collected from 14 sites in a major river catchment in northwest Western Australia. We found significant variation among populations in all three measures of eye morphology, but no effect of sex on eye size or eye position. Variation in eye diameter and eye position was best explained by the level of habitat complexity. Specifically, fish occurring in habitats with low complexity (i.e. open water) tended to have smaller, more dorsally located eyes than those occurring in more complex habitats (i.e. vegetation present). The size of the pupil relative to the size of the eye was most influenced by the presence of surrounding rock formations; fish living in gorge habitats had significantly smaller pupils (relative to eye size) than those occupying semi-gorge sites or open habitats. Our findings reveal that different ecological and environmental factors contribute to habitat-specific visual specialisations within a species.

Changes in key traits versus depth and latitude suggest energy-efficient locomotion, opportunistic feeding and light lead to adaptive morphologies of marine fishes.

Understanding patterns and processes governing biodiversity along broad-scale environmental gradients, such as depth or latitude, requires an assessment of not just taxonomic richness, but also morphological and functional traits of organisms. Studies of traits can help to identify major selective forces acting on morphology. Currently, little is known regarding patterns of variation in the traits of fishes at broad spatial scales. The aims of this study were (a) to identify a suite of key traits in marine fishes that would allow assessment of morphological variability across broad-scale depth (50-1200 m) and latitudinal (29.15-50.91°S) gradients, and (b) to characterize patterns in these traits across depth and latitude for 144 species of ray-finned fishes in New Zealand waters. Here, we describe three new morphological traits, namely fin-base-to-perimeter ratio, jaw-length-to-mouth-width ratio, and pectoral-fin-base-to-body-depth ratio. Four other morphological traits essential for locomotion and food acquisition that are commonly measured in fishes were also included in the study. Spatial ecological distributions of individual fish species were characterized in response to a standardized replicated sampling design, and morphological measurements were obtained for each species from preserved museum specimens. With increasing depth, fishes, on average, became larger and more elongate, with higher fin-base-to-perimeter ratio and larger jaw-length-to-mouth-width ratio, all of which translates into a more eel-like anguilliform morphology. Variation in mean trait values along the depth gradient was stronger at lower latitudes for fin-base-to-perimeter ratio, elongation and total body length. Average eye size peaked at intermediate depths (500-700 m) and increased with increasing latitude at 700 m. These findings suggest that, in increasingly extreme environments, fish morphology shifts towards a body shape that favours an energy-efficient undulatory swimming style and an increase in jaw-length vs. mouth width for opportunistic feeding. Furthermore, increases in eye size with both depth and latitude indicate that changes in both the average ambient light conditions as well as seasonal variations in day-length can act to select ecomorphological adaptations in fishes.

Predation selects for smaller eye size in a vertebrate: effects of environmental conditions and sex.

Increased eye size in animals results in a larger retinal image and thus improves visual acuity. Thus, larger eyes should aid both in finding food as well as detecting predators. On the other hand, eyes are usually very conspicuous and several studies have suggested that eye size is associated with predation risk. However, experimental evidence is scant. In this study, we address how predation affects variation in eye size by performing two experiments using Eurasian perch juveniles as prey and either larger perch or pike as predators. First, we used large outdoor tanks to compare selection due to predators on relative eye size in open and artificial vegetated habitats. Second, we studied the effects of both predation risk and resource levels on phenotypic plasticity in relative eye size in indoor aquaria experiments. In the first experiment, we found that habitat altered selection due to predators, since predators selected for smaller eye size in a non-vegetated habitat, but not in a vegetated habitat. In the plasticity experiment, we found that fish predators induced smaller eye size in males, but not in females, while resource levels had no effect on eye size plasticity. Our experiments provide evidence that predation risk could be one of the driving factors behind variation in eye size within species.

Eye and wing structure closely reflects the visual ecology of dung beetles.

An important resource partitioning strategy allowing dung beetles to coexist in the same habitat, while utilising the same food, is species' separation of activity times. After establishing the diel activity period of three closely related, co-occurring dung beetles, we examined their eye and wing morphology. Absolute and relative eye size, and facet size were greater in the nocturnal Escarabaeus satyrus, followed by the crepuscular Scarabaeus zambesianus and then the diurnal Kheper lamarcki. The diurnal K. lamarcki had the highest wing aspect ratio (long, narrow wings), followed by the crepuscular S. zambesianus and the nocturnal E. satyrus (short, broad wings), suggesting that dim-light active species fly slower than diurnal species. In addition, the two species active in dim light had a lower wing loading than the diurnal species, indicating the need for greater manoeuvrability in the dark. Analyses of wing shape revealed that the diurnal K. lamarcki wing had a proportionally larger jugal and anal region than both dim light species. Our results show that different species of dung beetles have a combination of optical and morphological wing adaptations to support their foraging activities in diverse light conditions.

Ocular size and shape in lens-induced Myopization in young Guinea pigs.

BACKGROUND: Lens-induced myopization in guinea pigs has been used as model for the process of myopization in humans. It has not been explored yet whether the change in globe shape in eyes undergoing myopization is similar in experimental myopia in guinea pigs and in clinical myopia in patients. METHODS: The study included 70 guinea pigs (age:2-3 weeks) equally divided into a study group with lens-induced myopization for 5 weeks, and a control group wearing goggles with no refractive power. The globe diameters were measured using a microcaliper after enucleation. RESULTS: The horizontal globe diameter (9.19 ± 0.15 mm versus 9.15 ± 0.18 mm; P = 0.25) and vertical globe diameter (9.02 ± 0.11 mm versus 8.99 ± 0.14 mm; P = 0.29) did not differ significantly between the study group and control group. The sagittal diameter was significantly longer in the study group (8.96 ± 0.15 mm versus 8.84 ± 0.14 mm; P = 0.001). While the vertical and horizontal globe diameters were correlated with each other in a ratio of approximately 1:1 (non-standardized regression coefficient B:0.94;95% confidence interval (CI):0.73,1.15), the steepness of the regression lines of the associations of both diameters with the sagittal diameter were flatter (horizontal to sagittal diameter: B: 0.64; 95% CI: 0.44,0.83; vertical to sagittal diameter:B:0.55;95% CI:0.41,0.69). Correspondingly, the ratios of horizontal-to-sagittal globe diameter and of vertical-to-sagittal globe diameter decreased (P < 0.001) with longer sagittal diameter. CONCLUSIONS: For each mm axial elongation in young guinea pigs the horizontal globe diameter increased by 0.64 mm (95%CI:0.44,0.83) and the vertical diameter by 0.55 mm (95% CI:0.41,0.69), indicating that the globe enlargement occurred predominantly in the sagittal direction. Axial elongation in guinea pigs led to a similar relative change in ocular shape as in humans.

The evolution of sexual signal modes and associated sensor morphology in fireflies (Lampyridae, Coleoptera).

Animals employ different sexual signal modes (e.g. visual, acoustic, chemical) in different environments and behavioural contexts. If sensory structures are costly, then evolutionary shifts in primary signal mode should be associated with changes in sensor morphology. Further, sex differences are expected if male and female signalling behaviours differ. Fireflies are known for their light displays, but many species communicate exclusively with pheromones, including species that recently lost their light signals. We performed phylogenetically controlled analyses of male eye and antenna size in 46 North American taxa, and found that light signals are associated with larger eyes and shorter antennae. In addition, following a transition from nocturnal light displays to diurnal pheromones, eye size reductions occur more rapidly than antenna size increases. In agreement with the North American taxa, across 101 worldwide firefly taxa in 32 genera, we found light displays are associated with larger eye and smaller antenna sizes in both males and females. For those taxa with both male and female data, we found sex differences in eye size and, for diurnal species, in antenna size.

Non-uniform evolutionary response of gecko eye size to changes in diel activity patterns.

Geckos feature a large range of eye sizes, but what drives this phenotypic diversity is currently unknown. Earlier studies point towards diel activity patterns (DAPs) and locomotory mode, but phylogenetic comparative studies in support of the proposed adaptive mode of eye evolution are lacking. Here, we test the hypothesis of DAPs as the driver of eye size evolution with a dataset on 99 species of gecko. Results from phylogenetic generalized least-square analysis (PGLS) and multivariate model-fitting reveal smaller eyes in diurnal geckos consistent with different phenotypic optima. However, Bayesian analyses of selective regime shifts demonstrate that only two of nine transitions from nocturnal to diurnal activity are coupled with decreases in eye size, and two other regime shifts are not associated with DAP transitions. This non-uniform evolutionary response suggests that eye size is not the only functionally relevant variable. Evolutionary adaptations may therefore include different combinations of several traits (e.g. photoreceptors), all with the same functional outcome. Our results further demonstrate that DAP only partially explains eye size diversity in geckos. As open habitats favour the evolution of large eyes while obstructed habitats favour small eyes, the degree of habitat clutter emerges as another potential axis of eye diversification.

Eye Size, Fovea, and Foraging Ecology in Accipitriform Raptors.

Birds with larger eyes are predicted to have higher spatial resolution because of their larger retinal image. Raptors are well known for their acute vision, mediated by their deep central fovea. Because foraging strategies may demand specific visual adaptations, eye size and fovea may differ between species with different foraging ecology. We tested whether predators (actively hunting mobile prey) and carrion eaters (eating dead prey) from the order Accipitriformes differ in eye size, foveal depth, and retinal thickness using spectral domain optical coherence tomography and comparative phylogenetic methods. We found that (1) all studied predators (except one) had a central and a temporal fovea, but all carrion eaters had only the central fovea; (2) eye size scaled with body mass both in predators and carrion eaters; (3) predators had larger eyes relative to body mass and a thicker retina at the edge of the fovea than carrion eaters, but there was no difference in the depth of the central fovea between the groups. Finally, we found that (4) larger eyes generally had a deeper central fovea. These results suggest that the visual system of raptors within the order Accipitriformes may be highly adapted to the foraging strategy, except for the foveal depth, which seems mostly dependent upon the eye size.

Association between axial length and horizontal and vertical globe diameters.

PURPOSE: To assess relationships between axial length and the horizontal and vertical globe diameters. MATERIAL AND METHODS: The study consisted of enucleated human eyes. The horizontal, vertical, and sagittal diameters were measured. RESULTS: The study included 135 globes removed because of malignant uveal melanoma (111 globes) or end-stage painful glaucoma (n = 24 eyes). Mean axial, horizontal, and vertical diameters were 24.6 ± 2.6 mm (range: 20-35 mm), 23.7 ± 1.4 mm (range: 21-29 mm) and 23.7 ± 1.4 mm (range: 20-29 mm) respectively. The horizontal diameter and vertical diameter did not differ significantly (P = 0.92), while both were significantly (P < 0.001) shorter than the axial diameter. The horizontal diameter was significantly and linearly correlated with the vertical globe diameter (P < 0.001; regression line: vertical globe diameter = 0.84 × horizontal globe diameter + 3.69). The axial diameter was significantly (P < 0.001) associated with the horizontal diameter and vertical diameters in a bipartite manner. In eyes with an axial length ≤24 mm, horizontal and vertical diameters increased by 0.44 and 0.51 mm, respectively, for each mm increase in axial diameter, while in eyes with an axial length >24 mm, the horizontal and vertical globe diameter increased by a lower amount of 0.19 and 0.21 mm, respectively, for each mm increase in axial diameter. CONCLUSIONS: Myopic enlargement of the globe beyond an axial length of 24 mm takes place predominantly in the sagittal axis, leading to a change in the globe form from a sphere to an elongated form. It fits with the notion that myopic elongation may occur by an elongation of the eye walls in regions close to the globe's equator.

Ocular biometry by computed tomography in different dog breeds.

OBJECTIVE: To (i) correlate B-mode ocular ultrasound (US) and computed tomography (CT) (prospective pilot study), (ii) establish a reliable method to measure the normal canine eye using CT, (iii) establish a reference guide for some dog breeds, (iv) compare eye size between different breeds and breed groups, and (v) investigate the correlation between eye dimensions and body weight, gender, and skull type (retrospective study). PROCEDURE: B-mode US and CT were performed on ten sheep cadaveric eyes. CT biometry involved 100 adult pure-bred dogs with nonocular and nonorbital disease, representing eleven breeds. Eye length, width, and height were each measured in two of three planes (horizontal, sagittal, and equatorial). RESULTS: B-mode US and CT measurements of sheep cadaveric eyes correlated well (0.70-0.71). The shape of the canine eye was found to be akin to an oblate spheroid (a flattened sphere). A reference guide was established for eleven breeds. Eyes of large breed dogs were significantly larger than those of medium and small breed dogs (P < 0.01), and eyes of medium breed dogs were significantly larger than those of small breed dogs (P < 0.01). Eye size correlated with body weight (0.74-0.82) but not gender or skull type. CONCLUSIONS: Computed tomography is a suitable method for biometry of the canine eye, and a reference guide was established for eleven breeds. Eye size correlated with breed size and body weight. Because correlation between B-mode US and CT was shown, the obtained values can be applied in the clinical setting, for example, for the diagnosis of microphthalmos and buphthalmos.

Microhabitat and body size effects on heat tolerance: implications for responses to climate change (army ants: Formicidae, Ecitoninae).

1. Models that predict organismal and population responses to climate change may be improved by considering ecological factors that affect species thermal tolerance. Species differences in microhabitat use can expose animals to diverse thermal selective environments at a given site and may cause sympatric species to evolve different thermal tolerances. 2. We tested the hypothesis that species differences in body size and microhabitat use (above- vs. below-ground activity) would correspond to differences in thermal tolerance (maximum critical temperatures: CTmax ). Thermal buffering effects of soil can reduce exposure to extreme high temperatures for below-ground active species. We predicted larger-bodied individuals and species would have higher CTmax and that species mean CTmax would covary positively with degree of above-ground activity. We used Neotropical army ants (Formicidae: Ecitoninae) as models. Army ants vary in microhabitat use from largely subterranean to largely above-ground active species and are highly size polymorphic. 3. We collected data on above- and below-ground temperatures in habitats used by army ants to test for microhabitat temperature differences, and we conducted CTmax assays for army ant species with varying degrees of surface activity and with different body sizes within and between species. We then tested whether microhabitat use was associated with species differences in CTmax and whether microhabitat was a better predictor of CTmax than body size for species that overlapped in size. 4. Microhabitat use was a highly significant predictor of species' upper thermal tolerance limits, both for raw data and after accounting for the effects of phylogeny. Below-ground species were more thermally sensitive, with lower maximum critical temperatures (CTmax ). The smallest workers within each species were the least heat tolerant, but the magnitude of CTmax change with body size was greater in below-ground species. Species-typical microhabitat was a stronger predictor of CTmax than body size for species that overlapped in size. Compared to the soil surface, 10-cm subsoil was a significantly moderated thermal environment for below-ground army ants, while maximum surface raid temperatures sometimes exceeded CTmax for the most thermally sensitive army ant castes. 5. We conclude sympatric species differences in thermal physiology correspond to microhabitat use. These patterns should be accounted for in models of species and community responses to thermal variation and climate change.

Predator-prey interactions, flight initiation distance and brain size.

Prey avoid being eaten by assessing the risk posed by approaching predators and responding accordingly. Such an assessment may result in prey-predator communication and signalling, which entail further monitoring of the predator by prey. An early antipredator response may provide potential prey with a selective advantage, although this benefit comes at the cost of disturbance in terms of lost foraging opportunities and increased energy expenditure. Therefore, it may pay prey to assess approaching predators and determine the likelihood of attack before fleeing. Given that many approaching potential predators are detected visually, we hypothesized that species with relatively large eyes would be able to detect an approaching predator from afar. Furthermore, we hypothesized that monitoring of predators by potential prey relies on evaluation through information processing by the brain. Therefore, species with relatively larger brains for their body size should be better able to monitor the intentions of a predator, delay flight for longer and hence have shorter flight initiation distances than species with smaller brains. Indeed, flight initiation distances increased with relative eye size and decreased with relative brain size in a comparative study of 107 species of birds. In addition, flight initiation distance increased independently with size of the cerebellum, which plays a key role in motor control. These results are consistent with cognitive monitoring as an antipredator behaviour that does not result in the fastest possible, but rather the least expensive escape flights. Therefore, antipredator behaviour may have coevolved with the size of sense organs, brains and compartments of the brain involved in responses to risk of predation.

Light conditions and the evolution of the visual system in birds.

Despite vision being an essential sense for many animals, the intuitively appealing notion that the visual system has been shaped by environmental light conditions is backed by insufficient evidence. Based on a comprehensive phylogenetic comparative analysis of birds, we investigate if exposure to different light conditions might have triggered evolutionary divergence in the visual system through pressures on light sensitivity, visual acuity, and neural processing capacity. Our analyses suggest that birds that have adopted nocturnal habits evolved eyes with larger corneal diameters and, to a lesser extent, longer axial length than diurnal species. However, we found no evidence that sensing and processing organs were selected together, as observed in diurnal birds. Rather than enlarging the processing centers, we found a tendency among nocturnal species to either reduce or maintain the size of the two main brain centers involved in vision-the optic tectum and the wulst. These results suggest a mosaic pattern of evolution, wherein optimization of the eye optics for efficient light capture in nocturnal species may have compromised visual acuity and central processing capacity.