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Santon, M.

Publications and source records attributed to Santon, M..

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Visual modelling validates prey detection by means of diurnal active photolocation in a small cryptobenthic fish

Active sensing has been well documented in animals that use echolocation and electrolocation. Active photolocation, or active sensing using light, has received much less attention, and only in bioluminescent nocturnal species. Recently, however, evidence has suggested the diurnal triplefin Tripterygion delaisi uses controlled iris radiance, termed ocular sparks, for prey-detection. While this form of diurnal active photolocation was behaviourally described, a study exploring the complete physical and theoretical process would provide a more compelling case supporting this mechanism. In this paper, we investigate the conditions under which diurnal active photolocation could assist T. delaisi in detecting potential prey items. In the field, we sampled prey gammarids (Genus Cheirocratus) from foraging substrates, and characterized the spectral properties of their body and eyes, which possess strong reflectors between the ommatidia. In the laboratory, we quantified ocular spark sizes and the angular dependence of their radiance. Together with environmental light measurements and the visual properties of T. delaisi, we modeled diurnal active photolocation under various scenarios. Our results corroborate that diurnal active photolocation can help T. delaisi detect gammarids at distances relevant to foraging, 4.5 cm under favourable conditions and up to 2.5 cm under average conditions. Because ocular sparks are widespread across many different fish species, diurnal active photolocation for micro-prey may be a common predation strategy.

zoology

Diurnal active photolocation enhances predator detection in a marine fish

Active sensing by means of light is rare. In vertebrates, it is known only from chemiluminescent fish with light organs below their pupils, an anatomical arrangement that is ideal to generate eyeshine in the pupils of nearby organisms. Here, we test whether diurnal fish can achieve the same by redirecting sunlight through reflection instead. We recently showed that small (< 5 cm), benthic, marine triplefin fish actively redirect downwelling light using their iris. We hypothesized that this mechanism allows triplefins to improve detection of a cryptic organism by generating eyeshine in its pupil. Here, we tested this by attaching small dark hats to triplefins to shade their iris from downwelling light. Two controls consisted of triplefins with a clear or no hat. These treatments test the prediction that light redirection increases the visual detection ability of triplefins. To this end, we placed treated fish in a tank with a display compartment containing either a stone as the control stimulus, or a scorpionfish, i.e. a cryptic, motionless triplefin predator with retroreflective eyes. After overnight acclimatization, we determined the average distance triplefins kept from the display compartment over two days. Both in the laboratory (n = 15 replicates per treatment) and in a similar field experiment at 15 m depth (n = 43 replicates per treatment) fish kept longer distances from the scorpionfish than from the stone. This response varied between hat treatments: shaded triplefins stayed significantly closer to the scorpionfish in the laboratory and in one of two orientations tested in the field. A follow-up field experiment at 10 m depth revealed the immediate response of triplefins to a scorpionfish. At first, many individuals (n = 80) moved towards it, with shaded triplefins getting significantly closer. All individuals then gradually moved to a safer distance at the opposite half of the tank. Visual modelling supported the experimental results by showing that triplefins can redirect enough light with their iris to increase a scorpionfishs pupil brightness above detection threshold at a distance of 7 cm under average field conditions and at more than 12 cm under favorable conditions. We conclude that triplefins are generally good in the visual detection of a cryptic predator, but can significantly improve this ability when able to redirect downwelling light with their iris and induce eyeshine in the predators pupil. We discuss the consequences of \"diurnal active photolocation\" for visual detection and camouflage among fish species.

zoology

Daytime eyeshine contributes to pupil camouflage in a cryptobenthic marine fish

Ocular reflectors enhance eye sensitivity in dim light, but can produce reflected eyeshine when illuminated. Most teleost fish occlude their reflectors during the day. The opposite is observed in cryptic sit-and-wait predators such as scorpionfish and toadfish, where reflectors are occluded at night and exposed during the day. This results in daytime eyeshine, proposed to enhance pupil camouflage by reducing the contrast between the otherwise black pupil and the surrounding tissue. In this study, we test this hypothesis in the scorpionfish Scorpaena porcus and show that eyeshine is the result of two mechanisms: the previously described Stratum Argenteum Reflected (SAR) eyeshine, and Pigment Epithelium Transmitted (PET) eyeshine, a newly described mechanism for this species. We confirm that the ocular reflector is exposed only when the eye is light-adapted, and present field measurements to show that eyeshine reduces pupil contrast against the iris. We then estimate the relative contribution of SAR and PET eyeshine to pupil brightness. Visual models for different light scenarios in the field show that daytime eyeshine enhances pupil camouflage from the perspective of a prey fish. We propose that the reversed occlusion mechanism of some cryptobenthic predators has evolved as a compromise between camouflage and vision.

evolutionary biology