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Adult jumping spiders are known for their extraordinary eyesight and complex, visually guided behaviors, including elaborate communicatory displays, navigational abilities, and prey-specific predatory strategies. Juvenile spiders also exhibit many of these behaviors, yet their visual systems are many times smaller. How do juveniles retain high visually guided performance despite severe size constraints on their visual systems? We investigated developmental changes in eye morphology and visual function in the jumping spider Phidippus audax using morphology, histology, ophthalmoscopy, and optical measurements. We find that juvenile spiders have proportionally larger lenses in relation to their body size than adults. This should alleviate some of the costs of small body size on visual function. However, photoreceptor number in the anterior lateral eyes (ALE) remains constant from early development onward, consistent with a developmental constraint on photoreceptor differentiation. To accommodate these photoreceptors within the diminutive volume of the spiderling cephalothorax, ALE rhabdoms in early life stages are more tightly packed and significantly smaller in diameter and length, properties that expand across development. Lens focal lengths increase as eyes and retinas grow, resulting in a remarkable maintenance of ALE spatial acuity and field-of-view across life stages. However, this maintenance of acuity comes at a sensitivity cost given the small rhabdomal volumes required by space constraints early in life. Taken together, our results indicate that young jumping spiders have eyes already equipped for high acuity vision, but these young spiders may struggle to perform visually demanding behaviors in low-light environments, a notion that warrants further testing.

Spiders are among the world's most species-rich animal lineages, and their visual systems are likewise highly diverse. These modular visual systems, composed of four pairs of image-forming "camera" eyes, have taken on a huge variety of forms, exhibiting variation in eye size, eye placement, image resolution, and field of view, as well as sensitivity to color, polarization, light levels, and motion cues. However, despite this conspicuous diversity, our understanding of the genetic underpinnings of these visual systems remains shallow. Here, we review the current literature, analyze publicly available transcriptomic data, and discuss hypotheses about the origins and development of spider eyes. Our efforts highlight that there are many new things to discover from spider eyes, and yet these opportunities are set against a backdrop of deep homology with other arthropod lineages. For example, many (but not all) of the genes that appear important for early eye development in spiders are familiar players known from the developmental networks of other model systems (e.g., Drosophila). Similarly, our analyses of opsins and related phototransduction genes suggest that spider photoreceptors employ many of the same genes and molecular mechanisms known from other arthropods, with a hypothesized ancestral spider set of four visual and four nonvisual opsins. This deep homology provides a number of useful footholds into new work on spider vision and the molecular basis of its extant variety. We therefore discuss what some of these first steps might be in the hopes of convincing others to join us in studying the vision of these fascinating creatures.

The study of colour in nature has generated insights intofundamental evolutionary and ecological processes, and researchinto colour traits is a rapidly growing field (Kelber & Osorio, 2010).Theongoinginterestinbiologicalcolorationhasinpartbeendrivenby the increased availability of key technologies, including spec-trometry and photography, and concurrent advances in methodsfor analysing colour data, such as visual models (e.g. Endler M Kelber, Vorobyev, & Osorio, 2003; Stevens, Parraga,Cuthill, Partridge, & Troscianko, 2007). While these developmentsare positive for the field, the increasingly complex analyses beingrun on ever greater amounts of data heighten the need forcomprehensive methods reporting and diligent data management(Alsheikh-Ali, Qureshi, Al-Mallah, & Ioannidis, 2011; Nekrutenko T Van Noorden, 2011; Whitlock, 2011). Astudy may be considered truly reproducible when it satisfies threebroad criteria: (1) methods are reported completely, (2) data arepubliclyavailable and archived,and (3) the chain of modification ofraw data is documented and preserved. While completely repro-ducible research (e.g. FitzJohn et al., 2014) is a laudable goal, theconsiderable demands it imposes on researchers means that it willoften, in practice, be unattainable. Nevertheless, even partialreproducibility through the relatively simple practices of completemethodsreportingandpublicdataarchivingisoftremendousvalue.Our aim was to explore the state of reproducibility in the studyofbiological coloration, and to suggest simpleways inwhich it maybe improved. We first outline common methods for studying bio-logical coloration and present guidelines for comprehensivemethods reporting. We then explore how well some of theseimportant criteria have been reported in the literature. We alsoquantify the availability of publicly archived data and code andsuggest some useful tools for increasing the reproducibility ofcolour trait research more broadly.

Insects live in a three-dimensional space, and need to be able to attach to different types of surfaces in a variety of environmental and behavioral contexts. Adult leaf beetles possess great attachment ability due to their hairy attachment pads. In contrast, their larvae depend on smooth pads to attach to the same host plant. We tested friction forces generated by larvae and adults of dock leaf beetles Gastrophysa viridula on different rough surfaces, and found that adults generate much higher attachment to various substrates than larvae, but are more susceptible to completely losing attachment ability on surfaces with “critical” roughness. Furthermore, sex-specific setal morphology has the effect that attachment forces of male adults are generally higher than those of females when adjusted for body weight. The results are discussed in the context of development, ecology, and changing behavioral strategies of successive life stages.

Jumping spiders (Salticidae) are renowned for their high performing visual system. In addition to their prominent forward-facing telescope-like principal eyes, salticids possess two or three pairs of secondary eyes used for wide-angle motion detection. Salticids orient towards relevant sources of motion detected by the secondary eyes, enabling them to inspect the stimulus with their spatially acute principal eyes. The anteriormost pair of secondary eyes, the anterior lateral (AL) eyes, also faces forward and has higher spatial acuity than the other, laterally-facing, secondary eyes. We used small computer-generated targets to elicit orienting saccades from tethered jumping spiders in order to examine the perceptual limits of the AL eyes. We describe the contrast thresholds of male and female spiders, investigate the reaction time between stimulus appearance and initiation of orientation, as well as the minimum distance a stimulus must travel before eliciting a saccade. Our results show that female spiders react to lower contrast stimuli than males and demonstrate that the secondary eyes can detect stimulus displacements considerably smaller than the inter-receptor angle.

SUMMARYJumping spiders, or salticids, sample their environment using a combination of two types of eyes. The forward-facing pair of ‘principal’ eyes have narrow fields of view, but exceptional spatial resolution, while the two or three pairs of ‘secondary’ eyes have wide fields of view and function especially well as motion analysers. Motion detected by the secondary eyes may elicit an orienting response, whereupon the object of interest is examined further using the high-acuity principal eyes. The anterior lateral (AL) eyes are particularly interesting, as they are the only forward-facing pair of secondary eyes. In this study, we aimed to determine characteristics of stimuli that elicit orienting responses mediated by the AL eyes. After covering all eyes except the AL eyes, we measured orienting responses to dot stimuli that varied in size and contrast, and moved at different speeds. We found that all stimulus parameters had significant effects on orientation propensity. When tethered flies were used as prey, we found that visual information from the AL eyes alone was sufficient to elicit stalking behaviour. These results suggest that, in terms of overall visual processing, the relevance of spatial vision in the AL eyes has been underestimated in the literature. Our results also show that female spiders are significantly more responsive than males. We found that hunger caused similar increases in orientation propensity in the two sexes, but females responded more often than males both when sated and when hungry. A higher propensity by females to orient toward moving objects may be related to females tending to experience higher nutritional demands than males.

While adult green dock leaf beetlesGastrophysa viridulause tarsal adhesive setae to attach to and walk on smooth vertical surfaces and ceilings, larvae apply different devices for similar purposes: pretarsal adhesive pads on thoracic legs and a retractable pygopod at the 10th abdominal segment. Both are soft smooth structures and capable of wet adhesion. We studied attachment ability of different larval instars, considering the relationship between body weight and real contact area between attachment devices and the substrate. Larval gait patterns were analysed using high-speed video recordings. Instead of the tripod gait of adults, larvae walked by swinging contralateral legs simultaneously while adhering by the pygopod. Attachment ability of larval instars was measured by centrifugation on a spinning drum, revealing that attachment force decreases relative to weight. Contributions of different attachment devices to total attachment ability were investigated by selective disabling of organs by covering them with melted wax. Despite their smaller overall contact area, tarsal pads contributed to a larger extent to total attachment ability, probably because of their distributed spacing. Furthermore, we observed different behaviour in adults and larvae when centrifuged: while adults gradually slipped outward on the centrifuge drum surface, larvae stayed at the initial position until sudden detachment.

High visual acuity allows parallel processing of distant environmental features, but only when photons are abundant enough. Diurnal tiger beetles (Carabidae: Cicindelinae) have acute vision for insects and visually pursue prey in open, flat habitats. Their fast running speed causes motion blur that degrades visual contrast, forces stop-and-go pursuit and potentially impairs obstacle detection. We demonstrate here that vision is insufficient for obstacle detection during running, and show instead that antennal touch is both necessary and sufficient for obstacle detection. While running, tiger beetle vision appears to be photon-limited in a way reminiscent of animals in low-light habitats. Such animals often acquire wide-field spatial information through mechanosensation mediated by longer, more mobile appendages. We show that a nocturnal tiger beetle species waves its antennae in elliptical patterns typical of poorly sighted insects. While antennae of diurnal species are also used for mechanosensation, they are rigidly held forward with the tips close to the substrate. This enables timely detection of path obstructions followed by an increase in body pitch to avoid collision. Our results demonstrate adaptive mechanosensory augmentation of blurred visual information during fast locomotion, and suggest that future studies may reveal non-visual sensory compensation in other fast-moving animals.

The modular visual system of jumping spiders (Salticidae) divides characteristics such as high spatial acuity and wide-field motion detection between different pairs of eyes. A large pair of telescope-like anterior-median (AM) eyes is supported by 2-3 pairs of 'secondary' eyes, which provide almost 360 degrees of visual coverage at lower resolution. The AM retinae are moveable and can be pointed at stimuli within their range of motion, but salticids have to turn to bring targets into this frontal zone in the first place. We describe how the front-facing pair of secondary eyes (anterior lateral, AL) mediates this through a series of whole-body 'tracking saccades' in response to computer-generated stimuli. We investigated the 'response area' of the AL eyes and show a clear correspondence between the physical margins of the retina and stimulus position at the onset of the first saccade. Saccade frequency is maximal at the margin of AL and AM fields of view. Furthermore, spiders markedly increase the velocity with which higher magnitude tracking saccades are carried out. This has the effect that the time during which vision is impaired due to motion blur is kept at an almost constant low level, even during saccades of large magnitude.

In dynamic locomotory contexts, visual cues often trigger adaptive behaviour by the viewer, yet studies investigating how animals determine impending collisions typically employ either stationary viewers or objects. Here, we describe a dynamic situation of visually guided prey pursuit in which both impending prey contact and escape elicit observable adaptive behaviours in the pursuer, a predatory beetle. We investigated which visual cues may independently control opening and closing of the beetle's jaws during chases of prey dummies. Jaw opening and closing typically occur when prey is within the 60° binocular field, but not at specific distances, angular sizes or time-to-collision. We show that a sign change in the expansion rate of the target image precedes jaw opening (16 ms) and closing (35 ms), signalling to the beetle that it is gaining on the target or that the target is getting away. We discuss the ‘sloppiness' of such variation in the lag of the behavioural response, especially jaw closing, as an adaptation to uncertainty about target position due to degradation of the target image by motion blur from the fast-running beetle.

Summary Jumping spiders (family Salticidae) are masters of miniature vision, achieving higher spatial resolution in relation to body size than any other animal [1]. While most members of this family do not use color in intraspecific communication, several genera serve as emerging examples of rapid evolutionary radiation in sexual display coloration [2]. These include the Australasian Maratus ‘peacock' spiders, and the American genus Habronattus . Males of these genera are often brilliantly colored on body surfaces they showcase to females during elaborate courtship dances (Figure 1A). However, molecular and electrophysiological data suggest that color vision in the acute ‘principal' eyes of most jumping spiders is based on only two types of photosensitive pigment, one sensitive to ultraviolet (UV) light, the other to green light [3–5]. We report here that Habronattus jumping spiders may achieve substantially better color vision via a mechanism previously unknown in spiders: the shifting of sensitivity of a subset of their photoreceptors from green to red via a long-pass filter positioned in their retina. Trichromatic vision resulting from this filter system should markedly enrich these animals' perception of color, including reds, oranges and yellows often found in their courtship displays.