Body size and sequence of host colonisation predict the presence of acoustic signalling in beetles.

Acoustic communication is widespread in beetles, is often sexually dimorphic, and plays a significant role in behaviours such as premating recognition, courtship, and copulation. However, the factors that determine the presence or absence of acoustic signalling in a given species remain unclear. We examined acoustic communication in bark beetles (Scolytinae) and pinhole borers (Platypodinae), which are two speciose groups with widespread sound production capabilities. We show that body size along with the sequence of host colonisation predict the presence of acoustic communication, and report, for the first time in the animal kingdom, a size limit-1.9 mm-below which acoustic signalling ceases to be present.

Adaptation and Survival of Marine-Associated Spiders (Araneae).

Aquatic environments are an unusual habitat for most arthropods. Nevertheless, many arthropod species that were once terrestrial dwelling have transitioned back to marine and freshwater environments, either as semiaquatic or, more rarely, as fully aquatic inhabitants. Transition to water from land is exceptional, and without respiratory modifications to allow for extended submergence and the associated hypoxic conditions, survival is limited. In this article, we review marine-associated species that have made this rare transition in a generally terrestrial group, spiders. We include several freshwater spider species for comparative purposes. Marine-associated spiders comprise less than 0.3% of spider species worldwide but are found in over 14% of all spider families. As we discuss, these spiders live in environments that, with tidal action, hydraulic forces, and saltwater, are more extreme than freshwater habitats, often requiring physiological and behavioral adaptations to survive. Spiders employ many methods to survive inundation from encroaching tides, such as air bubble respiration, airtight nests, hypoxic comas, and fleeing incoming tides. While airway protection is the primary survival strategy, further survival adaptations include saltwater-induced osmotic regulation, dietary composition, predator avoidance, reproduction, locomotory responses, and adaptation to extreme temperatures and hydrostatic pressures that challenge existence in marine environments.

Joyful by nature: approaches to investigate the evolution and function of joy in non-human animals.

The nature and evolution of positive emotion is a major question remaining unanswered in science and philosophy. The study of feelings and emotions in humans and animals is dominated by discussion of affective states that have negative valence. Given the clinical and social significance of negative affect, such as depression, it is unsurprising that these emotions have received more attention from scientists. Compared to negative emotions, such as fear that leads to fleeing or avoidance, positive emotions are less likely to result in specific, identifiable, behaviours being expressed by an animal. This makes it particularly challenging to quantify and study positive affect. However, bursts of intense positive emotion (joy) are more likely to be accompanied by externally visible markers, like vocalisations or movement patterns, which make it more amenable to scientific study and more resilient to concerns about anthropomorphism. We define joy as intense, brief, and event-driven (i.e. a response to something), which permits investigation into how animals react to a variety of situations that would provoke joy in humans. This means that behavioural correlates of joy are measurable, either through newly discovered 'laughter' vocalisations, increases in play behaviour, or reactions to cognitive bias tests that can be used across species. There are a range of potential situations that cause joy in humans that have not been studied in other animals, such as whether animals feel joy on sunny days, when they accomplish a difficult feat, or when they are reunited with a familiar companion after a prolonged absence. Observations of species-specific calls and play behaviour can be combined with biometric markers and reactions to ambiguous stimuli in order to enable comparisons of affect between phylogenetically distant taxonomic groups. Identifying positive affect is also important for animal welfare because knowledge of positive emotional states would allow us to monitor animal well-being better. Additionally, measuring if phylogenetically and ecologically distant animals play more, laugh more, or act more optimistically after certain kinds of experiences will also provide insight into the mechanisms underlying the evolution of joy and other positive emotions, and potentially even into the evolution of consciousness.

Experimental characterization and automatic identification of stridulatory sounds inside wood.

The propagation of animal vocalizations in water and in air is a well-studied phenomenon, but sound produced by bark and wood-boring insects, which feed and reproduce inside trees, is poorly understood. Often being confined to the dark and chemically saturated habitat of wood, many bark- and woodborers have developed stridulatory mechanisms to communicate acoustically. Despite their ecological and economic importance and the unusual medium used for acoustic communication, very little is known about sound production in these insects, or their acoustic interactions inside trees. Here, we use bark beetles (Scolytinae) as a model system to study the effects of wooden tissue on the propagation of insect stridulations and propose algorithms for their automatic identification. We characterize distance dependence of the spectral parameters of stridulatory sounds, propose data-based models for the power decay of the stridulations in both outer and inner bark, provide optimal spectral ranges for stridulation detectability and develop automatic methods for their detection and identification. We also discuss the acoustic discernibility of species cohabitating the same log. The species tested can be acoustically identified with 99% of accuracy at distances up to 20 cm and detected to the greatest extent in the 2-6 kHz frequency band. Phloem was a better medium for sound transmission than bark.

Jumping spiders do not seem fooled by texture gradient illusions.

Jumping spiders (Salticidae) use exceptional vision, largely mediated by their forward-facing anterior lateral (AL) and anterior medial (AM) eyes, to pounce on prey from a distance. We evaluated depth perception through the use of 'texture density' (depth estimation through surface texture comparisons, with greater distances having higher textural density) in the salticid Trite planiceps. In visual cliff experiments, spiders tended to choose an area with a false 'low drop' over a false 'high drop' with the same texture densities, but showed no preference for either area when presented with substrates with different texture densities at a constant height. This was corroborated when T. planiceps did not avoid jumping over an illusion resembling a trench compared to a no-illusion control pattern. We then selectively occluded both AL and 1 AM eye (monocular treatment), both AL eyes (ALE-occluded binocular treatment) or no eyes (control), and induced spiders to jump across a gap at different heights. Neither control spiders, spiders with binocular cues from the AM eyes or monocular treatment spiders exhibited a height preference. These results suggest that while T. planiceps accurately perceives depth, it does not appear to rely on texture gradients as a depth cue.

Spatial acuity-sensitivity trade-off in the principal eyes of a jumping spider: possible adaptations to a 'blended' lifestyle.

Jumping spiders (Salticidae) are diurnal visual predators known for elaborate, vision-mediated behaviour achieved through the coordinated work of four pairs of camera-type eyes. One pair ('principal' eyes) is responsible for colour and high spatial acuity vision, while three pairs ('secondary' eyes) are mostly responsible for motion detection. Based on its unusual capacity to visually discriminate specific prey in very low, but also under bright light settings, we investigated the structure of the principal and one pair of secondary eyes (antero-lateral eyes) of Cyrba algerina to determine how these eyes achieve the sensitivity, while maintaining spatial acuity, needed to sustain behaviour in low light. Compared to salticids that live in bright light, the principal eyes of C. algerina have a short focal length, and wide contiguous twin rhabdomeres that support optical pooling, overall favouring sensitivity (0.39 µm2), but without fully compromising acuity (12.4 arc min). The antero-lateral eye retinae have large receptors surrounded by pigment granules, providing effective shielding from scattered light. These adaptations may be beneficial for a xeric salticid species with a 'blended' lifestyle: generally living and hunting under stones in the dark, but sometimes venturing above them, in dramatically different light conditions.

Jumping spiders: An exceptional group for comparative cognition studies.

Several non-mutually exclusive hypotheses have been proposed to explain the evolution of cognition in animals. Broadly, these hypotheses fall under two categories: those that pertain to the selective pressures exerted either by sociality or by the ecological niche in which animals live. We review these ideas and then discuss why the highly visual jumping spiders (Salticidae) are excellent models for investigating how cognitive ability evolves. With few exceptions, these behaviorally complex spiders are non-social, making them ideal candidates to explore ideas pertaining to selection based on habitat complexity and selection based on predatory behavior (foraging niche hypotheses). With the exception of Antarctica, salticids are found in all habitats on Earth, ranging from very complex to barren and simple. While many species are generalist predators, a minority also have specialized predatory behavior and prey specialization on dangerous prey, which has been proposed as an explanation for advanced cognitive ability. As this large group has a diversity of habitats in which it lives, diverse predatory behavior, as well as some "social" species, we argue that salticids are ideal candidates for comparative studies to explore the myriad selection factors acting upon a group well known for their cognitive prowess, despite having miniature brains.

Distance assessment of detours by jumping spiders.

To take an indirect route (detour) in order to reach a specific target requires complex cognitive processes. Yet more demanding, from the cognitive point of view, is when the goal is only visible at the beginning of the detour. In spiders from the family Salticidae, vision is a key sensory modality mediating navigation and prey search. Their acute vision allows them to perform complicated detours, possibly as a consequence of the multitude of potential routes in their typically complex 3-dimensional habitats. We used a 4-route choice test, in which routes differed in being either short or long and in the presence or absence of a lure of a prey item, to investigate route assessment in 2 salticid species, Trite planiceps and Marpissa marina. Although both species showed evidence of motivation to follow lured-routes, judging by the number of times they re-oriented toward them while detouring, we found that Trite chose short routes in preference to long routes, but did not prefer the lured-routes. In contrast, Marpissa exhibited random route choice, although it oriented toward lured-routes more often than control routes (lure absent). Our results suggest that decision-making processes about which route to take occurs before embarking on a route, but this is cognitively challenging. Spiders exhibited cognitive limitations in which the lack of visibility of the goal affected success. However, the severity of cognitive limitations depended on species. We suggest that variability in spatial ability across the Salticidae may be related to the habitat complexity inhabited by each species.

Brown marmorated stink bug overwintering aggregations are not regulated through vibrational signals during autumn dispersal.

The brown marmorated stink bug, Halyomorpha halys (Heteroptera: Pentatomidae), is regarded as one of the world's most pernicious invasive pest species, as it feeds on a wide range of economically important crops. During the autumn dispersal period, H. halys ultimately moves to potential overwintering sites, such as human-made structures or trees where it will alight and seek out a final overwintering location, often aggregating with other adults. The cues used during this process are unknown, but may involve vibrational signals. We evaluated whether vibrational signals regulate cluster aggregation in H. halys in overwintering site selection. We collected acoustic data for six weeks during the autumn dispersal period and used it to quantify movement and detect vibrational communication of individuals colonizing overwintering shelters. Both movement and vibrational signal production increased after the second week, reaching their maxima in week four, before decaying again. We found that only males produced vibrations in this context, yet there was no correlation between movement and vibrational signals, which was confirmed through playback experiments. The cues regulating the formation of aggregations remain largely unknown, but vibrations may indicate group size.

Caffeine affects the vigilance decrement of Trite planiceps jumping spiders (Salticidae).

In jumping spiders (Salticidae), the vigilance decrement, or decrease in response to a repeated visual stimulus over time, directly parallels that found in humans. Explanations for the vigilance decrement in the human literature are heavily mentalistic and central nervous system (CNS) based, whereas response decrements in invertebrates are typically thought of as habituation at the sensory periphery. Here we explored whether the salticid vigilance decrement could be CNS modulated by using caffeine, which is a well-known CNS stimulant for both vertebrates and invertebrates. We used paired tests in which Trite planiceps Simon spiders were randomly given a drop of either caffeinated water or distilled water and were then shown dot stimuli presented on monitors. We measured both general "walking," or activity, as a control for physical fatigue, and optomotor responses to the stimuli. We found that the vigilance decrement was significantly shallower (i.e., spiders were more responsive for longer) when spiders were administered caffeine compared with water; furthermore, these spiders were also generally more active throughout the testing period and thus were not physically impaired. Our results suggest that, in at least some invertebrates, CNS modulation of the vigilance decrement is likely. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

First report of luminous stimuli eliciting sound production in weevils.

Light-based stimuli elicited acoustic responses in male Hylesinus aculeatus Say (Curculionidae: Scolytinae: Hylesinina) instantaneously, with 100% reliability. Stridulations were elicited with a white light beam in a dark environment and recorded with an ultrasonic microphone. Acoustic responses were consistent, and, when compared with sounds produced under stressful conditions (i.e. physical stimulation), no significant differences were found. Hylesinus aculeatus possess an elytro-tergal stridulatory organ and acoustic communication is only present in males. This is also the first report of acoustic communication for this species. Instantaneous light-elicited acoustic communication has potential applications in the development of electronic traps and real-time acoustic detection and identification of beetles, border biosecurity, and noise-reduction in acoustic data collection.

Dim-light vision in jumping spiders (Araneae, Salticidae): identification of prey and rivals.

Jumping spiders (family Salticidae) are known for their intricate vision-based behavior during encounters with prey and conspecific individuals. This is achieved using eyes specialized for discerning fine detail, but there has been minimal research on the capacity that salticids might have for visual performance under low ambient light levels. Here, we investigated the capacity of two salticid species, Cyrba algerina from Portugal and Cyrbaocellata from Kenya, to perform two specific visual tasks under low ambient light levels. We used lures made from spiders and midges in prey-identification experiments and mirror images (virtual conspecifics) in rival-identification experiments. These experiments were implemented under a range of ambient light levels (234, 1.35, 0.54, 0.24 cd m-2). In most instances, Calgerina and Cocellata were proficient at performing both of these visual tasks when ambient light was 234 and 1.35 cd m-2, and a minority performed these tasks at 0.54 cd m-2, but none succeeded when the light level was 0.24 cd m-2Cyrbaalgerina and C. ocellata showed vision-based discrimination under low ambient light levels previously associated with nocturnal species.

The effect of stimulus encounter rate on response decrement in jumping spiders.

The inability to maintain signal detection performance with time on task, or response decrement, has been widely studied. In animals with small brains, the ability to filter out repetitive, irrelevant stimuli may prevent the nervous system from being saturated with information. However, animals must be particular to which stimuli they attend and those to ignore, as mistakes may be costly. We explored the effect of inter-stimulus interval (ISI) between repeated presentations of a visual stimulus on the response decrement of the jumping spider Trite planiceps. Jumping spiders are active visual hunters, and moving visual stimuli trigger a readily observable optomotor response. We used either an ISI of 10 or 20 s between 160 stimulus presentations, predicting that a shorter ISI would result in a steeper decrement through cognitive overstimulation, according to the resource depletion theory. While a clear response decrement was seen for both conditions, the shorter ISI resulted in a more dramatic response decrement, aligning with the resource depletion theory posited in the human-based literature.

Vigilance all the way down: Vigilance decrement in jumping spiders resembles that of humans.

The inability to maintain signal detection performance with time on task, or vigilance decrement, is widely studied in people. Despite suggestions that limitations in sustained attention may be a fundamental characteristic of animal cognition, there has been limited research on the vigilance decrement in other animals. We conducted two experiments to explore vigilance in jumping spiders. Our first experiment established that the vigilance decrement, decline in signal detections with time on task, occurs in these spiders in laboratory settings. Our second experiment tested whether this phenomenon was simply the result of habituation of sensory receptors by employing two dishabituation manipulations. Neither dishabituation manipulation appeared to have an effect. Thus, the vigilance decrement in spiders appears to be due to something more than simply peripheral sensory habituation. We suggest that limitations in sustained attention may be a widespread phenomenon among animals that needs addressing when theorising about the vigilance decrement.

Psychophysical investigation of vigilance decrement in jumping spiders: overstimulation or understimulation?

The inability to maintain signal detection performance with time on task, or vigilance decrement, is widely studied in people because of its profound implications on attention-demanding tasks over sustained periods of time (e.g., air-traffic control). According to the resource depletion (overload) theory, a faster decrement is expected in tasks that are cognitively demanding or overstimulating, while the underload theory predicts steeper decrements in tasks that provide too little cognitive load, or understimulation. Using Trite planiceps, a jumping spider which is an active visual hunter, we investigated vigilance decrement to repetitive visual stimuli. Spiders were tethered in front of two stimulus presentation monitors and were given a polystyrene ball to hold. Movement of this ball indicates an attempt to turn towards a visual stimulus presented to a pair of laterally facing (anterior lateral) eyes for closer investigation with high acuity forward-facing (anterior median) eyes. Vigilance decrement is easily measured, as moving visual stimuli trigger clear optokinetic responses. We manipulated task difficulty by varying the contrast of the stimulus and the degree of 'noise' displayed on the screen over which the stimulus moved, thus affecting the signal:noise ratio. Additionally, we manipulated motivation by paired testing of hungry and sated spiders. All factors affected the vigilance decrement, but the key variable affecting decrement was stimulus contrast. Spiders exhibited a steeper decrement in the harder tasks, aligning with the resource depletion theory.

Positive emotional contagion in a New Zealand parrot.

Positive emotional contagions are outwardly emotive actions that spread from one individual to another, such as glee in preschool children [1] or laughter in humans of all ages [2]. The play vocalizations of some animals may also act as emotional contagions. For example, artificially deafened rats are less likely to play than their non-hearing-impaired conspecifics, while no such effect is found for blinded rats [3]. As rat play vocalizations are also produced in anticipation of play, they, rather than the play itself, may act as a contagion, leading to a hypothesis of evolutionary parallels between rat play vocalizations and human laughter [4]. The kea parrot (Nestor notabilis) has complex play behaviour and a distinct play vocalization [5]. We used acoustic playback to investigate the effect of play calls on wild kea, finding that play vocalizations increase the amount of play among both juveniles and adults, likely by acting as a positive emotional contagion.

Audiogram of the kea parrot, Nestor notabilis.

Vocal communication requires the sender to produce a sound, which transmits through the environment and is perceived by the receiver. Perception is dependent on the quality of the received signal and the receiver's frequency and amplitude sensitivity; hearing sensitivity of animals can be tested using behavioural detection tasks, showing the physical limitations of sound perception. Kea parrots (Nestor notabilis) were tested for their ability to hear sounds that varied in terms of both frequency and amplitude by means of a simple auditory detection task. Audiograms for three kea were similar, with the region of highest sensitivity (1-5 kHz) corresponding to the frequency of the highest amplitude in kea calls. Compared with other parrots and other bird taxa, the overall shape of the kea audiogram follows a similar pattern. However, two potentially interesting differences to the audiograms of other birds were found: an increase in sensitivity at approximately 12 kHz and a decreased sensitivity to frequencies below 1 kHz.

Innate pattern recognition and categorization in a jumping spider.

The East African jumping spider Evarcha culicivora feeds indirectly on vertebrate blood by preferentially preying upon blood-fed Anopheles mosquitoes, the vectors of human malaria1, using the distinct resting posture and engorged abdomen characteristic of these specific prey as key elements for their recognition. To understand perceptual categorization of objects by these spiders, we investigated their predatory behavior toward different digital stimuli--abstract 'stick figure' representations of Anopheles constructed solely by known key identification elements, disarranged versions of these, as well as non-prey items and detailed images of alternative prey. We hypothesized that the abstract images representing Anopheles would be perceived as potential prey, and would be preferred to those of non-preferred prey. Spiders perceived the abstract stick figures of Anopheles specifically as their preferred prey, attacking them significantly more often than non-preferred prey, even when the comprising elements of the Anopheles stick figures were disarranged and disconnected from each other. However, if the relative angles between the elements of the disconnected stick figures of Anopheles were altered, the otherwise identical set of elements was no longer perceived as prey. These data show that E. culicivora is capable of making discriminations based on abstract concepts, such as the hypothetical angle formed by discontinuous elements. It is this inter-element angle rather than resting posture that is important for correct identification of Anopheles. Our results provide a glimpse of the underlying processes of object recognition in animals with minute brains, and suggest that these spiders use a local processing approach for object recognition, rather than a holistic or global approach. This study provides an excellent basis for a comparative analysis on feature extraction and detection by animals as diverse as bees and mammals.

Mediation of a plant-spider association by specific volatile compounds.

Evarcha culicivora, an East African jumping spider (Salticidae), is the only spider for which there is evidence of innate olfactory affinity for particular plant species. Evarcha culicivora also actively chooses as preferred prey the females of Anopheles mosquitoes, and both sexes of Anopheles are known to visit plants for nectar meals. Here, we identified compounds present in the headspace of one of these species in Kenya, Lantana camara, and then used 11 of these compounds in olfactometer experiments. Our findings show that three terpenes [(E)-ß-caryophyllene, α-humulene and 1,8 cineole] can be discriminated by, and are salient to, E. culicivora. The spiders experienced no prior training with plants or the compounds we used. This is the first experimental demonstration of specific phytochemicals being innately attractive to a spider, a group normally characterized as predators.

Hyperacute motion detection by the lateral eyes of jumping spiders.

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.