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Responses of female wolf spiders (% of individuals responding with receptivity) of several Schizocosa spp. to isolated visual and vibratory signals from male courtship (see text for details). Based on data from multiple sources [Stratton and Uetz,1983; Scheffer et al., 1996; Hebets and Uetz, 1999; unpubl. obs.].

Responses of female wolf spiders (% of individuals responding with receptivity) of several Schizocosa spp. to isolated visual and vibratory signals from male courtship (see text for details). Based on data from multiple sources [Stratton and Uetz,1983; Scheffer et al., 1996; Hebets and Uetz, 1999; unpubl. obs.].

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Spiders perceive the world using multiple sensory modes, including vibration, vision, and chemical senses, for prey detection and communication. These sensory modes are used in many communication contexts, either individually or in multimodal signaling. Selection for effective signaler-receiver communication and species discrimination is especially...

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... cue-isola- tion studies on a number of species in the S. ocreata clade, and two outgroups (S. duplex and S. saltatrix), demon- strate that these species differ in their reliance on visual and vibratory communication during courtship [Stratton and Uetz, 1983;Scheffer et al., 1996;Hebets and Uetz, 1999]. Although females in both outgroups (saltatrix, duplex) and some of the species within the clade (rovneri, uetzi) respond predominately to vibration cues, females in other clade member species (stridulans, crassipes, ocreata) exhibit equal or greater receptivity to visual cues ( fig.1). In the latter three species, male forelegs have pig- mentation (stridulans, crassipes, ocreata) and/or decora- Responses of female wolf spiders (% of individuals re- sponding with receptivity) of several Schizocosa spp. to isolated visual and vibratory signals from male courtship (see text for details). ...

Citations

... Web spiders especially often transmit vibratory signals through web silk, which can be characterized as using a seismic channel, but spiders that lives away from webs rely on channels that are independent of web silk. For example, vision tends to be the dominant channel used by lycosid and salticid spiders that live away from webs (Uetz and Stratton 1983;Uetz and Roberts 2002;Elias et al. 2005). Several studies have assessed the effects of substrate changes on spider behavior and their communication signals on various substrates (Jackson and Blest 1982;Jackson and Brassington 1987;Michálek et al. 2019). ...
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Changes in the animals’ environment often impact on their behavior and the way they communicate. Particularly in spiders, living on webs or not is a determining factor. While the Lycosidae family predominantly comprises wandering spiders, only a few species inhabit webs. Aglaoctenus oblongus (C.L. Koch 1847) challenges this norm, as it has been found both in webs and wandering, suggesting a previously undocumented mixed strategy. This study aims to: (1) describe the sexual behavior of A. oblongus under laboratory conditions and (2) utilize phenological data to elucidate when and where mating occurs. Sexual encounters were observed within and outside webs, with minimal differences noted between them. Courtship was characterized by groping and leg shaking performed by the males. Copulation was lengthy (near to seven hours), featuring an irregular pattern of palpal insertions, and culminating in a backward dismount. Individuals showed a mixed life history, with the spiders most often found in webs being females with eggs. Both sexes were found throughout the year, with a peak in autumn. We compare the sexual characteristics of this species with that of A. lagotis (also belonging to the Sosippinae subfamily), the only other wolf spider studied to date that exclusively inhabits webs.
... Analyzing these communication patterns can provide insights into the complex web of ecological interactions within an ecosystem. Finally, the field of biomimicry could leverage these discoveries to design novel communication technologies inspired by the sophisticated language of spiders [53][54][55][56][57][58][59][60][61][62][63]. By mimicking how spiders encode complex messages into vibrations, researchers and scientists in the telecommunication, engineering, medical, and possibly other fields might unlock new possibilities for data transmission or even create biocompatible medical devices. ...
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Machine learning (ML) provides a strong way to unraveling the mysteries of spider communication , a complicated system mediated by vibrations. These vibrations include a multitude of information regarding a spider's behavior and state, as evidenced by sophisticated timing patterns and amplitude fluctuations. Traditional approaches, which rely on human feature extraction and categorization, fail to capture the entire richness of this data. Supervised learning systems, trained on labeled vibration data, may automatically categorize signals such as courting displays, territorial defense, and kin identification. Unsupervised learning can reveal hidden patterns in unlabeled data, which may lead to the discovery of novel communication signals or behaviors. Deep learning's capacity to understand complicated correlations from big datasets offers great promise for deciphering the intricate codes embedded in these vibrations. By combining these ML approaches, researchers can acquire unique insights into spider language. This not only transforms our knowledge of spider behavior and communication in behavioral ecology, but it also has the potential to spur advances in biomimetic technology inspired by these intriguing creatures. ARTICLE HISTORY
... Biotremology is indeed a whole field in biology that highlights the importance of multimodal signals in animal behaviour, with a particular attention to the neglected aspect of vibrational communication and orientation [21]. Again, many arachnid species are well known to use vibrations in their foraging [5,15,30,61]. Scorpions even possess an anatomical structure, namely the basitarsal compound slit sensilla that was shown to act as a vibrational receptor [67]. To our knowledge however, no similar structure has been described in V. destructor but it would be interesting to explore it further. ...
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Olfaction is a major sense in Varroa destructor. In natural conditions, it is known that this honey bee parasite relies on kairomones to detect its host or to reproduce. Yet, in artificial conditions, the parasite is able to feed and survive for a few days even though most honey bee pheromones are lacking. Other key cues are thus probably involved in V. destructor perception of its close environment. Here, we used several artificial feeding designs to explore the feeding behaviour of the parasite when it is deprived of olfactory cues. We found that V. destructor is still able to feed only guided by physical cues. The detection of the food source seems to be shape-related as a 3D membrane triggers arrestment and exploration more than a 2D membrane. The tactile sense of V. destructor could thus be essential to detect a feeding site, although further studies are needed to assess the importance of this sense combined with olfaction in natural conditions.
... In spiders, only a handful of studies have studied olfactory and tactile lures, with most studies focusing on chemical lures of bolas spiders (Zhu and Haines, 2004;Vereecken and McNeil 2010). Although non-visual lures might be widespread among spider taxa (Uetz and Roberts 2002;Hill and Wessel 2016;Virant-Doberlet et al. 2019), we currently have limited understanding of whether they are common hunting tactics in spiders or if they involve more complex strategies using multimodal mechanisms in combination with chemical and/or vibratory signals. ...
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Lures and other adaptations for prey attraction are particularly interesting from an evolutionary viewpoint because they are characterized by correlational selection, involve multicomponent signals, and likely reflect a compromise between maximizing conspicuousness to prey while avoiding drawing attention of enemies and predators. Therefore, investigating the evolution of lure and prey-attraction adaptations can help us understand a larger set of traits governing interactions among organisms. We review the literature focusing on spiders (Araneae), which is the most diverse animal group using prey attraction and show that the evolution of prey-attraction strategies must be driven by a trade-off between foraging and predator avoidance. This is because increasing detectability by potential prey often also results in increased detectability by predators higher in the food chain. Thus increasing prey attraction must come at a cost of increased risk of predation. Given this trade-off, we should expect lures and other prey-attraction traits to remain suboptimal despite a potential to reach an optimal level of attractiveness. We argue that the presence of this trade-off and the multivariate nature of prey-attraction traits are two important mechanisms that might maintain the diversity of prey-attraction strategies within and between species. Overall, we aim to stimulate research on this topic and progress in our general understanding of the diversity of predator and prey interactions.
... Playback studies using vibratory signals have been used successfully in an array of arthropod taxa (Cocroft and Rodríguez 2005;Eriksson et al. 2011;Fowler-Finn and Rodríguez 2012;Mazzoni et al. 2009;Rebar and Rodríguez 2013;Schmitt et al. 1994;Uetz and Roberts 2002). While vibratory playback techniques vary (see previously cited examples), we used piezoelectric disc benders (APC International), as used in other studies on S. ocreata Uetz 2016, 2019;Stoffer and Uetz 2017;Uetz et al. 2017;Stoffer et al. 2021). ...
... While vibratory playback techniques vary (see previously cited examples), we used piezoelectric disc benders (APC International), as used in other studies on S. ocreata Uetz 2016, 2019;Stoffer and Uetz 2017;Uetz et al. 2017;Stoffer et al. 2021). Female S. ocreata respond appropriately to vibratory playback of courting males and respond with receptivity displays, even if presented as a unimodal signal (Uetz and Roberts 2002;Uetz et al. 2013Uetz et al. , 2017Uetz 2016, 2019;Stoffer and Uetz 2017). An Apple iPod Touch (4th edition) with vibratory courtship recordings was connected to a Pyle Mini 2 × 40W Stereo Power Amplifier. ...
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While members of the choosier sex often prefer courting mates with bright, large, or loud phenotypes, social experience can result in variation in mate preferences. Fewer studies, though, have investigated how multiple social parameters might interact to affect such preferences. In the brush-legged wolf spider, Schizocosa ocreata, asynchrony of maturation between sexes provides a time period in which females might be exposed to male courtship prior to making a mating decision. We tested whether adult females demonstrated plasticity in their preferences for vibratory signal amplitude after experience with vibratory playback during their penultimate stage. Penultimate instar females were presented unimodal vibratory courtship signals via piezoelectric disc benders, manipulating the perceived encounter rate (every other day or once per day), the number of males (one or two), and/or the vibratory amplitude (low or high). As adults, each female was presented vibratory playback of a low- or high-amplitude courtship signal in both no-choice and two-choice designs. In no-choice trials, previous experience with different amplitude signals significantly affected adult preferences, while other social parameters did not. Specifically, female S. ocreata preferred high-amplitude signals to low-amplitude signals if previously exposed to high-amplitude signals, while those previously exposed to low-amplitude signals preferred low-amplitude signals. In two-choice trials, however, females preferred high-amplitude signals regardless of their previous social experience, suggesting that innate preferences for high-amplitude signals might outweigh any learned preferences in some contexts. Results from this study complement previous social experience studies in S. ocreata, by clearly demonstrating a second sensory modality through which social learning can occur. Significance statement Social experience affects subsequent mate preferences in a variety of taxa, but in many instances, single parameters are examined at a time. Furthermore, in species that use multimodal communication, there remain questions about whether a single sensory modality is sufficient to elicit such plasticity. In this study, we manipulated multiple parameters using vibratory playback to examine whether social experience during the brush-legged wolf spider’s juvenile stage affected preferences for low- and high-amplitude signals as an adult. Ultimately, only the amplitude of the male’s signals that they were exposed to as a juvenile impacted adult mate preference—not the number of perceived males or how often they encountered the males. These results, along with previous studies, demonstrate that S. ocreata is capable of plasticity in response to social cues in multiple modalities.
... Premating communication, however, often integrates two or more modalities. Multimodal courtship signals of spiders from the genus Schizocoza are combinations of chemical, visual, and vibrational communication (Uetz and Roberts 2002;Roberts and Uetz 2005;Moskalik and Uetz 2011;Uetz et al. 2013). A between-species comparison of Schizocoza female responses showed that all species used vibrational communication that was supplemented to a various extent by visual displays and chemicals (Uetz and Roberts 2002). ...
... Multimodal courtship signals of spiders from the genus Schizocoza are combinations of chemical, visual, and vibrational communication (Uetz and Roberts 2002;Roberts and Uetz 2005;Moskalik and Uetz 2011;Uetz et al. 2013). A between-species comparison of Schizocoza female responses showed that all species used vibrational communication that was supplemented to a various extent by visual displays and chemicals (Uetz and Roberts 2002). ...
Chapter
Cryptic species are organisms which look identical, but which represent distinct evolutionary lineages. They are an emerging trend in organismal biology across all groups, from flatworms, insects, amphibians, primates, to vascular plants. This book critically evaluates the phenomenon of cryptic species and demonstrates how they can play a valuable role in improving our understanding of evolution, in particular of morphological stasis. It also explores how the recognition of cryptic species is intrinsically linked to the so-called 'species problem', the lack of a unifying species concept in biology, and suggests alternative approaches. Bringing together a range of perspectives from practicing taxonomists, the book presents case studies of cryptic species across a range of animal and plant groups. It will be an invaluable text for all biologists interested in species and their delimitation, definition, and purpose, including undergraduate and graduate students and researchers.
... Male Schizocosa ocreata wolf spiders engage in courtship behaviours that involve waving and tapping their forelegs displaying tufts of bristles on their forelegs and producing seismic signals. Conspecific females are more likely to respond to males when both sensory modalities are present (Uetz et al., 2000;Uetz & Roberts, 2002) and detect such multimodal stimuli faster (Uetz, Roberts & Taylor, 2009), and as expected, these complex signals also increase risk of detection by predators. Predatory Phidippus clarus jumping spiders orient faster to S. ocreata males when signals include both visual and seismic components compared to only visual ones (Roberts, Taylor & Uetz, 2006). ...
Article
Eavesdropping predators, parasites and parasitoids exploit signals emitted by their prey and hosts for detection, assessment, localization and attack, and in the process impose strong selective pressures on the communication systems of the organisms they exploit. Signallers have evolved numerous anti‐eavesdropper strategies to mitigate the trade‐off between the costs imposed from signal exploitation and the need for conspecific communication. Eavesdropper strategies fall along a continuum from opportunistic to highly specialized, and the tightness of the eavesdropper–signaller relationship results in differential pressures on communication systems. A wide variety of anti‐eavesdropper strategies mitigate the trade‐off between eavesdropper exploitation and conspecific communication. Antagonistic selection from eavesdroppers can result in diverse outcomes including modulation of signalling displays, signal structure, and evolutionary loss or gain of a signal from a population. These strategies often result in reduced signal conspicuousness and in decreased signal ornamentation. Eavesdropping enemies, however, can also promote signal ornamentation. While less common, this alternative outcome offers a unique opportunity to dissect the factors that may lead to different evolutionary pathways. In addition, contrary to traditional assumptions, no sensory modality is completely ‘safe’ as eavesdroppers are ubiquitous and have a broad array of sensory filters that allow opportunity for signal exploitation. We discuss how anthropogenic change affects interactions between eavesdropping enemies and their victims as it rapidly modifies signalling environments and community composition. Drawing on diverse research from a range of taxa and sensory modalities, we synthesize current knowledge on anti‐eavesdropper strategies, discuss challenges in this field and highlight fruitful new directions for future research. Ultimately, this review offers a conceptual framework to understand the diverse strategies used by signallers to communicate under the pressure imposed by their eavesdropping enemies.
... Jumping spiders in the genus Habronattus, for example, are impressive in their coordination of sexually dimorphic ornamentation and associated dynamic visual displays with their complex vibratory songs that can consist of up to 20 elements (Elias et al., 2003(Elias et al., , 2012. Similarly, decades of research on wolf spiders in the genus Schizocosa have explored the form and function of visual (ornamentation and dynamic movements) and vibratory signaling (reviewed in Uetz and Roberts, 2002;Stratton, 2005;Hebets et al., 2013;Uetz et al., 2016;Hebets and McGinley, 2019;Starrett et al., 2022). Despite major advances in our understanding of bimodal signal interactions in select species [e.g., S. ocreata (Scheffer et al., 1996;Uetz et al., 2016); S. uetzi (Hebets, 2005); S. stridulans, ; S. floridana (Rosenthal and Hebets, 2012)], there remains one Schizocosa species where the visual and vibratory signaling appear non-functional from the female's perspective-Schizocosa. retrorsa (Banks, 1911). ...
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Wolf spiders within the genus Schizocosa have become a model system for exploring the form and function of multimodal communication. In terms of male signaling, much past research has focused on the role and importance of dynamic and static visual and substrate-borne vibratory communication. Studies on S. retrorsa, however, have found that female-male pairs were able to successfully mate in the absence of both visual and vibratory stimuli, suggesting a reduced or non-existent role of these signaling modalities in this species. Given these prior findings, it has been suggested that S. retrorsa males may utilize an additional signaling modality during courtship—air particle movement, often referred to as near-field sound—which they likely produce with rapid leg waving and receive using thin filiform sensory hairs called trichobothria. In this study, we tested the role of air-particle movement in mating success by conducting two independent sets of mating trials with randomly paired S. retrorsa females and males in the dark and on granite (i.e., without visual or vibratory signals) in two different signaling environments—(i) without (“No Noise”) and (ii) with (“Noise”) introduced air-particle movement intended to disrupt signaling in that modality. We also ran foraging trials in No Noise/Noise environments to explore the impact of our treatments on overall behavior. Across both mating experiments, our treatments significantly impacted mating success, with more mating in the No Noise signaling environments compared to the Noise environments. The rate of leg waving—a previously assumed visual dynamic movement that has also been shown to be able to produce air particle displacement—was higher in the No Noise than Noise environments. Across both treatments, males with higher rates of leg waving had higher mating success. In contrast to mating trials results, foraging success was not influenced by Noise. Our results indicate that artificially induced air particle movement disrupts successful mating and alters male courtship signaling but does not interfere with a female’s ability to receive and assess the rate of male leg waving.
... These results highlight that crossmodal effects can appear without a mammalian cortex. Given that multi-sensory integration occurs in invertebrates, we believe that cross-modal facilitation effects are likely, indicating that other, radically different, neural architectures can produce such effects [55][56][57]. ...
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Stimulation in one sensory modality can affect perception in a separate modality, resulting in diverse effects including illusions in humans. This can also result in cross-modal facilitation, a process where sensory performance in one modality is improved by stimulation in another modality. For instance, a simple sound can improve performance in a visual task in both humans and cats. However, the range of contexts and underlying mechanisms that evoke such facilitation effects remain poorly understood. Here, we demonstrated cross-modal stimulation in wild-caught túngara frogs, a species with well-studied acoustic preferences in females. We first identified that a combined visual and seismic cue (vocal sac movement and water ripple) was behaviourally relevant for females choosing between two courtship calls in a phonotaxis assay. We then found that this combined cross-modal stimulus rescued a species-typical acoustic preference in the presence of background noise that otherwise abolished the preference. These results highlight how cross-modal stimulation can prime attention in receivers to improve performance during decision-making. With this, we provide the foundation for future work uncovering the processes and conditions that promote cross-modal facilitation effects.
... Despite the lack of a well-supported phylogenetic hypothesis, Schizocosa has developed over the past decades as a model comparative system for studying spider communication and reproductive behavior (Herberstein and Hebets, 2013). Species exhibit extensive differences in morphological characters and behavioral signals used in courtship, making this group powerful for studying characters presumably experiencing strong sexual selection (Fowler-Finn et al., 2015;Hebets and Papaj, 2005;Kozak and Uetz, 2019;Rovner, 1975;Stratton, 2005;Stratron andUetz, 1981, 1986;Uetz and Denterlein, 1979;Uetz and Roberts, 2002;Uetz et al., 2016). Prior to the current study, males of fourteen described Schizocosa species were identified as having dark pigmentation on the first pair of legs, and seven species possess dense bristles on the tibia (Stratton, 2005). ...
... Despite high species diversity (2431; World Spider Catalog, 2020), abundance, ecological importance, and utility as behavioral models Just et al., 2019;Uetz and Roberts, 2002), the lycosid family has received remarkably little targeted attention in phylogenetic studies, particularly studies that employ molecular data (Ivanov et al., 2021;Murphy et al., 2006;Piacentini and Ramírez, 2019;Vink et al., 2002). Our sampling of Lycosidae is mostly restricted to Fig. 9. Ancestral state mapping of ornamentation index. ...
Article
Members of the Nearctic spider genus Schizocosa Chamberlin, 1904 have garnered much attention in behavioral studies and over many decades, a number of species have developed as model systems for investigating patterns of sexual selection and multimodal communication. Many of these studies have employed a comparative approach using putative, but not rigorously tested, sister species pairs that have distinctive morphological traits and attendant behaviors. Despite past emphasis on the efficacy of these presumably comparative-based studies of closely related species, generating a robust phylogenetic hypothesis for Schizocosa has been an ongoing challenge. Here, we apply a phylogenomic approach using anchored hybrid enrichment to generate a data set comprising over 400 loci representing a comprehensive taxonomic sample of 23 Nearctic Schizocosa. Our sampling also includes numerous outgroup lycosid genera that allow for a robust evaluation of genus monophyly. Based on analyses using concatenation and coalescent-based methods, we recover a well-supported phylogeny that infers the following: 1) The New World Schizocosa do not form a monophyletic group; 2) Previous hypotheses of North American species require reconsideration along with the composition of species groups; 3) Multiple longstanding model species are not genealogically exclusive and thus are not “good” species; 4) This updated phylogenetic framework establishes a new working paradigm for studying the evolution of characters associated with reproductive communication and mating. Ancestral character state reconstructions show a complex pattern of homoplasy that has likely obfuscated previous attempts to reconstruct relationships and delimit species. Important characters presumably related to sexual selection, such as foreleg pigmentation and dense bristle formation, have undergone repeated gain and loss events, many of which have led to increased morphological divergence between sister-species. Evaluation of these traits in a comparative framework illuminates how sexual selection and natural selection influence character evolution and provides a model for future studies of multimodal communication evolution and function.