Anne E. Wignall

Honeybees (Apis mellifera) holding on to memories: response competition causes retroactive interference effects

Five experiments on honeybees examined how the learning of a second task interferes with what was previously learned. Free flying bees were tested for landmark-based memory in variations on a paradigm of retroactive interference. Bees first learned Task 1, were tested on Task 1 (Test 1), then learned Task 2, and were tested again on Task 1 (Test 2). A 60-min delay (waiting in a box) before Test 2 caused no performance decrements. If the two tasks had conflicting response requirements, (e.g., target right of a green landmark in Task 1 and left of a blue landmark in Task 2), then a strong decrement on Test 2 was found (retroactive interference effect). When response competition was minimised during training or testing, however, the decrement on Test 2 was small or nonexistent. The results implicate response competition as a major contributor to the retroactive interference effect. The honeybee seems to hold on to memories; new memories do not wipe out old ones.

Assassin bug uses aggressive mimicry to lure spider prey

Assassin bugs (Stenolemus bituberus) hunt web-building spiders by invading the web and plucking the silk to generate vibrations that lure the resident spider into striking range. To test whether vibrations generated by bugs aggressively mimic the vibrations generated by insect prey, we compared the responses of spiders to bugs with how they responded to prey, courting male spiders and leaves falling into the web. We also analysed the associated vibrations. Similar spider orientation and approach behaviours were observed in response to vibrations from bugs and prey, whereas different behaviours were observed in response to vibrations from male spiders and leaves. Peak frequency and duration of vibrations generated by bugs were similar to those generated by prey and courting males. Further, vibrations from bugs had a temporal structure and amplitude that were similar to vibrations generated by leg and body movements of prey and distinctly different to vibrations from courting males or leaves, or prey beating their wings. To be an effective predator, bugs do not need to mimic the full range of prey vibrations. Instead bugs are general mimics of a subset of prey vibrations that fall within the range of vibrations classified by spiders as ‘prey’.

Male courtship vibrations delay predatory behaviour in female spiders

During courtship, individuals transfer information about identity, mating status and quality. However, male web-building spiders face a significant problem: how to begin courting female spiders without being mistaken for prey? Male Argiope spiders generate distinctive courtship vibrations (shudders) when entering a females web. We tested whether courtship shudders delay female predatory behaviour, even when live prey is present in the web. We presented a live cricket to females during playbacks of shudder vibrations, or white noise, and compared female responses to a control in which we presented a live cricket with no playback vibrations. Females were much slower to respond to crickets during playback of shudder vibrations. Shudder vibrations also delayed female predatory behaviour in a related spider species, showing that these vibrations do not simply function for species identity. These results suggest that male web-building spiders employ a phylogenetically conserved vibratory signal to ameliorate the risk of pre-copulatory cannibalism.

The Influence of Vibratory Courtship on Female Mating Behaviour in Orb-Web Spiders (Argiope keyserlingi, Karsch 1878)

Web-building spiders are important models for sexual selection. While our understanding of post-copulatory mechanisms including sperm competition and cryptic female choice is considerable, our knowledge of courtship and how it influences male and female mating decisions is still extremely poor. Here, we provide the first comprehensive description of male courtship behaviour and vibrations generated in the web by the orb-web spider, Argiope keyserlingi – a recognised model species. We identified three main elements of male courtship: shudders, abdominal wags and mating thread dances (including both plucks and bounces). The vibrations generated by these behaviours are described in detail. Male shuddering behaviour appears to have a strong influence on female latency to mate acceptance, with males that shudder at high rates without compromising shudder duration being preferred. Shuddering behaviour may also mediate female aggressive behaviour, with males that generate long shudders less likely to be cannibalised after copulation. Male abdominal wagging behaviour, however, appears to have only limited influence on female mating decisions. This study provides avenues for future work that synthesises pre- and post-copulatory mechanisms in web-building spiders to generate an all-encompassing model of how sexual selection operates.

Alternative predatory tactics of an araneophagic assassin bug (Stenolemus bituberus)

Predators of dangerous prey risk being injured or killed in counter-attacks and hence may use risk-reducing predatory tactics. Spiders are often dangerous predators to insects, but for a few, including Stenolemus bituberus assassin bugs, web-building spiders are prey. Despite the dangers of counter-attack when hunting spiders, there has been surprisingly little investigation of the predatory tactics used by araneophagic (spider-eating) insects. Here, we compare the pursuit tendency, outcome and predatory tactics of S. bituberus against five species of web-building spider. We found that S. bituberus were most likely to hunt and capture spiders from the genus Achaearanea, a particularly common prey in nature. Capture of Achaearanea sp. was more likely if the prey spider was relatively small, or if S. bituberus was in poor condition. S. bituberus used two distinct predatory tactics, ‘stalking’, in which they slowly approached the prey, and ‘luring’, in which they attracted spiders by manipulating the web to generate vibrations. Tactics were tailored to the prey species, with luring used more often against spiders from the genus Achaearanea, and stalking used more often against Pholcus phalangioides. The choice of hunting tactic used by S. bituberus may reduce the risk posed by the prey spider.

Biology and life history of the araneophagic assassin bug Stenolemus bituberus including a morphometric analysis of the instars (Heteroptera, Reduviidae)

Stenolemus bituberus is an araneophagic “assassin bug” (Heteroptera; Reduviidae; Emesinae) that typically is found living in spider webs. We documented the life history of S. bituberus in the field for 13 months, determining its developmental phenology and microhabitat characteristics as well as describing its mating and predatory behaviour. We also included a morphometric analysis of the instars. Our study revealed that S. bituberus is univoltine with five juvenile instars. Although S. bituberus is found in the webs of spiders from a wide range of genera, it is found most commonly in the webs of Achaearanea, Badumna, Pholcus, and Stiphidion. Multiple juveniles often were found on a single web, but adults tended to be more solitary. Stenolemus bituberus appears to have two distinct hunting strategies: “stalking”, in which they slowly approach the spider, make contact and then strike; and “luring”, in which they attract the spider within range by manipulating the silk with their legs.

Dangerous mating systems: signal complexity, signal content and neural capacity in spiders.

Spiders are highly efficient predators in possession of exquisite sensory capacities for ambushing prey, combined with machinery for launching rapid and determined attacks. As a consequence, any sexually motivated approach carries a risk of ending up as prey rather than as a mate. Sexual selection has shaped courtship to effectively communicate the presence, identity, motivation and/or quality of potential mates, which help ameliorate these risks. Spiders communicate this information via several sensory channels, including mechanical (e.g. vibrational), visual and/or chemical, with examples of multimodal signalling beginning to emerge in the literature. The diverse environments that spiders inhabit have further shaped courtship content and form. While our understanding of spider neurobiology remains in its infancy, recent studies are highlighting the unique and considerable capacities of spiders to process and respond to complex sexual signals. As a result, the dangerous mating systems of spiders are providing important insights into how ecology shapes the evolution of communication systems, with future work offering the potential to link this complex communication with its neural processes.

Predatory behaviour of an araneophagic assassin bug

Assassin bugs from the genus Stenolemus (Heteroptera, Reduviidae) are predators of web-building spiders. However, despite their fascinating lifestyle, little is known about how these insects hunt and catch their dangerous prey. Here we characterise in detail the behaviour adopted by Stenolemus bituberus (Stål) during encounters with web-building spiders, this being an important step toward understanding this species’ predatory strategy. These bugs employed two distinct predatory tactics, “stalking” and “luring”. When stalking their prey, bugs slowly approached the prey spider until within striking range, severing and stretching threads of silk that were in the way. When luring their prey, bugs attracted the resident spider by plucking and stretching the silk with their legs, generating vibrations in the web. Spiders approached the luring bug and were attacked when within range. The luring tactic of S. bituberus appears to exploit the tendency of spiders to approach the source of vibrations in the web, such as might be generated by struggling prey.

Introduction : spider biology

The introductory chapter has been written especially for readers unfamiliar with the finer details of spider systematics, terminology and biology. The introduction is by no means intended to be a complete account of spider biology, which can be found in the excellent Biology of Spiders by Rainer Foelix (1996). Instead, here we concentrate on those aspects of spider biology that prepare the reader for the behavioural chapters to follow. The sections on systematics, fossil record and evolutionary milestones will help place the various behaviours discussed into an evolutionary context. The biology section will familiarise the reader with the spider-specific terminology and reveal some of the peculiarities of spiders: did you know that in modern spiders females have two separate copulatory openings and that spiders can produce up to seven different types of silk? For readers already familiar with spiders, the introduction offers a succinct and up-to-date summary of spider biology. Scope of this book The aim of this book is to illustrate the incredible diversity and often bewildering complexity of spider behaviour. Researchers that regularly work with spiders are well aware of their behavioural potential, and yet spiders still surprise us constantly with behaviours and phenomena that are intriguing, often bizarre and uncommon in other animals. Here we aim to enthuse readers that may have not considered spiders as models for behavioural studies, perhaps assuming that they are limited in their behavioural repertoires.

The sterile male technique: Irradiation negatively affects male fertility but not male courtship

The sterile male technique is a common method to assign paternity, widely adopted due to its relative simplicity and low cost. Male sterility is induced by exposure to sub lethal doses of chemosterilants or irradiation, the dosage of which has to be calibrated for every species to provide successful male sterilisation, without affecting male physiology and behaviour. While the physiological effects of sterilisation are usually assessed for each study, the behavioural ones are rarely analysed in detail. Using the orb web spider Argiope keyserlingi as a model we first tested (1) the validity of the thread assay, which simulates male courtship behaviour in a standardised context, as a proxy representing courtship on a female web. We then investigated (2) the effectiveness of male sterilisation via irradiation and (3) its consequences on male courtship behaviour. Our results validate the thread assay and the sterile male technique as legitimate tools for the study of male courtship behaviour and fertilisation success. We show that these techniques are time and cost effective and reduce undesirable variation, thereby creating opportunities to study and understand the mechanisms underlying sexual selection.