Chemical investigation of aggregation behaviour in the two-spotted spider mite Tetranychus urticae.

The silk produced by the group-living mite Tetranychus urticae provides group protection and is used as an informative material during habitat settlement, egg laying, mating, and dispersal events. In this context, cues contained in the silk and other materials produced by mites [eggs, black faeces (BF) and white faeces + silk (WFS)] were investigated. Chemical compounds were extracted by hexane or methanol, and choice tests were used to determine the individual attractiveness of each extract. For both solvents, individuals did not respond to the extract from eggs and WFS. BF extracts were attractive for both solvents. After separating the BF methanol extract into four different chemical components using thin layer chromatography, no component was determined to be responsible for mite attraction. This work supports the evidence that the faeces of T. urticae do contain substances that promote behavioural changes. Not particular chemical compounds but combinations of them seem to induce the mites' preference. Moreover, the response of mites to chemicals seemed to be context dependent as mites belonging to populations with different densities differed in their attraction to BF extracts.

Reaching the ball or missing the flight? Collective dispersal in the two-spotted spider mite Tetranychus urticae.

The two-spotted spider mite is a worldwide phytophagous pest displaying a peculiar dispersal. At high density, when plants are exhausted, individuals gather at the plant apex to form a collective silk-ball. This structure can be dispersed by wind or phoresy. Individuals initiating the ball are enclosed in the centre and have a high risk to die. For the first time, the ultimate and proximate mechanisms leading to this group dispersal are examined. To explore if a particular mite genotype was involved in the ball formation, plants were infested with individuals of different genetic background. After the silk-ball formation, the mites in the ball and those remaining on the plant were collected and genotyped. The balls were harvested after 4h and 24h to determine the role of timing between the formation and dispersal on the mortality of mites. Mites do not segregate according to their degree of relatedness, stage, or sex. Mites parallel humans using public transportation: they climb up in the ball whatever their genetic background. Silk-balls composed of unrelated individuals may help avoiding inbreeding when colonizing a new plant. Our results also emphasize the importance of an adequate timing for efficient dispersal between the time spent between ball formation and dispersal.

Settlement decisions by the two-spotted spider mite Tetranychus urticae.

In silk-spinning arthropods, silk can be used for web building, protection, and communication. Silk is an informative material about the presence of conspecifics. It can therefore inform on habitat suitability and hence assist in habitat choice. In this context, we investigated the influence of silk on microhabitat choice by the two-spotted spider mite, Tetranychus urticae. Three factors that could potentially influence habitat choice were manipulated: the strain, number, and the stage of mites. Our study showed that these factors all influence the choice of microhabitat. The tendency of whether to settle on a silk-covered area was influenced by the origin of mites (strain effect). Adult females showed a higher tendency to settle on an area covered with the silk laid by numerous congeners (number effect). Moreover, larvae seemed to be more responsive to the presence of silk than adults (stage effect). This suggests that individuals use silk as a social cue in selecting their microhabitat and that the spatial organization and group behaviour seem to be shaped by the individuals' response to social cues, such as the amount of silk already present.

Should I lay or should I wait? Egg-laying in the two-spotted spider mite Tetranychus urticae Koch.

Optimality theory predicts that females tend to maximize their offspring survival by choosing the egg-laying site. In this context, the use of conspecific cues allows a more reliable assessment of the habitat quality. To test this hypothesis, Tetranychus urticae Koch is an appropriate biological model as it is a phytophagous mite living in group, protected against external aggression by a common web. Experiments were conducted to determine the respective influence of substrate (living substrate: bean leaf vs. non-living substrate: glass plate), silk and presence of conspecific eggs on the egg-laying behavior of T. urticae females. On both living and non-living substrates, the presence of silk positively influenced the probability of a female to lay an egg, but had no influence on the number of eggs deposited. The egg-laying behavior was mainly determined by the nature of the substrate with mites laying fewer eggs on a non-living substrate than on a living one. The presence of a conspecific egg had no impact on either the probability of laying an egg or on the oviposition rate. This study showed a high variability among females in their fecundity and egg-laying performance. The physiology of females (individual fecundity), the egg-laying substrate and to a lesser extent the presence of silk impacted on the decision of spider mites to lay eggs.

Discrimination through silk recognition: the case of the two-spotted spider mite Tetranychus urticae.

Choosing a suitable habitat is a main step in the settlement process, particularly for species having weak movement abilities. Reliable cues are thus needed for habitat selection. In silk-spinning arthropods, silk can be used as a social cue to select an appropriate location. Silk can also provide information on the presence of related or non-related individuals. In this article, we compare the settlement behaviour of two strains of the two-spotted spider mite, Tetranychus urticae, in response to the presence of silk woven by an individual of its own or another strain. We then examined how individuals behaved when confronted with both types of silk (own/another strain) simultaneously. Both strains were sensitive to related silk. Settlement decision for both strains did not differ according to the origin of the silk. Mites used the silk as a communication cue for habitat selection and strain discrimination. Our results provide experimental evidence for the use of multiple social cues in the settlement decision by weaving mites.

Testing for collective choices in the two-spotted spider mite.

Silk is a vector for collective behaviour in many spinning arthropods, including social spiders, social caterpillars, and some spider mites. In this study, the potential for silk-mediated collective choices is evaluated for the two-spotted spider mite Tetranychus urticae. This subsocial mite lives in large colonies on plants, sheltered under a collectively spun silk web. The silk has an attractive and arresting effect. We test whether the silk trails left by the spider mites can give rise to the collective choice of a path. The experiment consists in offering two identical paths to a group of migrating mites. Our results show that the presence of a silk trail influences the mites, but not sufficiently to systematically provoke a collective choice. In order to determine the trail following potential of T. urticae, we parameterise a theoretical trail following model to fit our experiments and those found in the literature. Our prediction is that even after a large number of mites have passed (200), a systematic collective choice of path should not be expected under the tested conditions. Our results, combined with what is known from the literature, allow us to propose a general scenario for the dispersal behaviour of T. urticae.

The silk road of Tetranychus urticae: is it a single or a double lane?

Tetranychus urticae (Acari: Tetranychidae) is a phytophagous mite that forms huge colonies. All active members of a colony (immatures and matures, females and males) spin silken threads. These mites construct a common web that protects the colony from external aggression. The silk coverage is well-known to provide advantages to the colony but very little is known about the characteristics of the threads themselves. Here is the first quantification of the diameter of silken threads spun by two different stages (adult females and larvae) and its relationship with body size of the spinning individuals. Moreover, we observed how silk was deposited on the substrate through their two pedipalps. Threads were observed by means of transmission electron and fluorescence microscopy. Silken threads spun by larvae (0.055 ± 0.018 µm) were significantly thinner than threads spun by adult females (0.111 ± 0.038 µm). In the first step of the silk depositing behaviour, the mite attached the thread to the substrate by putting its pedipalps in contact with the surface (adhesion, double silken threads). When walking, silken threads became detached from the substrate and spitted up (silken threads were free). Finally, silken threads adhered to the surface. The presence of single and double threads makes thread diameter highly variable.

The formation of collective silk balls in the spider mite Tetranychus urticae Koch.

Tetranychus urticae is a phytophagous mite that forms colonies of several thousand individuals. These mites construct a common web to protect the colony. When plants become overcrowded and food resources become scarce, individuals gather at the plant apex to form a ball composed of mites and their silk threads. This ball is a structure facilitating group dispersal by wind or animal transport. Until now, no quantitative study had been done on this collective form of migration. This is the first attempt to understand the mechanisms that underlie the emergence and growth of the ball. We studied this collective behaviour under laboratory conditions on standardized infested plants. Our results show that the collective displacement and the formation of balls result from a recruitment process: by depositing silk threads on their way up to the plant apex, mites favour and amplify the recruitment toward the balls. A critical threshold (quorum response) in the cumulative flow of mites must be reached to observe the emergence of a ball. At the beginning of the balls formation, mites form an aggregate. After 24 hours, the aggregated mites are trapped inside the silk balls by the complex network of silk threads and finally die, except for recently arrived individuals. The balls are mainly composed of immature stages. Our study reconstructs the key events that lead to the formation of silk balls. They suggest that the interplay between mites' density, plant morphology and plant density lead to different modes of dispersions (individual or collective) and under what conditions populations might adopt a collective strategy rather than one that is individually oriented. Moreover, our results lead to discuss two aspects of the cooperation and altruism: the importance of Allee effects during colonization of new plants and the importance of the size of a founding group.

Spatial distribution and inbreeding in Tetranychus urticae.

In group living, species spatial distribution results from responses to environmental heterogeneity and/or mutual interactions between individuals. These mutual interactions can be regulated by genetic and/or epigenetic factors. In this study, we focus on genetic factors and investigate how the spatial distribution of some individuals colonizing a new environment is influenced by inbreeding. Our biological model is Tetranychus urticae, a phytophagous mite considered as a major pest of many cultivated plants. Groups of T. urticae were composed of individuals from successive inbreeding (sister-brother sib-mating). Our results show that the inter-individual distances increase with inbreeding. Indeed, inbreeding level seems to be an important factor affecting the intra-plant spatial distribution of mites. These results confirm that mites have the capability to discriminate their kin and, moreover, that they are able to accurately perceive differences between close relatives from sib-mating lines.