Thierry Hance

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.

Microsatellite markers reveal spatial genetic structure of Tetranychus urticae (Acari : Tetranychidae) populations along a latitudinal gradient in Europe

The genetic structure of populations of the two-spotted spider mite Tetranychus urticae was investigated along a south–north European transect spanning from southern France to The Netherlands. Mites were collected on Urtica dioica in 6 sampling zones. Microsatellite variation at 5 loci revealed considerable genetic variation with an average heterozygozity of 0.49. Significant heterozygote deficiency was found in 7 populations out of the 18 samples analyzed and one of them was completely monomorphic. Tetranychus urticae populations show some level of genetic structuring. First, genetic differentiation between localities (FST estimates) was significant for all comparisons. Second, the analysis of molecular variance, AMOVA, indicates that there is an effect, albeit low (9%), of the locality in accounting for allele frequency variance. Geographic distance emerges as a factor responsible for this genetic structure. The results are discussed in relation to the biological features of the species and the known patterns of migration. Related agronomical issues are addressed.

Group effect on fertility, survival and silk production in the web spinner Tetranychus urticae (Acari: Tetranychidae) during colony foundation

In many vertebrates and invertebrates, individuals reared in isolation show biological modifications compared with those reared in groups of two of more. The spider mite Tetranychus urticae is characterised by a communal organization and displays some forms of cooperative behaviour (aggregation and common web spinning). To evaluate the potential fitness cost or gain of group living, we investigated the effect of being in group on life history parameters (silk production, fecundity, death rate, feeding rate). In this respect, virgin single females and grouped females (two to six individuals) were compared every day for 5 days. Grouped mites produced significantly more web/mite per day (since the second day of experiment) and more eggs/mite per day (since the fourth day of experiment) than single mites. Moreover, single mites had a higher death rate than mites living in groups. However, no difference was found concerning feeding rate. We assume that mites could benefit from the web production of other individuals and invested resources in other activities such as egg production. We showed that these group effects already exist in small groups. In natural conditions with a higher group size, the impact of group living might be stronger for the dynamics of T. urticae populations and the colony foundation.

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.

How to Visualize the Spider Mite Silk

Tetranychus urticae (Acari: Tetranychidae) is a phytophagous mite that forms colonies of several thousand individuals. Like spiders, every individual produces abundant silk strands and is able to construct a common web for the entire colony. Despite the importance of this silk for the biology of this worldwide species, only one previous study suggested how to visualize it. To analyze the web structuration, we developed a simple technique to dye T. urticaesilk on both inert and living substrates. Fluorescent brightener 28 (FB) (Sigma F3543) diluted in different solvents at different concentrations regarding the substrate was used to observe single strands of silk. On glass lenses, a 0.5% dimethyl sulfoxide solution was used and on bean leaves, a 0.1% aqueous solution. A difference of silk deposit was observed depending the substrate: rectilinear threads on glass lenses and more sinuous ones on bean leaves. This visualizing technique will help to carry out future studies about the web architecture and silk used by T. urticae. It might also be useful for the study of other silk‐spinning arthropods. Microsc. Res. Tech. 2009.

Impact of starvation on the silk attractiveness in a weaving mite, Tetranychus urticae (Acari: Tetranychidae)

The two-spotted spider mite Tetranychus urticae is a silk producer known to live in groups. Its silk production plays an important role in protection against external aggressions (predators, rains, etc.). It is also used for group dispersal through the formation of silkballs or as a thread during individual migration by walking. Until now, the role of silk in enhancing migration has been poorly studied. In this paper, the influence of the silken thread presence on T. urticae’s locomotor activity is studied. One virgin female is placed at the centre of a cover glass partially covered by silk. Moving and resting time are studied on the silk or the clean part of the cover glass as a function of the starvation level of mites (fed vs. starved) and the age of the silk (30 vs. 60 vs. 90 vs. 120 vs. 150xa0min). Results show that a fed female spends more time on the silk-covered part than on the clean part as long as the silk is fresh (less than 120xa0min). Thus, the age-related changes in chemical and/or physical properties of the silk influence the spatial distribution of fed mites. Whatever the age of the silk, starved mites spend more time on the clean part of the set-up. Indeed, the silk freshly laid by conspecifics is attractive only for fed mites; starved mites probably prefer sites without silk (and conspecifics). This study shows that that the silk influences the spatial distribution of T. urticae according to its level of starvation and that silk is probably is an indicator of the presence of conspecifics.

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.

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 locomotor activities on sites covered by silk produced by related and unrelated spider mites in Tetranychus urticae (Acari: Tetranychidae)

The two-spotted spider mite Tetranychus urticae is a silk producer known to live in groups with a common silk web that can cover entire plants and protect mites against predators, rain and wind. Silk also plays an important role during collective migration by aerial dispersal or by walking. In this context, we studied the locomotor activity i.e. time in movement, in resting, and in static exploration (probing behaviour) of an individual confronted to clean places and/or places covered by silk coming from its own population or from another. Two populations were tested, one of the red form (Tunisia) and another of the green form (Belgium). The experimental results showed that the presence or the absence of silk did not influence the relative proportion of each behavioural item either for the red or the green population. Individuals of the green form population spent more time moving and less time resting than individuals of the red form population and this, whatever the substrate (red/green silk, clean). Moreover, the silk from the red form population attracted individuals from both populations, whereas the silk produced by the green form population did not attract any individual either from the red or the green form. This surprising result might have been due to a difference in the quantity and/or quality of silk woven by the two forms. We discuss how the differences observed in the behaviour of these two populations may result from differences in their strategy to rapidly increase the population of the colony and reinforce the silk nest.