{"paper_id":"4d4d22af-702b-4c55-9be0-5702ea696884","body_text":"Ambulatory dispersal of Typhlodromus (Anthoseius) recki Wainstein (Acari: Phytoseiidae) along Solanceae stem | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Ambulatory dispersal of Typhlodromus (Anthoseius) recki Wainstein (Acari: Phytoseiidae) along Solanceae stem Marie-Stephane Tixier, Amandine Raeckelboom, Lou Tabary, Martial Douin, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4223917/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 01 Aug, 2024 Read the published version in Experimental and Applied Acarology → Version 1 posted You are reading this latest preprint version Abstract Tomato crops are attacked by several pests, including mites. While the main predatory mites are not effective enough to control mite pests due to problems with plant dispersal and establishment (associated with glandular trichomes - GT - on leaves and stems), recent studies have shown encouraging results with the European endemic phytoseiid Typhlodromus ( Anthoseius ) recki . The first objective of the present study was to assess the ability of this species to disperse along the tomato stem, considering six genotypes of Solanum lycopersicum , S. peruvianum and S. cheesmaniae with contrasted trichome numbers and types of stem trichomes, accuratetly characterised in a previous study. As morphological variation in body size has been observed within the species T . ( A. ) recki , the second objective was to determine how predator morphological traits can explain dispersal along the tomato stem. For this, ambulatory dispersal ability of females was tested in lab conditions on the eight Solanum genotypes. Then, the females were mounted on slides and body dimensions measured. No effect of the tomato genotypes was observed on the dispersal ability of the predator. However, specimens that succeeded in crossing the stem, had a higher mobility time than those that failed. Furthermore, body width at midbody and dorsal shield length were negatively correlated with dispersal ability. This suggests that the more slender and relatively small the specimens, the more are mobile and able to successfully cross the stem, regardless of the plant genotype considered. The number of glandular trichomes type (GT) VI and to a lesser extent GT I and IV, and non-glandular trichomes (NGT) II&III appear to limit dispersal. The GT VI seems to have a repellent effect, inducing mite escape in some cases. On the opposite, the number of NGT V were positively correlated with high mobility and stem crossing rates. Assuming that the main barrier to biological control efficiency is dispersal along tomato stems, the results obtained here should have implications for biological control success. The proportion of mites with 'optimal dimensions' appears to be low and further studies should be undertaken to better assess the proportion of mites with such ideal dimensions in different populations and also to determine whether these morphological traits are associated with different feeding abilities and/or abiotic conditions. Solanaceae mites biological control mite body size trichome Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction Tomato crops are attacked by several pests, including mite pests as Tetranychus urticae (Koch), T. evansi Baker & Pritchard and Aculops lycopersici (Tryon). Their control is still mainly based on pesticide application, especially sulfur (with partial efficiency), because of the inability of their main natural enemies (predatory mites of the family Phytoseiidae) to cope with tomato trichomes and thus to disperse and then feed on prey (Van Haren et al. 1987 ). Trichomes are well known plant defense mechanisms that limit herbivory (i.e., Glas et al. 2012 , Bar and Shtein 2019 , Karabourniotis et al. 2020 ). Among species of the genus Solanum as well as among genotypes of a given plant species, trichome types are very diverse (i.e., glandular (GT) vs non glandular (NGT), small vs large, secreting toxic and/or repellent compounds) (i.e., Simmons and Gurr 2005 ). Numerous studies have been carried out on the effect of tomato leaf trichomes on pests, generally showing negative effects of GTs, especially types I and IV, which affect pest survival both through physical effects (trapping on sticky trichome tips) and toxic and repellent effects due to the production of acylsucroses or terpenoids (Snyder et al. 1998 , Chatzivalsileiadis et al. 1999, Kroumova and Wagner 2003 , Fridman et al. 2005 , Simmons and Gurr 2005 , Economou et al. 2006 , Treutter 2006 , Gershenzon and Dudareva 2007 , Alba et al. 2009 , Glas el al. 2012, Dias et al. 2013 , Lucini et al. 2015 , Andrade et al. 2017 , Rakha et al. 2017 , Karabourniotis et al. 2020 ). Only a lower number studied the impact of such plant structures on natural enemies. Nevertheless, several authors have shown clear detrimental effects on several species of the predatory mite family Phytoseiidae. These authors emphasised the barrier effect of trichomes on the stem, preventing dispersal of predators throughout the plant and thus hampering biological control. GTs VI, which release toxic substances on contact with the mite, are the main types of trichomes that alter phytoseiid dispersal (Van Haren et al. 1987 , Nihoul 1994 , Cedola et al. 2001 , Kennedy 2003 , Sato et al. 2011 , Van Houten et al. 2013 , Davidson et al. 2016 , Paspati et al. 2021 , Castañé et al. 2022 , Pandey et al. 2023 ). As a result, none of the commercial Phytoseiidae mite species are effective enough to control mite pests on tomatoes, leading to numerous attempts to develop new biological control solutions. The Phytoseiidae mite family comprises more than 2,500 species world wide (Demite et al. 2024 ). Some of these species are currently reported in Solanaceae, particularly in the genus Solanum , suggesting that these mites may be adapted to plant defenses. Most of these predator species are reported in the Neotropical region, where the genus Solanum originated, but also where the diversity of Phytoseiidae is the highest (Tixier et al. 2020a ; Duarte et al. 2021 ). Because of potential problems related to ecological risks associated to the introduction of exotic species, studies have recently focused on endemic species from Europe collected on Solanaceae. Three species were particularly highly reported- Typhlodromus ( Typhlodromus ) athiasae Athias-Henriot, essentially occuring in Israel; Phytoseiulus persimilis Athias-Henriot, because of mass-release in tomato crops of this commercialised species; and Typhlodromus ( Anthoseius ) recki Wainstein widely reported all over Europe (Tixier et al. 2020a , Demite et al. 2024 ). Recent laboratory studies have shown that the latter species is able to feed on tomato mite pests and is therefore considered a promising predator (Tixier et al. 2020b , Ersin et al. 2021 ). The first objective of this study is to assess the ability of this species to disperse ambulatory along the tomato stem, considering six varieties of Solanum lycopersicum , S. peruvianum and S. cheesmaniae with contrasting types and densities of trichomes on stem. Some authors hypothezised that dispersal ability on tomato stem can be associated to mite size (Pijnakker et al. 2022 ). Observations of many specimens of T . ( A .) recki have revealed large variations in body size (Tixier et al. 2021 ), suggesting a large intraspecific polymorphism, as has been observed in other Phytoseiidae species whose body dimensions vary with season, temperature and diet (Chant 1955 , Tixier et al. 2003 , Walzer and Schausberger 2011 , Walzer et al. 2020 ). Therefore, the second objective of the present study was to determine whether the body size of T . ( A .) recki could be related to the success of the mite in crossing the plant stem, for the eight tomato genotypes considered. Material & methods Plant material. Eight genotypes were studied: Solanum cheesmaniae (LA1412), Solanum peruvianum (Peru CMV) and six varieties of S. lycopersicum (747, Wooly, Yellow Bells, Clomimbo, Lancaster, Hairless). These genotypes were the same as those previously used for assessing the effects of trichomes on leaf, stem and petiole on T. urticae and T . ( A .) recki (Tabary et al. 2024 ). To obtain the experimental plants, seeds were sown in plastic pots (8 x 8 x 8 cm) in a greenhouse (25 ± 2°C, 40 ± 30% RH) and watered twice a week until the beginning of the experiment (when plants reached three leaves). Mite Material. The mite stock colony of T. (A.) recki was initiated from populations (50–100 specimens) collected on Phlomis fruticosa L. (lamiaceae) at Saint Clément-de-Rivière (Hérault, France, 43.694144, 3.851792) on July 2021. Predatory mites were reared following the process described by McMurtry and Scriven ( 1965 ), in rearing units constituted of plastic arenas (10 x 15 cm) deposited on moistened sponge support, surrounded by water-saturated paper tissue for avoiding mite escape. The population was maintained in a climatic chamber (25 ± 2°C, 16D:8N photoperiod, 70 ± 30% RH) and fed with commercialized Typha angustifolia pollen (Nutrimite®) twice a week. Trichome characterization . The trichome characterization followed the protocol used by Tabary et al. ( 2024 ) on the same plant material. Briefly, a piece of 1 cm of stem between the second and third leaf was cut with a razor blade, and on two areas of 4 mm 2 the types and number of trichomes were evaluated using a stereomicroscope (Nikon SMZZ1500 (x20)). Two replications were performed per plant, and 5 plants of each genotype were evaluated (in total 10 replicates / genotype). The trichome type classification proposed by Luckwill ( 1943 ) and Channarayappa et al. (1992) were used. The trichomes types I, IV, V and VI were considered separately, while types II and III were grouped together due to their similarity and the lack of studies demonstrating their role in arthropod resistance (McDowell et al. 2011 ). Trichome types VII and VIII were not taken into account due to their rarity on the eight genotypes studied. Experimental design . A 5 cm stem piece, located between the second and third leaf, was cut with a razor blade and then disposed on wet cotton on a Petri dish. A bean leaf disc (2.5 cm diameter, cv. Contender) with pollen of T. angustifolia (Nutrimite®) was disposed at one extremity, whereas a small piece of black plastic was placed at the other extremity (Fig. 1 ). One young mated female of T . ( A .) recki was then disposed on the plastic piece. Once the female has dispersed on the stem, the plastic piece was removed. At least 10 replicates were carried out for each genotype. The age of predatory females was the same for each replicate. Observations were carried out at four different times: 10 minutes, 25, 55 and 100 minutes after the predator introduction. At each observation time, the predator was filmed for 5 minutes (camera Leica EZ4 W) and several parameters were assessed during the video observations: (i) the success of stem crossing : number of predators reaching the bean leaf and not turning back, (ii) hesitation behavior : the number of times the mite goes on and turns back on the stem without totally crossing it, (iii) escape behavior: number of predators retrieved on the wet cotton support and (iv) the mobility vs immobility duration. At the end of the experiment, the females were mounted on slides in Hoyer’s medium. The mites were then observed using a microscope (Leica DLMB, Leica Microsystèmes SAS, Rueil Malmaison, France) (400 x magnification), and body dimensions were measured. The Dorsal Shield Length (DSL) was assessed by measuring the distances between the setae j1 and J5 . Three Dorsal Shield Width (DSWs) were assessed: (i) at the fore hind part measuring the distances between the setae z4 (DSW1), (ii) at the median part measuring the distances between the setae s6 (DSW2) and (iii) at the backward part measuring the distances between the setae S5 (DSW3) (Fig. 2 ). Typhlodromus ( A . ) recki body size phenotypes . Tixier et al. ( 2021 ) carried measurements of 91 females of T . ( A .) recki collected on six plant species in France and in Italy. Among the characters considered, these authors measured the dorsal shield length and although the measurements were not taken at exactly the same positions on the body as in the present study, DSL (measured at the edge of the body tegument in Tixier et al. ( 2021 ) and between setae j1 and J5 in the present study) and DSW1 (measured at the edge of the body tegument at the level of setae z3 in Tixier et al. ( 2021 ) and between setae z4 in the present study), we decided to use this dataset to determine whether the populations under consideration had variable body sizes and the proportion of “small” individuals in each of these populations. Statistical analyses . All analyses were carried out using R Cote Team (2021). ANOVA were carried out to assess the mean differences in mobility time and hesitation numbers between the Solanum genotypes and between the females that succeed to cross or not the stem. ANOVA were also carried out to determine based on the dataset of Tixier et al. ( 2021 ) differences in DSL and DSW1 between the populations of T . ( A .) recki here considered. ANOVA was carried out using the Kruskal-Wallis test (package “rstatix”) followed by Dunn test when the data did not follow a Normal law and/or with different variances, overwise ANOVA tests (package “car”) followed by Newman & Keuls test (package “Agricolae”) were performed. A multifactorial analysis was used to assess the relationships between trichome numbers, mite body size and mobility time variables, using the packages “FactoMineR” and “factoextra”. Correlation tests (corr.test) were also carried out to determine the relationship between the three dorsal shield dimensions using the package “ggpubr”. Results Trichome characterization of the Solanum genotypes . The multifactorial analysis, which explains 35.2% and 23.6% of the variability for axes 1 and 2 respectively, shows that stem GT VI and NGT V differentiate the genotypes most (Fig. 3 a, b). On the axis 1, the density of these two trichome types were negatively correlated to the density of GT I, NGT II&III and GT IV on the axis 1. On the axis 2, the density of trichome GT IV, GT VI and NGT II&III was positively correlated to each other and differentiated from that of trichome GT I and NGT V. The genotypes were included into two main groups: ( i ) comprising Solanum peruvianum , S. cheesmaniae and S. lycopersicum cv. Hairless: characterized by high densities of NGT V, GT VI and low densities of trichomes GT I, NGT II&III and GT IV, and ( ii ) comprising S. lycopersicum cv. Wooly, cv. Yellow Bells and 747, characterized by opposite density of these latter trichome types. The cultivar Clomimbo and Lancaster had an intermediate position. Typhlodromus ( A . ) recki ambulatory dispersal ability . Predatory mites succeeded to cross the stem for five genotypes among the eight considered. However, the success crossing rate was low, except for S. peruvianum (Fig. 4 ). The mean numbers of “hesitation” was not different between genotypes ( P = 0.73). No significant difference was either observed in the numbers of “hesitation” between the females that succeed to cross or not the stem ( P = 0.56). The percentage of mobility duration was not significantly different between the eight genotypes, all along the experiment and for each date except at 10 minutes, where a higher mobility activity was observed on S. peruvianum . At all times measured, the time spent moving was longer for females that succeeded to cross the stem than for those that did not (Fig. 5 ). A multifactorial analysis was carried out, considering the mean values of trichome stem density of each genotype, dispersal predator parameters and its body size. The axes 1 and 2 represented 58.7% of the total variability of the system (Fig. 6 ). Mobility time variables correlated with each other, as well as with hesitation numbers and crossing success rates, suggesting that the more mobile females were, the more hesitation numbers (forwards and backwards) were high, and the more stem crossing success was important. These walking parameters were positively and negatively correlated with the number of trichome NGT V and GT VI, respectively, suggesting that the density of GT VI had a negative effect on dispersal, whereas the density of NGT V had no effect. In a lesser extent, the numbers of trichomes GT I, NGT II&III and GT IV were also negatively correlated with the mobility ability, suggesting that the numbers of these trichome types may also act as a barrier for predator ambulatory dispersal along the stem. The escape rate was positively associated to the number of GT VI, but not to the densities of GT I, NGT II&III and GT IV, suggesting that the presence of trichomes GT VI might be repellent for the predator. Predator mortality is not related to stem trichome characteristics and only little explains the variability in the system. Dispersal parameters were negatively correlated with dorsal shield length (DSL) and width, especially at the central body part (DSW2), suggesting that the smaller the mite length and width, the higher the mobility time, number of hesitations and crossing success. Finally, mites with a rather small and narrow body size were associated with stems with a low number of GT VI and, to a lesser extent, with GT I, NGT II&III and GT IV, suggesting that mites with such dimensions would be better able to cope with tomato trichome defenses. Variability of T . ( A . ) recki body dimensions. The analyses were conducted on the dataset produced by Tixier et al. ( 2021 ). The mean DSL and DSW were significantly different between the populations of T . ( A .) recki and the plants where they were collected. The lower dimensions were observed for specimens collected on Cirsium arvense L. (Compositae) and Echium vulgare L. (Boraginaceae), whereas the highest were observed from specimens collected on S. lycopersicum , Datura sp. (Solanceae) and Mentha suaveolens L. (Lamiaceae); the specimens collected on P. fruticosa showing an intermediate position (Fig. 7 ). These measurements were not exactly the same as those considered here, as they were taken at the boundary of the body and not between the setae, so their values are higher. Nevertheless, the Fig. 8 shows a correlation between these two measurements (R = 0.53, P < 0.0001). The proportion of small and narrow specimens was the highest for specimens collected on C. arvense and E. vulgare . Discussion The present study indicates that a relationship exists between types and numbers of trichomes on the stem and the ambulatory dispersal behavior of T . ( A .) recki , observed during a short period of time (110 minutes). The NGT V on the stem was not associated to plant defense and high numbers of this trichome type even favor the mobility of the predator, as already observed for various pest species (Onyambus et al. 2011 , Keskin and Kumral 2015 , Savi et al. 2019 ). The density of GT VI affected the most T . ( A .) recki mobility. This result is consistent with observations already reported for other Phytoseiidae mite species (Van Haren et al. 1987 , Paspati et al. 2021 ), whereas contrasted effects of the density of GT VI on the leaf were observed for the prey T. urticae (Onyambus et al. 2011 , Rakha et al. 2017 , Savi et al. 2019 , Kortbeek et al. 2021 ). For example, in Phytoseiidae mites, Paspati et al. ( 2021 ) showed detrimental effects of GT VI on the dispersal of the mite Amblyseius swirskii Athias-Henriot due to the toxic effect of acylsugars causing (i) suffocation due to acylsugar accumulation at mite cuticle openings and (ii) disruption of the subcuticular membrane (Puterka et al. 2003 ). Here, the mortality of the predator was not different between plant genotypes, regardless of the number of GT VI. The cuticle of T . ( A .) recki is quite sclerotized (Livshitz and Kuznetsov 1972 ) compared to that of other Phytoseiidae species, and especially A. swirskii . This morphological trait could explain the lack of effect of trichome exudates on the mortality of T . ( A .) recki , but the relatively short observation period could also explain the low mortality here observed. Further experiments will help to confirm if T . ( A .) recki can cope with the toxicity of acylsucrose. The density of GT VI was also correlated to the numbers of hesitation and escape rates. A repellent effect, causing the mites to move on and then turn back, could be hypothesized, as has already been observed for pest mites (Maluf et al. 2001 ) and Phytoseiidae mites (Sato et al. 2011 ). The density of the other trichome types (GT I and IV, and NGT II&III) was also associated with low mobility and stem crossing, but to a lesser extent than GT VI. The GT I and IV are reported to be detrimental for pests when present on leaves (i.e., Simmons and Gurr 2005 ). Here, GT I and IV do not seem to be related to escape. This suggests that, unlike trichome VI, they do not have a repellent effect. The effect of NGT II&III was rather unexpected, as studies even ignore this trichome type when examining the effects of trichomes density on arthropods. Most of these later studies concern the leaf surface, we can thus hypothesize a different effect of these trichomes when present on the stem. However, we cannot rule out the possibility that such an effect of NGT II&III may also be due to an artefact, since plants with high densities of GT I and IV also have high densities of NGT II&III. The present study provides new information on the dispersal behavior of predators, showing that mobility time and \"hesitation\" activity (walking forward and backward) are both related to stem crossing success. This suggests that the predator is trying to cross the stem in an incisive way, and the more it tries, the more it succeeds. Van Haren et al. ( 1987 ) showed that the dispersal of predators is facilitated when the pest mites have crossed the stem, as the passage of the prey causes trichome breakage and loss of toxicity over time. Van Houten et al. ( 2013 ) reported that dispersal of A. swirskii is favored on stems attacked by A. lycopersici , due to trichome collapse after the pest passage. To our knowledge, whether such trichome collapse and loss of toxicity occurs after predator passage is not known. It would therefore be interesting to determine whether the \"stubbornness\" of T . ( A .) recki individuals observed in this study could be linked to a strategy aimed at facilitating the dispersal of the rest of the colony, by carrying out similar experiments with a large number of specimens of the predator. Although only a small proportion of the total number of T . ( A .) recki examined managed to cross the stem (15%), there were correlations between mite size and crossing success / mobility activities. The mite size parameters that were the most related to stem crossing were the dorsal shield length and the mid-body with and to a lesser extent fore hind and back widths. These results seem to show that the smaller and narrower the mite specimens, the higher the stem crossing success. Several authors have in the past suggested a relationship between mite size and ability to squeeze through trichomes, especially for the tydeid mite Pronematus ubiquitus (McGregor) (Ilionidae), and for some mite pests as the eriophyid mite A. lycopersici (Pijnakker et al. 2022 ). However, these mite species are much smaller than T . ( A .) recki (even than specimens that succeeded to cross the stem). The Phytoseiidae mites studied until now are usually bigger than T . ( A .) recki , especially for A. swirskii and P. persimilis, P. longipes and P. macropilis . In addition, these predatory mites issued from commercial rearings are usually well fed, so their size is significant and their ability to spread through trichomes is likely to be difficult. It will therefore be necessary to test the success of stem crossing as a function of body size in other mite species and for other stages such as immatures and males. Tixier et al. ( 2021 ) showed different morphological phenotypes of T . ( A .) recki depending on the plants from which the populations were collected. Using this latter dataset, differences in the length and width of the dorsal shield were observed between these populations, with the smallest specimens found more frequently in the populations collected on E. vulgare and C. arvense , the population collected on P. fruticosa (the one used for the present experiments) having an intermediate position. It would therefore be interesting to carry out experiments similar to the present ones, using the other T . ( A .) recki populations, in order to validate the hypothesis that the size of the mites and the success of stem-crossing are related, i.e. that the mite crossing rate is higher for mites collected on E. vulgare and C. arvense and lower for mites collected on M. suaveolens , S. lycopersicum and Datura sp. Conclusion The present study confirms the detrimental role of stem trichome density in the dispersal of Phytoseiidae mites, especially GT VI, which also seems to have a repellent effect. It also shows that females try hard to cross the stem, and the more mobile they are, the higher the success rate. These results open new avenues for biocontrol, in particular to determine whether this behavior can be linked to trichome collapse, which subsequently might facilitate the passage of other specimens. Noteworthy relationships between predator body size and stem crossing success were also found. The proportion of stem-crossing specimens within the population studied here is quite low, but other populations, than the one currently considered, appear to have a higher proportion of small specimens. Thus, additional studies would be required to test the stem crossing success of specimens from these latter populations. Several factors can affect mite size, as climatic conditions and plant phenotypes (Chant 1955 , Tixier et al. 2003 , Walzer et al. 2020 ). Thus, although the present study highlights new potential traits for selecting the predator phenotype best adapted for dispersal along the tomato stem, further experiments are needed to determine how much predator size is labile and how much is determined by environmental factors. It would also be necessary to determine the extent to which feeding behavior differs between small and large predator phenotypes for issues of biological control efficacy. Finally, this study mainly focuses on mite size, but other characters may also be involved in crossing success, as mite dorsal sclerotization which is particularly well developed in T . ( A .) recki , in contrast to the other Phytoseiidae species tested so far. Declarations Author Contribution Marie-Stéphane Tixier, Maria Navajas and Denise Navia defined the experimental design. 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Kortbeek RWJ, Galland MD, Muras A, van der Kloet FM, André B, Heilijgers M, van Hijum SAFT, Haring MA, Schuurink RC, Bleeker PM (2021) Natural variation in wild tomato trichomes; selecting metabolites that contribute to insect resistance using a random forest approach. BMC Plant Biol 21(1):315. doi: 10.1186/s12870-021-03070-x Kroumova AB, Wagner GJ (2003) Different elongation pathways in the biosynthesis of acyl groups of trichome exudate sugar esters from various solanaceous plants. Planta 216: 1013–1021. Livshitz IZ, Kuznetsov NN (1972) Phytoseiid mites from Crimea (Parasitiformes: Phytoseiidae) [in Russian] – In : Pests and diseases of fruit and ornamental plants. Proceed. All-Union V. I. Lenin Academy of Agricultural Science, The State Nikita Botanical Gardens, 61: 13-64. Lucini T, Marcos VF, Cristhiane R, Juliano TVR, João RFO (2015) Acylsugar and the role of trichomes in tomato genotypes resistance to Tetranychus urticae . Arthropod-Plant Interact 9: 45–53. Luckwill LC (1943) The genus Lycopersicon : An historical biological and taxanomic survey of the wild and cultivated tomatoes. Aberdeen University Press, U.K., 44 pp. Maluf WR, Campos GA, Cardoso MDG (2001) Relationships between trichome types and spider mite ( Tetranychus evansi ) repellence in tomatoes with respect to foliar zingiberene contents. Euphytica121: 73–80. McDowell ET, Kapteyn J, Schmidt A, Li C, Kang J, Descour A, Shi F, Larson M, Schilmiller A, An L, Jones A, Pichersky E, Soderlund CA, Gang DR (2011) Comparative functional genomic analysis of Solanum glandular trichome types. Plant Physiol 155: 524–539. McMurtry JA, Scriven GT (1965) Insectary production of phytoseiid mites. J Econ Entomol 58: 282-284. doi:10.1093/jee/58.2.282 Nihoul P (1994) Phenology of glandular trichomes related to entrapment of Phytoseiulus persimilis A.-H. in the glasshouse tomato. J Hort Sci 69(5): 783-789. Onyambus GK, Maranga RO, Gitonga LM, Knapp M (2011) Host plant resistance among tomato accessions to the spider mite Tetranychus evansi in Kenya. Exp Appl Acarol 54: 385–393. Pandey S, Biscaia Ribeiro da Silva AL, Dutta B, Chong JH, Mutscher MA, Schmidt JA (2023) Acylsugar tomato lines suppress whiteflies and Amblyseius swirskii establishment. Entomol Exp Appl online first, doi:10.1111/eea.13342 Paspati A, Rambla Nebot JL, López-Gresa MP, Arbona V, Gómez-Cadenas A, Granell Richart A, González-Cabrera J, Urbaneja A (2021). Tomato trichomes are deadly hurdles limiting the establishment of Amblyseius swirskii Athias-Henriot (Acari: Phytoseiidae). Biol Contr 157: 1-9. doi:10.1016/j.biocontrol.2021.104572 Pijnakker J, Moerkens R, Vangansbeke D, Duarte M, Bellinkx S, Benavente A, Merckx J, Stevens I, Wäckers F (2022) Dual Protection: A Tydeoid Mite Effectively Controls Both a Problem Pest and a Key Pathogen in Tomato. Pest Manag Sci 78: 355–361. Puterka GJ, Farone W, Palmer T, Barrington A (2003) Structure-function relationships affecting the insecticidal and miticidal activity of sugar esters. J Econ Entomol 96: 636–644. R Core Team (2021) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/. Rakha M, Bouba N, Ramasamy S, Regnard J-L, Hanson P (2017 ). Evaluation of wild tomato accessions ( Solanum spp.) for resistance to two-spotted spider mite ( Tetranychus urticae Koch) based on trichome type and acylsugar content. Genet Res Crop Evol 64: 1011–1022. doi: 10.1007/s10722-016-0421-0 Sato MM, Moraes GJ, Haddad ML, Wekesa VW (2011) Effect of trichomes on the predation of Tetranychus urticae (Acari: Tetranychidae) by Phytoseiulus macropilis (Acari: Phytoseiidae) on tomato, and the interference of webbing. Exp Appl Acarol 54: 21–32. doi: 10.1007/s10493-011-9426-8 Savi PJ, Moraes GJ, Boiça AL Jr, Melville CC, Carvalho RF, Lourençao AL, Andrade DJ (2019) Impact of leaflet trichomes on settlement and oviposition of Tetranychus evansi (Acari: Tetranychidae) in African and South American tomatoes. Syst Appl Acarol 24(12): 2559-2576. Simmons AT, Gurr GM (2005) Trichomes of Lycopersicon species and their hybrids: effects on pests and natural enemies. Agric Forest Entomol 7: 265–276. Snyder JC, Simmons AM, Tracker RR (1998) Attractancy and oviposition response of type IV trichome density on leaves of Lycopersicon hirsutum grown in three day-lenght regimes. J Entomol Sci 33: 270-281. Tabary L, Navia D, Auger P, Migeon A, Navajas M, Tixier M-S (2024) Plant, pest and predator interplay: Tomato trichomes effects on Tetranychus urticae (Koch) and the predatory mite Typhlodromus (Anthoseius) recki Wainstein. Exp Appl Acarol - submitted Tixier M-S, Kreiter S, Cheval B, Auger P (2003) Morphometric variation between populations of Kampimodromus aberrans (Oudemans) (Acari: Phytoseiidae): Implications for the taxonomy of the genus. Inv syst 17: 349 – 358. Tixier M-S, Douin M, Kreiter S (2020a) Phytoseiidae (Acari: Mesostigmata) on Plants of the Family Solanaceae: Results of a Survey in the South of France and a Review of World Biodiversity. Exp Appl Acarol 81: 357-388. Tixier M-S, Douin M, Rocio O, Gonzalez L, Pount B, Kreiter S (2020b) Distribution and biological features of Typhlodromus ( Anthoseius ) recki (Acari: Phytoseiidae) on Tetranychus urticae , T. evansi (Acari: Tetranychidae) and Aculops lycopersici (Acari: Eriophyidae). Acarologia 60: 684–697. doi:10.24349/acarologia/20204396. Tixier M-S, Perez Martinez S, Douin M (2021)Markers of life history traits: variation in morphology, molecular and amino acid sequences within Typhlodromus ( Anthoseius ) recki Wainstein (Acari: Mesostigmata: Phytoseiidae). Biol J Linn Soc132: 53–73. https://doi.org/10.1093/biolinnean/blaa103 Treutter D (2006) Significance of flavonoids in plant resistance: A review. Environ. Chem Lett 4: 147–157. Doi : 10.1007/s10311-006-0068-8 Van Haren R, Steenhuis M, Sabelis MW, De Ponti O (1987) Tomato stem trichomes and dispersal success of Phytoseiulus persimilis relative to its prey Tetranychus urticae . Exp Appl Acarol 3: 115–121. van Houten YM, Glas JJ, Hoogerbrugge H, Rothe J, Bolckmans KJ, Simoni S, van Arkel J, Alba JM, Kant MR, Sabelis MW (2013) Herbivory-associated degradation of tomato trichomes and its impact on biological control of Aculops lycopersici . Exp Appl Acarol 60: 127-38. doi: 10.1007/s10493-012-9638-6 Walzer A, Schausberger P (2011) Sex-specific developmental plasticity of generalist andspecialist predatory mites (Acari: Phytoseiidae) inresponse to food stress. Biol J Linn Soc102: 650–660. Walzer A, Formayer H, Tixier M-S (2020) Evidence of trans-generational developmental modifications induced by simulated heat waves in an arthropod. Sci Rep 10: 4098. https://doi.org/10.1038/s41598-020-61040-z Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 01 Aug, 2024 Read the published version in Experimental and Applied Acarology → Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {\"props\":{\"pageProps\":{\"initialData\":{\"identity\":\"rs-4223917\",\"acceptedTermsAndConditions\":true,\"allowDirectSubmit\":true,\"archivedVersions\":[],\"articleType\":\"Research Article\",\"associatedPublications\":[],\"authors\":[{\"id\":289369780,\"identity\":\"dd6fef00-448f-43a0-ad6b-5d9827f35b7b\",\"order_by\":0,\"name\":\"Marie-Stephane Tixier\",\"email\":\"data:image/png;base64,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\",\"orcid\":\"\",\"institution\":\"L'Institut Agro Montpellier\",\"correspondingAuthor\":true,\"prefix\":\"\",\"firstName\":\"Marie-Stephane\",\"middleName\":\"\",\"lastName\":\"Tixier\",\"suffix\":\"\"},{\"id\":289369781,\"identity\":\"f2f3e3a3-304f-4cdc-a43f-df3c01d22e87\",\"order_by\":1,\"name\":\"Amandine Raeckelboom\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"National Research Institute for Agriculture, Food and Environment\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Amandine\",\"middleName\":\"\",\"lastName\":\"Raeckelboom\",\"suffix\":\"\"},{\"id\":289369782,\"identity\":\"4270ade2-a8ee-410b-9e50-e4b974d61697\",\"order_by\":2,\"name\":\"Lou Tabary\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"L'Institut Agro Montpellier\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Lou\",\"middleName\":\"\",\"lastName\":\"Tabary\",\"suffix\":\"\"},{\"id\":289369783,\"identity\":\"42ffc093-a952-4d8b-822f-4b32b69cfcdd\",\"order_by\":3,\"name\":\"Martial Douin\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"L'Institut Agro Montpellier\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Martial\",\"middleName\":\"\",\"lastName\":\"Douin\",\"suffix\":\"\"},{\"id\":289369784,\"identity\":\"8abd2009-3353-4bb8-b568-35f033e46b36\",\"order_by\":4,\"name\":\"Maria Navajas\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"National Research Institute for Agriculture, Food and Environment\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Maria\",\"middleName\":\"\",\"lastName\":\"Navajas\",\"suffix\":\"\"},{\"id\":289369785,\"identity\":\"12f7c018-f5b2-4782-9026-614b8d7f1ed0\",\"order_by\":5,\"name\":\"Denise Navia\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"National Research Institute for Agriculture, Food and Environment\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Denise\",\"middleName\":\"\",\"lastName\":\"Navia\",\"suffix\":\"\"}],\"badges\":[],\"createdAt\":\"2024-04-05 15:44:10\",\"currentVersionCode\":1,\"declarations\":\"\",\"doi\":\"10.21203/rs.3.rs-4223917/v1\",\"doiUrl\":\"https://doi.org/10.21203/rs.3.rs-4223917/v1\",\"draftVersion\":[],\"editorialEvents\":[{\"content\":\"https://doi.org/10.1007/s10493-024-00946-z\",\"type\":\"published\",\"date\":\"2024-08-01T15:57:09+00:00\"}],\"editorialNote\":\"\",\"failedWorkflow\":false,\"files\":[{\"id\":54519091,\"identity\":\"931f1e20-be80-4806-873c-14e3457c4e8b\",\"added_by\":\"auto\",\"created_at\":\"2024-04-11 17:36:21\",\"extension\":\"png\",\"order_by\":1,\"title\":\"Figure 1\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":129169,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eSchema showing the experimental design to study the movement of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003eon the stem of six varieties of \\u003cem\\u003eS. lycopersicum\\u003c/em\\u003e, and \\u003cem\\u003eS. cheesmaniae\\u003c/em\\u003eand \\u003cem\\u003eS. peruvianum\\u003c/em\\u003e\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"floatimage1.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4223917/v1/3247a89bbbc598ec16322c5d.png\"},{\"id\":54519094,\"identity\":\"b1a1926b-9a66-4c8e-9bda-83325dbc537a\",\"added_by\":\"auto\",\"created_at\":\"2024-04-11 17:36:21\",\"extension\":\"png\",\"order_by\":2,\"title\":\"Figure 2\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":1542533,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eSchema showing body dimensions measured on specimens of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"floatimage3.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4223917/v1/3554536d24a0090b4013ef56.png\"},{\"id\":54519294,\"identity\":\"55f4bf8a-86ae-49cb-853e-c8c00f36719c\",\"added_by\":\"auto\",\"created_at\":\"2024-04-11 17:44:21\",\"extension\":\"png\",\"order_by\":3,\"title\":\"Figure 3\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":353769,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eResults of the multifactorial analysis considering trichome density on stem of on the six \\u003cem\\u003eSolanum lycopersicum\\u003c/em\\u003evarieties, \\u003cem\\u003eS. peruvianum\\u003c/em\\u003e, \\u003cem\\u003eS. cheesmaniae\\u003c/em\\u003e : (a) correlation circle and (b) biplot showing both variables and the projection of the genotypes considered.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"floatimage5.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4223917/v1/877ec5c295be4958dc6804b2.png\"},{\"id\":54519092,\"identity\":\"61d8cee3-0d22-487e-beb1-f6fabb082f3e\",\"added_by\":\"auto\",\"created_at\":\"2024-04-11 17:36:21\",\"extension\":\"png\",\"order_by\":4,\"title\":\"Figure 4\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":105999,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePercentage of females having or not crossed the stem of the six varieties of \\u003cem\\u003eS. lycopersicum\\u003c/em\\u003e, and \\u003cem\\u003eS. cheesmaniae\\u003c/em\\u003e and \\u003cem\\u003eS. peruvianum\\u003c/em\\u003e\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"floatimage6.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4223917/v1/9d2dabeacae05cd6780ade2d.png\"},{\"id\":54519090,\"identity\":\"07fcaea1-1d07-441d-a9af-483abb2e1530\",\"added_by\":\"auto\",\"created_at\":\"2024-04-11 17:36:21\",\"extension\":\"png\",\"order_by\":5,\"title\":\"Figure 5\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":121727,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eMean number of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e crossing or not the stem during an observation period of: (a) 10 minutes, (b) 25 minutes, (c) 55 minutes and (d) 100 minutes after the predator introduction and (e) for the cumulated observation time (20 minutes).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"floatimage8.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4223917/v1/db62ed2b5ed8f9d72595a711.png\"},{\"id\":54519298,\"identity\":\"d3a9b21e-f9c1-4df5-bd7f-4b96def393ab\",\"added_by\":\"auto\",\"created_at\":\"2024-04-11 17:44:22\",\"extension\":\"png\",\"order_by\":6,\"title\":\"Figure 6\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":364618,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eResults of the multifactorial analysis considering the mean dispersal variables of \\u003cem\\u003eT. (A.) recki, \\u003c/em\\u003ethe mean size of\\u003cem\\u003e T. (A.) recki \\u003c/em\\u003especimens and the mean number of each trichome types on the stem on the six \\u003cem\\u003eSolanum lycopersicum\\u003c/em\\u003e varieties, and \\u003cem\\u003eS. peruvianum\\u003c/em\\u003e, \\u003cem\\u003eS. cheesmaniae\\u003c/em\\u003e (a) correlation circle and (b) projection of the mean data for the genotypes considered.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"floatimage10.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4223917/v1/d2a3de831a706d5111736dbd.png\"},{\"id\":54519297,\"identity\":\"3df06d63-9e9c-4dbf-b181-2fdca25e4576\",\"added_by\":\"auto\",\"created_at\":\"2024-04-11 17:44:21\",\"extension\":\"png\",\"order_by\":7,\"title\":\"Figure 7\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":138499,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eBoxplots showing (a) the mean number of dorsal shield length and (b) the dorsal shield width between specimens of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e collected on \\u003cem\\u003eMentha suaveolens\\u003c/em\\u003e, \\u003cem\\u003eSolanum lycopersicum\\u003c/em\\u003e, \\u003cem\\u003eDatura\\u003c/em\\u003e sp., \\u003cem\\u003ePhomis fruticosa\\u003c/em\\u003e, \\u003cem\\u003eEchium vulgare\\u003c/em\\u003e and \\u003cem\\u003eCirsium arvense\\u003c/em\\u003e (dataset issued from Tixier et al. 2021). Letters correspond to results of the Newman \\u0026amp; Keuls post-hoc test.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"floatimage12.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4223917/v1/ae0822585da1604c2bde1377.png\"},{\"id\":54519095,\"identity\":\"24763d0e-5e17-432a-ac52-be442a2c5488\",\"added_by\":\"auto\",\"created_at\":\"2024-04-11 17:36:21\",\"extension\":\"png\",\"order_by\":8,\"title\":\"Figure 8\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":121813,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eRelation between the Dorsal Shield Length (DSL) and Width (DSW) of specimens of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e collected on \\u003cem\\u003eMentha suaveolens\\u003c/em\\u003e, \\u003cem\\u003eSolanum lycopersicum\\u003c/em\\u003e, \\u003cem\\u003eDatura\\u003c/em\\u003e sp., \\u003cem\\u003ePhomis fruticosa\\u003c/em\\u003e, \\u003cem\\u003eEchium vulgare\\u003c/em\\u003eand \\u003cem\\u003eCirsium arvense\\u003c/em\\u003e (dataset issued from Tixier et al. 2021).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"floatimage14.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4223917/v1/e29953a4d9caf1d758568a24.png\"},{\"id\":61794324,\"identity\":\"ca7458f2-ff1b-4686-9e7a-eb541b9e86bf\",\"added_by\":\"auto\",\"created_at\":\"2024-08-05 16:18:05\",\"extension\":\"pdf\",\"order_by\":0,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"manuscript-pdf\",\"size\":3970368,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"manuscript.pdf\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4223917/v1/25307527-65ad-4cdb-be18-0366363c7b1b.pdf\"}],\"financialInterests\":\"No competing interests reported.\",\"formattedTitle\":\"Ambulatory dispersal of Typhlodromus (Anthoseius) recki Wainstein (Acari: Phytoseiidae) along Solanceae stem\",\"fulltext\":[{\"header\":\"Introduction\",\"content\":\"\\u003cp\\u003eTomato crops are attacked by several pests, including mite pests as \\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e (Koch), \\u003cem\\u003eT. evansi\\u003c/em\\u003e Baker \\u0026amp; Pritchard and \\u003cem\\u003eAculops lycopersici\\u003c/em\\u003e (Tryon). Their control is still mainly based on pesticide application, especially sulfur (with partial efficiency), because of the inability of their main natural enemies (predatory mites of the family Phytoseiidae) to cope with tomato trichomes and thus to disperse and then feed on prey (Van Haren et al. \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e1987\\u003c/span\\u003e). Trichomes are well known plant defense mechanisms that limit herbivory (i.e., Glas et al. \\u003cspan citationid=\\\"CR17\\\" class=\\\"CitationRef\\\"\\u003e2012\\u003c/span\\u003e, Bar and Shtein \\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e, Karabourniotis et al. \\u003cspan citationid=\\\"CR18\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Among species of the genus \\u003cem\\u003eSolanum\\u003c/em\\u003e as well as among genotypes of a given plant species, trichome types are very diverse (i.e., glandular (GT) \\u003cem\\u003evs\\u003c/em\\u003e non glandular (NGT), small \\u003cem\\u003evs\\u003c/em\\u003e large, secreting toxic and/or repellent compounds) (i.e., Simmons and Gurr \\u003cspan citationid=\\\"CR39\\\" class=\\\"CitationRef\\\"\\u003e2005\\u003c/span\\u003e). Numerous studies have been carried out on the effect of tomato leaf trichomes on pests, generally showing negative effects of GTs, especially types I and IV, which affect pest survival both through physical effects (trapping on sticky trichome tips) and toxic and repellent effects due to the production of acylsucroses or terpenoids (Snyder et al. \\u003cspan citationid=\\\"CR40\\\" class=\\\"CitationRef\\\"\\u003e1998\\u003c/span\\u003e, Chatzivalsileiadis et al. 1999, Kroumova and Wagner \\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e, Fridman et al. \\u003cspan citationid=\\\"CR15\\\" class=\\\"CitationRef\\\"\\u003e2005\\u003c/span\\u003e, Simmons and Gurr \\u003cspan citationid=\\\"CR39\\\" class=\\\"CitationRef\\\"\\u003e2005\\u003c/span\\u003e, Economou et al. \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e2006\\u003c/span\\u003e, Treutter \\u003cspan citationid=\\\"CR46\\\" class=\\\"CitationRef\\\"\\u003e2006\\u003c/span\\u003e, Gershenzon and Dudareva \\u003cspan citationid=\\\"CR16\\\" class=\\\"CitationRef\\\"\\u003e2007\\u003c/span\\u003e, Alba et al. \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2009\\u003c/span\\u003e, Glas el al. 2012, Dias et al. \\u003cspan citationid=\\\"CR11\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e, Lucini et al. \\u003cspan citationid=\\\"CR24\\\" class=\\\"CitationRef\\\"\\u003e2015\\u003c/span\\u003e, Andrade et al. \\u003cspan citationid=\\\"CR2\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e, Rakha et al. \\u003cspan citationid=\\\"CR36\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e, Karabourniotis et al. \\u003cspan citationid=\\\"CR18\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Only a lower number studied the impact of such plant structures on natural enemies. Nevertheless, several authors have shown clear detrimental effects on several species of the predatory mite family Phytoseiidae. These authors emphasised the barrier effect of trichomes on the stem, preventing dispersal of predators throughout the plant and thus hampering biological control. GTs VI, which release toxic substances on contact with the mite, are the main types of trichomes that alter phytoseiid dispersal (Van Haren et al. \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e1987\\u003c/span\\u003e, Nihoul \\u003cspan citationid=\\\"CR29\\\" class=\\\"CitationRef\\\"\\u003e1994\\u003c/span\\u003e, Cedola et al. \\u003cspan citationid=\\\"CR5\\\" class=\\\"CitationRef\\\"\\u003e2001\\u003c/span\\u003e, Kennedy \\u003cspan citationid=\\\"CR19\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e, Sato et al. \\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e, Van Houten et al. \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e, Davidson et al. \\u003cspan citationid=\\\"CR9\\\" class=\\\"CitationRef\\\"\\u003e2016\\u003c/span\\u003e, Paspati et al. \\u003cspan citationid=\\\"CR32\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e, Casta\\u0026ntilde;\\u0026eacute; et al. \\u003cspan citationid=\\\"CR4\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e, Pandey et al. \\u003cspan citationid=\\\"CR31\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). As a result, none of the commercial Phytoseiidae mite species are effective enough to control mite pests on tomatoes, leading to numerous attempts to develop new biological control solutions. The Phytoseiidae mite family comprises more than 2,500 species world wide (Demite et al. \\u003cspan citationid=\\\"CR10\\\" class=\\\"CitationRef\\\"\\u003e2024\\u003c/span\\u003e). Some of these species are currently reported in Solanaceae, particularly in the genus \\u003cem\\u003eSolanum\\u003c/em\\u003e, suggesting that these mites may be adapted to plant defenses. Most of these predator species are reported in the Neotropical region, where the genus \\u003cem\\u003eSolanum\\u003c/em\\u003e originated, but also where the diversity of Phytoseiidae is the highest (Tixier et al. \\u003cspan citationid=\\\"CR43\\\" class=\\\"CitationRef\\\"\\u003e2020a\\u003c/span\\u003e; Duarte et al. \\u003cspan citationid=\\\"CR12\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). Because of potential problems related to ecological risks associated to the introduction of exotic species, studies have recently focused on endemic species from Europe collected on Solanaceae. Three species were particularly highly reported- \\u003cem\\u003eTyphlodromus\\u003c/em\\u003e (\\u003cem\\u003eTyphlodromus\\u003c/em\\u003e) \\u003cem\\u003eathiasae\\u003c/em\\u003e Athias-Henriot, essentially occuring in Israel; \\u003cem\\u003ePhytoseiulus persimilis\\u003c/em\\u003e Athias-Henriot, because of mass-release in tomato crops of this commercialised species; and \\u003cem\\u003eTyphlodromus\\u003c/em\\u003e (\\u003cem\\u003eAnthoseius\\u003c/em\\u003e) \\u003cem\\u003erecki\\u003c/em\\u003e Wainstein widely reported all over Europe (Tixier et al. \\u003cspan citationid=\\\"CR43\\\" class=\\\"CitationRef\\\"\\u003e2020a\\u003c/span\\u003e, Demite et al. \\u003cspan citationid=\\\"CR10\\\" class=\\\"CitationRef\\\"\\u003e2024\\u003c/span\\u003e). Recent laboratory studies have shown that the latter species is able to feed on tomato mite pests and is therefore considered a promising predator (Tixier et al. \\u003cspan citationid=\\\"CR44\\\" class=\\\"CitationRef\\\"\\u003e2020b\\u003c/span\\u003e, Ersin et al. \\u003cspan citationid=\\\"CR14\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). The first objective of this study is to assess the ability of this species to disperse ambulatory along the tomato stem, considering six varieties of \\u003cem\\u003eSolanum lycopersicum\\u003c/em\\u003e, \\u003cem\\u003eS. peruvianum\\u003c/em\\u003e and S. \\u003cem\\u003echeesmaniae\\u003c/em\\u003e with contrasting types and densities of trichomes on stem. Some authors hypothezised that dispersal ability on tomato stem can be associated to mite size (Pijnakker et al. \\u003cspan citationid=\\\"CR33\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). Observations of many specimens of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e have revealed large variations in body size (Tixier et al. \\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e), suggesting a large intraspecific polymorphism, as has been observed in other Phytoseiidae species whose body dimensions vary with season, temperature and diet (Chant \\u003cspan citationid=\\\"CR7\\\" class=\\\"CitationRef\\\"\\u003e1955\\u003c/span\\u003e, Tixier et al. \\u003cspan citationid=\\\"CR42\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e, Walzer and Schausberger \\u003cspan citationid=\\\"CR49\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e, Walzer et al. \\u003cspan citationid=\\\"CR50\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Therefore, the second objective of the present study was to determine whether the body size of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e could be related to the success of the mite in crossing the plant stem, for the eight tomato genotypes considered.\\u003c/p\\u003e\"},{\"header\":\"Material \\u0026 methods\",\"content\":\"\\u003cp\\u003e \\u003cb\\u003ePlant material.\\u003c/b\\u003e Eight genotypes were studied: \\u003cem\\u003eSolanum cheesmaniae\\u003c/em\\u003e (LA1412), \\u003cem\\u003eSolanum peruvianum\\u003c/em\\u003e (Peru CMV) and six varieties of \\u003cem\\u003eS. lycopersicum\\u003c/em\\u003e (747, Wooly, Yellow Bells, Clomimbo, Lancaster, Hairless). These genotypes were the same as those previously used for assessing the effects of trichomes on leaf, stem and petiole on \\u003cem\\u003eT. urticae\\u003c/em\\u003e and \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e (Tabary et al. \\u003cspan citationid=\\\"CR41\\\" class=\\\"CitationRef\\\"\\u003e2024\\u003c/span\\u003e). To obtain the experimental plants, seeds were sown in plastic pots (8 x 8 x 8 cm) in a greenhouse (25\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;2\\u0026deg;C, 40\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;30% RH) and watered twice a week until the beginning of the experiment (when plants reached three leaves).\\u003c/p\\u003e \\u003cp\\u003e \\u003cb\\u003eMite Material.\\u003c/b\\u003e The mite stock colony of \\u003cem\\u003eT. (A.) recki\\u003c/em\\u003e was initiated from populations (50\\u0026ndash;100 specimens) collected on \\u003cem\\u003ePhlomis fruticosa\\u003c/em\\u003e L. (lamiaceae) at Saint Cl\\u0026eacute;ment-de-Rivi\\u0026egrave;re (H\\u0026eacute;rault, France, 43.694144, 3.851792) on July 2021. Predatory mites were reared following the process described by McMurtry and Scriven (\\u003cspan citationid=\\\"CR28\\\" class=\\\"CitationRef\\\"\\u003e1965\\u003c/span\\u003e), in rearing units constituted of plastic arenas (10 x 15 cm) deposited on moistened sponge support, surrounded by water-saturated paper tissue for avoiding mite escape. The population was maintained in a climatic chamber (25\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;2\\u0026deg;C, 16D:8N photoperiod, 70\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;30% RH) and fed with commercialized \\u003cem\\u003eTypha angustifolia\\u003c/em\\u003e pollen (Nutrimite\\u0026reg;) twice a week.\\u003c/p\\u003e \\u003cp\\u003e \\u003cb\\u003eTrichome characterization\\u003c/b\\u003e. The trichome characterization followed the protocol used by Tabary et al. (\\u003cspan citationid=\\\"CR41\\\" class=\\\"CitationRef\\\"\\u003e2024\\u003c/span\\u003e) on the same plant material. Briefly, a piece of 1 cm of stem between the second and third leaf was cut with a razor blade, and on two areas of 4 mm\\u003csup\\u003e2\\u003c/sup\\u003e the types and number of trichomes were evaluated using a stereomicroscope (Nikon SMZZ1500 (x20)). Two replications were performed per plant, and 5 plants of each genotype were evaluated (in total 10 replicates / genotype). The trichome type classification proposed by Luckwill (\\u003cspan citationid=\\\"CR25\\\" class=\\\"CitationRef\\\"\\u003e1943\\u003c/span\\u003e) and Channarayappa et al. (1992) were used. The trichomes types I, IV, V and VI were considered separately, while types II and III were grouped together due to their similarity and the lack of studies demonstrating their role in arthropod resistance (McDowell et al. \\u003cspan citationid=\\\"CR27\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e). Trichome types VII and VIII were not taken into account due to their rarity on the eight genotypes studied.\\u003c/p\\u003e \\u003cp\\u003e \\u003cb\\u003eExperimental design\\u003c/b\\u003e. A 5 cm stem piece, located between the second and third leaf, was cut with a razor blade and then disposed on wet cotton on a Petri dish. A bean leaf disc (2.5 cm diameter, cv. Contender) with pollen of \\u003cem\\u003eT. angustifolia\\u003c/em\\u003e (Nutrimite\\u0026reg;) was disposed at one extremity, whereas a small piece of black plastic was placed at the other extremity (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e). One young mated female of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e was then disposed on the plastic piece. Once the female has dispersed on the stem, the plastic piece was removed. At least 10 replicates were carried out for each genotype. The age of predatory females was the same for each replicate. Observations were carried out at four different times: 10 minutes, 25, 55 and 100 minutes after the predator introduction. At each observation time, the predator was filmed for 5 minutes (camera Leica EZ4 W) and several parameters were assessed during the video observations: (i) the success of stem crossing : number of predators reaching the bean leaf and not turning back, (ii) hesitation behavior : the number of times the mite goes on and turns back on the stem without totally crossing it, (iii) escape behavior: number of predators retrieved on the wet cotton support and (iv) the mobility \\u003cem\\u003evs\\u003c/em\\u003e immobility duration. At the end of the experiment, the females were mounted on slides in Hoyer\\u0026rsquo;s medium. The mites were then observed using a microscope (Leica DLMB, Leica Microsyst\\u0026egrave;mes SAS, Rueil Malmaison, France) (400 x magnification), and body dimensions were measured. The Dorsal Shield Length (DSL) was assessed by measuring the distances between the setae \\u003cem\\u003ej1\\u003c/em\\u003e and \\u003cem\\u003eJ5\\u003c/em\\u003e. Three Dorsal Shield Width (DSWs) were assessed: (i) at the fore hind part measuring the distances between the setae \\u003cem\\u003ez4\\u003c/em\\u003e (DSW1), (ii) at the median part measuring the distances between the setae \\u003cem\\u003es6\\u003c/em\\u003e (DSW2) and (iii) at the backward part measuring the distances between the setae \\u003cem\\u003eS5\\u003c/em\\u003e (DSW3) (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003cp\\u003e \\u003cb\\u003eTyphlodromus\\u003c/b\\u003e \\u003cb\\u003e(\\u003c/b\\u003e\\u003cb\\u003eA\\u003c/b\\u003e.\\u003cb\\u003e)\\u003c/b\\u003e \\u003cb\\u003erecki\\u003c/b\\u003e \\u003cb\\u003ebody size phenotypes\\u003c/b\\u003e. Tixier et al. (\\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e) carried measurements of 91 females of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e collected on six plant species in France and in Italy. Among the characters considered, these authors measured the dorsal shield length and although the measurements were not taken at exactly the same positions on the body as in the present study, DSL (measured at the edge of the body tegument in Tixier et al. (\\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e) and between setae \\u003cem\\u003ej1\\u003c/em\\u003e and \\u003cem\\u003eJ5\\u003c/em\\u003e in the present study) and DSW1 (measured at the edge of the body tegument at the level of setae \\u003cem\\u003ez3\\u003c/em\\u003e in Tixier et al. (\\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e) and between setae z4 in the present study), we decided to use this dataset to determine whether the populations under consideration had variable body sizes and the proportion of \\u0026ldquo;small\\u0026rdquo; individuals in each of these populations.\\u003c/p\\u003e \\u003cp\\u003e \\u003cb\\u003eStatistical analyses\\u003c/b\\u003e. All analyses were carried out using R Cote Team (2021). ANOVA were carried out to assess the mean differences in mobility time and hesitation numbers between the \\u003cem\\u003eSolanum\\u003c/em\\u003e genotypes and between the females that succeed to cross or not the stem. ANOVA were also carried out to determine based on the dataset of Tixier et al. (\\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e) differences in DSL and DSW1 between the populations of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e here considered. ANOVA was carried out using the Kruskal-Wallis test (package \\u0026ldquo;rstatix\\u0026rdquo;) followed by Dunn test when the data did not follow a Normal law and/or with different variances, overwise ANOVA tests (package \\u0026ldquo;car\\u0026rdquo;) followed by Newman \\u0026amp; Keuls test (package \\u0026ldquo;Agricolae\\u0026rdquo;) were performed. A multifactorial analysis was used to assess the relationships between trichome numbers, mite body size and mobility time variables, using the packages \\u0026ldquo;FactoMineR\\u0026rdquo; and \\u0026ldquo;factoextra\\u0026rdquo;. Correlation tests (corr.test) were also carried out to determine the relationship between the three dorsal shield dimensions using the package \\u0026ldquo;ggpubr\\u0026rdquo;.\\u003c/p\\u003e\"},{\"header\":\"Results\",\"content\":\"\\u003cp\\u003e \\u003cb\\u003eTrichome characterization of the\\u003c/b\\u003e \\u003cb\\u003eSolanum\\u003c/b\\u003e \\u003cb\\u003egenotypes\\u003c/b\\u003e. The multifactorial analysis, which explains 35.2% and 23.6% of the variability for axes 1 and 2 respectively, shows that stem GT VI and NGT V differentiate the genotypes most (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003ea, b). On the axis 1, the density of these two trichome types were negatively correlated to the density of GT I, NGT II\\u0026amp;III and GT IV on the axis 1. On the axis 2, the density of trichome GT IV, GT VI and NGT II\\u0026amp;III was positively correlated to each other and differentiated from that of trichome GT I and NGT V. The genotypes were included into two main groups: (\\u003cb\\u003ei\\u003c/b\\u003e) comprising \\u003cem\\u003eSolanum peruvianum\\u003c/em\\u003e, \\u003cem\\u003eS. cheesmaniae\\u003c/em\\u003e and \\u003cem\\u003eS. lycopersicum\\u003c/em\\u003e cv. Hairless: characterized by high densities of NGT V, GT VI and low densities of trichomes GT I, NGT II\\u0026amp;III and GT IV, and (\\u003cb\\u003eii\\u003c/b\\u003e) comprising \\u003cem\\u003eS. lycopersicum\\u003c/em\\u003e cv. Wooly, cv. Yellow Bells and 747, characterized by opposite density of these latter trichome types. The cultivar Clomimbo and Lancaster had an intermediate position.\\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003cp\\u003e \\u003cb\\u003eTyphlodromus\\u003c/b\\u003e \\u003cb\\u003e(\\u003c/b\\u003e\\u003cb\\u003eA\\u003c/b\\u003e.\\u003cb\\u003e)\\u003c/b\\u003e \\u003cb\\u003erecki\\u003c/b\\u003e \\u003cb\\u003eambulatory dispersal ability\\u003c/b\\u003e. Predatory mites succeeded to cross the stem for five genotypes among the eight considered. However, the success crossing rate was low, except for \\u003cem\\u003eS. peruvianum\\u003c/em\\u003e (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig4\\\" class=\\\"InternalRef\\\"\\u003e4\\u003c/span\\u003e). The mean numbers of \\u0026ldquo;hesitation\\u0026rdquo; was not different between genotypes (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.73). No significant difference was either observed in the numbers of \\u0026ldquo;hesitation\\u0026rdquo; between the females that succeed to cross or not the stem (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.56). The percentage of mobility duration was not significantly different between the eight genotypes, all along the experiment and for each date except at 10 minutes, where a higher mobility activity was observed on \\u003cem\\u003eS. peruvianum\\u003c/em\\u003e. At all times measured, the time spent moving was longer for females that succeeded to cross the stem than for those that did not (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003cp\\u003eA multifactorial analysis was carried out, considering the mean values of trichome stem density of each genotype, dispersal predator parameters and its body size. The axes 1 and 2 represented 58.7% of the total variability of the system (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig6\\\" class=\\\"InternalRef\\\"\\u003e6\\u003c/span\\u003e). Mobility time variables correlated with each other, as well as with hesitation numbers and crossing success rates, suggesting that the more mobile females were, the more hesitation numbers (forwards and backwards) were high, and the more stem crossing success was important. These walking parameters were positively and negatively correlated with the number of trichome NGT V and GT VI, respectively, suggesting that the density of GT VI had a negative effect on dispersal, whereas the density of NGT V had no effect. In a lesser extent, the numbers of trichomes GT I, NGT II\\u0026amp;III and GT IV were also negatively correlated with the mobility ability, suggesting that the numbers of these trichome types may also act as a barrier for predator ambulatory dispersal along the stem. The escape rate was positively associated to the number of GT VI, but not to the densities of GT I, NGT II\\u0026amp;III and GT IV, suggesting that the presence of trichomes GT VI might be repellent for the predator. Predator mortality is not related to stem trichome characteristics and only little explains the variability in the system. Dispersal parameters were negatively correlated with dorsal shield length (DSL) and width, especially at the central body part (DSW2), suggesting that the smaller the mite length and width, the higher the mobility time, number of hesitations and crossing success. Finally, mites with a rather small and narrow body size were associated with stems with a low number of GT VI and, to a lesser extent, with GT I, NGT II\\u0026amp;III and GT IV, suggesting that mites with such dimensions would be better able to cope with tomato trichome defenses.\\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003cp\\u003e \\u003cb\\u003eVariability of\\u003c/b\\u003e \\u003cb\\u003eT\\u003c/b\\u003e. \\u003cb\\u003e(\\u003c/b\\u003e\\u003cb\\u003eA\\u003c/b\\u003e.\\u003cb\\u003e)\\u003c/b\\u003e \\u003cb\\u003erecki\\u003c/b\\u003e \\u003cb\\u003ebody dimensions.\\u003c/b\\u003e The analyses were conducted on the dataset produced by Tixier et al. (\\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). The mean DSL and DSW were significantly different between the populations of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e and the plants where they were collected. The lower dimensions were observed for specimens collected on \\u003cem\\u003eCirsium arvense\\u003c/em\\u003e L. (Compositae) and \\u003cem\\u003eEchium vulgare\\u003c/em\\u003e L. (Boraginaceae), whereas the highest were observed from specimens collected on \\u003cem\\u003eS. lycopersicum\\u003c/em\\u003e, \\u003cem\\u003eDatura\\u003c/em\\u003e sp. (Solanceae) and \\u003cem\\u003eMentha suaveolens\\u003c/em\\u003e L. (Lamiaceae); the specimens collected on \\u003cem\\u003eP. fruticosa\\u003c/em\\u003e showing an intermediate position (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig7\\\" class=\\\"InternalRef\\\"\\u003e7\\u003c/span\\u003e). These measurements were not exactly the same as those considered here, as they were taken at the boundary of the body and not between the setae, so their values are higher. Nevertheless, the Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig8\\\" class=\\\"InternalRef\\\"\\u003e8\\u003c/span\\u003e shows a correlation between these two measurements (R\\u0026thinsp;=\\u0026thinsp;0.53, P\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.0001). The proportion of small and narrow specimens was the highest for specimens collected on \\u003cem\\u003eC. arvense\\u003c/em\\u003e and \\u003cem\\u003eE. vulgare\\u003c/em\\u003e.\\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e\"},{\"header\":\"Discussion\",\"content\":\"\\u003cp\\u003eThe present study indicates that a relationship exists between types and numbers of trichomes on the stem and the ambulatory dispersal behavior of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e, observed during a short period of time (110 minutes). The NGT V on the stem was not associated to plant defense and high numbers of this trichome type even favor the mobility of the predator, as already observed for various pest species (Onyambus et al. \\u003cspan citationid=\\\"CR30\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e, Keskin and Kumral \\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2015\\u003c/span\\u003e, Savi et al. \\u003cspan citationid=\\\"CR38\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e). The density of GT VI affected the most \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e mobility. This result is consistent with observations already reported for other Phytoseiidae mite species (Van Haren et al. \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e1987\\u003c/span\\u003e, Paspati et al. \\u003cspan citationid=\\\"CR32\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e), whereas contrasted effects of the density of GT VI on the leaf were observed for the prey \\u003cem\\u003eT. urticae\\u003c/em\\u003e (Onyambus et al. \\u003cspan citationid=\\\"CR30\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e, Rakha et al. \\u003cspan citationid=\\\"CR36\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e, Savi et al. \\u003cspan citationid=\\\"CR38\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e, Kortbeek et al. \\u003cspan citationid=\\\"CR21\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). For example, in Phytoseiidae mites, Paspati et al. (\\u003cspan citationid=\\\"CR32\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e) showed detrimental effects of GT VI on the dispersal of the mite \\u003cem\\u003eAmblyseius swirskii\\u003c/em\\u003e Athias-Henriot due to the toxic effect of acylsugars causing (i) suffocation due to acylsugar accumulation at mite cuticle openings and (ii) disruption of the subcuticular membrane (Puterka et al. \\u003cspan citationid=\\\"CR34\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e). Here, the mortality of the predator was not different between plant genotypes, regardless of the number of GT VI. The cuticle of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e is quite sclerotized (Livshitz and Kuznetsov \\u003cspan citationid=\\\"CR23\\\" class=\\\"CitationRef\\\"\\u003e1972\\u003c/span\\u003e) compared to that of other Phytoseiidae species, and especially \\u003cem\\u003eA. swirskii\\u003c/em\\u003e. This morphological trait could explain the lack of effect of trichome exudates on the mortality of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e, but the relatively short observation period could also explain the low mortality here observed. Further experiments will help to confirm if \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e can cope with the toxicity of acylsucrose. The density of GT VI was also correlated to the numbers of hesitation and escape rates. A repellent effect, causing the mites to move on and then turn back, could be hypothesized, as has already been observed for pest mites (Maluf et al. \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e2001\\u003c/span\\u003e) and Phytoseiidae mites (Sato et al. \\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e). The density of the other trichome types (GT I and IV, and NGT II\\u0026amp;III) was also associated with low mobility and stem crossing, but to a lesser extent than GT VI. The GT I and IV are reported to be detrimental for pests when present on leaves (i.e., Simmons and Gurr \\u003cspan citationid=\\\"CR39\\\" class=\\\"CitationRef\\\"\\u003e2005\\u003c/span\\u003e). Here, GT I and IV do not seem to be related to escape. This suggests that, unlike trichome VI, they do not have a repellent effect. The effect of NGT II\\u0026amp;III was rather unexpected, as studies even ignore this trichome type when examining the effects of trichomes density on arthropods. Most of these later studies concern the leaf surface, we can thus hypothesize a different effect of these trichomes when present on the stem. However, we cannot rule out the possibility that such an effect of NGT II\\u0026amp;III may also be due to an artefact, since plants with high densities of GT I and IV also have high densities of NGT II\\u0026amp;III.\\u003c/p\\u003e \\u003cp\\u003eThe present study provides new information on the dispersal behavior of predators, showing that mobility time and \\\"hesitation\\\" activity (walking forward and backward) are both related to stem crossing success. This suggests that the predator is trying to cross the stem in an incisive way, and the more it tries, the more it succeeds. Van Haren et al. (\\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e1987\\u003c/span\\u003e) showed that the dispersal of predators is facilitated when the pest mites have crossed the stem, as the passage of the prey causes trichome breakage and loss of toxicity over time. Van Houten et al. (\\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e) reported that dispersal of \\u003cem\\u003eA. swirskii\\u003c/em\\u003e is favored on stems attacked by \\u003cem\\u003eA. lycopersici\\u003c/em\\u003e, due to trichome collapse after the pest passage. To our knowledge, whether such trichome collapse and loss of toxicity occurs after predator passage is not known. It would therefore be interesting to determine whether the \\\"stubbornness\\\" of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e individuals observed in this study could be linked to a strategy aimed at facilitating the dispersal of the rest of the colony, by carrying out similar experiments with a large number of specimens of the predator.\\u003c/p\\u003e \\u003cp\\u003eAlthough only a small proportion of the total number of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e examined managed to cross the stem (15%), there were correlations between mite size and crossing success / mobility activities. The mite size parameters that were the most related to stem crossing were the dorsal shield length and the mid-body with and to a lesser extent fore hind and back widths. These results seem to show that the smaller and narrower the mite specimens, the higher the stem crossing success. Several authors have in the past suggested a relationship between mite size and ability to squeeze through trichomes, especially for the tydeid mite \\u003cem\\u003ePronematus ubiquitus\\u003c/em\\u003e (McGregor) (Ilionidae), and for some mite pests as the eriophyid mite \\u003cem\\u003eA. lycopersici\\u003c/em\\u003e (Pijnakker et al. \\u003cspan citationid=\\\"CR33\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). However, these mite species are much smaller than \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e (even than specimens that succeeded to cross the stem). The Phytoseiidae mites studied until now are usually bigger than \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e, especially for \\u003cem\\u003eA. swirskii\\u003c/em\\u003e and \\u003cem\\u003eP. persimilis, P. longipes\\u003c/em\\u003e and \\u003cem\\u003eP. macropilis\\u003c/em\\u003e. In addition, these predatory mites issued from commercial rearings are usually well fed, so their size is significant and their ability to spread through trichomes is likely to be difficult. It will therefore be necessary to test the success of stem crossing as a function of body size in other mite species and for other stages such as immatures and males. Tixier et al. (\\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e) showed different morphological phenotypes of \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e depending on the plants from which the populations were collected. Using this latter dataset, differences in the length and width of the dorsal shield were observed between these populations, with the smallest specimens found more frequently in the populations collected on \\u003cem\\u003eE. vulgare\\u003c/em\\u003e and \\u003cem\\u003eC. arvense\\u003c/em\\u003e, the population collected on \\u003cem\\u003eP. fruticosa\\u003c/em\\u003e (the one used for the present experiments) having an intermediate position. It would therefore be interesting to carry out experiments similar to the present ones, using the other \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e populations, in order to validate the hypothesis that the size of the mites and the success of stem-crossing are related, i.e. that the mite crossing rate is higher for mites collected on \\u003cem\\u003eE. vulgare\\u003c/em\\u003e and \\u003cem\\u003eC. arvense\\u003c/em\\u003e and lower for mites collected on \\u003cem\\u003eM. suaveolens\\u003c/em\\u003e, \\u003cem\\u003eS. lycopersicum\\u003c/em\\u003e and \\u003cem\\u003eDatura\\u003c/em\\u003e sp.\\u003c/p\\u003e\"},{\"header\":\"Conclusion\",\"content\":\"\\u003cp\\u003eThe present study confirms the detrimental role of stem trichome density in the dispersal of Phytoseiidae mites, especially GT VI, which also seems to have a repellent effect. It also shows that females try hard to cross the stem, and the more mobile they are, the higher the success rate. These results open new avenues for biocontrol, in particular to determine whether this behavior can be linked to trichome collapse, which subsequently might facilitate the passage of other specimens. Noteworthy relationships between predator body size and stem crossing success were also found. The proportion of stem-crossing specimens within the population studied here is quite low, but other populations, than the one currently considered, appear to have a higher proportion of small specimens. Thus, additional studies would be required to test the stem crossing success of specimens from these latter populations. Several factors can affect mite size, as climatic conditions and plant phenotypes (Chant \\u003cspan citationid=\\\"CR7\\\" class=\\\"CitationRef\\\"\\u003e1955\\u003c/span\\u003e, Tixier et al. \\u003cspan citationid=\\\"CR42\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e, Walzer et al. \\u003cspan citationid=\\\"CR50\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Thus, although the present study highlights new potential traits for selecting the predator phenotype best adapted for dispersal along the tomato stem, further experiments are needed to determine how much predator size is labile and how much is determined by environmental factors. It would also be necessary to determine the extent to which feeding behavior differs between small and large predator phenotypes for issues of biological control efficacy. Finally, this study mainly focuses on mite size, but other characters may also be involved in crossing success, as mite dorsal sclerotization which is particularly well developed in \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA\\u003c/em\\u003e.) \\u003cem\\u003erecki\\u003c/em\\u003e, in contrast to the other Phytoseiidae species tested so far.\\u003c/p\\u003e\"},{\"header\":\"Declarations\",\"content\":\"\\u003ch2\\u003eAuthor Contribution\\u003c/h2\\u003e\\u003cp\\u003eMarie-St\\u0026eacute;phane Tixier, Maria Navajas and Denise Navia defined the experimental design. Amandine Raeckelboom, Martial Douin and Lou Tabary carried out the experiments. Amandine Raeckelboom and Marie-St\\u0026eacute;phane Tixier carried out the statistical analyses. Marie-Stephane Tixier wrote the manuscript with Maria Navajas and Denise Navia.\\u003c/p\\u003e\"},{\"header\":\"References\",\"content\":\"\\u003col\\u003e\\n\\u003cli\\u003eAlba JM, Montserrat M, Fern\\u0026aacute;ndez-Mu\\u0026ntilde;oz R (2009) Resistance to the two-spotted spider mite (\\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e) by acylsucroses of wild tomato (\\u003cem\\u003eSolanum pimpinellifolium\\u003c/em\\u003e) trichomes studied in a recombinant inbred line population. \\u003cem\\u003eExp Appl Acarol\\u003c/em\\u003e 47: 35\\u0026ndash;47.\\u003c/li\\u003e\\n\\u003cli\\u003eAndrade MC, da Silva AA, Neiva IP, Oliveira IRC, De Castro EM, Francis DM, Maluf WR (2017)\\u003cem\\u003e \\u003c/em\\u003eInheritance of type IV glandular trichome density and its association with whitefly resistance from \\u003cem\\u003eSolanum galapagense\\u003c/em\\u003e accession LA1401. Euphytica 213: 52. doi:10.1007/s10681-016-1792-1\\u003c/li\\u003e\\n\\u003cli\\u003eBar M, Shtein A (2019) Plant trichomes and the biomechanics of defense in various systems, with Solanaceae as a model. Botany 97(12): 651-660. doi:10.1139/cjb-2019-0144\\u003c/li\\u003e\\n\\u003cli\\u003eCasta\\u0026ntilde;\\u0026eacute; C, Alomar O, Rocha A, Vila E, Riudavets J (2022) Control of \\u003cem\\u003eAculops lycopersici\\u003c/em\\u003e with the predatory mite \\u003cem\\u003eTranseius montdorensis\\u003c/em\\u003e. Insects 13: 1116. doi:10.3390/insects13121116 \\u003c/li\\u003e\\n\\u003cli\\u003eCedola CV, S\\u0026aacute;nchez NE, Liljesthr\\u0026ouml;m GG (2001) Effect of tomato leaf hairiness on functional and numerical response of \\u003cem\\u003eNeoseiulus californicus\\u003c/em\\u003e (Acari: Phytoseiidae). Exp Appl Acarol 25: 819 \\u0026ndash; 831.\\u003c/li\\u003e\\n\\u003cli\\u003eChannaryappa SG, Muniyappa V, Frist RH (1992) Resistance of \\u003cem\\u003eLycopersicon\\u003c/em\\u003e species to \\u003cem\\u003eBemisia tabaci\\u003c/em\\u003e, a tomato leaf curl virus vector. Can J Bot 70: 2184\\u0026ndash;2192.\\u003c/li\\u003e\\n\\u003cli\\u003eChant DA (1955) Notes on mites of the mites of the genus \\u003cem\\u003eTyphlodromus\\u003c/em\\u003e Scheuten 1857 (Acarina: Laelapidae) with description of the males of some species and the females of a new species. Can Entomol 87: 496-503. \\u003c/li\\u003e\\n\\u003cli\\u003eChatzivasileiadis EA, Boon JJ, Sabelis MW (1999) Accumulation and turnover of 2-tridecanone in \\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e and consequences for resistance of wild and cultivated tomatoes. Exp Appl Acarol 23: 1011-1021.\\u003c/li\\u003e\\n\\u003cli\\u003eDavidson MM, Nielsen M-C, Butler RC, Silberbauer RB (2016) Prey consumption and survival of the predatory mite, \\u003cem\\u003eAmblydromalus limonicus\\u003c/em\\u003e, on different prey and host plants. Biocontrol Sci 26: 722-726.\\u003c/li\\u003e\\n\\u003cli\\u003eDemite, P.R., Moraes, G.J. de, McMurtry, J.A., Denmark, H.A. \\u0026amp; Castilho, R. C. (2024) Phytoseiidae Database. Available from: www.lea.esalq.usp.br/phytoseiidae (accessed 30/03/2024)\\u003c/li\\u003e\\n\\u003cli\\u003eDias DM, Resende JTV, Faria MV, Camargo LKP, Lima IP (2013) Selection of processing tomato genotypes with high acyl sugar content that are resistant to the tomato pinworm. Genet Mol Res 12: 381\\u0026ndash;389. doi: 10.4238/2013.February.8.2\\u003c/li\\u003e\\n\\u003cli\\u003eDuarte ME, Demite PR, Mendon\\u0026ccedil;a RS, Michereff-Filho M, Santa Cruz de Mesquita Alves ML, Peixoto JR, Navia D (2021) Phytoseiidae mites associated with native and cultivated solanaceous in Central-West Brazil. Sys Appl Acarol 26(12), 2358-2384. doi : 10.11158/saa.26.12.13\\u003c/li\\u003e\\n\\u003cli\\u003eEconomou L, Lykouressis D, Barbetaki A (2006) Time allocation of activities of two heteropteran predators on the leaves of three tomato cultivars with variable glandular trichome density. Environ Entomol 35: 387\\u0026ndash;393. doi:10.1603/0046-225X-35.2.387 \\u003c/li\\u003e\\n\\u003cli\\u003eErsin F, Turanli F, Cakmak I (2021) Development and life history parameters of \\u003cem\\u003eTyphlodromus recki\\u003c/em\\u003e (Acari: Phytoseiidae) feeding on \\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e (Acari: Tetranychidae) at different temperatures. Syst Appl Acarol 26: 496-508. doi:10.11158/saa.26.2.12\\u003c/li\\u003e\\n\\u003cli\\u003eFridman E, Wang J, Iijima Y, Froehlich JE, Gang DR, Ohlrogge J, Pichersky E (2005) Metabolic, genomic, and biochemical analyses of glandular trichomes from the wild tomato species \\u003cem\\u003eLycopersicon hirsutum\\u003c/em\\u003e identify a key enzyme in the biosynthesis of methylketones. Plant Cell 17(4): 1252\\u0026ndash;1267. doi: 10.1105/tpc.104.029736\\u003c/li\\u003e\\n\\u003cli\\u003eGershenzon J, Dudareva N (2007) The function of terpene natural products in the natural world. Nat Chem Biol 3: 408\\u0026ndash;414. doi: 10.1038/nchembio.2007.5\\u003c/li\\u003e\\n\\u003cli\\u003eGlas JJ, Schimmel BC, Alba JM, Escobar-Bravo R, Schuurink RC, Kant MR (2012) Plant glandular trichomes as targets for breeding or engineering of resistance to herbivores. Int J Mol Sci 13 (12): 17077\\u0026ndash;17103. doi:10.3390/ijms131217077\\u003c/li\\u003e\\n\\u003cli\\u003eKarabourniotis G, Liakopoulos G, Nikolopoulos D, Bresta A (2020) Protective and defensive roles of non-glandular trichomes against multiple stresses: structure\\u0026ndash;function coordination. J For Res 31: 1\\u0026ndash;12. doi:10.1007/s11676-019-01034-4\\u003c/li\\u003e\\n\\u003cli\\u003eKennedy GG (2003) Tomato, pests, parasitoids, and predators: tritrophic interactions involving the genus \\u003cem\\u003eLycopersicon\\u003c/em\\u003e. \\u003cem\\u003eAnnu Rev Entomol\\u003c/em\\u003e 48: 51\\u0026ndash;72.\\u003c/li\\u003e\\n\\u003cli\\u003eKeskin N, Kumral NA (2015) Screening tomato varietal resistance against the two-spotted spider mite [\\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e (Koch)]. \\u003cem\\u003eIntern J Acarol\\u003c/em\\u003e 41: 300\\u0026ndash;309.\\u003c/li\\u003e\\n\\u003cli\\u003eKortbeek RWJ, Galland MD, Muras A, van der Kloet FM, Andr\\u0026eacute; B, Heilijgers M, van Hijum SAFT, Haring MA, Schuurink RC, Bleeker PM (2021) Natural variation in wild tomato trichomes; selecting metabolites that contribute to insect resistance using a random forest approach. BMC Plant Biol 21(1):315. doi: 10.1186/s12870-021-03070-x\\u003c/li\\u003e\\n\\u003cli\\u003eKroumova AB, Wagner GJ (2003) Different elongation pathways in the biosynthesis of acyl groups of trichome exudate sugar esters from various solanaceous plants. Planta 216: 1013\\u0026ndash;1021.\\u003c/li\\u003e\\n\\u003cli\\u003eLivshitz IZ, Kuznetsov NN (1972) Phytoseiid mites from Crimea (Parasitiformes: Phytoseiidae) [in Russian] \\u0026ndash; \\u003cem\\u003eIn\\u003c/em\\u003e: Pests and diseases of fruit and ornamental plants. Proceed. All-Union V. I. Lenin Academy of Agricultural Science, The State Nikita Botanical Gardens, 61: 13-64.\\u003c/li\\u003e\\n\\u003cli\\u003eLucini T, Marcos VF, Cristhiane R, Juliano TVR, Jo\\u0026atilde;o RFO (2015) Acylsugar and the role of trichomes in tomato genotypes resistance to \\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e. Arthropod-Plant Interact 9: 45\\u0026ndash;53. \\u003c/li\\u003e\\n\\u003cli\\u003eLuckwill LC (1943) The genus \\u003cem\\u003eLycopersicon\\u003c/em\\u003e: An historical biological and taxanomic survey of the wild and cultivated tomatoes. Aberdeen University Press, U.K., 44 pp.\\u003c/li\\u003e\\n\\u003cli\\u003eMaluf WR, Campos GA, Cardoso MDG (2001) Relationships between trichome types and spider mite (\\u003cem\\u003eTetranychus evansi\\u003c/em\\u003e) repellence in tomatoes with respect to foliar zingiberene contents. Euphytica121: 73\\u0026ndash;80.\\u003c/li\\u003e\\n\\u003cli\\u003eMcDowell ET, Kapteyn J, Schmidt A, Li C, Kang J, Descour A, Shi F, Larson M, Schilmiller A, An L, Jones A, Pichersky E, Soderlund CA, Gang DR (2011) Comparative functional genomic analysis of \\u003cem\\u003eSolanum\\u003c/em\\u003e\\u003cem\\u003e \\u003c/em\\u003eglandular trichome types. Plant Physiol 155: 524\\u0026ndash;539.\\u003c/li\\u003e\\n\\u003cli\\u003eMcMurtry JA, Scriven GT (1965) Insectary production of phytoseiid mites. J Econ Entomol 58: 282-284. doi:10.1093/jee/58.2.282 \\u003c/li\\u003e\\n\\u003cli\\u003eNihoul P (1994) Phenology of glandular trichomes related to entrapment of \\u003cem\\u003ePhytoseiulus persimilis\\u003c/em\\u003e A.-H. in the glasshouse tomato. J Hort Sci 69(5): 783-789.\\u003c/li\\u003e\\n\\u003cli\\u003eOnyambus GK, Maranga RO, Gitonga LM, Knapp M (2011) Host plant resistance among tomato accessions to the spider mite \\u003cem\\u003eTetranychus evansi\\u003c/em\\u003e in Kenya. Exp Appl Acarol 54: 385\\u0026ndash;393.\\u003c/li\\u003e\\n\\u003cli\\u003ePandey S, Biscaia Ribeiro da Silva AL, Dutta B, Chong JH, Mutscher MA, Schmidt JA (2023) Acylsugar tomato lines suppress whiteflies and \\u003cem\\u003eAmblyseius swirskii\\u003c/em\\u003e establishment. Entomol Exp Appl online first, doi:10.1111/eea.13342\\u003c/li\\u003e\\n\\u003cli\\u003ePaspati A, Rambla Nebot JL, L\\u0026oacute;pez-Gresa MP, Arbona V, G\\u0026oacute;mez-Cadenas A, Granell Richart A, Gonz\\u0026aacute;lez-Cabrera J, Urbaneja A (2021). Tomato trichomes are deadly hurdles limiting the establishment of \\u003cem\\u003eAmblyseius swirskii\\u003c/em\\u003e Athias-Henriot (Acari: Phytoseiidae). Biol Contr 157: 1-9. doi:10.1016/j.biocontrol.2021.104572\\u003c/li\\u003e\\n\\u003cli\\u003ePijnakker J, Moerkens R, Vangansbeke D, Duarte M, Bellinkx S, Benavente A, Merckx J, Stevens I, W\\u0026auml;ckers F (2022) Dual Protection: A Tydeoid Mite Effectively Controls Both a Problem Pest and a Key Pathogen in Tomato. Pest Manag Sci 78: 355\\u0026ndash;361.\\u003c/li\\u003e\\n\\u003cli\\u003ePuterka GJ, Farone W, Palmer T, Barrington A (2003) Structure-function relationships affecting the insecticidal and miticidal activity of sugar esters. J Econ Entomol 96: 636\\u0026ndash;644.\\u003c/li\\u003e\\n\\u003cli\\u003eR Core Team (2021) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/.\\u003c/li\\u003e\\n\\u003cli\\u003eRakha M, Bouba N, Ramasamy S, Regnard J-L, Hanson P (2017\\u003cem\\u003e).\\u003c/em\\u003e Evaluation of wild tomato accessions (\\u003cem\\u003eSolanum\\u003c/em\\u003e spp.) for resistance to two-spotted spider mite (\\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e Koch) based on trichome type and acylsugar content. Genet Res Crop Evol 64: 1011\\u0026ndash;1022. doi: 10.1007/s10722-016-0421-0 \\u003c/li\\u003e\\n\\u003cli\\u003eSato MM, Moraes GJ, Haddad ML, Wekesa VW (2011) Effect of trichomes on the predation of \\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e (Acari: Tetranychidae) by \\u003cem\\u003ePhytoseiulus macropilis\\u003c/em\\u003e (Acari: Phytoseiidae) on tomato, and the interference of webbing. Exp Appl Acarol 54: 21\\u0026ndash;32. doi: 10.1007/s10493-011-9426-8\\u003c/li\\u003e\\n\\u003cli\\u003eSavi PJ, Moraes GJ, Boi\\u0026ccedil;a AL Jr, Melville CC, Carvalho RF, Louren\\u0026ccedil;ao AL, Andrade DJ (2019) Impact of leaflet trichomes on settlement and oviposition of \\u003cem\\u003eTetranychus evansi\\u003c/em\\u003e (Acari: Tetranychidae) in African and South American tomatoes. 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Exp Appl Acarol - submitted\\u003c/li\\u003e\\n\\u003cli\\u003eTixier M-S, Kreiter S, Cheval B, Auger P (2003) Morphometric variation between populations of \\u003cem\\u003eKampimodromus\\u003c/em\\u003e \\u003cem\\u003eaberrans\\u003c/em\\u003e (Oudemans) (Acari: Phytoseiidae): Implications for the taxonomy of the genus. Inv syst 17: 349 \\u0026ndash; 358.\\u003c/li\\u003e\\n\\u003cli\\u003eTixier M-S, Douin M, Kreiter S (2020a) Phytoseiidae (Acari: Mesostigmata) on Plants of the Family Solanaceae: Results of a Survey in the South of France and a Review of World Biodiversity. Exp Appl Acarol 81: 357-388.\\u003c/li\\u003e\\n\\u003cli\\u003eTixier M-S, Douin M, Rocio O, Gonzalez L, Pount B, Kreiter S (2020b) Distribution and biological features of \\u003cem\\u003eTyphlodromus\\u003c/em\\u003e (\\u003cem\\u003eAnthoseius\\u003c/em\\u003e) \\u003cem\\u003erecki\\u003c/em\\u003e (Acari: Phytoseiidae) on \\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e, \\u003cem\\u003eT. evansi\\u003c/em\\u003e (Acari: Tetranychidae) and \\u003cem\\u003eAculops lycopersici\\u003c/em\\u003e (Acari: Eriophyidae). Acarologia 60: 684\\u0026ndash;697. doi:10.24349/acarologia/20204396.\\u003c/li\\u003e\\n\\u003cli\\u003eTixier M-S, Perez Martinez S, Douin M (2021)Markers of life history traits: variation in morphology, molecular and amino acid sequences within \\u003cem\\u003eTyphlodromus\\u003c/em\\u003e (\\u003cem\\u003eAnthoseius\\u003c/em\\u003e) \\u003cem\\u003erecki\\u003c/em\\u003e Wainstein (Acari: Mesostigmata: Phytoseiidae). Biol J Linn Soc132: 53\\u0026ndash;73. https://doi.org/10.1093/biolinnean/blaa103\\u003c/li\\u003e\\n\\u003cli\\u003eTreutter D (2006) Significance of flavonoids in plant resistance: A review. Environ. Chem Lett 4: 147\\u0026ndash;157. Doi : 10.1007/s10311-006-0068-8\\u003c/li\\u003e\\n\\u003cli\\u003eVan Haren R, Steenhuis M, Sabelis MW, De Ponti O (1987) Tomato stem trichomes and dispersal success of \\u003cem\\u003ePhytoseiulus persimilis\\u003c/em\\u003e relative to its prey \\u003cem\\u003eTetranychus urticae\\u003c/em\\u003e. Exp Appl Acarol 3: 115\\u0026ndash;121.\\u003c/li\\u003e\\n\\u003cli\\u003evan Houten YM, Glas JJ, Hoogerbrugge H, Rothe J, Bolckmans KJ, Simoni S, van Arkel J, Alba JM, Kant MR, Sabelis MW (2013) Herbivory-associated degradation of tomato trichomes and its impact on biological control of \\u003cem\\u003eAculops lycopersici\\u003c/em\\u003e. Exp Appl Acarol 60: 127-38. doi: 10.1007/s10493-012-9638-6 \\u003c/li\\u003e\\n\\u003cli\\u003eWalzer A, Schausberger P (2011) Sex-specific developmental plasticity of generalist andspecialist predatory mites (Acari: Phytoseiidae) inresponse to food stress. Biol J Linn Soc102: 650\\u0026ndash;660.\\u003c/li\\u003e\\n\\u003cli\\u003eWalzer A, Formayer H, Tixier M-S (2020) Evidence of trans-generational developmental modifications induced by simulated heat waves in an arthropod. Sci Rep 10: 4098. https://doi.org/10.1038/s41598-020-61040-z\\u003c/li\\u003e\\n\\u003c/ol\\u003e\"}],\"fulltextSource\":\"\",\"fullText\":\"\",\"funders\":[],\"hasAdminPriorityOnWorkflow\":false,\"hasManuscriptDocX\":true,\"hasOptedInToPreprint\":true,\"hasPassedJournalQc\":\"\",\"hasAnyPriority\":false,\"hideJournal\":true,\"highlight\":\"\",\"institution\":\"\",\"isAcceptedByJournal\":true,\"isAuthorSuppliedPdf\":false,\"isDeskRejected\":\"\",\"isHiddenFromSearch\":false,\"isInQc\":false,\"isInWorkflow\":false,\"isPdf\":false,\"isPdfUpToDate\":true,\"isWithdrawnOrRetracted\":false,\"journal\":{\"display\":true,\"email\":\"info@researchsquare.com\",\"identity\":\"researchsquare\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":true,\"externalIdentity\":\"\",\"sideBox\":\"\",\"snPcode\":\"\",\"submissionUrl\":\"/submission\",\"title\":\"Research Square\",\"twitterHandle\":\"researchsquare\",\"acdcEnabled\":true,\"dfaEnabled\":false,\"editorialSystem\":\"\",\"reportingPortfolio\":\"\",\"inReviewEnabled\":false,\"inReviewRevisionsEnabled\":true},\"keywords\":\"Solanaceae, mites, biological control, mite body size, trichome \",\"lastPublishedDoi\":\"10.21203/rs.3.rs-4223917/v1\",\"lastPublishedDoiUrl\":\"https://doi.org/10.21203/rs.3.rs-4223917/v1\",\"license\":{\"name\":\"CC BY 4.0\",\"url\":\"https://creativecommons.org/licenses/by/4.0/\"},\"manuscriptAbstract\":\"\\u003cp\\u003eTomato crops are attacked by several pests, including mites. While the main predatory mites are not effective enough to control mite pests due to problems with plant dispersal and establishment (associated with glandular trichomes - GT - on leaves and stems), recent studies have shown encouraging results with the European endemic phytoseiid \\u003cem\\u003eTyphlodromus\\u003c/em\\u003e (\\u003cem\\u003eAnthoseius\\u003c/em\\u003e) \\u003cem\\u003erecki\\u003c/em\\u003e. The first objective of the present study was to assess the ability of this species to disperse along the tomato stem, considering six genotypes of \\u003cem\\u003eSolanum lycopersicum\\u003c/em\\u003e, \\u003cem\\u003eS. peruvianum\\u003c/em\\u003e and \\u003cem\\u003eS. cheesmaniae\\u003c/em\\u003e with contrasted trichome numbers and types of stem trichomes, accuratetly characterised in a previous study. As morphological variation in body size has been observed within the species \\u003cem\\u003eT\\u003c/em\\u003e. (\\u003cem\\u003eA.\\u003c/em\\u003e) \\u003cem\\u003erecki\\u003c/em\\u003e, the second objective was to determine how predator morphological traits can explain dispersal along the tomato stem. For this, ambulatory dispersal ability of females was tested in lab conditions on the eight \\u003cem\\u003eSolanum\\u003c/em\\u003e genotypes. Then, the females were mounted on slides and body dimensions measured. No effect of the tomato genotypes was observed on the dispersal ability of the predator. However, specimens that succeeded in crossing the stem, had a higher mobility time than those that failed. Furthermore, body width at midbody and dorsal shield length were negatively correlated with dispersal ability. This suggests that the more slender and relatively small the specimens, the more are mobile and able to successfully cross the stem, regardless of the plant genotype considered. The number of glandular trichomes type (GT) VI and to a lesser extent GT I and IV, and non-glandular trichomes (NGT) II\\u0026amp;III appear to limit dispersal. The GT VI seems to have a repellent effect, inducing mite escape in some cases. On the opposite, the number of NGT V were positively correlated with high mobility and stem crossing rates. Assuming that the main barrier to biological control efficiency is dispersal along tomato stems, the results obtained here should have implications for biological control success. The proportion of mites with 'optimal dimensions' appears to be low and further studies should be undertaken to better assess the proportion of mites with such ideal dimensions in different populations and also to determine whether these morphological traits are associated with different feeding abilities and/or abiotic conditions.\\u003c/p\\u003e\",\"manuscriptTitle\":\"Ambulatory dispersal of Typhlodromus (Anthoseius) recki Wainstein (Acari: Phytoseiidae) along Solanceae stem\",\"msid\":\"\",\"msnumber\":\"\",\"nonDraftVersions\":[{\"code\":1,\"date\":\"2024-04-11 17:36:16\",\"doi\":\"10.21203/rs.3.rs-4223917/v1\",\"editorialEvents\":[{\"type\":\"communityComments\",\"content\":0}],\"status\":\"published\",\"journal\":{\"display\":true,\"email\":\"info@researchsquare.com\",\"identity\":\"researchsquare\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":true,\"externalIdentity\":\"\",\"sideBox\":\"\",\"snPcode\":\"\",\"submissionUrl\":\"/submission\",\"title\":\"Research Square\",\"twitterHandle\":\"researchsquare\",\"acdcEnabled\":true,\"dfaEnabled\":false,\"editorialSystem\":\"\",\"reportingPortfolio\":\"\",\"inReviewEnabled\":false,\"inReviewRevisionsEnabled\":true}}],\"origin\":\"\",\"ownerIdentity\":\"3987cda2-2588-492b-97df-7cfc36b10c09\",\"owner\":[],\"postedDate\":\"April 11th, 2024\",\"published\":true,\"recentEditorialEvents\":[],\"rejectedJournal\":[],\"revision\":\"\",\"amendment\":\"\",\"status\":\"posted\",\"subjectAreas\":[],\"tags\":[],\"updatedAt\":\"2024-08-05T16:12:17+00:00\",\"versionOfRecord\":{\"articleIdentity\":\"rs-4223917\",\"link\":\"https://doi.org/10.1007/s10493-024-00946-z\",\"journal\":{\"identity\":\"experimental-and-applied-acarology\",\"isVorOnly\":false,\"title\":\"Experimental and Applied Acarology\"},\"publishedOn\":\"2024-08-01 15:57:09\",\"publishedOnDateReadable\":\"August 1st, 2024\"},\"versionCreatedAt\":\"2024-04-11 17:36:16\",\"video\":\"\",\"vorDoi\":\"10.1007/s10493-024-00946-z\",\"vorDoiUrl\":\"https://doi.org/10.1007/s10493-024-00946-z\",\"workflowStages\":[]},\"version\":\"v1\",\"identity\":\"rs-4223917\",\"journalConfig\":\"researchsquare\"},\"__N_SSP\":true},\"page\":\"/article/[identity]/[[...version]]\",\"query\":{\"redirect\":\"/article/rs-4223917\",\"identity\":\"rs-4223917\",\"version\":[\"v1\"]},\"buildId\":\"qtupq5eGEP_6zYnWcrvyt\",\"isFallback\":false,\"isExperimentalCompile\":false,\"dynamicIds\":[84888],\"gssp\":true,\"scriptLoader\":[]}","source_license":"CC-BY-4.0","license_restricted":false}