Host plant quality mediates dispersal, oviposition, and sex allocation in a Tetranychus spider mite

Host plant quality mediates dispersal, oviposition, and sex allocation in a Tetranychus spider mite | 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 Host plant quality mediates dispersal, oviposition, and sex allocation in a Tetranychus spider mite Jhaman Kundun, Resona Simkhada, Svetla Sofkova-Bobcheva, Xiong Zhao He This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6772534/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Dispersal is a key adaptative strategy to escape deteriorating environments, and habitat selection by dispersers is critical to their own and offspring fitness. Using the haplodiploid spider mite Tetranychus ludeni Zacher as a model species, this study investigated how host plant quality influenced dispersal probability, habitat selection, and subsequent reproductive performances of mated females. We designed two dispersal scenarios, i.e., females were allowed to disperse from low-quality (LQ) or high-quality (HQ) habitat and select between LQ and HQ habitats. Results show that significantly more females dispersed from LQ habitats than from HQ habitats, and dispersers significantly preferring and settling in HQ habitats regardless of the dispersal scenarios. However, aggregating in HQ habitats resulted in higher number of eggs cumulated but also increased immature mortality. Individual females restrained reproductive output under the deteriorating environments. Egg size had no significant effect on egg hatching or immature survival. Females dispersed from LQ habitats produced significantly smaller eggs but maintained similar proportion of daughters compared to those remaining in LQ habitats. Females dispersed from HQ habitats produced eggs of similar size but significantly higher proportion of daughters. These results suggest that dispersing females might manipulate offspring sex ratio by lowing the fertilization threshold to fertilise relatively smaller eggs. Population density has less impact on egg size and offspring sex ratio. This study delivers insights into the dispersal and reproductive strategies of a haplodiploid spider mite, highlighting how host quality shapes adaptive responses in challenging environments. Habitat quality Habitat selection Population density Reproductive performance Tetranychus ludeni Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Introduction In herbivore-plant systems, host plants serve as the habitats for phytophagous animal development, growth, and reproduction (Jermy 1984 ; Meiners 2015 ). As both an essential environmental factor and a primary food resource, host plants significantly influence the occurrence, distribution, and population density of herbivores (Awmack and Leather 2002 ; Knolhoff and Heckel 2014 ). However, host plant quality often decline due to the increase of animal population density during the growing seasons (Li and Margolies 1993 ; Denno et al. 2000 ; Sandeson et al. 2002 ; Rhainds and Shipp 2003 ; Rhainds et al. 2005 ; Reisig et al. 2010 ), which will reduce the carrying capacity, the theoretical limit of population size sustained by the given resources of a particular habitat (Cadet et al. 2003 ; Chapman and Byron 2018 ). In such dynamic environments, phytophagous insects are subjected to strong selection pressure to maximize or maintain their fitness by avoiding unsuitable habitats and locating suitable ones (Bruce et al. 2005 ; Bruce and Pickett 2011 ). Previous theoretical studies (McPeek and Holt 1992 ; Parvinen and Egas 2004 ) suggest that dispersal is a key adaptation to the temporal variability of habitat quality. Biological dispersal, referring to the movement of individuals from their birth site to their breeding site (natal dispersal) (Johnson and Horvitz 2005 ; Ruf et al. 2011 ) or between breeding sites (breeding dispersal) (McCauley 2010 ) or both (Dieckmann et al. 1999 ; Clobert et al. 2012 ), is the primary mechanism enabling them to find suitable habitats that can support their survival and reproduction (den Boer 1990 ; Dieckmann et al. 1999 ; Clobert et al. 2012 ). Dispersal is driven by multiple selective forces, including the avoidance of competition with relatives, reduction of inbreeding risks, or spreading risk in spatially and temporally varying environments or demography (Duputié and Massol 2013 ). Dispersal involves three distinct stages, i.e., departure, transience, and settlement (Clobert et al. 2009 ; Bonte and Dahirel 2017 ), and each stage is subjected to selection pressures that minimize the overall costs of dispersal and maximize the fitness of dispersers and their offspring (Bonte et al. 2012 ; Travis et al. 2012 ). Among these stages, settlement associated with habitat selection is critical for the survival and reproduction success and thus the establishment and subsequent expansion of populations in new habitats (Kot et al. 1996 ; Phillips et al. 2010 ; Ronce and Clobert 2012 ; Renault et al. 2018 ; Williams et al. 2019 ). Phytophagous animals, through long-term adaptation and co-evolution, have developed sophisticated sensory and nervous systems (Martin et al. 2011 ; Bruce 2015 ) to detect and evaluate the quality of host plants (Bernays 2001 ; Bruce 2015 ; Burgueño et al. 2024 ; Karlsson Green et al. 2024 ). This ability to assess host suitability is crucial for future survival and reproductive success, as phytophagous insects usually have limited opportunities to make settlement decisions (Burgueño et al. 2024 ; Karlsson Green et al. 2024 ). Previous studies on phytophagous animal dispersal mostly focus on the determinations of mechanisms or factors affecting dispersal (e.g., Li and Margolies 1993 ; Sandeson et al. 2002 ; Poethke and Hovestadt 2002 ; Sloggett and Weisser 2002 ; Lombaert et al. 2006 ; Yano 2008 , Bowler and Benton 2009 ; McCauley 2010 ; Poethke et al. 2011 ; Bitume et al. 2011 , 2014 ; Pepi et al. 2016 ; Puzin et al. 2018 ; Plazio et al. 2020 ; Hewison et al. 2021 ), few research has examined the subsequent reproduction of dispersers in the new habitats (e.g., Khuhro et al. 2014 ; Nasu and Tokuda 2021 ; Zhou et al. 2021 , 2024 ). To date, how dispersers adjust their behaviours and reproductive strategies to adapt to the new habitats is largely unknown. Spider mites (Acari: Tetranychidae) consisting of over 1300 species (Migeon and Dorkeld 2024 ), and some species, such as Tetranychus urticae Koch, T. evansi Baker and Pritchard, T. kanzawai Kishida, and T. ludeni Zacher, are serious pests damaging many economic crops worldwide (Bolland et al. 1998 ; Zhang 2003 ). These mites feed by piercing the leaf tissues and sucking out plant contents with their sap-sucking mouthparts, leading to the reduced photosynthetic activities and visible damage such as yellow and white spots on leaves (Dhooria 2016 ). Spider mites often live in groups (Strong et al. 1997 ; Dhooria 2016 ), which benefit individuals from modifying plant biochemistry, such as breaking down the plant defense system, resulting in more favourable nutritional quality of the shared host plants (Kant et al. 2008 ; Rioja et al. 2017 ). However, Tetranychus species may build up the local populations quickly due to their short lifecycle and high fecundity, for example T. ludeni could complete development from egg to adult stage in about 10 days at 27 ºC and lay up to 110 eggs on suitable hosts (Adango et al. 2006 ). These life-history traits may result in scramble competition for food resources between individuals (Krips et al. 1998 ). When the population becomes large and dense, spider mites, especially the young, mated females, tend to disperse to search for new habitats due to food deficiency and habitat deterioration (Azandémè-Hounmalon et al. 2014 ; Schausberger et al. 2021 ; Zhou et al. 2021 ). Dispersal in spider mites mainly occurs in young females after mating (Mitchell 1973 ; Li and Margolies 1993 ) and primarily through active movement, either individually and collectively by walking (Kondo and Takafuji 1985 ; Yano 2008 ), although spider mites may disperse solitarily by walking (Yano 2008 ), although passive dispersal via wind is also observed (Brandenburg and Kennedy 1982 ; Jung 2005 ; Yano 2008 ; Clotuche et al. 2011 , 2013 ). Collective dispersal of spider mite females may benefit group members because it enables the establishment of new colonies, and the immediate cooperative construction of webs which would effectively protect them from predators (McMurtry et al. 1970 ; Sabelis and Bakker 1992 ; Schausberger et al. 2021 ) and from wind and rain (Davis 1952 ; Linke 1953 ; Schausberger et al. 2021 ). Moreover, habitat heterogeneity in quality is common in nature. Unlike the passively dispersing organisms who have no opportunity to choose where they settle, active dispersal allows them to select among breeding habitats that differ in key characteristics, such as size, quality, and conspecific density (Stamps, 2001 ). Both dispersal behaviours and subsequent reproduction of spider mites, for example the T. ludeni , have been partially assessed (Zhou et al. 2021 , 2024 ); however, those authors detect the subsequent reproduction of dispersing females by maintaining them individually on a leaf disc, how collective dispersers select habitats of various quality and whether they adjust the reproductive strategies in response to the environmental conditions of post-dispersal habitats (e.g., habitat quality and population density) need further investigations. In this study, we investigated the collective dispersal, habitat selection, and subsequent reproduction of the European native spider mite T. ludeni . We simulated scenarios in nature by allowing the mated females to disperse from the low- or high-quality habitats, and choose between the low- and high-quality habitats. We then recorded the reproductive performances of collective dispersing females in terms of number and size of eggs, immature survival, and offspring sex ratio. We tested the hypotheses that (1) females are more likely to disperse from the low-quality habitats compared to the high-quality ones; (2) females would select and settle in high-quality habitats rather than the low-quality ones, and (3) dispersing females would lay more and large eggs, and generate more female offspring in the high-quality habitats than in the low-quality ones. This study will deliver knowledge to the strategies of spider mite dispersal and reproduction in response to social environment changes. Materials and methods Mite colony and experimental conditions The colony of spider mite T. ludeni started from female adults collected from a garden in Palmerston North, New Zealand. Mites were reared on kidney bean plants ( Phaseolus vulgaris L.) grown in potting mix (19:9:10 NPK at 400 g/100 L) in plastic pots (8 cm bottom diameter × 10 cm top diameter × 9 cm height). The mite colony was maintained in nylon-framed cages (65 cm length × 50 cm width × 50 cm height, aperture size = 0.075 mm length × 0.075 mm width) in a bioassay room in the Entomology and IPM laboratory, Massey University, New Zealand, at 25 ± 1 ºC, 40 ± 10% RH and 14:10 h (L:D) photoperiod. Three- to 5-week-old plants were used to maintain the colony, while first expanded plant leaves (1 to 2 weeks old) were used for experiments. Mites were reared for at least three successive generations prior to experiments. Experiments were carried out under the above-mentioned environmental conditions. Dispersal and habitat selection To prepare T. ludeni female adults for experiments, five female adults randomly selected from the colony were introduced onto a bean leaf square (2 cm length × 2 cm width) placed upside down on a water saturated cotton pad in a Petri dish (1.0 cm height × 9.5 cm diameter) with a mesh-covered opening (1.0 cm diameter) on the lid for ventilation. Females were allowed to lay eggs for 24 h, after which they were removed to another leaf square for egg laying. Thirty eggs remained on the leaf square were allowed to develop to deutonymphal stage. Although some males would emerge from the 30 eggs, we introduced 10 male adults developing from eggs laid by virgin females onto the leaf squares before deutonymphs emerged to adults, which ensured females mated at emergence. The 1-day-old mated females were used for experiments. To prepare low-quality (LQ) habitats for experiments, 10 female adults were randomly selected from the colony and introduced onto a bean leaf square (1 cm length × 1 cm width) in a Petri dish as did above. Mites were allowed to feed on the leaf square for five days. After which time, all mites, eggs, silks/webs, and faeces were removed using a fine brush, and the leaf square was washed by tap water to eliminate or at least reduce the traces left by the previous mites. The leaf squares were air-dried under laboratory conditions. Fresh plant leaf squares without mites infested were treated as high-quality (HQ) habitats. To investigate the dispersal and habitat selection behaviors of mites, two treatments were set up: (1) females dispersed from LQ habitat to select between the LQ and HQ habitats, and (2) females dispersed from HQ habitat to select between the LQ and HQ habitats. There were 15 replicates for each treatment. For each replicate, 15 1-day-old mated females prepared above were introduced onto a LQ or HQ habitat placed on the centre of a cotton pad in a Petri dish and bridged to one LQ and one HQ habitat by two parafilm strips (2 cm length × 0.5 cm width). The number of individuals residing on or dispersing to a habitat was counted after 24 hours. Oviposition and sex allocation of resident and dispersing mites (dup: abstract ?) To test the consequent effects of habitat selection on reproductive performance of female mites, the three leaf squares along with the cotton pad in a Petri dish were separated and individually transferred onto a new Petri dish. The resident and dispersing females were allowed to lay eggs in situ for five days, for two reasons, (1) maintaining the consistent social environments (i.e., habitat, and population size and density) for the resident or dispersed individuals, and (2) mortality occurring on sixth day of adult life. As some eggs might hatch within four days, the total number of eggs per habitat was counted as the sum of newly laid unhatched eggs and emerged larvae/nymphs, and the number of eggs per female was also calculated as total number of eggs /number of females. As egg size does not change before hatching (three days after oviposition under the experimental conditions) (Weerawansha et al. 2024 ), all or a maximum of 20 eggs were randomly selected from a habitat on the 3rd day of oviposition and the egg diameter as measured under a stereomicroscope (Leica MZ12, Germany) connected to a digital camera (Olympus SC30, Japan) and an imaging software (CellSens® GS-ST-V1.7, Olympus, Japan). The spider mite eggs are sphere shape, and their size was then calculated as: volume = 4/3π r 3 , where r is the radius (= diameter/2). All resident and dispersing females were removed from the habitats on the 6th day. The egg hatching rate and immature survival rate were also calculated. The newly emerged adults were sexed, and the proportion of daughters was calculated. Statistical analysis Data were analysed using SAS 9.13 (SAS Institute Inc, USA), with a significant level of α = 0.05. A generalized linear model (GLIMMIX Procedure) with a binomial distribution and a logit-link function in the model was used to compare the difference in dispersal behaviour, egg hatch and immature survival rate, and proportion of offspring daughters between the resident and selected habitats and to determine proportion of offspring daughters over the female density. Another generalized linear model (GLIMMIX Procedure) with a poisson or gamma distribution and a logit-link function in the model was applied to compare the difference in total number of eggs laid by all residing/dispersing females, number of eggs laid by each female, and egg size between the resided and selected habitats and to test their relationship to female density. A Tukey-Kramer test was applied to perform the multiple comparisons. A linear regression (GLM Procedure) as applied to assess the relationships between immature survival rate and larva number, between immature survival rate and egg size, and between proportion of daughters and egg size. Results Dispersal and habitat selection Female mites exhibited significantly higher dispersal probability when they initially settled in low-quality (LQ) habitats compared to that in high-quality (HQ) habitats (64.67 ± 3.10% vs. 29.58 ± 3.10%) ( F 1,30 = 56.94, P < 0.0001). When initially settled in LQ habitats, mites were significantly more likely to disperse to HQ habitats rather than to reside in LQ habitats or to disperse to LQ habitats ( F 2,45 = 39.52, P < 0.0001) (Fig. 1 a). When initially settled in HQ habitats, mites were significantly more likely to reside in HQ habitats rather than to disperse to HQ or LQ habitats ( F 2,45 = 92.65, P < 0.0001) (Fig. 1 b). Oviposition and sex allocation of resident and dispersing mites When dispersed from LQ habitats, the total number of eggs laid by females dispersing to HQ habitats was significantly higher than that laid by females residing in the LQ habitats, with a significant lower number of eggs laid by females dispersing to LQ habitats ( F 2,39 = 310.68, P < 0.0001) (Fig. 2 a); when dispersed from HQ habitats, the total number of eggs laid by the resident females was significantly higher than those laid by females dispersing to the HQ habitats, with a significant lower number of eggs laid by females dispersing to LQ habitats ( F 2,39 = 622.87, P < 0.0001) (Fig. 2 b). The egg number increased significantly with the increasing number of mites on a habitat regardless of the dispersing scenario except females dispersing from LQ to HQ habitats ( F 1,13 = 143.45, P < 0.0001 for LQ-Resident; F 1,14 = 2.73, P = 0.12 for LQ-HQ; F 1,9 = 10.13, P = 0.0111 for LQ-LQ; F 1,14 = 14.65, P = 0.0018 for HQ-Resident; F 1,13 = 51.26, P < 0.0001 for HQ-HQ; F 1,9 = 200.97, P < 0.0001 for HQ-LQ) (Fig. 2 c, 2 d); however, for the scenario of dispersing from LQ habitats the rate of increase (i.e., slope) was significantly greater for females residing in the LQ habitats than those dispersing to HQ or LQ habitats ( F 2,36 = 21.51, P < 0.0001) (Fig. 2 c), while for the scenario of dispersing from HQ habitats the rate of increase was significantly greater for mites dispersing to the LQ habitats than those dispersing to HQ habitats, with a significant slower increasing rate detected for mites dispersing to HQ habitats ( F 2,36 = 89.29, P < 0.0001) (Fig. 2 d). When females resided in or dispersed from LQ habitats to HQ or LQ habitats, they laid similar number of eggs ( F 2,39 = 3.10, P = 0.06) (Fig. 3 a); while the number of eggs laid by each female was significantly higher when females dispersed from HQ habitats to LQ habitats than that laid by that dispersing from HQ habitats to HQ habitats with a significant fewer eggs laid by the resident mites ( F 2,39 = 27.60, P < 0.0001) (Fig. 3 b). The egg number by each female decreased with the increasing number of mites, but significance was found for females that resided in the LQ habitats, dispersed from LQ habitats to LQ habitats, or from HQ habitats to HQ habitats ( F 1,13 = 12.59, P = 0.0036 for LQ-Resident; F 1,14 = 3.76, P = 0.07 for LQ-HQ; F 1,9 = 49.52, P < 0.0001 for LQ-LQ; F 1,14 = 2.71, P = 0.12 for HQ-Resident; F 1,13 = 4.86, P = 0.046 for HQ-HQ; F 1,9 = 0.09, P = 0.77 for HQ-LQ) (Fig. 3 c, 3 d). For the scenario of dispersing from LQ habitats, the decrease of eggs laid (i.e., slope) was significantly faster for females dispersing from LQ habitats to LQ habitats ( F 2,36 = 8.92, P = 0.0007) (Fig. 3 c), while for the scenario of dispersing from HQ habitats, the rate of decrease was not significantly different between the residing and dispersing females ( F 2,36 = 0.78, P = 0.47) (Fig. 3 d). Females residing in LQ habitats produced significantly larger eggs than the dispersing females, with a significantly smaller egg size detected for females that dispersed from LQ to LQ habitats ( F 2,39 = 4.39, P < 0.0191) (Fig. 4 a); however, when females initially settled in HQ habitats, there was no significant difference in egg size between the resident and dispersing females ( F 2,39 = 0.64, P = 0.54) (Fig. 4 b). Female density in a habitat had no significant effect on egg size, except when females resided in HQ habitats the egg size significantly increased with female number ( F 1,13 = 0.56, P = 0.47 for LQ-Resident; F 1,14 = 0.64, P = 0.35 for LQ-HQ; F 1,9 = 0.01, P = 0.93 for LQ-LQ; F 1,14 = 14.00, P = 0.0022 for HQ-Resident; F 1,13 = 0.01, P = 0.91 for HQ-HQ; F 1,9 = 0.65, P = 0.44 for HQ-LQ) (Fig. 4 c, 4 d). There was no significant difference in regression slop between habitats in each dispersing scenario ( F 2,36 = 0.22, P = 0.81, Fig. 4c; F 2,36 = 0.78, P = 0.47; Fig. 4 d). There was no significant difference in egg hatch rate between the resident and dispersing females in either dispersal scenario ( F 2,39 = 1.71, P = 0.32 for dispersing from LQ habitat; F 2,39 = 1.60, P = 0.21 for dispersing from HQ habitat) (Fig. 5 a). However, when mites dispersed from LQ habitats, immature survival was significantly lower for those dispersing to HQ habitats compared to those residing in the LQ habitats, with a significant higher immature survival for those dispersing to LQ habitats ( F 2,39 = 16.51, P < 0.0001). In contrast, when mites dispersed from HQ habitats, immature survival was significantly lower for those residing in HQ habitats compared to those dispersing to the HQ or LQ habitats ( F 2,39 = 22.91, P < 0.0001) (Fig. 5 b). When mites dispersed from LQ habitats, there was no significant difference in offspring sex ratio between the resident and dispersing females ( F 2,39 = 1.69, P = 0.20) (Fig. 6 a); while when mites dispersed from HQ habitats, the proportion of daughters was significantly higher for females dispersing to HQ and LQ habitats than for resident females ( F 2,38 = 9.85, P = 0.0004) (Fig. 6 b). Female density in a habitat had no significant effect on sex allocation, except when females resided in HQ habitats the proportion of daughters significantly increased with female number ( F 1,13 = 2.22, P = 0.16 for LQ-Resident; F 1,14 = 3.81, P = 0.07 for LQ-HQ; F 1,9 = 1.12, P = 0.32 for LQ-LQ; F 1,14 = 15.39, P = 0.0015 for HQ-Resident; F 1,13 = 3.45, P = 0.09 for HQ-HQ; F 1,8 = 0.01, P = 0.92 for HQ-LQ) (Fig. 6 c, 6 d). There was no significant difference in regression slop between habitats in each dispersing scenario ( F 2,36 = 0.34, P = 0.72, Fig. 6c; F 2,36 = 0.73, P = 0.49; Fig. 6 d). As shown in Fig. 7 a, immature survival significantly decreased with increasing population density of mite larvae ( F 1,82 = 11.20, P = 0.0012). Egg size did not have significant impact on immature survival ( F 1,82 = 0.01, P = 0.93) (Fig. 7 b) and proportion of daughters ( F 1,81 = 0.21, P = 0.15) (Fig. 7 c). Discussion Dispersal and habitat selection Phytophagous organisms are subject to various selective pressures in nature due to the spatial and temporal variability of habitat quality. Spider mites can expand their local populations rapidly due to their specific life history traits, e.g., group living (Strong et al. 1997 ; Le Goff et al. 2010 ; Clotuche et al. 2011 ; Yano 2012 ), short lifecycle (Shih et al. 1976 ; Adango et al. 2006 ; Bounfour and Tanigoshi 2001 ; Gotoh et al. 2015 ; Tuan et al. 2016 ), high fecundity (Adango et al. 2006 ; Bounfour and Tanigoshi 2001 ; Gotoh et al. 2015 ; Tuan et al. 2016 ), and extremely high female-biased offspring sex ratio (Carey and Bradley 1982 ; Macke et al. 2011a ; Weerawansha et al. 2023a ). However, these life-history traits may result in scramble competition between individuals, inducing an overexploitation of and quick cumulative excrement on host plants, and a fast depletion of resource and decrease of habitat quality (Hussey and Parr 1963 ; Krips et al. 1998 ; Weerawansha 2023b). Our results demonstrate that T. ludeni females were more likely to disperse from low-quality (LQ) habitats (64.7%) than from the high-quality (HQ) habitats (29.6%), indicating that declining resource quality triggers dispersal in spider mites (Jeppson et al. 1975 ; Boykin and Campbell 1984 ; Azandémè-Hounmalon et al. 2014 ). Dispersing from deteriorating environments to more favourable habitats is an adaptive strategy for spider mites to maximize their own and offspring fitness (Azandémè-Hounmalon et al. 2014 ; Schausberger et al. 2021 ; Zhou et al. 2021 ). Habitat selection is a key determination of dispersal success. Spider mites usually prefer young leaves with a high level of nutrients (e.g., nitrogen) for feeding and oviposition (Watson 1964 ; Suski and Badowska 1975 ; Mellors and Propts 1983 ; Wermelinger et al. 1985 ; Wilson 1994 ). As expected, dispersing T. ludeni females significantly preferred and settled in the HQ habitats compared to the LQ ones (Fig. 1 ). The results may have two implications. First, spider mites are able to assess plant quality and alter their behavior accordingly (Wilson 1994 ), and the HQ habitats are expected to slow density-dependent declines in fitness (Avgar et al. 2020 ). Second, dispersal in spider mites is a non-random process. In many web-spinning arthropods, such as the social spiders, social caterpillars, and spider mites, silk is a vector for collective behaviour (Fernández Ferrari et al. 2013). Yano ( 2008 ) shows evidence that in T. urticae , silk is strongly attractive, and dispersing females could distinguish between silk trails laid by the preceding solitary or grouping females, which provides positive feedback to the dispersers and results in collective dispersal by following the path with more silk on it. Our results further reveal that even though dispersing mites had opportunities to choose between the HQ and LQ habitats, some individuals still settled in LQ habitats regardless of the dispersal scenarios (Fig. 1 ). Bruce ( 2015 ) argues that dispersers may make ‘mistakes’ and settle in poor quality hosts, although phytophagous species are under selection pressure to find quality hosts and have evolved a fine-tuned sensory system for the detection of host cues and avoidance of unsuitable hosts (Martin et al. 2011 ; Bruce 2015 ). Rushing et al. ( 2021 ) further hypothesize that dispersers who can reliably assess habitats and are free to settle should select the highest quality habitats until density-dependent mechanisms reduce expected reproductive success to the point, after which individuals would have higher fitness by settling in lower quality patches. The hypothesis may explain the probability of some T. ludeni females selecting the LQ habitats, especially when significantly more individuals dispersed from the LQ habitats (Fig. 1 ). Oviposition and sex allocation of resident and dispersing mites Spider mites live in groups (Strong et al. 1997 ; Dhooria 2016 ), which may benefit individuals from modifying plant biochemistry by breaking down the plant defense system and resulting in more favourable nutritional quality of the shared host plants (Kant et al. 2008 ; Rioja et al. 2017 ), and reducing the intensity of individual web production (thinner, shorter, and/or fewer threads) and thus saving energy and nutrients (protein, and amino acids) in web production that can be invested in reproduction (Hazan et al. 1974 ; Le Goff et al. 2010 ). However, our results show that individual females remained in HQ habitats significantly reduced the number of eggs laid, and when T. ludeni females dispersed from LQ habitats to the HQ ones, they did not significantly increase reproduction (Fig. 3 a, 3 b). These may be attributed to two reasons. First, the high population density (Fig. 1 ) and subsequent high egg density (Fig. 2 a, 2 b) in the LQ-HQ and resident-HQ habitats might have approached the carrying capacity; therefore, individual females restrained reproductive output at crowded environments, which may be an adaptive strategy to reduce future food competition and ensure offspring survival (Weerawansha et al. 2024 ). Second, residing in a large group will induce costs on foraging and feeding efficiency due to the higher interference among group members (Bilde et al. 2007 ; Estevez et al. 2007 ; Grove 2012 ; Wong et al. 2013 ; Li and Zhang 2021 ; Tinsley Johnson et al. 2021 ), which may restrain of female fecundity (Krips et al. 1998 ; Clotuche 2011 ; Bitume et al. 2013 ). Furthermore, individual females also restrained reproduction in response to the increasing population density (Fig. 3 c, 3 d), especially in LQ-LQ habitats females had a significant faster decrease of reproduction with increase population density (Fig. 3 c) despite a lower population density (Fig. 1 a), indicating that the population density-dependent reproductive constraint in T. ludeni is regulated by habitat quality. A trade-off between egg number and egg size is frequently reported (Smith and Fretwell 1974 ; Parker and Begon 1986 ; Stearns 1989 ; Fox and Czesak 2000 ; Macke et al. 2012 ; Walzer and Schausberger 2015 ; Maenoa et al. 2020), and larger egg size may facilitate survival of immature stages (Goulden et al. 1987 ; Fox 1994 ; Johnston and Leggett 2002 ; Pick et al. 2016 ; Xu et al. 2019 ). Unfortunately, in this study, reproductive constraint in LQ-HQ habitats (Fig. 3 a) or oviposition reduction in HQ-Resident habitats (Fig. 3 b) did not significantly increased egg size (Fig. 4 a, 4 b), and increasing egg size did not significantly promoted egg hatch rate (Fig. 5 a), and immature survival rate (Figs. 4 a, 7 b). Previous studies also reported that when females are shifted between habitats of high to low population density, a trade-off between egg number and size, and egg size had little impact on reproductive fitness in terms of immature survival (Weerawansha et al. 2022a , b ). Therefore, it may be concluded that when future environmental conditions are uncertain or unpredictable, egg size may not be a reliable indicator of offspring fitness (Wiklund and Persson 1983 ; Karlsson and Wiklund 1985 ; McEdward and Carson 1987 ; Lalonde 2005 ; Morrongiello et al. 2012 ; Weerawansha et al. 2022a , b ). While our results reveal that increasing population of active immature individuals significantly decreased immature survival rate (Fig. 7 a), because the nature of scramble resource competition between individual spider mites in habitats with high population density will result in a fast depletion of local resource and decrease of habitat quality (Hussey and Parr 1963 ; Krips et al. 1998 ; Weerawansha 2023a), which may induce the high mortality before they develop to adults due to the deficient nutrients. Therefore, in this study, the carrying capacity of habitats is the major factor affecting the reproductive performances of T. ludeni females and fitness of offspring. In spider mites, the mated females are capable of manipulating offspring sex ratio by selectively fertilizing (Young et al. 1986 ; Roeder et al. 1996 ; Macke et al. 2011a ; Weerawansha et al. 2022b ) when the egg size exceeds a threshold value (Macke et al. 2011b ). However, results of this study show that increasing egg size did not significantly facilitate the production of daughters (Fig. 7 c). Alternatively, in the scenario of mites dispersing from LQ habitats, dispersing females laid significantly smaller eggs (Fig. 4 a) but produced similar proportion of daughters regardless of the quality of selected habitats (Fig. 6 a); while in the scenario of mites dispersing from HQ habitats, dispersing females laid eggs of similar size (Fig. 4 b) but produced significantly higher proportion of daughters regardless the quality of selected habitats (Fig. 6 b). These results suggest that the dispersing females could adjust the fertilization threshold to a lower level and fertilise relatively smaller eggs. Weerawansha et al. ( 2022a , b ) also demonstrated that when T. ludeni females aggregate into a large group, they lay significantly smaller eggs but produced a significantly higher female-biased sex ratio, which is attributed to the flexibility of egg fertilization. Furthermore, our results show that compared to the habitat quality, female density had less impact on egg size (Fig. 4 a, 4 b) and daughter production (Fig. 6 a, 6 b); therefore, the impact of habitat quality was superior to that of population density on the adjustment of fertilization threshold that determines the offspring sex ratio in T. ludeni . Many studies have demonstrated that dispersal is dependent on both disperser phenotype and the local environment (Bowler and Benton 2005 ; Clobert et al. 2009 ; Bonte et al. 2014 ; Hollander et al. 2014 ; Baines et al. 2019 ). In spider mites, when the local resources are depleted, the reproductive females will disperse to seek new habitats for the next generation either by ambulatory dispersal within a habitat (Tien et al. 2011 ; Azandémè-Hounmalon et al. 2014 ; Li and Zhang 2021 ; Schausberger et al. 2021 ; Zhou et al. 2021 , 2024 ) or aerial dispersal for a long distance (Boykin and Campbell 1984 ; Smitley and Kennedy 1988 ; Li and Margolies 1993 ; Osakabe et al. 2008 ; Clotuche et al. 2011 ). We reveal that relatively higher proportion of T. ludeni females (35%) still resided in the low-quality habitats. Whether spider mites have evolved phenotypic plasticity for dispersal, and whether the philopatric and dispersing phenotypes have developed different strategies in oviposition and sex allocation remain further investigations. Conclusion Tetranychus ludeni females are more likely to disperse from the low-quality (LQ) habitats than from the high-quality (HQ) ones, and the dispersing females significantly prefer and settle in HQ habitats regardless the dispersal scenarios. The significantly high population density and total number of eggs laid in HQ habitats result in restrained oviposition of females and higher mortality of immature offspring. Egg size has no significant effect on egg hatching and immature survival. Females that disperse from LQ habitats to the HQ or LQ habitats produce significantly smaller eggs but similar proportion of daughters, and females that disperse from HQ habitats to the HQ or LQ habitats produce eggs of similar size but significantly higher proportion of daughters, suggesting that dispersing females may manipulate offspring sex ratio by lowing the fertilization threshold to fertilise relatively smaller eggs. However, population density has less impact on egg size and offspring sex ratio. Our results indicate that the dispersal and habitat selection of T. ludeni females are mediated by host plant quality, and the dispersing females could adjust their reproductive strategies to maximise their own and offspring fitness. Declarations Acknowledgements We thank Professor Z.-Q. Zhang for identification of this spider mite to species, and Plant Growth Unit (PGU), Massey University for providing potting mix for bean plant growth. Funding This work was jointly funded by the Manaaki New Zealand Scholarships. Competing interests The authors declare no competing interests. Ethical approval Not applicable. Consent for publication Not applicable. Availability of data and materials The datasets used and/or analysed during the current study are available from the corresponding author on reasonable request. Author contributions JK and XZH conceived and designed the experiments. JK and RS collected the data. JK and XZH analysed the data. JK prepared the first draft. All authors contributed to manuscript revision. References Adango E, Onzo A, Hanna R, James B, Atachi P (2006) Comparative demography of the spider mite, Tetranychus ludeni , on two host plants in West Africa. J Inse Sci 6:1–9. https://doi.org/10.1673/031.006.4901 Avgar T, Betini GS, Fryxell JM (2020) Habitat selection patterns are density dependent under the ideal free distribution. J Anim Ecol 89:2777–2787. https://doi.org/10.1111/1365-2656.13352 Awmack CS, Leather SR (2002) Host plant quality and fecundity in herbivorous insects. Annu Rev Entomol 47:817–844. https://doi.org/10.1146/annurev.ento.47.091201.145300 Azandémè-Hounmalon GY, Fellous S, Kreiter S, Fiaboe KK, Subramanian S, Kungu M, Martin T (2014) Dispersal behavior of Tetranychus evansi and T. urticae on tomato at several spatial scales and densities: implications for integrated pest management. PLoS ONE 9:e95071. https://doi.org/10.1371/journal.pone.0095071 Baines CB, Ferzoco IMC, McCauley SJ (2019) Phenotype‐by‐environment interactions influence dispersal. J Anim Ecol 88:1263–1274. https://doi.org/10.1111/1365-2656.13008 Bernays EA (2001) Neural limitations in phytophagous insects: implications for diet breadth and evolution of host affiliation. Annu Rev Entomol 46:703–727. https://doi.org/10.1146/annurev.ento.46.1.703 Bilde T, Coates KS, Birkhofer K, Bird T, Maklakov AA, Lubin Y, Avilés L (2007) Survival benefits select for group living in a social spider despite reproductive costs. J Evol Biol 20:2412–2426. https://doi.org/10.1111/j.1420-9101.2007.01407.x Bitume EV, Bonte D, Magalhães S, Martin GS, Van Dongen S, Bach F, Anderson JM, Olivieri I, Nieberding CM (2011) Heritability and artificial selection on ambulatory dispersal distance in Tetranychus urticae : Effects of density and maternal effects. PLoS ONE 6:1–9. https://doi.org/10.1371/journal.pone.0026927 Bitume EV, Bonte D, Ronce O, Olivieri I, Nieberding CM (2014) Dispersal distance is influenced by parental and grand-parental density. Proc R Soc B 281:1–8. https://doi.org/10.1098/rspb.2014.1061 Bitume EV, Nieberding CM, Ronce O, Bach F, Flaven E, Olivieri I, Bonte D (2013) Density and genetic relatedness increase dispersal distance in a subsocial organism. Ecol Lett 16:430–437. https://doi.org/10.1111/ele.12057 Bolland HR, Gutierrez J, Flechtmann CH (1998) World catalogue of the spider mite family (Acari: Tetranychidae). Brill Academic Publishers, Leiden, Boston. Bonte D, Dahirel M (2017) Dispersal: A central and independent trait in life history. Oikos 126:472–479. https://doi.org/10.1111/oik.03801 Bonte D, De Roissart A, Wybouw N, Van Leeuwen T (2014) Fitness maximization by dispersal: Evidence from an invasion experiment. Ecology 95:3104–3111. https://doi.org/10.1890/13-2269.1 Bonte D, Van Dyck H, Bullock JM et al. (2012) Costs of dispersal. Biol Rev 87:290–312. https://doi.org/10.1111/j.1469-185X.2011.00201.x Bounfour M, Tanigoshi LK (2001) Effect of temperature on development and demographic parameters of Tetranychus urticae and Eotetranychus carpini borealis (Acari: Tetranychidae). Ann Entomol Soc Am 94:400–404. https://doi.org/10.1603/0013-8746(2001)094[0400:EOTODA]2.0.CO;2 Bowler DE, Benton TG (2005) Causes and consequences of animal dispersal strategies: relating individual behaviour to spatial dynamics. Biol Rev 80:205–225. https://doi.org/10.1017/S1464793104006645 Bowler DE, Benton TG (2009) Variation in dispersal mortality and dispersal propensity among individuals: the effects of age, sex and resource availability. J Anim Ecol 78:1234–1241. https://doi.org/10.1111/j.1365-2656.2009.01580.x Boykin L, Campbell W (1984) Wind dispersal of the twospotted spider mite (Acari: Tetranychidae) in North Carolina peanut fields. Environ Entomol 13:221–227. https://doi.org/10.1093/ee/13.1.221 Brandenburg R, Kennedy G (1982) Intercrop relationships and spider mite dispersal in a corn/peanut agro‐ecosystem. Entomol Exp Appl 32:269–276. https://doi.org/10.1111/j.1570-7458.1982.tb03217.x Bruce TJA (2015) Interplay between insects and plants: Dynamic and complex interactions that have coevolved over millions of years but act in milliseconds. J Exp Bot 66:455–465. https://doi.org/10.1093/jxb/eru391 Bruce TJA, Pickett JA (2011) Perception of plant volatile blends by herbivorous insects – Finding the right mix. Phytochemistry 72:1605–1611. https://doi.org/10.1016/j.phytochem.2011.04.011 Bruce TJA, Wadhams LJ, Woodcock CM (2005) Insect host location: A volatile situation. Trends Plant Sci 10:269–274. https://doi.org/10.1016/j.tplants.2005.04.003 Burgueño AP, Amorós ME, Deagosto E, Davyt B, Díaz M, González A, Rossini C (2024) Preference and performance in an herbivorous coccinellid beetle: A comparative study of host plant defensive traits, insect preference, and survival. Arthropod-Plant Interact 18:617–636. https://doi.org/10.1007/s11829-023-10004-x Cadet C, Ferrière R, Metz JAJ, van Baalen M (2003) The evolution of dispersal under demographic stochasticity. Am Nat 162:427–441. https://doi.org/10.1086/378213 Carey J, Bradley J (1982) Developmental rates, vital schedules, sex ratios and life tables for Tetranychus urticae , T. turkestani and T. pacificus (Acarina: Tetranychidae) on cotton. Acarologia 23:333–345. Chapman EJ, Byron CJ (2018) The flexible application of carrying capacity in ecology. Glob Ecol Conserv 13:e00365. https://doi.org/10.1016/j.gecco.2017.e00365 Clobert J, Baguette M, Benton TG, Bullock JM (2012) Dispersal Ecology and Evolution. Oxford University Press. Clobert J, Le Galliard JF, Cote J, Meylan S, Massot M (2009) Informed dispersal, heterogeneity in animal dispersal syndromes and the dynamics of spatially structured populations. Ecol Lett 12:197–209. https://doi.org/10.1111/j.1461-0248.2008.01267.x Clotuche G (2011) The silk as a thread to understand social behaviour in the weaving mite Tetranychus urticae . Thesis, UCL-Université Catholique de Louvain, Belgium. Clotuche G, Mailleux AC, Astudillo Fernández A, Deneubourg JL, Detrain C, Hance T (2011) The formation of collective silk balls in the spider mite Tetranychus urticae Koch. PLoS ONE 6:e18854. https://doi.org/10.1371/journal.pone.0018854 Clotuche G, Navajas M, Mailleux AC, Hance T (2013) Reaching the ball or missing the flight? Collective dispersal in the two-spotted spider mite Tetranychus urticae . PLoS ONE 8:e77573. https://doi.org/10.1371/journal.pone.0077573 Davis DW (1952) Influence of population density on Tetranychus multisetis . J Econ Entomol 45:652–652. https://doi.org/10.1093/jee/45.4.652 den Boer PJ (1990) The survival value of dispersal in terrestrial arthropods. Biol Conserv 54:175–192. https://doi.org/10.1016/0006-3207(90)90050-Y Denno RF, Peterson MA, Gratton C, Cheng J, Langellotto GA, Huberty AF, Finke DL (2000) Feeding‐induced changes in plant quality mediate interspecific competition between sap‐feeding herbivores. Ecology 81:1814–1827. https://doi.org/10.1890/0012-9658(2000)081[1814:FICIPQ]2.0.CO;2 Dhooria MS (2016) Fundamentals of Applied Acarology. Springer, Singapore. Dieckmann U, O'Hara B, Weisser W (1999) The evolutionary ecology of dispersal. Trends Ecol Evol 14:88–90. https://doi.org/10.1016/S0169-5347(98)01571-7 Duputié A, Massol F (2013) An empiricist's guide to theoretical predictions on the evolution of dispersal. Interface Focus 3:20130028. https://doi.org/10.1098/rsfs.2013.0028 Estevez I, Andersen IL, Naevdal E (2007) Group size, density and social dynamics in farm animals. Appl Anim Behav Sci 103:185–204. https://doi.org/10.1016/j.applanim.2006.05.025 Fernández Ferrari MC, Schausberger P (2013) From repulsion to attraction: Species- and spatial context-dependent threat sensitive response of the spider mite Tetranychus urticae to predatory mite cues. Naturwissenschaften 100:541–549. https://doi.org/10.1007/s00114-013-1050-5 Fox CW (1994) The influence of egg size on offspring performance in the seed beetle, Callosobruchus maculatus . Oikos 71:321–325. https://doi.org/10.2307/3546280 Fox CW, Czesak ME (2000) Evolutionary ecology of progeny size in arthropods. Ann Rev Entomol 45:341–369. https://doi.org/10.1146/annurev.ento.45.1.341 Gotoh T, Moriya D, Nachman G (2015) Development and reproduction of five Tetranychus species (Acari: Tetranychidae): Do they all have the potential to become major pests? Exp Appl Acarol 66:453–479. https://doi.org/10.1007/s10493-015-9919-y Goulden CE, Henry L, Berrigan D (1987) Egg size, postembryonic yolk, and survival ability. Oecologia 72:28–31. https://doi.org/10.1007/BF00385040 Grove M (2012) Space, time, and group size: A model of constraints on primate social foraging. Anim Behav 83:411–419. https://doi.org/10.1016/j.anbehav.2011.11.011 Hazan A, Gerson U, Tahori A (1974) Spider mite webbing. I. The production of webbing under various environmental conditions. Acarologia 16:68–84. Hewison AJM, Morellet N, Debeffe L, Cargnelutti B, Gaillard JM, Cagnacci F, Gehr B, Kröschel M, Heurich M, Coulon A, Kjellander P, Börger L, Focardi S (2021) Sex differences in condition dependence of natal dispersal in a large herbivore: Dispersal propensity and distance are decoupled. Proc R Soc B Biol Sci 288:20202947. https://doi.org/10.1098/rspb.2020.2947 Hollander J, Verzijden M, Svensson E, Brönmark C (2014) Animal movement across scales. In: Hansson LA, Åkesson S (eds) Animal Movement Across Scales. Oxford University Press, pp 110–125. Hussey NW, Parr WJ (1963) Dispersal of the glasshouse red spider mite Tetranychus urticae Koch (Acarina, Tetranychidae). Entomol Exp Appl 6:207–214. https://doi.org/10.1111/j.1570-7458.1963.tb00619.x Jeppson LR, Keifer HH, Baker EW (1975) Mites Injurious to Economic Plants. University of California Press. Jermy T, 1984. Evolution of insect/host plant relationships. Am Nat 124:609–630. http://www.jstor.org/stable/2461372 Johnson DM, Horvitz CC (2005) Estimating postnatal dispersal: Tracking the unseen dispersers. Ecology 86:1185–1190. https://doi.org/10.1890/04-0974 Johnston TA, Leggett WC (2002) Maternal and environmental gradients in the egg size of an iteroparous fish. Ecology 83:1777–1791. https://doi.org/10.1890/0012-9658(2002)083[1777:MAEGIT]2.0.CO;2 Jung C (2005) Some evidences of aerial dispersal of twospotted spider mites from an apple orchard into a soybean field. J Asia-Pac Entomol 8:279–283. https://doi.org/10.1016/S1226-8615(08)60246-0 Kant MR, Sabelis MW, Haring MA, Schuurink RC (2008) Intraspecific variation in a generalist herbivore accounts for differential induction and impact of host plant defences. Proc R Soc B Biol Sci 275:443–452. https://doi.org/10.1098/rspb.2007.1277 Karlsson Green K, Chiara De P, Maria L, Peter A (2024) Population comparison of innate and plastic host plant preference and performance in a polyphagous insect. Front Ecol Evol 12:1426923. https://doi.org/10.3389/fevo.2024.1426923 Karlsson B, Wiklund C (1985) Egg weight variation in relation to egg mortality and starvation endurance of newly hatched larvae in some satyrid butterflies. Ecol Entomol 10:205–211. https://doi.org/10.1111/j.1365-2311.1985.tb00549.x Khuhro NH, Biondi A, Desneux N, Zhang L, Zhang Y, Chen H (2014) Trade-off between flight activity and life-history components in Chrysoperla sinica . BioControl 59:219–227. https://doi.org/10.1007/s10526-014-9560-4 Knolhoff LM, Heckel DG (2014) Behavioral assays for studies of host plant choice and adaptation in herbivorous insects. Ann Rev Entomol 59:263–278. https://doi.org/10.1146/annurev-ento-011613-161945 Kondo A, Takafuji A (1985) Resource utilization pattern of two species of tetranychid mites (Acarina: Tetranychidae). Res Popul Ecol 27:145–157. https://doi.org/10.1007/BF02515487 Kot M, Lewis MA, van den Driessche P (1996) Dispersal data and the spread of invading organisms. Ecology 77:2027–2042. https://doi.org/10.2307/2265698 Krips OE, Witul A, Willems PEL, Dicke M (1998) Intrinsic rate of population increase of the spider mite Tetranychus urticae on the ornamental crop gerbera: Intraspecific variation in host plant and herbivore. Entomol Exp Appl 89:159–168. https://doi.org/10.1046/j.1570-7458.1998.00395.x Lalonde R (2005) Egg size variation does not affect offspring performance under intraspecific competition in Nasonia vitripennis , a gregarious parasitoid. J Anim Ecol 74:630–635. https://doi.org/10.1111/j.1365-2656.2005.00958.x Le Goff GJ, Mailleux AC, Detrain C, Deneubourg JL, Clotuche G, Hance T (2010) Group effect on fertility, survival and silk production in the web spinner Tetranychus urticae (Acari: Tetranychidae) during colony foundation. Behaviour 147:1169–1184. https://doi.org/10.1163/000579510X510980 Li G-Y, Zhang Z-Q (2021) The costs of social interaction on survival and reproduction of arrhenotokous spider mite Tetranychus urticae . Entomol Gen 41:49–57. https://doi.org/10.1127/entomologia/2020/0911 Li J, Margolies DC (1993) Effects of mite age, mite density, and host quality on aerial dispersal behavior in the twospotted spider mite. Entomol Exp Appl 68:79–86. https://doi.org/10.1111/j.1570-7458.1993.tb01691.x Linke W (1953) Investigation of the biology and epidemiology of the common spider mite, Tetranychus althaeae v. Hanst. with particular consideration of the hop as the host. Hoefchen-Briefe Bayer Pflanz. Nachr. 6:181–232. Lombaert E, Boll R, Lapchin L (2006) Dispersal strategies of phytophagous insects at a local scale: Adaptive potential of aphids in an agricultural environment. BMC Evol Biol 6:1–13. https://doi.org/10.1186/1471-2148-6-75 Macke E, Magalhães S, Bach F, Olivieri I (2011a) Experimental evolution of reduced sex ratio adjustment under local mate competition. Science 334:1127–1129. https://doi.org/10.1126/science.1212177 Macke E, Magalhaes S, Khan HDT, Luciano A, Frantz A, Facon B, Olivieri I (2011b) Sex allocation in haplodiploids is mediated by egg size: Evidence in the spider mite Tetranychus urticae Koch. Proc R Soc B Biol Sci 278:1054–1063. https://doi.org/10.1098/rspb.2010.1706 Macke E, Magalhaes S, Khanh Do-Thi H, Frantz A, Facon B, Olivieri I (2012) Mating modifies female life history in a haplodiploid spider mite. Am Nat 179:147–162. https://doi.org/10.1086/665002 Maeno KO, Piou C, Ghaout S (2020) The desert locust, Schistocerca gregaria , plastically manipulates egg size by regulating both egg numbers and production rate according to population density. J Insect Physiol 122:104020. https://doi.org/10.1016/j.jinsphys.2020.104020 Martin JP, Beyerlein A, Dacks AM, Reisenman CE, Riffell JA, Lei H, Hildebrand JG (2011) The neurobiology of insect olfaction: Sensory processing in a comparative context. Prog Neurobiol 95:427–447. https://doi.org/10.1016/j.pneurobio.2011.09.007 McCauley SJ (2010) Body size and social dominance influence breeding dispersal in male Pachydiplax longipennis (Odonata). Ecol Entomol 35:377–385. https://doi.org/10.1111/j.1365-2311.2010.01191.x McEdward LR, Carson SF (1987) Variation in egg organic content and its relationship with egg size in the starfish Solaster stimpsoni . Mar Ecol Prog Ser 37:159–169. McMurtry J, Huffaker C, van de Vrie M (1970) Ecology of tetranychid mites and their natural enemies: A review: I. Tetranychid enemies: Their biological characters and the impact of spray practices. Hilgardia 40:331–390. https://doi.org/10.3733/hilg.v40n11p331 McPeek MA, Holt RD (1992) The evolution of dispersal in spatially and temporally varying environments. Am Nat 140:1010–1027. https://doi.org/10.1111/evo.12699 Meiners T (2015) Chemical ecology and evolution of plant-insect interactions: A multitrophic perspective. Curr Opin Insect Sci 8:22–28. https://doi.org/10.1016/j.cois.2015.02.003 Mellors WK, Propts SE (1983) Effects of fertilizer level, fertility balance, and soil moisture on the interaction of two-spotted spider mites (Acarina: Tetranychidae) with radish plants. Environ Entomol 12:1239–1239. https://doi.org/10.1093/ee/12.4.1239 Migeon A, Dorkeld F (2024) Spider mites web: A comprehensive database for the Tetranychidae. https://www1.montpellier.inra.fr/CBGP/spmweb (Accessed 30 October 2024). Mitchell R (1973) Growth and population dynamics of a spider mite ( Tetranychus urticae K., Acarina: Tetranychidae). Ecology 54:1349–1355. https://doi.org/10.2307/1934198 Morrongiello JR, Bond NR, Crook DA, Wong BBM (2012) Spatial variation in egg size and egg number reflects trade-offs and bet-hedging in a freshwater fish. J Anim Ecol 81:806–817. https://doi.org/10.1111/j.1365-2656.2012.01961.x Nasu S, Tokuda M (2021) Dispersal–reproduction trade-off in the leaf beetle Galerucella grisescens . Entomol Exp Appl 169:542–549. https://doi.org/10.1111/eea.13042 Osakabe M, Isobe H, Kasai A, Masuda R, Kubota S, Umeda M (2008) Aerodynamic advantages of upside down take-off for aerial dispersal in Tetranychus spider mites. Exp Appl Acarol 44:165–183. https://doi.org/10.1007/s10493-008-9141-2 Parker GA, Begon M (1986) Optimal egg size and clutch size: Effects of environment and maternal phenotype. Am Nat 128:573–592. https://doi.org/10.1086/284589 Parvinen K, Egas M (2004) Dispersal and the evolution of specialisation in a two-habitat type metapopulation. Theor Popul Biol 66:233–248. https://doi.org/10.1016/j.tpb.2004.06.002 Pepi AA, Broadley HJ, Elkinton JS (2016) Density-dependent effects of larval dispersal mediated by host plant quality on populations of an invasive insect. Oecologia 182:499–509. https://doi.org/10.1007/s00442-016-3689-z Phillips BL, Brown GP, Shine R (2010) Life-history evolution in range-shifting populations. Ecology 91:1617–1627. https://doi.org/10.1890/09-0910.1 Pick JL, Hutter P, Tschirren B (2016) In search of genetic constraints limiting the evolution of egg size: Direct and correlated responses to artificial selection on a prenatal maternal effector. Heredity 116:542–549. https://doi.org/10.1038/hdy.2016.16 Plazio E, Margol T, Nowicki P (2020) Intersexual differences in density-dependent dispersal and their evolutionary drivers. J Evol Biol 33:1495–1506. https://doi.org/10.1111/jeb.13688 Poethke HJ, Gros A, Hovestadt T (2011) The ability of individuals to assess population density influences the evolution of emigration propensity and dispersal distance. J Theor Biol 282:93–99. https://doi.org/10.1016/j.jtbi.2011.05.012 Poethke HJ, Hovestadt T (2002) Evolution of density- and patch-size-dependent dispersal rates. Proc R Soc B Biol Sci 269:637–645. https://doi.org/10.1098/rspb.2001.1936 Puzin C, Pétillon J, Bonte D (2018) Influence of individual density and habitat availability on long-distance dispersal in a salt-marsh spider. Ethol Ecol Evol 31:28–37. https://doi.org/10.1080/03949370.2018.1486888 Reisig DD, Godfrey LD, Marcum DB (2010) Plant quality and conspecific density effects on Anaphothrips obscurus (Thysanoptera: Thripidae) wing diphenism and population ecology. Environ Entomol 39:685–694. https://doi.org/10.1603/EN09332 Renault D, Laparie M, McCauley SJ, Bonte D (2018) Environmental adaptations, ecological filtering, and dispersal central to insect invasions. Annu Rev Entomol 63:345–368. https://doi.org/10.1146/annurev-ento-020117-043315 Rhainds M, Shipp L (2003) Dispersal of adult western flower thrips (Thysanoptera: Thripidae) on Chrysanthemum plants: Impact of feeding-induced senescence of inflorescences. Environ Entomol 32:1056–1065. https://doi.org/10.1603/0046-225X-32.5.1056 Rhainds M, Shipp L, Woodrow L, Anderson D (2005) Density, dispersal, and feeding impact of western flower thrips (Thysanoptera: Thripidae) on flowering chrysanthemum at different spatial scales. Ecol Entomol 30:96–104. https://doi.org/10.1111/j.0307-6946.2005.00663.x Rioja C, Zhurov V, Bruinsma K, Grbic M, Grbic V (2017) Plant-Herbivore interactions: A case of an extreme generalist, the two-spotted spider mite Tetranychus urticae . MPMI 30:935–945. https://doi.org/10.1094/MPMI-07-17-0168-CR Roeder C, Harmsen R, Mouldey S (1996) The effects of relatedness on progeny sex ratio in spider mites. J Evol Biol 9:143–151. https://doi.org/10.1046/j.1420-9101.1996.9020143.x Ronce O, Clobert J (2012) Dispersal syndromes. In: Clobert J, Baguette M, Benton TG, Bullock JM (eds) Dispersal Ecology and Evolution. Oxford University Press, pp. 119–138. Ruf D, Dorn S, Mazzi D (2011) Females leave home for sex: Natal dispersal in a parasitoid with complementary sex determination. Anim Behav 81:1083–1089. https://doi.org/10.1016/j.anbehav.2011.02.028 Rushing CS, Brandt Ryder T, Valente JJ, Scott Sillett T, Marra PP (2021) Empirical tests of habitat selection theory reveal that conspecific density and patch quality, but not habitat amount, drive long‐distance immigration in a wild bird. Ecol Lett 24:1167–1177. https://doi.org/10.1111/ele.13729 Sabelis MW, Bakker FM (1992) How predatory mites cope with the web of their tetranychid prey: A functional view on dorsal chaetotaxy in the Phytoseiidae. Exp Appl Acarol 16:203–225. https://doi.org/10.1007/BF01193804 Sandeson PD, Boiteau G, Le Blanc J PR (2002) Adult density and the rate of colorado potato beetle (Coleoptera: Chrysomelidae) flight take-off. Environ Entomol 31:533–537. https://doi.org/10.1603/0046-225X-31.3.533 Schausberger P, Yano S, Sato Y (2021) Cooperative behaviors in group-living spider mites. Front Ecol Evol 9:745036. https://doi.org/10.3389/fevo.2021.745036 Shih CT, Poe SL, Cromroy HL (1976) Biology, life table, and intrinsic rate of increase of Tetranychus urticae . Ann Entomol Soc Am 69:362–364. https://doi.org/10.1093/aesa/69.2.362 Sloggett JJ, Weisser WW (2002) Parasitoids induce production of the dispersal morph of the pea aphid, Acyrthosiphon pisum . Oikos 98:323–333. https://doi.org/10.1034/j.1600-0706.2002.980213.x Smith CC, Fretwell SD (1974) The optimal balance between size and number of offspring. Am Nat 108:499–506. https://www.jstor.org/stable/2459681 Smitley DR, Kennedy GG (1988) Aerial dispersal of the two-spotted spider mite ( Tetranychus urticae ) from field corn. Exp Appl Acarol 5:33–46. https://doi.org/10.1007/BF02053815 Stamps J (2001) Habitat selection by dispersers: integrating proximate and ultimate approaches. In: Clobert J, Danchin E, Dhondt AA, Nichols JD (eds) Dispersal. Oxford University Press, pp 110–122. https://doi.org/10.1093/oso/9780198506607.003.0018 Stearns SC (1989) Trade-offs in life-history evolution. Funct Ecol 3:259–268. https://doi.org/10.2307/2389364 Strong WB, Croft BA, Slone DH (1997) Spatial aggregation and refugia of the mites Tetranychus urticae and Neoseiulus fallacis (Acari: Tetranychidae, Phytoseiidae) on Hop. Environ Entomol 26:859–865. https://doi.org/10.1093/ee/26.4.859 Suski Z, Badowska T (1975) Effect of the host plant nutrition on the population of the two spotted spider mite, Tetranychus urticae Koch (Acarina, Tetranychidae). Ekologia Polska 23:185–209. Tien NS, Sabelis MW, Egas M (2011) Ambulatory dispersal in Tetranychus urticae : An artificial selection experiment on propensity to disperse yields no response. Exp Appl Acarol 53:349–360. https://doi.org/10.1007/s10493-010-9411-7 Tinsley Johnson E, Feder JA, Lu A, Bergman TJ, Beehner JC, Snyder-Mackler N (2021) The goldilocks effect: Female geladas in mid-sized groups have higher fitness. Proc R Soc B 288:20210820. https://doi.org/10.1098/rspb.2021.0820 Travis JMJ, Mustin K, Palmer SCF, Bartoń KA, Hovestadt T, Benton TG, Clobert J, Delgado MM, Dytham C, Van Dyck H, Bonte D (2012) Modelling dispersal: An eco-evolutionary framework incorporating emigration, movement, settlement behaviour and the multiple costs involved. Methods Ecol Evol 3:628–641. https://doi.org/10.1111/j.2041-210X.2012.00193.x Tuan SJ, Lin YH, Yang CM, Atlihan R, Saska P, Chi H (2016) Survival and reproductive strategies in two-spotted spider mites: demographic analysis of arrhenotokous parthenogenesis of Tetranychus urticae (Acari: Tetranychidae). J Econ Entomol 109:502–509. https://doi.org/10.1093/jee/tov386 Walzer A, Schausberger P (2015) Food stress causes sex-specific maternal effects in mites. J Exp Biol 218:2603–2609. https://doi.org/10.1242/jeb.123752 Watson TF (1964) Influence of host plant condition on population increase of Tetranychus telarius (Linnaeus) (Acarina: Tetranychidae). Hilgardia 35:273–322. https://doi.org/10.3733/hilg.v35n11p273 Weerawansha N, Wang Q, He XZ (2022a) A haplodiploid mite adjusts fecundity and sex ratio in response to density changes during the reproductive period. Exp Appl Acarol 88:277–288. https://doi.org/10.1007/s10493-022-00749-0 Weerawansha N, Wang Q, He XZ (2022b) Comparing the effects of social environments and life history traits on sex allocation in a haplodiploid spider mite. Syst Appl Acarol 27:2123–2130. https://doi.org/10.11158/saa.27.10.20 Weerawansha N, Wang Q, He XZ (2023a) Local mate competition model alone cannot predict the offspring sex ratio in large and dense populations of a haplodiploid arthropod. Curr Zool 69:219–221. https://doi.org/10.1093/cz/zoac022 Weerawansha N, Wang Q, He XZ (2023b) Reproductive plasticity in response to the changing cluster size during the breeding period: A case study in a spider mite. Exp Appl Acarol 91:237–250. https://doi.org/10.1007/s10493-023-00834-y Weerawansha N, Wang Q, He XZ (2024) Conspecific cues mediate habitat selection and reproductive performance in a haplodiploid spider mite. Curr Zool 70:795–802. https://doi.org/10.1093/cz/zoae013 Wermelinger B, Oertli JJ, Delucchi V (1985) Effect of host plant nitrogen fertilization on the biology of the two‐spotted spider mite, Tetranychus urticae . Entomol Exp Appl 38:23–28. https://doi.org/10.1111/j.1570-7458.1985.tb03493.x Wiklund C, Persson A (1983) Fecundity, and the relation of egg weight variation to offspring fitness in the speckled wood butterfly Pararge aegeria , or why don't butterfly females lay more eggs? Oikos 40:53–63. https://doi.org/10.2307/3544198 Williams JL, Hufbauer RA, Miller TEX (2019) How evolution modifies the variability of range expansion. Trends Ecol Evol 34:903–913. https://doi.org/10.1016/j.tree.2019.05.012 Wilson LJ (1994) Plant-quality effect on life-history parameters of the twospotted spider mite (Acari: Tetranychidae) on cotton. J Econ Entomol 87:1665–1673. https://doi.org/10.1093/jee/87.6.1665 Wong JWY, Meunier J, Koelliker M (2013) The evolution of parental care in insects: The roles of ecology, life history and the social environment. Ecol Entomol 38:123–137. https://doi.org/10.1111/een.12000 Xu F, Yang W, Li Y (2019) Enlarged egg size increases offspring fitness of a frog species on the Zhoushan Archipelago of China. Sci Rep 9:11653. https://doi.org/10.1038/s41598-019-48147-8 Yano S (2008) Collective and solitary behaviors of twospotted spider mite (Acari: Tetranychidae) are induced by trail following. Ann Entomol Soc Am 101:247–252. https://doi.org/10.1603/0013-8746(2008)101[247:CASBOT]2.0.CO;2 Yano S (2012) Cooperative web sharing against predators promotes group living in spider mites. Behav Ecol Sociobiol 66:845–853. https://doi.org/10.1007/s00265-012-1332-5 Young SSY, Wrensch DL, Kongchuensin M (1986) Control of sex ratio by female spider mites. Entomol Exp Appl 40:53–60. https://doi.org/10.1111/j.1570-7458.1986.tb02155.x Zhang Z-Q (2003) Mites of Greenhouses: Identification, Biology and Control. CABI Publishing, UK. Zhou P, He XZ, Chen C, Wang Q (2021) Resource relocations in relation to dispersal in Tetranychus ludeni Zacher. Syst Appl Acarol 26:2018–2026. https://doi.org/10.11158/saa.26.11.3 Zhou P, He XZ, Chen C, Wang Q (2024) Age and density of mated females affect dispersal strategies in spider mite Tetranychus ludeni Zacher. Insects 15:387. https://doi.org/10.3390/insects15060387 Cite Share Download PDF Status: Posted 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. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-6772534","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":472544624,"identity":"9546d327-a0f9-42c5-9d48-017db7bcdb4e","order_by":0,"name":"Jhaman Kundun","email":"","orcid":"","institution":"Massey University - Manawatu Campus: Massey University","correspondingAuthor":false,"prefix":"","firstName":"Jhaman","middleName":"","lastName":"Kundun","suffix":""},{"id":472544625,"identity":"0a734941-3690-4f1a-b644-abe131cfc6ab","order_by":1,"name":"Resona Simkhada","email":"","orcid":"","institution":"Massey University - Manawatu Campus: Massey University","correspondingAuthor":false,"prefix":"","firstName":"Resona","middleName":"","lastName":"Simkhada","suffix":""},{"id":472544626,"identity":"8621d355-fccf-4261-af6b-2b5b17a370fc","order_by":2,"name":"Svetla Sofkova-Bobcheva","email":"","orcid":"","institution":"Massey University - Manawatu Campus: Massey University","correspondingAuthor":false,"prefix":"","firstName":"Svetla","middleName":"","lastName":"Sofkova-Bobcheva","suffix":""},{"id":472544627,"identity":"deb57c98-2b26-4276-8eb4-ef2a54dd69ba","order_by":3,"name":"Xiong Zhao He","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA3klEQVRIiWNgGAWjYJACZgYGG9K1pJGu5TAJyg0OsD9gLqg4H83ff/zhYx4GO3kG6fYHeLVINjAkMM84czt3xo0cY2MehmTDBpkDCXi18DMwHGDmbbud23CDh02ah4E5gUEi4QBeLWwMjA3MvP/O5c4/f/z5bx6GeqCWxAYCtjAzMPM2HMjdcCDBjJmH4TBQSzJeHQySzWwMh3mOJeduBPpFco7BccM2CQIhbnC8HRhQNXa5884ff/jhTUW1PL9E+gP8eoCRcgDGZuIxAPmOFMD4gyTlo2AUjIJRMFIAALELPNGjCUAPAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0003-3956-4498","institution":"Massey University - Manawatu Campus: Massey University","correspondingAuthor":true,"prefix":"","firstName":"Xiong","middleName":"Zhao","lastName":"He","suffix":""}],"badges":[],"createdAt":"2025-05-29 04:14:11","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6772534/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6772534/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":84910233,"identity":"bd5747fd-f376-4974-bec4-4c48dec7b7e8","added_by":"auto","created_at":"2025-06-18 16:51:47","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":22012,"visible":true,"origin":"","legend":"\u003cp\u003eMean (± SE) probability of resident or dispersing females of \u003cem\u003eTetranychus ludeni\u003c/em\u003e from LQ habitat (\u003cstrong\u003ea\u003c/strong\u003e) and HQ habitat (\u003cstrong\u003eb\u003c/strong\u003e). LQ-Resident – resided in low-quality (LQ) habitat; LQ-HQ – dispersed from low- to high-quality habitat; LQ-LQ – dispersed from low- to low-quality habitat; HQ-Resident – resided in high-quality (HQ) habitat; HQ-HQ – dispersed from high- to high-quality habitat; and HQ-LQ – dispersed from high- to low-quality habitat. Brown columns, resided in or dispersed to LQ habitats; green columns, resided in or dispersed to HQ habitats. In each dispersal scenario, columns with different letters are significantly different (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-6772534/v1/00a74d4532f198f1367931d5.png"},{"id":84911066,"identity":"4a574940-d007-4c35-9bd9-1c76ccd6d0ff","added_by":"auto","created_at":"2025-06-18 16:59:47","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":56915,"visible":true,"origin":"","legend":"\u003cp\u003eMean (± SE) total number of eggs laid by all \u003cem\u003eTetranychus ludeni\u003c/em\u003e females residing in or dispersing from LQ habitat (\u003cstrong\u003ea\u003c/strong\u003e) and HQ habitat (\u003cstrong\u003eb\u003c/strong\u003e), and its relation to female density (FD) (\u003cstrong\u003ec\u003c/strong\u003e, \u003cstrong\u003ed\u003c/strong\u003e): LQ-Resident, Eggs = exp(0.1786 + 3.5238 FD); LQ-HQ, Eggs = exp(0.0288 + 4.7609 FD); LQ-LQ, Eggs = exp(0.0934 + 3.5570 FD); HQ-Resident, Eggs = exp(0.0369 + 4.8865 FD); HQ-HQ, Eggs = exp(0.1972 + 3.2791 FD); and HQ-LQ, Eggs = exp(0.4887 + 3.0616 FD). Brown columns or lines, resided in or dispersed to LQ habitats; green columns or lines, resided in or dispersed to HQ habitats. In Fig. 2a and 2b, columns with different letters indicate there is a significant difference in total number of eggs laid in different habitats (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05); in Fig. 2c and 2d, lines with different letters indicate there is a significant difference in slope between different habitats (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-6772534/v1/3f0efdf5fe2fc4e15d3feff5.png"},{"id":84910235,"identity":"2b812bd0-2515-484f-969a-cbf9a2ea90fd","added_by":"auto","created_at":"2025-06-18 16:51:47","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":56913,"visible":true,"origin":"","legend":"\u003cp\u003eMean (± SE) number of eggs laid by each \u003cem\u003eTetranychus ludeni\u003c/em\u003e female residing in or dispersing from LQ habitat (\u003cstrong\u003ea\u003c/strong\u003e) and HQ habitat (\u003cstrong\u003eb\u003c/strong\u003e), and its relation to female density (FD) (\u003cstrong\u003ec\u003c/strong\u003e, \u003cstrong\u003ed\u003c/strong\u003e): LQ-Resident, Eggs = exp(– 0.1215 + 3.6352 FD); LQ-HQ, Eggs = exp(– 0.0897 + 3.6415 FD); LQ-LQ, Eggs = exp(– 0.3470 + 3.9335 FD); HQ-Resident, Eggs = exp(– 0.0550 + 3.5130 FD); HQ-HQ, Eggs = exp(– 0.1118 + 3.6052 FD); and HQ-LQ, Eggs = exp(– 0.0183 + 3.5410 FD). Brown columns or lines, resided in or dispersed to LQ habitats; green columns or lines, resided in or dispersed to HQ habitats. In Fig. 3a and 3b, columns with different letters indicate there is a significant difference in number of eggs laid per female in different habitats (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05); in Fig. 3c and 3d, lines with different letters indicate there is a significant difference in slope between different habitats (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-6772534/v1/aba5790de206091b1663a5aa.png"},{"id":84910238,"identity":"27300b22-5043-4108-8a14-d1d48e2e4b8e","added_by":"auto","created_at":"2025-06-18 16:51:47","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":49822,"visible":true,"origin":"","legend":"\u003cp\u003eMean (± SE) size of eggs laid by each \u003cem\u003eTetranychus ludeni\u003c/em\u003e female residing in or dispersing from LQ habitat (\u003cstrong\u003ea\u003c/strong\u003e) and HQ habitat (\u003cstrong\u003eb\u003c/strong\u003e), and its relation to female density (FD) (\u003cstrong\u003ec\u003c/strong\u003e, \u003cstrong\u003ed\u003c/strong\u003e): LQ-Resident, Egg size = exp(– 0.0056 + 0.2431 FD); LQ-HQ, Egg size = exp(– 0.0075 + 0.2409 FD); LQ-LQ, Egg size = exp(0.0008 + 0.1515 FD); HQ-Resident, Egg size = exp(0.0148 + 0.0152 FD); HQ-HQ, Egg size = exp(– 0.0009 + 0.1599 FD); and HQ-LQ, Egg size = exp(0.0123 + 0.1510 FD). Brown columns or lines, resided in or dispersed to LQ habitats; green columns or lines, resided in or dispersed to HQ habitats. In Fig. 4a and 4b, columns with different letters indicate there is a significant difference in egg size in different habitats (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05); in Fig. 4c and 4d, lines with different letters indicate there is a significant difference in slope between different habitats (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-6772534/v1/90b542f1ea926521bfed61b0.png"},{"id":84911067,"identity":"1943c4d8-b4fb-43ff-a8b2-b9aa12d0e886","added_by":"auto","created_at":"2025-06-18 16:59:47","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":31859,"visible":true,"origin":"","legend":"\u003cp\u003eMean (± SE) egg hatch rate (\u003cstrong\u003ea\u003c/strong\u003e) and immature (larvae and nymphs) survival rate (\u003cstrong\u003eb\u003c/strong\u003e) for all resident/dispersing females of \u003cem\u003eTetranychus ludeni\u003c/em\u003e in different habitats: Resident – resided in a primary habitat; LQ-HQ – dispersed from a low- to a high-quality habitat; LQ-LQ – dispersed from a low- to a low-quality habitat; HQ-HQ – dispersed from a high- to a high-quality habitat; and HQ-LQ – dispersed from a high- to a low-quality habitat. Brown columns, resided in or dispersed to LQ habitats; green columns, resided in or dispersed to HQ habitats. In each dispersal scenario, columns with different letters are significantly different (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-6772534/v1/d2b91db94de1021a515fb14d.png"},{"id":84910244,"identity":"77f28eef-e859-4a10-b1ef-fd36dd2cc050","added_by":"auto","created_at":"2025-06-18 16:51:47","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":58126,"visible":true,"origin":"","legend":"\u003cp\u003eMean (± SE) proportion of daughters of \u003cem\u003eTetranychus ludeni\u003c/em\u003e females residing in or dispersing from LQ habitat (\u003cstrong\u003ea\u003c/strong\u003e) and HQ habitat (\u003cstrong\u003eb\u003c/strong\u003e), and its relation to female density (FD) (\u003cstrong\u003ec\u003c/strong\u003e, \u003cstrong\u003ed\u003c/strong\u003e): LQ-Resident, Daughters%= exp1.2892 – 0.0624 FD); LQ-HQ, Daughters%= exp(2.1151 – 0.1231 FD); LQ-LQ, Daughters% = exp(1.3915 – 0.0992 FD); HQ-Resident, Daughters%= exp(– 0.4011 + 0.1236 FD); HQ-HQ, Daughters%= exp(0.6555 + 0.1609 FD); and HQ-LQ, Daughters%= exp(01.4591 – 0.0128 FD). Brown columns or lines, resided in or dispersed to LQ habitats; green columns or lines, resided in or dispersed to HQ habitats. In Fig. 6a and 6b, columns with different letters indicate there is a significant difference in proportion of daughters in different habitats (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05); in Fig. 6c and 6d, lines with different letters indicate there is a significant difference in slope between different habitats (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-6772534/v1/515ed15bc45af0eb4e8251a1.png"},{"id":84910237,"identity":"fd6c26c0-1a95-4d6d-80f4-41ff306d05ef","added_by":"auto","created_at":"2025-06-18 16:51:47","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":88464,"visible":true,"origin":"","legend":"\u003cp\u003eRelationships between immature mortality rate and larva number (\u003cstrong\u003ea\u003c/strong\u003e), immature mortality rate and egg size (\u003cstrong\u003eb\u003c/strong\u003e), and proportion of daughters and egg size (\u003cstrong\u003ec\u003c/strong\u003e) in\u003cem\u003e Tetranychus ludeni\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"7.png","url":"https://assets-eu.researchsquare.com/files/rs-6772534/v1/5510fd9f6378b18010098dd2.png"},{"id":87887562,"identity":"a37ea1c8-24dc-42a8-8e39-181bc651dbca","added_by":"auto","created_at":"2025-07-30 05:38:37","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1143788,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6772534/v1/c69074c7-5def-427e-9494-083055e9522d.pdf"}],"financialInterests":"","formattedTitle":"Host plant quality mediates dispersal, oviposition, and sex allocation in a Tetranychus spider mite","fulltext":[{"header":"Introduction","content":"\u003cp\u003eIn herbivore-plant systems, host plants serve as the habitats for phytophagous animal development, growth, and reproduction (Jermy \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Meiners \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). As both an essential environmental factor and a primary food resource, host plants significantly influence the occurrence, distribution, and population density of herbivores (Awmack and Leather \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Knolhoff and Heckel \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). However, host plant quality often decline due to the increase of animal population density during the growing seasons (Li and Margolies \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Denno et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Sandeson et al. \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Rhainds and Shipp \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Rhainds et al. \u003cspan citationid=\"CR96\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Reisig et al. \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), which will reduce the carrying capacity, the theoretical limit of population size sustained by the given resources of a particular habitat (Cadet et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Chapman and Byron \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In such dynamic environments, phytophagous insects are subjected to strong selection pressure to maximize or maintain their fitness by avoiding unsuitable habitats and locating suitable ones (Bruce et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Bruce and Pickett \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Previous theoretical studies (McPeek and Holt \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Parvinen and Egas \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) suggest that dispersal is a key adaptation to the temporal variability of habitat quality.\u003c/p\u003e \u003cp\u003eBiological dispersal, referring to the movement of individuals from their birth site to their breeding site (natal dispersal) (Johnson and Horvitz \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Ruf et al. \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) or between breeding sites (breeding dispersal) (McCauley \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) or both (Dieckmann et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Clobert et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), is the primary mechanism enabling them to find suitable habitats that can support their survival and reproduction (den Boer \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Dieckmann et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Clobert et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Dispersal is driven by multiple selective forces, including the avoidance of competition with relatives, reduction of inbreeding risks, or spreading risk in spatially and temporally varying environments or demography (Duputi\u0026eacute; and Massol \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Dispersal involves three distinct stages, i.e., departure, transience, and settlement (Clobert et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Bonte and Dahirel \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and each stage is subjected to selection pressures that minimize the overall costs of dispersal and maximize the fitness of dispersers and their offspring (Bonte et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Travis et al. \u003cspan citationid=\"CR115\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Among these stages, settlement associated with habitat selection is critical for the survival and reproduction success and thus the establishment and subsequent expansion of populations in new habitats (Kot et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Phillips et al. \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Ronce and Clobert \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Renault et al. \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Williams et al. \u003cspan citationid=\"CR126\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePhytophagous animals, through long-term adaptation and co-evolution, have developed sophisticated sensory and nervous systems (Martin et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Bruce \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) to detect and evaluate the quality of host plants (Bernays \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Bruce \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Burgue\u0026ntilde;o et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Karlsson Green et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). This ability to assess host suitability is crucial for future survival and reproductive success, as phytophagous insects usually have limited opportunities to make settlement decisions (Burgue\u0026ntilde;o et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Karlsson Green et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Previous studies on phytophagous animal dispersal mostly focus on the determinations of mechanisms or factors affecting dispersal (e.g., Li and Margolies \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Sandeson et al. \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Poethke and Hovestadt \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Sloggett and Weisser \u003cspan citationid=\"CR106\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Lombaert et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Yano \u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e2008\u003c/span\u003e, Bowler and Benton \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; McCauley \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Poethke et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Bitume et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2011\u003c/span\u003e, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Pepi et al. \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Puzin et al. \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Plazio et al. \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Hewison et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), few research has examined the subsequent reproduction of dispersers in the new habitats (e.g., Khuhro et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Nasu and Tokuda \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zhou et al. \u003cspan citationid=\"CR134\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR135\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). To date, how dispersers adjust their behaviours and reproductive strategies to adapt to the new habitats is largely unknown.\u003c/p\u003e \u003cp\u003eSpider mites (Acari: Tetranychidae) consisting of over 1300 species (Migeon and Dorkeld \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), and some species, such as \u003cem\u003eTetranychus urticae\u003c/em\u003e Koch, \u003cem\u003eT. evansi\u003c/em\u003e Baker and Pritchard, \u003cem\u003eT. kanzawai\u003c/em\u003e Kishida, and \u003cem\u003eT. ludeni\u003c/em\u003e Zacher, are serious pests damaging many economic crops worldwide (Bolland et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Zhang \u003cspan citationid=\"CR133\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). These mites feed by piercing the leaf tissues and sucking out plant contents with their sap-sucking mouthparts, leading to the reduced photosynthetic activities and visible damage such as yellow and white spots on leaves (Dhooria \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Spider mites often live in groups (Strong et al. \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Dhooria \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), which benefit individuals from modifying plant biochemistry, such as breaking down the plant defense system, resulting in more favourable nutritional quality of the shared host plants (Kant et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Rioja et al. \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). However, \u003cem\u003eTetranychus\u003c/em\u003e species may build up the local populations quickly due to their short lifecycle and high fecundity, for example \u003cem\u003eT. ludeni\u003c/em\u003e could complete development from egg to adult stage in about 10 days at 27 \u0026ordm;C and lay up to 110 eggs on suitable hosts (Adango et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). These life-history traits may result in scramble competition for food resources between individuals (Krips et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). When the population becomes large and dense, spider mites, especially the young, mated females, tend to disperse to search for new habitats due to food deficiency and habitat deterioration (Azand\u0026eacute;m\u0026egrave;-Hounmalon et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Schausberger et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zhou et al. \u003cspan citationid=\"CR134\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDispersal in spider mites mainly occurs in young females after mating (Mitchell \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e1973\u003c/span\u003e; Li and Margolies \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1993\u003c/span\u003e) and primarily through active movement, either individually and collectively by walking (Kondo and Takafuji \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1985\u003c/span\u003e; Yano \u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), although spider mites may disperse solitarily by walking (Yano \u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), although passive dispersal via wind is also observed (Brandenburg and Kennedy \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Jung \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Yano \u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Clotuche et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2011\u003c/span\u003e, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Collective dispersal of spider mite females may benefit group members because it enables the establishment of new colonies, and the immediate cooperative construction of webs which would effectively protect them from predators (McMurtry et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1970\u003c/span\u003e; Sabelis and Bakker \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Schausberger et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and from wind and rain (Davis \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1952\u003c/span\u003e; Linke \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e1953\u003c/span\u003e; Schausberger et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Moreover, habitat heterogeneity in quality is common in nature. Unlike the passively dispersing organisms who have no opportunity to choose where they settle, active dispersal allows them to select among breeding habitats that differ in key characteristics, such as size, quality, and conspecific density (Stamps, \u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Both dispersal behaviours and subsequent reproduction of spider mites, for example the \u003cem\u003eT. ludeni\u003c/em\u003e, have been partially assessed (Zhou et al. \u003cspan citationid=\"CR134\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR135\" class=\"CitationRef\"\u003e2024\u003c/span\u003e); however, those authors detect the subsequent reproduction of dispersing females by maintaining them individually on a leaf disc, how collective dispersers select habitats of various quality and whether they adjust the reproductive strategies in response to the environmental conditions of post-dispersal habitats (e.g., habitat quality and population density) need further investigations.\u003c/p\u003e \u003cp\u003eIn this study, we investigated the collective dispersal, habitat selection, and subsequent reproduction of the European native spider mite \u003cem\u003eT. ludeni\u003c/em\u003e. We simulated scenarios in nature by allowing the mated females to disperse from the low- or high-quality habitats, and choose between the low- and high-quality habitats. We then recorded the reproductive performances of collective dispersing females in terms of number and size of eggs, immature survival, and offspring sex ratio. We tested the hypotheses that (1) females are more likely to disperse from the low-quality habitats compared to the high-quality ones; (2) females would select and settle in high-quality habitats rather than the low-quality ones, and (3) dispersing females would lay more and large eggs, and generate more female offspring in the high-quality habitats than in the low-quality ones. This study will deliver knowledge to the strategies of spider mite dispersal and reproduction in response to social environment changes.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eMite colony and experimental conditions\u003c/h2\u003e \u003cp\u003eThe colony of spider mite \u003cem\u003eT. ludeni\u003c/em\u003e started from female adults collected from a garden in Palmerston North, New Zealand. Mites were reared on kidney bean plants (\u003cem\u003ePhaseolus vulgaris\u003c/em\u003e L.) grown in potting mix (19:9:10 NPK at 400 g/100 L) in plastic pots (8 cm bottom diameter \u0026times; 10 cm top diameter \u0026times; 9 cm height). The mite colony was maintained in nylon-framed cages (65 cm length \u0026times; 50 cm width \u0026times; 50 cm height, aperture size\u0026thinsp;=\u0026thinsp;0.075 mm length \u0026times; 0.075 mm width) in a bioassay room in the Entomology and IPM laboratory, Massey University, New Zealand, at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;1 \u0026ordm;C, 40\u0026thinsp;\u0026plusmn;\u0026thinsp;10% RH and 14:10 h (L:D) photoperiod. Three- to 5-week-old plants were used to maintain the colony, while first expanded plant leaves (1 to 2 weeks old) were used for experiments. Mites were reared for at least three successive generations prior to experiments. Experiments were carried out under the above-mentioned environmental conditions.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eDispersal and habitat selection\u003c/h3\u003e\n\u003cp\u003eTo prepare \u003cem\u003eT. ludeni\u003c/em\u003e female adults for experiments, five female adults randomly selected from the colony were introduced onto a bean leaf square (2 cm length \u0026times; 2 cm width) placed upside down on a water saturated cotton pad in a Petri dish (1.0 cm height \u0026times; 9.5 cm diameter) with a mesh-covered opening (1.0 cm diameter) on the lid for ventilation. Females were allowed to lay eggs for 24 h, after which they were removed to another leaf square for egg laying. Thirty eggs remained on the leaf square were allowed to develop to deutonymphal stage. Although some males would emerge from the 30 eggs, we introduced 10 male adults developing from eggs laid by virgin females onto the leaf squares before deutonymphs emerged to adults, which ensured females mated at emergence. The 1-day-old mated females were used for experiments.\u003c/p\u003e \u003cp\u003eTo prepare low-quality (LQ) habitats for experiments, 10 female adults were randomly selected from the colony and introduced onto a bean leaf square (1 cm length \u0026times; 1 cm width) in a Petri dish as did above. Mites were allowed to feed on the leaf square for five days. After which time, all mites, eggs, silks/webs, and faeces were removed using a fine brush, and the leaf square was washed by tap water to eliminate or at least reduce the traces left by the previous mites. The leaf squares were air-dried under laboratory conditions. Fresh plant leaf squares without mites infested were treated as high-quality (HQ) habitats.\u003c/p\u003e \u003cp\u003eTo investigate the dispersal and habitat selection behaviors of mites, two treatments were set up: (1) females dispersed from LQ habitat to select between the LQ and HQ habitats, and (2) females dispersed from HQ habitat to select between the LQ and HQ habitats. There were 15 replicates for each treatment. For each replicate, 15 1-day-old mated females prepared above were introduced onto a LQ or HQ habitat placed on the centre of a cotton pad in a Petri dish and bridged to one LQ and one HQ habitat by two parafilm strips (2 cm length \u0026times; 0.5 cm width). The number of individuals residing on or dispersing to a habitat was counted after 24 hours.\u003c/p\u003e\n\u003ch3\u003eOviposition and sex allocation of resident and dispersing mites (dup: abstract ?)\u003c/h3\u003e\n\u003cp\u003eTo test the consequent effects of habitat selection on reproductive performance of female mites, the three leaf squares along with the cotton pad in a Petri dish were separated and individually transferred onto a new Petri dish. The resident and dispersing females were allowed to lay eggs \u003cem\u003ein situ\u003c/em\u003e for five days, for two reasons, (1) maintaining the consistent social environments (i.e., habitat, and population size and density) for the resident or dispersed individuals, and (2) mortality occurring on sixth day of adult life. As some eggs might hatch within four days, the total number of eggs per habitat was counted as the sum of newly laid unhatched eggs and emerged larvae/nymphs, and the number of eggs per female was also calculated as total number of eggs /number of females. As egg size does not change before hatching (three days after oviposition under the experimental conditions) (Weerawansha et al. \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), all or a maximum of 20 eggs were randomly selected from a habitat on the 3rd day of oviposition and the egg diameter as measured under a stereomicroscope (Leica MZ12, Germany) connected to a digital camera (Olympus SC30, Japan) and an imaging software (CellSens\u0026reg; GS-ST-V1.7, Olympus, Japan). The spider mite eggs are sphere shape, and their size was then calculated as: volume\u0026thinsp;=\u0026thinsp;4/3π\u003cem\u003er\u003c/em\u003e\u003csup\u003e3\u003c/sup\u003e, where \u003cem\u003er\u003c/em\u003e is the radius (=\u0026thinsp;diameter/2). All resident and dispersing females were removed from the habitats on the 6th day. The egg hatching rate and immature survival rate were also calculated. The newly emerged adults were sexed, and the proportion of daughters was calculated.\u003c/p\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eData were analysed using SAS 9.13 (SAS Institute Inc, USA), with a significant level of α\u0026thinsp;=\u0026thinsp;0.05. A generalized linear model (GLIMMIX Procedure) with a binomial distribution and a logit-link function in the model was used to compare the difference in dispersal behaviour, egg hatch and immature survival rate, and proportion of offspring daughters between the resident and selected habitats and to determine proportion of offspring daughters over the female density. Another generalized linear model (GLIMMIX Procedure) with a poisson or gamma distribution and a logit-link function in the model was applied to compare the difference in total number of eggs laid by all residing/dispersing females, number of eggs laid by each female, and egg size between the resided and selected habitats and to test their relationship to female density. A Tukey-Kramer test was applied to perform the multiple comparisons. A linear regression (GLM Procedure) as applied to assess the relationships between immature survival rate and larva number, between immature survival rate and egg size, and between proportion of daughters and egg size.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eDispersal and habitat selection\u003c/h2\u003e \u003cp\u003eFemale mites exhibited significantly higher dispersal probability when they initially settled in low-quality (LQ) habitats compared to that in high-quality (HQ) habitats (64.67\u0026thinsp;\u0026plusmn;\u0026thinsp;3.10% vs. 29.58\u0026thinsp;\u0026plusmn;\u0026thinsp;3.10%) (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,30\u003c/sub\u003e = 56.94, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001). When initially settled in LQ habitats, mites were significantly more likely to disperse to HQ habitats rather than to reside in LQ habitats or to disperse to LQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,45\u003c/sub\u003e = 39.52, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea). When initially settled in HQ habitats, mites were significantly more likely to reside in HQ habitats rather than to disperse to HQ or LQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,45\u003c/sub\u003e = 92.65, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eOviposition and sex allocation of resident and dispersing mites\u003c/h3\u003e\n\u003cp\u003eWhen dispersed from LQ habitats, the total number of eggs laid by females dispersing to HQ habitats was significantly higher than that laid by females residing in the LQ habitats, with a significant lower number of eggs laid by females dispersing to LQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 310.68, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea); when dispersed from HQ habitats, the total number of eggs laid by the resident females was significantly higher than those laid by females dispersing to the HQ habitats, with a significant lower number of eggs laid by females dispersing to LQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 622.87, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb). The egg number increased significantly with the increasing number of mites on a habitat regardless of the dispersing scenario except females dispersing from LQ to HQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,13\u003c/sub\u003e = 143.45, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001 for LQ-Resident; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,14\u003c/sub\u003e = 2.73, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.12 for LQ-HQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,9\u003c/sub\u003e = 10.13, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0111 for LQ-LQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,14\u003c/sub\u003e = 14.65, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0018 for HQ-Resident; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,13\u003c/sub\u003e = 51.26, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001 for HQ-HQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,9\u003c/sub\u003e = 200.97, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001 for HQ-LQ) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec, \u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed); however, for the scenario of dispersing from LQ habitats the rate of increase (i.e., slope) was significantly greater for females residing in the LQ habitats than those dispersing to HQ or LQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,36\u003c/sub\u003e = 21.51, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec), while for the scenario of dispersing from HQ habitats the rate of increase was significantly greater for mites dispersing to the LQ habitats than those dispersing to HQ habitats, with a significant slower increasing rate detected for mites dispersing to HQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,36\u003c/sub\u003e = 89.29, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eWhen females resided in or dispersed from LQ habitats to HQ or LQ habitats, they laid similar number of eggs (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 3.10, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.06) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea); while the number of eggs laid by each female was significantly higher when females dispersed from HQ habitats to LQ habitats than that laid by that dispersing from HQ habitats to HQ habitats with a significant fewer eggs laid by the resident mites (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 27.60, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb). The egg number by each female decreased with the increasing number of mites, but significance was found for females that resided in the LQ habitats, dispersed from LQ habitats to LQ habitats, or from HQ habitats to HQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,13\u003c/sub\u003e = 12.59, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0036 for LQ-Resident; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,14\u003c/sub\u003e = 3.76, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.07 for LQ-HQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,9\u003c/sub\u003e = 49.52, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001 for LQ-LQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,14\u003c/sub\u003e = 2.71, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.12 for HQ-Resident; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,13\u003c/sub\u003e = 4.86, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.046 for HQ-HQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,9\u003c/sub\u003e = 0.09, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.77 for HQ-LQ) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec, \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed). For the scenario of dispersing from LQ habitats, the decrease of eggs laid (i.e., slope) was significantly faster for females dispersing from LQ habitats to LQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,36\u003c/sub\u003e = 8.92, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0007) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec), while for the scenario of dispersing from HQ habitats, the rate of decrease was not significantly different between the residing and dispersing females (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,36\u003c/sub\u003e = 0.78, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.47) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFemales residing in LQ habitats produced significantly larger eggs than the dispersing females, with a significantly smaller egg size detected for females that dispersed from LQ to LQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 4.39, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0191) (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea); however, when females initially settled in HQ habitats, there was no significant difference in egg size between the resident and dispersing females (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 0.64, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.54) (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb). Female density in a habitat had no significant effect on egg size, except when females resided in HQ habitats the egg size significantly increased with female number (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,13\u003c/sub\u003e = 0.56, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.47 for LQ-Resident; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,14\u003c/sub\u003e = 0.64, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.35 for LQ-HQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,9\u003c/sub\u003e = 0.01, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.93 for LQ-LQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,14\u003c/sub\u003e = 14.00, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0022 for HQ-Resident; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,13\u003c/sub\u003e = 0.01, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.91 for HQ-HQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,9\u003c/sub\u003e = 0.65, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.44 for HQ-LQ) (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec, \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ed). There was no significant difference in regression slop between habitats in each dispersing scenario (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,36\u003c/sub\u003e = 0.22, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.81, Fig.\u0026nbsp;4c; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,36\u003c/sub\u003e = 0.78, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.47; Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ed).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThere was no significant difference in egg hatch rate between the resident and dispersing females in either dispersal scenario (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 1.71, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.32 for dispersing from LQ habitat; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 1.60, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.21 for dispersing from HQ habitat) (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea). However, when mites dispersed from LQ habitats, immature survival was significantly lower for those dispersing to HQ habitats compared to those residing in the LQ habitats, with a significant higher immature survival for those dispersing to LQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 16.51, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001). In contrast, when mites dispersed from HQ habitats, immature survival was significantly lower for those residing in HQ habitats compared to those dispersing to the HQ or LQ habitats (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 22.91, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eb).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eWhen mites dispersed from LQ habitats, there was no significant difference in offspring sex ratio between the resident and dispersing females (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,39\u003c/sub\u003e = 1.69, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.20) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea); while when mites dispersed from HQ habitats, the proportion of daughters was significantly higher for females dispersing to HQ and LQ habitats than for resident females (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,38\u003c/sub\u003e = 9.85, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0004) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eb). Female density in a habitat had no significant effect on sex allocation, except when females resided in HQ habitats the proportion of daughters significantly increased with female number (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,13\u003c/sub\u003e = 2.22, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.16 for LQ-Resident; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,14\u003c/sub\u003e = 3.81, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.07 for LQ-HQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,9\u003c/sub\u003e = 1.12, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.32 for LQ-LQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,14\u003c/sub\u003e = 15.39, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0015 for HQ-Resident; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,13\u003c/sub\u003e = 3.45, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.09 for HQ-HQ; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,8\u003c/sub\u003e = 0.01, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.92 for HQ-LQ) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ec, \u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ed). There was no significant difference in regression slop between habitats in each dispersing scenario (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,36\u003c/sub\u003e = 0.34, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.72, Fig.\u0026nbsp;6c; \u003cem\u003eF\u003c/em\u003e\u003csub\u003e2,36\u003c/sub\u003e = 0.73, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.49; Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ed).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eAs shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea, immature survival significantly decreased with increasing population density of mite larvae (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,82\u003c/sub\u003e = 11.20, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0012). Egg size did not have significant impact on immature survival (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,82\u003c/sub\u003e = 0.01, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.93) (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eb) and proportion of daughters (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1,81\u003c/sub\u003e = 0.21, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.15) (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ec).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eDispersal and habitat selection\u003c/h2\u003e \u003cp\u003ePhytophagous organisms are subject to various selective pressures in nature due to the spatial and temporal variability of habitat quality. Spider mites can expand their local populations rapidly due to their specific life history traits, e.g., group living (Strong et al. \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Le Goff et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Clotuche et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Yano \u003cspan citationid=\"CR131\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), short lifecycle (Shih et al. \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e1976\u003c/span\u003e; Adango et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Bounfour and Tanigoshi \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Gotoh et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Tuan et al. \u003cspan citationid=\"CR116\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), high fecundity (Adango et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Bounfour and Tanigoshi \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Gotoh et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Tuan et al. \u003cspan citationid=\"CR116\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), and extremely high female-biased offspring sex ratio (Carey and Bradley \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Macke et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2011a\u003c/span\u003e; Weerawansha et al. \u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e2023a\u003c/span\u003e). However, these life-history traits may result in scramble competition between individuals, inducing an overexploitation of and quick cumulative excrement on host plants, and a fast depletion of resource and decrease of habitat quality (Hussey and Parr \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1963\u003c/span\u003e; Krips et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Weerawansha 2023b). Our results demonstrate that \u003cem\u003eT. ludeni\u003c/em\u003e females were more likely to disperse from low-quality (LQ) habitats (64.7%) than from the high-quality (HQ) habitats (29.6%), indicating that declining resource quality triggers dispersal in spider mites (Jeppson et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1975\u003c/span\u003e; Boykin and Campbell \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Azand\u0026eacute;m\u0026egrave;-Hounmalon et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Dispersing from deteriorating environments to more favourable habitats is an adaptive strategy for spider mites to maximize their own and offspring fitness (Azand\u0026eacute;m\u0026egrave;-Hounmalon et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Schausberger et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zhou et al. \u003cspan citationid=\"CR134\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eHabitat selection is a key determination of dispersal success. Spider mites usually prefer young leaves with a high level of nutrients (e.g., nitrogen) for feeding and oviposition (Watson \u003cspan citationid=\"CR118\" class=\"CitationRef\"\u003e1964\u003c/span\u003e; Suski and Badowska \u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e1975\u003c/span\u003e; Mellors and Propts \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e1983\u003c/span\u003e; Wermelinger et al. \u003cspan citationid=\"CR124\" class=\"CitationRef\"\u003e1985\u003c/span\u003e; Wilson \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003e1994\u003c/span\u003e). As expected, dispersing \u003cem\u003eT. ludeni\u003c/em\u003e females significantly preferred and settled in the HQ habitats compared to the LQ ones (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The results may have two implications. First, spider mites are able to assess plant quality and alter their behavior accordingly (Wilson \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003e1994\u003c/span\u003e), and the HQ habitats are expected to slow density-dependent declines in fitness (Avgar et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Second, dispersal in spider mites is a non-random process. In many web-spinning arthropods, such as the social spiders, social caterpillars, and spider mites, silk is a vector for collective behaviour (Fern\u0026aacute;ndez Ferrari et al. 2013). Yano (\u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) shows evidence that in \u003cem\u003eT. urticae\u003c/em\u003e, silk is strongly attractive, and dispersing females could distinguish between silk trails laid by the preceding solitary or grouping females, which provides positive feedback to the dispersers and results in collective dispersal by following the path with more silk on it.\u003c/p\u003e \u003cp\u003eOur results further reveal that even though dispersing mites had opportunities to choose between the HQ and LQ habitats, some individuals still settled in LQ habitats regardless of the dispersal scenarios (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Bruce (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) argues that dispersers may make \u0026lsquo;mistakes\u0026rsquo; and settle in poor quality hosts, although phytophagous species are under selection pressure to find quality hosts and have evolved a fine-tuned sensory system for the detection of host cues and avoidance of unsuitable hosts (Martin et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Bruce \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Rushing et al. (\u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) further hypothesize that dispersers who can reliably assess habitats and are free to settle should select the highest quality habitats until density-dependent mechanisms reduce expected reproductive success to the point, after which individuals would have higher fitness by settling in lower quality patches. The hypothesis may explain the probability of some \u003cem\u003eT. ludeni\u003c/em\u003e females selecting the LQ habitats, especially when significantly more individuals dispersed from the LQ habitats (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eOviposition and sex allocation of resident and dispersing mites\u003c/h2\u003e \u003cp\u003eSpider mites live in groups (Strong et al. \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Dhooria \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), which may benefit individuals from modifying plant biochemistry by breaking down the plant defense system and resulting in more favourable nutritional quality of the shared host plants (Kant et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Rioja et al. \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and reducing the intensity of individual web production (thinner, shorter, and/or fewer threads) and thus saving energy and nutrients (protein, and amino acids) in web production that can be invested in reproduction (Hazan et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1974\u003c/span\u003e; Le Goff et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). However, our results show that individual females remained in HQ habitats significantly reduced the number of eggs laid, and when \u003cem\u003eT. ludeni\u003c/em\u003e females dispersed from LQ habitats to the HQ ones, they did not significantly increase reproduction (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea, \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb). These may be attributed to two reasons. First, the high population density (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) and subsequent high egg density (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea, \u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb) in the LQ-HQ and resident-HQ habitats might have approached the carrying capacity; therefore, individual females restrained reproductive output at crowded environments, which may be an adaptive strategy to reduce future food competition and ensure offspring survival (Weerawansha et al. \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Second, residing in a large group will induce costs on foraging and feeding efficiency due to the higher interference among group members (Bilde et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Estevez et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Grove \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Wong et al. \u003cspan citationid=\"CR128\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Li and Zhang \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Tinsley Johnson et al. \u003cspan citationid=\"CR114\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), which may restrain of female fecundity (Krips et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Clotuche \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Bitume et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Furthermore, individual females also restrained reproduction in response to the increasing population density (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec, \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed), especially in LQ-LQ habitats females had a significant faster decrease of reproduction with increase population density (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec) despite a lower population density (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea), indicating that the population density-dependent reproductive constraint in \u003cem\u003eT. ludeni\u003c/em\u003e is regulated by habitat quality.\u003c/p\u003e \u003cp\u003eA trade-off between egg number and egg size is frequently reported (Smith and Fretwell \u003cspan citationid=\"CR107\" class=\"CitationRef\"\u003e1974\u003c/span\u003e; Parker and Begon \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Stearns \u003cspan citationid=\"CR110\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Fox and Czesak \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Macke et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Walzer and Schausberger \u003cspan citationid=\"CR117\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Maenoa et al. 2020), and larger egg size may facilitate survival of immature stages (Goulden et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Fox \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Johnston and Leggett \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Pick et al. \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Xu et al. \u003cspan citationid=\"CR129\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Unfortunately, in this study, reproductive constraint in LQ-HQ habitats (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea) or oviposition reduction in HQ-Resident habitats (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb) did not significantly increased egg size (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea, \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb), and increasing egg size did not significantly promoted egg hatch rate (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea), and immature survival rate (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea, \u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eb). Previous studies also reported that when females are shifted between habitats of high to low population density, a trade-off between egg number and size, and egg size had little impact on reproductive fitness in terms of immature survival (Weerawansha et al. \u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e, \u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003eb\u003c/span\u003e). Therefore, it may be concluded that when future environmental conditions are uncertain or unpredictable, egg size may not be a reliable indicator of offspring fitness (Wiklund and Persson \u003cspan citationid=\"CR125\" class=\"CitationRef\"\u003e1983\u003c/span\u003e; Karlsson and Wiklund \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e1985\u003c/span\u003e; McEdward and Carson \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Lalonde \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Morrongiello et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Weerawansha et al. \u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e, \u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003eb\u003c/span\u003e). While our results reveal that increasing population of active immature individuals significantly decreased immature survival rate (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea), because the nature of scramble resource competition between individual spider mites in habitats with high population density will result in a fast depletion of local resource and decrease of habitat quality (Hussey and Parr \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1963\u003c/span\u003e; Krips et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Weerawansha 2023a), which may induce the high mortality before they develop to adults due to the deficient nutrients. Therefore, in this study, the carrying capacity of habitats is the major factor affecting the reproductive performances of \u003cem\u003eT. ludeni\u003c/em\u003e females and fitness of offspring.\u003c/p\u003e \u003cp\u003eIn spider mites, the mated females are capable of manipulating offspring sex ratio by selectively fertilizing (Young et al. \u003cspan citationid=\"CR132\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Roeder et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Macke et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2011a\u003c/span\u003e; Weerawansha et al. \u003cspan citationid=\"CR120\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e) when the egg size exceeds a threshold value (Macke et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e). However, results of this study show that increasing egg size did not significantly facilitate the production of daughters (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ec). Alternatively, in the scenario of mites dispersing from LQ habitats, dispersing females laid significantly smaller eggs (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea) but produced similar proportion of daughters regardless of the quality of selected habitats (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea); while in the scenario of mites dispersing from HQ habitats, dispersing females laid eggs of similar size (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb) but produced significantly higher proportion of daughters regardless the quality of selected habitats (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eb). These results suggest that the dispersing females could adjust the fertilization threshold to a lower level and fertilise relatively smaller eggs. Weerawansha et al. (\u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e, \u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003eb\u003c/span\u003e) also demonstrated that when \u003cem\u003eT. ludeni\u003c/em\u003e females aggregate into a large group, they lay significantly smaller eggs but produced a significantly higher female-biased sex ratio, which is attributed to the flexibility of egg fertilization. Furthermore, our results show that compared to the habitat quality, female density had less impact on egg size (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea, \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb) and daughter production (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea, \u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eb); therefore, the impact of habitat quality was superior to that of population density on the adjustment of fertilization threshold that determines the offspring sex ratio in \u003cem\u003eT. ludeni\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eMany studies have demonstrated that dispersal is dependent on both disperser phenotype and the local environment (Bowler and Benton \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Clobert et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Bonte et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Hollander et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Baines et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In spider mites, when the local resources are depleted, the reproductive females will disperse to seek new habitats for the next generation either by ambulatory dispersal within a habitat (Tien et al. \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Azand\u0026eacute;m\u0026egrave;-Hounmalon et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Li and Zhang \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Schausberger et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zhou et al. \u003cspan citationid=\"CR134\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR135\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) or aerial dispersal for a long distance (Boykin and Campbell \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Smitley and Kennedy \u003cspan citationid=\"CR108\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Li and Margolies \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Osakabe et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Clotuche et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). We reveal that relatively higher proportion of \u003cem\u003eT. ludeni\u003c/em\u003e females (35%) still resided in the low-quality habitats. Whether spider mites have evolved phenotypic plasticity for dispersal, and whether the philopatric and dispersing phenotypes have developed different strategies in oviposition and sex allocation remain further investigations.\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003e \u003cem\u003eTetranychus ludeni\u003c/em\u003e females are more likely to disperse from the low-quality (LQ) habitats than from the high-quality (HQ) ones, and the dispersing females significantly prefer and settle in HQ habitats regardless the dispersal scenarios. The significantly high population density and total number of eggs laid in HQ habitats result in restrained oviposition of females and higher mortality of immature offspring. Egg size has no significant effect on egg hatching and immature survival. Females that disperse from LQ habitats to the HQ or LQ habitats produce significantly smaller eggs but similar proportion of daughters, and females that disperse from HQ habitats to the HQ or LQ habitats produce eggs of similar size but significantly higher proportion of daughters, suggesting that dispersing females may manipulate offspring sex ratio by lowing the fertilization threshold to fertilise relatively smaller eggs. However, population density has less impact on egg size and offspring sex ratio. Our results indicate that the dispersal and habitat selection of \u003cem\u003eT. ludeni\u003c/em\u003e females are mediated by host plant quality, and the dispersing females could adjust their reproductive strategies to maximise their own and offspring fitness.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank Professor Z.-Q. Zhang for identification of this spider mite to species, and Plant Growth Unit (PGU), Massey University for providing potting mix for bean plant growth.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was jointly funded by the Manaaki New Zealand Scholarships.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthical approval\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets used and/or analysed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eJK and XZH conceived and designed the experiments. JK and RS collected the data. JK and XZH analysed the data. JK prepared the first draft. All authors contributed to manuscript revision.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAdango E, Onzo A, Hanna R, James B, Atachi P (2006) Comparative demography of the spider mite, \u003cem\u003eTetranychus ludeni\u003c/em\u003e, on two host plants in West Africa. J Inse Sci 6:1\u0026ndash;9. https://doi.org/10.1673/031.006.4901\u003c/li\u003e\n\u003cli\u003eAvgar T, Betini GS, Fryxell JM (2020) Habitat selection patterns are density dependent under the ideal free distribution. J Anim Ecol 89:2777\u0026ndash;2787. https://doi.org/10.1111/1365-2656.13352 \u003c/li\u003e\n\u003cli\u003eAwmack CS, Leather SR (2002) Host plant quality and fecundity in herbivorous insects. Annu Rev Entomol 47:817\u0026ndash;844. https://doi.org/10.1146/annurev.ento.47.091201.145300 \u003c/li\u003e\n\u003cli\u003eAzand\u0026eacute;m\u0026egrave;-Hounmalon GY, Fellous S, Kreiter S, Fiaboe KK, Subramanian S, Kungu M, Martin T (2014) Dispersal behavior of \u003cem\u003eTetranychus evansi \u003c/em\u003eand\u003cem\u003e T. urticae\u003c/em\u003e on tomato at several spatial scales and densities: implications for integrated pest management. PLoS ONE 9:e95071. https://doi.org/10.1371/journal.pone.0095071\u003c/li\u003e\n\u003cli\u003eBaines CB, Ferzoco IMC, McCauley SJ (2019) Phenotype‐by‐environment interactions influence dispersal. J Anim Ecol 88:1263\u0026ndash;1274. https://doi.org/10.1111/1365-2656.13008\u003c/li\u003e\n\u003cli\u003eBernays EA (2001) Neural limitations in phytophagous insects: implications for diet breadth and evolution of host affiliation. Annu Rev Entomol 46:703\u0026ndash;727. https://doi.org/10.1146/annurev.ento.46.1.703 \u003c/li\u003e\n\u003cli\u003eBilde T, Coates KS, Birkhofer K, Bird T, Maklakov AA, Lubin Y, Avil\u0026eacute;s L (2007) Survival benefits select for group living in a social spider despite reproductive costs. J Evol Biol 20:2412\u0026ndash;2426. https://doi.org/10.1111/j.1420-9101.2007.01407.x\u003c/li\u003e\n\u003cli\u003eBitume EV, Bonte D, Magalh\u0026atilde;es S, Martin GS, Van Dongen S, Bach F, Anderson JM, Olivieri I, Nieberding CM (2011) Heritability and artificial selection on ambulatory dispersal distance in \u003cem\u003eTetranychus urticae\u003c/em\u003e: Effects of density and maternal effects. PLoS ONE 6:1\u0026ndash;9. https://doi.org/10.1371/journal.pone.0026927\u003c/li\u003e\n\u003cli\u003eBitume EV, Bonte D, Ronce O, Olivieri I, Nieberding CM (2014) Dispersal distance is influenced by parental and grand-parental density. Proc R Soc B 281:1\u0026ndash;8. https://doi.org/10.1098/rspb.2014.1061\u003c/li\u003e\n\u003cli\u003eBitume EV, Nieberding CM, Ronce O, Bach F, Flaven E, Olivieri I, Bonte D (2013) Density and genetic relatedness increase dispersal distance in a subsocial organism. Ecol Lett 16:430\u0026ndash;437. https://doi.org/10.1111/ele.12057\u003c/li\u003e\n\u003cli\u003eBolland HR, Gutierrez J, Flechtmann CH (1998) World catalogue of the spider mite family (Acari: Tetranychidae). Brill Academic Publishers, Leiden, Boston.\u003c/li\u003e\n\u003cli\u003eBonte D, Dahirel M (2017) Dispersal: A central and independent trait in life history. Oikos 126:472\u0026ndash;479. https://doi.org/10.1111/oik.03801\u003c/li\u003e\n\u003cli\u003eBonte D, De Roissart A, Wybouw N, Van Leeuwen T (2014) Fitness maximization by dispersal: Evidence from an invasion experiment. Ecology 95:3104\u0026ndash;3111. https://doi.org/10.1890/13-2269.1\u003c/li\u003e\n\u003cli\u003eBonte D, Van Dyck H, Bullock JM et al. (2012) Costs of dispersal. Biol Rev 87:290\u0026ndash;312. https://doi.org/10.1111/j.1469-185X.2011.00201.x\u003c/li\u003e\n\u003cli\u003eBounfour M, Tanigoshi LK (2001) Effect of temperature on development and demographic parameters of \u003cem\u003eTetranychus urticae \u003c/em\u003eand\u003cem\u003e Eotetranychus carpini borealis\u003c/em\u003e (Acari: Tetranychidae). Ann Entomol Soc Am 94:400\u0026ndash;404. https://doi.org/10.1603/0013-8746(2001)094[0400:EOTODA]2.0.CO;2\u003c/li\u003e\n\u003cli\u003eBowler DE, Benton TG (2005) Causes and consequences of animal dispersal strategies: relating individual behaviour to spatial dynamics. Biol Rev 80:205\u0026ndash;225. https://doi.org/10.1017/S1464793104006645\u003c/li\u003e\n\u003cli\u003eBowler DE, Benton TG (2009) Variation in dispersal mortality and dispersal propensity among individuals: the effects of age, sex and resource availability. J Anim Ecol 78:1234\u0026ndash;1241. https://doi.org/10.1111/j.1365-2656.2009.01580.x\u003c/li\u003e\n\u003cli\u003eBoykin L, Campbell W (1984) Wind dispersal of the twospotted spider mite (Acari: Tetranychidae) in North Carolina peanut fields. Environ Entomol 13:221\u0026ndash;227. https://doi.org/10.1093/ee/13.1.221\u003c/li\u003e\n\u003cli\u003eBrandenburg R, Kennedy G (1982) Intercrop relationships and spider mite dispersal in a corn/peanut agro‐ecosystem. Entomol Exp Appl 32:269\u0026ndash;276. https://doi.org/10.1111/j.1570-7458.1982.tb03217.x\u003c/li\u003e\n\u003cli\u003eBruce TJA (2015) Interplay between insects and plants: Dynamic and complex interactions that have coevolved over millions of years but act in milliseconds. J Exp Bot 66:455\u0026ndash;465. https://doi.org/10.1093/jxb/eru391\u003c/li\u003e\n\u003cli\u003eBruce TJA, Pickett JA (2011) Perception of plant volatile blends by herbivorous insects \u0026ndash; Finding the right mix. Phytochemistry 72:1605\u0026ndash;1611. https://doi.org/10.1016/j.phytochem.2011.04.011\u003c/li\u003e\n\u003cli\u003eBruce TJA, Wadhams LJ, Woodcock CM (2005) Insect host location: A volatile situation. Trends Plant Sci 10:269\u0026ndash;274. https://doi.org/10.1016/j.tplants.2005.04.003\u003c/li\u003e\n\u003cli\u003eBurgue\u0026ntilde;o AP, Amor\u0026oacute;s ME, Deagosto E, Davyt B, D\u0026iacute;az M, Gonz\u0026aacute;lez A, Rossini C (2024) Preference and performance in an herbivorous coccinellid beetle: A comparative study of host plant defensive traits, insect preference, and survival. Arthropod-Plant Interact 18:617\u0026ndash;636. https://doi.org/10.1007/s11829-023-10004-x\u003c/li\u003e\n\u003cli\u003eCadet C, Ferri\u0026egrave;re R, Metz JAJ, van Baalen M (2003) The evolution of dispersal under demographic stochasticity. Am Nat 162:427\u0026ndash;441. https://doi.org/10.1086/378213\u003c/li\u003e\n\u003cli\u003eCarey J, Bradley J (1982) Developmental rates, vital schedules, sex ratios and life tables for \u003cem\u003eTetranychus urticae\u003c/em\u003e, \u003cem\u003eT. turkestani \u003c/em\u003eand \u003cem\u003eT. pacificus\u003c/em\u003e (Acarina: Tetranychidae) on cotton. Acarologia 23:333\u0026ndash;345.\u003c/li\u003e\n\u003cli\u003eChapman EJ, Byron CJ (2018) The flexible application of carrying capacity in ecology. Glob Ecol Conserv 13:e00365. https://doi.org/10.1016/j.gecco.2017.e00365\u003c/li\u003e\n\u003cli\u003eClobert J, Baguette M, Benton TG, Bullock JM (2012) Dispersal Ecology and Evolution. Oxford University Press.\u003c/li\u003e\n\u003cli\u003eClobert J, Le Galliard JF, Cote J, Meylan S, Massot M (2009) Informed dispersal, heterogeneity in animal dispersal syndromes and the dynamics of spatially structured populations. Ecol Lett 12:197\u0026ndash;209. https://doi.org/10.1111/j.1461-0248.2008.01267.x\u003c/li\u003e\n\u003cli\u003eClotuche G (2011) The silk as a thread to understand social behaviour in the weaving mite \u003cem\u003eTetranychus urticae\u003c/em\u003e. Thesis, UCL-Universit\u0026eacute; Catholique de Louvain, Belgium.\u003c/li\u003e\n\u003cli\u003eClotuche G, Mailleux AC, Astudillo Fern\u0026aacute;ndez A, Deneubourg JL, Detrain C, Hance T (2011) The formation of collective silk balls in the spider mite \u003cem\u003eTetranychus urticae\u003c/em\u003e Koch. PLoS ONE 6:e18854. https://doi.org/10.1371/journal.pone.0018854\u003c/li\u003e\n\u003cli\u003eClotuche G, Navajas M, Mailleux AC, Hance T (2013) Reaching the ball or missing the flight? Collective dispersal in the two-spotted spider mite \u003cem\u003eTetranychus urticae\u003c/em\u003e. PLoS ONE 8:e77573. https://doi.org/10.1371/journal.pone.0077573\u003c/li\u003e\n\u003cli\u003eDavis DW (1952) Influence of population density on \u003cem\u003eTetranychus multisetis\u003c/em\u003e. J Econ Entomol 45:652\u0026ndash;652. https://doi.org/10.1093/jee/45.4.652\u003c/li\u003e\n\u003cli\u003eden Boer PJ (1990) The survival value of dispersal in terrestrial arthropods. Biol Conserv 54:175\u0026ndash;192. https://doi.org/10.1016/0006-3207(90)90050-Y\u003c/li\u003e\n\u003cli\u003eDenno RF, Peterson MA, Gratton C, Cheng J, Langellotto GA, Huberty AF, Finke DL (2000) Feeding‐induced changes in plant quality mediate interspecific competition between sap‐feeding herbivores. Ecology 81:1814\u0026ndash;1827. https://doi.org/10.1890/0012-9658(2000)081[1814:FICIPQ]2.0.CO;2 \u003c/li\u003e\n\u003cli\u003eDhooria MS (2016) Fundamentals of Applied Acarology. Springer, Singapore.\u003c/li\u003e\n\u003cli\u003eDieckmann U, O\u0026apos;Hara B, Weisser W (1999) The evolutionary ecology of dispersal. Trends Ecol Evol 14:88\u0026ndash;90. https://doi.org/10.1016/S0169-5347(98)01571-7 \u003c/li\u003e\n\u003cli\u003eDuputi\u0026eacute; A, Massol F (2013) An empiricist\u0026apos;s guide to theoretical predictions on the evolution of dispersal. Interface Focus 3:20130028. https://doi.org/10.1098/rsfs.2013.0028\u003c/li\u003e\n\u003cli\u003eEstevez I, Andersen IL, Naevdal E (2007) Group size, density and social dynamics in farm animals. Appl Anim Behav Sci 103:185\u0026ndash;204. https://doi.org/10.1016/j.applanim.2006.05.025\u003c/li\u003e\n\u003cli\u003eFern\u0026aacute;ndez Ferrari MC, Schausberger P (2013) From repulsion to attraction: Species- and spatial context-dependent threat sensitive response of the spider mite \u003cem\u003eTetranychus urticae\u003c/em\u003e to predatory mite cues. Naturwissenschaften 100:541\u0026ndash;549. https://doi.org/10.1007/s00114-013-1050-5\u003c/li\u003e\n\u003cli\u003eFox CW (1994) The influence of egg size on offspring performance in the seed beetle, \u003cem\u003eCallosobruchus maculatus\u003c/em\u003e. Oikos 71:321\u0026ndash;325. https://doi.org/10.2307/3546280\u003c/li\u003e\n\u003cli\u003eFox CW, Czesak ME (2000) Evolutionary ecology of progeny size in arthropods. Ann Rev Entomol 45:341\u0026ndash;369. https://doi.org/10.1146/annurev.ento.45.1.341\u003c/li\u003e\n\u003cli\u003eGotoh T, Moriya D, Nachman G (2015) Development and reproduction of five \u003cem\u003eTetranychus\u003c/em\u003e species (Acari: Tetranychidae): Do they all have the potential to become major pests? Exp Appl Acarol 66:453\u0026ndash;479. https://doi.org/10.1007/s10493-015-9919-y \u003c/li\u003e\n\u003cli\u003eGoulden CE, Henry L, Berrigan D (1987) Egg size, postembryonic yolk, and survival ability. Oecologia 72:28\u0026ndash;31. https://doi.org/10.1007/BF00385040 \u003c/li\u003e\n\u003cli\u003eGrove M (2012) Space, time, and group size: A model of constraints on primate social foraging. Anim Behav 83:411\u0026ndash;419. https://doi.org/10.1016/j.anbehav.2011.11.011\u003c/li\u003e\n\u003cli\u003eHazan A, Gerson U, Tahori A (1974) Spider mite webbing. I. The production of webbing under various environmental conditions. Acarologia 16:68\u0026ndash;84.\u003c/li\u003e\n\u003cli\u003eHewison AJM, Morellet N, Debeffe L, Cargnelutti B, Gaillard JM, Cagnacci F, Gehr B, Kr\u0026ouml;schel M, Heurich M, Coulon A, Kjellander P, B\u0026ouml;rger L, Focardi S (2021) Sex differences in condition dependence of natal dispersal in a large herbivore: Dispersal propensity and distance are decoupled. Proc R Soc B Biol Sci 288:20202947. https://doi.org/10.1098/rspb.2020.2947\u003c/li\u003e\n\u003cli\u003eHollander J, Verzijden M, Svensson E, Br\u0026ouml;nmark C (2014) Animal movement across scales. In: Hansson LA, \u0026Aring;kesson S (eds) Animal Movement Across Scales. Oxford University Press, pp 110\u0026ndash;125.\u003c/li\u003e\n\u003cli\u003eHussey NW, Parr WJ (1963) Dispersal of the glasshouse red spider mite \u003cem\u003eTetranychus urticae\u003c/em\u003e Koch (Acarina, Tetranychidae). Entomol Exp Appl 6:207\u0026ndash;214. https://doi.org/10.1111/j.1570-7458.1963.tb00619.x\u003c/li\u003e\n\u003cli\u003eJeppson LR, Keifer HH, Baker EW (1975) Mites Injurious to Economic Plants. University of California Press.\u003c/li\u003e\n\u003cli\u003eJermy T, 1984. Evolution of insect/host plant relationships. Am Nat 124:609\u0026ndash;630. http://www.jstor.org/stable/2461372 \u003c/li\u003e\n\u003cli\u003eJohnson DM, Horvitz CC (2005) Estimating postnatal dispersal: Tracking the unseen dispersers. Ecology 86:1185\u0026ndash;1190. https://doi.org/10.1890/04-0974\u003c/li\u003e\n\u003cli\u003eJohnston TA, Leggett WC (2002) Maternal and environmental gradients in the egg size of an iteroparous fish. Ecology 83:1777\u0026ndash;1791. https://doi.org/10.1890/0012-9658(2002)083[1777:MAEGIT]2.0.CO;2\u003c/li\u003e\n\u003cli\u003eJung C (2005) Some evidences of aerial dispersal of twospotted spider mites from an apple orchard into a soybean field. J Asia-Pac Entomol 8:279\u0026ndash;283. https://doi.org/10.1016/S1226-8615(08)60246-0\u003c/li\u003e\n\u003cli\u003eKant MR, Sabelis MW, Haring MA, Schuurink RC (2008) Intraspecific variation in a generalist herbivore accounts for differential induction and impact of host plant defences. Proc R Soc B Biol Sci 275:443\u0026ndash;452. https://doi.org/10.1098/rspb.2007.1277\u003c/li\u003e\n\u003cli\u003eKarlsson Green K, Chiara De P, Maria L, Peter A (2024) Population comparison of innate and plastic host plant preference and performance in a polyphagous insect. Front Ecol Evol 12:1426923. https://doi.org/10.3389/fevo.2024.1426923\u003c/li\u003e\n\u003cli\u003eKarlsson B, Wiklund C (1985) Egg weight variation in relation to egg mortality and starvation endurance of newly hatched larvae in some satyrid butterflies. Ecol Entomol 10:205\u0026ndash;211. https://doi.org/10.1111/j.1365-2311.1985.tb00549.x\u003c/li\u003e\n\u003cli\u003eKhuhro NH, Biondi A, Desneux N, Zhang L, Zhang Y, Chen H (2014) Trade-off between flight activity and life-history components in \u003cem\u003eChrysoperla sinica\u003c/em\u003e. BioControl 59:219\u0026ndash;227. https://doi.org/10.1007/s10526-014-9560-4 \u003c/li\u003e\n\u003cli\u003eKnolhoff LM, Heckel DG (2014) Behavioral assays for studies of host plant choice and adaptation in herbivorous insects. Ann Rev Entomol 59:263\u0026ndash;278. https://doi.org/10.1146/annurev-ento-011613-161945 \u003c/li\u003e\n\u003cli\u003eKondo A, Takafuji A (1985) Resource utilization pattern of two species of tetranychid mites (Acarina: Tetranychidae). Res Popul Ecol 27:145\u0026ndash;157. https://doi.org/10.1007/BF02515487 \u003c/li\u003e\n\u003cli\u003eKot M, Lewis MA, van den Driessche P (1996) Dispersal data and the spread of invading organisms. Ecology 77:2027\u0026ndash;2042. https://doi.org/10.2307/2265698\u003c/li\u003e\n\u003cli\u003eKrips OE, Witul A, Willems PEL, Dicke M (1998) Intrinsic rate of population increase of the spider mite \u003cem\u003eTetranychus urticae\u003c/em\u003e on the ornamental crop gerbera: Intraspecific variation in host plant and herbivore. Entomol Exp Appl 89:159\u0026ndash;168. https://doi.org/10.1046/j.1570-7458.1998.00395.x\u003c/li\u003e\n\u003cli\u003eLalonde R (2005) Egg size variation does not affect offspring performance under intraspecific competition in \u003cem\u003eNasonia vitripennis\u003c/em\u003e, a gregarious parasitoid. J Anim Ecol 74:630\u0026ndash;635. https://doi.org/10.1111/j.1365-2656.2005.00958.x\u003c/li\u003e\n\u003cli\u003eLe Goff GJ, Mailleux AC, Detrain C, Deneubourg JL, Clotuche G, Hance T (2010) Group effect on fertility, survival and silk production in the web spinner \u003cem\u003eTetranychus urticae\u003c/em\u003e (Acari: Tetranychidae) during colony foundation. Behaviour 147:1169\u0026ndash;1184. https://doi.org/10.1163/000579510X510980\u003c/li\u003e\n\u003cli\u003eLi G-Y, Zhang Z-Q (2021) The costs of social interaction on survival and reproduction of arrhenotokous spider mite \u003cem\u003eTetranychus urticae\u003c/em\u003e. Entomol Gen 41:49\u0026ndash;57. https://doi.org/10.1127/entomologia/2020/0911 \u003c/li\u003e\n\u003cli\u003eLi J, Margolies DC (1993) Effects of mite age, mite density, and host quality on aerial dispersal behavior in the twospotted spider mite. Entomol Exp Appl 68:79\u0026ndash;86. https://doi.org/10.1111/j.1570-7458.1993.tb01691.x\u003c/li\u003e\n\u003cli\u003eLinke W (1953) Investigation of the biology and epidemiology of the common spider mite, \u003cem\u003eTetranychus althaeae\u003c/em\u003e v. Hanst. with particular consideration of the hop as the host. Hoefchen-Briefe Bayer Pflanz. Nachr. 6:181\u0026ndash;232.\u003c/li\u003e\n\u003cli\u003eLombaert E, Boll R, Lapchin L (2006) Dispersal strategies of phytophagous insects at a local scale: Adaptive potential of aphids in an agricultural environment. BMC Evol Biol 6:1\u0026ndash;13. https://doi.org/10.1186/1471-2148-6-75\u003c/li\u003e\n\u003cli\u003eMacke E, Magalh\u0026atilde;es S, Bach F, Olivieri I (2011a) Experimental evolution of reduced sex ratio adjustment under local mate competition. Science 334:1127\u0026ndash;1129. https://doi.org/10.1126/science.1212177\u003c/li\u003e\n\u003cli\u003eMacke E, Magalhaes S, Khan HDT, Luciano A, Frantz A, Facon B, Olivieri I (2011b) Sex allocation in haplodiploids is mediated by egg size: Evidence in the spider mite \u003cem\u003eTetranychus urticae\u003c/em\u003e Koch. Proc R Soc B Biol Sci 278:1054\u0026ndash;1063. https://doi.org/10.1098/rspb.2010.1706\u003c/li\u003e\n\u003cli\u003eMacke E, Magalhaes S, Khanh Do-Thi H, Frantz A, Facon B, Olivieri I (2012) Mating modifies female life history in a haplodiploid spider mite. Am Nat 179:147\u0026ndash;162. https://doi.org/10.1086/665002\u003c/li\u003e\n\u003cli\u003eMaeno KO, Piou C, Ghaout S (2020) The desert locust, \u003cem\u003eSchistocerca gregaria\u003c/em\u003e, plastically manipulates egg size by regulating both egg numbers and production rate according to population density. J Insect Physiol 122:104020. https://doi.org/10.1016/j.jinsphys.2020.104020\u003c/li\u003e\n\u003cli\u003eMartin JP, Beyerlein A, Dacks AM, Reisenman CE, Riffell JA, Lei H, Hildebrand JG (2011) The neurobiology of insect olfaction: Sensory processing in a comparative context. Prog Neurobiol 95:427\u0026ndash;447. https://doi.org/10.1016/j.pneurobio.2011.09.007\u003c/li\u003e\n\u003cli\u003eMcCauley SJ (2010) Body size and social dominance influence breeding dispersal in male \u003cem\u003ePachydiplax longipennis\u003c/em\u003e (Odonata). Ecol Entomol 35:377\u0026ndash;385. https://doi.org/10.1111/j.1365-2311.2010.01191.x\u003c/li\u003e\n\u003cli\u003eMcEdward LR, Carson SF (1987) Variation in egg organic content and its relationship with egg size in the starfish \u003cem\u003eSolaster stimpsoni\u003c/em\u003e. Mar Ecol Prog Ser 37:159\u0026ndash;169.\u003c/li\u003e\n\u003cli\u003eMcMurtry J, Huffaker C, van de Vrie M (1970) Ecology of tetranychid mites and their natural enemies: A review: I. Tetranychid enemies: Their biological characters and the impact of spray practices. Hilgardia 40:331\u0026ndash;390. https://doi.org/10.3733/hilg.v40n11p331\u003c/li\u003e\n\u003cli\u003eMcPeek MA, Holt RD (1992) The evolution of dispersal in spatially and temporally varying environments. Am Nat 140:1010\u0026ndash;1027. https://doi.org/10.1111/evo.12699\u003c/li\u003e\n\u003cli\u003eMeiners T (2015) Chemical ecology and evolution of plant-insect interactions: A multitrophic perspective. Curr Opin Insect Sci 8:22\u0026ndash;28. https://doi.org/10.1016/j.cois.2015.02.003\u003c/li\u003e\n\u003cli\u003eMellors WK, Propts SE (1983) Effects of fertilizer level, fertility balance, and soil moisture on the interaction of two-spotted spider mites (Acarina: Tetranychidae) with radish plants. Environ Entomol 12:1239\u0026ndash;1239. https://doi.org/10.1093/ee/12.4.1239\u003c/li\u003e\n\u003cli\u003eMigeon A, Dorkeld F (2024) Spider mites web: A comprehensive database for the Tetranychidae. https://www1.montpellier.inra.fr/CBGP/spmweb (Accessed 30 October 2024).\u003c/li\u003e\n\u003cli\u003eMitchell R (1973) Growth and population dynamics of a spider mite (\u003cem\u003eTetranychus urticae\u003c/em\u003e K., Acarina: Tetranychidae). Ecology 54:1349\u0026ndash;1355. https://doi.org/10.2307/1934198\u003c/li\u003e\n\u003cli\u003eMorrongiello JR, Bond NR, Crook DA, Wong BBM (2012) Spatial variation in egg size and egg number reflects trade-offs and bet-hedging in a freshwater fish. J Anim Ecol 81:806\u0026ndash;817. https://doi.org/10.1111/j.1365-2656.2012.01961.x\u003c/li\u003e\n\u003cli\u003eNasu S, Tokuda M (2021) Dispersal\u0026ndash;reproduction trade-off in the leaf beetle \u003cem\u003eGalerucella grisescens\u003c/em\u003e. Entomol Exp Appl 169:542\u0026ndash;549. https://doi.org/10.1111/eea.13042\u003c/li\u003e\n\u003cli\u003eOsakabe M, Isobe H, Kasai A, Masuda R, Kubota S, Umeda M (2008) Aerodynamic advantages of upside down take-off for aerial dispersal in \u003cem\u003eTetranychus\u003c/em\u003e spider mites. Exp Appl Acarol 44:165\u0026ndash;183. https://doi.org/10.1007/s10493-008-9141-2 \u003c/li\u003e\n\u003cli\u003eParker GA, Begon M (1986) Optimal egg size and clutch size: Effects of environment and maternal phenotype. Am Nat 128:573\u0026ndash;592. https://doi.org/10.1086/284589\u003c/li\u003e\n\u003cli\u003eParvinen K, Egas M (2004) Dispersal and the evolution of specialisation in a two-habitat type metapopulation. Theor Popul Biol 66:233\u0026ndash;248. https://doi.org/10.1016/j.tpb.2004.06.002\u003c/li\u003e\n\u003cli\u003ePepi AA, Broadley HJ, Elkinton JS (2016) Density-dependent effects of larval dispersal mediated by host plant quality on populations of an invasive insect. Oecologia 182:499\u0026ndash;509. https://doi.org/10.1007/s00442-016-3689-z \u003c/li\u003e\n\u003cli\u003ePhillips BL, Brown GP, Shine R (2010) Life-history evolution in range-shifting populations. Ecology 91:1617\u0026ndash;1627. https://doi.org/10.1890/09-0910.1\u003c/li\u003e\n\u003cli\u003ePick JL, Hutter P, Tschirren B (2016) In search of genetic constraints limiting the evolution of egg size: Direct and correlated responses to artificial selection on a prenatal maternal effector. Heredity 116:542\u0026ndash;549. https://doi.org/10.1038/hdy.2016.16 \u003c/li\u003e\n\u003cli\u003ePlazio E, Margol T, Nowicki P (2020) Intersexual differences in density-dependent dispersal and their evolutionary drivers. J Evol Biol 33:1495\u0026ndash;1506. https://doi.org/10.1111/jeb.13688\u003c/li\u003e\n\u003cli\u003ePoethke HJ, Gros A, Hovestadt T (2011) The ability of individuals to assess population density influences the evolution of emigration propensity and dispersal distance. J Theor Biol 282:93\u0026ndash;99. https://doi.org/10.1016/j.jtbi.2011.05.012\u003c/li\u003e\n\u003cli\u003ePoethke HJ, Hovestadt T (2002) Evolution of density- and patch-size-dependent dispersal rates. Proc R Soc B Biol Sci 269:637\u0026ndash;645. https://doi.org/10.1098/rspb.2001.1936\u003c/li\u003e\n\u003cli\u003ePuzin C, P\u0026eacute;tillon J, Bonte D (2018) Influence of individual density and habitat availability on long-distance dispersal in a salt-marsh spider. Ethol Ecol Evol 31:28\u0026ndash;37. https://doi.org/10.1080/03949370.2018.1486888\u003c/li\u003e\n\u003cli\u003eReisig DD, Godfrey LD, Marcum DB (2010) Plant quality and conspecific density effects on \u003cem\u003eAnaphothrips obscurus\u003c/em\u003e (Thysanoptera: Thripidae) wing diphenism and population ecology. Environ Entomol 39:685\u0026ndash;694. https://doi.org/10.1603/EN09332\u003c/li\u003e\n\u003cli\u003eRenault D, Laparie M, McCauley SJ, Bonte D (2018) Environmental adaptations, ecological filtering, and dispersal central to insect invasions. Annu Rev Entomol 63:345\u0026ndash;368. https://doi.org/10.1146/annurev-ento-020117-043315\u003c/li\u003e\n\u003cli\u003eRhainds M, Shipp L (2003) Dispersal of adult western flower thrips (Thysanoptera: Thripidae) on Chrysanthemum plants: Impact of feeding-induced senescence of inflorescences. Environ Entomol 32:1056\u0026ndash;1065. https://doi.org/10.1603/0046-225X-32.5.1056\u003c/li\u003e\n\u003cli\u003eRhainds M, Shipp L, Woodrow L, Anderson D (2005) Density, dispersal, and feeding impact of western flower thrips (Thysanoptera: Thripidae) on flowering chrysanthemum at different spatial scales. Ecol Entomol 30:96\u0026ndash;104. https://doi.org/10.1111/j.0307-6946.2005.00663.x\u003c/li\u003e\n\u003cli\u003eRioja C, Zhurov V, Bruinsma K, Grbic M, Grbic V (2017) Plant-Herbivore interactions: A case of an extreme generalist, the two-spotted spider mite \u003cem\u003eTetranychus urticae\u003c/em\u003e. MPMI 30:935\u0026ndash;945. https://doi.org/10.1094/MPMI-07-17-0168-CR\u003c/li\u003e\n\u003cli\u003eRoeder C, Harmsen R, Mouldey S (1996) The effects of relatedness on progeny sex ratio in spider mites. J Evol Biol 9:143\u0026ndash;151. https://doi.org/10.1046/j.1420-9101.1996.9020143.x\u003c/li\u003e\n\u003cli\u003eRonce O, Clobert J (2012) Dispersal syndromes. In: Clobert J, Baguette M, Benton TG, Bullock JM (eds) Dispersal Ecology and Evolution. Oxford University Press, pp. 119\u0026ndash;138.\u003c/li\u003e\n\u003cli\u003eRuf D, Dorn S, Mazzi D (2011) Females leave home for sex: Natal dispersal in a parasitoid with complementary sex determination. Anim Behav 81:1083\u0026ndash;1089. https://doi.org/10.1016/j.anbehav.2011.02.028\u003c/li\u003e\n\u003cli\u003eRushing CS, Brandt Ryder T, Valente JJ, Scott Sillett T, Marra PP (2021) Empirical tests of habitat selection theory reveal that conspecific density and patch quality, but not habitat amount, drive long‐distance immigration in a wild bird. Ecol Lett 24:1167\u0026ndash;1177. https://doi.org/10.1111/ele.13729\u003c/li\u003e\n\u003cli\u003eSabelis MW, Bakker FM (1992) How predatory mites cope with the web of their tetranychid prey: A functional view on dorsal chaetotaxy in the Phytoseiidae. Exp Appl Acarol 16:203\u0026ndash;225. https://doi.org/10.1007/BF01193804 \u003c/li\u003e\n\u003cli\u003eSandeson PD, Boiteau G, Le Blanc J PR (2002) Adult density and the rate of colorado potato beetle (Coleoptera: Chrysomelidae) flight take-off. Environ Entomol 31:533\u0026ndash;537. https://doi.org/10.1603/0046-225X-31.3.533\u003c/li\u003e\n\u003cli\u003eSchausberger P, Yano S, Sato Y (2021) Cooperative behaviors in group-living spider mites. Front Ecol Evol 9:745036. https://doi.org/10.3389/fevo.2021.745036\u003c/li\u003e\n\u003cli\u003eShih CT, Poe SL, Cromroy HL (1976) Biology, life table, and intrinsic rate of increase of \u003cem\u003eTetranychus urticae\u003c/em\u003e. Ann Entomol Soc Am 69:362\u0026ndash;364. https://doi.org/10.1093/aesa/69.2.362\u003c/li\u003e\n\u003cli\u003eSloggett JJ, Weisser WW (2002) Parasitoids induce production of the dispersal morph of the pea aphid, \u003cem\u003eAcyrthosiphon pisum\u003c/em\u003e. Oikos 98:323\u0026ndash;333. https://doi.org/10.1034/j.1600-0706.2002.980213.x\u003c/li\u003e\n\u003cli\u003eSmith CC, Fretwell SD (1974) The optimal balance between size and number of offspring. Am Nat 108:499\u0026ndash;506. https://www.jstor.org/stable/2459681 \u003c/li\u003e\n\u003cli\u003eSmitley DR, Kennedy GG (1988) Aerial dispersal of the two-spotted spider mite (\u003cem\u003eTetranychus urticae\u003c/em\u003e) from field corn. Exp Appl Acarol 5:33\u0026ndash;46. https://doi.org/10.1007/BF02053815 \u003c/li\u003e\n\u003cli\u003eStamps J (2001) Habitat selection by dispersers: integrating proximate and ultimate approaches. In: Clobert J, Danchin E, Dhondt AA, Nichols JD (eds) Dispersal. Oxford University Press, pp 110\u0026ndash;122. https://doi.org/10.1093/oso/9780198506607.003.0018\u003c/li\u003e\n\u003cli\u003eStearns SC (1989) Trade-offs in life-history evolution. Funct Ecol 3:259\u0026ndash;268. https://doi.org/10.2307/2389364 \u003c/li\u003e\n\u003cli\u003eStrong WB, Croft BA, Slone DH (1997) Spatial aggregation and refugia of the mites \u003cem\u003eTetranychus urticae\u003c/em\u003e and \u003cem\u003eNeoseiulus fallacis\u003c/em\u003e (Acari: Tetranychidae, Phytoseiidae) on Hop. Environ Entomol 26:859\u0026ndash;865. https://doi.org/10.1093/ee/26.4.859\u003c/li\u003e\n\u003cli\u003eSuski Z, Badowska T (1975) Effect of the host plant nutrition on the population of the two spotted spider mite, \u003cem\u003eTetranychus urticae\u003c/em\u003e Koch (Acarina, Tetranychidae). Ekologia Polska 23:185\u0026ndash;209.\u003c/li\u003e\n\u003cli\u003eTien NS, Sabelis MW, Egas M (2011) Ambulatory dispersal in \u003cem\u003eTetranychus urticae\u003c/em\u003e: An artificial selection experiment on propensity to disperse yields no response. Exp Appl Acarol 53:349\u0026ndash;360. https://doi.org/10.1007/s10493-010-9411-7 \u003c/li\u003e\n\u003cli\u003eTinsley Johnson E, Feder JA, Lu A, Bergman TJ, Beehner JC, Snyder-Mackler N (2021) The goldilocks effect: Female geladas in mid-sized groups have higher fitness. Proc R Soc B 288:20210820. https://doi.org/10.1098/rspb.2021.0820\u003c/li\u003e\n\u003cli\u003eTravis JMJ, Mustin K, Palmer SCF, Bartoń KA, Hovestadt T, Benton TG, Clobert J, Delgado MM, Dytham C, Van Dyck H, Bonte D (2012) Modelling dispersal: An eco-evolutionary framework incorporating emigration, movement, settlement behaviour and the multiple costs involved. Methods Ecol Evol 3:628\u0026ndash;641. https://doi.org/10.1111/j.2041-210X.2012.00193.x\u003c/li\u003e\n\u003cli\u003eTuan SJ, Lin YH, Yang CM, Atlihan R, Saska P, Chi H (2016) Survival and reproductive strategies in two-spotted spider mites: demographic analysis of arrhenotokous parthenogenesis of \u003cem\u003eTetranychus urticae\u003c/em\u003e (Acari: Tetranychidae). J Econ Entomol 109:502\u0026ndash;509. https://doi.org/10.1093/jee/tov386\u003c/li\u003e\n\u003cli\u003eWalzer A, Schausberger P (2015) Food stress causes sex-specific maternal effects in mites. J Exp Biol 218:2603\u0026ndash;2609. https://doi.org/10.1242/jeb.123752\u003c/li\u003e\n\u003cli\u003eWatson TF (1964) Influence of host plant condition on population increase of \u003cem\u003eTetranychus telarius\u003c/em\u003e (Linnaeus) (Acarina: Tetranychidae). Hilgardia 35:273\u0026ndash;322. https://doi.org/10.3733/hilg.v35n11p273 \u003c/li\u003e\n\u003cli\u003eWeerawansha N, Wang Q, He XZ (2022a) A haplodiploid mite adjusts fecundity and sex ratio in response to density changes during the reproductive period. Exp Appl Acarol 88:277\u0026ndash;288. https://doi.org/10.1007/s10493-022-00749-0 \u003c/li\u003e\n\u003cli\u003eWeerawansha N, Wang Q, He XZ (2022b) Comparing the effects of social environments and life history traits on sex allocation in a haplodiploid spider mite. Syst Appl Acarol 27:2123\u0026ndash;2130. https://doi.org/10.11158/saa.27.10.20\u003c/li\u003e\n\u003cli\u003eWeerawansha N, Wang Q, He XZ (2023a) Local mate competition model alone cannot predict the offspring sex ratio in large and dense populations of a haplodiploid arthropod. Curr Zool 69:219\u0026ndash;221. https://doi.org/10.1093/cz/zoac022\u003c/li\u003e\n\u003cli\u003eWeerawansha N, Wang Q, He XZ (2023b) Reproductive plasticity in response to the changing cluster size during the breeding period: A case study in a spider mite. Exp Appl Acarol 91:237\u0026ndash;250. https://doi.org/10.1007/s10493-023-00834-y \u003c/li\u003e\n\u003cli\u003eWeerawansha N, Wang Q, He XZ (2024) Conspecific cues mediate habitat selection and reproductive performance in a haplodiploid spider mite. Curr Zool 70:795\u0026ndash;802. https://doi.org/10.1093/cz/zoae013\u003c/li\u003e\n\u003cli\u003eWermelinger B, Oertli JJ, Delucchi V (1985) Effect of host plant nitrogen fertilization on the biology of the two‐spotted spider mite, \u003cem\u003eTetranychus urticae\u003c/em\u003e. Entomol Exp Appl 38:23\u0026ndash;28. https://doi.org/10.1111/j.1570-7458.1985.tb03493.x\u003c/li\u003e\n\u003cli\u003eWiklund C, Persson A (1983) Fecundity, and the relation of egg weight variation to offspring fitness in the speckled wood butterfly \u003cem\u003ePararge aegeria\u003c/em\u003e, or why don\u0026apos;t butterfly females lay more eggs? Oikos 40:53\u0026ndash;63. https://doi.org/10.2307/3544198 \u003c/li\u003e\n\u003cli\u003eWilliams JL, Hufbauer RA, Miller TEX (2019) How evolution modifies the variability of range expansion. Trends Ecol Evol 34:903\u0026ndash;913. https://doi.org/10.1016/j.tree.2019.05.012\u003c/li\u003e\n\u003cli\u003eWilson LJ (1994) Plant-quality effect on life-history parameters of the twospotted spider mite (Acari: Tetranychidae) on cotton. J Econ Entomol 87:1665\u0026ndash;1673. https://doi.org/10.1093/jee/87.6.1665\u003c/li\u003e\n\u003cli\u003eWong JWY, Meunier J, Koelliker M (2013) The evolution of parental care in insects: The roles of ecology, life history and the social environment. Ecol Entomol 38:123\u0026ndash;137. https://doi.org/10.1111/een.12000\u003c/li\u003e\n\u003cli\u003eXu F, Yang W, Li Y (2019) Enlarged egg size increases offspring fitness of a frog species on the Zhoushan Archipelago of China. Sci Rep 9:11653. https://doi.org/10.1038/s41598-019-48147-8 \u003c/li\u003e\n\u003cli\u003eYano S (2008) Collective and solitary behaviors of twospotted spider mite (Acari: Tetranychidae) are induced by trail following. Ann Entomol Soc Am 101:247\u0026ndash;252. https://doi.org/10.1603/0013-8746(2008)101[247:CASBOT]2.0.CO;2\u003c/li\u003e\n\u003cli\u003eYano S (2012) Cooperative web sharing against predators promotes group living in spider mites. Behav Ecol Sociobiol 66:845\u0026ndash;853. https://doi.org/10.1007/s00265-012-1332-5 \u003c/li\u003e\n\u003cli\u003eYoung SSY, Wrensch DL, Kongchuensin M (1986) Control of sex ratio by female spider mites. Entomol Exp Appl 40:53\u0026ndash;60. https://doi.org/10.1111/j.1570-7458.1986.tb02155.x\u003c/li\u003e\n\u003cli\u003eZhang Z-Q (2003) Mites of Greenhouses: Identification, Biology and Control. CABI Publishing, UK.\u003c/li\u003e\n\u003cli\u003eZhou P, He XZ, Chen C, Wang Q (2021) Resource relocations in relation to dispersal in \u003cem\u003eTetranychus ludeni\u003c/em\u003e Zacher. Syst Appl Acarol 26:2018\u0026ndash;2026. https://doi.org/10.11158/saa.26.11.3\u003c/li\u003e\n\u003cli\u003eZhou P, He XZ, Chen C, Wang Q (2024) Age and density of mated females affect dispersal strategies in spider mite \u003cem\u003eTetranychus ludeni\u003c/em\u003e Zacher. Insects 15:387. https://doi.org/10.3390/insects15060387\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","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":"Habitat quality, Habitat selection, Population density, Reproductive performance, Tetranychus ludeni","lastPublishedDoi":"10.21203/rs.3.rs-6772534/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6772534/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eDispersal is a key adaptative strategy to escape deteriorating environments, and habitat selection by dispersers is critical to their own and offspring fitness. Using the haplodiploid spider mite \u003cem\u003eTetranychus ludeni\u003c/em\u003e Zacher as a model species, this study investigated how host plant quality influenced dispersal probability, habitat selection, and subsequent reproductive performances of mated females. We designed two dispersal scenarios, i.e., females were allowed to disperse from low-quality (LQ) or high-quality (HQ) habitat and select between LQ and HQ habitats. Results show that significantly more females dispersed from LQ habitats than from HQ habitats, and dispersers significantly preferring and settling in HQ habitats regardless of the dispersal scenarios. However, aggregating in HQ habitats resulted in higher number of eggs cumulated but also increased immature mortality. Individual females restrained reproductive output under the deteriorating environments. Egg size had no significant effect on egg hatching or immature survival. Females dispersed from LQ habitats produced significantly smaller eggs but maintained similar proportion of daughters compared to those remaining in LQ habitats. Females dispersed from HQ habitats produced eggs of similar size but significantly higher proportion of daughters. These results suggest that dispersing females might manipulate offspring sex ratio by lowing the fertilization threshold to fertilise relatively smaller eggs. Population density has less impact on egg size and offspring sex ratio. This study delivers insights into the dispersal and reproductive strategies of a haplodiploid spider mite, highlighting how host quality shapes adaptive responses in challenging environments.\u003c/p\u003e","manuscriptTitle":"Host plant quality mediates dispersal, oviposition, and sex allocation in a Tetranychus spider mite","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-18 16:51:42","doi":"10.21203/rs.3.rs-6772534/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","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":"17b5cc6f-838a-44c7-91a7-2029dac33740","owner":[],"postedDate":"June 18th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-09-09T11:19:57+00:00","versionOfRecord":[],"versionCreatedAt":"2025-06-18 16:51:42","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6772534","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6772534","identity":"rs-6772534","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}