{"paper_id":"4bf66985-4c54-450a-8d62-1ac7d7f0f7e4","body_text":"A test of trade-offs in dispersal and reproduction within and between a sister species pair of specialist insect herbivores | 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 A test of trade-offs in dispersal and reproduction within and between a sister species pair of specialist insect herbivores Amy M. Roush, Linyi Zhang, Glen Ray Hood, James R. Ott, Scott P. Egan This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3171363/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 07 Feb, 2024 Read the published version in Oecologia → Version 1 posted 4 You are reading this latest preprint version Abstract A critical question in understanding the origin of trade-offs and their impact on trait evolution is whether trade-offs between species originate from within-species variation. Despite their importance, studies linking interspecific divergence between closely related species to intraspecific trait variation are still rare. In this study, we describe a trade-off between dispersal and reproductive effort between two sympatric sister species of wasps in the genus Belonocnema (Hymenoptera: Cynipini: Cynipidae) that form galls on live oaks: B. fossoria , which specializes on Quercus geminata , and B. treatae , which specializes on Q. virginiana . Specifically, our results suggest that B. fossoria has evolved reduced flight capability and smaller wings, but a larger abdomen and greater total reproductive effort than B. treatae , which has larger wings and is a stronger flier, but a smaller abdomen and reduced total reproductive effort. Despite these significant morphological and reproductive differences, these traits remain unchanged when transplanting B. fossoria and B. treatae onto the alternative host plant, suggesting that divergence in these traits is likely genetic as opposed to a plastic response to the different rearing environments. However, we did not find evidence of intraspecific trade-offs between wing length and reproductive traits within either B. fossoria or B. treatae , indicating that trade-offs in life history traits between the two species is a result of independent adaptations in response to different environments. Our study informs our understanding of the evolution of trade-offs among life history traits by examining trade-offs at different biological organizations. Belonocnema gall wasp host-plant adaptation life history traits Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Introduction Trade-offs in life history traits are commonly observed among organisms adapting to different environmental conditions (Roff and Fairbairn 2007). Classic examples include trade-offs between growth rate and reproduction (e.g., Stearns 1989), survival and reproduction (e.g., Partridge & Farquhar 1981), and dispersal and reproduction (e.g., Zera and Denno 1997 , Carroll et al. 2003 ). One trait that is critical in mediating local adaptation to variable environments and regulating population connectivity is dispersal capacity (Stevens et al. 2013 ). For example, insect populations with higher dispersal ability are often associated with taller plants or ephemeral resources, as flight ability allows insects to better navigate complex habitats and/or access additional resources (Denno et al. 1996 ; Zera and Denno 1997 ; Duthie et al. 2015 ; Dell’Aglio et al. 2022 ). Thus, dispersal capacity ultimately can influence the dynamics of population divergence and speciation (Claramunt et al. 2012; Stevens et al. 2013 ). However, the evolution of increased dispersal capacity is often accompanied by a trade-off in reproductive output due to the constraint of limited energy resources (Zera and Denno 1997 ; Zera and Harshman 2001 ; Ghalambor et al. 2004 ; Vorburger 2005 ). Such trade-offs between dispersal and reproduction can be critical in maintaining population sizes under fluctuating environments, promoting species coexistence, and/or affecting range expansion (Stevens et al. 2013 ; Duthie et al. 2015 ). Trade-offs between dispersal and reproduction are often found within wing-dimorphic insect species (Roff 1986 ; Guerra 2011 ), where flight-capable, long-winged ‘macropterous’ morphs and reduced-flight, short-winged ‘brachypterous’ morphs are each present within a species and can be expressed as plastic responses to fluctuating environments (Zera and Rankin 1989 ; Carroll et al. 2003 ; Nasu and Tokuda 2021 ). However, divergence in trade-offs between dispersal and reproduction among closely related species within a genus are less prevalent (Tigreros and Davidowitz 2019 ). Investigating whether such interspecific trade-offs scale from intraspecific variation is particularly interesting with respect to understanding how divergence across a suite of traits between species can evolve (Worley et al. 2000 ; Sargent et al. 2007 ; Agrawal 2020 ). For example, if trade-offs between traits are found at both the intra- and interspecific levels, trait divergence between species are likely constrained by trade-offs from intraspecific variation (Agrawal 2020 ). Alternatively, if interspecies differences in flight capacity and any associated trade-off(s) do not scale from intraspecific variation, this may reflect independent evolutionary trajectories between species in response to different environmental conditions. Another possibility is that the lack of trade-offs between life history traits within species is because resource acquisition varies greatly among individuals, thus difference in life history traits due to resource availability masks the evolutionary trade-offs among different traits (van Noordwijk et al. 1986). To date, however, few studies have examined trade-offs between dispersal and reproduction between closely related species, while simultaneously exploring whether evidence of trade-offs exist within each species (Agrawal 2020 ). Sympatric sister species of gall wasps in the genus Belonocnema (Hymenoptera: Cynipidae) provide an opportunity to test the hypothesis that trade-offs between dispersal and reproduction are similar at each level of biological orgainization. First, the two sister species, B. fossoria and B. treatae , differ in flight capacity; asexual generation B. fossoria , specialized to develop on shorter sand live oak, Quercus geminata , are flightless, while asexual generation B. treatae , specialized on taller southern live oak, Q. virginiana , are capable of flight (Zhang et al. 2021c ). Second, Belonocnema do not feed as adults, thus, the pool of resources available for allocation to dispersal (flight capacity) and reproduction (egg number and egg size) is finite, with those resources derived from stores acquired during the earlier larval feeding period (Lund et al. 1998 ; Zera and Harshman 2001 ; Bonnet et al. 2002 ). Third, the two species are sympatric in a large portion of their range, but occupy different host plants that represent different environmental challenges (Cavender-Bares and Pahlich 2009 ; Cavender‐Bares et al. 2015). Finally, the species are near-completely reproductively isolated because of multiple pre- and postzygotic reproductive barriers (Egan et al. 2012a , b ; Zhang et al. 2017 , 2021a , b ; Hood et al. 2019 ) that result in sympatric host-associated populations that are genomically distinct (Driscoe et al. 2019 ; Zhang et al. 2021c ). Here, we first test whether the difference in flight capacity of B. fossoria and B. treatae is associated with between-species differences in wing length, body size and/or reproductive traits, as predicted by a dispersal/reproduction trade-off. Second, we test the nature of potential trade-offs in life history traits between species by extending the framework proposed by Hood and Ott ( 2017 ) and illustrated in Fig. 1 to test for independent adjustments of wing length and reproductive traits against body size between B. fossoria and B. treatae . Third, we conduct a reciprocal transplant experiment to test whether such the observed trade-offs in life history traits between species are genetic or a plastic response to the different host plant rearing environments. Finally, we test whether the trade-off in dispersal and reproduction observed between species also exists within either ecologically divergent sister species. Methods Biology of Belonocnema wasps Wasps in the genus Belonocnema alternate between sexual and asexual generations that form galls that develop on different plant tissues of their respective host plants to complete a yearly bivoltine life cycle (Lund et al. 1998 ; Zhang et al. 2021c ). In both B. fossoria and B. treatae , the sexual generation develops within multi-chambered galls, formed on oak rootlets, from which adults emerge in the spring. Upon emergence, individuals mate, and females immediately begin ovipositing into the lateral veins on the underside of newly flushed leaves, inducing single chambered leaf galls within which the asexual generation develops (Lund et al. 1998 ; Hood et al. 2019 ). In this study, we compared the asexual generation of B. fossoria and B. treatae following emergence from leaf galls in late fall. Belonocnema fossoria and B. treatae are specialized on Quercus geminata and Q. virginiana , respectively, which are two sister species of American live oaks (subsection Virentes ) that overlap broadly in their distribution along the Gulf and eastern coasts of the southeastern United States. Despite their overlap, each host-plant species occupies different microhabitats (Fig. 2 , Cavender-Bares and Pahlich 2009 ; Cavender‐Bares et al. 2015). The sand live oak, Q. geminata , occupies drier, sandy soils with higher pH compared to the southern live oak, Q. virginiana , which lives in more mesic soils. The two host plants also differ in aspects of leaf morphology, physiology, phenology, and canopy height, with adult Q. virginiana (mean height = ~ 21 m) being more than twice as tall as mature Q. geminata (mean height = ~ 10 m) (Cavender‐Bares and Pahlich 2009). Sample collection To examine morphological and life history traits among asexual Belonocnema species, leaf galls containing late pupal stage asexual Belonocnema were collected from multiple locations for both species throughout the southeastern USA in October and November in the fall of 2017 and 2018 (Fig. 3 , Table S1 ). Galls produced by each species collected from each site were placed inside one liter glass jars covered with a coffee filter secured by a rubber band and housed in a greenhouse at Rice University (Houston, Texas) under ambient conditions (~ 23°C). The jars were monitored once every two days, and newly emerged adults were collected and preserved in 95% ethanol. To measure life history traits, we randomly sampled up to 30 individuals at four B. fossoria sites and five B. treatae sites. For the sites with fewer than 30 wasps, we measured all available individuals yielding a total of 105 B. fossoria and 93 B. treatae (see Table S2 for additional details). Measurements of morphological and life history traits For each individual wasp, we measured right hind tibia length, right forewing length, and the length, width, and height of the abdomen. In Hymenoptera, the length of the right hind tibia has previously been shown to be correlated with overall body size (Rogers et al. 1976 ; Honěk 1993 ; Hood et al. 2011, 2017). To measure tibias and forewings accurately, they were removed from the body and slide mounted prior to measuring. After removal, the size of the abdomen was then estimated by measuring length (the longest distance between anterior and posterior ends), width (widest point perpendicular to length), and height (tallest point when viewed from the side, perpendicular to length). Abdomen volume (AV) per individual was then estimated by assuming that abdomen shape was best described as an ellipsoid using the equation AV = 4/3*π*(1/2*L)*(1/2*W)*(1/2*H), where L = length, W = width, and H = height. After its size was assessed, the abdomen was dissected following the methods of Hood and Ott ( 2011 , 2017 ) to measure two direct components of potential reproductive effort: egg number and egg size. Species of Belonocnema are pro-ovigenic, where the ovaries contained within the abdomen house the full complement of eggs at the time of adult emergence (Hood and Ott 2010 ). Hence, a count of the number of eggs per female provides an estimate of potential maximum lifetime fecundity while measurements of individual egg size provide an estimate of reproductive investment per egg (Hood and Ott 2011 ). The product of total egg number × mean egg size provides an estimate of potential ‘total reproductive effort’, which we estimated for each female. We measured the size of the abdomen for two reasons. First, we tested whether abdomen volume is predictive of total reproductive effort since the abdomen contains the ovaries, which are filled with mature eggs, and abdomen size correlates with egg number in many insect species (Honěk 1993 ). Second, changes in abdomen size (and shape) may relate to how observed differences in reproductive effort between the species are achieved. All traits described above were digitally measured to the nearest 0.01 mm at 20x magnification using a LEICA M125 microscope. To count the number of eggs and measure egg size, the abdomen was removed and placed in a depression slide containing a 1:1:13 acetic acid, glycerol, and water solution for at least 20 minutes to loosen ovarian follicular tissue binding eggs to the ovaries and abdomen wall (Hood and Ott 2011 ). At this time, the exoskeleton was removed, then the abdomen was dyed with a single drop of 1% methylene blue to better visualize the eggs. Potential fecundity was determined by counting the total number of eggs contained in the abdomen of each female. Mean egg size per female was estimated by randomly selecting five eggs from each female, and then measuring the length (the longest axis) and width (the widest point perpendicular to the length) following the methods of Hood and Ott ( 2011 ). Both measurements were made using a LEICA M125 microscope at 100x magnification. Egg volume (EV) was estimated using the formula for a prolate spheroid following Hood and Ott ( 2011 , 2017 ): EV = 4/3*π*(1/2*L)*(1/2*W) 2 , where L = length and W = width. Total reproductive effort per female was then estimated by multiplying the estimated average egg volume by the observed total number of eggs for each female (Hood and Ott 2011 ). Test of divergence of morphological and life history traits To compare differences in body size, wing length, number of eggs, egg size, abdomen volume, and total reproductive effort between B. fossoria and B. treatae , we performed linear mixed model analysis with each of these six traits as the response variables, ‘wasp species’ as a fixed factor, and ‘collection site’ as a random factor. This and all other analyses were conducted in R version 4.0.2 (R Core Team 2020). Test of independent adjustment of life history traits To examine if and to what extent trait divergence is dependent on body size, we assessed the relationships between (a) body size and wing length, (b) body size and abdomen volume, (c) body size and number of eggs, (d) body size and egg size, and (e) body size and total reproductive effort using the comparative framework outlined by Hood and Ott ( 2017 ) see modified version in Fig. 1 . This modified framework outlines four scenarios for how differences in body size and life history traits between species can be achieved by comparing changes in the species-specific slope and/or intercept of the linear relationship between the traits (see Fig. 1 for details). To statistically test whether and how these linear relationships (slope and/or intercept) between life history traits and body size differ between the two wasp species (Fig. 1 ), linear mixed model analysis was applied with each of the five traits listed above as the response variable, and ‘body size’, ‘wasp species’, and the interaction term ‘body size × wasp species’ as fixed factors, and collection site as the random factor. To reduce multicollinearity among interaction terms, all response variables and body size were standardized using the function scale in R. Divergence in linear relationship of response variables with body size between species is indicated by a significant wasp species term (i.e., different intercepts) and/or a significant interaction term between body size and wasp species (i.e., different slopes) in the model. Test of environmental effects of morphological and life history traits To examine whether phenotypic difference between species has a genetic basis or is a product of the rearing host plant environment (i.e., plasticity), we conducted a reciprocal transplant experiment. Details of the reciprocal transplant experiment are given in Hood et al. ( 2019 ) and Zhang et al. ( 2021a ), respectively. Briefly, in the spring of 2017, newly emerged and mated sexual generation females of each Belonocnema species were separately bagged onto the newly flushed leaves of four- to five-year-old saplings (~ 1.5 m in height) of each oak species to create four treatments: asexual generation B. fossoria developing on its natal ( Q. geminata ) or non-natal ( Q. virginiana ) host plant, and asexual generation B. treatae developing on its natal ( Q. virginiana) or non-natal ( Q. geminata ) host plant. Following oviposition, saplings with wasp eggs deposited within their leaves were transferred to a greenhouse at Rice University held at ambient conditions (~ 23°C), and watered once every other day until gall formers matured in fall 2017. At this time, galls containing Belonocnema from each replicate of each treatment were transferred to rearing containers and monitored for emergence as described above. In total, 70 Q. virginiana saplings and 66 Q. geminata saplings were used to rear asexual generation adults from the reciprocal transplant experiment. We measured body size (right hind tibia length), wing length, and abdomen size as a proxy of reproductive effort for wasps reared from each of the four experimental treatments using the methods described above (see Table S2 for a summary of the number of asexual females measured per treatment). Based on the patterns in our data, the length of the abdomen is a sufficient estimator of the total predicted reproductive effort per female (correlation coefficient r = 0.84, see Fig. S1 ). To determine whether the relationship between body size and each of the two morphological traits, wing length (i.e., a proxy for dispersal ability) and abdomen length (i.e., a proxy for reproductive effort), differed between wasp species reared on natal versus non-natal host plants, we conducted a linear mixed model with each life history trait as the response variable, ‘body size’, ‘wasp species’, ‘rearing host ’ (natal or non-natal host plant), and interaction among these three factors as fixed factors, and ‘collection site’ treated as a random factor. In each mixed model analysis, we used the function lme in R package “ nlme ” and calculated the least square means of each of the phenotypic traits using function emmeans . Test of within-species trade-offs between dispersal and reproduction We tested the relationship between the following traits: wing size, abdomen volume, egg size, egg number, and reproductive effort. Many of these traits covary with body size. To remove the effect of body size variation masking the trade-offs between wing size and reproductive traits within species, we first calculated the residuals of the correlation between wing size against body size. Then we assessed the linear correlations between the residuals of wing size against body size with reproductive traits including abdomen volume, egg number, egg size, and total reproductive effort. If intraspecific trade-offs exist, a significant negative correlation coefficient of these linear relationships is expected. These analyses were conducted for B. treatae and B. fossoria datasets separately. Results Divergence of morphological and life history traits between species On average, B. fossoria and B. treatae do not differ in body size as indexed by hind tibia length (least square mean ± SE: B. fossoria = 1.09 ± 0.031 mm, B. treatae = 1.11 ± 0.031 mm, t = 0.569, P = 0.587, Fig. 4 A). However, B. fossoria had significantly shorter wings and larger abdomens than B. treatae (wing length: B. fossoria = 2.21 ± 0.082 mm, B. treatae = 2.94 ± 0.083 mm, t = -6.240, P < 0.001, Fig. 4 B; abdomen volume: B. fossoria = 1.391 ± 0.156 mm 3 , B. treatae = 0.745 ± 0.162 mm 3 , t = 2.873, P = 0.028, Fig. 4 C). While B. fossoria tended to produce more eggs per individual than B. treatae , the difference was not significant ( B. fossoria = 244 ± 17.1, B. treatae = 199 ± 17.6, t = -1.826, P = 0.111; Fig. 4 D). However, B. fossoria did allocate significantly more resources towards reproduction than B. treatae (Fig. 4 E-F) based on individual egg volume ( B. fossoria = 6.6 ± 0.179 x 10 − 4 mm 3 , B. treatae = 5.5 ± 1.89 x 10 − 4 mm 3 , t = -4.148, P = 0.004) and total reproductive effort ( B. fossoria = 0.162 ± 0.010 mm 3 , B. treatae = 0.11 ± 0.01 mm 3 , t = -3.643, P = 0.008). Divergence in relationship between life history traits and body size within species The association between body size and wing length for B. fossoria and B. treatae was consistent with the second scenario in Fig. 1 where wing length increased with body size in each species ( t = 5.43, P < 0.001), but only the y-intercept differed significantly between the two species ( t = 12.46, P < 0.001; Table 1 ; Fig. 5 a). For each reproductive trait, the linear relationships with body size followed the fourth scenario in Fig. 1 where abdomen volume ( t = 9.12, P < 0.001), potential fecundity ( t = 11.85, P < 0.001), egg volume ( t = 3.59, P < 0.001) and total reproductive effort ( t = 11.76, P < 0.001) increased with increasing body size (Fig. 5 B-E). The y-intercept differed significantly between the two wasp species (abdomen volume: t = 4.83, P = 0.020; potential fecundity: t = 2.55, P = 0.038; egg volume: t = 4.37, P = 0.003; total reproductive effort: t = 4.85, P = 0.002; Table 1 ). Similarly, the slope of the linear relationship in each comparison differed between species (abdomen volume: t = 1.957, P = 0.052; potential fecundity: t = 2.959, P = 0.004; egg volume: t = 2.36; P = 0.020; predicted reproductive effort: t = 4.31, P < 0.001; Table 1 ). Table 1 Linear mixed model analysis of the relationship between body size and five life history traits: wing length, abdomen volume, potential fecundity, egg volume and reproductive effort (the product of egg number and mean egg volume per female) for B. treatae and B. fossoria . Responsible variable Factors Estimate SE t P Wing length (mm) Body size 0.31 0.06 5.43 < 0.001 Species 1.4 0.11 12.46 < 0.001 Body size × Species 0.06 0.09 0.63 0.532 Abdomen volume (mm 3 ) Body size 0.62 0.06 9.49 < 0.001 Species -1.22 0.24 -5.03 0.002 Body size × Species -0.42 0.09 -4.61 < 0.001 Potential fecundity Body size 0.73 0.06 11.85 < 0.001 Species -0.66 0.26 -2.55 0.038 Body size × Species -0.32 0.11 -2.96 0.004 Egg volume (mm 3 ) Body size 0.29 0.08 3.59 < 0.001 Species -0.94 0.22 -4.37 0.003 Body size × Species -0.33 0.14 -2.36 0.020 Reproductive effort Body size 0.72 0.06 11.76 < 0.001 Species -0.9 0.18 -4.85 0.002 Body size × Species -0.46 0.11 -4.31 < 0.001 * P -values in bold: P < 0.05. Test for environmental effects on morphological and life history traits due to different host plants The reciprocal transplant experiments demonstrated that the observed differences between species in the linear relationship between (a) body size and wing length (Fig. 6 A) and (b) body size and abdomen length (Fig. 6 B) did not change when wasps are reared in their non-natal host environment. The y-intercept of the linear relationship between wing length and body size differed significantly between B. treatae and B. fossoria ( t = 9.50, P < 0.001; Table 2 ), but not between alternative rearing hosts ( t = 0. 820, P = 0.414; Table 2 ). Similarly, the slope did not change between different rearing hosts ( t = -0.19, P = 0.846; Table 2 ). For the linear relationship between abdomen length and body size, the y-intercept differed significantly between the two wasp species ( t = -2.91, P = 0.044; Table 2 ), but not between alternative rearing host environments ( t = 0.99, P = 0.325; Table 2 ). Similarly, the slope did not change between different rearing hosts (body size ´ rearing host: t = -1.43, P = 0.156; Table 2 ). Table 2 Linear mixed model analysis of the reciprocal transplant experiment testing the potential effect of host plant rearing environment on abdomen size (length) and forewing length of B. treatae (N = 105) and B. fossoria (N = 93). Responsible variable Factors Estimate SE df t P Abdomen size Body size 0.74 0.08 126 9.86 < 0.001 Wasp species -0.82 0.28 4 -2.91 0.044 Rearing host 0.13 0.13 126 0.99 0.325 Body size × Wasp species -0.16 0.11 126 -1.43 0.156 Body size × Rearing host 0.26 0.12 126 2.21 0.029 Wasp species × Rearing host 0.14 0.24 126 0.57 0.568 Body size × Wasp species × Rearing host -0.32 0.21 126 -1.51 0.132 Wing length Body size 0.71 0.07 120 10.47 < 0.001 Wasp species 1.25 0.13 4 9.5 0.001 Rearing host 0.09 0.1 120 0.82 0.414 Body size × Wasp species -0.12 0.09 120 -1.31 0.193 Body size × Rearing host -0.02 0.1 120 -0.19 0.846 Wasp species × Rearing host 0.3 0.19 120 1.57 0.119 Body size × Wasp species × Rearing host 0.13 0.18 120 0.73 0.466 * P -values in bold: P < 0.05. Test of within-species trade-off between wing length and reproductive effort After controlling for the effect of body size, we found no evidence of trade-offs between wing length and reproductive effort within B. fossoria or B. treatae (Fig. 7 ). This conclusion is supported by a lack of significant correlations between wing length and abdomen volume, egg number, egg size, and total reproductive effort in B. fossoria and in B. treatae summarized in Table 3 . Table 3 Intraspecific correlations between wing size (associated with dispersal capacity) and four reproductive traits (abdomen volume, egg number, egg volume, and reproductive effort) after correcting for body size for B. fossoria and B. treatae. B. fossoria B. treatae r P r P Abdomen volume 0.06 0.627 0.05 0.612 Egg number -0.05 0.679 0.07 0.495 Egg volume -0.02 0.859 -0.147 0.202 Reproductive effort -0.13 0.344 -0.05 0.689 Discussion Trade-offs among life history traits are of key interest in evolutionary biology due to their potential impacts on maintaining phenotypic variation or constraining adaptation under changing environments (Reznick and Bryga 1987 ; Futuyma and Moreno 1988 ). On one hand, trade-offs may act as evolutionary constraints for adaptation among individuals; on the other hand, long-term selection may overcome such constraints and reshape trait combinations (Agrawal 2020 ). Therefore, investigating whether trade-offs scale across different levels of biological organization, from individuals to populations and species, will shed light on the evolutionary processes underlying life history traits (Bolnick et al. 2003 ; Agrawal 2020 ). For example, if trait divergence between species originates from trade-offs at the intraspecific level, we would expect to find trade-offs among traits both within and between species. Alternatively, if recently diverged species evolve independently in response to divergent environmental conditions, trade-offs might only be detected at the interspecific level due to environmental constraints while not being evident at the intraspecific level. Our study addresses this potential connection between inter- and intraspecific variation by testing whether there is a trade-off between dispersal capacity and reproductive output in two sister species of gall wasps belonging to the genus Belonocnema , which exhibit different dispersal capabilities. We also examined whether such trade-offs exist within each monomorphic species. We discovered that trade-offs between flight capability and reproductive output are evident as a consistent difference between the two wasp species, but no trade-offs were detected at the intraspecific level. Specifically, B. fossoria exhibits reduced flight capability and smaller wing size, while demonstrating significantly higher reproductive effort compared to B. treatae (Fig. 4 ). Our finding of an interspecific trade-off, but the absence of intraspecific trade-offs, aligns with previous studies exploring trade-offs between life history traits such as flower size and flower number (Worley et al. 2000 ; Sargent et al. 2007 ; Hahn and Maron 2016 ). One possible explanation for the lack of intraspecific trade-offs in these studies is that individual variation in resource acquisition masks the correlation among traits. Individuals with greater resource availability may develop both larger and more numerous flowers, whereas individuals with limited resources but larger flowers would have fewer flowers (van Noordwijk et al. 1986; King et al. 2011 ; Agrawal 2020 ). However, in the case of our wasp species, individual resource acquisition variation is unlikely to explain the absence of trade-offs between flight-related traits and reproductive output. This is because our analysis already partially controls for the effect of body size, which indirectly accounts for individual resource variation (Fig. 7 ). An alternative explanation is that these two wasp species have recently diverged and specialized within a narrow niche (only one or two host plant species), resulting in long-term and strong selection that has shaped divergence in resource allocation between the species. Within each species, considering the narrow niche, further specialization along these life history dimensions may not be expected, thereby leading to the absence of trade-offs among life history traits (Futuyma and Moreno 1988 ). We posit that the discrepancy in trade-off patterns between intra- and interspecific levels is more likely to occur among closely related species inhabiting different environments compared to species inhabiting similar environments. This is based on the common assumption that phylogenetically closely related species tend to share similar life history traits and evolutionary constraints, resulting in a consistent trade-off pattern within and among sister species (Webb et al. 2002 ). However, if a particular life history trait plays a crucial role in adaptation to divergent environments, long-term and intense selection pressure could disrupt the genetic constraints on the evolution of life history traits (Agrawal 2020 ). Further studies employing a comparative framework similar to this study are needed to systematically test this hypothesis. For the two wasp species studied here, B. fossoria and B. treatae , the divergence in life history traits is not solely due to changes in body size but also involves alterations in the relationship between body size and multiple traits, including wing size, abdomen size, and total reproductive effort (Fig. 5 ). This means that the adjustments in flight capability and reproductive effort, as depicted by Scenario 4 in Fig. 1 , are independent of body size. Specifically, the independent adjustment of flight capability between the sister species is reflected in variation in wing size, while changes in reproductive effort are achieved through modifications in abdomen size. The independent adjustment of these life history traits is further supported by a common garden rearing experiment: the divergence in the linear relationship between body size and wing size, as well as between body size and reproductive effort, is maintained even when the wasps are reared on a non-native host (Fig. 6 ). Therefore, the divergence in life history traits is likely attributable, at least in part, to genetic differences between the two wasp species rather than being solely a plastic response to the rearing host plants (i.e., environmental effects and phenotypic plasticity). Interestingly, the linear relationship between body size and wing size differs in intercepts, while the linear relationship between body size and other reproductive-related traits, including abdomen volume, potential fecundity, total reproductive effort, differs in both intercepts and slopes (Table 1 , Fig. 5 ). According to quantitative genetic theory, linear relationships between traits are more prone to change in intercepts under selection, while stronger and longer-term selection is required for changes in slopes (Roff et al. 2002 ). Thus, it is possible that reproductive traits experience stronger selection pressure than flight-related traits. Notably, similar changes in both intercepts and slopes between potential fecundity and body size have been observed in the alternating asexual and sexual generations of another species in the genus, B. kinsey (Hood and Ott 2017 ), where each generation faces unique environmental challenges. Hence, both Hood and Ott ( 2017 ) and the present study provide evidence that reproductive traits are highly adaptable phenotype under selection in this group of Belonocnema gall wasps. Since dispersal and reproduction often exhibit trade-offs either within species or among species across many study systems (Carroll et al. 2003 ; Guerra 2011 ; King et al. 2011 ; Nasu and Tokuda 2021 ), more studies should link changes in linear relationship among these life history traits to the variation of selection pressures each of these traits experience (e.g., populations at the range edge). We argue that B. fossoria individuals with higher reproductive output but lower flight capability may gain a fitness advantage in the Q. geminata environment, leading to selection favoring the divergent phenotypes between B. fossoria and B. treatae. This is because B. fossoria 's host plant, Q. geminata , is typically found as short shrubs no taller than 10 m in small clusters in sandy soil, whereas B. treatae 's host plant, Q. virginiana , is a much taller tree species (with a mean height of ~ 21m) and has a patchier distribution (Cavender-Bares and Pahlich 2009 ). Dense, short trees may require less flight capability for the asexual generation of B. fossoria to locate suitable sites for gall induction on the roots, which host the next generation (Zera and Denno 1997 ). The reduced flight capability observed in B. fossoria , is likely a product of relaxed selection on flight associated with more persistent, shorter, and smaller host trees, as suggested by other studies highlighting the role of habitat structure in the evolution of flight capability (Roff 1990 ; Roff 1994; Denno et al. 1996 ; Zera and Denno 1997 ; Dell'Aglio et al. 2022). Additionally, a third allopatric species in this genus, B. kinseyi , which shares host plant Q. virginina with B. treatae , exhibits life history traits similar in size to the B. treatae populations analyzed here. These include similar flight capability (Zhang et al. 2021c ) and correlations between body size and wing length (wasp species: t = -0.03, p = 0.981; body size x wasp species: t = -1.04, p = 0.303) and between body size and abdomen length (wasp species: t = -2.88, p = 0.103; body size x wasp species: t = 0.56, p = 0.581; see Supplement). Therefore, the divergent phenotypes between B. treatae and B. fossoria are likely the result of adaptation to divergent host plant-related environments. Future studies should apply a similar comparative work to other gall wasp species that also utilized the same set of host plant species Q. virginiana and Q. geminata . If similar divergent phenotypes are observed repeatedly between host-associated populations in other gall wasp species, this would strongly suggest the similar host plant-related environments play a critical role in shaping the divergence among these traits (Egan et al. 2013 ; Zhang et al. 2019 , 2022 ). Declarations Acknowledgements We thank Elaine Hu, Isaac Carroo and Hannah Towbin for assistance with sample collection and morphological measurements. Funding was provided to AMR by the Spurlino Summer Undergraduate Research Fellowship, to LZ by student research award from Society for Integrative and Comparative Biology, Rosemary Grant Award from the Society for the Study of Evolution, and the Diana McSherry and Patrick Poe research award, to SPE from the Department of BioSciences at Rice University and to GRH from the Rice University Academy of Fellows. Author Contributions: LZ, SPE, GRH, and JRO conceived and designed the experiments. AMR performed the experiments. AMR and LZ analyzed the data. AMR and LZ wrote the manuscript; other authors edited the manuscript. References Agrawal AA (2020) A scale-dependent framework for trade-offs, syndromes, and specialization in organismal biology. 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Zool J Linn Soc. 10.1093/zoolinnean/zlab001 Supplementary Files 2023OecologiaSupplementalmaterial.docx B.treataereciprocaltransplantmorphologydata.csv Reproductive.Strategy.Data.csv Cite Share Download PDF Status: Published Journal Publication published 07 Feb, 2024 Read the published version in Oecologia → Version 1 posted Reviewers agreed at journal 02 Aug, 2023 Reviewers invited by journal 31 Jul, 2023 Editor assigned by journal 17 Jul, 2023 First submitted to journal 15 Jul, 2023 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-3171363\",\"acceptedTermsAndConditions\":true,\"allowDirectSubmit\":false,\"archivedVersions\":[],\"articleType\":\"Research Article\",\"associatedPublications\":[],\"authors\":[{\"id\":222899530,\"identity\":\"141ebdfc-dc36-4c89-b3c5-1d27c213f5fa\",\"order_by\":0,\"name\":\"Amy M. Roush\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"Rice University\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Amy\",\"middleName\":\"M.\",\"lastName\":\"Roush\",\"suffix\":\"\"},{\"id\":222899531,\"identity\":\"04664aff-ca33-4e23-8c22-677c42291a7f\",\"order_by\":1,\"name\":\"Linyi Zhang\",\"email\":\"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA20lEQVRIiWNgGAWjYBACPmYgkQBGDIwPIGIJ+LWwIWlhNiBOCwNCGZsEcVrYeY9JPGCoy+Of3X6t8mvbYQZ+9hwDAg7jS5NIYGArlrhzpuy2LFCLZM8bQlp4zIBaeBIbbuSk3ZbcdpjB4AZBW8BaJBLnA7UUg7TYE6nFIHHDjfRjjB9BtkgQ1mJskWCQkLjxRg6zNOO/dB6JM88K8Grh5z9jePNHRV3ivBvpDz/+OGMtx9+evAGvFiBgkWAAu4THgJkHSBJSDgLMHyA0+wPGH8SoHwWjYBSMghEHAKjwQOt56s0aAAAAAElFTkSuQmCC\",\"orcid\":\"https://orcid.org/0000-0001-9401-0282\",\"institution\":\"The George Washington University Columbian College of Arts and Sciences\",\"correspondingAuthor\":true,\"prefix\":\"\",\"firstName\":\"Linyi\",\"middleName\":\"\",\"lastName\":\"Zhang\",\"suffix\":\"\"},{\"id\":222899532,\"identity\":\"6ec349ae-b60c-4ece-bfa1-e88570082192\",\"order_by\":2,\"name\":\"Glen Ray Hood\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"Wayne State University\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Glen\",\"middleName\":\"Ray\",\"lastName\":\"Hood\",\"suffix\":\"\"},{\"id\":222899533,\"identity\":\"76bf9db7-b036-4af1-885e-e4f9fa7a287f\",\"order_by\":3,\"name\":\"James R. Ott\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"Texas State University\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"James\",\"middleName\":\"R.\",\"lastName\":\"Ott\",\"suffix\":\"\"},{\"id\":222899534,\"identity\":\"0931b3e1-dff6-42a1-8b82-3188eb7a939d\",\"order_by\":4,\"name\":\"Scott P. Egan\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"Rice University\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Scott\",\"middleName\":\"P.\",\"lastName\":\"Egan\",\"suffix\":\"\"}],\"badges\":[],\"createdAt\":\"2023-07-14 18:11:46\",\"currentVersionCode\":1,\"declarations\":\"\",\"doi\":\"10.21203/rs.3.rs-3171363/v1\",\"doiUrl\":\"https://doi.org/10.21203/rs.3.rs-3171363/v1\",\"draftVersion\":[],\"editorialEvents\":[{\"content\":\"https://doi.org/10.1007/s00442-024-05512-3\",\"type\":\"published\",\"date\":\"2024-02-07T15:01:35+00:00\"}],\"editorialNote\":\"\",\"failedWorkflow\":false,\"files\":[{\"id\":41097598,\"identity\":\"af34fa6f-b44c-426b-93b7-e18b5420bd34\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:34:42\",\"extension\":\"png\",\"order_by\":1,\"title\":\"Figure 1\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":29545,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eFour scenarios for how differences in body size and life history traits between species arise [modified from Hood and Ott (2017)]. \\u003cem\\u003eScenario 1:\\u003c/em\\u003e Linear relationship between body size and life history traits (e.g., fecundity) do not differ between two species. Here differences in trait value result from changes in body size between species. \\u003cem\\u003eScenario 2:\\u003c/em\\u003eDifferences in life history trait values occur due to a difference in the y-intercepts of the same linear relationship between body size and life history traits. \\u003cem\\u003eScenario 3:\\u003c/em\\u003e Differences in life history trait values are due to a difference in the slopes of the linear relationship between body size and life history traits with no change in average body size between species. In this scenario, the y-intercepts of the relationship between the life history trait value and body size do not differ between species. \\u003cem\\u003eScenario 4:\\u003c/em\\u003eDifferences in life history trait values are due to changes in both the y-intercept and slopes of the linear relationship between body size and life history traits (i.e., independent adjustments within each species). Black line represents species 1 (S1); gray line represents species 2 (S2).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Onlinefloatimage1.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/1a47180a5a868247467423cb.png\"},{\"id\":41097600,\"identity\":\"6c5a1f46-281d-40ae-8eaa-3dafd6132315\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:34:43\",\"extension\":\"png\",\"order_by\":2,\"title\":\"Figure 2\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":1045816,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003e(A) Large southern live oak tree, \\u003cem\\u003eQuercus virginiana,\\u003c/em\\u003e including\\u003cem\\u003e \\u003c/em\\u003e(B) branches and leaves galled by \\u003cem\\u003eBelonocnema treatae\\u003c/em\\u003e contrasted with (C) the smaller, shrubbier sand live oak, \\u003cem\\u003eQ. geminate\\u003c/em\\u003e,\\u003cem\\u003e \\u003c/em\\u003eincluding (D) branches and leaves galled by \\u003cem\\u003eB. fossoria\\u003c/em\\u003e. Asexual generation adult (E) \\u003cem\\u003eB. treatae\\u003c/em\\u003e and (F) \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and (G) eggs of asexual generation \\u003cem\\u003eBelonocnema \\u003c/em\\u003eat 100´ magnification.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Onlinefloatimage2.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/194f6f275be5a0090b0c1488.png\"},{\"id\":41097599,\"identity\":\"8b5d9863-7355-4bb2-8192-bbef9659ab59\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:34:43\",\"extension\":\"png\",\"order_by\":3,\"title\":\"Figure 3\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":71263,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eCollection sites in Florida, USA, for \\u003cem\\u003eB. fossoria\\u003c/em\\u003e (black circles) and \\u003cem\\u003eB. treatae\\u003c/em\\u003e (open circles). See Table S1 for location abbreviations.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Onlinefloatimage3.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/1915b50e5afec0a7ae0c0040.png\"},{\"id\":41099762,\"identity\":\"ef25ff8d-17d7-4576-a186-7ed058198bea\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:50:43\",\"extension\":\"png\",\"order_by\":4,\"title\":\"Figure 4\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":470786,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eBoxplots comparing life history traits of \\u003cem\\u003eB. fossoria\\u003c/em\\u003e sampled from \\u003cem\\u003eQ. geminata\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003esampled from \\u003cem\\u003eQ. virginiana\\u003c/em\\u003e: (A) body size (tibia length), (B) forewing length, (C) abdomen volume, (D) potential fecundity (number of eggs), (E) individual egg size (volume), and (F) total reproductive effort (calculated as the produce of potential fecundity and egg size). The full distribution of trait values for both \\u003cem\\u003eB. fossoria\\u003c/em\\u003e (dark grey) and \\u003cem\\u003eB. treatae\\u003c/em\\u003e(light grey) are displayed on the right side of each panel. (* \\u003cem\\u003eP\\u003c/em\\u003e \\u0026lt; 0.05; ** \\u003cem\\u003eP\\u003c/em\\u003e \\u0026lt; 0.01, *** \\u003cem\\u003eP \\u003c/em\\u003e\\u0026lt; 0.001)\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Onlinefloatimage4.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/d9beedf02fce7ae8048b100c.png\"},{\"id\":41099429,\"identity\":\"12845a95-b337-4598-b31a-b103b35705df\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:42:43\",\"extension\":\"png\",\"order_by\":5,\"title\":\"Figure 5\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":707335,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eLinear relationships between body size (tibia length) and (A) wing length, (B) abdomen volume, (C)potential fecundity, (D) egg volume, and (E) reproductive effort for both \\u003cem\\u003eB. fossoria\\u003c/em\\u003e (solid lines, filled circles) and \\u003cem\\u003eB. treatae\\u003c/em\\u003e (dashed lines, open circles).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Onlinefloatimage5.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/415f6a9a66f2e91f948495f5.png\"},{\"id\":41099426,\"identity\":\"9a0ef10f-08ca-46e0-90d7-ebad74ab68c6\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:42:43\",\"extension\":\"png\",\"order_by\":6,\"title\":\"Figure 6\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":361381,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eLinear relationships between (A) body size and wing length and (B) body size and abdomen length for \\u003cem\\u003eB. fossoria\\u003c/em\\u003e (Bf; solid lines) and \\u003cem\\u003eB. treatae\\u003c/em\\u003e (Bt; dashed lines) reared on both native (circles) and non-native (squares) host plants.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Onlinefloatimage6.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/a7c497dc3dc6214d4cb79b1f.png\"},{\"id\":41097601,\"identity\":\"3000b1df-cdd0-446b-97cb-cc6c496ed6da\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:34:43\",\"extension\":\"png\",\"order_by\":7,\"title\":\"Figure 7\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":393595,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eScatterplot between wing length after controlling for the effect of body size and (A) abdomen volume, (B) egg number, (C) egg volume, and (D) total reproductive effort for both \\u003cem\\u003eB. fossoria\\u003c/em\\u003e(filled circles) and \\u003cem\\u003eB. treatae\\u003c/em\\u003e (open circles).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Onlinefloatimage7.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/d21ba48eb835b3ad7c8ab121.png\"},{\"id\":51005728,\"identity\":\"7659eb0a-9ab5-4b93-9bb9-43ef7c2d728f\",\"added_by\":\"auto\",\"created_at\":\"2024-02-12 15:12:03\",\"extension\":\"pdf\",\"order_by\":0,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"manuscript-pdf\",\"size\":2626162,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"manuscript.pdf\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/b0a0d9d9-4ee2-48f3-b800-26f654560e9c.pdf\"},{\"id\":41099761,\"identity\":\"a4a2dbff-d259-4201-a260-b190d0e6a3e5\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:50:43\",\"extension\":\"docx\",\"order_by\":1,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"supplement\",\"size\":134841,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"2023OecologiaSupplementalmaterial.docx\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/62ef97ef2e6b4f9ab7e5d7bd.docx\"},{\"id\":41097597,\"identity\":\"8915c1b3-0506-4430-956d-a74588e0f45a\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:34:42\",\"extension\":\"csv\",\"order_by\":2,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"supplement\",\"size\":12005,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"B.treataereciprocaltransplantmorphologydata.csv\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/ccb6de6a8217043a7c1c70f8.csv\"},{\"id\":41097606,\"identity\":\"c4fbdcc8-eae1-4ead-9cd5-5fbc10d01fac\",\"added_by\":\"auto\",\"created_at\":\"2023-08-04 23:34:43\",\"extension\":\"csv\",\"order_by\":3,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"supplement\",\"size\":29363,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"Reproductive.Strategy.Data.csv\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3171363/v1/9549d5d854f21d0b455432f4.csv\"}],\"financialInterests\":\"\",\"formattedTitle\":\"A test of trade-offs in dispersal and reproduction within and between a sister species pair of specialist insect herbivores\",\"fulltext\":[{\"header\":\"Introduction\",\"content\":\"\\u003cp\\u003eTrade-offs in life history traits are commonly observed among organisms adapting to different environmental conditions (Roff and Fairbairn 2007). Classic examples include trade-offs between growth rate and reproduction (e.g., Stearns 1989), survival and reproduction (e.g., Partridge \\u0026amp; Farquhar 1981), and dispersal and reproduction (e.g., Zera and Denno \\u003cspan citationid=\\\"CR43\\\" class=\\\"CitationRef\\\"\\u003e1997\\u003c/span\\u003e, Carroll et al. \\u003cspan citationid=\\\"CR4\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e). One trait that is critical in mediating local adaptation to variable environments and regulating population connectivity is dispersal capacity (Stevens et al. \\u003cspan citationid=\\\"CR36\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e). For example, insect populations with higher dispersal ability are often associated with taller plants or ephemeral resources, as flight ability allows insects to better navigate complex habitats and/or access additional resources (Denno et al. \\u003cspan citationid=\\\"CR9\\\" class=\\\"CitationRef\\\"\\u003e1996\\u003c/span\\u003e; Zera and Denno \\u003cspan citationid=\\\"CR43\\\" class=\\\"CitationRef\\\"\\u003e1997\\u003c/span\\u003e; Duthie et al. \\u003cspan citationid=\\\"CR11\\\" class=\\\"CitationRef\\\"\\u003e2015\\u003c/span\\u003e; Dell\\u0026rsquo;Aglio et al. \\u003cspan citationid=\\\"CR8\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). Thus, dispersal capacity ultimately can influence the dynamics of population divergence and speciation (Claramunt et al. 2012; Stevens et al. \\u003cspan citationid=\\\"CR36\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e). However, the evolution of increased dispersal capacity is often accompanied by a trade-off in reproductive output due to the constraint of limited energy resources (Zera and Denno \\u003cspan citationid=\\\"CR43\\\" class=\\\"CitationRef\\\"\\u003e1997\\u003c/span\\u003e; Zera and Harshman \\u003cspan citationid=\\\"CR44\\\" class=\\\"CitationRef\\\"\\u003e2001\\u003c/span\\u003e; Ghalambor et al. \\u003cspan citationid=\\\"CR16\\\" class=\\\"CitationRef\\\"\\u003e2004\\u003c/span\\u003e; Vorburger \\u003cspan citationid=\\\"CR40\\\" class=\\\"CitationRef\\\"\\u003e2005\\u003c/span\\u003e). Such trade-offs between dispersal and reproduction can be critical in maintaining population sizes under fluctuating environments, promoting species coexistence, and/or affecting range expansion (Stevens et al. \\u003cspan citationid=\\\"CR36\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e; Duthie et al. \\u003cspan citationid=\\\"CR11\\\" class=\\\"CitationRef\\\"\\u003e2015\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003eTrade-offs between dispersal and reproduction are often found within wing-dimorphic insect species (Roff \\u003cspan citationid=\\\"CR31\\\" class=\\\"CitationRef\\\"\\u003e1986\\u003c/span\\u003e; Guerra \\u003cspan citationid=\\\"CR17\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e), where flight-capable, long-winged \\u0026lsquo;macropterous\\u0026rsquo; morphs and reduced-flight, short-winged \\u0026lsquo;brachypterous\\u0026rsquo; morphs are each present within a species and can be expressed as plastic responses to fluctuating environments (Zera and Rankin \\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e1989\\u003c/span\\u003e; Carroll et al. \\u003cspan citationid=\\\"CR4\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e; Nasu and Tokuda \\u003cspan citationid=\\\"CR27\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). However, divergence in trade-offs between dispersal and reproduction among closely related species within a genus are less prevalent (Tigreros and Davidowitz \\u003cspan citationid=\\\"CR38\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e). Investigating whether such interspecific trade-offs scale from intraspecific variation is particularly interesting with respect to understanding how divergence across a suite of traits between species can evolve (Worley et al. \\u003cspan citationid=\\\"CR42\\\" class=\\\"CitationRef\\\"\\u003e2000\\u003c/span\\u003e; Sargent et al. \\u003cspan citationid=\\\"CR35\\\" class=\\\"CitationRef\\\"\\u003e2007\\u003c/span\\u003e; Agrawal \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). For example, if trade-offs between traits are found at both the intra- and interspecific levels, trait divergence between species are likely constrained by trade-offs from intraspecific variation (Agrawal \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Alternatively, if interspecies differences in flight capacity and any associated trade-off(s) do not scale from intraspecific variation, this may reflect independent evolutionary trajectories between species in response to different environmental conditions. Another possibility is that the lack of trade-offs between life history traits within species is because resource acquisition varies greatly among individuals, thus difference in life history traits due to resource availability masks the evolutionary trade-offs among different traits (van Noordwijk et al. 1986). To date, however, few studies have examined trade-offs between dispersal and reproduction between closely related species, while simultaneously exploring whether evidence of trade-offs exist within each species (Agrawal \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003eSympatric sister species of gall wasps in the genus \\u003cem\\u003eBelonocnema\\u003c/em\\u003e (Hymenoptera: Cynipidae) provide an opportunity to test the hypothesis that trade-offs between dispersal and reproduction are similar at each level of biological orgainization. First, the two sister species, \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e, differ in flight capacity; asexual generation \\u003cem\\u003eB. fossoria\\u003c/em\\u003e, specialized to develop on shorter sand live oak, \\u003cem\\u003eQuercus geminata\\u003c/em\\u003e, are flightless, while asexual generation \\u003cem\\u003eB. treatae\\u003c/em\\u003e, specialized on taller southern live oak, \\u003cem\\u003eQ. virginiana\\u003c/em\\u003e, are capable of flight (Zhang et al. \\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e2021c\\u003c/span\\u003e). Second, \\u003cem\\u003eBelonocnema\\u003c/em\\u003e do not feed as adults, thus, the pool of resources available for allocation to dispersal (flight capacity) and reproduction (egg number and egg size) is finite, with those resources derived from stores acquired during the earlier larval feeding period (Lund et al. \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e1998\\u003c/span\\u003e; Zera and Harshman \\u003cspan citationid=\\\"CR44\\\" class=\\\"CitationRef\\\"\\u003e2001\\u003c/span\\u003e; Bonnet et al. \\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e2002\\u003c/span\\u003e). Third, the two species are sympatric in a large portion of their range, but occupy different host plants that represent different environmental challenges (Cavender-Bares and Pahlich \\u003cspan citationid=\\\"CR6\\\" class=\\\"CitationRef\\\"\\u003e2009\\u003c/span\\u003e; Cavender‐Bares et al. 2015). Finally, the species are near-completely reproductively isolated because of multiple pre- and postzygotic reproductive barriers (Egan et al. \\u003cspan citationid=\\\"CR12\\\" class=\\\"CitationRef\\\"\\u003e2012a\\u003c/span\\u003e,\\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003eb\\u003c/span\\u003e; Zhang et al. \\u003cspan citationid=\\\"CR46\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2021a\\u003c/span\\u003e,\\u003cspan citationid=\\\"CR49\\\" class=\\\"CitationRef\\\"\\u003eb\\u003c/span\\u003e; Hood et al. \\u003cspan citationid=\\\"CR23\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e) that result in sympatric host-associated populations that are genomically distinct (Driscoe et al. \\u003cspan citationid=\\\"CR10\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e; Zhang et al. \\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e2021c\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003eHere, we first test whether the difference in flight capacity of \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e is associated with between-species differences in wing length, body size and/or reproductive traits, as predicted by a dispersal/reproduction trade-off. Second, we test the nature of potential trade-offs in life history traits between species by extending the framework proposed by Hood and Ott (\\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e) and illustrated in Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e to test for independent adjustments of wing length and reproductive traits against body size between \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e. Third, we conduct a reciprocal transplant experiment to test whether such the observed trade-offs in life history traits between species are genetic or a plastic response to the different host plant rearing environments. Finally, we test whether the trade-off in dispersal and reproduction observed between species also exists within either ecologically divergent sister species.\\u003c/p\\u003e\"},{\"header\":\"Methods\",\"content\":\"\\u003cp\\u003e \\u003cb\\u003eBiology of\\u003c/b\\u003e \\u003cb\\u003eBelonocnema\\u003c/b\\u003e \\u003cb\\u003ewasps\\u003c/b\\u003e \\u003c/p\\u003e \\u003cp\\u003eWasps in the genus \\u003cem\\u003eBelonocnema\\u003c/em\\u003e alternate between sexual and asexual generations that form galls that develop on different plant tissues of their respective host plants to complete a yearly bivoltine life cycle (Lund et al. \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e1998\\u003c/span\\u003e; Zhang et al. \\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e2021c\\u003c/span\\u003e). In both \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e, the sexual generation develops within multi-chambered galls, formed on oak rootlets, from which adults emerge in the spring. Upon emergence, individuals mate, and females immediately begin ovipositing into the lateral veins on the underside of newly flushed leaves, inducing single chambered leaf galls within which the asexual generation develops (Lund et al. \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e1998\\u003c/span\\u003e; Hood et al. \\u003cspan citationid=\\\"CR23\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e). In this study, we compared the asexual generation of \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e following emergence from leaf galls in late fall.\\u003c/p\\u003e \\u003cp\\u003e \\u003cem\\u003eBelonocnema fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e are specialized on \\u003cem\\u003eQuercus geminata\\u003c/em\\u003e and \\u003cem\\u003eQ. virginiana\\u003c/em\\u003e, respectively, which are two sister species of American live oaks (subsection \\u003cem\\u003eVirentes\\u003c/em\\u003e) that overlap broadly in their distribution along the Gulf and eastern coasts of the southeastern United States. Despite their overlap, each host-plant species occupies different microhabitats (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e, Cavender-Bares and Pahlich \\u003cspan citationid=\\\"CR6\\\" class=\\\"CitationRef\\\"\\u003e2009\\u003c/span\\u003e; Cavender‐Bares et al. 2015). The sand live oak, \\u003cem\\u003eQ. geminata\\u003c/em\\u003e, occupies drier, sandy soils with higher pH compared to the southern live oak, \\u003cem\\u003eQ. virginiana\\u003c/em\\u003e, which lives in more mesic soils. The two host plants also differ in aspects of leaf morphology, physiology, phenology, and canopy height, with adult \\u003cem\\u003eQ. virginiana\\u003c/em\\u003e (mean height\\u0026thinsp;=\\u0026thinsp;~\\u0026thinsp;21 m) being more than twice as tall as mature \\u003cem\\u003eQ. geminata\\u003c/em\\u003e (mean height\\u0026thinsp;=\\u0026thinsp;~\\u0026thinsp;10 m) (Cavender‐Bares and Pahlich 2009).\\u003c/p\\u003e \\u003cdiv id=\\\"Sec3\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eSample collection\\u003c/h2\\u003e \\u003cp\\u003eTo examine morphological and life history traits among asexual \\u003cem\\u003eBelonocnema\\u003c/em\\u003e species, leaf galls containing late pupal stage asexual \\u003cem\\u003eBelonocnema\\u003c/em\\u003e were collected from multiple locations for both species throughout the southeastern USA in October and November in the fall of 2017 and 2018 (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003e, Table \\u003cspan refid=\\\"MOESM1\\\" class=\\\"InternalRef\\\"\\u003eS1\\u003c/span\\u003e). Galls produced by each species collected from each site were placed inside one liter glass jars covered with a coffee filter secured by a rubber band and housed in a greenhouse at Rice University (Houston, Texas) under ambient conditions (~\\u0026thinsp;23\\u0026deg;C). The jars were monitored once every two days, and newly emerged adults were collected and preserved in 95% ethanol. To measure life history traits, we randomly sampled up to 30 individuals at four \\u003cem\\u003eB. fossoria\\u003c/em\\u003e sites and five \\u003cem\\u003eB. treatae\\u003c/em\\u003e sites. For the sites with fewer than 30 wasps, we measured all available individuals yielding a total of 105 \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and 93 \\u003cem\\u003eB. treatae\\u003c/em\\u003e (see Table \\u003cspan refid=\\\"MOESM2\\\" class=\\\"InternalRef\\\"\\u003eS2\\u003c/span\\u003e for additional details).\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec4\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eMeasurements of morphological and life history traits\\u003c/h2\\u003e \\u003cp\\u003eFor each individual wasp, we measured right hind tibia length, right forewing length, and the length, width, and height of the abdomen. In Hymenoptera, the length of the right hind tibia has previously been shown to be correlated with overall body size (Rogers et al. \\u003cspan citationid=\\\"CR34\\\" class=\\\"CitationRef\\\"\\u003e1976\\u003c/span\\u003e; Honěk \\u003cspan citationid=\\\"CR19\\\" class=\\\"CitationRef\\\"\\u003e1993\\u003c/span\\u003e; Hood et al. 2011, 2017). To measure tibias and forewings accurately, they were removed from the body and slide mounted prior to measuring. After removal, the size of the abdomen was then estimated by measuring length (the longest distance between anterior and posterior ends), width (widest point perpendicular to length), and height (tallest point when viewed from the side, perpendicular to length). Abdomen volume (AV) per individual was then estimated by assuming that abdomen shape was best described as an ellipsoid using the equation AV\\u0026thinsp;=\\u0026thinsp;4/3*π*(1/2*L)*(1/2*W)*(1/2*H), where L\\u0026thinsp;=\\u0026thinsp;length, W\\u0026thinsp;=\\u0026thinsp;width, and H\\u0026thinsp;=\\u0026thinsp;height.\\u003c/p\\u003e \\u003cp\\u003eAfter its size was assessed, the abdomen was dissected following the methods of Hood and Ott (\\u003cspan citationid=\\\"CR21\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e) to measure two direct components of potential reproductive effort: egg number and egg size. Species of \\u003cem\\u003eBelonocnema\\u003c/em\\u003e are pro-ovigenic, where the ovaries contained within the abdomen house the full complement of eggs at the time of adult emergence (Hood and Ott \\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2010\\u003c/span\\u003e). Hence, a count of the number of eggs per female provides an estimate of potential maximum lifetime fecundity while measurements of individual egg size provide an estimate of reproductive investment per egg (Hood and Ott \\u003cspan citationid=\\\"CR21\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e). The product of total egg number \\u0026times; mean egg size provides an estimate of potential \\u0026lsquo;total reproductive effort\\u0026rsquo;, which we estimated for each female. We measured the size of the abdomen for two reasons. First, we tested whether abdomen volume is predictive of total reproductive effort since the abdomen contains the ovaries, which are filled with mature eggs, and abdomen size correlates with egg number in many insect species (Honěk \\u003cspan citationid=\\\"CR19\\\" class=\\\"CitationRef\\\"\\u003e1993\\u003c/span\\u003e). Second, changes in abdomen size (and shape) may relate to how observed differences in reproductive effort between the species are achieved. All traits described above were digitally measured to the nearest 0.01 mm at 20x magnification using a LEICA M125 microscope.\\u003c/p\\u003e \\u003cp\\u003eTo count the number of eggs and measure egg size, the abdomen was removed and placed in a depression slide containing a 1:1:13 acetic acid, glycerol, and water solution for at least 20 minutes to loosen ovarian follicular tissue binding eggs to the ovaries and abdomen wall (Hood and Ott \\u003cspan citationid=\\\"CR21\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e). At this time, the exoskeleton was removed, then the abdomen was dyed with a single drop of 1% methylene blue to better visualize the eggs. Potential fecundity was determined by counting the total number of eggs contained in the abdomen of each female. Mean egg size per female was estimated by randomly selecting five eggs from each female, and then measuring the length (the longest axis) and width (the widest point perpendicular to the length) following the methods of Hood and Ott (\\u003cspan citationid=\\\"CR21\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e). Both measurements were made using a LEICA M125 microscope at 100x magnification. Egg volume (EV) was estimated using the formula for a prolate spheroid following Hood and Ott (\\u003cspan citationid=\\\"CR21\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e): EV\\u0026thinsp;=\\u0026thinsp;4/3*π*(1/2*L)*(1/2*W)\\u003csup\\u003e2\\u003c/sup\\u003e, where L\\u0026thinsp;=\\u0026thinsp;length and W\\u0026thinsp;=\\u0026thinsp;width. Total reproductive effort per female was then estimated by multiplying the estimated average egg volume by the observed total number of eggs for each female (Hood and Ott \\u003cspan citationid=\\\"CR21\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e).\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec5\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eTest of divergence of morphological and life history traits\\u003c/h2\\u003e \\u003cp\\u003eTo compare differences in body size, wing length, number of eggs, egg size, abdomen volume, and total reproductive effort between \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e, we performed linear mixed model analysis with each of these six traits as the response variables, \\u0026lsquo;wasp species\\u0026rsquo; as a fixed factor, and \\u0026lsquo;collection site\\u0026rsquo; as a random factor. This and all other analyses were conducted in R version 4.0.2 (R Core Team 2020).\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec6\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eTest of independent adjustment of life history traits\\u003c/h2\\u003e \\u003cp\\u003eTo examine if and to what extent trait divergence is dependent on body size, we assessed the relationships between (a) body size and wing length, (b) body size and abdomen volume, (c) body size and number of eggs, (d) body size and egg size, and (e) body size and total reproductive effort using the comparative framework outlined by Hood and Ott (\\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e) see modified version in Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e. This modified framework outlines four scenarios for how differences in body size and life history traits between species can be achieved by comparing changes in the species-specific slope and/or intercept of the linear relationship between the traits (see Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e for details).\\u003c/p\\u003e \\u003cp\\u003eTo statistically test whether and how these linear relationships (slope and/or intercept) between life history traits and body size differ between the two wasp species (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e), linear mixed model analysis was applied with each of the five traits listed above as the response variable, and \\u0026lsquo;body size\\u0026rsquo;, \\u0026lsquo;wasp species\\u0026rsquo;, and the interaction term \\u0026lsquo;body size \\u0026times; wasp species\\u0026rsquo; as fixed factors, and collection site as the random factor. To reduce multicollinearity among interaction terms, all response variables and body size were standardized using the function \\u003cem\\u003escale\\u003c/em\\u003e in R. Divergence in linear relationship of response variables with body size between species is indicated by a significant wasp species term (i.e., different intercepts) and/or a significant interaction term between body size and wasp species (i.e., different slopes) in the model.\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec7\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eTest of environmental effects of morphological and life history traits\\u003c/h2\\u003e \\u003cp\\u003eTo examine whether phenotypic difference between species has a genetic basis or is a product of the rearing host plant environment (i.e., plasticity), we conducted a reciprocal transplant experiment. Details of the reciprocal transplant experiment are given in Hood et al. (\\u003cspan citationid=\\\"CR23\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e) and Zhang et al. (\\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2021a\\u003c/span\\u003e), respectively. Briefly, in the spring of 2017, newly emerged and mated sexual generation females of each \\u003cem\\u003eBelonocnema\\u003c/em\\u003e species were separately bagged onto the newly flushed leaves of four- to five-year-old saplings (~\\u0026thinsp;1.5 m in height) of each oak species to create four treatments: asexual generation \\u003cem\\u003eB. fossoria\\u003c/em\\u003e developing on its natal (\\u003cem\\u003eQ. geminata\\u003c/em\\u003e) or non-natal (\\u003cem\\u003eQ. virginiana\\u003c/em\\u003e) host plant, and asexual generation \\u003cem\\u003eB. treatae\\u003c/em\\u003e developing on its natal (\\u003cem\\u003eQ. virginiana)\\u003c/em\\u003e or non-natal (\\u003cem\\u003eQ. geminata\\u003c/em\\u003e) host plant. Following oviposition, saplings with wasp eggs deposited within their leaves were transferred to a greenhouse at Rice University held at ambient conditions (~\\u0026thinsp;23\\u0026deg;C), and watered once every other day until gall formers matured in fall 2017. At this time, galls containing \\u003cem\\u003eBelonocnema\\u003c/em\\u003e from each replicate of each treatment were transferred to rearing containers and monitored for emergence as described above. In total, 70 \\u003cem\\u003eQ. virginiana\\u003c/em\\u003e saplings and 66 \\u003cem\\u003eQ. geminata\\u003c/em\\u003e saplings were used to rear asexual generation adults from the reciprocal transplant experiment.\\u003c/p\\u003e \\u003cp\\u003eWe measured body size (right hind tibia length), wing length, and abdomen size as a proxy of reproductive effort for wasps reared from each of the four experimental treatments using the methods described above (see Table \\u003cspan refid=\\\"MOESM2\\\" class=\\\"InternalRef\\\"\\u003eS2\\u003c/span\\u003e for a summary of the number of asexual females measured per treatment). Based on the patterns in our data, the length of the abdomen is a sufficient estimator of the total predicted reproductive effort per female (correlation coefficient r\\u0026thinsp;=\\u0026thinsp;0.84, see Fig. \\u003cspan refid=\\\"MOESM1\\\" class=\\\"InternalRef\\\"\\u003eS1\\u003c/span\\u003e). To determine whether the relationship between body size and each of the two morphological traits, wing length (i.e., a proxy for dispersal ability) and abdomen length (i.e., a proxy for reproductive effort), differed between wasp species reared on natal versus non-natal host plants, we conducted a linear mixed model with each life history trait as the response variable, \\u0026lsquo;body size\\u0026rsquo;, \\u0026lsquo;wasp species\\u0026rsquo;, \\u0026lsquo;rearing host\\u003cem\\u003e\\u0026rsquo;\\u003c/em\\u003e (natal or non-natal host plant), and interaction among these three factors as fixed factors, and \\u0026lsquo;collection site\\u0026rsquo; treated as a random factor. In each mixed model analysis, we used the function \\u003cem\\u003elme\\u003c/em\\u003e in R package \\u0026ldquo;\\u003cem\\u003enlme\\u003c/em\\u003e\\u0026rdquo; and calculated the least square means of each of the phenotypic traits using function \\u003cem\\u003eemmeans\\u003c/em\\u003e.\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec8\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eTest of within-species trade-offs between dispersal and reproduction\\u003c/h2\\u003e \\u003cp\\u003eWe tested the relationship between the following traits: wing size, abdomen volume, egg size, egg number, and reproductive effort. Many of these traits covary with body size. To remove the effect of body size variation masking the trade-offs between wing size and reproductive traits within species, we first calculated the residuals of the correlation between wing size against body size. Then we assessed the linear correlations between the residuals of wing size against body size with reproductive traits including abdomen volume, egg number, egg size, and total reproductive effort. If intraspecific trade-offs exist, a significant negative correlation coefficient of these linear relationships is expected. These analyses were conducted for \\u003cem\\u003eB. treatae\\u003c/em\\u003e and \\u003cem\\u003eB. fossoria\\u003c/em\\u003e datasets separately.\\u003c/p\\u003e \\u003c/div\\u003e\"},{\"header\":\"Results\",\"content\":\"\\u003cdiv id=\\\"Sec10\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eDivergence of morphological and life history traits between species\\u003c/h2\\u003e \\u003cp\\u003eOn average, \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e do not differ in body size as indexed by hind tibia length (least square mean\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;SE: \\u003cem\\u003eB. fossoria\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;1.09\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.031 mm, \\u003cem\\u003eB. treatae\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;1.11\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.031 mm, \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.569, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.587, Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig4\\\" class=\\\"InternalRef\\\"\\u003e4\\u003c/span\\u003eA). However, \\u003cem\\u003eB. fossoria\\u003c/em\\u003e had significantly shorter wings and larger abdomens than \\u003cem\\u003eB. treatae\\u003c/em\\u003e (wing length: \\u003cem\\u003eB. fossoria\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;2.21\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.082 mm, \\u003cem\\u003eB. treatae\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;2.94\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.083 mm, \\u003cem\\u003et\\u003c/em\\u003e = -6.240, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001, Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig4\\\" class=\\\"InternalRef\\\"\\u003e4\\u003c/span\\u003eB; abdomen volume: \\u003cem\\u003eB. fossoria\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;1.391\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.156 mm\\u003csup\\u003e3\\u003c/sup\\u003e, \\u003cem\\u003eB. treatae\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.745\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.162 mm\\u003csup\\u003e3\\u003c/sup\\u003e, \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;2.873, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.028, Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig4\\\" class=\\\"InternalRef\\\"\\u003e4\\u003c/span\\u003eC). While \\u003cem\\u003eB. fossoria\\u003c/em\\u003e tended to produce more eggs per individual than \\u003cem\\u003eB. treatae\\u003c/em\\u003e, the difference was not significant (\\u003cem\\u003eB. fossoria\\u0026thinsp;=\\u003c/em\\u003e\\u0026thinsp;244\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;17.1, \\u003cem\\u003eB. treatae\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;199\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;17.6, \\u003cem\\u003et\\u003c/em\\u003e = -1.826, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.111; Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig4\\\" class=\\\"InternalRef\\\"\\u003e4\\u003c/span\\u003eD). However, \\u003cem\\u003eB. fossoria\\u003c/em\\u003e did allocate significantly more resources towards reproduction than \\u003cem\\u003eB. treatae\\u003c/em\\u003e (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig4\\\" class=\\\"InternalRef\\\"\\u003e4\\u003c/span\\u003eE-F) based on individual egg volume (\\u003cem\\u003eB. fossoria\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;6.6\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.179 x 10\\u003csup\\u003e\\u0026minus;\\u0026thinsp;4\\u003c/sup\\u003e mm\\u003csup\\u003e3\\u003c/sup\\u003e, \\u003cem\\u003eB. treatae\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;5.5\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;1.89 x 10\\u003csup\\u003e\\u0026minus;\\u0026thinsp;4\\u003c/sup\\u003e mm\\u003csup\\u003e3\\u003c/sup\\u003e, \\u003cem\\u003et\\u003c/em\\u003e = -4.148, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.004) and total reproductive effort (\\u003cem\\u003eB. fossoria\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.162\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.010 mm\\u003csup\\u003e3\\u003c/sup\\u003e, \\u003cem\\u003eB. treatae\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.11\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.01 mm\\u003csup\\u003e3\\u003c/sup\\u003e, \\u003cem\\u003et\\u003c/em\\u003e = -3.643, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.008).\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec11\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eDivergence in relationship between life history traits and body size within species\\u003c/h2\\u003e \\u003cp\\u003eThe association between body size and wing length for \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e was consistent with the second scenario in Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e where wing length increased with body size in each species (\\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;5.43, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001), but only the y-intercept differed significantly between the two species (\\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;12.46, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001; Table\\u0026nbsp;\\u003cspan refid=\\\"Tab1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e; Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003ea). For each reproductive trait, the linear relationships with body size followed the fourth scenario in Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e where abdomen volume (\\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;9.12, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001), potential fecundity (\\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;11.85, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001), egg volume (\\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;3.59, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001) and total reproductive effort (\\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;11.76, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001) increased with increasing body size (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003eB-E). The y-intercept differed significantly between the two wasp species (abdomen volume: \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;4.83, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.020; potential fecundity: \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;2.55, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.038; egg volume: \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;4.37, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.003; total reproductive effort: \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;4.85, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.002; Table\\u0026nbsp;\\u003cspan refid=\\\"Tab1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e). Similarly, the slope of the linear relationship in each comparison differed between species (abdomen volume: \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;1.957, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.052; potential fecundity: \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;2.959, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.004; egg volume: \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;2.36; \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.020; predicted reproductive effort: \\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;4.31, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001; Table\\u0026nbsp;\\u003cspan refid=\\\"Tab1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003e \\u003cdiv class=\\\"gridtable\\\"\\u003e\\u003ctable float=\\\"Yes\\\" id=\\\"Tab1\\\" border=\\\"1\\\"\\u003e \\u003ccaption language=\\\"En\\\"\\u003e \\u003cdiv class=\\\"CaptionNumber\\\"\\u003eTable 1\\u003c/div\\u003e \\u003cdiv class=\\\"CaptionContent\\\"\\u003e \\u003cp\\u003eLinear mixed model analysis of the relationship between body size and five life history traits: wing length, abdomen volume, potential fecundity, egg volume and reproductive effort (the product of egg number and mean egg volume per female) for \\u003cem\\u003eB. treatae\\u003c/em\\u003e and \\u003cem\\u003eB. fossoria\\u003c/em\\u003e.\\u003c/p\\u003e \\u003c/div\\u003e \\u003c/caption\\u003e \\u003ccolgroup cols=\\\"6\\\"\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c1\\\" colnum=\\\"1\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c2\\\" colnum=\\\"2\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"char\\\" char=\\\".\\\" class=\\\"colspec\\\" colname=\\\"c3\\\" colnum=\\\"3\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"char\\\" char=\\\".\\\" class=\\\"colspec\\\" colname=\\\"c4\\\" colnum=\\\"4\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"char\\\" char=\\\".\\\" class=\\\"colspec\\\" colname=\\\"c5\\\" colnum=\\\"5\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"char\\\" char=\\\".\\\" class=\\\"colspec\\\" colname=\\\"c6\\\" colnum=\\\"6\\\"\\u003e\\u003c/div\\u003e \\u003cthead\\u003e \\u003ctr\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c1\\\"\\u003e \\u003cp\\u003eResponsible variable\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eFactors\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003eEstimate\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003eSE\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003et\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003eP\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/th\\u003e \\u003c/tr\\u003e \\u003c/thead\\u003e \\u003ctbody\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\" morerows=\\\"2\\\" rowspan=\\\"3\\\"\\u003e \\u003cp\\u003eWing length (mm)\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.31\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.06\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e5.43\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eSpecies\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e1.4\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.11\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e12.46\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Species\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.06\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.09\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e0.63\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e0.532\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\" morerows=\\\"2\\\" rowspan=\\\"3\\\"\\u003e \\u003cp\\u003eAbdomen volume (mm\\u003csup\\u003e3\\u003c/sup\\u003e)\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.62\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.06\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e9.49\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eSpecies\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-1.22\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.24\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-5.03\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e0.002\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Species\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.42\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.09\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-4.61\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\" morerows=\\\"2\\\" rowspan=\\\"3\\\"\\u003e \\u003cp\\u003ePotential fecundity\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.73\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.06\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e11.85\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eSpecies\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.66\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.26\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-2.55\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e0.038\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Species\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.32\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.11\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-2.96\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e0.004\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\" morerows=\\\"2\\\" rowspan=\\\"3\\\"\\u003e \\u003cp\\u003eEgg volume (mm\\u003csup\\u003e3\\u003c/sup\\u003e)\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.29\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.08\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e3.59\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eSpecies\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.94\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.22\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-4.37\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e0.003\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Species\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.33\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.14\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-2.36\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e0.020\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\" morerows=\\\"2\\\" rowspan=\\\"3\\\"\\u003e \\u003cp\\u003eReproductive effort\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.72\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.06\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e11.76\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eSpecies\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.9\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.18\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-4.85\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e0.002\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Species\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.46\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.11\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-4.31\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003c/tbody\\u003e \\u003c/colgroup\\u003e \\u003ctfoot\\u003e \\u003ctr\\u003e\\u003ctd colspan=\\\"6\\\"\\u003e* \\u003cem\\u003eP\\u003c/em\\u003e-values in bold: P\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05.\\u003c/td\\u003e\\u003c/tr\\u003e \\u003c/tfoot\\u003e \\u003c/table\\u003e\\u003c/div\\u003e \\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec12\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eTest for environmental effects on morphological and life history traits due to different host plants\\u003c/h2\\u003e \\u003cp\\u003eThe reciprocal transplant experiments demonstrated that the observed differences between species in the linear relationship between (a) body size and wing length (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig6\\\" class=\\\"InternalRef\\\"\\u003e6\\u003c/span\\u003eA) and (b) body size and abdomen length (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig6\\\" class=\\\"InternalRef\\\"\\u003e6\\u003c/span\\u003eB) did not change when wasps are reared in their non-natal host environment. The y-intercept of the linear relationship between wing length and body size differed significantly between \\u003cem\\u003eB. treatae\\u003c/em\\u003e and \\u003cem\\u003eB. fossoria\\u003c/em\\u003e (\\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;9.50, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001; Table\\u0026nbsp;\\u003cspan refid=\\\"Tab2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e), but not between alternative rearing hosts (\\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0. 820, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.414; Table\\u0026nbsp;\\u003cspan refid=\\\"Tab2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e). Similarly, the slope did not change between different rearing hosts (\\u003cem\\u003et\\u003c/em\\u003e = -0.19, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.846; Table\\u0026nbsp;\\u003cspan refid=\\\"Tab2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e). For the linear relationship between abdomen length and body size, the y-intercept differed significantly between the two wasp species (\\u003cem\\u003et\\u003c/em\\u003e = -2.91, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.044; Table\\u0026nbsp;\\u003cspan refid=\\\"Tab2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e), but not between alternative rearing host environments (\\u003cem\\u003et\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.99, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.325; Table\\u0026nbsp;\\u003cspan refid=\\\"Tab2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e). Similarly, the slope did not change between different rearing hosts (body size \\u0026acute; rearing host: \\u003cem\\u003et\\u003c/em\\u003e = -1.43, \\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.156; Table\\u0026nbsp;\\u003cspan refid=\\\"Tab2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003e \\u003cdiv class=\\\"gridtable\\\"\\u003e\\u003ctable float=\\\"Yes\\\" id=\\\"Tab2\\\" border=\\\"1\\\"\\u003e \\u003ccaption language=\\\"En\\\"\\u003e \\u003cdiv class=\\\"CaptionNumber\\\"\\u003eTable 2\\u003c/div\\u003e \\u003cdiv class=\\\"CaptionContent\\\"\\u003e \\u003cp\\u003eLinear mixed model analysis of the reciprocal transplant experiment testing the potential effect of host plant rearing environment on abdomen size (length) and forewing length of \\u003cem\\u003eB. treatae\\u003c/em\\u003e (N\\u0026thinsp;=\\u0026thinsp;105) and \\u003cem\\u003eB. fossoria\\u003c/em\\u003e (N\\u0026thinsp;=\\u0026thinsp;93).\\u003c/p\\u003e \\u003c/div\\u003e \\u003c/caption\\u003e \\u003ccolgroup cols=\\\"7\\\"\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c1\\\" colnum=\\\"1\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c2\\\" colnum=\\\"2\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"char\\\" char=\\\".\\\" class=\\\"colspec\\\" colname=\\\"c3\\\" colnum=\\\"3\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"char\\\" char=\\\".\\\" class=\\\"colspec\\\" colname=\\\"c4\\\" colnum=\\\"4\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"char\\\" char=\\\".\\\" class=\\\"colspec\\\" colname=\\\"c5\\\" colnum=\\\"5\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"char\\\" char=\\\".\\\" class=\\\"colspec\\\" colname=\\\"c6\\\" colnum=\\\"6\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"char\\\" char=\\\".\\\" class=\\\"colspec\\\" colname=\\\"c7\\\" colnum=\\\"7\\\"\\u003e\\u003c/div\\u003e \\u003cthead\\u003e \\u003ctr\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c1\\\"\\u003e \\u003cp\\u003eResponsible variable\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eFactors\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003eEstimate\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003eSE\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003edf\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003et\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003eP\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/th\\u003e \\u003c/tr\\u003e \\u003c/thead\\u003e \\u003ctbody\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\" morerows=\\\"6\\\" rowspan=\\\"7\\\"\\u003e \\u003cp\\u003eAbdomen size\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.74\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.08\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e126\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e9.86\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eWasp species\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.82\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.28\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e4\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e-2.91\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e0.044\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eRearing host\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.13\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.13\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e126\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e0.99\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e0.325\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Wasp species\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.16\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.11\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e126\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e-1.43\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e0.156\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Rearing host\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.26\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.12\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e126\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e2.21\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e0.029\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eWasp species \\u0026times; Rearing host\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.14\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.24\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e126\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e0.57\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e0.568\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Wasp species \\u0026times; Rearing host\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.32\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.21\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e126\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e-1.51\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e0.132\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\" morerows=\\\"6\\\" rowspan=\\\"7\\\"\\u003e \\u003cp\\u003eWing length\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.71\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.07\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e120\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e10.47\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e\\u0026lt;\\u0026thinsp;0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eWasp species\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e1.25\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.13\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e4\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e9.5\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e\\u003cb\\u003e0.001\\u003c/b\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eRearing host\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.09\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.1\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e120\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e0.82\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e0.414\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Wasp species\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.12\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.09\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e120\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e-1.31\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e0.193\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Rearing host\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e-0.02\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.1\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e120\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e-0.19\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e0.846\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eWasp species \\u0026times; Rearing host\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.3\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.19\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e120\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e1.57\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e0.119\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003eBody size \\u0026times; Wasp species \\u0026times; Rearing host\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.13\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c4\\\"\\u003e \\u003cp\\u003e0.18\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e120\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e0.73\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"char\\\" char=\\\".\\\" colname=\\\"c7\\\"\\u003e \\u003cp\\u003e0.466\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003c/tbody\\u003e \\u003c/colgroup\\u003e \\u003ctfoot\\u003e \\u003ctr\\u003e\\u003ctd colspan=\\\"7\\\"\\u003e* \\u003cem\\u003eP\\u003c/em\\u003e-values in bold: P\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05.\\u003c/td\\u003e\\u003c/tr\\u003e \\u003c/tfoot\\u003e \\u003c/table\\u003e\\u003c/div\\u003e \\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec13\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eTest of within-species trade-off between wing length and reproductive effort\\u003c/h2\\u003e \\u003cp\\u003eAfter controlling for the effect of body size, we found no evidence of trade-offs between wing length and reproductive effort within \\u003cem\\u003eB. fossoria\\u003c/em\\u003e or \\u003cem\\u003eB. treatae\\u003c/em\\u003e (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig7\\\" class=\\\"InternalRef\\\"\\u003e7\\u003c/span\\u003e). This conclusion is supported by a lack of significant correlations between wing length and abdomen volume, egg number, egg size, and total reproductive effort in \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and in \\u003cem\\u003eB. treatae\\u003c/em\\u003e summarized in Table\\u0026nbsp;\\u003cspan refid=\\\"Tab3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003e.\\u003c/p\\u003e \\u003cp\\u003e \\u003cdiv class=\\\"gridtable\\\"\\u003e\\u003ctable float=\\\"Yes\\\" id=\\\"Tab3\\\" border=\\\"1\\\"\\u003e \\u003ccaption language=\\\"En\\\"\\u003e \\u003cdiv class=\\\"CaptionNumber\\\"\\u003eTable 3\\u003c/div\\u003e \\u003cdiv class=\\\"CaptionContent\\\"\\u003e \\u003cp\\u003eIntraspecific correlations between wing size (associated with dispersal capacity) and four reproductive traits (abdomen volume, egg number, egg volume, and reproductive effort) after correcting for body size for \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae.\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/div\\u003e \\u003c/caption\\u003e \\u003ccolgroup cols=\\\"6\\\"\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c1\\\" colnum=\\\"1\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c2\\\" colnum=\\\"2\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c3\\\" colnum=\\\"3\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c4\\\" colnum=\\\"4\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c5\\\" colnum=\\\"5\\\"\\u003e\\u003c/div\\u003e \\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c6\\\" colnum=\\\"6\\\"\\u003e\\u003c/div\\u003e \\u003cthead\\u003e \\u003ctr\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u0026nbsp;\\u003c/th\\u003e \\u003cth align=\\\"left\\\" colspan=\\\"2\\\" nameend=\\\"c3\\\" namest=\\\"c2\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003eB. fossoria\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/th\\u003e \\u003cth align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u0026nbsp;\\u003c/th\\u003e \\u003cth align=\\\"left\\\" colspan=\\\"2\\\" nameend=\\\"c6\\\" namest=\\\"c5\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003eB. treatae\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/th\\u003e \\u003c/tr\\u003e \\u003c/thead\\u003e \\u003ctbody\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u0026nbsp;\\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003er\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003eP\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u0026nbsp;\\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003er\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e\\u003cem\\u003eP\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e \\u003cp\\u003eAbdomen volume\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003e0.06\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.627\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u0026nbsp;\\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e0.05\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e0.612\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e \\u003cp\\u003eEgg number\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003e-0.05\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.679\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u0026nbsp;\\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e0.07\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e0.495\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e \\u003cp\\u003eEgg volume\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003e-0.02\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.859\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u0026nbsp;\\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-0.147\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e0.202\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003ctr\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e \\u003cp\\u003eReproductive effort\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e \\u003cp\\u003e-0.13\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e \\u003cp\\u003e0.344\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u0026nbsp;\\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e \\u003cp\\u003e-0.05\\u003c/p\\u003e \\u003c/td\\u003e \\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e \\u003cp\\u003e0.689\\u003c/p\\u003e \\u003c/td\\u003e \\u003c/tr\\u003e \\u003c/tbody\\u003e \\u003c/colgroup\\u003e \\u003c/table\\u003e\\u003c/div\\u003e \\u003c/p\\u003e \\u003c/div\\u003e\"},{\"header\":\"Discussion\",\"content\":\"\\u003cp\\u003eTrade-offs among life history traits are of key interest in evolutionary biology due to their potential impacts on maintaining phenotypic variation or constraining adaptation under changing environments (Reznick and Bryga \\u003cspan citationid=\\\"CR29\\\" class=\\\"CitationRef\\\"\\u003e1987\\u003c/span\\u003e; Futuyma and Moreno \\u003cspan citationid=\\\"CR15\\\" class=\\\"CitationRef\\\"\\u003e1988\\u003c/span\\u003e). On one hand, trade-offs may act as evolutionary constraints for adaptation among individuals; on the other hand, long-term selection may overcome such constraints and reshape trait combinations (Agrawal \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Therefore, investigating whether trade-offs scale across different levels of biological organization, from individuals to populations and species, will shed light on the evolutionary processes underlying life history traits (Bolnick et al. \\u003cspan citationid=\\\"CR2\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e; Agrawal \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). For example, if trait divergence between species originates from trade-offs at the intraspecific level, we would expect to find trade-offs among traits both within and between species. Alternatively, if recently diverged species evolve independently in response to divergent environmental conditions, trade-offs might only be detected at the interspecific level due to environmental constraints while not being evident at the intraspecific level. Our study addresses this potential connection between inter- and intraspecific variation by testing whether there is a trade-off between dispersal capacity and reproductive output in two sister species of gall wasps belonging to the genus \\u003cem\\u003eBelonocnema\\u003c/em\\u003e, which exhibit different dispersal capabilities. We also examined whether such trade-offs exist within each monomorphic species.\\u003c/p\\u003e \\u003cp\\u003eWe discovered that trade-offs between flight capability and reproductive output are evident as a consistent difference between the two wasp species, but no trade-offs were detected at the intraspecific level. Specifically, \\u003cem\\u003eB. fossoria\\u003c/em\\u003e exhibits reduced flight capability and smaller wing size, while demonstrating significantly higher reproductive effort compared to \\u003cem\\u003eB. treatae\\u003c/em\\u003e (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig4\\\" class=\\\"InternalRef\\\"\\u003e4\\u003c/span\\u003e). Our finding of an interspecific trade-off, but the absence of intraspecific trade-offs, aligns with previous studies exploring trade-offs between life history traits such as flower size and flower number (Worley et al. \\u003cspan citationid=\\\"CR42\\\" class=\\\"CitationRef\\\"\\u003e2000\\u003c/span\\u003e; Sargent et al. \\u003cspan citationid=\\\"CR35\\\" class=\\\"CitationRef\\\"\\u003e2007\\u003c/span\\u003e; Hahn and Maron \\u003cspan citationid=\\\"CR18\\\" class=\\\"CitationRef\\\"\\u003e2016\\u003c/span\\u003e). One possible explanation for the lack of intraspecific trade-offs in these studies is that individual variation in resource acquisition masks the correlation among traits. Individuals with greater resource availability may develop both larger and more numerous flowers, whereas individuals with limited resources but larger flowers would have fewer flowers (van Noordwijk et al. 1986; King et al. \\u003cspan citationid=\\\"CR25\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; Agrawal \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). However, in the case of our wasp species, individual resource acquisition variation is unlikely to explain the absence of trade-offs between flight-related traits and reproductive output. This is because our analysis already partially controls for the effect of body size, which indirectly accounts for individual resource variation (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig7\\\" class=\\\"InternalRef\\\"\\u003e7\\u003c/span\\u003e). An alternative explanation is that these two wasp species have recently diverged and specialized within a narrow niche (only one or two host plant species), resulting in long-term and strong selection that has shaped divergence in resource allocation between the species. Within each species, considering the narrow niche, further specialization along these life history dimensions may not be expected, thereby leading to the absence of trade-offs among life history traits (Futuyma and Moreno \\u003cspan citationid=\\\"CR15\\\" class=\\\"CitationRef\\\"\\u003e1988\\u003c/span\\u003e). We posit that the discrepancy in trade-off patterns between intra- and interspecific levels is more likely to occur among closely related species inhabiting different environments compared to species inhabiting similar environments. This is based on the common assumption that phylogenetically closely related species tend to share similar life history traits and evolutionary constraints, resulting in a consistent trade-off pattern within and among sister species (Webb et al. \\u003cspan citationid=\\\"CR41\\\" class=\\\"CitationRef\\\"\\u003e2002\\u003c/span\\u003e). However, if a particular life history trait plays a crucial role in adaptation to divergent environments, long-term and intense selection pressure could disrupt the genetic constraints on the evolution of life history traits (Agrawal \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Further studies employing a comparative framework similar to this study are needed to systematically test this hypothesis.\\u003c/p\\u003e \\u003cp\\u003eFor the two wasp species studied here, \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e, the divergence in life history traits is not solely due to changes in body size but also involves alterations in the relationship between body size and multiple traits, including wing size, abdomen size, and total reproductive effort (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003e). This means that the adjustments in flight capability and reproductive effort, as depicted by Scenario 4 in Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e, are independent of body size. Specifically, the independent adjustment of flight capability between the sister species is reflected in variation in wing size, while changes in reproductive effort are achieved through modifications in abdomen size. The independent adjustment of these life history traits is further supported by a common garden rearing experiment: the divergence in the linear relationship between body size and wing size, as well as between body size and reproductive effort, is maintained even when the wasps are reared on a non-native host (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig6\\\" class=\\\"InternalRef\\\"\\u003e6\\u003c/span\\u003e). Therefore, the divergence in life history traits is likely attributable, at least in part, to genetic differences between the two wasp species rather than being solely a plastic response to the rearing host plants (i.e., environmental effects and phenotypic plasticity).\\u003c/p\\u003e \\u003cp\\u003eInterestingly, the linear relationship between body size and wing size differs in intercepts, while the linear relationship between body size and other reproductive-related traits, including abdomen volume, potential fecundity, total reproductive effort, differs in both intercepts and slopes (Table\\u0026nbsp;\\u003cspan refid=\\\"Tab1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e, Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003e). According to quantitative genetic theory, linear relationships between traits are more prone to change in intercepts under selection, while stronger and longer-term selection is required for changes in slopes (Roff et al. \\u003cspan citationid=\\\"CR33\\\" class=\\\"CitationRef\\\"\\u003e2002\\u003c/span\\u003e). Thus, it is possible that reproductive traits experience stronger selection pressure than flight-related traits. Notably, similar changes in both intercepts and slopes between potential fecundity and body size have been observed in the alternating asexual and sexual generations of another species in the genus, \\u003cem\\u003eB. kinsey\\u003c/em\\u003e (Hood and Ott \\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e), where each generation faces unique environmental challenges. Hence, both Hood and Ott (\\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e) and the present study provide evidence that reproductive traits are highly adaptable phenotype under selection in this group of \\u003cem\\u003eBelonocnema\\u003c/em\\u003e gall wasps. Since dispersal and reproduction often exhibit trade-offs either within species or among species across many study systems (Carroll et al. \\u003cspan citationid=\\\"CR4\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e; Guerra \\u003cspan citationid=\\\"CR17\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; King et al. \\u003cspan citationid=\\\"CR25\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; Nasu and Tokuda \\u003cspan citationid=\\\"CR27\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e), more studies should link changes in linear relationship among these life history traits to the variation of selection pressures each of these traits experience (e.g., populations at the range edge).\\u003c/p\\u003e \\u003cp\\u003eWe argue that \\u003cem\\u003eB. fossoria\\u003c/em\\u003e individuals with higher reproductive output but lower flight capability may gain a fitness advantage in the \\u003cem\\u003eQ. geminata\\u003c/em\\u003e environment, leading to selection favoring the divergent phenotypes between \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae.\\u003c/em\\u003e This is because \\u003cem\\u003eB. fossoria\\u003c/em\\u003e's host plant, \\u003cem\\u003eQ. geminata\\u003c/em\\u003e, is typically found as short shrubs no taller than 10 m in small clusters in sandy soil, whereas \\u003cem\\u003eB. treatae\\u003c/em\\u003e's host plant, \\u003cem\\u003eQ. virginiana\\u003c/em\\u003e, is a much taller tree species (with a mean height of ~\\u0026thinsp;21m) and has a patchier distribution (Cavender-Bares and Pahlich \\u003cspan citationid=\\\"CR6\\\" class=\\\"CitationRef\\\"\\u003e2009\\u003c/span\\u003e). Dense, short trees may require less flight capability for the asexual generation of \\u003cem\\u003eB. fossoria\\u003c/em\\u003e to locate suitable sites for gall induction on the roots, which host the next generation (Zera and Denno \\u003cspan citationid=\\\"CR43\\\" class=\\\"CitationRef\\\"\\u003e1997\\u003c/span\\u003e). The reduced flight capability observed in \\u003cem\\u003eB. fossoria\\u003c/em\\u003e, is likely a product of relaxed selection on flight associated with more persistent, shorter, and smaller host trees, as suggested by other studies highlighting the role of habitat structure in the evolution of flight capability (Roff \\u003cspan citationid=\\\"CR32\\\" class=\\\"CitationRef\\\"\\u003e1990\\u003c/span\\u003e; Roff 1994; Denno et al. \\u003cspan citationid=\\\"CR9\\\" class=\\\"CitationRef\\\"\\u003e1996\\u003c/span\\u003e; Zera and Denno \\u003cspan citationid=\\\"CR43\\\" class=\\\"CitationRef\\\"\\u003e1997\\u003c/span\\u003e; Dell'Aglio et al. 2022). Additionally, a third allopatric species in this genus, \\u003cem\\u003eB. kinseyi\\u003c/em\\u003e, which shares host plant \\u003cem\\u003eQ. virginina\\u003c/em\\u003e with \\u003cem\\u003eB. treatae\\u003c/em\\u003e, exhibits life history traits similar in size to the \\u003cem\\u003eB. treatae\\u003c/em\\u003e populations analyzed here. These include similar flight capability (Zhang et al. \\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e2021c\\u003c/span\\u003e) and correlations between body size and wing length (wasp species: t = -0.03, p\\u0026thinsp;=\\u0026thinsp;0.981; body size x wasp species: t = -1.04, p\\u0026thinsp;=\\u0026thinsp;0.303) and between body size and abdomen length (wasp species: t = -2.88, p\\u0026thinsp;=\\u0026thinsp;0.103; body size x wasp species: t\\u0026thinsp;=\\u0026thinsp;0.56, p\\u0026thinsp;=\\u0026thinsp;0.581; see Supplement). Therefore, the divergent phenotypes between \\u003cem\\u003eB. treatae\\u003c/em\\u003e and \\u003cem\\u003eB. fossoria\\u003c/em\\u003e are likely the result of adaptation to divergent host plant-related environments. Future studies should apply a similar comparative work to other gall wasp species that also utilized the same set of host plant species \\u003cem\\u003eQ. virginiana\\u003c/em\\u003e and \\u003cem\\u003eQ. geminata\\u003c/em\\u003e. If similar divergent phenotypes are observed repeatedly between host-associated populations in other gall wasp species, this would strongly suggest the similar host plant-related environments play a critical role in shaping the divergence among these traits (Egan et al. \\u003cspan citationid=\\\"CR14\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e; Zhang et al. \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR50\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e).\\u003c/p\\u003e\"},{\"header\":\"Declarations\",\"content\":\"\\u003cp\\u003e\\u003cstrong\\u003eAcknowledgements\\u003c/strong\\u003e\\u003c/p\\u003e\\n\\u003cp\\u003eWe thank Elaine Hu, Isaac Carroo and Hannah Towbin for assistance with sample collection and morphological measurements. Funding was provided to AMR by the Spurlino Summer Undergraduate Research Fellowship, to LZ by student research award from Society for Integrative and Comparative Biology, Rosemary Grant Award from the Society for the Study of Evolution, and the Diana McSherry and Patrick Poe research award, to SPE from the Department of BioSciences at Rice University and to GRH from the Rice University Academy of Fellows.\\u003c/p\\u003e\\n\\u003cp\\u003e\\u003cstrong\\u003eAuthor Contributions:\\u003c/strong\\u003e LZ, SPE, GRH, and JRO conceived and designed the experiments. AMR performed the experiments. AMR and LZ analyzed the data. AMR and LZ wrote the manuscript; other authors edited the manuscript.\\u003c/p\\u003e\"},{\"header\":\"References\",\"content\":\"\\u003col\\u003e\\u003cli\\u003e\\u003cspan\\u003eAgrawal AA (2020) A scale-dependent framework for trade-offs, syndromes, and specialization in organismal biology. 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Annu Rev Ecol Syst 33:475\\u0026ndash;505\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eWorley AC, Baker AM, Thompson JD, Barrett SC (2000) Floral display in \\u003cem\\u003eNarcissus\\u003c/em\\u003e: variation in flower size and number at the species, population, and individual levels. Int J Plant Sci 161:69\\u0026ndash;79\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eZera AJ, Denno RF (1997) Physiology and ecology of dispersal polymorphism in insects. Ann Rev Entomol 42:207\\u0026ndash;230\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eZera AJ, Harshman LG (2001) The physiology of life history trade-offs in animals. Annu Rev Ecol Syst 32:95\\u0026ndash;126\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eZera AJ, Rankin MA (1989) Wing dimorphism in \\u003cem\\u003eGryllus rubens\\u003c/em\\u003e: genetic basis of morph determination and fertility differences between morphs. 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Am Nat 197:732\\u0026ndash;739\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eZhang L, Hood GR, Roush AM, Shzu SA, Comerford MS, Ott JR, Egan SP (2021b) Asymmetric, but opposing reductions in immigrant viability and fecundity promote reproductive isolation among host-associated populations of an insect herbivore. Evolution 75:476\\u0026ndash;489\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eZhang L, Mullin BF, Shzu S, Davis CK, Hu EG, Carroo IJ, Egan SP (2022) Parallel host-plant-associated differences in an extended phenotype between populations of six species of gall-forming insects. Ecol Entomol 47:323\\u0026ndash;330\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eZhang YM, Egan SP, Driscoe AL, Ott JR (2021c) One hundred and sixty years of taxonomic confusion resolved: \\u003cem\\u003eBelonocnema\\u003c/em\\u003e (Hymenoptera: Cynipidae: Cynipini) gall wasps associated with live oaks in the USA. Zool J Linn Soc. \\u003cspan class=\\\"ExternalRef\\\"\\u003e\\u003cspan class=\\\"RefSource\\\"\\u003e10.1093/zoolinnean/zlab001\\u003c/span\\u003e\\u003cspan address=\\\"10.1093/zoolinnean/zlab001\\\" targettype=\\\"DOI\\\" class=\\\"RefTarget\\\"\\u003e\\u003c/span\\u003e\\u003c/span\\u003e\\u003c/span\\u003e\\u003c/li\\u003e\\u003c/ol\\u003e\"}],\"fulltextSource\":\"\",\"fullText\":\"\",\"funders\":[],\"hasAdminPriorityOnWorkflow\":false,\"hasManuscriptDocX\":true,\"hasOptedInToPreprint\":true,\"hasPassedJournalQc\":\"\",\"hasAnyPriority\":false,\"hideJournal\":false,\"highlight\":\"\",\"institution\":\"\",\"isAcceptedByJournal\":true,\"isAuthorSuppliedPdf\":false,\"isDeskRejected\":\"\",\"isHiddenFromSearch\":false,\"isInQc\":false,\"isInWorkflow\":true,\"isPdf\":false,\"isPdfUpToDate\":true,\"isWithdrawnOrRetracted\":false,\"journal\":{\"display\":true,\"email\":\"info@researchsquare.com\",\"identity\":\"oecologia\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":false,\"externalIdentity\":\"oeco\",\"sideBox\":\"Learn more about [Oecologia](https://www.springer.com/journal/442)\",\"snPcode\":\"442\",\"submissionUrl\":\"https://submission.nature.com/new-submission/442/3\",\"title\":\"Oecologia\",\"twitterHandle\":\"\",\"acdcEnabled\":true,\"dfaEnabled\":true,\"editorialSystem\":\"em\",\"reportingPortfolio\":\"Springer Hybrid\",\"inReviewEnabled\":true,\"inReviewRevisionsEnabled\":false},\"keywords\":\"Belonocnema, gall wasp, host-plant adaptation, life history traits\",\"lastPublishedDoi\":\"10.21203/rs.3.rs-3171363/v1\",\"lastPublishedDoiUrl\":\"https://doi.org/10.21203/rs.3.rs-3171363/v1\",\"license\":{\"name\":\"CC BY 4.0\",\"url\":\"https://creativecommons.org/licenses/by/4.0/\"},\"manuscriptAbstract\":\"\\u003cp\\u003eA critical question in understanding the origin of trade-offs and their impact on trait evolution is whether trade-offs between species originate from within-species variation. Despite their importance, studies linking interspecific divergence between closely related species to intraspecific trait variation are still rare. In this study, we describe a trade-off between dispersal and reproductive effort between two sympatric sister species of wasps in the genus \\u003cem\\u003eBelonocnema\\u003c/em\\u003e (Hymenoptera: Cynipini: Cynipidae) that form galls on live oaks: \\u003cem\\u003eB. fossoria\\u003c/em\\u003e, which specializes on \\u003cem\\u003eQuercus geminata\\u003c/em\\u003e, and \\u003cem\\u003eB. treatae\\u003c/em\\u003e, which specializes on \\u003cem\\u003eQ. virginiana\\u003c/em\\u003e. Specifically, our results suggest that \\u003cem\\u003eB. fossoria\\u003c/em\\u003e has evolved reduced flight capability and smaller wings, but a larger abdomen and greater total reproductive effort than \\u003cem\\u003eB. treatae\\u003c/em\\u003e, which has larger wings and is a stronger flier, but a smaller abdomen and reduced total reproductive effort. Despite these significant morphological and reproductive differences, these traits remain unchanged when transplanting \\u003cem\\u003eB. fossoria\\u003c/em\\u003e and \\u003cem\\u003eB. treatae\\u003c/em\\u003e onto the alternative host plant, suggesting that divergence in these traits is likely genetic as opposed to a plastic response to the different rearing environments. However, we did not find evidence of intraspecific trade-offs between wing length and reproductive traits within either \\u003cem\\u003eB. fossoria\\u003c/em\\u003e or \\u003cem\\u003eB. treatae\\u003c/em\\u003e, indicating that trade-offs in life history traits between the two species is a result of independent adaptations in response to different environments. Our study informs our understanding of the evolution of trade-offs among life history traits by examining trade-offs at different biological organizations.\\u003c/p\\u003e\",\"manuscriptTitle\":\"A test of trade-offs in dispersal and reproduction within and between a sister species pair of specialist insect herbivores\",\"msid\":\"\",\"msnumber\":\"\",\"nonDraftVersions\":[{\"code\":1,\"date\":\"2023-08-04 23:34:38\",\"doi\":\"10.21203/rs.3.rs-3171363/v1\",\"editorialEvents\":[{\"type\":\"communityComments\",\"content\":0},{\"type\":\"reviewerAgreed\",\"content\":\"\",\"date\":\"2023-08-02T19:39:50+00:00\",\"index\":0,\"fulltext\":\"\"},{\"type\":\"reviewersInvited\",\"content\":\"\",\"date\":\"2023-07-31T13:20:23+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"editorAssigned\",\"content\":\"\",\"date\":\"2023-07-17T12:28:03+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"submitted\",\"content\":\"Oecologia\",\"date\":\"2023-07-15T11:00:20+00:00\",\"index\":\"\",\"fulltext\":\"\"}],\"status\":\"published\",\"journal\":{\"display\":true,\"email\":\"info@researchsquare.com\",\"identity\":\"oecologia\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":false,\"externalIdentity\":\"oeco\",\"sideBox\":\"Learn more about [Oecologia](https://www.springer.com/journal/442)\",\"snPcode\":\"442\",\"submissionUrl\":\"https://submission.nature.com/new-submission/442/3\",\"title\":\"Oecologia\",\"twitterHandle\":\"\",\"acdcEnabled\":true,\"dfaEnabled\":true,\"editorialSystem\":\"em\",\"reportingPortfolio\":\"Springer Hybrid\",\"inReviewEnabled\":true,\"inReviewRevisionsEnabled\":false}}],\"origin\":\"\",\"ownerIdentity\":\"8c168490-643b-48ba-8c03-9972eb672904\",\"owner\":[],\"postedDate\":\"August 4th, 2023\",\"published\":true,\"recentEditorialEvents\":[],\"rejectedJournal\":[],\"revision\":\"\",\"amendment\":\"\",\"status\":\"published-in-journal\",\"subjectAreas\":[],\"tags\":[],\"updatedAt\":\"2024-02-12T15:06:37+00:00\",\"versionOfRecord\":{\"articleIdentity\":\"rs-3171363\",\"link\":\"https://doi.org/10.1007/s00442-024-05512-3\",\"journal\":{\"identity\":\"oecologia\",\"isVorOnly\":false,\"title\":\"Oecologia\"},\"publishedOn\":\"2024-02-07 15:01:35\",\"publishedOnDateReadable\":\"February 7th, 2024\"},\"versionCreatedAt\":\"2023-08-04 23:34:38\",\"video\":\"\",\"vorDoi\":\"10.1007/s00442-024-05512-3\",\"vorDoiUrl\":\"https://doi.org/10.1007/s00442-024-05512-3\",\"workflowStages\":[]},\"version\":\"v1\",\"identity\":\"rs-3171363\",\"journalConfig\":\"researchsquare\"},\"__N_SSP\":true},\"page\":\"/article/[identity]/[[...version]]\",\"query\":{\"redirect\":\"/article/rs-3171363\",\"identity\":\"rs-3171363\",\"version\":[\"v1\"]},\"buildId\":\"_2-kVJe1T_tPrBINL-cwx\",\"isFallback\":false,\"isExperimentalCompile\":false,\"dynamicIds\":[84888],\"gssp\":true,\"scriptLoader\":[]}","source_license":"CC-BY-4.0","license_restricted":false}