Landscape fragmentation and agricultural context impact pollination dynamics of native annual plants

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Walker, Margaret M. Mayfield This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4756788/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 04 Feb, 2026 Read the published version in Landscape Ecology → Version 1 posted 9 You are reading this latest preprint version Abstract Context Agricultural intensification leads to habitat loss and fragmentation, disrupting plant-pollinator interactions directly, through changes in landscape configuration, and indirectly through altered land-use practices. This has detrimental consequences for the persistence of plants, pollinators, and the ecosystem services they provide. Objectives We investigated the mechanisms by which environmental and agricultural context impact pollination dynamics and the reproductive success of native plant species in remnant vegetation within an agricultural mosaic. Specifically, we evaluate the direct and indirect effects of landscape fragmentation (patch size and edge effect) and agricultural practices (crop type adjacent to natural remnants) on bee communities and native plants seed production. Methods We sampled the pollinator community and conducted pollination experiments on four native annual plant species in the core and edge of nine natural remnants. For each site, we recorded remnant size, adjacent crop type (canola or wheat), and local environmental and biological conditions. We then assessed the relationships between these landscape features, bee communities, pollination services, and the reproductive success of native annual forb species. Results Bee abundance was higher in reserves adjacent to canola compared to wheat. However, bee abundance decreased from the core to the edge of remnants adjacent to canola, suggesting a possible pollinator dilution effect. Canola directly and indirectly increased seed production of the focal plant species, mediated by changes in pollinator abundance. Conclusions Adjacent crop type, edge effects, and patch size shape plant-pollinator interactions through changes in pollinator abundance, whereas local-scale floral abundance influence pollination dynamics. Our findings indicate that agricultural practices impact the reproductive success of native plants persisting in remnants within an intensively managed agricultural landscape. Further, we show that this effect is mediated by the abundance of generalist insect pollinators in remnant vegetation. landscape fragmentation agricultural practices pollination dynamics plant population York gum-Jam woodlands Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Land-use intensification and landscape fragmentation negatively impact biodiversity (Fahrig 2003 ). This has been shown to occur through direct processes such as population reduction and isolation (Honnay et al. 2005 ; Lino et al. 2018 ) and indirect processes such as disruptions to species interactions (Mustajärvi et al. 2001 ; Ferreira et al. 2013 ). While these direct processes are well understood (Fahrig 2003 ; Wilson et al. 2016 ) and we have a theoretical understanding of indirect effects (Püttker et al. 2020 ), we still lack a strong empirical understanding of how indirect mechanisms drive specific consequences of land-use intensification on native species and their persistence in habitat remnants (but see Lázaro et al. 2020 ). Multi-trophic and mutualistic interactions, such as plant-pollinator interactions, are critical for maintaining biodiversity (Allen-Wardell et al. 1998 ; Wei et al. 2021 ). They are well known to break down in isolated habitat remnants and their disruption can lead to changes in ecosystem function and services in fragmented landscapes (Viana et al. 2012 ; Hadley and Betts 2012 ). Natural vegetation remnants within agricultural landscapes are critical for biodiversity conservation and ecosystem services (Soga et al. 2014 ; Decocq et al. 2016 ). Resident plants and insect pollinators in remnants are often highly dependent on each other, yet fragmentation isolates them from other populations, limiting the extent of pollination services in fragmented landscapes (Harris and Johnson 2004 ; Aguilar et al. 2006 ; Kolb 2008 ). Disruptions to plant-pollinator interactions can occur in remnants when the temporal and spatial distribution of interacting partners are restricted (e.g., due to isolation) or when the interacting species respond differently to the novel environmental conditions (e.g., edge effects; Hadley and Betts 2012 ). Pollinator spillover from remnant vegetation into agricultural fields can greatly benefit crop production (Garibaldi et al. 2013 ). Pollinators, particularly bees, nest in native vegetation patches and forage on crop flowers in adjacent farmlands and native plants within remnants (Potts et al. 2005 ). This pollination service mediates seed and fruit production for both crops and native plants (Garibaldi et al. 2013 ). While the beneficial effect of pollinator spillover on crop systems is well understood, we have limited knowledge of how this spillover affects within-remnant species interactions and native plant reproduction, despite its known importance for maintaining the conservation value of these remnants (Blitzer et al. 2012 ; Geslin et al. 2017 ). Insect-attracting mass flowering crops, such as canola ( Brassica napus , Brassicaceae), share pollinators with co-flowering native plants in many agricultural systems (Stanley and Stout 2014 ; Reynolds et al. 2022 ). Canola crops are known to sometimes increase the abundance of pollinators in adjacent remnants due to the presence of this abundant but ephemeral food resource (Holzschuh et al. 2013 ; Bailey et al. 2014 ). Other studies, however, have shown that the presence of abundant canola flowers can alternatively lead to pollinator dilution, with pollinators attracted to crop flowers over native flowers, especially at peak flowering when canola is the most abundant, if not the highest quality, resource available (Holzschuh et al. 2016 ; Reynolds et al. 2022 ). This shift of pollinator foraging is known to lower the reproductive output of co-flowering native plants due to deficient pollinator services (Knight et al. 2005 ). Indeed, in some landscapes dominated by canola, canola has been shown to have negative effects on native plant reproduction despite increased pollinator abundances in the landscape as a whole (Holzschuh et al. 2011 ; Van Reeth et al. 2019 ). While pollinator dilution is often invoked to explain this effect, few studies have attempted to determine the mechanisms by which mass flowering crops like canola affect native plant seed production in vegetation remnants in agricultural landscapes, with most assuming rather than showing a relationship (but see Stanley and Stout 2014 ; Ekroos et al. 2015 ). Alternatively, interacting landscape and local scale factors, such as plant community composition and abundance, can buffer plant reproductive success from the negative impact of pollinator competition with flowering crops and landscape fragmentation (Kovács-Hostyánszki et al. 2013 ; Ekroos et al. 2015 ). Our study aims to determine the biological consequences of land-use intensification on native plant population dynamics as mediated by insect pollinators. We explore this in the critically endangered York gum-Jam woodlands (YGJW) of SW Western Australia. These woodlands have lost more than 93% of their original extent due to land clearing and agricultural intensification since European invasion (Prober and Smith 2009 ; DAWE, 2023 ). Native vegetation is now found only in small isolated patches embedded within an agricultural mosaic of wheat and canola fields (Yates and Hobbs 1997 ). Understanding the pollination dynamics of wildflower species in such a system is critical for effective management and conservation. In this study, we explore how environmental and agricultural context impacts pollinator abundance, pollination services, and the reproductive success of four native annual forb species. Specifically, we ask (i) how the pollinator community and plant diversity are affected by landscape context (patch size, edge effect, and type of neighbouring crops)? and (ii) does the interplay between landscape context, pollinator abundance, and plant community structure modulate the reproductive success of plant species in remnant woodlands? Methods Study system The York gum-Jam woodlands (YGJW) of SW Western Australia are a threatened ecological community within the global biodiversity hot spot of the Southwest Australian floristic region (DAWE 2023 ). These woodlands occur throughout the agricultural wheatbelt region and have experienced widespread clearing and fragmentation (Prober and Smith 2009 ; DAWE 2023 ). Remnant YGJW typically occur in small, geographically isolated patches within an agricultural matrix. These woodlands are dominated by an overstorey of York gum ( Eucalyptus loxophleba Beth. Subsp. Loxophleba) and Jam ( Acacia acuminata Benth) and comprise a diverse understory of shrubs, perennial grasses, and annual and perennial exotic and native forbs (Dwyer et al. 2015 ). The climate in this region is semi-arid (350 to 450 mm rainfall per year) with cool, wet winters and warm, dry summers (Hopper and Gioia 2004 ). These woodlands generally have an open canopy and phosphorous-poor sandy soils (Lambers et al. 2008 ), creating diverse microhabitats across gradients of light, litter, and soil nutrients (Dwyer et al. 2015 ). The annual wildflower community primarily consists of Asteraceae, Goodeniaceae, and Araliaceae species that germinate during the winter (July-August) and flower and seed in spring (August-November). This study was conducted across nine YGJW remnants (Fig. 1 a) adjacent to either an insect pollinator-attracting crop (canola) or a non-insect-attracting crop (wheat or pasture). Both Wheat and canola bloom from July to late August in this part of Western Australia, flowering at the same time as many of the native wildflowers that live in the YGJW remnants. Sites were selected based on remnant size, accessibility, and neighbouring crop identity. Not all species were found in all reserves, and some sites did not have adequate plant abundances to perform edge and core transects in the same remnants (Table S1 ). Two sites were established in the same remnant but with sites positioned more than 500 m apart. In both cases, the two sites in the same reserves were adjacent to different crop types (either canola or wheat/pasture). These sites were assumed to have independence (despite occurring in the same remnant) because the native pollinating insects in this system have small home ranges and typically travel less than 500 m to forage (Gathmann and Tscharntke 2002 ). Further, the YGJW vegetation is patchy within remnants (Fig. 1 b) and sites were not positioned in contiguous patches of YJGW (i.e., they are separated by patches of other woodland types that do not have wildflower understories). Study species Four native annual plant species were selected for this study (Fig. 2 ): Goodenia rosea (S.Moore) K.A.Sheph. (‘GORO’, Goodeniaceae), Lawrencella rosea Lindl. (‘LARO’, Asteraceae), Podolepis aristata Benth. (‘POAR’, Asteraceae), and Podotheca gnaphaloides Graham (‘POGN’, Asteraceae). These species were selected because they common within YGJW remnants, were found in most sites (Table S1 ) and have overlapping peak flowering with canola in the region. These species have contrasting floral morphology (Fig. 2 ) and are visited by a diverse group of pollinators, including feral honeybees, native bees, flies, and beetles (personal observation). Pollination treatments and experimental design We performed pollen limitation experiments in edge and core transects in all resident remnants to determine how landscape context impacted reproductive success mediated by insect pollinators. These experiments took place from late July to late October in 2022, during the complete flowering period of each focal species. To do this, we set up two 50 m x 5 m long transects in each remnant parallel to an adjacent crop field: one next to the remnant edge and a second at the core of the remnant, defined as at least 150 m from the crop field edge (Fig. 1 b). Past studies have shown this distance to be sufficient to observe differences in insect pollinators in this system (Reynolds et al. 2022 ). Along each transect, ten 30 x 30 cm plots were established for each species. This resulted in a total of 520 plots and 2080 individual plants across all four species. For LARO, 640 individual plants were assigned to one of the four pollination treatments (160 per treatment) in a total of 8 sites, and the rest of the species (GORO, POAR, POGN) had each 480 individuals selected in a total of six different reserves each (120 per treatment). Within each plot, four individual plants of the focal species were selected. Each plant was randomly assigned one of the four pollination treatments: (i) ‘open’ – flowers remained unmanipulated and open to wild pollinators; (ii) ‘open-supplemented’ – flowers remained open to wild pollinators and were hand pollinated; (iii) ‘bagged’ – flowers were covered with a mesh bag before flowering; and (iv) ‘bagged-supplemented’ – flowers were bagged before opening and hand pollinated once flowers opened. Pollinator exclusion involved covering a flower with a mesh bag to prevent pollinators from accessing or interacting with the floral reproductive parts, and pollen supplementation consisted of manually adding pollen from different conspecific individuals in the immediate area to the focal plant. Pollen supplementation was performed by rubbing the focal plant inflorescence with three different conspecific inflorescences for the Asteraceae species or rubbing indusium (pollen-cup structure) in the case of GORO. Pollen supplementation was conducted every four to five days, and it was applied to all open flowers of each focal plant in both open and bagged-supplemented treatment. Focal plants in these two treatments were hand-pollinated at least three times for LARO and POGN (shorter flowering period) and six times for GORO and POAR (longer flowering period). At the end of the flowering season, all fruits were protected with a mesh bag and collected when matured. Across the 2080 focal plants, we collected 310,880 viable seeds. We calculated the pollen limitation index ( PLi ) for each plot using Eckert et al., ( 2010 ); \(\:PLi=\frac{Ssup-Sop}{Sop}\) , where \(\:Ssup\) is the average individual seed production of the focal plant that was open to natural pollination, and hand-supplemented (‘open supplemented’) and \(\:Sop\) is the average individual seed production of naturally pollinated flowers (open). This index calculates the percentage increase in seed production due to the supplemented treatment ranging from - ∞ to 1. Positive values indicate higher reproductive success in pollen-supplemented plants (higher rates of pollen limitation), and negative values indicate that control plants (open) had higher reproductive success (low or no pollen limitation). Any pollen limitation index value below − 1 was removed from the analysis (8% of the data). Pollen limitation values below − 1 are likely due to low seed production of the open-supplemented focal individual due to plant damage during the pollen supplementation process (Larson and Barrett 2000 ). Due to variations in the number of flowers per plant and the ability of many plant species to reallocate resources based on flower numbers (Knight et al. 2005 ), we determined the average individual seed production by dividing the total viable seed production by the number of individual flowers. However, the number of seeds per flower could not be quantified for GORO, as its fruits opened once collected inside the paper collection bag. We used the uncorrected total number of seeds produced per individual for this species to calculate the pollen limitation index. Landscape context To determine how different landscape features impact seed production and pollen limitation, we examined how plant reproduction varies with landscape variables such as patch size (area of each woodland remnant calculated from satellite imagery in QGIS 3.22; km 2 ), adjacent crop type (canola or wheat/pasture), and landscape position of each transect inside the reserve (edge or core; to determine an edge effect). Local variables We conducted neighbourhood floral and microhabitat surveys to explore how different local environmental factors can affect focal plant species’ fecundity and pollination dynamics. At a plot level, the identity and abundance of all flowering plants were recorded, as well as the percentage cover by grasses, leaf litter, woody debris, bare ground, and shade. Floral sampling was conducted once during the peak flowering period (e.g., late August to mid September). Using these variables, we generate a principal component analysis (PCA) per focal species to identify critical environmental axes that best explained variation in each species immediate neighbourhood. By using local environmental PCAs, we also reduced the number of collinearities between local environmental factors in our analyses. The first two principal component axes (PCA1 32% and PCA2 25%) explained 57% of the variation in the local environmental factors (Fig. S1 ). PCA1 describes a gradient from high leaf litter cover (negative values) to exposed bare ground (positive values). PCA2 describes of environmental conditions with high canopy shade (positive values) and high woody debris cover (negative values). Pollinator diversity Potential pollinators were sampled using blue vane traps to determine the abundance and diversity of potential pollinators available to plant species at each transect. This passive insect trapping method has effectively attracted native bee and non-bee insects like flies, wasps, and beetles (Hall 2018 ; Hall and Reboud 2019 ). As they are passive, they also vastly increase the sampling time possible compared to hand netting. Vane traps consist of a collecting jar, a top funnel and two interconnected blue ‘vanes’. Traps are suspended at 1m from the ground and filled with a soapy water solution. Our traps were placed in the middle of each transect for two weeks during the canola’s peak bloom. After sampling, insects were collected and preserved in 70% alcohol for (morphospecies) identification. The transect scale pollinator diversity index was calculated using the Hill-Shannon diversity index (Roswell et al. 2021 ). This diversity index assesses the community by leveraging species richness by abundance to give a less biased assessment of diversity than just richness. Plant diversity To assess plant diversity in each site, a 5 x 5 m square plot was established in the middle of each study transect, resulting in two community-level neighbourhood samples per natural remnant (edge and core). For the two remnants that were bordering the two different crop types (Buntine and Unnamed WA12427), we had four transects (two edge and core sites). The identity and percentage cover of each plant species were recorded, as well as the same variables as in the plots-level scale. The plant diversity index was calculated using the Hill-Shannon diversity index per transect. Statistical analysis All statistical analyses were conducted in R Studio using R version 4.2.3 (R Core Team, 2023). To answer our core research questions, we first checked that our focal plant species rely on insect pollinators for seed production. To determine this we conducted a negative binomial generalised linear model with log-link function glmer.nb using the MASS package (Ripley et al. 2023 ). The number of seeds per plant was the response variable, and pollination treatment (open, open and supplemented, bagged, and bagged and supplemented) was the explanatory variable. Site was included as a random factor to account for natural variance between sites. This was followed by a multiple comparison analysis between treatments with glht function of the multcomp package (Hothorn et al. 2023 ). How is bee community and community composition affected by the landscape context (remnant size, proximity to crops, and type of neighbouring crops)? Given that the majority of insects and potential pollinators captured in the study were bees (60%; both native and exotic European honeybees), the following analyses were conducted considering only the abundance and diversity of bee species. To understand how native bees, honeybees, and all bees (combined native bee and honeybee) abundance are influenced by landscape context, we used a generalised linear model glm function with a Poisson distribution. We included bee abundance as the response variable, and neighbouring crop (canola vs wheat), landscape position (core vs edge), and remnant size as explanatory variables. We used a separate generalised linear model with Gaussian distribution, utilising the same landscape variables, to assess the relationship between bee and plant diversity within landscape context. Lastly, we assessed bee composition by calculating the dissimilarity of native bee species identities and abundances between remnants using beta.multi (using presence-absence) and beta.multi.abund (species abundance) functions from the betapart package (Baselga and Orme 2012 ). Does the interplay between landscape context, pollinator community and plant community structure modulate the reproduction of native annual wildflower species in York Gum-Jam remnants? Seed production and landscape context, local factors, and pollinators To evaluate the direction and strength of the relationships between seed production (response) and multiple landscape and local predictors, we designed a species-specific piecewise structural equation model (SEM) using psem function in “picewiseSEM” package (Lefcheck 2016 ). SEMs comprise linear models organised in a causal network, assessing direct and indirect effects among multiple variables within complex systems. First, a unique SEM was developed for each focal species from a conceptual model including all possible predictors (Fig. 3 ): landscape variables (neighbouring crop, landscape position, the interaction of the previous two variables, and patch size), local variables (PCA1, PCA2, floral abundance, plant diversity), and all bee abundance. We used bee abundance instead of bee diversity because it responds to the different landscape components, but not bee diversity (see results section). This SEM was composed of two foundational models; (i) a negative binomial generalised linear model with log-link function ( glm.nb) evaluating the relationship between seed production with the species-specific model, and secondly and (ii) a generalised linear model ( glm ) with a Poisson distribution linking bee abundance with landscape components and their effect on seed production. To simplify the general conceptual model, we used a model selection process to identify and the most relevant local variables for each focal plant species. We did this using the dredge function of the package “MuMIn” (Bartoń 2023 ). This automated model selection process ranks all alternative models, including subsets of explanatory variables and different levels of complexity from an intercept-only model to the global model, using AIC (Akaike Information Criterion) values. We also explored whether bee diversity better predicted seed production than bee abundance. Except for POGN, bee abundance was a better predictor than bee diversity (AIC < 2), so we focus on results from the bee abundance models in our results. To evaluate the removal or addition of any variable to the main SEM and model fit, we tested the model fit using X 2 -test , Fisher's C statistic, and AIC values. Individual model assumptions and collinearity between variables (variation inflation factors; VIF) were checked outside the SEM with the check_model function in the “performance” package (Lüdecke et al. 2021 ). Pollen limitation and landscape context, local factors, and pollinators To test whether landscape and local context affected pollen limitation, we designed a model with the landscape context variables (crop type, landscape position, and patch size), bee abundance, plot-level plant diversity and floral density (number of total flowers inside the plot). Even though bee diversity is important for pollination dynamics (Albrecht et al. 2012 ), it was not included in the model because it is highly correlated with bee abundance ( r 18 = 0.56, p = 0.01). Crop and transect interaction was not included in the model because it was not significant, and models excluding this interaction had lower AIC values (< 2). Results Pollinator dependency For all focal species, restricting pollinator access to flowers (i.e. bagged treatment) resulted in reduced seed set per plant compared to those with access to natural pollination (open treatment; Table S2). The pattern of lower seed production in the absence of insect pollinators was more prominent for the three Asteraceae species, with seed production 76.4% lower for L awrencella rosea , 82% lower for Podolepis aristata and 71.9% lower for Podotheca gnaphaloides when pollinators were excluded. Goodenia rosea produced 26.6% fewer seeds on average than when pollinators were present (Table S3). Hand pollen supplementation increased seed production in GORO by 14%, LARO by 7.5%, POGN by 6.3%, but decreased seed production in POAR by 9.6% (Table S3). Open and open-supplemented treatments were not significantly different for any focal species, a pattern that was driven by high variation among individuals in these treatments. Focal plants in the bagged-supplemented treatment produced an intermediate, yet significantly higher number of seeds compared to bagged and open plants (Table S2), showing the receptiveness of focal plants to our hand pollination treatment. Relationship between bee diversity and abundance, plant diversity and landscape context A total of 597 insects were captured using the blue vane traps. This included 358 bees, 119 flies, 107 beetles, and 12 wasps (Table S4). Native bee species comprised only 30% of all bee captures (107 individuals), with honeybees ( Apis mellifera ) accounting for 70% of captured bees (251 individuals) and representing the most common potential pollinator across all study reserves. In total, 32 native bee morphospecies were collected across all remnants. The genus Leiproctus (Colletidae) was the most abundant and species rich group of bees. Bee and plant diversity (Hill-Shannon index) were not affected by the crop type growing adjacent to each remnant, landscape position, or patch size (Table S5). Bee and plant diversity were not correlated ( p = 0.895). The beta-diversity of native bee species across sampled remnants was very high (0.899, 0.933; using presence or abundance data), with very low nestedness of bee species (0.056, 0.026), and very high species turnover (0.955, 0.959). This indicates that native bee community composition varies substantially between sites with very few shared species between remnants. Overall, all bee abundance (native bees and honeybees pooled) was higher in remnants adjacent to canola, especially at the core of the remnants. Both native bees and honeybees had similar responses to landscape composition and position (Fig. 4 ; Table S6), with the abundance of both groups higher in reserves adjacent to canola than adjacent to wheat. This pattern was strongest for core transects. We found a significant interaction between crop and transect for all bees, indicating that bee abundance was greater at the core than at the edge, adjacent to canola. Bee abundance was similar between core and edge when measured adjacent to wheat/pasture. Patch size had a marginally negative effect on all bee abundance ( p = 0.063; Table 6). Seed production and landscape context, local predictor, and pollinator abundance Across all species, seed production was influenced by landscape components, local-level predictors, and bee abundance. Although plant species' responses were species-specific, they exhibited similar trends. GORO and POGN had a higher seed set in canola-adjacent remnants located at the core (Fig. 5 ). Both species had a similar reproductive response to crop and landscape position interaction, with higher seed set at the edge in canola-adjacent remnants, but a higher seed set at the core in wheat-adjacent remnants. LARO and POAR reproductive output was indirectly influenced by crop type and their position in the remnant, mediated by changes in bee abundance (Fig. 5 ). The seed set of these two species was positively influenced by bee abundance. GORO’s SEM did not support the direct link between bee abundance and seed production, but no significant effect was found when tested outside the SEM ( p = 0.820). Patch size (area) had direct and indirect negative relations with seed set on three of the four species: GORO, LARO and POAR, where seed set per plant was higher in smaller reserves. The link between patch size and seed production was not supported in POGN’s SEM, but patch size did not affect seed production ( p = 0.920) when tested outside the SEM. Various local-scale factors influenced seed production of the focal species (Fig. 5 ); LARO reproductive output declined with increasing PCA1 (bare ground cover), showing higher seed sets in leaf litter-covered plots. POGN fecundity was positively correlated with floral abundance, exhibiting an increased seed set in plots with more floral abundance. GORO's best model included a non-significant but positive effect with PCA2 (shade), suggesting better reproductive output in less sun-exposed plots. POAR best explanatory model also included a non-significant but positive trend between seed production and PCA1, with focal plants producing more seeds in plots with higher bare ground percentages. Pollen limitation and landscape context, local predictor, and pollinator abundance When controlling for flower number, experimental pollen supplementation increased seed set of all focal species when compared with plants only exposed to natural pollination. All species experienced a similar degree of pollen limitation; with open-supplemented plants producing 9 ~ 15% more seeds per flower than open-treatment plants ( PLi : LARO 0.09, POAR 0.14, POGN 0.14, and GORO 0.15). Different environmental variables influenced pollen limitation for each species (Table 1 ); LARO focal plants located at the edge of remnants and within dense floral plots experienced stronger pollen limitation. POAR exhibited stronger pollen limitation in larger remnants and in those with lower bee abundance. POGN’s pollen limitation decreased in plots with higher floral abundance. GORO pollination dynamics were not affected by any of the landscape or pollinator predictors. Table 1 Results of the GLM of each species with pollen limitation index as response variable and crop (canola), location in the reserve (transect), patch size, plant diversity index (Hill-Shannon index; HSI), bee abundance and floral abundance as independent predictors. Focal species Predictor Estimate Std. error Z p GORO Canola -0.03 0.16 -0.18 0.861 Transect (Core) -0.01 0.09 -0.15 0.884 Patch size -0.03 0.03 -0.92 0.358 Plant diversity (HIS) -0.06 0.06 -0.94 0.349 Bee abundance 0.001 0.01 0.58 0.565 Floral abundance 0.001 0.001 0.30 0.763 LARO Canola 0.15 0.1 1.52 0.130 Transect (Core) -0.20 0.08 -2.38 0.019 Patch size 0.001 0.002 0.88 0.379 Plant diversity (HIS) 0.01 0.04 0.23 0.816 Bee abundance 0.001 0.005 -0.58 0.561 Floral abundance 0.001 0.001 2.39 0.018 POAR Canola 0.14 0.11 1.29 0.200 Transect (Core) 0.11 0.08 1.3 0.197 Patch size 0.01 0.003 2.19 0.032 Plant diversity (HIS) -0.01 0.04 -0.19 0.852 Bee abundance -0.01 0.005 -2.3 0.024 Floral abundance -0.001 0.002 -0.74 0.461 POGN Canola -0.08 0.08 -1.09 0.278 Transect (Core) -0.07 0.08 -0.82 0.414 Patch size -0.002 0.002 -0.98 0.332 Plant diversity (HIS) -0.04 0.04 -0.96 0.340 Bee abundance 0.001 0.001 -0.12 0.907 Floral abundance 0.001 0.001 -2.5 0.014 Discussion We show that agricultural practices influence plant reproductive success through multiple mechanisms driven by insect pollinators and the environment. Crop type adjacent to woodland remnants substantially alters bee abundance, independent of bee diversity, indirectly impacting native annual plant seed production. Complex interactions between local and landscape factors and the pollinator community were important in mediating seed output but with the direction and strength of the effect being species specific. Our results provide valuable insights into the mechanisms by which multiple co-occurring environmental factors characterised at different spatial scales can influence pollinators, pollination dynamics, and plant reproduction in heavily managed agricultural landscapes. Direct and indirect effects of landscape components on bee abundance and diversity Crop type mediates the abundance but not diversity of bees in adjacent natural remnants. Bee abundance was greater in remnants adjacent to canola than those adjacent to wheat, being attracted to the rich floral resource provided by the canola during bloom (Holzschuh et al. 2013 ). This shows that large-scale canola crops act as pollinator magnets in this system, attracting bees to the adjacent remnants. Changes in bee abundance between crop types were mainly driven by the abundance of one species, feral European honeybees (there are no domestic managed hives in this landscape), in the different remnants, with native bee abundance less affected by adjacent crop type (Reynolds et al. 2022 ). Interestingly, bee diversity was similar across all sites and varied independently of both local and landscape floral resources. This indicates that although native bees utilize floral resources from canola crops during the flowering period, they are constrained to reserves where they are found and mostly influenced by within reserve resources and features (Kennedy et al. 2013 ; Martínez-Núñez et al. 2022 ). In contrast, honeybees have a high dispersal capacity with large foraging ranges (Beekman and Ratnieks 2000 ; Greenleaf et al. 2007 ; Danner et al. 2016 ). Extensive crop areas, like those in the landscapes where this study was conducted, are less likely to act as a dispersal barrier for honeybees. Therefore, native bees are more dependent on local floral resources and cannot respond as strongly as honeybees to spatially-explicit temporal crop resources such as canola (Gathmann and Tscharntke 2002 ; Grundel et al. 2010 ; Kennedy et al. 2013 ). The degree of crop type impacts on bee abundance is dependent on proximity to the remnant edge, with higher bee abundance at the core of canola-adjacent remnants compared to the edge. In contrast, bee abundance remained similar in both transect locations in wheat-adjacent remnants. We show that canola fields attract pollinators away from remnant edges (Holzschuh et al. 2016 ). The core of remnants, however, seemed to be buffered from this pollinator dilution. Compared to the edge, the remnant core might provide more abundant and diverse native floral resources (Hofmeister et al. 2013 ; Schöpke et al. 2023 ), attracting and concentrating bees and other pollinators (Nicholson et al. 2019 ). This pollination dilution and pollinator competition between resident plants and crops has complex implications for native plants in agricultural landscapes (Cussans et al. 2010 ; Diekötter et al. 2010 ; Holzschuh et al. 2011 ; Qiu et al. 2023 ). Direct and indirect effects of landscape components and bee abundance on plant reproduction Plant reproductive responses to agricultural and landscape components can emerge from diverse mechanisms, as illustrated by our structural equation models. Despite the dilution of pollinators at reserve edges, canola increased seed production of our focal annual plants in adjacent remnants compared to wheat adjacent remnants. Canola indirectly increased seed set mediated by changes in bee abundance for two of our plant focal species. Lawrencella rosea and P. aristata seed production was directly linked to bee abundance, resulting in increased seed set in remnants with higher bee abundance. Our results are consistent with the idea that canola increases the attractiveness of a landscape to many bees, increasing overall bee abundance in adjacent areas (Hanley et al. 2011 ; Holzschuh et al. 2016 ). These results align with prior research looking at the effect of crop type on native plants in semi-natural habitats (Cussans et al. 2010 ; Kovács-Hostyánszki et al. 2013 ). Insect-attracting crops such as canola can facilitate pollination of co-flowering native plant species in adjacent remnants, compared with non-insect attracting crops such as cereals (e.g., wheat) (Cussans et al. 2010 ; Kovács-Hostyánszki et al. 2013 ). For our other two focal plant species ( G. rosea and P. gnaphaloides ), canola directly increased reproductive success through mechanisms other than changes in bee abundance, as the link from crop to bee abundance to seeds was not significant or supported by the SEMs. Despite the absence of a direct link, the increase in seed production in these two species likely resulted from changes in pollination behaviour and visitation in canola-adjacent remnants (Bänsch et al. 2020 ; Yourstone et al. 2021 ; Qiu et al. 2023 ). Although G. rosea had a high capacity for self-fertilization, its reproductive success did significantly increase when it had access to pollinators. Thus, even low pollinator densities may have provided sufficient pollination services due to self-fertilization mechanisms (Aarssen 2000 ). Conversely, mass-flowering canola crops might be facilitating P. gnaphaloides pollination through pollinator spill-over from crops to natural remnants, as has been noted for some shrub species (Kovács-Hostyánszki et al. 2013 ) and a perennial herb (Qiu et al. 2023 ). While the mechanisms for these direct links between crop type and seed production were not identified by our study, likely drivers are interactions with herbivores, parasites, or competition for pollinators with other plant species in this system (Blitzer et al. 2012 ). This result presents an avenue for further investigation between different mutualistic and antagonistic interactions between crops and native plants in agricultural landscapes (e.g. Chamberlain et al. 2013 ). In contrast with our findings, the presence of large-scale canola has previously been found to have competitive effects on neighbouring plants near remnants and reserves in canola landscapes elsewhere in the world (Van Reeth et al. 2019 ). For instance, Holzschuh et al. ( 2011 ) found a landscape-scale competitive effect of canola on Primula veris with the pollination services of this species reduced when canola crops were presence in the landscape in Germany. Despite pollinator abundances increasing in the remnants adjacent to canola, the seed set of P. veris was reduced due to pollinator dilution of the main pollinator, bumblebees ( Bombus spp. ). Interestingly, despite our study revealing similar pollinator trends, the outcomes of these trends in our system were different. Differences in plant seed production linked to adjacent crop type can be attributed to ecosystem type, the identity of pollinators, and the life strategies of our focal plants. Unlike previous studies, ours is the first to explore the effects of crop type on the seed production and pollination services of native annual plants in remnant reserves, while also linking these effects with different environmental factors at a landscape and local context. Most annual plants have generalist pollination reproductive strategies and are able to maximise seed production from a small number of pollinator visits. This is essential, given their single, short flowering period and the stochastic nature of the pollination environment between years (Munoz et al. 2016 ). Because of this life history strategy, annual plants may be more likely to benefit from temporary increases in pollinator abundances caused by mass flowering crops. This effect may explain the strong indirect benefits canola was found to have in our system (Qiu et al. 2023 ). Patch size has been reported to have a positive relationship with bee abundance and plant reproduction in other systems (Hadley et al. 2014 ; Blaauw and Isaacs 2014 ; Rahimi et al. 2022 ), but our study revealed a different trend. We found that plant reproduction was lower in larger remnants than in small remnants. This counterintuitive finding was also identified by (Lázaro et al. 2020 ), who found that two perennial plant species had lower seed set in larger remnants. This negative relationship may be caused by a decline in pollinator visits across larger areas, due to floral resources scattered over larger areas in larger remnants, and potential barriers to pollinator movement in large remnants, as suggested by Lázaro et al. ( 2020 ). Pollinator populations are typically limited by nesting habitats and floral resources, which are commonly heterogeneously distributed across habitat patches (Reverté et al. 2019 ). In our study system, floral resources are highly concentrated in patches of certain woodland types (e.g., York Gum-Jam Woodlands) that typically occupy only a part of the whole remnant (see Fig. 1 b). This results in a heterogeneous distribution of pollinators. These pollinator clusters provide uneven pollination services throughout these remnants (Reverté et al. 2019 ; Escobedo-Kenefic et al. 2020 ; Martínez-Núñez et al. 2022 ). In smaller remnants, pollinators might be spatially constrained and have fewer floral options, increasing the visitation rate of local flowers. Effect of local-scale components on pollination dynamics Canola flowers are acting as pollinator magnets, bringing pollinators into adjacent reserves where pollinators can nest. However, local conditions and resources influence their foraging behaviours. We show that local factors influence pollination dynamics, with local bee and floral abundances having mixed consequences for plant pollen limitation. As plants are well known to vary in their reliance on insect pollinators, this species-level variation is not a surprise. However, the evidence of consequences for annual plant seed production across four arbitrarily selected species in this system suggests that across these specious wildflower communities, adjacent crop type is likely an important mechanism for plant fitness in vegetation remnants in these landscapes. This may then have further trickle-down effects on community diversity and structure. This study shows the complex dynamics of multi-scale factors in plant-pollinator interactions. While our work explores aspects of landscape composition, our results highlight that more work is needed to resolve how interannual climatic variation affects the population dynamics of native bees and plants. Fragmentation appears to have disrupted native pollinator communities in this region, with low diversity in each reserve and high species turnover between reserves. However, we currently lack a baseline to understand the population and diversity of native bees throughout this floral biodiversity hotspot. In particular, further research is needed to understand how landscape-level factors and competition for floral resources with honeybees affects the diversity, abundance and dispersion of native bees (e.g., Prendergast et al. 2021 ). Despite depauperate native bee communities, we found that pollination services of both crops and native flora are being complemented by the feral honeybee (Hung et al. 2018 ). This may become even more important with the recent introduction of varroa mites to Australia which are likely to decimate managed and feral honeybee populations (Iwasaki et al. 2015 ). As the mites spread across the country, the reliance on native bees for pollination services in natural vegetation is likely to increase, introducing a new type of disturbance to this vulnerable ecosystem. Conclusion Despite the vast research on the benefits of pollinator spillover from natural remnants to crop fields, little is known about the consequence of pollinator spillover from agricultural fields to remnants of native vegetation. This study represents one of the first to explore the complex reproductive responses of annual native plants to co-occurring landscape fragmentation, agricultural practices, and local plant community attributes. We found that crop identity can directly and indirectly modify plant-pollinator interactions in adjacent remnants, and these effects extend beyond the remnant edge to the core. Large-scale canola crops adjacent to remnants resulted in a direct increase in bee abundance but not bee richness, and an indirect increase in native plant seed production, compared to non-insect rewarding crops such as wheat. Yet, canola's effect on bee abundance depended on the proximity to the remnant edge and pollinator group, mainly increasing feral honeybees’ abundance. We explored the possible mechanisms in which crop type adjacent to the remnant affects pollination dynamics through pollination experiments, suggesting that, while landscape-scale factors directly influenced bee abundance and plant reproduction, local-scale landscapes elements also influence pollination dynamics and bee diversity. Our study helps unravel how different aspects of the landscape simultaneously affect the abundance of pollinators, their behaviour, and the implications for the persistence of native plant populations in isolated remnants. Declarations Competing Interests The authors declare no competing interests. Funding This project was funded by the Australian Research Council (DP210100913) and Botany Foundation at the University of Melbourne (G.A.M. Scott Research Award, 2022). Author Contribution MS and MMM contributed to the study's conceptualisation, development, and experimental design. MS collected the field data with assistance from ZCW. MS conducted the analysis with assistance from ZCW and MMM. All authors contributed to the interpretation of the results. MS wrote the first draft of the manuscript, and ZCW and MMM contributed significantly to its revision. Acknowledgement We acknowledge the Yamatji People as the traditional owners of the land on which we conducted this study. We pay our respects to their Elders past and present. Thank you to Lisa Buche and Hanlun Liu for their help with fieldwork, statistical advice, and manuscript comments. We also thank various Mayfield Lab members for manuscript comments and Yulin He for technical support. Data Availability The data used in this study will be made available in an online repository following manuscript acceptance. References Aarssen LW (2000) Why are most selfers annuals? 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4756788","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":340681227,"identity":"210b2e43-478d-470d-8069-a20310ae7ad7","order_by":0,"name":"Manuel Sevenello","email":"data:image/png;base64,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","orcid":"","institution":"University of Melbourne","correspondingAuthor":true,"prefix":"","firstName":"Manuel","middleName":"","lastName":"Sevenello","suffix":""},{"id":340681228,"identity":"94b60314-ade6-491f-adbf-73803dd4904f","order_by":1,"name":"Zac C. Walker","email":"","orcid":"","institution":"University of Melbourne","correspondingAuthor":false,"prefix":"","firstName":"Zac","middleName":"C.","lastName":"Walker","suffix":""},{"id":340681229,"identity":"c3a47505-1a19-4ddb-9204-e8d762bfa820","order_by":2,"name":"Margaret M. Mayfield","email":"","orcid":"","institution":"University of Melbourne","correspondingAuthor":false,"prefix":"","firstName":"Margaret","middleName":"M.","lastName":"Mayfield","suffix":""}],"badges":[],"createdAt":"2024-07-17 14:05:18","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4756788/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4756788/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10980-026-02305-2","type":"published","date":"2026-02-04T15:59:18+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":62642309,"identity":"d4d8a740-a5bb-4cfd-bf65-cf9eb2869c0e","added_by":"auto","created_at":"2024-08-16 19:22:31","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":809133,"visible":true,"origin":"","legend":"\u003cp\u003eMap of the study region, with a close-up of one of the study remnants and examples of environmental conditions at the two study landscape positions: edge and core. \u003cstrong\u003ea)\u003c/strong\u003eMap showing the location of the eleven sites across nine YGJW remnants in southwest Western Australia. The map inset shows the location of the study region within Australia. Yellow dots represent remnants adjacent to canola and green dots represent remnants adjacent to wheat or pasture. The white square shows a close-up of Milton Nature Reserve. \u003cstrong\u003eb)\u003c/strong\u003e Map of the Milton Nature Reserve, different colors within the remnants correspond to different woodland types. The light green rectangle shows the position of the transect located at the edge and the orange rectangle shows the position at the core of the remnant.\u003c/p\u003e","description":"","filename":"Fig.1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4756788/v1/281ae17c569b2083098ad1bb.jpg"},{"id":62642626,"identity":"9785bade-820b-45fc-add1-fff467e8e334","added_by":"auto","created_at":"2024-08-16 19:30:31","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1521738,"visible":true,"origin":"","legend":"\u003cp\u003eFocal plant species a) \u003cem\u003eGoodenia rosea\u003c/em\u003e (GORO), b) L\u003cem\u003eawrencella rosea\u003c/em\u003e(LARO), c) \u003cem\u003ePodolepis aristata\u003c/em\u003e (POAR), and d) \u003cem\u003ePodotheca gnaphaloides\u003c/em\u003e(POGN).\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-4756788/v1/b413b71c338b6dcac4c8aa6b.png"},{"id":62642306,"identity":"dda2290f-ea19-4840-b4b3-491abac871a9","added_by":"auto","created_at":"2024-08-16 19:22:31","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":132839,"visible":true,"origin":"","legend":"\u003cp\u003eThe conceptual structural equation model (SEM) showing all possible relationships between seed production and all examined predictors. The two underlying models are specified in different colours, black arrows depicting the relationships between seed production, environmental factors, and bee abundance. Blue lines show relationships between bee abundance and landscape components. The thicker line shows the relation between bee abundance and seed set. The direction of the arrows indicates the direction of the relationships.\u003c/p\u003e","description":"","filename":"Fig.3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4756788/v1/0acc064cd467e759f117a788.jpg"},{"id":62642971,"identity":"2a4bb196-8ed1-46cf-81b3-670ad9c6e03d","added_by":"auto","created_at":"2024-08-16 19:38:31","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":76802,"visible":true,"origin":"","legend":"\u003cp\u003eAbundance responses of all bees (orange; native and honeybees pooled together), honeybees (yellow), and native bees (red) to crop type and location in the reserve. The asterisks above the bars indicate significant differences between transects or crop types.\u003c/p\u003e","description":"","filename":"Figure4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4756788/v1/cf917e0488dda84c2deeefb0.jpg"},{"id":62642310,"identity":"3ff221d9-eec7-4429-b271-606fe0d57382","added_by":"auto","created_at":"2024-08-16 19:22:31","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":621858,"visible":true,"origin":"","legend":"\u003cp\u003eResults of the structural equation model (SEM) for each focal species showing the positive and negative interactions between landscape context (crop: canola, position: core), local predictors, pollinator abundance and plant reproduction. Solid lines represent significative interactions while dashed lines non-significant. Black and red show positive and negative, respectively. The thickness of each arrow shows the relationship strength (standardized path coefficients). Crop and transect interaction effect on seed production is presented next to each species SEM. The crop x position graph shows the effect (adjusted marginal mean and 95% confidence intervals) of the interaction between crop and transect, with the y-axes representing the seed production and the x-axes crop type and transect location.\u003c/p\u003e","description":"","filename":"Fig.5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4756788/v1/d1b97dba851b6e89cac8c6d2.jpg"},{"id":102235915,"identity":"11466d36-6f92-470a-a048-ce629db8ed73","added_by":"auto","created_at":"2026-02-09 16:18:21","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4847809,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4756788/v1/3b194666-27be-42af-81f8-f2c5abfd011c.pdf"},{"id":62642311,"identity":"ac6eff51-8eca-41f0-bf90-f12c7b5506dd","added_by":"auto","created_at":"2024-08-16 19:22:31","extension":"docx","order_by":7,"title":"","display":"","copyAsset":false,"role":"supplement","size":287327,"visible":true,"origin":"","legend":"","description":"","filename":"landscapeecologysupplementaryMS.docx","url":"https://assets-eu.researchsquare.com/files/rs-4756788/v1/4e306b8f68cd94a10f97b1b3.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Landscape fragmentation and agricultural context impact pollination dynamics of native annual plants","fulltext":[{"header":"Introduction","content":"\u003cp\u003eLand-use intensification and landscape fragmentation negatively impact biodiversity (Fahrig \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). This has been shown to occur through direct processes such as population reduction and isolation (Honnay et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Lino et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and indirect processes such as disruptions to species interactions (Mustaj\u0026auml;rvi et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Ferreira et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). While these direct processes are well understood (Fahrig \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Wilson et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and we have a theoretical understanding of indirect effects (P\u0026uuml;ttker et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), we still lack a strong empirical understanding of how indirect mechanisms drive specific consequences of land-use intensification on native species and their persistence in habitat remnants (but see L\u0026aacute;zaro et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Multi-trophic and mutualistic interactions, such as plant-pollinator interactions, are critical for maintaining biodiversity (Allen-Wardell et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Wei et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). They are well known to break down in isolated habitat remnants and their disruption can lead to changes in ecosystem function and services in fragmented landscapes (Viana et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Hadley and Betts \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eNatural vegetation remnants within agricultural landscapes are critical for biodiversity conservation and ecosystem services (Soga et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Decocq et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Resident plants and insect pollinators in remnants are often highly dependent on each other, yet fragmentation isolates them from other populations, limiting the extent of pollination services in fragmented landscapes (Harris and Johnson \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Aguilar et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Kolb \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Disruptions to plant-pollinator interactions can occur in remnants when the temporal and spatial distribution of interacting partners are restricted (e.g., due to isolation) or when the interacting species respond differently to the novel environmental conditions (e.g., edge effects; Hadley and Betts \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePollinator spillover from remnant vegetation into agricultural fields can greatly benefit crop production (Garibaldi et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Pollinators, particularly bees, nest in native vegetation patches and forage on crop flowers in adjacent farmlands and native plants within remnants (Potts et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). This pollination service mediates seed and fruit production for both crops and native plants (Garibaldi et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). While the beneficial effect of pollinator spillover on crop systems is well understood, we have limited knowledge of how this spillover affects within-remnant species interactions and native plant reproduction, despite its known importance for maintaining the conservation value of these remnants (Blitzer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Geslin et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eInsect-attracting mass flowering crops, such as canola (\u003cem\u003eBrassica napus\u003c/em\u003e, Brassicaceae), share pollinators with co-flowering native plants in many agricultural systems (Stanley and Stout \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Reynolds et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Canola crops are known to sometimes increase the abundance of pollinators in adjacent remnants due to the presence of this abundant but ephemeral food resource (Holzschuh et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Bailey et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Other studies, however, have shown that the presence of abundant canola flowers can alternatively lead to pollinator dilution, with pollinators attracted to crop flowers over native flowers, especially at peak flowering when canola is the most abundant, if not the highest quality, resource available (Holzschuh et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Reynolds et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This shift of pollinator foraging is known to lower the reproductive output of co-flowering native plants due to deficient pollinator services (Knight et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Indeed, in some landscapes dominated by canola, canola has been shown to have negative effects on native plant reproduction despite increased pollinator abundances in the landscape as a whole (Holzschuh et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Van Reeth et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). While pollinator dilution is often invoked to explain this effect, few studies have attempted to determine the mechanisms by which mass flowering crops like canola affect native plant seed production in vegetation remnants in agricultural landscapes, with most assuming rather than showing a relationship (but see Stanley and Stout \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Ekroos et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Alternatively, interacting landscape and local scale factors, such as plant community composition and abundance, can buffer plant reproductive success from the negative impact of pollinator competition with flowering crops and landscape fragmentation (Kov\u0026aacute;cs-Hosty\u0026aacute;nszki et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Ekroos et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOur study aims to determine the biological consequences of land-use intensification on native plant population dynamics as mediated by insect pollinators. We explore this in the critically endangered York gum-Jam woodlands (YGJW) of SW Western Australia. These woodlands have lost more than 93% of their original extent due to land clearing and agricultural intensification since European invasion (Prober and Smith \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; DAWE, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Native vegetation is now found only in small isolated patches embedded within an agricultural mosaic of wheat and canola fields (Yates and Hobbs \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). Understanding the pollination dynamics of wildflower species in such a system is critical for effective management and conservation. In this study, we explore how environmental and agricultural context impacts pollinator abundance, pollination services, and the reproductive success of four native annual forb species. Specifically, we ask (i) how the pollinator community and plant diversity are affected by landscape context (patch size, edge effect, and type of neighbouring crops)? and (ii) does the interplay between landscape context, pollinator abundance, and plant community structure modulate the reproductive success of plant species in remnant woodlands?\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy system\u003c/h2\u003e \u003cp\u003eThe York gum-Jam woodlands (YGJW) of SW Western Australia are a threatened ecological community within the global biodiversity hot spot of the Southwest Australian floristic region (DAWE \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). These woodlands occur throughout the agricultural wheatbelt region and have experienced widespread clearing and fragmentation (Prober and Smith \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; DAWE \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Remnant YGJW typically occur in small, geographically isolated patches within an agricultural matrix. These woodlands are dominated by an overstorey of York gum (\u003cem\u003eEucalyptus loxophleba\u003c/em\u003e Beth. Subsp. Loxophleba) and Jam (\u003cem\u003eAcacia acuminata\u003c/em\u003e Benth) and comprise a diverse understory of shrubs, perennial grasses, and annual and perennial exotic and native forbs (Dwyer et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The climate in this region is semi-arid (350 to 450 mm rainfall per year) with cool, wet winters and warm, dry summers (Hopper and Gioia \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). These woodlands generally have an open canopy and phosphorous-poor sandy soils (Lambers et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), creating diverse microhabitats across gradients of light, litter, and soil nutrients (Dwyer et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The annual wildflower community primarily consists of Asteraceae, Goodeniaceae, and Araliaceae species that germinate during the winter (July-August) and flower and seed in spring (August-November).\u003c/p\u003e \u003cp\u003eThis study was conducted across nine YGJW remnants (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea) adjacent to either an insect pollinator-attracting crop (canola) or a non-insect-attracting crop (wheat or pasture). Both Wheat and canola bloom from July to late August in this part of Western Australia, flowering at the same time as many of the native wildflowers that live in the YGJW remnants. Sites were selected based on remnant size, accessibility, and neighbouring crop identity. Not all species were found in all reserves, and some sites did not have adequate plant abundances to perform edge and core transects in the same remnants (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). Two sites were established in the same remnant but with sites positioned more than 500 m apart. In both cases, the two sites in the same reserves were adjacent to different crop types (either canola or wheat/pasture). These sites were assumed to have independence (despite occurring in the same remnant) because the native pollinating insects in this system have small home ranges and typically travel less than 500 m to forage (Gathmann and Tscharntke \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Further, the YGJW vegetation is patchy within remnants (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb) and sites were not positioned in contiguous patches of YJGW (i.e., they are separated by patches of other woodland types that do not have wildflower understories).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eStudy species\u003c/h2\u003e \u003cp\u003eFour native annual plant species were selected for this study (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e): \u003cem\u003eGoodenia rosea\u003c/em\u003e (S.Moore) K.A.Sheph. (\u0026lsquo;GORO\u0026rsquo;, Goodeniaceae), \u003cem\u003eLawrencella rosea\u003c/em\u003e Lindl. (\u0026lsquo;LARO\u0026rsquo;, Asteraceae), \u003cem\u003ePodolepis aristata\u003c/em\u003e Benth. (\u0026lsquo;POAR\u0026rsquo;, Asteraceae), and \u003cem\u003ePodotheca gnaphaloides\u003c/em\u003e Graham (\u0026lsquo;POGN\u0026rsquo;, Asteraceae). These species were selected because they common within YGJW remnants, were found in most sites (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e) and have overlapping peak flowering with canola in the region. These species have contrasting floral morphology (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) and are visited by a diverse group of pollinators, including feral honeybees, native bees, flies, and beetles (personal observation).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003ePollination treatments and experimental design\u003c/h2\u003e \u003cp\u003eWe performed pollen limitation experiments in edge and core transects in all resident remnants to determine how landscape context impacted reproductive success mediated by insect pollinators. These experiments took place from late July to late October in 2022, during the complete flowering period of each focal species. To do this, we set up two 50 m x 5 m long transects in each remnant parallel to an adjacent crop field: one next to the remnant edge and a second at the core of the remnant, defined as at least 150 m from the crop field edge (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb). Past studies have shown this distance to be sufficient to observe differences in insect pollinators in this system (Reynolds et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Along each transect, ten 30 x 30 cm plots were established for each species. This resulted in a total of 520 plots and 2080 individual plants across all four species. For LARO, 640 individual plants were assigned to one of the four pollination treatments (160 per treatment) in a total of 8 sites, and the rest of the species (GORO, POAR, POGN) had each 480 individuals selected in a total of six different reserves each (120 per treatment).\u003c/p\u003e \u003cp\u003eWithin each plot, four individual plants of the focal species were selected. Each plant was randomly assigned one of the four pollination treatments: (i) \u0026lsquo;open\u0026rsquo; \u0026ndash; flowers remained unmanipulated and open to wild pollinators; (ii) \u0026lsquo;open-supplemented\u0026rsquo; \u0026ndash; flowers remained open to wild pollinators and were hand pollinated; (iii) \u0026lsquo;bagged\u0026rsquo; \u0026ndash; flowers were covered with a mesh bag before flowering; and (iv) \u0026lsquo;bagged-supplemented\u0026rsquo; \u0026ndash; flowers were bagged before opening and hand pollinated once flowers opened. Pollinator exclusion involved covering a flower with a mesh bag to prevent pollinators from accessing or interacting with the floral reproductive parts, and pollen supplementation consisted of manually adding pollen from different conspecific individuals in the immediate area to the focal plant. Pollen supplementation was performed by rubbing the focal plant inflorescence with three different conspecific inflorescences for the Asteraceae species or rubbing indusium (pollen-cup structure) in the case of GORO. Pollen supplementation was conducted every four to five days, and it was applied to all open flowers of each focal plant in both open and bagged-supplemented treatment. Focal plants in these two treatments were hand-pollinated at least three times for LARO and POGN (shorter flowering period) and six times for GORO and POAR (longer flowering period). At the end of the flowering season, all fruits were protected with a mesh bag and collected when matured. Across the 2080 focal plants, we collected 310,880 viable seeds.\u003c/p\u003e \u003cp\u003eWe calculated the pollen limitation index (\u003cem\u003ePLi\u003c/em\u003e) for each plot using Eckert et al., (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2010\u003c/span\u003e); \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:PLi=\\frac{Ssup-Sop}{Sop}\\)\u003c/span\u003e\u003c/span\u003e, where \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:Ssup\\)\u003c/span\u003e\u003c/span\u003e is the average individual seed production of the focal plant that was open to natural pollination, and hand-supplemented (\u0026lsquo;open supplemented\u0026rsquo;) and \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:Sop\\)\u003c/span\u003e\u003c/span\u003e is the average individual seed production of naturally pollinated flowers (open). This index calculates the percentage increase in seed production due to the supplemented treatment ranging from - \u0026infin; to 1. Positive values indicate higher reproductive success in pollen-supplemented plants (higher rates of pollen limitation), and negative values indicate that control plants (open) had higher reproductive success (low or no pollen limitation). Any pollen limitation index value below \u0026minus;\u0026thinsp;1 was removed from the analysis (8% of the data). Pollen limitation values below \u0026minus;\u0026thinsp;1 are likely due to low seed production of the open-supplemented focal individual due to plant damage during the pollen supplementation process (Larson and Barrett \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Due to variations in the number of flowers per plant and the ability of many plant species to reallocate resources based on flower numbers (Knight et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), we determined the average individual seed production by dividing the total viable seed production by the number of individual flowers. However, the number of seeds per flower could not be quantified for GORO, as its fruits opened once collected inside the paper collection bag. We used the uncorrected total number of seeds produced per individual for this species to calculate the pollen limitation index.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eLandscape context\u003c/h2\u003e \u003cp\u003eTo determine how different landscape features impact seed production and pollen limitation, we examined how plant reproduction varies with landscape variables such as patch size (area of each woodland remnant calculated from satellite imagery in QGIS 3.22; km\u003csup\u003e2\u003c/sup\u003e), adjacent crop type (canola or wheat/pasture), and landscape position of each transect inside the reserve (edge or core; to determine an edge effect).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eLocal variables\u003c/h2\u003e \u003cp\u003eWe conducted neighbourhood floral and microhabitat surveys to explore how different local environmental factors can affect focal plant species\u0026rsquo; fecundity and pollination dynamics. At a plot level, the identity and abundance of all flowering plants were recorded, as well as the percentage cover by grasses, leaf litter, woody debris, bare ground, and shade. Floral sampling was conducted once during the peak flowering period (e.g., late August to mid September). Using these variables, we generate a principal component analysis (PCA) per focal species to identify critical environmental axes that best explained variation in each species immediate neighbourhood. By using local environmental PCAs, we also reduced the number of collinearities between local environmental factors in our analyses. The first two principal component axes (PCA1 32% and PCA2 25%) explained 57% of the variation in the local environmental factors (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). PCA1 describes a gradient from high leaf litter cover (negative values) to exposed bare ground (positive values). PCA2 describes of environmental conditions with high canopy shade (positive values) and high woody debris cover (negative values).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003ePollinator diversity\u003c/h2\u003e \u003cp\u003ePotential pollinators were sampled using blue vane traps to determine the abundance and diversity of potential pollinators available to plant species at each transect. This passive insect trapping method has effectively attracted native bee and non-bee insects like flies, wasps, and beetles (Hall \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Hall and Reboud \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). As they are passive, they also vastly increase the sampling time possible compared to hand netting. Vane traps consist of a collecting jar, a top funnel and two interconnected blue \u0026lsquo;vanes\u0026rsquo;. Traps are suspended at 1m from the ground and filled with a soapy water solution. Our traps were placed in the middle of each transect for two weeks during the canola\u0026rsquo;s peak bloom. After sampling, insects were collected and preserved in 70% alcohol for (morphospecies) identification. The transect scale pollinator diversity index was calculated using the Hill-Shannon diversity index (Roswell et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This diversity index assesses the community by leveraging species richness by abundance to give a less biased assessment of diversity than just richness.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003ePlant diversity\u003c/h2\u003e \u003cp\u003eTo assess plant diversity in each site, a 5 x 5 m square plot was established in the middle of each study transect, resulting in two community-level neighbourhood samples per natural remnant (edge and core). For the two remnants that were bordering the two different crop types (Buntine and Unnamed WA12427), we had four transects (two edge and core sites). The identity and percentage cover of each plant species were recorded, as well as the same variables as in the plots-level scale. The plant diversity index was calculated using the Hill-Shannon diversity index per transect.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eAll statistical analyses were conducted in R Studio using R version 4.2.3 (R Core Team, 2023). To answer our core research questions, we first checked that our focal plant species rely on insect pollinators for seed production. To determine this we conducted a negative binomial generalised linear model with log-link function \u003cem\u003eglmer.nb\u003c/em\u003e using the MASS package (Ripley et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The number of seeds per plant was the response variable, and pollination treatment (open, open and supplemented, bagged, and bagged and supplemented) was the explanatory variable. Site was included as a random factor to account for natural variance between sites. This was followed by a multiple comparison analysis between treatments with \u003cem\u003eglht\u003c/em\u003e function of the multcomp package (Hothorn et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eHow is bee community and community composition affected by the landscape context (remnant size, proximity to crops, and type of neighbouring crops)?\u003c/b\u003e \u003c/p\u003e \u003cp\u003eGiven that the majority of insects and potential pollinators captured in the study were bees (60%; both native and exotic European honeybees), the following analyses were conducted considering only the abundance and diversity of bee species. To understand how native bees, honeybees, and all bees (combined native bee and honeybee) abundance are influenced by landscape context, we used a generalised linear model \u003cem\u003eglm\u003c/em\u003e function with a Poisson distribution. We included bee abundance as the response variable, and neighbouring crop (canola \u003cem\u003evs\u003c/em\u003e wheat), landscape position (core \u003cem\u003evs\u003c/em\u003e edge), and remnant size as explanatory variables. We used a separate generalised linear model with Gaussian distribution, utilising the same landscape variables, to assess the relationship between bee and plant diversity within landscape context. Lastly, we assessed bee composition by calculating the dissimilarity of native bee species identities and abundances between remnants using \u003cem\u003ebeta.multi\u003c/em\u003e (using presence-absence) and \u003cem\u003ebeta.multi.abund\u003c/em\u003e (species abundance) functions from the betapart package (Baselga and Orme \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eDoes the interplay between landscape context, pollinator community and plant community structure modulate the reproduction of native annual wildflower species in York Gum-Jam remnants?\u003c/b\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eSeed production and landscape context, local factors, and pollinators\u003c/h2\u003e \u003cp\u003eTo evaluate the direction and strength of the relationships between seed production (response) and multiple landscape and local predictors, we designed a species-specific piecewise structural equation model (SEM) using \u003cem\u003epsem\u003c/em\u003e function in \u0026ldquo;picewiseSEM\u0026rdquo; package (Lefcheck \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). SEMs comprise linear models organised in a causal network, assessing direct and indirect effects among multiple variables within complex systems. First, a unique SEM was developed for each focal species from a conceptual model including all possible predictors (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e): landscape variables (neighbouring crop, landscape position, the interaction of the previous two variables, and patch size), local variables (PCA1, PCA2, floral abundance, plant diversity), and all bee abundance. We used bee abundance instead of bee diversity because it responds to the different landscape components, but not bee diversity (see results section). This SEM was composed of two foundational models; (i) a negative binomial generalised linear model with log-link function (\u003cem\u003eglm.nb)\u003c/em\u003e evaluating the relationship between seed production with the species-specific model, and secondly and (ii) a generalised linear model (\u003cem\u003eglm\u003c/em\u003e) with a Poisson distribution linking bee abundance with landscape components and their effect on seed production.\u003c/p\u003e \u003cp\u003eTo simplify the general conceptual model, we used a model selection process to identify and the most relevant local variables for each focal plant species. We did this using the \u003cem\u003edredge\u003c/em\u003e function of the package \u0026ldquo;MuMIn\u0026rdquo; (Bartoń \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This automated model selection process ranks all alternative models, including subsets of explanatory variables and different levels of complexity from an intercept-only model to the global model, using AIC (Akaike Information Criterion) values. We also explored whether bee diversity better predicted seed production than bee abundance. Except for POGN, bee abundance was a better predictor than bee diversity (AIC\u0026thinsp;\u0026lt;\u0026thinsp;2), so we focus on results from the bee abundance models in our results. To evaluate the removal or addition of any variable to the main SEM and model fit, we tested the model fit using \u003cem\u003eX\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003cem\u003e-test\u003c/em\u003e, Fisher's C statistic, and AIC values. Individual model assumptions and collinearity between variables (variation inflation factors; VIF) were checked outside the SEM with the \u003cem\u003echeck_model\u003c/em\u003e function in the \u0026ldquo;performance\u0026rdquo; package (L\u0026uuml;decke et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003ePollen limitation and landscape context, local factors, and pollinators\u003c/h2\u003e \u003cp\u003eTo test whether landscape and local context affected pollen limitation, we designed a model with the landscape context variables (crop type, landscape position, and patch size), bee abundance, plot-level plant diversity and floral density (number of total flowers inside the plot). Even though bee diversity is important for pollination dynamics (Albrecht et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), it was not included in the model because it is highly correlated with bee abundance (\u003cem\u003er\u003c/em\u003e\u003csub\u003e18\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;0.56, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.01). Crop and transect interaction was not included in the model because it was not significant, and models excluding this interaction had lower AIC values (\u0026lt;\u0026thinsp;2).\u003c/p\u003e "},{"header":"Results","content":" \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003ePollinator dependency\u003c/h2\u003e \u003cp\u003eFor all focal species, restricting pollinator access to flowers (i.e. bagged treatment) resulted in reduced seed set per plant compared to those with access to natural pollination (open treatment; Table S2). The pattern of lower seed production in the absence of insect pollinators was more prominent for the three Asteraceae species, with seed production 76.4% lower for L\u003cem\u003eawrencella rosea\u003c/em\u003e, 82% lower for \u003cem\u003ePodolepis aristata\u003c/em\u003e and 71.9% lower for \u003cem\u003ePodotheca gnaphaloides\u003c/em\u003e when pollinators were excluded. \u003cem\u003eGoodenia rosea\u003c/em\u003e produced 26.6% fewer seeds on average than when pollinators were present (Table S3). Hand pollen supplementation increased seed production in GORO by 14%, LARO by 7.5%, POGN by 6.3%, but decreased seed production in POAR by 9.6% (Table S3). Open and open-supplemented treatments were not significantly different for any focal species, a pattern that was driven by high variation among individuals in these treatments. Focal plants in the bagged-supplemented treatment produced an intermediate, yet significantly higher number of seeds compared to bagged and open plants (Table S2), showing the receptiveness of focal plants to our hand pollination treatment.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eRelationship between bee diversity and abundance, plant diversity and landscape context\u003c/h2\u003e \u003cp\u003eA total of 597 insects were captured using the blue vane traps. This included 358 bees, 119 flies, 107 beetles, and 12 wasps (Table S4). Native bee species comprised only 30% of all bee captures (107 individuals), with honeybees (\u003cem\u003eApis mellifera\u003c/em\u003e) accounting for 70% of captured bees (251 individuals) and representing the most common potential pollinator across all study reserves. In total, 32 native bee morphospecies were collected across all remnants. The genus \u003cem\u003eLeiproctus\u003c/em\u003e (Colletidae) was the most abundant and species rich group of bees. Bee and plant diversity (Hill-Shannon index) were not affected by the crop type growing adjacent to each remnant, landscape position, or patch size (Table S5). Bee and plant diversity were not correlated (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.895). The beta-diversity of native bee species across sampled remnants was very high (0.899, 0.933; using presence or abundance data), with very low nestedness of bee species (0.056, 0.026), and very high species turnover (0.955, 0.959). This indicates that native bee community composition varies substantially between sites with very few shared species between remnants.\u003c/p\u003e \u003cp\u003eOverall, all bee abundance (native bees and honeybees pooled) was higher in remnants adjacent to canola, especially at the core of the remnants. Both native bees and honeybees had similar responses to landscape composition and position (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e; Table S6), with the abundance of both groups higher in reserves adjacent to canola than adjacent to wheat. This pattern was strongest for core transects. We found a significant interaction between crop and transect for all bees, indicating that bee abundance was greater at the core than at the edge, adjacent to canola. Bee abundance was similar between core and edge when measured adjacent to wheat/pasture. Patch size had a marginally negative effect on all bee abundance (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.063; Table\u0026nbsp;6).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eSeed production and landscape context, local predictor, and pollinator abundance\u003c/h2\u003e \u003cp\u003eAcross all species, seed production was influenced by landscape components, local-level predictors, and bee abundance. Although plant species' responses were species-specific, they exhibited similar trends. GORO and POGN had a higher seed set in canola-adjacent remnants located at the core (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Both species had a similar reproductive response to crop and landscape position interaction, with higher seed set at the edge in canola-adjacent remnants, but a higher seed set at the core in wheat-adjacent remnants. LARO and POAR reproductive output was indirectly influenced by crop type and their position in the remnant, mediated by changes in bee abundance (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). The seed set of these two species was positively influenced by bee abundance. GORO\u0026rsquo;s SEM did not support the direct link between bee abundance and seed production, but no significant effect was found when tested outside the SEM (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.820). Patch size (area) had direct and indirect negative relations with seed set on three of the four species: GORO, LARO and POAR, where seed set per plant was higher in smaller reserves. The link between patch size and seed production was not supported in POGN\u0026rsquo;s SEM, but patch size did not affect seed production (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.920) when tested outside the SEM.\u003c/p\u003e \u003cp\u003eVarious local-scale factors influenced seed production of the focal species (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e); LARO reproductive output declined with increasing PCA1 (bare ground cover), showing higher seed sets in leaf litter-covered plots. POGN fecundity was positively correlated with floral abundance, exhibiting an increased seed set in plots with more floral abundance. GORO's best model included a non-significant but positive effect with PCA2 (shade), suggesting better reproductive output in less sun-exposed plots. POAR best explanatory model also included a non-significant but positive trend between seed production and PCA1, with focal plants producing more seeds in plots with higher bare ground percentages.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003ePollen limitation and landscape context, local predictor, and pollinator abundance\u003c/h2\u003e \u003cp\u003eWhen controlling for flower number, experimental pollen supplementation increased seed set of all focal species when compared with plants only exposed to natural pollination. All species experienced a similar degree of pollen limitation; with open-supplemented plants producing 9\u0026thinsp;~\u0026thinsp;15% more seeds per flower than open-treatment plants (\u003cem\u003ePLi\u003c/em\u003e: LARO 0.09, POAR 0.14, POGN 0.14, and GORO 0.15). Different environmental variables influenced pollen limitation for each species (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e); LARO focal plants located at the edge of remnants and within dense floral plots experienced stronger pollen limitation. POAR exhibited stronger pollen limitation in larger remnants and in those with lower bee abundance. POGN\u0026rsquo;s pollen limitation decreased in plots with higher floral abundance. GORO pollination dynamics were not affected by any of the landscape or pollinator predictors.\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\u003eResults of the GLM of each species with pollen limitation index as response variable and crop (canola), location in the reserve (transect), patch size, plant diversity index (Hill-Shannon index; HSI), bee abundance and floral abundance as independent predictors.\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\u003eFocal species\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePredictor\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\u003eStd. error\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eZ\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\"\u003e \u003cp\u003e\u003cb\u003eGORO\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCanola\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.861\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTransect (Core)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.01\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-0.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.884\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePatch size\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.358\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePlant diversity (HIS)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.06\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\u003e-0.94\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.349\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBee abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.565\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFloral abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.763\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eLARO\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCanola\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.15\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\u003e1.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.130\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTransect (Core)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.20\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\u003e-2.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.019\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePatch size\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.379\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePlant diversity (HIS)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.816\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBee abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.005\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.561\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFloral abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.018\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ePOAR\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCanola\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\u003e0.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.200\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTransect (Core)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.11\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\u003e1.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.197\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePatch size\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.003\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.032\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePlant diversity (HIS)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.852\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBee abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.005\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-2.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.024\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFloral abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.74\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.461\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ePOGN\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCanola\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.08\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\u003e-1.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.278\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTransect (Core)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.07\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\u003e-0.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.414\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePatch size\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.332\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePlant diversity (HIS)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.96\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.340\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBee abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.907\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFloral abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-2.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.014\u003c/b\u003e\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\u003eWe show that agricultural practices influence plant reproductive success through multiple mechanisms driven by insect pollinators and the environment. Crop type adjacent to woodland remnants substantially alters bee abundance, independent of bee diversity, indirectly impacting native annual plant seed production. Complex interactions between local and landscape factors and the pollinator community were important in mediating seed output but with the direction and strength of the effect being species specific. Our results provide valuable insights into the mechanisms by which multiple co-occurring environmental factors characterised at different spatial scales can influence pollinators, pollination dynamics, and plant reproduction in heavily managed agricultural landscapes.\u003c/p\u003e \u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003eDirect and indirect effects of landscape components on bee abundance and diversity\u003c/h2\u003e \u003cp\u003eCrop type mediates the abundance but not diversity of bees in adjacent natural remnants. Bee abundance was greater in remnants adjacent to canola than those adjacent to wheat, being attracted to the rich floral resource provided by the canola during bloom (Holzschuh et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). This shows that large-scale canola crops act as pollinator magnets in this system, attracting bees to the adjacent remnants. Changes in bee abundance between crop types were mainly driven by the abundance of one species, feral European honeybees (there are no domestic managed hives in this landscape), in the different remnants, with native bee abundance less affected by adjacent crop type (Reynolds et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Interestingly, bee diversity was similar across all sites and varied independently of both local and landscape floral resources. This indicates that although native bees utilize floral resources from canola crops during the flowering period, they are constrained to reserves where they are found and mostly influenced by within reserve resources and features (Kennedy et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Mart\u0026iacute;nez-N\u0026uacute;\u0026ntilde;ez et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In contrast, honeybees have a high dispersal capacity with large foraging ranges (Beekman and Ratnieks \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Greenleaf et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Danner et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Extensive crop areas, like those in the landscapes where this study was conducted, are less likely to act as a dispersal barrier for honeybees. Therefore, native bees are more dependent on local floral resources and cannot respond as strongly as honeybees to spatially-explicit temporal crop resources such as canola (Gathmann and Tscharntke \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Grundel et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Kennedy et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe degree of crop type impacts on bee abundance is dependent on proximity to the remnant edge, with higher bee abundance at the core of canola-adjacent remnants compared to the edge. In contrast, bee abundance remained similar in both transect locations in wheat-adjacent remnants. We show that canola fields attract pollinators away from remnant edges (Holzschuh et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The core of remnants, however, seemed to be buffered from this pollinator dilution. Compared to the edge, the remnant core might provide more abundant and diverse native floral resources (Hofmeister et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Sch\u0026ouml;pke et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), attracting and concentrating bees and other pollinators (Nicholson et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). This pollination dilution and pollinator competition between resident plants and crops has complex implications for native plants in agricultural landscapes (Cussans et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Diek\u0026ouml;tter et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Holzschuh et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Qiu et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003eDirect and indirect effects of landscape components and bee abundance on plant reproduction\u003c/h2\u003e \u003cp\u003ePlant reproductive responses to agricultural and landscape components can emerge from diverse mechanisms, as illustrated by our structural equation models. Despite the dilution of pollinators at reserve edges, canola increased seed production of our focal annual plants in adjacent remnants compared to wheat adjacent remnants. Canola indirectly increased seed set mediated by changes in bee abundance for two of our plant focal species. \u003cem\u003eLawrencella rosea\u003c/em\u003e and \u003cem\u003eP. aristata\u003c/em\u003e seed production was directly linked to bee abundance, resulting in increased seed set in remnants with higher bee abundance. Our results are consistent with the idea that canola increases the attractiveness of a landscape to many bees, increasing overall bee abundance in adjacent areas (Hanley et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Holzschuh et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). These results align with prior research looking at the effect of crop type on native plants in semi-natural habitats (Cussans et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Kov\u0026aacute;cs-Hosty\u0026aacute;nszki et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Insect-attracting crops such as canola can facilitate pollination of co-flowering native plant species in adjacent remnants, compared with non-insect attracting crops such as cereals (e.g., wheat) (Cussans et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Kov\u0026aacute;cs-Hosty\u0026aacute;nszki et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFor our other two focal plant species (\u003cem\u003eG. rosea\u003c/em\u003e and \u003cem\u003eP. gnaphaloides\u003c/em\u003e), canola directly increased reproductive success through mechanisms other than changes in bee abundance, as the link from crop to bee abundance to seeds was not significant or supported by the SEMs. Despite the absence of a direct link, the increase in seed production in these two species likely resulted from changes in pollination behaviour and visitation in canola-adjacent remnants (B\u0026auml;nsch et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Yourstone et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Qiu et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Although \u003cem\u003eG. rosea\u003c/em\u003e had a high capacity for self-fertilization, its reproductive success did significantly increase when it had access to pollinators. Thus, even low pollinator densities may have provided sufficient pollination services due to self-fertilization mechanisms (Aarssen \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Conversely, mass-flowering canola crops might be facilitating \u003cem\u003eP. gnaphaloides\u003c/em\u003e pollination through pollinator spill-over from crops to natural remnants, as has been noted for some shrub species (Kov\u0026aacute;cs-Hosty\u0026aacute;nszki et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and a perennial herb (Qiu et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). While the mechanisms for these direct links between crop type and seed production were not identified by our study, likely drivers are interactions with herbivores, parasites, or competition for pollinators with other plant species in this system (Blitzer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). This result presents an avenue for further investigation between different mutualistic and antagonistic interactions between crops and native plants in agricultural landscapes (e.g. Chamberlain et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn contrast with our findings, the presence of large-scale canola has previously been found to have competitive effects on neighbouring plants near remnants and reserves in canola landscapes elsewhere in the world (Van Reeth et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). For instance, Holzschuh et al. (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) found a landscape-scale competitive effect of canola on \u003cem\u003ePrimula veris\u003c/em\u003e with the pollination services of this species reduced when canola crops were presence in the landscape in Germany. Despite pollinator abundances increasing in the remnants adjacent to canola, the seed set of \u003cem\u003eP. veris\u003c/em\u003e was reduced due to pollinator dilution of the main pollinator, bumblebees (\u003cem\u003eBombus spp.\u003c/em\u003e). Interestingly, despite our study revealing similar pollinator trends, the outcomes of these trends in our system were different. Differences in plant seed production linked to adjacent crop type can be attributed to ecosystem type, the identity of pollinators, and the life strategies of our focal plants. Unlike previous studies, ours is the first to explore the effects of crop type on the seed production and pollination services of native annual plants in remnant reserves, while also linking these effects with different environmental factors at a landscape and local context. Most annual plants have generalist pollination reproductive strategies and are able to maximise seed production from a small number of pollinator visits. This is essential, given their single, short flowering period and the stochastic nature of the pollination environment between years (Munoz et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Because of this life history strategy, annual plants may be more likely to benefit from temporary increases in pollinator abundances caused by mass flowering crops. This effect may explain the strong indirect benefits canola was found to have in our system (Qiu et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePatch size has been reported to have a positive relationship with bee abundance and plant reproduction in other systems (Hadley et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Blaauw and Isaacs \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Rahimi et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), but our study revealed a different trend. We found that plant reproduction was lower in larger remnants than in small remnants. This counterintuitive finding was also identified by (L\u0026aacute;zaro et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), who found that two perennial plant species had lower seed set in larger remnants. This negative relationship may be caused by a decline in pollinator visits across larger areas, due to floral resources scattered over larger areas in larger remnants, and potential barriers to pollinator movement in large remnants, as suggested by L\u0026aacute;zaro et al. (\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePollinator populations are typically limited by nesting habitats and floral resources, which are commonly heterogeneously distributed across habitat patches (Revert\u0026eacute; et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In our study system, floral resources are highly concentrated in patches of certain woodland types (e.g., York Gum-Jam Woodlands) that typically occupy only a part of the whole remnant (see Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb). This results in a heterogeneous distribution of pollinators. These pollinator clusters provide uneven pollination services throughout these remnants (Revert\u0026eacute; et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Escobedo-Kenefic et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mart\u0026iacute;nez-N\u0026uacute;\u0026ntilde;ez et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In smaller remnants, pollinators might be spatially constrained and have fewer floral options, increasing the visitation rate of local flowers.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003eEffect of local-scale components on pollination dynamics\u003c/h2\u003e \u003cp\u003eCanola flowers are acting as pollinator magnets, bringing pollinators into adjacent reserves where pollinators can nest. However, local conditions and resources influence their foraging behaviours. We show that local factors influence pollination dynamics, with local bee and floral abundances having mixed consequences for plant pollen limitation. As plants are well known to vary in their reliance on insect pollinators, this species-level variation is not a surprise. However, the evidence of consequences for annual plant seed production across four arbitrarily selected species in this system suggests that across these specious wildflower communities, adjacent crop type is likely an important mechanism for plant fitness in vegetation remnants in these landscapes. This may then have further trickle-down effects on community diversity and structure.\u003c/p\u003e \u003cp\u003eThis study shows the complex dynamics of multi-scale factors in plant-pollinator interactions. While our work explores aspects of landscape composition, our results highlight that more work is needed to resolve how interannual climatic variation affects the population dynamics of native bees and plants. Fragmentation appears to have disrupted native pollinator communities in this region, with low diversity in each reserve and high species turnover between reserves. However, we currently lack a baseline to understand the population and diversity of native bees throughout this floral biodiversity hotspot. In particular, further research is needed to understand how landscape-level factors and competition for floral resources with honeybees affects the diversity, abundance and dispersion of native bees (e.g., Prendergast et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Despite depauperate native bee communities, we found that pollination services of both crops and native flora are being complemented by the feral honeybee (Hung et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). This may become even more important with the recent introduction of varroa mites to Australia which are likely to decimate managed and feral honeybee populations (Iwasaki et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). As the mites spread across the country, the reliance on native bees for pollination services in natural vegetation is likely to increase, introducing a new type of disturbance to this vulnerable ecosystem.\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eDespite the vast research on the benefits of pollinator spillover from natural remnants to crop fields, little is known about the consequence of pollinator spillover from agricultural fields to remnants of native vegetation. This study represents one of the first to explore the complex reproductive responses of annual native plants to co-occurring landscape fragmentation, agricultural practices, and local plant community attributes. We found that crop identity can directly and indirectly modify plant-pollinator interactions in adjacent remnants, and these effects extend beyond the remnant edge to the core. Large-scale canola crops adjacent to remnants resulted in a direct increase in bee abundance but not bee richness, and an indirect increase in native plant seed production, compared to non-insect rewarding crops such as wheat. Yet, canola's effect on bee abundance depended on the proximity to the remnant edge and pollinator group, mainly increasing feral honeybees\u0026rsquo; abundance. We explored the possible mechanisms in which crop type adjacent to the remnant affects pollination dynamics through pollination experiments, suggesting that, while landscape-scale factors directly influenced bee abundance and plant reproduction, local-scale landscapes elements also influence pollination dynamics and bee diversity. Our study helps unravel how different aspects of the landscape simultaneously affect the abundance of pollinators, their behaviour, and the implications for the persistence of native plant populations in isolated remnants.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eCompeting Interests\u003c/h2\u003e \u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis project was funded by the Australian Research Council (DP210100913) and Botany Foundation at the University of Melbourne (G.A.M. Scott Research Award, 2022).\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eMS and MMM contributed to the study's conceptualisation, development, and experimental design. MS collected the field data with assistance from ZCW. MS conducted the analysis with assistance from ZCW and MMM. All authors contributed to the interpretation of the results. MS wrote the first draft of the manuscript, and ZCW and MMM contributed significantly to its revision.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe acknowledge the Yamatji People as the traditional owners of the land on which we conducted this study. We pay our respects to their Elders past and present. Thank you to Lisa Buche and Hanlun Liu for their help with fieldwork, statistical advice, and manuscript comments. We also thank various Mayfield Lab members for manuscript comments and Yulin He for technical support.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eThe data used in this study will be made available in an online repository following manuscript acceptance.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAarssen LW (2000) Why are most selfers annuals? A new hypothesis for the fitness benefit of selfing. 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Biol Conserv 261:109249. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1016/J.BIOCON.2021.109249\u003c/span\u003e\u003cspan address=\"10.1016/J.BIOCON.2021.109249\" 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":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"landscape-ecology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"land","sideBox":"Learn more about [Landscape Ecology](https://www.springer.com/journal/10980)","snPcode":"10980","submissionUrl":"https://submission.nature.com/new-submission/10980/3","title":"Landscape Ecology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"landscape fragmentation, agricultural practices, pollination dynamics, plant population, York gum-Jam woodlands","lastPublishedDoi":"10.21203/rs.3.rs-4756788/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4756788/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eContext\u003c/h2\u003e \u003cp\u003eAgricultural intensification leads to habitat loss and fragmentation, disrupting plant-pollinator interactions directly, through changes in landscape configuration, and indirectly through altered land-use practices. This has detrimental consequences for the persistence of plants, pollinators, and the ecosystem services they provide.\u003c/p\u003e\u003ch2\u003eObjectives\u003c/h2\u003e \u003cp\u003eWe investigated the mechanisms by which environmental and agricultural context impact pollination dynamics and the reproductive success of native plant species in remnant vegetation within an agricultural mosaic. Specifically, we evaluate the direct and indirect effects of landscape fragmentation (patch size and edge effect) and agricultural practices (crop type adjacent to natural remnants) on bee communities and native plants seed production.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e \u003cp\u003eWe sampled the pollinator community and conducted pollination experiments on four native annual plant species in the core and edge of nine natural remnants. For each site, we recorded remnant size, adjacent crop type (canola or wheat), and local environmental and biological conditions. We then assessed the relationships between these landscape features, bee communities, pollination services, and the reproductive success of native annual forb species.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e \u003cp\u003eBee abundance was higher in reserves adjacent to canola compared to wheat. However, bee abundance decreased from the core to the edge of remnants adjacent to canola, suggesting a possible pollinator dilution effect. Canola directly and indirectly increased seed production of the focal plant species, mediated by changes in pollinator abundance.\u003c/p\u003e\u003ch2\u003eConclusions\u003c/h2\u003e \u003cp\u003eAdjacent crop type, edge effects, and patch size shape plant-pollinator interactions through changes in pollinator abundance, whereas local-scale floral abundance influence pollination dynamics. Our findings indicate that agricultural practices impact the reproductive success of native plants persisting in remnants within an intensively managed agricultural landscape. Further, we show that this effect is mediated by the abundance of generalist insect pollinators in remnant vegetation.\u003c/p\u003e","manuscriptTitle":"Landscape fragmentation and agricultural context impact pollination dynamics of native annual plants","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-08-16 19:22:26","doi":"10.21203/rs.3.rs-4756788/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-07-28T15:04:52+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-22T14:28:21+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-10T16:51:45+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"141367675391637121536844616397054064909","date":"2025-07-08T13:29:40+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"205104830625579704039701682260753365956","date":"2025-06-11T09:30:09+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-08-10T11:56:44+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-07-17T17:08:48+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-07-17T17:08:42+00:00","index":"","fulltext":""},{"type":"submitted","content":"Landscape Ecology","date":"2024-07-17T14:03:53+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"landscape-ecology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"land","sideBox":"Learn more about [Landscape Ecology](https://www.springer.com/journal/10980)","snPcode":"10980","submissionUrl":"https://submission.nature.com/new-submission/10980/3","title":"Landscape Ecology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"5f3cb76f-b466-4c05-b67f-da0dda664f88","owner":[],"postedDate":"August 16th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-02-09T16:16:57+00:00","versionOfRecord":{"articleIdentity":"rs-4756788","link":"https://doi.org/10.1007/s10980-026-02305-2","journal":{"identity":"landscape-ecology","isVorOnly":false,"title":"Landscape Ecology"},"publishedOn":"2026-02-04 15:59:18","publishedOnDateReadable":"February 4th, 2026"},"versionCreatedAt":"2024-08-16 19:22:26","video":"","vorDoi":"10.1007/s10980-026-02305-2","vorDoiUrl":"https://doi.org/10.1007/s10980-026-02305-2","workflowStages":[]},"version":"v1","identity":"rs-4756788","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4756788","identity":"rs-4756788","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}