Establishment of most grassland species was not more suppressed by invasive Sporobolus cryptandrus litter than by native grass litter

Establishment of most grassland species was not more suppressed by invasive Sporobolus cryptandrus litter than by native grass litter | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 25 July 2025 V1 Latest version Share on Establishment of most grassland species was not more suppressed by invasive Sporobolus cryptandrus litter than by native grass litter Authors : Patricia Elizabeth Díaz Cando 0000-0001-9455-5362 , Judit Sonkoly 0000-0002-4301-5240 [email protected] , Annamária Fenesi , Luis Roberto Guallichico Suntaxi 0009-0003-7508-3998 , Gergely Kovacsics-Vári , Luca Di Vita , Francis David Espinoza Ami , … Show All … , Szilvia Madar , Evelin Károlyi , Andrea McIntosh-Buday , Viktória Törő-Szijgyártó , Béla Tóthmérész , and Peter Török 0000-0002-4428-3327 Show Fewer Authors Info & Affiliations https://doi.org/10.22541/au.175345756.69073301/v1 226 views 192 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Sporobolus cryptandrus, a North American C4 grass, is a recent invader in some European sandy grasslands, particularly in Central Europe (Hungary), where it severely threatens native plant communities. As allelopathy has been documented for other Sporobolus species, we tested whether litter from S. cryptandrus has a different effect on the germination and seedling emergence of native grassland species compared to native grass litter. The germination and seedling growth of nine native grassland species and S. cryptandrus were examined in three treatments: without litter (control), with native litter and with S. cryptandrus litter. We hypothesized that: (i) litter has an overall negative effect on seedling germination and establishment compared to no litter; (ii) Sporobolus litter has a significantly more negative effect compared to native grass litter on seedling germination and establishment; and that (iii) the effect of litter type is highly species-specific. Our results showed that the presence of litter did not negatively affect germination and establishment across species, in fact, seedling length even increased in the presence of litter. Contrary to our expectation, Sporobolus litter negatively affected the germination and seedling growth only in the case of Bromus tectorum. For most of the other species, the effects of native and Sporobolus litter were highly similar, suggesting that litter influenced seedlings primarily through physical rather than chemical mechanisms, such as allelopathy. However, the clear suppression of B. tectorum by Sporobolus litter also suggests a potential allelopathic effect. According to our findings, litter effects are species-specific. Moreover, the suppression of B. tectorum by Sporobolus litter could offer novel insights for managing B. tectorum, a problematic invader in North America. Future research should explore the long-term effects of the litter of Sporobolus species in guiding restoration actions and invasive species management. jabbrv-ltwa-all.ldf jabbrv-ltwa-en.ldf Introduction Accumulated plant litter is widely recognized to be among the key drivers of ecosystem processes and vegetation change (Wardle et al. 1997; Loydi et al. 2013; Giweta 2020). Plant litter can impact plant communities in multiple ways by affecting germination, seedling establishment (Carson and Peterson 1990; Hovstad and Ohlson, 2008), species composition, and above-ground biomass production (Kelemen et al. 2013). These effects operate through various mechanisms, including shading, changing soil moisture, formation of a mechanical barrier, influencing nutrient cycling, releasing bioactive chemical compounds, and altering the structure and composition of the soil biota (Weltzin et al. 2005; Letts et al. 2015; Shen et al. 2016). However, distinguishing between these mechanisms is often challenging, as they mostly act in concert. This challenge becomes even more pronounced when dealing with the litter of invasive species which often differs in its quantity and chemical composition compared to native litter (Hassan et al. 2021). Invasive species can fundamentally alter plant community composition and ecosystem processes, not only through direct competition but also by modifying the environment through their litter inputs (Prescott and Zukswert 2016). Litter from invasive plants may physically obstruct seedlings, chemically interfere with germination, and influence soil nutrient cycling, while also affecting native plant growth and microbial communities (Farrer and Goldberg, 2009; Souza et al. 2023). The impact of invasive species’ litter may be even stronger when the invasive species produce a higher amount of biomass or possess leaf traits that differ functionally from those of native species, like higher lignin content or slower decomposition rates (Chiba De Castro et al. 2020; Souza et al. 2023). For example, Bromus diandrus, an invasive species in North America, produces a persistent and thick litter layer inhibiting the seedling establishment of both native and invasive species, with varying effects depending on species and seed size (Chen et al. 2018). Similarly, the invasive grass Microstegium vimineum produces litter that not only suppresses the growth and establishment of the native grass Elymus virginicus but also increases the incidence of fungal disease on the native species. Although both species are negatively affected, the litter disproportionately suppresses E. virginicus , ultimately favouring M. vimineum in competitive interactions (Benitez et al. 2022). These cases demonstrate that invasive species’ litter can reinforce invasive dominance and impact native biodiversity and ecosystem stability, through physical, chemical, or biotic mechanisms (Chen et al. 2018; Benitez et al. 2022; Vujanović et al. 2022). Among the aforementioned mechanisms, allelopathy, i.e., the release of chemical compounds that inhibit the growth of neighbouring plants, has been extensively studied in invasive species research (Ruprecht et al. 2010; Zhang and Fu, 2010; Samedani et al. 2013; Greer et al. 2014; Guido et al. 2020; Hassan et al. 2021; Yuan, et al. 2021; Menicagli et al. 2023; Talmot et al. 2024). While allelopathy represents just one of several mechanisms through which litter can influence plant community dynamics and competitive interaction (Hierro 2003; Kalisz et al. 2021), it may provide an important advantage to invasive species. For example, Imperata cylindrica , Sorghum bicolor and Cynodon dactylon are known to release allelochemicals and suppress native species (Macdonald 2004; Dayan et al. 2009; Mahmoodzadeh 2010; Parker 2022). Allelopathy appears to play a significant role in shaping plant interaction, particularly in grass-dominated ecosystems. Sporobolus cryptandrus , an invasive C4 grass, has become an important concern in some European countries, particularly in Hungary, where it invades disturbed and semi-natural dry grasslands, by outcompeting native species (Török et al. 2021, 2024). Its litter may contribute to its successful invasion through its presumed negative effects on the germination and seedling establishment of native species. These effects may occur through several mechanism, including physical obstruction, altered microclimatic conditions, and allelopathy. While allelopathy has been documented for other species in the same genus, such as Sporobolus pyramidatus ( Rasmussen and Rice, 1971), its potential role in the case of S. cryptandrus remained unexplored so far. In this study, we aimed to analyse whether the litter of S. cryptandrus might affect the germination and early establishment of native species through its physical effects or via allelopathic effects caused by chemical compounds leaching from the litter. We studied the germination and seedling establishment of nine native sand grassland species of Central Europe and S. cryptandrus under three conditions: no litter addition, addition of native species’ litter, or addition of Sporobolus cryptandrus litter. We hypothesized that (i) litter addition has an overall negative effect on seed germination and seedling establishment compared to the ‘no litter’ treatment; (ii) Sporobolus litter has a significantly more negative effect on the germination and early establishment of the studied species compared to native grass litter; and that (iii) the effect of litter type is highly species-specific. Addressing these hypotheses can provide valuable information for designing more effective strategies to control this invasive grass species and guide the proper management of invaded grasslands. Materials and methods The effect of different litter types on the emergence of sand grassland species and early seedling growth were studied using the seeds of 10 species: nine grassland species native to Central Europe and one invasive species, Sporobolus cryptandrus . The species were selected to represent a relatively broad range of functional traits and seed masses (Table 1). The species’ seeds were collected from natural populations located in the Great Hungarian Plain in Eastern Hungary, Central Europe during the summer of 2021. The seeds were cleaned, and stored in a dry, dark environment at room temperature (max. 21°C) until their use. The standing litter (dead plant material still attached to the upright stems) was collected from June to September of 2021 from semi-natural grasslands. The standing litter was collected from three species: the invasive S. cryptandrus and from two native species: Festuca vaginata and Corynephorus canescens. The litter was dried and stored in a dark environment at room temperature. In a greenhouse experiment, plastic pots (9 cm × 9 cm × 9 cm) were filled with steam-sterilized potting soil, and 25 seeds from each one of the ten species were evenly placed on the soil surface in each pot. The seeds (1) were either left uncovered ( no litter treatment); (2) covered with 2.43 g (corresponding to 300 g/m 2 , see Sonkoly et al. 2020) of a 1:1 mixture of F. vaginata and C. canescens litter ( native litter treatment), or (3) covered with 2.43 g of dry litter of S. cryptandrus ( Sporobolus litter treatment). The native litter treatment was a mixture of F. vaginata and C. canescens litter because these native species are co-occurring and co-dominant grass species of sand grasslands and are responsible for most of the accumulated litter in these habitats. The experimental design consisted of 150 pots in total (10 species × 3 treatments × 5 replications). The pots were arranged randomly in a greenhouse (and frequently re-arranged), watered daily with tap water to ensure optimal germination conditions. The germination experiment started on 25th April 2022 (mid spring), and both seed germination and seedling emergence were monitored for five weeks. This duration was chosen based on previous studies that show most seedlings emerge within 7–14 days under greenhouse conditions (Török et al. 2021) and are well established within 4–6 weeks (Török et al. 2024; UMCES 2025). On 26th May 2022 (late spring) all seedlings were counted and removed. The shoot length of seven randomly selected seedlings per pot was measured and the total aboveground dry biomass of seedlings in each pot was weighed, with an accuracy of 0.0001 g. Only seedlings that successfully emerged above the litter layer were considered. Statistical analyses The effect of the litter treatments on the seedling emergence and early seedling growth of nine native sand grasslands species and Sporobolus cryptandrus were analysed using analysis of variance (one-way ANOVAs) followed by Tukey’s test for post hoc comparisons. The dependent variables were germination rate, mean seedling length, and mean seedling dry weight, while the predictor variable was litter treatment with three nominal categories (no litter , native litter, and Sporobolus litter). Shapiro tests, histograms and model diagnostic plots were used to check the assumptions of ANOVA. If the model did not meet the assumptions, logarithmic or square root transformation of the dependent variable was applied. When this did not sufficiently improve the model fit, the non-parametric Kruskal-Wallis test, followed by Dunn’s test with Bonferroni correction for post hoc comparisons was used. To analyse the litter effects across all species in a comparable way, germination rate, mean seedling length, and mean seedling dry weight were standardized by expressing each value as a percentage of the mean value for the same species in the control treatment (‘no litter’). This standardization eliminated the potential confounding influence of species identity. Thereafter, germination rate, seedling length, and dry weight in the three treatments were also analysed for each of the studied species separately. Results When analysed across species, we observed that litter treatment had no significant effect on germination rate ( F 2,147 = 0.822, p = 0.442; Fig. 1A), and on dry weight per seedling ( F 2,147 = 2.574, p = 0.080; Fig. 1C). However, litter treatment had a significant effect on seedling length ( F 2,147 = 8.032, p < 0.001; Fig. 1B); with both native and Sporobolus litter, seedling length was higher compared to the no litter treatment (see Supplemental Information 1). When species were analysed separately, we found that the effect of litter treatment on germination rate was highly species-specific. Only the germination rates of Bromus tectorum and Jasione montana were significantly influenced by litter treatment (Fig. 2), while no significant effects were observed for the remaining eight species (see Table S1 and Figure S1 in Supplemental Information 2). Specifically, Sporobolus litter had a significant negative effect on the germination of Bromus tectorum compared to native litter and no litter treatment (Fig. 2A), but not on the germination of Jasione montana compared to the native litter treatment (Fig. 2B). Litter treatment significantly affected the seedling length of six of the ten species (except for Erysimum diffusum, Festuca pseudovina, J. montana, and S. cryptandrus , see Table S2 and Figure S2 in Supplemental Information 2). In five of these six species, both native and Sporobolus litter significantly increased seedling length compared to the no litter treatment (Fig. 3A, 3C–F). B. tectorum was an exception, for which only native litter significantly increased seedling length compared to the no litter treatment, while Sporobolus litter had no significant effect (Fig. 3B). Litter treatment also significantly affected the seedling dry weight of seven of the ten species, excluding B. tectorum, F. pseudovina, and S. cryptandrus (see Table S3 and Figure S3 in Supplemental Information 2). However, in those seven species, the effect of native and Sporobolus litter did not differ significantly (Fig. 4). Discussion Our hypothesis that added litter reduces seedling germination and establishment compared to the no litter treatment was partially supported: litter moderately affected germination and had a strong impact on the seedling length of some species but this was not true for all of the studied species. The detected pattern supported the findings of some previous studies showing that litter effects depend on both species and litter types (Donath and Eckstein 2008; Ruprecht et al. 2010). However, our results showed that compared with no litter treatment, both litter types (native and Sporobolus litter) had a significant effect on seedling length. This may be because both type of litters come from grasses which are adapted to dry, nutrient-poor environment and tend to produce litter with similar structural characteristics, such as high carbon to nitrogen ratios and slower decomposition rates (Cornwell et al. 2008; Szabó et al. 2017; Seres et al. 2022). Some studies reported that using litter from distinct plant groups like forbs or sedges can have different physical or chemical effects on seedling emergence and growth compared to grass litter (Xiong and Nilsson 1999). Some studies suggest that allelopathy may play a key role in supporting alien species in invading plant communities (Callaway and Aschehoug 2000; Ruprecht et al. 2008; Loydi et al. 2015). In addition, two recent reviews found that most of the invasive species they considered produce allelochemicals with the potential to negatively affect native plant performance (Kalisz et al. 2021) and that native plants suffered more from leachates of alien plants than from leachates of other natives (Zhang et al. 2020). For instance, allelopathic effects have been documented for some Sporobolus species, indicating the potential of this mechanism in the genus more broadly (Rasmussen and Rice 1971). Besides, it is important to note that allelopathic effects may arise not only from compounds actively produced by living plants, but also from the microbial or abiotic decomposition of litter, as observed in species like Juglans regia or Eucalyptus species (Jose and Gillespie 1998; Zhang and Fu 2010). Despite this, our study found no significant effect of either native litter or Sporobolus litter on seed germination and seedling weight when analysed across all species. Our findings are in contrast with previous studies (e.g., Loydi et al. 2013; 2015) which found that the non-native litter reduced the native species’ germination rate while increased the biomass of successfully established seedlings, compared to control conditions without litter. Regarding our second hypothesis, we expected that Sporobolus litter would have a stronger negative effect on the germination and early establishment of native species compared to native grass litter. Our results provided a partial support: Sporobolus litter had a strong negative effect on Bromus tectorum , by suppressing its germination and seedling growth. On the other hand, the effects of Sporobolus litter on other species were similar to that of native litter. This suggests that the litter’s physical structure may exert a general effect across species. The lack of a consistent or considerable difference between the effects of native and Sporobolus litter on most species indicates that any allelopathic effects of S. cryptandrus are probably mild or not as strong as the general mechanical effects of litter. Therefore, allelopathy may not be a dominant mechanism driving the success of S. cryptandrus as an invader. However, the stronger suppression of B. tectorum under Sporobolus litter indicates a potential species-specific chemical interaction, possibly involving allelochemicals, as observed for another species in the genus ( Sporobolus pyramidatus ; Rasmussen and Rice 1971). Additionally, B. tectorum is known to be sensitive to allelopathic effects, especially under stressed conditions like shading or nutrient competition (Machado 2007; Blank and Morgan 2012; Nesrine et al. 2012). While our study did not directly assess chemical composition, this known sensitivity may help explain the stronger response of this species compared to the other studied ones. Further research is needed to clarify the mechanisms driving these interactions, particularly the role of allelochemicals released during litter decomposition and their species-specific effects. The third hypothesis was supported by the results: the effect of litter type was found to be highly species-specific. Litter can have different effects on plants because of various factors (Loydi et al. 2013; 20215; Kortessis et al. 2022). While it can retain soil moisture and facilitate germination and seedling emergence (Letts et al. 2015; Shen et al. 2016), litter can also inhibit growth through shading or by acting as a physical barrier that limits seedling emergence (Vázquez-Yanes et al. 1990; Olson and Wallander 2002; Tormo et al. 2020; Wang et al. 2022). In our study, the responses to litter were highly variable across species. The presence of both native and Sporobolus litter significantly increased the seedling length of six species (except for E. diffusum , F. pseudovina , J. montana , and S. cryptandrus ) and significantly increased the seedling weight of seven species (except for B. tectorum , F. pseudovina , and S. cryptandrus ). While litter promoted seedling elongation, this did not necessarily lead to an increase in seedling dry weight. Instead, elongation may be a stress response rather than an indicator of enhanced growth. Although seedlings grow taller, they may not necessarily be more robust or healthier, as suggested by their biomass (Liu et al. 2017). This trade-off between elongation and accumulating biomass is particularly relevant in short-term experiments, such as ours (lasting five weeks), where seedlings may still be in an early stage of resource allocation. For instance, B. tectorum displayed significant elongation but did not accumulate more biomass, indicating that its response was driven by the shading of litter rather than improved conditions for growth. The results reinforce the idea that litter acts as a physical barrier influencing species differently depending on their functional traits and phenotypic plasticity (Quested and Eriksson 2006; Kortessis et al. 2022; Sparks and Rasmussen 2023). Additionally, the highly variable effects on seedling length and dry weight across different species confirmed the hypothesis that litter effects are species-specific. While native litter often facilitates germination and seedling growth, the effect of Sporobolus litter is less consistent. Contrary to a general suppressive effect from other Sporobolus species, Sporobolus cryptandrus litter did not inhibit growth across all species, as certain plants have developed adaptations that allow them to survive and even thrive under different litter conditions (Facelli and Pickett 1991). Litter reduces light availability, which can negatively impact both seed germination and seedling emergence, with its effects being stronger in species with smaller seeds compared to species with larger ones (Ruprecht et al. 2010; Loydi et al. 2013; Molinari and D’Antonio 2014). The stronger inhibitory effect of litter on the germination of small-seeded species compared to large-seeded species is likely due to the lower energy reserves and higher dependence on light availability for germination in small-seeded species (Jensen and Gutekunst 2003; Eckstein and Donath 2005; Loydi et al. 2013; Sonkoly et al. 2020). Similarly, litter acts as a physical barrier during emergence, making it more challenging for small-seeded species to successfully establish compared to species with larger seeds (Wang et al. 2022). These findings are supported by our data; the small-seeded species J. montana exhibited lower germination rates in the presence of native litter than larger-seeded species such as B. tectorum . Moreover, it is important to consider that grasses used in this study are larger-seeded species with long, narrow coleoptile and vertically arranged leaves. Thus, it is easier for these grasses to grow through deep litter than smaller-seeded dicots with flat leaves growing more horizontally (Grime 2006). For instance, the three grass species B. tectorum , F. pseudovina and F. vaginata showed higher emergence rates in the presence of litter, possibly because their narrow leaf blades facilitated penetration through the litter layer (Germino et al. 2016; Szabó et al. 2017). It is interesting to note that in contrast to the other grasses in this study, S. cryptandrus has relatively small seeds. However, it did not show significant differences in germination, seedling length or biomass among the three-litter treatment (see Supplemental information). This implies that, despite its small seed size, its grass-like morphology – particularly its growth and narrow, erect leaves – may have a compensatory effect against the potential limitations associated with small seed size. These findings show how seed size and seedling morphology interact to influence how species react to litter and may help explain the strong establishment ability of S. cryptandrus which may contribute to its successful invasion. Furthermore, our findings suggest that the physical effects of litter, particularly shading and moisture retention, interact with species-specific traits such as seed size and germination requirements, thereby creating microenvironmental conditions that differently affect species. Although our experimental conditions were designed to optimize watering which may minimize the influence of moisture variations, the importance of light limitation and mechanical disturbance were most important in determining the change in emergence pattern. For example, larger-seeded species may tolerate shading better, whereas smaller-seeded species might experience reduced emergence due to light limitation (Scarpa and Valio 2008; Wang et al. 2022). This aligns with previous research showing that litter can act as both a physical barrier and a protective microenvironment, depending on species traits (Ming-Zhang et al. 2003; Makkonen et al. 2013; Dias et al. 2017). The differential effect of Sporobolus litter on B. tectorum has interesting implications for the interaction between native and invasive species. In North American prairies, where S. cryptandrus is native, these results may offer an additional possible option for managing invasive species such as B. tectorum (Mack, 1981; Leger and Goergen, 2017). Although our results only provide indirect evidence for a presumable allelopathic effect of S. cryptandrus on B. tectorum , our findings imply that the issue deserves additional investigation, especially the identification of the compounds involved and the ecological relevance. In summary, our findings highlight that litter influences germination, seedling emergence and early seedling growth in a species-specific way, with Sporobolus litter exerting stronger negative effects on certain species, particularly B. tectorum . Native and Sporobolus litter can have contrasting impacts, but these effects are highly dependent on the species in question. The similar effects of Sporobolus and native litter on most species suggest that Sporobolus litter primarily exerts the same physical effect on germinating seeds as native litter. On the other hand, Sporobolus litter had a significantly greater negative effect on B. tectorum germination and seedling length than native litter, suggesting a specific, presumably chemical rather than physical effect. Conclusions and Outlook Contrary to our expectations, we found that compared to native litter, the litter of Sporobolus cryptandrus only significantly suppressed the establishment of B. tectorum . To our knowledge, this is the first experimental study addressing the potential allelopathic effects of S. cryptandrus , as previous allelopathy studies in the genus have focused on other species like Sporobolus pyramidatus (Rasmussen and Rice, 1971). Our results suggest that S. cryptandrus litter does not have a broad allelopathic effect, but the results reveal that it can negatively affect certain species. Thus, our results offer novel information on the possible potential role by the presence of litter interaction in the invasiveness of S. cryptandrus. Although we found no general allelopathic effect, the observed suppression of B. tectorum by S. cryptandrus litter may have important implications. This finding may offer a natural, low-impact option for controlling the spread of B. tectorum in North American prairie ecosystems where B. tectorum is an invasive plant and S. cryptandrus is native. Additionally, S. cryptandrus itself was unaffected by any of the litter treatments. Its germination and seedling growth were not significantly affected by the presence of either its own litter or native litter. It may indicate high litter tolerance that could contribute to its rapid establishment and hence competitive advantage in communities with high litter accumulation. This tolerance of S. cryptandrus to litter accumulation may contribute to its successful invasion as much as its potential allelopathic effects. Future studies should explore whether S. cryptandrus influences neighbouring species through other mechanism like root exudates, and whether its litter can suppress invasive B. tectorum populations in North America under field conditions. References Benitez, L., Kendig, A. E., Adhikari, A., Clay, K., Harmon, P. F., Holt, R. D., Goss, E. M., and Flory, S. L. 2022. Invasive grass litter suppresses a native grass species and promotes disease. Ecosphere . 13: e3907. https://doi.org/10.1002/ecs2.3907 Blank, R., and Morgan, T. 2012. Suppression of Bromus tectorum L. by established perennial grasses: potential mechanisms—part one. Appl. Environ. Soil Sci. 2012: 632172. https://doi.org/10.1155/2012/632172 Callaway, R. M., and Aschehoug, E. T. 2000. Invasive plants versus their new and old neighbors: a mechanism for exotic invasion. Science . 290: 521–523. https://doi.org/10.1126/science.290.5491.521 Carson, W. P., and Peterson, C. J. 1990. The role of litter in an old-field community: impact of litter quantity in different seasons on plant species richness and abundance. Oecologia. 85: 8–13. https://doi.org/10.1007/BF00317337 Chen, B. M., D’Antonio, C. M., Molinari, N., and Peng, S. L. 2018. Mechanisms of influence of invasive grass litter on germination and growth of coexisting species in California. Biol. Invas. 20: 1881–1897. https://doi.org/10.1007/s10530-018-1668-5 Chiba De Castro, W. A., Almeida, R. V., Xavier, R. O., Bianchini, I., Moya, H., and Silva Matos, D. M. 2020. Litter accumulation and biomass dynamics in riparian zones in tropical South America of the Asian invasive plant Hedychium coronarium J. König (Zingiberaceae). Plant Ecol. Divers . 13: 47–59. https://doi.org/10.1080/17550874.2019.1673496 Prescott, C. E., and Zukswert, J. M. 2016. Invasive plant species and litter decomposition: time to challenge assumptions. New Phytol . 209: 5–7. https://www.jstor.org/stable/newphytologist.209.1.5 Cornwell, W. K., Cornelissen, J. H., Amatangelo, K., Dorrepaal, E., Eviner, V. T., Godoy, O., Hobbie, S. E., Hoorens, B., Kurokawa, H., Pérez-Harguindeguy, N., Quested, H. M., Santiago, L. S., Wardle, D. A., Wright, I. J., Aerts, R., Allison, S. D., Van Bodegom, P., Brovkin, V., Chatain, A., Callaghan, T. V., Díaz, S., Garnier, E., Gurvich, D. E., Kazakou, E., Klein, J. A., Read, J., Reich, P. B., Soudzilovskaia, N. A., Vaieretti, M. V., and Westoby, M. 2008. Plant species traits are the predominant control on litter decomposition rates within biomes worldwide. Ecol. Lett . 11: 1065–1071. https://doi.org/10.1111/j.1461-0248.2008.01219.x Dayan, F. E., Howell, J. L., and Weidenhamer, J. D. 2009. Dynamic root exudation of sorgoleone and its in planta mechanism of action. J. Exp. Bot. 60: 2107–2117. https://doi.org/10.1093/jxb/erp082 Donath, T. W., and Eckstein, R. L. 2008. Grass and oak litter exert different effects on seedling emergence of herbaceous perennials from grasslands and woodlands. J. Ecol. 96: 272– 280. https://doi.org/10.1111/j.1365-2745.2007.01338.x Dias, A. T. C., Cornelissen, J. H., and Berg, M. P. 2017. Litter for life: assessing the multifunctional legacy of plant traits. J. Ecol. 105: 1163–1168. https://doi.org/10.1111/1365-2745.12763 Vujanović, D., Losapio, G., Milić, S., and Milić, D. 2022. The impact of multiple species invasion on soil and plant communities increases with invasive species co-occurrence. Front. Plant Sci . 13 : 875824. https://doi.org/10.3389/fpls.2022.875824 Eckstein, R. L., and Donath, T. W. 2005. Interactions between litter and water availability affect seedling emergence in four familial pairs of floodplain species. J. Ecol . 93: 807–816. https://doi.org/10.1111/j.1365-2745.2005.01015.x Facelli, J. M., Pickett, S.T. 1991. Plant litter: Its dynamics and effects on plant community structure. Bot. Rev. 57: 1–32. https://doi.org/10.1007/BF02858763 Farrer, E. C., and Goldberg, D. E. 2009. Litter drives ecosystem and plant community changes in cattail invasion. Ecol. Appl , 19: 398–412. https://doi.org/10.1890/08-0485.1 Germino, M. J., Belnap, J., Stark, J. M., Allen, E. B., and Rau, B. M. 2016. “Ecosystem impacts of exotic annual invaders in the genus Bromus”. In Exotic brome-grasses in arid and semiarid ecosystems of the western US: Causes, consequences, and management implications , edited by Germino, M., Chambers, J., Brown, C. 61–95. Switzerland, Springer, Cham. https://doi.org/10.1007/978-3-319-24930-8_3 Giweta, M. 2020. Role of litter production and its decomposition, and factors affecting the processes in a tropical forest ecosystem: a review. J. Ecol. Environ . 44: 11. https://doi.org/10.1186/s41610-020-0151-2 Greer, M. J., Wilson, G. W., Hickman, K. R., and Wilson, S. M. 2014. Experimental evidence that invasive grasses use allelopathic biochemicals as a potential mechanism for invasion: chemical warfare in nature. Plant soil . 385: 165–179. https://doi.org/10.1007/s11104-014-2209-3 Grime, J. P. 2006. Plant strategies, vegetation processes, and ecosystem properties. John Wiley & Sons Ltd., Chichester-New York-Brisbane-Toronto. 222 pp. Guido, A., Quiñones, A., Pereira, A. L., and Da Silva, E. R. 2020. Are the invasive grasses Cynodon dactylon and Eragrostis plana more phytotoxic than a co-occurring native? Ecol. Austral. 30: 295–303. https://doi.org/10.25260/EA.20.30.2.0.1090 Hassan, N., Sher, K., Rab, A., Abdullah, I., Zeb, U., Naeem, I., Shuaib, M., Khan, H., Khan, W., and Khan, A. 2021. Effects and mechanism of plant litter on grassland ecosystem: A review. Acta Ecol. Sin . 41: 341–345. https://doi.org/10.1016/j.chnaes.2021.02.006 Hierro, J. L., and Callaway, R. M. 2003. Allelopathy and exotic plant invasion. Plant Soil .256: 29–39. https://doi.org/10.1023/A:1026208327014 Hierro, J. L., and Callaway, R. M. 2021. The ecological importance of allelopathy. Annu. Rev. of Ecol. Evol. Syst . 52: 25–45. https://doi.org/10.1146/annurev-ecolsys-051120-030619 Hovstad, K. A., and Ohlson, M. 2008. Physical and chemical effects of litter on plant establishment in semi-natural grasslands. Plant Ecol . 196: 251–260. https://doi.org/10.1007/s11258-007-9349-y Jensen, K., and Gutekunst, K. 2003. Effects of litter on establishment of grassland plant species: the role of seed size and successional status. Basic Appl. Ecol . 4: 579–587. https://doi.org/10.1078/1439-1791-00179 Jose, S., and Gillespie, A. R. 1998. Allelopathy in black walnut ( Juglans nigra L.) alley cropping. II. Effects of juglone on hydroponically grown corn ( Zea mays L.) and soybean ( Glycine max L. Merr.) growth and physiology. Plant Soil . 203 : 199–206. https://doi.org/10.1023/A:1004353326835 Kalisz, S., Kivlin, S. N., and Bialic-Murphy, L. 2021. Allelopathy is pervasive in invasive plants. Biol. Invasions . 23 : 367–371. https://doi.org/10.1007/s10530-020-02383-6 Kelemen, A., Török, P., Valkó, O., Miglécz, T., and Tóthmérész, B. 2013. Mechanisms shaping plant biomass and species richness: plant strategies and litter effect in alkali and loess grasslands. J. Veg. Sci . 24: 1195–1203. https://doi.org/10.1111/jvs.12027 Kleyer, M., Bekker, R. M., Knevel, I. C., Bakker, J. P., Thompson, K., Sonnenschein, M., Poschlod, P., van Groenendael, J. M., Klimes, L., Klimesová, J., Klotz, S., Rusch, G. M., Hermy, M., Adriaens, D., Boedeltje, G., Bossuyt, B., Dannemann, A., Endels, P., Götzenberger, L., Hodgson, J. G., Jackel, A.-K., Kühn, I., Kunzmann, D., Ozinga, W. A., Römermann, C., Stadler, M., Schlegelmilch, J., Steendam, H. J., Tackenberg, O., Wilmann, B., Cornelissen, J. H. C., Eriksson, O., Garnier, E. and Peco, B. 2008. The LEDA Traitbase: a database of life-history traits of Northwest European flora. J. Ecol. 96: 1266–1274. https://doi.org/10.1111/j.1365-2745.2008.01430.x Kortessis, N., Kendig, A. E., Barfield, M., Flory, S. L., Simon, M. W., and Holt, R. D. 2022. Litter, plant competition, and ecosystem dynamics: a theoretical perspective. Am. Nat . 200: 739–754. https://doi.org/10.1086/721438 Leger, E. A., and Goergen, E. M. 2017. Invasive Bromus tectorum alters natural selection in arid systems. J. Ecol . 105: 1509–1520. https://doi.org/10.1111/1365-2745.12852 Letts, B., Lamb, E. G., Mischkolz, J. M., and Romo, J. T. 2015. Litter accumulation drives grassland plant community composition and functional diversity via leaf traits. Plant Ecol . 216: 357–370. https://doi.org/10.1007/s11258-014-0436-6 Liu, B., Daryanto, S., Wang, L., Li, Y., Liu, Q., Zhao, C., and Wang, Z. 2017. Excessive accumulation of Chinese fir litter inhibits its own seedling emergence and early growth—A greenhouse perspective. Forests . 8: 341. https://doi.org/10.3390/f8090341 Loydi, A., Eckstein, R. L., Otte, A., and Donath, T. W. 2013. Effects of litter on seedling establishment in natural and semi‐natural grasslands: a meta‐analysis. J. Ecol . 101: 454–464. https://doi.org/10.1111/1365-2745.12033 Loydi, A., Donath, T.W., Eckstein, R.L., and Otte, A. 2015. Non-native species litter reduces germination and growth of resident forbs and grasses: allelopathic, osmotic or mechanical effects? Biol. Invasions . 17: 581–595 https://doi.org/10.1007/s10530-014-0750-x Machado, S. 2007. Allelopathic potential of various plant species on downy brome: implications for weed control in wheat production. Agron. J . 99: 127–132. https://doi.org/10.2134/agronj2006.0122 MacDonald, G. E. 2004. Cogongrass ( Imperata cylindrica )—biology, ecology, and management. Crit. Rev. Plant Sci . 23: 367–380. https://doi.org/10.1080/07352680490505114 Mack, R. N. 1981. Invasion of Bromus tectorum L. into western North America: an ecological chronicle. Agroecosystems . 7: 145–165. https://doi.org/10.1016/0304-3746(81)90027-5 Mahmoodzadeh, H. 2010. Allelopathic Plants 23. Cynodon dactylon (L.) Pers . Allelopathy J . 25: 227–237 Makkonen, M., Berg, M. P., van Logtestijn, R. S., van Hal, J. R., and Aerts, R. 2013. Do physical plant litter traits explain non‐additivity in litter mixtures? A test of the improved microenvironmental conditions theory. Oikos . 122: 987–997. https://doi.org/10.1111/j.1600-0706.2012.20750.x Menicagli, V., Balestri, E., Giommoni, F., Vannini, C., and Lardicci, C. 2023. Plastic litter changes the rhizosphere bacterial community of coastal dune plants. Sci. Total Environ . 880: 163293. https://doi.org/10.1016/j.scitotenv.2023.163293 Ming-Zhang, W. E. N., Dan, Y. U., and Ji-Xun, G. U. O. 2003. Influence of litter layer on microenvironment in northeast Leymus chinensis grassland. Plant Sci. J . 21: 395–400. Molinari, N. A., and D’Antonio, C. M. 2014. Structural, compositional and trait differences between native‐and non‐native‐dominated grassland patches. Funct. Ecol. 28: 745–754. https://doi.org/10.1111/1365-2435.12206 Nesrine, S., El-Darier, S. M., and Taher, H. M. 2012. The allelochemicals effect of Zygophyllum album on control of Bromus tectorum . J. Life Sci. 6: 182–186 Olson, B. E., and Wallander, R. T. 2002. Effects of invasive forb litter on seed germination, seedling growth and survival. Basic Appl. Ecol . 3: 309–317. https://doi.org/10.1078/1439-1791-00127 Parker, C. 2022. Imperata cylindrica (cogon grass). CABI Compendium. https://doi.org/10.1079/cpc.28580.20210100324 Pladias 2025. Pladias Database of the Czech Flora and Vegetation. Published on the Internet. www.pladias.cz [Accessed June 8 2025]. POWO. 2025. Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. Published on the Internet. http://www.plantsoftheworldonline.org/ [Accessed July 8, 2025]. Quested, H., and Eriksson, O. 2006. Litter species composition influences the performance of seedlings of grassland herbs. Funct. Ecol. 20: 522–532. https://www.jstor.org/stable/3806544 Rasmussen, J. A., and Rice, E. L. 1971. Allelopathic effects of Sporobolus pyramidatus on vegetational patterning. Am. Mindl. Nat. 86: 309–326. https://doi.org/10.2307/2423626 Ruprecht, E., Donath, T. W., Otte, A., and Eckstein, R. L. 2008. Chemical effects of a dominant grass on seed germination of four familial pairs of dry grassland species. Seed Sci. Res . 18: 239–248. https://doi.org/10.1017/S0960258508096190 Ruprecht, E., Józsa, J., Ölvedi, T. B., and Simon, J. 2010. Differential effects of several “litter” types on the germination of dry grassland species. J. Veg. Sci. 21: 1069–1081. https://doi.org/10.1111/j.1654-1103.2010.01206.x Samedani, B., Juraimi, A. S., Rafii, M. Y., Anuar, A. R., Sheikh Awadz, S. A., and Anwar, M. P. 2013. Allelopathic effects of litter Axonopus compressus against two weedy species and its persistence in soil. Sci World J . 2013: 695404. https://doi.org/10.1155/2013/695404 Scarpa, F. M., and Valio, I. F. M. 2008. Relationship between seed size and litter effects on early seedling establishment of 15 tropical tree species. J. Trop. Ecol . 24: 569–573. https://doi.org/10.1017/S0266467408005300 Seres, A., Kröel‐Dulay, G., Szakálas, J., Nagy, P. I., Boros, G., Ónodi, G., Kertész, M., Szitár, K., and Mojzes, A. 2022. The response of litter decomposition to extreme drought modified by plant species, plant part, and soil depth in a temperate grassland. Ecol. Evol . 12 : e9652. https://doi.org/10.1002/ece3.9652 Shen, Y., Chen, W., Yang, G., Yang, X., Liu, N., Sun, X., Chen, J., and Zhang, Y. 2016. Can litter addition mediate plant productivity responses to increased precipitation and nitrogen deposition in a typical steppe? Ecol. Res. 31: 579–587. https://doi.org/10.1007/s11284-016-1368-5 Sonkoly, J., Valkó, O., Balogh, N., Godó, L., Kelemen, A., Kiss, R., Miglécz, T., Tóth, E., Tóth, K., Tóthmérész, B., and Török, P. 2020. Germination response of invasive plants to soil burial depth and litter accumulation is species‐specific. J. Veg. Sci . 31: 1079–1087. https://doi.org/10.1111/jvs.12891 Sonkoly, J., Tóth, E., Balogh, N., Balogh, L., Bartha, D., Csendesné Bata, K., Bátori, Z., Békefi, N., Botta-Dukát, Z., Bölöni, J., Csecserits, A., Csiky, J., Csontos, P., Dancza, I., Deák, B., Dobolyi, Z. K., Vojtkó, A., Gyulai, F., Hábenczyus, A. A., Henn, T., Horváth, F., Höhn, M., Jakab, G., Kelemen, A., Király, G., Kis, S., Kovacsics-Vári, G., Kun, A., Lehoczky, É., Lengyel, A., Lhotsky, B., Löki, V., Lukács, B. A., Matus, G., McIntosh-Buday, A., Mesterházy, A., Miglécz, T., Molnár, V, A., Molnár, Z., Morschhauser, T., Papp, L., Pósa, P., Rédei, T., Schmidt, D., Szmorad, F., Takács, A., Tamás, J., Tiborcz, V., Tölgyesi, C., Tóth, K., Tóthmérész, B., Valkó, O., Virók, V., Wirth, T., and Török, P. 2023. PADAPT 1.0–the Pannonian dataset of plant traits. Sci. Data , 10: 742. https://doi.org/10.1038/s41597-023-02619-9 Souza, T., Lucena, E. O. D., de Andrade, L. A., da Silva, L. J. R., Nascimento, G. D. S., and Freitas, H. 2023. Litter deposition and nutrient cycling of invaded environments by Cryptostegia madagascariensis at tropical cambisols from northeastern Brazil. Int. J. Plant Biol . 14: 254–265. https://doi.org/10.3390/ijpb14010021 Sparks, E. E., and Rasmussen, A. 2023. Trade‐offs in plant responses to the environment. Plant Cell Environ. 46: 2943–2945. https://doi.org/10.1111/pce.14689 Szabó, G., Zimmermann, Z., Catorci, A., Csontos, P., Wichmann, B., Szentes, S., Barczi, S., and Penksza, K. 2017. Comparative study on grasslands dominated by Festuca vaginata and F-pseudovaginata in the Carpathian Basin. Tuexenia . 37: 415–429. https://doi.org/10.14471/2017.37.018 Talmot, V., Hardion, L., Sexton, A., Jumeau, J., Jugieau, E., and Staentzel, C. 2024. Environmental conditions affect the allelopathic potential of three invasive alien plants species in North‐Eastern France. J. Ecol . 113: 155–167. https://doi.org/10.1111/1365-2745.14447 Tormo, J., Amat, B., and Cortina, J. 2020. Litter as a filter for germination in semi-arid Stipa tenacissima steppes. J. Arid. Environ . 183: 104258. https://doi.org/10.1016/j.jaridenv.2020.104258 Török, P., Schmidt, D., Bátori, Z., Aradi, E., Kelemen, A., Hábenczyus, A. A., Díaz Cando, P., Tölgyesi, C., Pál, R. W., Balog. N., Tóth, E., Matus, G., Tánorská, J., Sramkó, G., Laczkó, L., Jordán, S., McIntosh‐Buday, A., Kovacsics-Vári, G., and Sonkoly, J. 2021. Invasion of the North American sand dropseed ( Sporobolus cryptandrus )–A new pest in Eurasian sand areas? Glob. Ecol. Conserv. 32: e01942. https://doi.org/10.1016/j.gecco.2021.e01942 Török, P., Espinoza Ami, F. D., Szél‐Tóth, K., Díaz Cando, P., Guallichico Suntaxi, L. R., McIntosh‐Buday, A., Hábenczyus, A. A., Törő-Szijgyártó, V., Kovacsics-Vári, G., Tölgyesi, C., Tóthmérész, B., and Sonkoly, J. 2024. Accumulated soil seed bank of the invasive sand dropseed ( Sporobolus cryptandrus ) poses a challenge for its suppression. Land Degrad. Dev . 35: 4105–4120. https://doi.org/10.1002/ldr.5208 UMCES. University of Maryland Center for Environmental Science (UMCES). 2025 Sand, Prairie and Rough dropseed ( Sporobolus ‐summary) – University of Maryland Center for Environmental Science. Available at: https://www.umces.edu/sites/default/files/Sporobolus-summary.pdf [accessed 21 Jul. 2025]. Vázquez-Yanes, C., Orozco-Segovia, A., Rincon, E. E. A., Sanchez-Coronado, M. E., Huante, P., Toledo, J. R., and Barradas, V. L. 1990. Light beneath the litter in a tropical forest: effect on seed germination. Ecology . 71: 1952–1958. https://doi.org/10.2307/1937603 Wang, Z., Wang, D., Liu, Q., Xing, X., Liu, B., Jin, S., and Tigabu, M. 2022. Meta-analysis of effects of forest litter on seedling establishment. Forests. 13: 644. https://doi.org/10.3390/f13050644 Wardle, D. A., Bonner, K. I., and Nicholson, K. S. 1997. Biodiversity and plant litter: experimental evidence which does not support the view that enhanced species richness improves ecosystem function. Oikos , 79: 247–258. https://www.jstor.org/stable/3546010 Weltzin, J. F., Keller, J. K., Bridgham, S. D., Pastor, J., Allen, P. B., and Chen, J. 2005. Litter controls plant community composition in a northern fen. Oikos . 110: 537–546. https://doi.org/10.1111/j.0030-1299.2005.13718.x Xiong, S., and Nilsson, C. 1999. The effects of plant litter on vegetation: a meta‐analysis. J. Ecol. 87: 984–994. https://doi.org/10.1046/j.1365-2745.1999.00414.x Yuan, L., Li, J. M., Yu, F. H., Oduor, A. M., and van Kleunen, M. 2021. Allelopathic and competitive interactions between native and alien plants. Biol. Invasions. 23: 3077–3090. https://doi.org/10.1007/s10530-021-02565-w Zhang, C., and Fu, S. 2010. Allelopathic effects of leaf litter and live roots exudates of Eucalyptus species on crops. Allelopathy J. 26: 91–99. Zhang, Z., Liu, Y., Yuan, L., Weber, E., and van Kleunen, M. 2020. Effect of allelopathy on plant performance: a meta‐analysis. Ecol. Lett . 24: 348–362. https://doi.org/10.1111/ele.13627 jabbrv-ltwa-all.ldf jabbrv-ltwa-en.ldf Tables Table 1. List of the ten 10 studied species, including their native distribution (POWO 2025), thousand-seed weight (TSW) in grams (Sonkoly et al. 2023), and habitat types where they occur (Török et al. 2021; Pladias 2025). Tait data were collected from published source (LEDA Traitbase: Kleyer et al. 2008) Sporobolus cryptandrus Poaceae North America Perennial grass Grass 0.090 Dry grasslands and other open areas, on sandy to rocky soils Arenaria leptoclados Caryophyllaceae Europe, Asia Annual forb Forb 0.020 Sandy and rocky habitats Bromus tectorum Poaceae Europe, Asia Annual grass 3.696 Disturbed areas (pastures, along roadsides) Crepis rhoeadifolia Asteraceae Europe, Asia Annual forb Forb 0.295 Meadows, open woodlands, and disturbed areas Erysimum diffusum Brassicaceae Central Europe, Asia short-lived perennial Forb 0.187 Rocky slopes, various grasslands, and open woodlands Festuca pseudovina Poaceae Central Europe, Asia Perennial Grass 0.263 Dry grasslands, meadows, and open woodlands Festuca vaginata Poaceae Central Europe Perennial grass Grass 0.647 Dry grasslands, meadows, and open woodlands Jasione montana Campanulaceae Europe, Africa Short-lived perennial Forb 0.016 Grasslands, heathlands, and rocky slopes. Plantago indica Plantaginaceae Europe, Africa Annual Forb 0.805 Disturbed areas, along roadsides, and in grasslands. Petrorhagia prolifera Caryophyllaceae Europe, Africa Annual Forb 0.278 Sandy and rocky habitats Figure 1. The effect of litter treatment on germination rate (A), seedling length (B), and dry weight per seedling (C) analysed across species (one-way ANOVA and Tukey tests). Standardised values for germination rate, seedling length and dry weight were given compared to the mean value of the given variable in the no litter (control) treatment for every species to avoid the confounding effect of species identity. Significant differences (p<0.05) were indicated by superscripted letters. Figure 2. The effect of litter treatment on the germination rate of Bromus tectorum (A) and Jasione montana (B) (one-way ANOVA and Tukey tests, only species with significant differences are shown). Figure 3. The effect of litter treatment on seedling length analysed separately for each species using one-way ANOVA and Tukey tests, only species with significant differences are shown: Arenaria leptoclados ( A), Bromus tectorum ( B), Crepis rhoeadifolia ( C), Festuca vaginata ( D) , Petrorhagia prolifera (E) , and Plantago indica ( F) . Figure 4. The effect of litter treatment on dry weight per seedling analysed separately for each species using one-way ANOVAs and Tukey tests, only species with significant differences are shown (indicated by superscripted letters): Erysimum diffusum ( A) , Festuca vaginata ( B) , Arenaria leptoclados (C) , Crepis rhoeadifolia ( D) , Jasione montana ( E) , Petrorhagia prolifera ( F), and Plantago indica ( G). jabbrv-ltwa-all.ldf jabbrv-ltwa-en.ldf Figure 4. continued Information & Authors Information Version history V1 Version 1 25 July 2025 Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords allelopathy germination grasslands invasive species litter Authors Affiliations Patricia Elizabeth Díaz Cando 0000-0001-9455-5362 University of Debrecen View all articles by this author Judit Sonkoly 0000-0002-4301-5240 [email protected] University of Debrecen View all articles by this author Annamária Fenesi Babeş-Bolyai University View all articles by this author Luis Roberto Guallichico Suntaxi 0009-0003-7508-3998 University of Debrecen View all articles by this author Gergely Kovacsics-Vári University of Debrecen View all articles by this author Luca Di Vita University of Palermo View all articles by this author Francis David Espinoza Ami University of Debrecen View all articles by this author Szilvia Madar HUN-REN-UD Functional and Restoration Ecology Research Group View all articles by this author Evelin Károlyi HUN-REN-UD Functional and Restoration Ecology Research Group View all articles by this author Andrea McIntosh-Buday University of Debrecen View all articles by this author Viktória Törő-Szijgyártó University of Debrecen View all articles by this author Béla Tóthmérész MTA-DE View all articles by this author Peter Török 0000-0002-4428-3327 University of Debrecen View all articles by this author Metrics & Citations Metrics Article Usage 226 views 192 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Patricia Elizabeth Díaz Cando, Judit Sonkoly, Annamária Fenesi, et al. Establishment of most grassland species was not more suppressed by invasive Sporobolus cryptandrus litter than by native grass litter. Authorea . 25 July 2025. DOI: https://doi.org/10.22541/au.175345756.69073301/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu . 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