Decisions to allocate management resources should be underpinned by estimates of the impacts of biological invasions that are comparable across species and locations. For the same reason, it is important to assess what type of impacts are likely to occur where, and if such patterns can be generalised. In this paper, we aim to understand factors shaping patterns in the type and magnitude of impacts of a subset of alien grasses. We used the Generic Impact Scoring System (GISS) to review and quantify published impact records of 58 grass species that are alien to South Africa and to at least one other biogeographical realm. Based on the GISS scores, we investigated how impact magnitudes varied across habitats, regions and impact mechanisms using multiple regression. We found impact records for 48 species. Cortaderia selloana had the highest overall impact score, although in contrast to five other species (Glyceria maxima, Nassella trichotoma, Phalaris aquatica, Polypogon monspeliensis, and Sorghum halepense) it did not score the highest possible impact score for any specific impact mechanism. Consistent with other studies, we found that the most frequent environmental impact was through competition with native plant species (with 75% of cases). Socio-economic impacts were recorded more often and tended to be greater in magnitude than environmental impacts, with impacts recorded particularly often on agricultural and animal production (57% and 51% of cases respectively). There was variation across different regions and habitats in impact magnitude, but the differences were not statistically significant. In conclusion, alien grasses present in South Africa have caused a wide range of negative impacts across most habitats and regions of the world. Reviewing impacts from around the world has provided important information for the management of alien grasses in South Africa, and, we believe, is an important component of management prioritisation processes in general.

alien grasses, environmental impact, GISS, impact assessment, impact magnitude, impact mechanism, socio-economic impact.

Grasses (family Poaceae) are among the most introduced species around the world; they occur on every continent and in various habitat types (Linder et al. 2018, van Kleunen et al. 2015, Visser et al. 2016). Alien grasses are often introduced for their high economic value. They are the source for the most consumed staple foods in the world (cereal grains) (Prescott-Allen and Prescott-Allen 1990), pasturage for livestock in agriculture (Boval and Dixon 2012), energy through biofuels (Pimentel and Patzek 2005), and they are used in alcoholic beverages such as beer and whisky (Solange et al. 2014). Alien grasses have also, however, been introduced to new areas as transport contaminants and stowaways. For example, a study by Whinam et al. (2005) found that the major source of alien grass (such as Agrostis stolonifera) introductions into sub-Antarctic islands was the transport used for ship to shore food transfers.

Whether such introductions were accidental or deliberate, and regardless of the many benefits they provide, the introduction of alien grasses can result in invasions that cause substantial negative environmental and socio-economic impacts (Early et al. 2016, D’Antonio and Vitousek 1992, Driscoll et al. 2014). Grasses such as Andropogon gayanus have been reported to increase fire frequencies and intensity in fire-prone ecosystems (Rossiter-Rachor et al. 2004, Rossiter-Rachor et al. 2009, Setterfield et al. 2010). Arundo donax is known to change community structure, thereby causing habitat loss for birds and small mammals in the USA (Bell 1997). And in China, Avena fatua is reported to cause economic losses of US$500 million annually by invading agricultural land and reducing crop yields (Willenborg et al. 2005).

Less is known about how these impacts vary across different introduced ranges, but it has been suggested that some introduced ranges experience fewer recorded impacts from alien grasses due to context-dependent factors (Hulme et al. 2013); e.g. the level of grass invasions might track variation in fire regimes, or might be an artifact of how well studied invasions are (Visser et al. 2016). Either way, impacts of alien grasses are most likely still increasing due to factors such as climate change and propagule pressure (Chuine et al. 2012, Fensham et al. 2013). We therefore need to understand these impacts and take precautionary measures in order to prevent or reduce them (Hulme 2003, 2006, Keller and Perrings 2011). Impact assessments are cost-effective tools used to estimate the impacts of alien species and help in the decision-making process during the prioritization of limited resources (Jeschke et al. 2014, Kumschick et al. 2012, Kumschick and Richardson 2013). Impact assessments have also been used to try to identify factors that predict impacts. Studies have found that traits such as a high fecundity, a habitat generalist strategy, a wide native range, a large body size and a large clutch size are associated with high environmental impacts for mammals, birds, and amphibians (Kumschick et al. 2013, Measey et al. 2016), and traits such as height, life form and life history are associated with greater impacts for plant species (Pyšek et al. 2012, Rumlerová et al. 2016). However, traits have generally been much more successful in predicting invasion success than in predicting impact magnitude. Moreover, impact magnitude has been found to be independent of invasion success (Ricciardi and Cohen 2007).

Similar to the ‘invasive elsewhere’ strategy of predicting invasion (Gordon et al. 2010), is the use of records of ‘impact elsewhere’ to quantify the potential impacts of alien species (Kumschick et al. 2015, Ricciardi 2003). This approach can be useful in predicting the impacts of species such as grasses with biased impact records, i.e. uneven research effort across their introduced ranges. This is because it allows species with limited information to be assessed, compared against other species, and be included in management strategies. Furthermore, the approach also facilitates the search for patterns related to the impact mechanisms and magnitudes, which can ultimately lead to a more predictive understanding of invasions.

Here we assess the environmental and socio-economic impacts of selected alien grasses occurring in South Africa by consolidating their impact records across their introduced ranges (e.g. see Kumschick et al. 2015 for examples of this for alien plants and animals in Europe, and Measey et al. 2016 for amphibians). We do this with the aim of providing quantitative estimates in order to determine which alien grasses have the greatest impacts, and to therefore assist decision makers when prioritising which alien grasses to manage. Furthermore, in order to improve our understanding of the likely impacts, we assess which factors contribute to an increased magnitude of impact in alien grasses by investigating habitats impacted by the species across different regions and determining the mechanisms through which impacts occur.

This study is the first environmental and socio-economic impact assessment to focus specifically on alien grasses. Using the GISS we were able to quantify the impacts of alien grasses using information from across the globe. This study, therefore, provides a useful overview of the literature on evidence-based impacts of alien grasses and highlights potential risks to South Africa. Furthermore, it shows gaps in the available literature as some species could not be assessed due to a lack of impact studies.

We found that alien grasses generally scored higher for socio-economic than environmental impacts. Grass impact scores were particularly high for agricultural and animal production. This might reflect the large number of agricultural weeds that are grasses (Daehler 1998) or their initial introduction for agricultural purposes (Hancock 2012). Alien grasses scored the lowest for impacts caused via transmission of diseases or parasites to native species, with a maximum score of 2, which represents a minor impact (Nentwig et al. 2016), while the frequency under this mechanism was larger. On the contrary, mechanisms with scarce literature, such as impacts on native animals, obtained higher impact scores. This could be because impacts through the transmission of disease or parasites between plant species are not readily observed in the wild, most of the literature under this mechanism is form small-scale laboratory studies which do not report impacts on the overall population.

Despite most grasses not having very high overall impact scores compared to other species (e.g., Kumschick et al. 2015), many alien grasses scored high across the full range of impact mechanisms (i.e. alien grasses can cause a wide range of environmental and socio-economic impacts) and so had high total impact scores. For example, Cortaderia selloana did not have any individual mechanism score over 3 but has the highest overall score (Table 1) due to the many different mechanisms through which it causes impacts. In contrast, Polypogon monspeliensis and Phalaris aquatica scored the highest impact (5) in certain impact mechanisms, but their overall score is lower. This trend is not observed in other studies, such as the one conducted on alien aquatics by Laverty et al. (2015), where the species with the highest overall score also obtained an impact score of 5 for two different mechanisms. Grasses thus provide an interesting case to explore whether we should be more concerned with invasive species that cause a range of different types of impacts or invasive species that only cause a few types of impacts but with greater magnitude.

Grasses are one of the most cosmopolitan plant families in the world and are present in almost all terrestrial habitats. They also impact a wide range of habitats, as demonstrated in this study. Knowledge about which habitats are most severely impacted by alien grasses is essential for their management. Grasses can cause rapid and dramatic transformation of non-grassy habitats into grass-dominated communities. For example, Bromus rubens and B. madritensis have caused widespread transformation of shrubby systems in the Mojave Desert (DeFalco et al. 2007, Jurand et al. 2013). With regards to regions, we found that Antarctica (sub-Antarctic islands mostly) on average has the highest alien grass impact scores. Grasses such as Agrostis stolonifera reduce moss diversity, liverwort populations, and replace the rosaceous dwarf shrub (Acaena magellanica) with dense grassland patches on Marion Island (Gremmen et al. 1998). It is not clear, however, whether this trend is due to differences in sampling effort or a greater susceptibility of sub-Antarctic islands to impacts than the mainland (Hagen and Kumschick 2018).

However, neither habitat nor region were found to be significant predictors of impact magnitude. This could suggest that the impacts are the same across habitats and regions, but the lack of signal likely also reflects the low sample sizes for most habitat types and some regions. Furthermore, it will be interesting to repeat this study based on a more representative global sample of species (the bias in this current analysis towards grasses alien to South Africa was simply for applied reasons).

When we compare impacts scores of alien grasses with impact scores of studies that assessed other plant taxa (Kumschick et al. 2015, Rumlerová et al. 2016), our results also show that the competition with native plant species is the most frequent mechanism through which alien grasses cause impacts. Four species from our list were previously assessed in those studies (Kumschick et al. 2015, Rumlerová et al. 2016), and our results were similar to them for two of the species (Arundo donax and Paspalum dilatatum), each with a difference of less than 5 between the overall impact scores. However, we obtained higher overall impacts than Kumschick et al. (2015) and Rumlerová et al. (2016) for the other two species (Cortaderia selloana and Hordeum jubatum), each with a difference of 9 and 8 respectively. These differences can be explained by the broader search criteria applied; for example, authors of the above-mentioned studies used keywords such as ‘‘invas* or exot* or weed*’’ in addition to the species name, while we only used the species name as a search term.

Although impacts of alien grasses are poorly studied when compared to other species, such as birds and mammals, we were able to find impact records for more than 80% of the grass species selected for the assessment, which is higher than for other species, such as amphibians (41.3%) (Measey et al. 2016). The average number of papers (5.7) used to score impacts of alien grasses across the globe was also higher than the amphibians and other species (Kumschick et al. 2015, Measey et al. 2016). Similar to the mammals and other plants (Kumschick and Nentwig 2010, Kumschick et al. 2015), alien grasses were also reported to cause impact across all impact mechanisms. This might be because grasses occur across a wide range of sectors and habitats, which allows them to exert impact across all mechanisms. When prioritising management of all alien species, our list can be compared to other assessments conducted for other species, such as birds, amphibians, mammals, and aquatic species (Kumschick and Nentwig 2010, Laverty et al. 2015, Measey et al. 2016, Nentwig et al. 2010). However, it is important to note that impact assessments of some of those species are based on impacts recorded only in Europe and not globally, which may cause a bias to the overall impact scores. More impact studies are still needed for alien grass species, especially when it comes to species with no impact records across all introduced ranges, but with taxonomic characteristics of invaders (such as Bambusa balcooa, Canavan et al. 2016). It will be interesting to see if the findings of Canavan et al. (2018a), that bamboos have similar impacts in their native and alien ranges are the same for other grasses or perhaps only other tall-statured grasses (Canavan et al. 2018b). However, we suspect there are qualitative differences between the impacts in the native and alien ranges, for the grasses studied here, as the impacts observed are not primarily a response to human disturbance.

Two species were scored as causing very high impacts (4 or 5) outside of South Africa, but only low levels of impact (1 or 2) in South Africa. For instance, Glyceria maxima obtained a score of 5 because it is associated with the death of livestock through poisoning in Australia (Barton et al. 1983), but such impacts have not (yet) been recorded in South Africa. This can flag species that could potentially cause high impacts in South Africa and which should therefore be monitored, or preventative measures put in place to limit such impacts occurring in future. In most other cases the impact elsewhere was either the same or slightly higher than that recorded in South Afica, except for Agrostis stolonifera, Hordeum murinum, and Nassella trichotoma. This included two species (Nassella trichotoma and Hordeum murinum) whose impacts in South Africa were one level higher than elsewhere. For example, Nassella trichotoma obtained a score of 5 in South Africa and 4 elsewhere (in Australia) for impacts on animal production by reducing livestock carrying capacity and pasture production (Klepeis et al. 2009). The lack of correlation between impacts found in South Africa and elsewhere should, however, be assessed with caution – it is indicative of a research gap. Records of impacts are generally fewer in South Africa (with a maximum of five sources per species and an average of 1.9) and even lacking for most species. Alternatively, it could indicate that there is an impact debt (Rouget et al. 2016), i.e. species have not reached their full impact potential in South Africa (yet), as species with more information in South Africa did not show higher similarities in impact magnitudes to elsewhere. Finally, South Africa might be more resilient to grass invasions, and impacts are actually lower here (Visser et al. 2017). These hypotheses warrant more research and can only be disentangled once more data become available.

In summary, the lack of statistically significant differences in impact magnitudes across habitats and regions for alien grasses suggests that impact in this group is not habitat or region specific as in other groups (cf. Hulme et al. 2013, Pyšek et al. 2011). As such, we recommend that different habitats should be equally considered for alien grass impact management. While we recommend that impact scoring schemes, such as the one used in this study, should be incorporated in the decision-making processes for alien species management, we caution that extrapolations from other invaded regions indicate potential and not actual impacts.

We acknowledge the financial support from the Department of Environmental Affairs through their funding of the South African National Biodiversity Institute. We also acknowledge the DST-NRF Centre of Excellence for Invasion Biology for their support.

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