Sand Dropseed (Sporobolus Cryptandrus) – a New Pest in Eurasian Sand Areas? 3 4 *Török P.1,2, Schmidt D.3, Bátori Z.4, Aradi E.5, Kelemen A.5,6, Hábenczyus A
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bioRxiv preprint doi: https://doi.org/10.1101/2021.07.05.451115; this version posted July 5, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. Sand dropseed - a new pest in Eurasia 1 2 Sand dropseed (Sporobolus cryptandrus) – A new pest in Eurasian sand areas? 3 4 *Török P.1,2, Schmidt D.3, Bátori Z.4, Aradi E.5, Kelemen A.5,6, Hábenczyus A. A.4, Diaz C. P.2, 5 Tölgyesi C.1,4, Pál R.W.7, Balogh N.1, Tóth E.1,2, Matus G.8, Táborská J.9, Sramkó G.8,10, Laczkó 6 L.8,10, Jordán S.8,10, Sonkoly J.1,2 7 8 9 1MTA-DE Lendület Functional and Restoration Ecology Research Group, 4032 Debrecen, 1 10 Egyetem sqr., Hungary 11 2Department of Ecology, University of Debrecen, 4032 Debrecen, 1 Egyetem sqr., Hungary 12 3University of Sopron, Institute of Botany and Nature Conservation, 9700 Sopron, 4 Bajcsy-Zs. str., 13 Hungary 14 4Department of Ecology, University of Szeged, 6726 Szeged, 52 Közép boulevard, Hungary 15 5Kiskunság National Park Directorate, 19 Liszt Ferenc str., 6000 Kecskemét, Hungary 16 6Lendület Seed Ecology Research Group, Institute of Ecology and Botany, Centre for Ecological 17 Research, 2-4 Alkotmány Street, 2163 Vácrátót, Hungary 18 7Department of Biological Sciences, Montana Technological University, 1300 W Park Street, Butte, 19 MT 59701, USA 20 8Department of Botany, University of Debrecen, 4032 Debrecen, 1 Egyetem sqr., Hungary 21 9Department of Botany and Plant Physiology, Eszterházy Károly University, 3300 Eger, 6 Leányka 22 str., Hungary. 23 10MTA-DE "Lendület" Evolutionary Phylogenomics Research Group, 4032 Debrecen, 1 Egyetem 24 sqr., Hungary 25 26 *Corresponding author: [email protected] 27 28 Running head: Sand dropseed – a new pest in Eurasia 29 30 Abstract 31 32 For the effective control of an invasive species, gathering as much information as possible on its 33 ecology, establishment and persistence in the subjected communities is of utmost importance. We 34 aimed to review the current distribution and characteristics of Sporobolus cryptandrus (sand 35 dropseed), an invasive C4 grass species of North American origin recently discovered in Hungary. 36 We aimed to provide information on (i) its current distribution paying special attention to its 37 invasion in Eurasia; (ii) the characteristics of the invaded habitats in Central Europe; (iii) seed bank 38 formation and germination characteristics, crucial factors in early establishment; and (iv) the effects 39 of its increasing cover on vegetation composition. Finally, we aimed to (v) point out further 40 research directions that could enable us to understand the invasion success of this potential invasive 41 species. Field surveys uncovered large stands of the species in Central and Eastern Hungary with 42 most of the locations in the former, especially the Kiskunság region. The species invaded disturbed 43 stands of dry and open sand grasslands, closed dune slack grasslands and it also penetrates into 44 natural open sand grasslands from neighbouring disturbed habitats. Increasing cover of Sporobolus 45 cryptandrus caused a decline in species richness and abundance of subordinate species both in the 46 vegetation and seed banks, but a low density of Sporobolus cryptandrus can even have a weak 47 positive effect on these characteristics. Viable seeds of Sporobolus were detected from all soil 48 layers (2.5 cm layers measured from the surface to 10 cm in depth), which indicates that the species 1 bioRxiv preprint doi: https://doi.org/10.1101/2021.07.05.451115; this version posted July 5, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. Sand dropseed - a new pest in Eurasia 49 is able to form a persistent seed bank (1,114 to 3,077 seeds/m2 with increasing scores towards 50 higher abundance of the species in vegetation). Germination of Sporobolus cryptandrus was 51 negatively affected by both litter cover and 1 cm deep soil burial. To sum up, Sporobolus 52 cryptandrus can be considered as a transformer invasive species, whose spread forms a high risk for 53 dry sand and steppe grasslands in Eurasia. We can conclude that for the effective suppression of the 54 species it is necessary: (i) to clarify the origin of the detected populations; (ii) to assess its 55 competitive ability including its potential allelopathic effects; (iii) to assess its seed bank formation 56 potential in habitats with different abiotic conditions; and (iv) to assess the possibility of its 57 suppression by natural enemies and management techniques such as mowing or livestock grazing. 58 59 Keywords: plant invasion; C4 grass; seed bank; germination; sand grassland; steppe; grazing; 60 transformer species 61 2 bioRxiv preprint doi: https://doi.org/10.1101/2021.07.05.451115; this version posted July 5, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. Sand dropseed - a new pest in Eurasia 62 63 Introduction 64 65 The distribution range and abundance of invasive plants have dramatically increased in recent 66 decades providing a serious challenge for protection, conservation, and restoration of natural and 67 semi-natural habitats worldwide (van Kleunen et al. 2019, Pyšek et al. 2020). While casual 68 establishment of alien species in various natural ecosystems became a relatively frequent 69 phenomenon as a consequence of increased human influence, transformer invasive species form one 70 of the most serious threats for natural communities and ecosystems (Richardson et al. 2000). 71 Transformer invasive species often reduce biodiversity, alter disturbance regimes, and affect 72 ecosystem structure and functions in the subjected communities (Richardson et al. 2000, Byers et al. 73 2010, Catford et al. 2011). 74 75 For the effective control of an invasive species, it is crucial to collect as much information as 76 possible on (i) their ecology, especially establishment and persistence characteristics, and on (ii) 77 communities potentially threatened by its invasion. It is also crucial to detect the plant invasion in 78 an early stage, when the distribution of the invasive species is still limited to one or a few isolated 79 locations, where its eradication still might be possible. However, this is rather challenging for 80 inconspicuous species (such as certain grasses), which are usually difficult to determine and to 81 detect (Jarić et al. 2019). Members of the Poaceae and Asteraceae families contribute most of the 82 aggressive invasive plant species across the globe (Pyšek et al. 2017). Invasions of many short-lived 83 and perennial grasses present serious problems worldwide including grasses characterised either by 84 C3 or C4 photosynthetic pathway (D’Antonio & Vitousek 1992, Fusco et al. 2019, van Kleunen et 85 al. 2019). The C4 photosynthetic pathway provides many advantages over the C3 one in arid and 86 warm climate, such as carbon fixation with a lower water cost, higher temperature optimum for 87 carbon fixation, lower sensitivity to water stress and high fire resistance (Johnston 1996). 3 bioRxiv preprint doi: https://doi.org/10.1101/2021.07.05.451115; this version posted July 5, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. Sand dropseed - a new pest in Eurasia 88 89 Thermophilic neophytes include numerous species of Poaceae with the C4 photosynthetic pathway, 90 which have become constant elements of some warm ruderal communities in Eurasia (Leuschner & 91 Ellenberg, 2017). Introduced C4 grasses have already caused dramatic losses of biodiversity in the 92 Americas (e.g., savanna and forest ecosystems in central and South America or desert grasslands 93 and dry woodlands in the North America, Williams & Brauch 2000) or in Australia (e.g., tropical 94 grasslands in Australia, Brooks et al. 2010). Similarly, the Eurasian steppe zone is characterised by 95 the prevailing dominance of C3 grasses with only some notable exceptions of C4 species (e.g., 96 Botriochloa ischaemum, Cynodon dactylon), which are thought to be introduced in historical times 97 (Hurka et al. 2019). According to the projected climate change scenarios, global temperatures will 98 increase in the future, likely resulting in an increased expansion of C4 grasses in plant communities 99 of arid environments of Eurasia. Direct and indirect effects of climate change include the increase 100 of minimum and maximum temperatures, the increasing frequency and magnitude of droughts in 101 the vegetation period, and the changing annual distribution of precipitation shifting the peak of 102 precipitation from the vegetation period to the dormant period (in Central Europe shifting from the 103 summer period to winter, IPCC 2013). This means that increased levels of aridity, increased 104 likeliness of weather extremities, and the associated increased risk of extreme fire events favour the 105 formation and spread of more drought-adapted communities and species. The decline of dominant 106 native C3 grasses like Festuca species increases the risk of invasion via the colonization of drought- 107 adapted non-native C4 species. The effects of climate change may be amplified at the regional scale 108 by large-scale water regulation works and the increased demand for irrigation in agricultural areas, 109 or by the high transpiration rate of established non-native tree plantations (Tölgyesi et al.