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Impacts of Bracteole Removal and Seeding Rate on Seedling Emergence of Halophyte Shrubs: Implications for Rangeland Rehabilitation in Arid Environments

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Impacts of Bracteole Removal and Seeding Rate on Seedling Emergence of Halophyte Shrubs: Implications for Rangeland Rehabilitation in Arid Environments CSIRO PUBLISHING The Rangeland Journal, 2019, 41,33–41 https://doi.org/10.1071/RJ18064 Impacts of bracteole removal and seeding rate on seedling emergence of halophyte shrubs: implications for rangeland rehabilitation in arid environments Mounir Louhaichi A,B,H, Sawsan Hassan A, Ali Mekki Missaoui C, Serkan Ates B, Steven L. Petersen D, Abdoul Aziz Niane E, Slim SlimF and Azaiez Ouled BelgacemG AInternational Centre for Agricultural Research in the Dry Areas (ICARDA), PO Box 950764 – Amman, Jordan. BDepartment of Animal and Rangeland Sciences, Oregon State University, Corvallis, OR 97330, USA. CCentre for Applied Genetic Technologies, University of Georgia, 111 Riverbend Road, Athens, GA 30602, USA. DDepartment of Plant and Wildlife Sciences, Brigham Young University, Provo, UT 84602, USA. EInternational Centre for Agricultural Research in the Dry Areas (ICARDA), PO Box 114/5055 – Beirut, Lebanon. FHigher School of Agriculture of Mateur, University of Carthage, Mateur, Tunisia. GICARDA, Arabian Peninsula Regional Program, PO Box 13979 – Dubai, UAE. HCorresponding author. Email: [email protected] Abstract. Direct seeding techniques often result in unsatisfactory outcomes in rangeland rehabilitation, primarily because of low seedling emergence and poor establishment. Seed processing techniques aimed at improving seedling emergence have gained interest by pasture managers. The purpose of this study was to investigate the combined effects of bracteole removal and seeding rate on seedling emergence in seven halophytic species: Atriplex halimus, A. canescens, A. leucoclada, A. nummularia, A. lentiformis, Salsola vermiculata and Haloxylon aphyllum under semi-arid conditions in Tel Hadya (Syria). Each of these species was evaluated for seedling emergence under two seed treatments (bracteoles removed and non-removed bracteoles) with three seeding rates (10, 30 and 60 seeds per pot), in a completely randomised block design. The results showed a positive effect of seed treatment on seedling emergence for all studied species. The native A. halimus had the highest emergence percentages whereas the introduced A. mummularia, had the lowest. However, there were no significant effects of seeding rates on seedling emergence. These results showed that bracteole removal could improve germination and seedling emergence, and potentially increase the rate of establishment of the species studied. Therefore, when implementing rangeland rehabilitation projects, bracteole removal needs to be considered. The native S. vermiculata should be recommended for direct seeding in the West Asia and North Africa region given its high seedling emergence, known high palatability, nutritive value, and high auto-regeneration performance. Additional keywords: arid and semi-arid ecosystems, direct seeding, indigenous plants, rangeland restoration, seed scarification treatments. Received 4 December 2017, accepted 8 January 2019, published online 12 February 2019 Introduction (Louhaichi et al. 2016a). Therefore, cost-effective techniques Most rangelands in the West Asia and North Africa (WANA) for slowing down and eventually reversing the trend of region are poorly managed, resulting in land degradation, rangeland degradation are needed. reduced productivity, and loss of plant biodiversity (Hudson Transplanting shrub species has been extensively used et al. 2017). Overgrazing, conversion of rangelands into rainfed for rangeland rehabilitation in the WANA region due to high cropping systems, land tenure issues, and climate change are transplant survival, effective soil stabilisation, and reduced soil major drivers for this degradation (Ates and Louhaichi 2012; erosion (Louhaichi and Tastad 2010; Louhaichi et al. 2016b). Ouled Belgacem and Louhaichi 2013). Several government To improve establishment rates, shrub seedlings are usually agencies have becoming increasingly aware of the magnitude grown in nurseries and then transplanted in the field (Franco of the problem and strive to provide solutions to address et al. 2006). However, this process is often expensive and rangeland degradation. However, given the low and slow return time consuming (Douglas et al. 2007) as it requires facilities on investment, governments are not able to finance large-scale and labour to grow seedlings for several weeks or months in projects to effectively restore and develop rangeland resources nurseries. Furthermore, establishing shrubs from nursery-grown Journal compilation Ó Australian Rangeland Society 2019 Open Access CC BY-NC-ND www.publish.csiro.au/journals/trj 34 The Rangeland Journal M. Louhaichi et al. seedlings is particularly challenging in arid environments organic sheep manure. The average annual rainfall over two and/or remote rangeland sites due to lack of irrigation and decades in the study site is 340 mm (ICARDA headquarters adequate site preparation (e.g. establishment of contour meteorological station), occurring predominantly from October furrows). Besides the irrigation cost, maintenance is needed to May. to replace dead seedlings during the first year of establishment Seeds of Atriplex halimus, A. canescens, A. leucoclada, (Palmerlee and Young 2010). A. nummularia, A. lentiformis, Salsola vermiculata and Direct seeding might be a more effective alternative to Haloxylon aphyllum were collected in the autumn of 2011 transplantation of seedlings to improve plant species richness from the ICARDA pastoretum (Table 1). and the productivity of rangelands (Fernández et al. 2012). Direct seeding techniques are common practices for the Experimental procedure establishment of most herbaceous pasture species and have The pots were arranged in a randomised complete block design been used to successfully revegetate large areas (Malcolm with five replications. The treatments consisted of removal 1994). Although direct seeding is a cost-effective technique and non-removal of bracteoles. Bracteoles were manually to implement large-scale restoration projects, its success in removed on half of the seeds (termed seeds without bracteoles) arid regions is limited by harsh environmental factors and poor of all species by scarification using sand paper 212, P80, and soil properties, in addition to seed characteristics such as aluminum oxide. The other half of each seed sample retained seed dormancy and physical structures (e.g. bracteoles) that its bracteoles (termed seeds with bracteoles) and were interfere with seed germination (Ungar and Khan 2001; considered as the control treatments. García-Fayos et al. 2010; Busso et al. 2012; Raizada and Juyal The recommended rate for direct seeding (drill-seeding) 2012; Prats et al. 2016). – of most Atriplex species is between 25 and 35 kg ha 1 (seeds Many halophytes, in particular Atriplex species, have with bracteoles) (Watson and O’Leary 1993; Booth 2002). The chemical and morphological characteristics that regulate seed average weight of 1000 seeds without bracteoles was 0.50 g for dormancy and germination (Ungar 1978; Khan and Ungar A. leucoclada, 0.64 g for A. lentiformis, 0.93 g for A. halimus, 1985). Atriplex seeds are enclosed within bracteoles that 0.72 g for A. nummularia, 1.02 g for S. vermiculata, 1.81 g for control the timing of water imbibition, a crucial step in the H. aphyllum and 2.82 g for A. canescens. Three seeding rates seed germination process (Ungar and Khan 2001). The bracts consisting of 10 (low), 30 (medium), and 60 (high) seeds per pot surrounding the seeds may contain chemical compounds that for each species were compared. This resulted in 21 treatment interfere with germination (Young et al. 1980; Jefferson and combinations for seeds without bracteoles and seeds with Pennacchio 2003). Removal of these bracts has been suggested bracteoles. These combinations resulted in 42 three-way as a mean of improving seed germination of Atriplex halimus treatments (species  bracteole removal  seeding rate). and Salsola vermiculata (Osman and Ghassali 1997). Seeds without bracteoles and seeds with bracteoles were In most rehabilitation projects, managers tend to increase sown directly in the pots at a depth of 0.5 to 1 cm and patted seeding rate assuming that seeds could have a low germination down firmly on 27 January 2012. Five randomly selected rate or could be subject to predation by wildlife (Florentine pots were assigned for each treatment combination, for a total et al. 2013). Direct seeding with a higher seeding rate increases of 210 pots. The pots were kept under natural environmental input costs and may lead to inter-plant competition for conditions of rainfall, temperature, and light. During the nutrients and space (Gordon et al. 1989). However, using low course of this experiment (27 January to 10 March 2012), the seeding rate greatly decreases plant population and results in site received 121.6 mm of rainfall and the average minimum low productivity (Florentine et al. 2013). and maximum temperature varied from 1.98C to 12.58C. Understanding the factors that affect seedling emergence The long average of rainfall and temperature is presented in and plant establishment under rangeland conditions is critical Table 2. for improving our ability to address the important rehabilitation problems such as low succession and poor spatial distribution of perennial plants (Lesica and Allendorf 1999; Ghassali et al. Table 1. Geographic origin of the seven evaluated halophytic rangeland shrub species. Three of the species (A. halimus, A. leucoclada, 2012; Kemp et al. 2013; Porqueddu et al. 2016). Therefore, the S. vermiculata) are indigenous
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