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Environmental Effects on Germination of Carex Utriculata and Carex Nebrascensis Relative to Riparian Restoration

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Environmental Effects on Germination of Carex Utriculata and Carex Nebrascensis Relative to Riparian Restoration WETLANDS, Vol. 24, No. 2, June 2004, pp. 467±479 q 2004, The Society of Wetland Scientists ENVIRONMENTAL EFFECTS ON GERMINATION OF CAREX UTRICULATA AND CAREX NEBRASCENSIS RELATIVE TO RIPARIAN RESTORATION Kimberly L. Jones1, Bruce A. Roundy1,3, Nancy L. Shaw2, and Jeffrey R. Taylor1 1 Department of Integrative Biology 401 WIDB Brigham Young University Provo, Utah, USA 84602 2 Aquatic Sciences Laboratory U. S. Department of Agriculture, Forest Service Rocky Mountain Research Station 316 E. Myrtle Boise, Idaho, USA 83702 3 Corresponding author Abstract: Seasonal riparian seedbed temperatures were measured and germination of Carex utriculata and C. nebrascensis seeds was tested in relation to chilling, perigynia removal, incubation temperature, and light to help guide propagation and direct seeding of these species for riparian restoration. Diurnal temperatures of riparian seedbeds at two sites in Strawberry Valley, Utah, USA ranged from 3.1 to 11.28 C in May to 9.5 to 24.18 C in August when water was generally available for seed germination. Pre-incubation treatments of chilling at 58 C for 7 to 150 days and perigynia removal increased germination of 2-year-old seeds of these species but were not necessary for high germination percentages (.89%) when seeds were incubated in light under a summer temperature regime (10 to 248 C). Seeds aged 0.5 and 1.5 years had lower germi- nation percentages than 2-year-old seed but also had adequate germination percentages (. 20%) for green- house propagation without pre-treatments when incubated in light at the summer temperature regime. After 5.5 years of storage at room temperatures, germination of C. utriculata was negligible, but that of C. nebrascensis was . 35%. Seeds of both species that were overwintered in seed bags in riparian microsites had high germination percentages (.80%) when retrieved the following summer and incubated at a summer temperature regime in light. Although direct seeding in fall would allow natural chilling and potentially high germination percentages the following spring or summer, the risk of seed loss or excessive burial is great during high spring stream ¯ows. A better strategy is to surface-seed wet seedbeds in early summer after peak ¯ows have receded and temperature and light conditions are conducive to high germination percentages. Key Words: seed dormancy, sedge, revegetation, wetlands, chilling, perigynia, temperature, light, over- wintering INTRODUCTION parian and wetland areas in the Intermountain area, Carex species are commonly reintroduced in wet- USA (Frandsen 1995, Dearden 1998). Determination land and riparian restoration by transplanting sod of their germination requirements would better guide plugs, rhizomes, or intact plants from an adjacent area both nursery propagation and direct seeding strategies or planting greenhouse-propagated seedlings (Ratliff for these species. 1985, Nelson and Williams 1986, Hoag et al. 2001). Pre-incubation treatments required to break dorman- For large areas, direct seeding is potentially more cost- cy, incubation conditions that maximize germination, effective and less disruptive than transplantation. How- and the effects of seed age on germination must be ever, revegetation to restore Carex meadows and wet- understood to guide ef®cient greenhouse propagation lands by direct seeding has failed due to lack of seed (Table 1). Propagators can then weigh the necessity, viability, low germination, or seedling failure (van der costs, and returns of speci®c treatments. For direct Valk et al. 1999). Carex utriculata Boott (now rec- seeding, germination responses can guide the pre-treat- ognized as distinct from C. rostrata Stokes) and C. ments necessary, the season in which to seed, and nebrascensis Dewey are commonly used to restore ri- method of seeding (Table 1). Direct seeding is most 467 468 WETLANDS, Volume 24, No. 2, 2004 Table 1. Research questions and experiments to determine strategies for propagation or ®eld seeding of two Carex species. Experiment number Question Effects tested Greenhouse Propagation 2, 5 Are pre-incubation treatments necessary to break Chilling and perigynia removal on germination seed dormancy? 2, 3, 5 What incubation conditions enhance germination? Incubation temperatures and light on germination 2, 3, 5 How does seed age affect germination? Chilling, incubation temperatures, and light on differ- ent seed ages Field Seeding 1 What are the seasonal diurnal temperatures of ripari- Seasonal temperatures and water potential in riparian an seedbeds when water is available? seedbeds Should we seed in the fall? 2, 5 1. Is chilling necessary for germination? Chilling on germination 2 2. After chilling, how does incubation temperature Spring and summer incubation temperatures on ger- affect germination? mination with or without chilling 4 3. How does overwintering in riparian seedbed mi- Over wintering on germination of seeds in seed bags crosites affect germination? buried and retrieved from different microsites 2, 3, 5 Can we seed after peak ¯ow in summer? Summer seedbed temperatures on germination Practical considerations 2, 4, 5 1. Should we bury seeds or surface sow? Light on germination, burial of overwintered seeds on in-place germination 2 2. Should we remove the perigynia? Perigynia removal on germination 2, 3, 5 3. What seed ages can we use? Seed age on germination ef®cient for wetland species if pretreatments are not For direct seeding, seed burial is preferred to in- required and germination requirements can be met crease the time of available water to seeds and to avoid when conditions are optimal for seedling establish- seed loss by predation or ¯owing water. However, a ment. light requirement for germination necessitates surface Germination of some Carex species is increased by sowing. Surface-sown seeds would be less at risk of chilling, light, removal of the sac-like perigynia cov- washing away if sown in summer rather than fall, but ering the seed, speci®c alternating incubation temper- the area of seedbed where water is available for ger- atures, and certain combinations of these factors (Bu- mination may be restricted. Seeds could be surface- delsky and Galatowitsch 1999, van der Valk et al. sown repetitively over riparian seedbeds that become 1999, SchuÈtz 2000, Hoag et al. 2001). If chilling is exposed in spring as ¯ows diminish. If seeds can ger- required for germination, either fall seeding or chill minate without light, broadcasting them and burying treatments prior to spring or summer sowing are nec- them by raking or harrowing might reduce the number essary. Fall- sowing when stream ¯ows are low can of seeds that wash away in spring and allow sowing potentially cover a larger area of exposed seedbed than in fall over a potentially larger area than in summer. in spring when ¯ows are high. Fall-sown seeds have Germination responses to light under different season- the advantage of being in place when water is available al temperature regimes are required to suggest direct as temperatures warm suf®ciently in spring for ger- seeding options. mination. However, fall-sown seeds are at risk of Because wildland seed production and harvest can washing away in spring ¯oods or being buried too vary annually, seed vendors store Carex seed for re- deep for seedling emergence. Seeding in early summer vegetation projects for 1 to 5 years prior to sale. van when stream ¯ows have stabilized could avoid seed der Valk et al. (1999) recommended that fresh seed be and seedling loss and increase direct seeding success, used in restoration of C. atherodes Spreng., C. lacus- if germination requirements are met. If these Carex tris Willd., and C. stricta Lam. based on reduced ger- species germinate at spring and summer riparian seed- minability and viability after 6 to 18 months of stor- bed temperatures, restorationists can weigh the advan- age. However, the moisture and temperature environ- tages and risks of fall versus spring or summer seeding ment of storage affects germination of various Carex for their particular sites. Seasonal riparian seedbed species differently (Budelsky and Galatowitsch 1999), temperatures when water is available for germination and some species have increased germination after have not been reported. storage, presumably due to after-ripening loss of dor- Jones et al., GERMINATION OF TWO CAREX SPECIES 469 mancy (SchuÈtz 2000). Our purpose was to determine calculated for the population of seeds that germinated germination responses of C. utriculata and C. nebras- in each petri dish. censis to guide strategies for restoration. Field Study Sites METHODS AND MATERIALS Two stream sites differing in geomorphologyÐCo- We conducted ®ve experiments to address speci®c op Creek and Trail Hollow Creek in the Strawberry questions relevant to greenhouse propagation and ®eld Basin, Utah were selected for soil moisture and tem- sowing of C. utriculata and C. nebrascensis (Table 1). perature measurements (Experiment 1) and an over- First, we measured actual riparian seedbed tempera- wintering experiment (Experiment 4). The two differ- tures and water potentials (Experiment 1). We then ent streams were chosen to measure a range of tem- tested germination of 2-year-old seed in response to perature regimes and germination responses after ex- chilling, perigynia removal, light, and incubation tem- posure to riparian seedbed microsites. Co-op Creek is peratures based on ®eld or commonly used propaga- located on the north side of Strawberry Reservoir tion temperatures (Experiment 2).
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