Phenotypic variation of side-oats grama grass [Bouteloua curtipendula (Michx.) Torr.

] collections from the Canadian prairie

M. P. Schellenberg, B. Biligetu1, G. J. McLeod, and Z. Wang
Semiarid Prairie Agricultural Research Centre, Agriculture and Agri-Food Canada, Swift Current, Saskatchewan, Canada S9H 3X2. Received 29 July 2011, accepted 25 December 2011.
Schellenberg, M. P., Biligetu, B., McLeod, G. J. and Wang, Z. 2012. Phenotypic variation of side-oats grama grass [Bouteloua curtipendula (Michx.) Torr.] collections from the Canadian prairie. Can. J. Plant Sci. 92: 10431048. Native prairie grasses are adapted to local climates and have the potential for development as forage grass cultivars for semiarid environments. Side-oats grama grass [Bouteloua curtipendula (Michx.) Torr.] is a drought-tolerant grass with desirable forage characteristics, and is distributed across the Canadian prairies and throughout the Great Plains. Understanding ecotypic variability of this species is a prerequisite for developing populations suitable for drier regions of western Canada. A randomized complete block field plot was established using nine seed collections from Manitoba and Saskatchewan in 2006 near Swift Current (lat. 50825?N, long. 107844?W), Canada. Seed yield, tiller number, plant height, foliage diameter, and days to flower of individual plants were measured for each collection in the summers of 2007 and 2010. The Sidney and Wolseley collections had taller plants with greater basal diameter. Plants from these two collections also had higher tiller number, and required fewer days to reach flowering stage. However, there was no clear ranking of these variables among other collections. Averaged flowering date was Aug. 15 for Alexander, Sidney, and Wolseley collections, Aug. 21 for Wellwood, Glenboro, and Pratt collections, and Aug. 26 for Minto and Coulter collections, respectively. Taller plants with greater tiller number for the Sidney and Wolseley collections showed a potential for further cultivar development. Key words: Phenotype, ecotype, genetic variation, grama grass Schellenberg, M. P., Biligetu, B., McLeod, G. J. et Wang, Z. 2012. Variation phe notypique du grand boutelou [Bouteloua curtipendula (Michx.) Torr.] dans les collections venant des Prairies canadiennes. Can. J. Plant Sci. 92: 10431048. Les gramine es indige ` nes des Prairies se sont adapte es au climat local et on pourrait les ame liorer en vue dobtenir des cultivars de gramine es fourrage ` res pour les milieux semi-arides. Le grand boutelou [Bouteloua curtipendula (Michx.) Torr.] est une gramine e qui tole ` re la se cheresse et pre sente des caracte ristiques fourrage ` res souhaitables. Elle pousse un peu partout dans les Prairies canadiennes et les grandes plaines. Avant de cre er des populations convenant aux re gions plus se ` ches du Canada, il importe de comprendre la variabilite des e cotypes de lespe ` ce. En 2006, les auteurs ont ame nage une parcelle en blocs randomise s complets pre ` s de Swift Current (50825?N, 107844?O), au Canada, en recourant a ` neuf collections de semences issues du Manitoba et de la Saskatchewan. Ils ont note le rendement grainier, le nombre de talles, la hauteur du plant, le diame ` tre du feuillage et le nombre de jours avant la floraison des plants issus de chaque collection durant le te de 2007 et de 2010. Les collections Sidney et Wolseley comptaient les plants les plus hauts, dun diame ` tre plus grand a ` la base. Les plants de ces deux collections se caracte risaient aussi par des talles plus nombreuses et prenaient moins de temps pour parvenir a ` la floraison. Ne anmoins, ces variables nont pas permis de classer clairement les plants des autres collections. En moyenne, la floraison survient respectivement le 15 aou t pour les plants des collections Alexander, Sidney et Wolseley, le 21 aou t pour ceux des collections Wellwood, Glenboro et Pratt, et le 26 aou t pour ceux des collections Minto et Coulter. Les plants plus hauts et aux talles plus nombreuses des collections Sidney et Wolseley laissent entrevoir la possibilite du de veloppement de cultivars. s: Phe Mots cle notype, e cotype, variation ge ne tique, grand boutelou

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In the semiarid prairies of western Canada, low and erratic precipitation in combination with high transpiration rate contributes to extremely limited water availability most of the growing season. The semiarid region is also one of the main beef cattle production areas in Canada. Soil moisture limitation for forage production is evident during the summer months under this climate.
1

Corresponding author (e-mail: Biligetu@agr.gc.ca).

The prairie region is predicted to be impacted by the changing climate, with Saskatchewan potentially experiencing a 3.58C increase in temperature (World Resources 1990). Genetic enhancement of available forage species or introduction of new forage species is necessary to meet the new climate constraints on forage production. In the prairie provinces of Canada, the majority of grass species are cool-season grasses (Budd et al. 1979), that exhibit the three-carbon (C3) photosynthetic biochemistry. However, many warm-season grasses that
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Can. J. Plant Sci. (2012) 92: 10431048 doi:10.4141/CJPS2011-142

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exhibit the four-carbon (C4) photosynthetic biochemistry are also found in the region, particularly in southern Manitoba and south-eastern Saskatchewan (Jefferson et al. 2002). Cool-season grasses complete most developmental phases during periods of cool weather in the spring and the fall of the year; however, warm-season grass developmental phases occur during the hottest time of the year (Lauenroth and Whitman 1977). The offsetting growth cycles between cool- and warm-season species may be a strategy for reducing inter-specific competition in native prairie habitats (Williams 1974) and increasing plant community production (Lorenz and Rogler 1972). Evaluation of suitable warm-season grasses that are productive during hot, dry summer months may increase forage production in this region. Side-oats grama grass [Bouteloua curtipendula (Michx.) Torr.] is a native, warm-season grass widely distributed in native grasslands of the Great Plains (Newell et al. 1962). It belongs to the Festucoideae subfamily and Chlorideae tribe (Budd et al. 1979). In Canada, a great number of side-oats grama grass populations are distributed in southeast Saskatchewan, western Alberta and southwest Manitoba (Budd et al. 1979). It is a perennial, 70100 cm tall, sod-forming grass with short, scaly rhizomes. It grows well on dry south-facing slopes or sandy plains. Due to its drought tolerance and easy establishment characteristics, side-oats grama grass was widely used for land reclamation during the 1930s drought (Weaver and Albertson 1944). It is also considered to be excellent forage for livestock and wildlife (Stubbendieck et al. 1982). Side-oats grama grass grows well in combinations with other warm-season grasses, and is considered to be an important component of grass mixtures planted for summer forage on fine-textured upland soils (Newell et al. 1962). In addition, side-oats grama grass populations from the Canadian prairies have a high final germination and can germinate under low water availability (Biligetu et al. 2011). The use of side-oats grama grass is presently limited in western Canada due to the limited availability of an adapted seed source. The evaluation of different ecotypes collected from various Canadian prairie regions may provide a great opportunity to select superior lines. The objective of the present study was to compare phenotypic variations of side-oats

grama grass collections from various locations of the Canadian prairie grown in a common nursery. MATERIALS AND METHODS Seeds of side-oats grama grass were collected from nine sites in Manitoba and Saskatchewan, Canada, in 2005 by the field staff of Ducks Unlimited Canada (Table 1). Collection site data included latitude, longitude, soil type, soil moisture condition, and site position (Table 1). In May 2006, a common field nursery was established at the Semiarid Prairie Agricultural Research Centre near Swift Current (lat. 50825?N, long. 107844?W), SK, Canada, from the nine seed collections using seedlings grown in the greenhouse. The soil type was an Orthic Brown Chernozem in the Canadian soil classification (Ayers et al. 1985). Prior to transplanting the seedlings, the field plot was covered by landscape fabric to control weeds. The field plots were arranged on the landscape fabric as a randomized complete block design (RCBD) with four replicates. Row spacing was 60 cm between any two individual plants. In each replicate, 30 plants were included from each of the nine seed collections. Hand weeding was used for weed control. No fertilizer was applied during the study. All plants were harvested to a 5-cm stubble height each September from 2007. Climate data were obtained from Agriculture and Agri-Food Canada real time weather network (Fig. 1). Data Collection At anthesis, plant height, foliage diameter and tiller count data were collected (Moore and Moser 1995). Plant height was measured as the height of the tallest reproductive culm near the center of the plant. Foliage diameter was determined from the average width of two measurements of foliage taken at right angles. Tiller count was made on each individual plant at anthesis stage. The days to flower was recorded as the number of days from May 01 to emergence of the first flower on each plant. Seeds were hand-harvested at maturity, airdried in paper bags, threshed, and cleaned to determine seed yield for individual plants. Seed harvest dates were early September of each year. Data were not collected in 2008 and 2009 due to insufficient technical staffs.

Table 1. Geographic location and site characteristics of site-oats grama grass [Bouteloua curtipendula (Michx.) Torr.] collections in the Canadian prairies Location Alexander, MB Coulter, MB Glenboro, MB Minto, MB Pratt, MB Wolseley, SK Sidney, MB Wawanesa, MB Wellwood, MB Latitude 50829?N 49852?N 49833?N 49824?N 49848?N 50825?N 49853?N 49835?N 50802?N Longitude 96803?W 99821?W 99817?W 10081?W 98854?W 10381?W 99807?W 99841?W 99819?W Year 2005 2005 2005 2005 2005 2005 2005 2005 2005 Soil texture Sandy loam Till loam Sandy loam Sandy loam Sandy loam Gravely loam Sandy loam Sandy loam Sandy loam Soil moisture Dry Mesic Dry Dry Dry Dry Dry Dry Dry Site position South east slope East slope West slope West slope South slope South slope South slope South slope South slope

SCHELLENBERG ET AL. * PHENOTYPIC VARIATION OF SIDE-OATS GRAMA GRASS

160 140 120 2007 2010 30-year means 2007 2010 30-year means 20 25

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Rainfall (mm)

100 80 60 40

15

10

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20 0 Apr May Jun Jul Aug Sep 0

Month

Fig. 1. Monthly mean air temperature (lines) and rainfall (bars) received during the studies in 2007 and 2010 and long-term average at the experiment site near Swift Current, Canada.

Statistical Analysis The analysis of variance (ANOVA) was carried out using the MIXED procedure of SAS (Littell et al. 1996) with collection, year, and their interaction considered as fixed effects and replications considered as a random effect and significance declared at a 0.05. In the model, year was considered a fixed effect because measurements were made on the same plants that continued to grow over the study period, which would cause a difference between two measurements. If no significant year collection interaction (P0.05) was detected for measured variables, values were averaged across 2 yr. When the F-tests from the ANOVA indicated significant differences among collections (P50.05), the means were separated using the Least Square Means comparison. Principal component analysis (PCA) was conducted on means of phenotypic variable data using PROC PRINCOMP procedure of SAS software. Cluster analysis was conducted using PROC CLUSTER procedure of SAS software. RESULTS Environmental Conditions Monthly rainfall during the study was below average in 2007, but it was above average in 2010 (Fig. 1). Compared with the long-term average, growing season total rainfall (AprilSeptember) was 44% less in 2007 and 121% more in 2010. Monthly mean air temperature was above average in 2007, but it was below average in 2010 (Fig. 1). Growing season total growing degree days (base temperature 58C) was 1768 and 1420 in 2007 and 2010, respectively. Plant Height and Foliage Diameter The collection year interaction was significant for plant height and foliage diameter. Therefore, they were

not combined across 2 yr (Table 2). Collections differed significantly for plant height, ranging from 33.8 to 50.9 cm in 2007 and from 31.9 to 62.9 cm in 2010. The Sidney and Wolseley collections had the tallest plants, while the Minto collection had the shortest plants in both study years. Collections also differed significantly for foliage diameter, ranging from 8.3 to 12 cm in 2007 and from 18.4 to 32.4 cm in 2010 (Table 2). The Sidney collection had the widest diameter, followed by the Wolseley collection. In 2010, the Wellwood and Glenboro collections had the narrowest plants. Tiller Number Compared with 2007, the tiller number of side-oats grama grass increased in 2010 (Table 2). The Sidney collection produced the highest tiller number in 2007, followed by Wolseley and Alexander. Collections from Minto, Coulter, and Wellwood had the lowest tiller number in 2007. In 2010, tiller number was highest for the Sidney, Wolseley, Alexander, and Wawanesa collections and lowest for the Wellwood and Glenboro collections. Tiller number for the Minto collection in 2010 increased about eightfold from the 2007 tiller count. Seed Yield and Days to Flower Collections also differed for seed yield plant 1. Seed yield was the highest for the Sidney, Wolseley, Alexander, and Wawanesa collections in 2007. Collections from Wolseley, Alexander, Wawanesa, and Minto had the highest seed yield in 2010 (Table 2). The collection year interaction was not significant for days to flower; thus, means were calculated across 2 yr. Days to flower were least for the Sidney, Wolseley, and Alexander collections, followed by Wawanesa and Wellwood

Air temperature (C)

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Table 2. Phenotypic characteristics of side-oats grama grass [Bouteloua curtipendula (Michx.) Torr.] collections in 2007 and 2010 in a nursery near Swift Current, Canada Foliage width (cm) Location Sidney Wolseley Alexander Wawanesa Coulter Minto Pratt Wellwood Glenboro P SEMx
z

Plant height (cm) 2007 50.9a 50.7a 41.1bc 48.6a 40.6bc 33.8d 46.5ab 41.9bc 39.2cd B0.0001 2.1 2010 62.9a 62.8a 52.8b 44.4cd 31.9f 35.9ef 53.2b 42.6de 51.8bc B0.0001 2.8

Tiller number plant  1 2007 86.3a 61.4b 45.9bc 25.4de 21.9e 14.4e 30.9cde 22.9e 41.4cd B0.0001 6.7 2010 169a 142.6abc 156.7ab 136.3abcd 106.0cde 120.3bcde 119.9bcde 89.5de 71.9e 0.0024 17.1

Seed yield plant  1 (g) 2007 1.1ab 1.5a 1.5a 0.9ab 0.7b 0.6b 0.3b 0.4b 0.7b 0.012 0.3 2010 2.2c 4.1a 4.1a 3.6ab 2.9bc 4.3a 1.8cd 2.2c 0.9d B0.0001 0.4

Days to flower (d)z 2-yr meany 76.2d 76.5d 74.8d 79.2c 84.9a 86.5a 81.4b 80.1bc 81.8b B0.0001 0.8

2007 12.0a 11.5a 9.3b 8.0bc 7.3c 9.5b 8.3bc 9.0bc 8.5bc B0.0001 0.6

2010 32.4a 28.1b 26.7b 23.6c 22.7c 22.3c 21.4cd 18.8de 18.4e B0.0001 1.0

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Days to flower was counted from May 01 in each year. Two-year data for days to flower were averaged due to no significant year collection effect (P0.05). SEM, standard error of means. af Means in a column with the same lower case letter do not differ significantly (P0.05).
y x

collections in both years. The Coulter and Minto collections required more days to reach anthesis stage than Wawanesa and Wellwood. Principal Component Analysis and Cluster Analysis of Phenotypic Variables The first two eigenvalues of the PCA explained more than 91% of the among-collection variation, with the first eigenvalue explaining 72%, and the second explaining 19% (Table 3). The remainder of the variation was explained by the next three eigenvalues. Tiller number (0.51), foliage diameter (0.50) had closer correlations with the first principal component, and seed yield (0.77) and plant height ( 0.55) had closer correlation with the second principal component. The scatter plot based on the first and second principal component scores showed three clustering patterns (Fig. 2). The first group included the Sidney, Wolseley and Alexander collections, the second group included the Minto and Coulter collections, and the Wellwood, Glenboro, and Pratt collections were included in the third group. In the cluster analysis, based on average distance between clusters, the first group consisted of the Sidney, Wolseley and Alexander collections, and the rest
Table 3. Correlation of phenotypic variables with the rst two principal components in principal component analysis Phenotypic variables Foliage diameter (cm) Plant height (cm) Days to flower (d) Seed weight plant  1 (g) Tiller plant  1 Eigenvalue Proportionz
z

was in the second group (Fig. 3), which is in agreement with the principal components analysis results. DISCUSSION Successful stand establishment is a critical step for native prairie plant seeding. The location of the field study (lat. 50825?N, long. 107844?W) is considered suitable to grow warm-season grasses (Jefferson et al. 2002). The latitude of collection sites ranged from 49824?N to 50829?N, which was similar to the field study site. In the present study, seedlings were transplanted to establish a field nursery, and seedling survival was high in spring 2007 and thereafter. Considerable variation was detected among side-oats grama grass collections for all measured variables, which provides an oppotunity for further selection of superior genetic material. Genetic improvement for certain traits such as seed yield, plant vigour, and local adaptation would reduce seed cost and enhance stand establishment. The use of locally adapted seed sources could minimize the risk of establishment failures (Jefferson et al. 2002). The Sidney and Wolseley collections had taller plants with greater basal diameter. Plants from these two collections also had the highest tiller numbers. In a forage grass, these characteristics are always associated with higher forage biomass. In addition, side-oat grama grass possesses relatively high forage quality (Stubbendieck et al. 1982), which also demonstrates its value as a forage grass. The order of ranking for the Sidney and Wolseley collections is relatively consistent even though contrasting growth conditions occurred in 2007 and 2010. Plants from these collections could be used as a base population for future development of forage type side-oats grama cultivar. In our study, side-oats grama grass collections varied for flowering date. The differences in latitude among

Prin 1 0.5 0.43 0.45 0.32 0.51 3.58 0.72

Prin 2 0.08 0.55 0.27 0.77 0.14 0.97 0.19

The proportion of each principal component in the total variation.

SCHELLENBERG ET AL. * PHENOTYPIC VARIATION OF SIDE-OATS GRAMA GRASS

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Fig. 2. Scatter plots of the rst principal component score (Prin1, represent 72% variation) with the second principal component score (Prin2, represent 19% variation) obtained from phenotypic data of side-oats grama grass collections.

collection sites was relatively small, but the geographic distance (data not shown) between two sites ranged from 20 km (minimum) to 326 km (maximum). The geographic distance, however, may not be a major factor affecting phenotypic variation in this study. For example, the Sidney and Wolseley collections had

similar maturity although distance between these two sites is more than 326 km. Phan and Smith (2000) found that warm-season grass collections from higher latitudes tend to reach anthesis stage earlier. Flowering date was positively correlated with temperature and negatively correlated with elevation (Bhattarai et al. 2010).

Fig. 3. A dendrogram of nine side-oats grama collections constructed from average distance of collections based on phenotypic data.

1048 CANADIAN JOURNAL OF PLANT SCIENCE

High-elevation sites typically are cooler than lowelevation sites; thus, plants from high elevations are generally adapted to lower temperature (Bhattarai et al. 2010). Flowering date for the collections ranged from mid- to late-July in both study years. In western Canada, cool-season grasses reach anthesis and peak biomass by the end of June (Jefferson et al. 2002). This timing of the growth of side-oats grama grass can provide an alternative forage source during hot summer months. CONCLUSIONS The Sidney and Wolseley collections were superior in characteristics related to high forage production and were early maturing. These would be appropriate base populations in which to select forage types of side-oats grama grass adapted to a more northern climate. ACKNOWLEDGEMENTS We thank R. Muri and P. Coward at the Semiarid Prairie Agricultural Research Centre (SPARC) of Agriculture and Agri-Food Canada (AAFC), Swift Current, Canada, for their technical assistance during plot establishment and management. Thanks to M. Kehler at SPARC-AAFC for her help during editing of this article. We also thank Agriculture and Agri-Food Canada and the Beef Industry Science Cluster for funding support.
Ayers, K. W., Acton, D. F. and Ellis, J. G. 1985. The soils of the Swift Current map area 72J Saskatchewan. Saskatchewan Institute of Pedology, Extension Division, University of Saskatchewan, Extension Publication 86. Saskatoon, SK. 48 pp. Biligetu, B., Schellenberg, M. P. and McLeod, G. J. 2011. The effect of temperature and water potential on seed germination of poly-cross side-oats grama [Bouteloua curtipendula (Michx.) Torr.] population of Canadian prairie. Seed Sci. Technol. 39: 7481.

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Bhattarai, K., Bushman, B. S., Johnson, D. A. and Carman, J. G. 2010. Phenotypic and genetic characterization of western prairie clover collections from the western United States. Rangeland Ecol. Manage. 63: 696706. Budd, A. C., Looman, J. and Best, K. F. 1979. Budds ora of the Canadian prairie provinces. Research Branch, Agriculture Canada, Ottawa, ON. Publication 1662. 863 pp. Jefferson, P. G., McCaughey, W. P., May, K., Woosaree, J., MaFarlane, L. and Wright, S. M. B. 2002. Performance of American native grass cultivars in the Canadian prairie provinces. J. Native Plant. 3: 2433. Lauenroth, W. K. and Whitman, W. C. 1977. Dynamics of dry matter production in a mixed- grass prairie in western North Dakota. Oecol. (Berl.) 27: 339351. Littell, R. C., Milliken, G. A., Stroup, W. W. and Wolnger, R. D. 1996. SAS system for mixed models. SAS Institute, Inc., Cary, NC. Lorenz, R. J. and Rogler, G. A. 1972. Forage production and botanical composition of mixed prairie as inuenced by nitrogen and phosphorus fertilization. Agron. J. 64: 244249. Moore, K. J. and Moser, L. E. 1995. Quantifying developmental morphology of perennial grasses. Crop Sci. 35: 3743. Newell, L. C., Staten, R. D., Jackson, E. B. and Conard, E. C. 1962. Side-oats grama in the central Great Plains. Nebr. Agric. Res. Bull. 207, Lincoln, NE. Phan, A. T. and Smith, S. R. 2000. Seed yield variation in blue grama and little bluestem plant collections in southem Manitoba, Canada. Crop Sci. 40: 555561. Stubbendieck, J., Hatch, S. L. and Kju, K. J. 1982. North American range plants. 2nd ed. University of Nebraska Press, Lincoln, NE. 464 pp. Weaver, J. E. and Albertson, F. W. 1944. Nature and degree of recovery of grassland from the great droutht of 1933 to 1940. Ecol. Monogr. 14: 393479. Williams III, G. J. 1974. Photosynthetic adaptation to temperature in C3 and C4 grasses. Plant Physiol. 54: 709711. World Resources 1990 1991. A guide to the global environment. Oxford University Press, New York, NY. 97 p.