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Morphological and Genetic Variation Among Four High Desert

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Morphological and Genetic Variation Among Four High Desert HORTSCIENCE 47(6):715–720. 2012. Drought-tolerant plants, including native species adapted to arid or semiarid regions, are key elements for successful low-water Morphological and Genetic landscapes because they require little main- tenance, provide a natural look to the land- Variation among Four High scape (Cane and Kervin, 2009; Kjelgren et al., 2009; McKinney, 2002), and honor Desert Sphaeralcea Species natural habitats. Native plant landscaping also supports local economies, facilitating sus- Chalita Sriladda1 tainability in urban systems at multiple levels. Department of Plants, Soils, and Climate, Utah State University, 4820 Old However, lack of native plant availability, Main Hill, Logan, UT 84322 cost of native plant materials, inconsistent and unreliable demand, and lack of production Heidi A. Kratsch knowledge have minimalized the native plant University of Nevada Cooperative Extension, University of Nevada, 4955 market (Peppin et al., 2010). The U.S. Intermountain West (IMW) is arid Energy Way, Reno, NV 89502 to semiarid and has an abundance of woody Steven R. Larson and herbaceous perennial native plants that are drought-tolerant and ornamentally attractive Forage and Range Research Laboratory, USDA-ARS, Utah State University, (Intermountain Native Plant Growers Associa- Logan, UT 84322 tion, 2012; Kratsch, 2011; Mee et al., 2003; Meyer et al., 2009). The genus Sphaeralcea Roger K. Kjelgren (Malvaceae), commonly known as globemal- Department of Plants, Soils, and Climate, Utah State University, 4820 Old low, is an annual–perennial herb or shrub Main Hill, Logan, UT 84322 wildflower characterized by brilliant, largely orange, flowers. Approximately 40 species of Additional index words. Sphaeralcea coccinea, Sphaeralcea grossulariifolia, Sphaeralcea the genus are found throughout temperate munroana, Sphaeralcea parvifolia, globemallow, AFLP North and South America (Holmgren et al., Abstract. The herbaceous perennial species in the genus Sphaeralcea have desirable 2005). The genus Sphaeralcea includes 27 drought tolerance and aesthetics with potential for low-water use landscapes in the species in North America, from southern Intermountain West. However, taxonomy of these species is ambiguous, which leads to Canada through the western United States decreased consumer confidence in the native plant nursery industry. The goal of this to northern Mexico, with disjunct popula- study was to test and clarify morphological and genetic differentiation among four tions in temperate South America (Holmgren putative Sphaeralcea species. Morphological characteristics of the type specimens were et al., 2005). Several of these species appear used as species references in canonical variate analysis to generate a classification model. to have both aesthetic and drought-tolerance This model was then used to assign putative species names to herbarium voucher qualities that lend them well to low-water specimens and to field-collected voucher specimens to clarify genetic variation among landscaping (Intermountain Native Plant species. Field specimens were also classified using Bayesian cluster analyses of amplified Growers Association, 2012; Mee et al., fragment length polymorphism (AFLP) genotypes. Sphaeralcea coccinea (Nutt.) Rydb. 2003); however, they are ostensibly closely and S. grossulariifolia (Hook. & Arn.) Rydb. formed a composite group morphologically related and difficult to distinguish. Four and genetically distinct from the S. munroana (Douglas) Spach and S. parvifolia A. Nelson promising low-water landscape species, S. composite group. Each composite group displayed genetic isolation by geographic coccinea, S. grossulariifolia, S. munroana, distance. Also, morphological traits of S. munroana and S. parvifolia correlated to and S. parvifolia, are commonly found in the geographic distance. Taken together these results suggest that our samples represent two IMW (Holmgren et al., 2005; Ring, 2005; sympatric yet reproductively isolated groups. Distinguishing between these two Sphaeralcea Ring and Cully, 2007; Ring et al., 2009), and composite groups can create greater consumer confidence in plant material developed they are particularly difficult to distinguish for use in Intermountain West low-water landscaping. from one another (Holmgren et al., 2005). Polyploidy and hybridization have been reported as factors contributing to weak mor- phological differences among species in the Globally, urbanization has dramatically facing the additional challenge of absolute genus (La Duke and Northington, 1978). Tate increased with nearly half of the world’s water scarcity (Vorosmarty et al., 2000). (2002) suggested the possible importance of population currently living in urban areas; it Demand for urban green spaces, including geographical isolation for speciation. Analy- is expected that approximately three-fourths urban landscapes, is increasing with popula- sis of nuclear ribosomal DNA internal tran- of the world’s population will be living in tion growth because those spaces provide scribed spacer (ITS) sequences showed that urban areas by the year 2025 (United Nations, aesthetic and mental health benefits and a the North American Sphaeralcea species 2012). Substantial increases in urban popu- positive association with the perceived gen- [S. angustifolia (Cav.) G. Don and S. wrightii lation lead to increased demand for water in eral health of residents (Maas et al., 2006). A. Gray] cluster together as do the two South commercial, industrial, and residential sectors Urban green spaces also mitigate a number American taxa (S. crispa Hook. ex Baker f. (Foster and Beattie, 1979). This increased de- of undesirable environmental effects such as and S. philippiana Krapov.) (Tate, 2002). mand in arid and semiarid urban regions means urban heat islands (Deloya, 1993). Many urban Intergradation among species challenged green spaces require irrigation, particularly in early taxonomists in their efforts to clearly arid and semiarid areas. Thus, increasing irri- identify Sphaeralcea species using only mor- gation for urban landscapes is causing an in- phological distinctions. Received for publication 16 Feb. 2012. Accepted creased demand for more efficient watering These four IMW species are being pro- for publication 10 Apr. 2012. systems. Low-water landscaping describes moted by the nursery industry (Intermountain We thank Linnea Johnson for help in genetic laboratory works. We thank Mary Barkworth landscapes full of plants that use substantially Native Plant Growers Association, 2012), but and Michael Piep for facilitating the loan of type less water. Therefore, low-water landscaping it is difficult to determine which species are specimens. is a key tool for conserving water in irrigated actually being sold. Difficulty in determining 1To whom reprint requests should be addressed; urban landscapes, particularly in arid and species decreases consumer confidence in e-mail [email protected]. semiarid areas (Kjelgren et al., 2009). these species. Leaf morphology was used as HORTSCIENCE VOL. 47(6) JUNE 2012 715 the first taxonomic key to separate these species into two groups (Holmgren et al., 2005). Shallowly lobed leaves putatively sep- arate S. munroana and S. parvifolia from S. grossulariifolia and S. coccinea, whose leaves are deeply lobed, cleft more than halfway to the base, and, in many, cleft to the base. However, S. munroana is often difficult to distinguish from S. parvifolia, whereas S. grossulariifolia can be easily confused with S. coccinea and some specimens of S. munroana (Holmgren et al., 2005). Besides leaf morphology, the mature fruiting carpel characteristics, density of leaf hairs, and hair ray orientation sometimes are used as keys for taxonomists to identify specimens (Atwood and Welsh, 2002). How- ever, these characters are seldom used in practical settings and are not efficient for field use or for non-taxonomist collectors. Theinter- grading morphology of these species might be a result of genetic overlap and makes selection for superior forms difficult. In terms of the geological distribution of the four species (United States Department of Agriculture, 2012), S. coccinea is found extensively throughout the IMW, and S. grossulariifolia is found throughout the Great Basin, generally toward the western part of the IMW. Distribution of S. munroana and S. parvifolia overlaps in Nevada, Utah, and Colorado, where S. parvifolia is distributed further south and S. munroana distributed further north. In Utah, where the distributions of the four species overlap, geographical differences seem to play an important role in putative species distribution, particularly for S. munroana and S. parvifolia, which occur in the north and south, respectively Fig. 1. Geographical distribution of Sphaeralcea species in Utah used in this study; letters were used to (Shultz et al., 2012). The distribution has indicate species names; capital letters represent herbarium specimens (C = S. coccinea;G=S. sometimes been used as a key for species grossulariifolia;M=S. munroana;P=S. parvifolia); lower-case letters represent field-collected identification because of identification diffi- specimens (c = S. coccinea;g=S. grossulariifolia;m=S. munroana;p=S. parvifolia). culty resulting from morphological overlap among these species. Difficulty in their accurate identification creates concerns for those collecting Sphaeralcea populations were no longer present at many specimens to one of four putative Sphaer- seed for nursery production and reduces con- of the field sites. We were
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