20.5 Seashore Paspalum

From PlantBreeding
Revision as of 13:23, 15 December 2013 by Plantbreedingadmin (Talk | contribs)

(diff) ← Older revision | Latest revision (diff) | Newer revision → (diff)
Jump to: navigation, search

by Clinton J. Steketee, Institute of Plant Breeding, Genetics & Genomics, University of Georgia, Athens, GA; Edited by Paul Raymer, Institute of Plant Breeding, Genetics & Genomics, University of Georgia - Griffin Campus, Griffin, GA

Abstract

Seashore paspalum (Paspalum vaginatum Swartz) is a warm-season perennial turfgrass used primarily as a fine-turf in recreational areas such as golf courses and athletic fields. This species is inherently tolerant to drought and salt conditions, and has been touted as an environmentally friendly turfgrass that is cheaper to manage compared to salt-sensitive turfgrass species in similar saline environments. Grown only in coastal areas or salt-laden soils until recent years, Paspalum vaginatum is currently being used on a variety of turfgrass sites, and has been shown to have turfgrass quality similar to, or better than other warm-season turfgrass species. Seashore paspalum was once considered a niche turfgrass and marketed as such, but is now becoming more popular in the turfgrass industry, as the focus on sustainability has grown. As water quantity and quality issues continue to affect turfgrass managers, the demand for Paspalum vaginatum is expected to increase as it is accepted as a solution to these water concerns. Seashore paspalum breeding and the development of improved cultivars will be of paramount importance for the turfgrass industry in the coming years.


Introduction

Seashore paspalum (Paspalum vaginatum Swartz) is a warm-season perennial turfgrass (Morton, 1973) that is used primarily as a fine-turf in recreational turfgrass systems (Figure 1). This littoral, C4 (Chapman and Peat, 1992) species of turfgrass is typically found in subtropical and tropical to warm temperate regions near 30 degrees N-S latitudes (Skerman and Riveros, 1990). It is naturally found on the banks of coastal areas and sandy beaches, but can also be found inland growing in saline environments (Duncan and Carrow, 1999). Seashore paspalum possesses several morphological characteristics that make it a suitable turfgrass, such as tolerance to mowing, dark green color, and both a rhizomatous and stoloniferous spreading growth habit (Webster, 1987). Most importantly, it also has high tolerance to salt and is well adapted to saline growing conditions (Morton, 1973; Duncan and Carrow, 1999; Lee et al., 2005). Seashore paspalum is considered to be the most salt-tolerant of all the warm-season turfgrass species (Carrow and Duncan, 1998); salt-tolerant grasses are considered an important component of the solution to growing water quantity and quality issues that will face turfgrass managers. As alternative water sources such as seawater blends, effluent, and saline ground water have become more widely used on recreational turfgrass systems, the demand for and focus on this turfgrass species has increased dramatically.

Figure 1 Seashore paspalum used on a golf course putting green. Photo credit: Paul Raymer, unpublished.

Seashore paspalum has been touted as an environmentally friendly turfgrass, and can cost half as much to manage as salt-sensitive turfgrass species in similar saline environments (Duncan and Carrow, 1999). This turfgrass species has high tolerance to multiple environmental stresses, including salinity, drought (Huang et al., 1997), low light intensity (Jiang et al., 2004), and soils with hypoxia conditions (Duncan and Carrow, 1999). In addition, Duncan and Carrow (1999) stated that its turfgrass quality (combination of density, uniformity, color, density, texture, and disease or environmental stress tolerance (Morris and Shearman, 2008)) traits are similar to, or better than alternative turfgrass species, especially when grown in high stress environments. This species was once considered a niche turfgrass for coastal environments, but is now becoming increasingly environmentally and economically significant in the turfgrass industry. Seashore paspalum breeding and the development of improved cultivars will be of great importance for the turfgrass industry in the coming years.


Background and History of the Species

Botanists do not agree upon the center of origin for Paspalum vaginatum. Some individuals consider it native to Asia, Africa, and Europe (Judd, 1979), while other botanists consider it indigenous to the New World (Americas), and then later adapted to the Asia, Africa, and Europe (Bovo et al., 1988). Chen et al. (2005) supports the theory that South Africa is the center of origin, as their studies show that accessions from this country have the most genetic diversity. Seashore paspalum was introduced throughout the world as a result of its use as bedding on ships used during the slave trade between the Americas and African countries in the eighteenth and early nineteenth centuries (Gray, 1933). It has also spread due to its ability to reclaim salt-affected areas in other parts of the world (Casler and Duncan, 2003).

Fine-leaf seashore paspalum is a popular choice for golf courses, athletic fields, and home lawns or landscape areas. It has also been used for forage production, reclamation of salt-affected areas, soil stabilization, and erosion control in other instances. Paspalum vaginatum is especially prevalent in sites that utilize alternative irrigation sources or are in close proximity to a coastal area or the sea (Duncan and Carrow, 1999).


Taxonomy and Morphology

Paspalum vaginatum Swartz is commonly known as seashore paspalum, or most simply paspalum. Other common names that are less frequently used include sand knotgrass, siltgrass (Morton, 1973), and saltwater couch (Duncan and Carrow, 1999). Several other scientific names, primarily Paspalum distichum, have also been used in publications since 1788 (Casler and Duncan, 2003). The nomenclature committee for Spermatophyta selected Paspalum vaginatum Swartz as the official scientific name in 1983 (Brummitt, 1983). Seashore paspalum is classified taxonomically as Poaceae (Family), Panicoideae (Subfamily), Panicodae (Supertribe), Paniceae (Tribe), Setariinae (Subtribe), Paspalum (Genus), Disticha (Group), vaginatum (Species), and Swartz (Authority) (Duncan and Carrow, 1999).

Paspalum vaginatum has a chromosome number that is predominately diploid (2n = 2x = 20) (Bashaw et al., 1970; Burson, 1983). However, several reports of tetraploid (2n = 4x = 40) and hexaploid (2n = 6x = 60) accessions have been documented in literature (Casler and Duncan, 2003). Seashore paspalum has the D genome (Bashaw et al., 1970; Burson, 1983), making interspecific crosses with other Paspalum species, and production of viable seed very challenging.

Morphologically, seashore paspalum has both stoloniferous and rhizomatous growth habits (Webster, 1987), with the stolon nodes being noticeably pubescent. It also possesses conduplicate (folded) vernation (Green et al., 2012), and auricles are absent on the plant. Flowering culms (stems) are basally decumbent or erect, and can vary in height from 8 to 60 cm with 5-13 hairless nodes. The leaves of the plant arise on opposite sides of the stem; two vertical rows of leaves are present, and are approximately 50-220 mm long by 1-4 mm wide. The ligule is around 1 mm in diameter, membranous, and hairy. The inflorescence (seed head) commonly consists of two racemes (branches) that are 20-60 mm in length; each of these branches has around 16-32 twin-rowed spikelets. Anthers are 1.2 to 1.6 mm in length, and are yellow and purple striped, while stigmas are feathery and purple (Figures 2 and 3). Paspalum vaginatum seeds (caryopses) are 1.5 mm wide, and 2.5-3.0 mm long, ovate with the narrower end at the base, and concave on one side, while convex on the other (Casler and Duncan, 2003). The weight of one gram of seed is equal to approximately 1760 seeds (Raymer, personal communication, 2012). Seashore paspalum leaf textures range from very coarse (forage types) to fine-leaf types (golf course types).


Figure 2 Mature seed head (inflorescence) with two racemes and both anthers (white) and stigmas (purple) emerged. Photo credit: Paul Raymer, unpublished.
Figure 3 Inflorescence of Paspalum vaginatum in early anthesis, prior to the anther development. Making crosses is ideal during this stage. Photo credit: Clinton Steketee, unpublished.


Germplasm and Cultivars

In 1993, Dr. R. R. Duncan initiated the first seashore paspalum breeding program at the University of Georgia Griffin Campus (Raymer et al., 2008), and continued his work until 2003. During his tenure with the breeding program, he amassed a collection of approximately 250 accessions and began to intensely breed these ecotypes for use in turfgrass. Dr. P. L. Raymer has continued this work since 2003, and maintains what is currently the largest collection of seashore paspalum germplasm (Raymer, personal communication, 2012). Several smaller collections exist in Argentina, Brazil, the University of Florida, and the National Genetic Resources Program (NGRP) of the USDA-ARS in Griffin, Georgia (Casler and Duncan, 2003).

The University of Georgia seashore paspalum breeding program has released several improved cultivars of seashore paspalum since its inception in 1993. Duncan (2003) provides a complete list of seashore paspalum cultivars and their origin prior to 2001. Differentiating these varieties is often difficult, because most commercially grown cultivars are fairly morphologically similar. Amplified fragment length polymorphism (AFLP) is used as a tool to identify cultivars for breeding programs, and to protect intellectual property rights (IPR). The use of simple sequence repeats (SSR) is also increasing being used and provides an additional tool for genotype identification (Raymer et al., 2008).


Reproduction Characteristics and Propagation

Seashore paspalum is a sexually reproducing, cross-pollinating diploid that possesses a degree of self-incompatibility (Carpenter, 1958). Espinoza and Quarin (1997) determined the self-incompatibility is gametophytic, when their studies showed that pollen germinated and penetrated the stigma, but failed to grow in the style. Paspalum vaginatum is heterozygous and rarely cross-pollinates with other paspalum species, because of its D genome. In some instances, crossing closely related species can be accomplished during the early morning hours (0500-0730) at a temperature of around 18-21°C (Burson, 1985). These crosses, with Paspalum vaginatum as the pollen parent, often are not successful, and typically produce sterile hybrids. However, Paspalum vaginatum is capable of successful cross-pollination with other ecotypes and accessions of seashore paspalum, creating genetic recombination that can be utilized in a breeding program (Shin et al., 2006).

Paspalum vaginatum is normally vegetatively propagated from sod or sprigs, but one commercially marketed seeded cultivar is available. Spriging rates can vary from 19-76 m3/ha (5 to 20 bushels per 1000 ft2) (Raymer, personal communication, 2012). In general, seashore paspalum establishes slightly slower than bermudagrass, but is comparable to most other warm-season turfgrasses due to it stoloniferous and rhizomatous growth habit (Duncan and Carrow, 1999). Seaspray® is the only commercially available seeded cultivar at this time and should be seeded at rates between 4.9-9.8 g/m2 (1 to 2 lbs./1,000 ft2) (Raymer, personal communication, 2012).


Breeding Strategies, Techniques, and Hybridization

Until recent years, warm-season turfgrass cultivar development, including seashore paspalum improvement, was rather difficult due to a lack of information on how to make successful crosses. However, laboratory research has helped to determine the proper conditions to make sexually compatible seashore paspalum crosses that produce seeds that lack dormancy and are viable (Shin et al., 2006). Given this information, The University of Georgia seashore paspalum breeding program is able to utilize traditional breeding techniques to hybridize ecotypes, and create genetic diversity that can be used in the development of new varieties.

The creation of F1 hybrids can be done using hand pollination or by using open-pollinated polycrosses. Hybridization usually takes place in a greenhouse rather than in a field setting. Hand pollinations are performed by following these steps: 1) cover racemes (flowers) with small glassine bags supported by wire stakes prior to stigma appearance, 2) transfer pollen from male plant flowers to desired female parent after stigmas have emerged, 3) replace bags over racemes after hand pollination (Figure 4), and 4) wait for viable seed to develop (approximately 1 month) (Raymer, personal communication, 2012). Successful crosses will yield around 2-10 F1 seed that are then harvested by cutting off flower racemes, and threshing seeds after they have dried down. These seeds are then germinated (Figure 5) and grown into mature seashore paspalum plants. The potted plants are then placed in nylon mesh cages (Figure 6), and full-sib mated to produce hundreds of unique individuals per family. Placing flowering mature Paspalum vaginatum plants near one another for approximately 6 to 8 weeks creates F1 hybrids from polycrosses with an unknown male parent. Mass selection is then used to identify desirable phenotypes (Raymer, personal communication, 2012).


Figure 4 Seashore paspalum crosses after hand pollination. Photo credit: Paul Raymer, unpublished.


Figure 5 Germination of Paspalum vaginatum lines after initial crosses using germination boxes (stacked on left side of table). Photo credit: Paul Raymer, unpublished.


Figure 6 Nylon mesh cages used for full-sib mating. Photo credit: Paul Raymer, unpublished.


In vitro propagation has been used recently to obtain sethoxydim resistance in Paspalum vaginatum (Heckart et. al, 2010). Callus (Figure 7) was plated on an induction medium that contained sethoxydim to select for mutant herbicide resistant cells. This method provides an alterative to genetic modification (GM) to manage bermudagrass and other weedy grasses with herbicide resistance technology. Very little molecular marker technology is currently utilized in seashore paspalum breeding, but the use of simple sequence repeats (SSRs) will likely increase in the near future (Wang et al., 2006), and should help make development of marker assisted selection possible for major traits of interest. At this time, genetic modification is not used in seashore paspalum breeding programs due to pollen escape concerns, and regulatory issues.


Figure 7 Seashore paspalum callus tissue used with in vitro propagation. Photo credit: Paul Raymer, unpublished.

Breeding for Abiotic Stress Resistance

Seashore paspalum’s coastal origin exposed the species to several abiotic stresses during its evolution that enabled it to develop numerous stress tolerance and resistance mechanisms. Close approximation to coastal areas, seawater sprays, and some instances of salt waterlogging has made Paspalum vaginatum inherently more salt tolerant than their warm-season turfgrass counterparts in most cases (Duncan and Carrow, 1999). However, Lee et al. (2004) demonstrated that different accessions of seashore paspalum vary greatly in their tolerance to salt with some lines being highly tolerant and others performing no better than a typical hybrid bermudagrass. The trend for turfgrasses to be increasingly irrigated with water that has higher salinity concentrations, and development of golf courses closer to the ocean will drive the need for the next generation of salt tolerant seashore paspalum varieties (Raymer et al., 2008). Screening for salt tolerance (Figure 8) early in the breeding program is a necessary component of breeding for multiple stress resistance and development of improved cultivars. Raymer et al. (2005) describes an effective and efficient procedure to screen for this abiotic stress.


Figure 8 Comparison of two salt sensitive (left) and tolerant (right) seashore paspalum plants following high salt level screening. Photo credit: Paul Raymer, unpublished.


Wear and traffic tolerance is also an important abiotic stress that needs to be selected for in Paspalum vaginatum. Wear and traffic injury tolerance consists of how well a turfgrass stand responds to plant tissue tearing, or physical destruction of an area of turf from events such as divots taken by a golf club on a fairway, cleat marks on an athletic field, and maintenance vehicles driving repeatedly over the same area of seashore paspalum (Duncan and Carrow, 1999). Carrow and Petrovic (1992) showed that turfgrasses with higher shoot and root densities in turn have the best tolerance to traffic and wear stresses, and fill in damaged areas quicker than slow growing, less dense turfgrasses. To test for these stresses, equipment such as the Brinkman traffic simulator can be used to simulate traffic and wear on a turfgrass stand at varying levels of intensity (Cockerham et al., 1990).

Breeding for drought resistance and avoidance in Paspalum vaginatum will be necessary with escalating water quality and quantity concerns (Carrow et al., 1990). According to Huang et al. (1997), seashore paspalum lines have significant genetic diversity in respect to their drought tolerance, and can be assessed using a stress index developed prior to these studies. Other future breeding objectives could focus on cold temperature and shade tolerance of different Paspalum vaginatum ecotypes. Tolerance to mowing at heights that range from greens (3-5 mm) to roughs (25-50 mm) is also a necessary trait that an improved seashore paspalum cultivar must possess (Duncan and Carrow, 1999).


Breeding for Biotic Stress Resistance

Seashore paspalum is planted and maintained perennially using a single commercially available cultivar on golf courses, athletic fields, and for other recreational and aesthetic uses. This uniformity, in conjunction with stresses induced from routine mowing, and the high amounts of clippings and litter present in the turfgrass system from this management practice, can lead to disease problems and epidemics in Paspalum vaginatum (Casler and Duncan, 2003). However, seashore paspalum is not as susceptible to diseases as many other warm-season turfgrasses, likely due to its littoral evolvement that exposed the species to multiple pathogens in a moist environment (Duncan and Carrow, 1999). Seashore paspalum is also no longer exclusively grown on these coastal areas, and when grown inland, may not have the advantage of using salt to manage pathogens and weeds (Duncan and Carrow, 1999). Dollar spot, caused by Sclerotinia homoeocarpa F.T. Bennett, is a major fungal disease that causes turf quality and playability issues. Screening by Raymer et al. (2008) showed substantial differences in dollar spot resistance among eight standard seashore paspalum varieties. Germplasm with resistance can be used as parents in the breeding program to develop improved cultivars with better resistance than what is currently available. Screening for pathogens, insects, and nematodes early in cultivar development, as well as in regional and National Turfgrass Evaluation Program (NTEP) trials, are necessary to produce Paspalum vaginatum lines with resistance to multiple biotic stresses (Raymer, personal communication, 2012).

Experimental Design for Breeding Efforts

Experiments are conducted to select grass genotypes that could later be released as a variety. Most of these trials involve screening for a particular trait of Paspalum vaginatum. Plot size can vary from a pot containing a single clonal propagation early in cultivar development, to larger (6 m2) plots during collaborative, regional evaluation. In the initial field trials, only one replication may be available or used, but several (at least 4 replications) should be planted of the best lines closer to release. Testing improved lines in NTEP also provides replication across multiple locations. Industry standard cultivars of seashore paspalum and other warm-season turfgrass species should be planted with experimental lines as a control (Raymer, personal communication, 2012).

Phenotyping and Data Collection

Phenotyping is often done using visual ratings throughout the course of a breeding program or study according to NTEP guidelines. Before making any ratings, it may be helpful to identify the best and worst performing seashore paspalum lines as a reference. Visual ratings are taken for most aspects of turf on a 1 to 9 scale, with one performing the poorest and nine being the highest rating. Many raters only use whole numbers, as it may be difficult to differentiate similar performing lines with the human eye. Turfgrass quality ratings (6 is acceptable) must take into account the functional and aesthetic aspects of the turfgrass stand, and what the seashore paspalum cultivar will be used for upon commercial release. Turfgrass density reflects the number of plants or Paspalum vaginatum tillers in a given area, with a rating of 9 equaling maximum density. Pests in a seashore paspalum line can also be measured on the 1 to 9 scale, with 1 reflecting 100% injury from the disease or insect, and 9 reflecting 0% injury and a possible resistant line (Morris and Shearman, 2008). Other ratings can be taken for color, percent cover, or traffic and drought tolerance in a similar fashion.

Data from salt screening at various levels of saline water concentration can be obtained by trimming the Paspalum vaginatum lines to a specified height, collecting and drying clippings, recording the dry weight, and then comparing the results to other breeding lines. Greater clipping dry weights is generally a good indication that a possible cultivar performs well in various salt laden conditions over time, but visual ratings, much like those described by NTEP, can be valuable data as well (Raymer, personal communication, 2012).

Richardson et al. (2001) introduced a way to quantify turfgrass cover and color (Karcher and Richardson, 2003) using digital image analysis (DIA). This technique utilizes four incandescent light bulbs inside a light box that are powered by a portable battery or generator (Figure 9), with all components attached to a dolly cart. A small circular hole is on top of the box that allows a camera to take pictures using only the light provided by the light bulbs inside the box. These images are then processed using different macros in SigmaScan Pro (SPSS Inc., 1998) to provide turfgrass breeders with unbiased ratings of their plots for several different facets of turfgrass quality. Numerous other phenotyping equipment options are available for use in a turfgrass breeding program, but are not included in this guidebook for simplicity reasons.


Figure 9 Light box, camera, and portable power source used to collect images for digital image analysis. Photo credit: Paul Raymer, unpublished.


Cultivar Development

Promising seashore paspalum lines are screened in several phases for multiple abiotic and biotic stress resistances in environmental conditions that emulate where they will be utilized. These screening phases occur after initial F1 populations are created from hand pollination or poly-crosses. After the full-sib mating, approximately 5,000 individual plants are tested for their tolerance to saline water. The visual quality of unreplicated experimental Paspalum vaginatum lines that survive high concentrations of salt are assessed, and then around 2,000 lines are planted into a single plot (space dependent, but no less than 5 m2) at a single location. These plots (Figure 10) are managed rather intensively to find lines that perform well under typical turfgrass growing conditions (routine mowing, as well as regular irrigation and fertilizer applications). Each individual plot is rated monthly for overall turfgrass quality and other factors, depending on the main breeding objectives for creating a new cultivar. Approximately 3-5% (60 to 100) of the best individuals are then selected for further evaluation. These high performing lines are then clonally propagated, tested in replicated greenhouse salt screens, and the best 30 to 50 lines are tested in larger replicated plots (6 m2), with specific management protocols at multiple locations. After two to three years of intensive evaluation, one to three lines may be selected for more extensive evaluation, and under various management protocols. In this stage, prospective cultivars might be entered in NTEP trials, and established on golf courses for evaluation and demonstration. Breeder stock of candidates for release as cultivars begins with a single stolon clonally propagated in the greenhouse, until sufficient material is available to plant a field block of breeder’s material. Plants in this block are closely monitored for off-types and contaminates of other species, and eventually used to develop a larger planting of foundation material, no less than one hectare in size. Upon successful release of the cultivar, foundation plant material is used to establish certified commercial production fields on a sod producer’s farm, which is licensed to then sell the new cultivar to golf courses and other customers. (Raymer, personal communication, 2012).


Figure 10 Single plant nursery of promising seashore paspalum lines. Photo credit: Paul Raymer, unpublished.


References

Bashaw, E., A. Hovin, and E. Holt. 1970. Apomixis, its evolutionary significance and ulitization in plant breeding. Proceedings of the XI International Grassland Congress held at Surfers Paradise, Queensland, Australia, 13-23 April 1970. pp. 245-48.

Bovo, O., L. Mroginski, C. Quarin, and Y. Bajaj. 1988. Paspalum spp. Biotechnology in agriculture and forestry 6. Crops II.:495-503.

Brummitt, R. 1983. Report of the Committee for Spermatophyta: 25. Taxon 32:279-284.

Burson, B. 1983. Phylogenetic investigations of Paspalum dilatatum and related species, 14º Int. Grassland Congress p. pp. 170-173.

Burson, B.L. 1985. Cytology of Paspalum chacoence and P. durifolium and their relationship to P. dilatatum. Bot. Gaz.:124-129.

Cardona, C. and R. Duncan. 1997. Callus induction and high efficiency plant regeneration via somatic embryogenesis in paspalum. Crop Sci. 37:1297-1302.

Carpenter, J. 1958. Production and use of seed in seashore paspalum. Journal of the Australian Institute of Agricultural Science 24:252-256.

Carrow, R.N. and R.R. Duncan. 1998. Salt-affected turfgrass sites: assessment and management Ann Arbor Press Chelsea, MI.

Carrow, R. N. and A.M. Petrovic,. 1992. Effects of traffic on turfgrasses. Agronomy. A series of monographs-American Society of Agronomy (32), 285.

Carrow, R.N., R.C. Shearman, and J.R. Watson. 1990. Turfgrass. Agronomy.

Casler, M.D. and R.R. Duncan. 2003 Turfgrass biology, genetics, and breeding Wiley.

Chapman, G.P. and W. Peat. 1992. An introduction to the grasses (including bamboos and cereals) CAB International.

Chen, Z., M. Newman, K. Wook, M. Wang, and P. Raymer. 2005. Molecular characterization of genetic diversity in the USDA Seashore Paspalum germplasm collection. ITS Research Journal 10:543-549.

Cockerham, S.T., V.A. Gibeault, J. Van Dam, and M.K. Leonard. 1990. Tolerance of several cool-season turfgrasses to simulated sports traffic. Natural and Artificial Playing Fields: Characteristics and Playing Features. ASTM STP 1073:85-89.

Duncan, R.R., and R.N. Carrow. 1999. Seashore paspalum: the environmental turfgrass Wiley.

Espinoza, F. and C.L. Quarín. 1997. Cytoembryology of Paspalum chaseanum and sexual diploid biotypes of two apomictic Paspalum species. Aust. J. Bot. 45:871-877.

Gray, L.C. 1933. History of agriculture in the southern United States to 1860 AM Kelley.

Green, T., J. Dunne, and J. Rogers III. 2012. A clarification of seashore paspalum vernation description. Applied Turfgrass Science.

Heckart, D.L., W.A. Parrott, and P.L. Raymer. 2010. Obtaining sethoxydim resistance in seashore paspalum. Crop Sci. 50:2632-2640.

Huang, B., R. Duncan, and R. Carrow. 1997. Drought-resistance mechanisms of seven warm-season turfgrasses under surface soil drying: II. Root aspects. Crop Sci. 37:1863-1869.

Jiang, Y., R.R. Duncan, and R.N. Carrow. 2004. Assessment of low light tolerance of seashore paspalum and bermudagrass. Crop Sci. 44:587-594.

Judd, B.I. 1979. Handbook of tropical forage grasses.

Karcher, D.E. and M.D. Richardson. 2003. Quantifying turfgrass color using digital image analysis. Crop Sci. 43:943-951. DOI: 10.2135/cropsci2003.9430.

Lee, G., R.N. Carrow, and R.R. Duncan. 2004. Salinity tolerance of selected seashore paspalums and bermudagrasses: Root and verdure responses and criteria. HortScience 39:1143-1147.

Lee, G., R.N. Carrow, and R.R. Duncan. 2005. Criteria for assessing salinity tolerance of the halophytic turfgrass seashore paspalum. Crop Sci. 45:251-258.

Lee, G., R.R. Duncan, and R.N. Carrow. 2004. Salinity tolerance of seashore paspalum ecotypes: Shoot growth responses and criteria. HortScience 39:1138-1142.

Morris, K.N. and R. Shearman. 2008. NTEP turfgrass evaluation guidelines. Online. Nat. Turfgrass Eval. Prog., Beltsville, MD.

Morton, J.F. 1974. Salt-tolerant silt grass (Paspalum vaginatum Sw.). Proc Fla State Hortic Soc 86, P 482-490, 1974.

Raymer, P., S. Braman, L. Burpee, R. Carrow, Z. Chen, and T. Murphy. 2008. Seashore paspalum: breeding a turfgrass for the future. Green Section Record. January/February:22-26.

Raymer, P., R. Carrow, and D. Wyatt. 2005. Screening for salt tolerance in seashore paspalum, Proc. Inter. Salinity Forum. pp. 25-27.

Raymer, P.L. 2006. Salt Tolerance in Seashore Paspalum. Turfgrass trends.

Richardson, M., D. Karcher, and L. Purcell. 2001. Quantifying turfgrass cover using digital image analysis. Crop Sci. 41:1884-1888.

Shin, J., P. Raymer, and W. Kim. 2006. Environmental factors influencing germination in seeded seashore paspalum. HortScience 41:1330-1331.

Skerman, P.J. and F. Riveros. 1990. Tropical grasses. Food & Agriculture Org.

Wang, M., Z. Chen, N. Barkley, M. Newman, W. Kim, P. Raymer, and G. Pederson. 2006. Characterization of Seashore Paspalum (Paspalum vaginatum Swartz) Germplasm by Transferred SSRs from Wheat, Maize and Sorghum. Genetic Resources and Crop Evolution 53:779-791. DOI: 10.1007/s10722-004-5540-3.

Webster, R.D. 1987. Australian Paniceae (Poaceae) J. Cramer.