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Texas Wild Rice (Zizania Texana) As a Model

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Texas Wild Rice (Zizania Texana) As a Model Genet Resour Crop Evol (2007) 54:837–848 DOI 10.1007/s10722-006-9167-4 ORIGINAL PAPER Capturing genetic diversity of wild populations for ex situ conservation: Texas wild rice (Zizania texana) as a model Christopher M. Richards Æ Michael F. Antolin Æ Ann Reilley Æ Jackie Poole Æ Christina Walters Received: 19 November 2005 / Accepted: 13 April 2006 / Published online: 7 December 2006 Ó Springer Science+Business Media B.V. 2006 Abstract Genebanks complement other conser- only 45 individuals, while random sampling along vation programs because they preserve genetic the river captured much less diversity. The early diversity needed for future breeding and resto- conservation collection captured as much diver- ration. We evaluated efficiency of capturing sity as expected from random sampling. Texas genetic diversity, using endangered Zizania wild rice stands resemble a mainland-island texana (Texas wild rice) as a model for plants with metapopulation; our analyses suggest that strati- recalcitrant seeds. This perennial aquatic grass is fied sampling maximizes genetic diversity for this restricted to 4 km of the San Marcos River in population dynamic. Demographic and genetic Texas. An early conservation collection included information is important for validating the design plants from stands throughout the river, based on of efficient ex situ collections and guiding in situ the assumption stands would be unique geno- conservation. types. Using microsatellite markers, we found that genetic diversity was concentrated in five of Key words Stratified sampling Æ Allelic richness Æ 15 large, demographically stable stands; 96 stands Conservation genetics Æ Genetic structure Æ smaller than 2 m2 contributed no unique alleles. Zizania texana High heterozygosity and few duplicate genotypes suggested that sexual reproduction occurs more often than presumed. Simulations of stratified Introduction sampling of large stands captured all alleles in Assembling ex situ collections from wild popula- tions is an important component of wider con- C. M. Richards (&) Æ A. Reilley Æ C. Walters USDA/ARS National Center for Genetic Resources servation goals (Schoen and Brown 2001). Preservation, 1111 South Mason Street, Fort Collins, Conservation collections in botanical gardens, CO 80521, USA zoos, or genebanks are designed to safeguard e-mail: [email protected] genetic diversity, enhance current population size, M. F. Antolin keep useful traits accessible, and re-establish Department of Biology, Colorado State University, populations in restoration projects. As such, col- Fort Collins, CO 80523-1878, USA lection strategies must efficiently capture a high percentage of the extant genetic diversity, as J. Poole Texas Parks & Wildlife, 3000 I-35 South, Austin, TX measured directly by variation in molecular or 78704, USA quantitative traits (Brown and Hardner 2000) or 123 838 Genet Resour Crop Evol (2007) 54:837–848 indirectly through a geographically stratified A conservation collection of 48 Texas wild rice sampling scheme (Shands 1991). Without direct plants from different locations throughout its measurements of genetic variation of phenotypic range was established from 1986 to 1999 at a fish traits or molecular markers, indirect methods hatchery in San Marcos, Texas in raceways with a often provide excellent first approximations of continuous flow-through of water from the river. genetic variation when they take into account the However, this collection of plants was both costly target species’ life history characteristics, repro- to maintain and fraught with inherent risks: by ductive system, and long-term population changes 1999, mortality reduced the conservation collec- (Brown and Briggs 1991; Schoen and Brown tion to 35 plants (for which we have DNA sam- 1991). For instance, if populations are abundant ples and genotypes), and the collection continues but are comprised mainly of a few genotypes that to decline. arise by asexual reproduction, total genetic It is possible to establish genebanks using seeds diversity could be found in relatively small of Z. texana. We have developed cryogenic samples collected from each population. methods to preserve Texas wild rice seeds and Efficiency in collecting is important not just demonstrated that these methods do not reduce because risk of eminent extinction, but also genetic diversity in stored samples for a set of because maintaining large, redundant collections variable microsatellite loci (Walters et al. 2002; may divert resources from other conservation Richards et al. 2004). Further, growth conditions priorities such as habitat protection. Seedbanks that induce flowering and seed production in can provide an important conservation tool controlled environments are known (Power and because the viability of hundreds or thousands of Fonteyn 1995; C. Walters unpublished data). Al- individuals can be maintained for decades with- though cryogenic storage protocols for Texas wild out regeneration (Walters 2004). However, rice are time consuming and costly, developing an recalcitrant seeds, which cannot be stored under ex situ collection seems particularly useful as a the standard cold and dry conditions used for complementary tool for other conservation pro- most seeds, require specialized protocols to jects for Texas wild rice because human induced ensure their viability in genebanks. Recalcitrant pressures on this species’ habitat, especially by seeds are common in many of the riparian and increased diversions of water, has accelerated in tropical species (Hong et al. 1998) that are the last few decades (Vaughan 1986). currently under threat because of harvesting or In this study we examined genetic diversity and habitat loss. Developing cryogenic methods for population structure of stands of Texas wild rice storage of recalcitrant seeds requires significant along its entire range in the San Marcos River, investment of time and resources. It is thus critical using six highly variable microsatellite markers. to design efficient collections that optimize the Sampling over three years was designed to sample capture of genetic diversity from wild. among and within stands, test whether reproduction Population genetic analysis of the aquatic within the river is primarily asexual, and evaluate Texas wild rice (Zizania texana Hitchcock) pro- temporal changes in genetic variation and struc- vides a case study for targeted sampling for a ture. We follow this analysis with a sampling species with a narrow geographic range: four km simulation to identify collecting strategies that of the San Marcos River in south central Texas. most efficiently capture the genetic diversity of Texas wild rice seeds are recalcitrant and do not Texas wild rice for ex-situ genebanks. survive in conventional storage. Monitoring of stands since the 1960s has revealed little or no flowering in the river and implied that the pri- Natural history of Z. texana mary mode of reproduction of this perennial grass is asexual (Emery 1977; Power 1996). These Texas wild rice is an aquatic perennial grass observations suggested one approach for ex situ endemic to a four km stretch of the San Marcos conservation was to dig plants from the river and River in Hays CO, TX (Emery 1977) (Fig. 1). The grow them in pots in an artificial aquatic habitat. river’s clear, spring-fed headwaters emerge from 123 Genet Resour Crop Evol (2007) 54:837–848 839 Fig. 1 Map of stands of Z. texana in the San Marcos River in Texas, showing river sections in annual surveys (left) the Edwards Aquifer through the permeable karst comparisons. Estimates of the percent coverage in the Balcones Fault of Texas at an average flow within polygons (m2) were used as relative mea- rate of 157 cu ft/s, and the river flows south to the sures of stand size. Two prominent features confluence of the Blanco River (Terrell et al. emerge from this long term study. First, while 1978). Below this confluence, where the water be- sizes of stands varied from 0.01 to 335 m2 (see comes heavily silted, no Z. texana have been re- Table 3 below), most stands were small (mean corded. The habitat requirements of Z. texana size is 8.5 m2). Second, the distribution and local appear to be confined to the upper reaches of the abundance of this species was highly dynamic river, which varies from 10 to 25 m wide and is up (Poole unpublished data 2003). Over the years, to four m deep in some channels. Texas wild rice the number of stands fluctuated markedly along grows mostly in large vegetative stands that are the river (mean = 206, SD = 45). Small stands submerged in mid-river in relatively swift and experienced higher rates of turnover (extinction shallow water (less than one m) and rooted in and colonization) than larger stands, with several sandy or gravely bottoms (Poole and Bowles 1999). of the largest stands being stable throughout the The spatial distribution of stands has been survey period (Poole unpublished data 2004). In mapped annually for 15 years by the Texas Parks addition, the census revealed that the spatial and Wildlife Department to monitor the abun- arrangement of stands was more clumped in the dance of this species and to describe its habitat upper reaches of the river, in segments A and B, requirements (Poole unpublished data). Occupied and progressively became more dispersed down- reaches of the river were designated as segments stream of segment F. (Emery, Southwest Texas State University Texas wild rice rarely flowers in the river unpublished data 1978) that correspond to phys- (Emery 1977; Power 1996) but it is capable of ical features of the river such as bridges, dams and flowering when grown in fish-rearing raceways other structures (Fig. 1). Stands, which are de- located at the National Fish Hatchery and Tech- fined as continuous masses of Z. texana plants, nology Center in San Marcos, TX, in an old fish were delineated by polygons that enclosed the hatchery on the campus of Texas State University- irregularly-shaped stands, mapped using survey- San Marcos (Power 1996), and in greenhouses at ing equipment, and recorded in GIS for yearly Colorado State University (Walters unpublished 123 840 Genet Resour Crop Evol (2007) 54:837–848 data 2004).
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