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Oxytropis Lambertii</Emp P1: GFU/GVM Journal of Chemical Ecology [joec] pp438-joec-369022 April 5, 2002 8:41 Style file version Nov. 19th, 1999 Journal of Chemical Ecology, Vol. 28, No. 4, April 2002 (C 2002) DISTRIBUTION OF LOCOWEED TOXIN SWAINSONINE IN POPULATIONS OF Oxytropis lambertii M. H. RALPHS,1, S. L. WELSH,2 and D. R. GARDNER1 1USDA/ARS Poisonous Plant Laboratory 1150 E. 1400 N., Logan, Utah 84341 2Bean Natural History Museum Brigham Young University Provo, Utah 84602 (Received October 10, 2001; accepted December 17, 2001) Abstract—Oxytropis lambertii has been considered to be one of the major lo- coweeds responsible for livestock poisoning on rangelands, but there has been much confusion as to its taxonomic identity. The objective of this study was to conduct a field survey of several populations of each of the three varieties [var. lambertii Pursh; var bigelovii A. Gray; var. articulata (E. Greene) Barneby] to document the presence or absence of the locoweed toxin, swainsonine. Swain- sonine was found at detectable levels (>0.001% dry weight) in only five popula- tions of var. bigelovii in the southwest portion of its distribution in southern Utah, Arizona, and southwestern New Mexico, USA. No swainsonine was detected in populations in the northeast areas of its distribution (eastern Utah, Colorado, northeastern New Mexico, USA). The other varieties, articulata and lambertii, also did not contain swainsonine. It is suspected that a plant fungal endophyte may be responsible for the high variability in swainsonine content in populations of O. lambertii. Key Words—Lambert locoweed, crazyweed, Oxytropis lambertii, swainsonine, livestock poisoning, toxic plant, poisonous plant. INTRODUCTION Locoweeds (Astragalus and Oxytropis spp.) cause widespread poisoning of live- stock in the western United States (Kingsbury, 1964). Much of the early toxico- logical research was conducted on the complex of species then called Aragallus lambertii (Marsh, 1909), which included both the white-flowered and the lavender- flowered groups of plants. Later Marsh (1919) referred to the white locoweed as To whom correspondence should be addressed. E-mail: [email protected] 701 0098-0331/02/0400-0701/0 C 2002 Plenum Publishing Corporation P1: GFU/GVM Journal of Chemical Ecology [joec] pp438-joec-369022 April 5, 2002 8:41 Style file version Nov. 19th, 1999 702 RALPHS,WELSH, AND GARDNER Astragalus lambertii. Butcher et al. (1953) were first to point out that the plant with which Marsh conducted most of his early toxicology research was indeed the white flowered O. sericea. Taxonomists have separated the two species based on the color of their flowers (O. sericea is white to slightly yellow; O. lambertii is purple violet or lavender) and the dolabriform (ox head-like) pubescence that is unique to O. lambertii (Barneby, 1952; Welsh, 2001). The confusion in taxonomy complicates the question as to whether O. lambertii is really a locoweed and whether it has been responsible for poi- soning livestock. Literature reviews of the locoweeds all include O. lambertii as one of the major locoweeds (Marsh, 1919; Kingsbury, 1964; James et al., 1981; Ralphs and James, 1999). However, there is no empirical evidence of poisoning attributable to O. lambertii. The objective of this research was to document the presence of the locoweed toxin swainsonine in O. lambertii. METHODS AND MATERIALS A literature review was conducted of the toxicology and chemistry literature for any reference to swainsonine in O. lambertii. A widespread field survey of O. lambertii populations was conducted and laboratory analysis performed to determine the presence or absence of swainsonine. Voucher specimens are held in the S. L. Welsh Herbarium in the Bean Natural History Museum, Brigham Young University, Provo, Utah. Known populations of each of three varieties representing the entire geo- graphical distribution of that variety were sampled in June 2000 (Table 1). Ten individual plants were collected at each site. Half of each plant was pressed for identification. The other half was air dried in a forced air oven for 48 hr at 60C and then ground through a cyclone grinder. Samples were analyzed for the locoweed toxin swainsonine by the procedures described by Gardner et al. (2001). Detec- tion limit of swainsonine was 0.001% of dry weight. Swainsonine concentration was compared by one-way analysis of variance to determine differences between locations. RESULTS AND DISCUSSION There are few reports in the literature of O. lambertii containing swainso- nine (Table 2). Molyneux et al. (1989, 1991) reported the presence of swainsonine in O. lambertii by using a thin-layer chromotography technique. Later samples from the Henry Mountains in southeast Utah, and from Livermore, Colorado, showed swainsonine levels of 0.031 and 0.007%, respectively (Molyneux, unpub- lished data). Fox et al. (1998) reported O. lambertii from western and northern New Mexico contained swainsonine. Braun (1999) also reported swainsonine in P1: GFU/GVM Journal of Chemical Ecology [joec] pp438-joec-369022 April 5, 2002 8:41 Style file version Nov. 19th, 1999 SWAINSONINE IN LOCOWEED 703 TABLE 1. VARIETIES OF Oxytropis lambertii VAR. bigelovii,STAGE OF GROWTH AND SAMPLING SITES Variety/ voucher Stage Location GPS Soil/Site Plant community bigelovii 27661a Seed Kanab, UT 370601900N Sandy hills Desert shrub 1115102800W 27665a Veg Flagstaff, AZ 370604100N Sandy loam Ponderosa pine 1115102800W foothills openings 27667a Veg Springerville, AZ 340004900N Gravely hills Pinyon/juniper 1111004800W woodland 27668a Veg Kingston, NM 325205100N Rocky, mountain Ponderosa pine/alligator 1075105500W slopes juniper forest 27669a Bud Winston, NM 332104300Nl´oamy valley Blue grama 1073404100W grassland 27672 Flo Ocate, NM 361501100N Gravely hills Ponderosa pine 1050203200W 440981 Flo Capulin, NM 304102500N Rocky loam Shortgrass prairie 1040803500W sidehill 440982 Flo Sophia, NM 362800600NL´oamy flats Shortgrass prairie 1035905400W 440980 Flo Ft. Collins, CO 405603900N Gravely foothills Mixed grass prairie 1051503300W 440983 Flo Ferron, UT 390605700N Sandy mountain Low sage, bluebunch 1111703600W bench wheatgrass articulata 27689 Pod Knowles, OK 365501500N Rolling hills Tall grass prairie 1001802300W 27697 Pod Buffalo, OK 364804400N Rolling hills Tall grass prairie 994602200W 27698 Pod Meade, KA 371000900N Calcareous Tall grass prairie 1002300300W breaks lambertii 27704 Flo Sidney, NB 410901800N Limestone bluff Mixed grass prairie 1030501800W 27717 Flo Hot Spring, SD 432403500N Gravely hills Ponderosa pine, 1032002300W mixed grass prairie 27721 Flo Lusk, WY 43 0501200N Sandy/gravel Mixed grass prairies 1041903600W hills a Populations containing swainsonine. O. lambertii in Grant County, New Mexico, that was associated with a fungal endophyte. A recent grazing study comparing the relative palatability of O. lambertii with O. sericea growing together on short-grass prairie in north-central Colorado, showed that swainsonine was not present in detectable amounts in samples of P1: GFU/GVM Journal of Chemical Ecology [joec] pp438-joec-369022 April 5, 2002 8:41 Style file version Nov. 19th, 1999 704 RALPHS,WELSH, AND GARDNER TABLE 2. REPORTED ACCOUNTS OF O. lambertii CONTAINING SWAINSONINE Date Location Reference Swainsonine (%) 1986 Henry Mt., Wayne Co., UT Molyneux (unpubl.) 0.031 n.d. Livermore, Larimer Co., CO Molyneux (unpubl.) 0.007 1989 no location given Molyneux et al. (1991) >0.01a 1987 Winston, Sierra Co., NM Fox et al. (1998) 0.099 1991 Winston, Sierra Co., NM Fox et al. (1998) 0.021 1991 Ocate, Moro Co., NM Fox et al. (1998) 0.026 1999 Grant Co., NM Braun (1999) 0.576 1999 Colfax Co., NM Braun (1999) 0.0006 a Presence detected by thin-layer chromotography. O. lambertii taken from different pastures in 1998 and 1999 (Ralphs et al., 2001). Samples of O. lambertii collected from another research location in northeastern New Mexico also did not contain swainsonine (Ralphs, unpublished data). There are three varieties of O. lambertii. Variety bigelovii A. Gray is separated from the other two by having stipitate pods (slender stalklike base). It occurs from southeastern Wyoming, southward through the foothills and mountains of Colorado and New Mexico, and westward into eastern Utah and Arizona. Variety lambertii Pursh is separated from var. articulata (E. Greene) Barneby by its longer calyx teeth, about one third as long as the tube and shorter flowers, <18 mm. It occurs on prairies from North Dakota to Kansas with a few populations in eastern Wyoming. Variety articulata has short calyx teeth, about one fifth as long as the tube, and flowers 18–25 mm long. It occurs in southwestern Kansas, western Oklahoma, the Texas panhandle, and northeastern New Mexico (Welsh, 2001). Variety bigelovii is the most widespread of the three varieties. Ten popula- tions were sampled in Colorado, Utah, Arizona, and New Mexico (Table 1). Three populations of var. articulata were sampled in Oklahoma and Kansas. Three pop- ulations of var. lambertii were sampled in Wyoming, South Dakota, and Nebraska. The locoweed toxin is a trihydroxy indolizidine alkaloid, swainsonine (Figure 1) (Molyneux and James, 1982). Detectable levels of swainsonine were found in only five of the 10 populations of bigelovii located in the southwestern geographical areas of its range (Table 3). The other five populations in its eastern range did not contain detectable levels. The other two varieties also did not contain detectable levels of swainsonine. FIG. 1. The toxic alkaloid swainsonine found in locoweeds. P1: GFU/GVM Journal of Chemical Ecology [joec] pp438-joec-369022 April 5, 2002 8:41 Style file version Nov. 19th, 1999 SWAINSONINE IN LOCOWEED 705 TABLE 3. LOCATIONS IN 2000 SURVEY WHERE O. lambertii CONTAINED SWAINSONINE (MEAN SE) Location Mean (%) Range (%) Flagstaff, AZ 0.054 0.027 0.022–0.106 Winston, NM 0.038 0.035 0–0.068 Springerville, AZ 0.026 0.021 0–0.065 Kingston, NM 0.016 0.013 0–0.043 Kanab, UT 0.008 0.016 0–0.047 Four of the five southwestern populations of bigelovii containing swainsonine were in the vegetative or bud stage of growth when they were harvested in June (Table 1), indicating they flowered later in the summer.
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