Geographic Distribution of Races The seven geographic regions of North America (fig. 4) are determined based on geographic barriers, prevailing winds and other weather, crop management, and occurrence and virulence of races of Puccinia striiformis. Region 1 has the greatest number of races (fig. 4) and the races with the greatest range of virulence, i.e., virulence on the greatest number of differential cultivars in North America (table 4). Of the 39 races identified in North America, 31 occur in region 1; of those 31, 26 were first detected in the region. Of the five races that were first detected in other regions, CDL-8, CDL-16, and CDL-18 were detected in region 5, CDL-25 was detected in region 4, and CDL-6 was detected in region 6. Based on the characteristics of the regions, it is deduced that CDL-6, CDL-8, CDL-16, and CDL-18 probably evolved independently in region 1 and were not introduced from other regions. CDL-25 could have been introduced from region 4, but all available evidence indicates that it evolved independently in region 1. Because region 1 is bordered on the west and east by mountains and the prevailing winds are from the southwest, the probability of the introduction of spores from other regions is low. Region 1 is an important wheat-growing region with an environment that is favorable for rust survival in winter and rust development in spring. During the last 30 years, 3 out of every 4 years have been favorable for severe rust epidemics in fields of susceptible cultivars in region 1. In addition, cropping practices favor the survival and development of rust in region 1. All but one of the major classes of wheat are grown in region 1. Soft white winter wheat and soft white spring wheat are grown primarily in the eastern third of the region (which has the region's highest rainfall) and in irrigated fields throughout the region. Hard red winter wheat is grown primarily in the lower-rainfall areas in the central part of the region. Hard red spring wheat is grown in both the low-rainfall areas and under irrigation in the central and western parts of region 1. Club wheats are grown primarily in intermediate-rainfall areas bordered on the west by the hard red winter wheat and on the east by the soft white winter wheat. Spring durum wheat is grown by a few farmers, usually under contract. Under drier conditions, winter wheat is planted in August and September; under wetter conditions, it is planted in October and November. Spring wheat is planted in March or April. Thus management of these wheats provides a continuous green host for the survival and development of rust in region 1. The distribution and prevalence of the races of P. striiformis in region 1 are primarily related to susceptibility of the cultivars. The races that are virulent on Moro, Paha, Lee, and Tyee have been consistently most prevalent in the low-to-intermediate rainfall areas where susceptible club wheats and hard red winter wheats are grown. Races virulent on Fielder, Druchamp, Produra, and Stephens have been consistently most prevalent in the southern half of region 1, where Stephens is most commonly grown. Races virulent on Fielder but not Druchamp, Produra, and Stephens are prevalent in the northern half of the region, especially where Daws is grown. Races virulent on Lemhi and Heines VII remain common throughout the region. Races virulent on Yamhill are occasionally detected, and races virulent on Chinese 166 are very rare. Table 4. Virulence and avirulence of races of Puccinia striiformis from seven regions of North America on differential wheat cultivars. _____________________________________________________________________________ North Differential wheat cultivar number America _____________________________________________________________________ region1 1 2 3 4 5 6 7 8 9 10 11 12 13 _____________________________________________________________________________ 1 V3 V V V V V A3 V V V V V V 2 V V V V A V A V V V A V A 3 V A V A A V A V A V A V A 4 V V V V A V A V V V V V A 5 V V V V A V A V V V V V A 6 V V V A A V A V A V A V A 7 V A V A A A A A V A A A A _____________________________________________________________________________ 1 See fig. 4 for map of regions. 2 1 = Lemhi, 2 = Chinese 166, 3 = Heines VII, 4 = Paha, 6 = Druchamp, 7 = Riebesel 47-51, 8 = Produra, 9 = Yamhill, 10 = Stephens, 11 = Lee, 12 = Fielder, 13 = Tyee. 3 V = virulent, A = avirulent. Regions 2 and 3 are somewhat isolated from other regions to the west, and generally their winters are usually unfavorable for the survival of stripe rust. Consequently, severe epidemics are less frequent in those regions. All races of P. striiformis identified from 1968 to 1987 in regions 2 and 3 were first detected in region 1. Races virulent on Lemhi, Heines VII, Druchamp, Produra, Stephens, and Fielder (table 4) are prevalent in both regions. Virulences on Chinese 166, Moro, and Yamhill are present in region 2 but not region 3. CDL-5 was prevalent in region 2 in the late 1960's and early 1970's, but its prevalence decreased when Crest wheat was no longer grown. CDL-5 or other races that are virulent on Moro have never been detected in region 3 even though many of the hard red winter wheat cultivars in that region are susceptible to them. All data on the races indicate that the races in regions 2 and 3 originated in region 1. When rust is severe in region 1, rust epidemics often occur later in the season in region 2, and the races that are detected are the same as those that occurred earlier in the eastern and northern sections of region 1. Because of the late development of epidemics, spring wheat cultivars, such as Fielder, Fieldwin, and Sterling (all susceptible to the same races), are the most vulnerable to stripe rust, especially in Alberta, Canada. Races that are virulent on Fielder are common in that area. The geographic barriers, direction of prevailing winds, favorable temperature and precipitation, and virulence of the races of P. striiformis all provide evidence that stripe rust races in region 4 probably originate locally and that the introduction of inoculum from other regions, except by humans, seldom (if ever) occurs. Races virulent on Yamhill and Stephens, the major cultivars grown in region 4, are most prevalent in the region. All races in the region are virulent on Lemhi, and virulence on Heines VII, Druchamp, Produra, and Fielder is common. In the early 1970's, experimental plots were inoculated with races virulent on Chinese 166 and Moro, but those races did not spread very far from the inoculation site, probably because there was and is no selective pressure for the virulences. Because of mountain barriers and prevailing winds, stripe rust is indigenous to region 5 and rust inoculum does not enter from other regions. Further, because of mild winters and cool springs and summers, stripe rust in region 5 has been severe annually for the last 16 years. Eighteen of the thirty-nine races have been detected in region 5, and nine of the races were first detected there. Seven races (CDL-8, CDL-10, CDL16, CDL-17, CDL-18, CDL-23, CDL-37) are virulent on Yamhill, and three races (CDL15, CDL-23, CDL-37) are virulent on Stephens. Yamhill and Stephens are the two major cultivars grown in region 5. One race (CDL-11) is virulent on only Lemhi. The most prevalent races have been and currently are those virulent on Heines VII, Yamhill, Druchamp, and Stephens. Races virulent on Yamhill are also virulent on Heines VII, and races virulent on Stephens are also virulent on Druchamp. Virulence on Druchamp was probably present in the region for years. Druchamp has been grown in the region and was still grown in the 1970's on Lopez Island in region 5. Races virulent on Yamhill were prevalent in the 1970's, when Yamhill made up more than 95% of the wheat grown in region 5. Most farmers in the region changed to Stephens in 1978. Since then, more than 95% of the wheat has been Stephens, and races virulent on Stephens have been most prevalent. Virulence on Heines VII and Druchamp may be explained by their common association with virulence on Yamhill or Stephens. However, the occurrence of some of the other races cannot be explained by the susceptibility of local cultivars to those races. The presence of races that are virulent on Chinese 166, Moro, Lee, Produra, or Fielder is not related to cultivars grown in the region. Region 5 is a center of major rust diversity, and stripe rust has existed there for more than 100 years (Humphrey et al. 1924). Stripe rust also occurs on several species of wild grasses and on cultivated rye, and those hosts may contribute to the diversity of the races. However, predominant rust collections from those species have been identical to collections from adjacent wheat fields in the region. Region 6, which is also separated from the other regions, has a favorable environment for rust during the growing season (October to June) but a less favorable environment during summer, when wheat and triticale are not grown as crops. However, those hosts are present as volunteer plants in other crops, along the roadsides, and on irrigation dikes (Line 1976). Eleven races have been detected in region 6, of which four were first detected in the region. Virulences on Lemhi, Heines VII, Druchamp, Produra, Stephens, and Fielder are common (table 4), and virulence on Chinese 166 is more common in region 6 than in other regions. CDL-21 and CDL-35 have been collected from only region 6; CDL-11, which occurs only in regions 5 and 6, is most common in region 6. Each of these three races has a narrow range of virulence (table 3). CDL-21 and CDL-35 appear to have a closer association with triticale than to wheat. The rust occurs on various grasses, especially Hordeum sp., and these additional hosts may contribute to the uniqueness of the races in region 6. Region 7 includes all wheat-producing areas east of the Rocky Mountains. The winter weather in the south is favorable for the survival and increase of stripe rust, but the hot spring and summer weather in the south and hot summers in the north often limit the development of severe epidemics. However, stripe rust has become more prevalent in region 7 since 1984, especially in the South-Central States. Virulence of the races is consistent throughout the region, probably because of prevailing winds and the absence of geographic barriers. Isolates collected from that region and northern Mexico during the period before 1985 were all identified as CDL-3. Since then, a closely related race, CDL-8, has been found in region 7. There may be other virulences that cannot be detected by use of the current differentials. However, of the many cultivars from region 7 that have been tested, none have detected virulences that have not been detected by the current differentials.