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.