The karyology of the Common shrew, Sorex araneus Linné, 1758 (Insectivora, Soricidae) in southwestern Germany

Presented is the karyotype of Sorex araneus from 4 trapping sites in southwestern Germany. From this species nearly 20 chromosomal races have so far been described. In the study area autosomal numbers of 2n_a= 22–24 were recorded. The characteristical metacentrics jl, hi, gm, kr and the acrocentrics n, o, p, q were identified by G-banding. The element kr shows Robertsonian polymorphism. The metacentrics only allow a classification of the studied populations as chromosomal race “Vaud” from Switzerland. The postglacial recolonization and the possible presence of other chromosomal races in the central and northern parts of Germany are briefly discussed. Different selection pressures act on metacentrics and acrocentrics in central populations of a chromosomal race in contrast to those living in contact zones of different chromosomal races. It is suggested that the smaller autosomal arms are subject to weaker selection pressure to be fixed as metacentrics.     

Craniometric study of the common shrew (Sorex araneus L. 1758) from different localities and chromosomal races across Germany and Europe

A total of 440 skulls of common shrews, Sorex araneus, from Germany and Europe (Croatia, Hungary, Austria, and Norway) were studied. The material represented six chromosomal races (Ulm, Laska, Drnholec, Mooswald, Jütland, and Abisko) assignable to the Western European and Northern European karyotypic groups. The race of a few samples was not determined. Twenty-one linear measurements were taken on skulls and mandibles and used in this study. Pearson correlations and multiple linear regressions were used to see the relationship of the cranial variables to altitude, latitude, the chromosomal race, and the geographic location. The results from the tested samples differed; the most negative correlations to latitude were found in the samples assigned to the Western European karyotypic group (WEK), the least negative ones in the samples of the Ulm race. These results indicate the converse of Bergmann's rule. But taking into consideration all the samples of the different karyotypic groups across Europe, the correlations to latitude included positive ones, which would indicate that Sorex follows Bergmann's rule in some of the variables. The studied material of different karyotypic groups could not be differentiated in discriminant analyses. The separation of the studied races within the WEK alone was slightly better, but about 30 % of ungrouped cases remained. Only the separation of the regional samples within one chromosomal race revealed better results but was still very different between the races. This indicates that within the races, regional differences might be strong enough for a separation of the samples and that within a karyotypic group, and even more so across karyotypic groups, regional differences conceal racial differences.     

New Races of Sunflower Downy Mildew (Plasmopara Halstedii) in Germany

Plasmopara halstedii, the causal agent of downy mildew of cultivated sunflower (Helianthus annuus), was documented in Germany for the first time in commercial fields. The pathogen was first observed in the Württemberg area, where races 1 and 4 were identified using a set of differential lines. Later, commercial fields near Baden were found to be infected by race 5, which is the first occurrence of that race outside of North America. With the discovery of race 5, there are now eight races of the sunflower downy mildew fungus that have been found in Europe. The sunflower cultivars most frequently grown in Germany were investigated for resistance to race 1, 4 and 5; while all were resistant to race 1, none were resistant to either race 4 or 5.     

New Races of Sunflower Downy Mildew (Plasmopara Halstedii) in Germany

Plasmopara halstedii, the causal agent of downy mildew of cultivated sunflower (Helianthus annuus), was documented in Germany for the first time in commercial fields. The pathogen was first observed in the Württemberg area, where races 1 and 4 were identified using a set of differential lines. Later, commericial fields near Baden were found to be infected by race 5, which is the first occurrence of that race outside of North America. Withthe discovery of race 5, there are now eight races of the sunflower downy mildew fungus that have been found in Europe. The sunflower cultivars most frequently grown in Germany were investigated for resistance to race 1, 4 and 5; while all were resistant to race 1, none were resistant to either race, 4 or 5.     

The geographic distribution of flavone-glycosylating genes in Silene pratensis (Rafn.) Godron and Gren. (Caryophyllaceae)

In Silene pratensis three loci (g, gl and fg) control the glycosylation of isovitexin. Three alleles are known for both the g-locus (g, g ^G and g ^X ) and the gl-locus (gl, gl ^A and gl ^R ); for the fg-locus there are only two alleles (fg and Fg). The distribution of these alleles over 285 European populations of S. pratensis has been investigated. It was concluded that there are three different chemical races within S. pratensis in Europe. The first race contains the populations in western and southern Europe and displays high frequencies of g ^G , gl and fg. The frequencies of g ^G and gl ^R are very high in the second chemical race, which can be found in the USSR, Scandinavia and eastern Poland. The third chemical race occurs in central Europe and in this race the frequencies of both g and gl ^R are high, Fg has low to moderate frequencies in the second and third groups. The alleles gl ^A and g ^X are seldom found in S. pratensis, but are present in the closely related S. dioica. They do occur with low frequencies in some populations of S. pratensis, possibly as a result of hybridization with S. dioica.     

Comparative chiasma analysis using a computerised optical digitiser

A new computerised technique has been devised for measuring the distribution of chiasmata along diplotene bivalents. The method involves the introduction into the field of view of the microscope, of a fine light spot which can be accurately manipulated along the chromosomes of each bivalent. The data recorded include (a) the positions of the chiasmata along the bivalent in terms of their relative distances from the centromere and (b) the individual bivalent and cellular chiasma frequencies. — The method has been applied to the analysis of chiasma distribution patterns in the two known species of the genus Caledia, C. species nova 1 and C. captiva and in two chromosomal races of the latter. Statistical tests indicate that within bivalents at least 40% of the comparative distribution patterns of chiasmata between races and species are significantly different. Similar comparisons between populations within races reveal only 18% significant differences. — The observed distribution patterns of chiasmata in this genus suggest that chiasma formation is sequential from centromere to telomere. — The variation in the frequency and distribution of chiasmata between races and species suggests that the interference distances between successive chiasmata are, at least partially, independent of chiasma frequency and position. — The interracial and interspecific differences in chromosome structure are correlated with changes in chiasma pattern.     

The cytology of the genus Briza L. (Gramineae)

Meiosis has been studied in five European and four South American species of Briza. The various species have different chiasma frequencies and different patterns of chiasma localisation. Population studies show that there are significant differences in mean plant chiasma frequency between populations of B. maxima whereas B. media and B. spicata do not show interpopulation differences. B. media is shown to have diploid and autotetraploid races and the distribution of these chromosome races in Europe has been plotted. The South American species studied are all bivalent forming tetraploids and would therefore appear to have an allopolyploid origin. Interchange heterozygosity has been found in B. media and B. elatior, different populations of B. media have been shown to be heterozygous for different interchanges. These topics are discussed in relation to the regulation of recombination.     

Molecular mapping of genes for race-specific overall resistance to stripe rust in wheat cultivar Express

‘Express’, a hard red spring wheat cultivar that has been widely grown in the western United States, is used to differentiate races of Puccinia striiformis f. sp. tritici, the causal fungal pathogen of wheat stripe rust. To identify genes conferring race-specific, overall resistance to stripe rust, Express was crossed with ‘Avocet S’. The parents and F₁, F₂, F₃ and F₅ populations were tested with races PST-1, PST-21, PST-43, and PST-45 of P. striiformis f. sp. tritici in the seedling stage under controlled greenhouse conditions. Two dominant genes for resistance to stripe rust were identified, one conferring resistance to PST-1 and PST-21, and the other conferring resistance to all four races. Linkage groups were constructed for the resistance genes using 146 F₅ lines to establish resistance gene analog and chromosome-specific simple sequence repeat marker polymorphisms. The gene for resistance to races PST-1 and PST-21 was mapped on the long arm of chromosome 1B, and that conferring resistance to all four races was mapped on the long arm of chromosome 5B. We temporarily designate the gene on 1BL as YrExp1 and the gene on 5BL as YrExp2. Polymorphism of at least one of the two markers flanking YrExp2 was detected in 91% of the 44 tested wheat genotypes, suggesting that they would be useful in marker-assisted selection for combining the gene with other resistance genes into many other wheat cultivars. Knowledge of these genes will be useful to understand recent virulence changes in the pathogen populations.     

A comparative study of the chromosomes in the incipient species of the Drosophila paulistorum complex

Summary Drosophila paulistorum Dobzhansky et Pavan is a complex of six races or incipient species. The races are mostly allopatric, but they are reproductively isolated sufficiently to permit them to exist also sympatrically in some places. The gene arrangements in the chromosomes of the races have been compared by means of examination of the giant chromosomes in the larval salivary glands; 28 strains of all races, and about an equal number of interracial hybrids have been studied.Chromosomal inversion polymorphism has been discovered in all races, even in the Guianan race of which only a single strain is available. Inversion heterozygotes are found in every one of the five chromosomal strands which the species has. Interracial hybrids tend to be heterozygous for more inversions than are present in the strains of the parental races. The Transitional race has however much the same gene arrangements as the widespread Andean — South Brazilian race.With the exception of the Transitional race, and of three other possible exceptions, each race has a collection of its own race-specific inversion polymorphs, not found in the other races. This very striking finding is discussed in connection with the hypothesis which envisages the origin of new species from marginal colonies at the periphery of the geographic distribution area of the ancestral species.The work reported in this article has been carried under Contract No. AT-(30-1)-1151, U.S. Atomic Energy Commission, mostly at the Department of Zoology, Columbia University, New York.     

The complex hybrid zone between the Abisko and Sidensjö chromosome races of Sorex araneus in Sweden

Six chromosome races of the common shrew occur in Sweden, each with its characteristic arm combination of metacentric chromosomes. G-banded karyotypes were analysed from 201 common shrews in 14 localities of the northern hybrid zone in Sweden. Analyses from another 64 shrews from seven localities outside the hybrid zone w ere included for comparison. The shrews were classified with respect to karyotype into any of five categories: (1) Abisko race, (2) Sidensjö race, (3) hybrids between the parental races, (4) pseudohybrids (a type of hybrid), and (5) AT with all race-specific chromosomes (h, i, n, o, p, r) present as telocentrics. Hybrids occurred at a frequency close to Hardy-Weinberg expectation in the centre of the hybrid zone. Chromosome polymorphism of Robertsonian type was common and 43 different karyotypes were found among the specimens studied. The polymorphism involved six metacentric pairs in the Abisko and three in the Sidensjö race. The frequency of the Sidensjö race-specific metacentric hi decreased and the frequency of the Abisko race-specific hn increased from south-west to north-east along a transect across the hybrid zone. The number of race-specific telocentrics reached a peak 13 km north-east of the hi-hn cline centre. The estimated standardized cline width for chromosomes hi and hn was 16.0 km. The extension of the Sidensjö race is comparatively narrow [c. 50 km in the region of the investigation), and it is regarded to be a 'hybrid race' between the Uppsala race, which colonized Sweden from the south-west, and the Abisko race which arrived from the north-east after the most recent glaciation. The origin of the Sidensjö race is thus less than 10 000 years old, because earlier this area was covered by glacial ice.     

Standing geographic variation in eclosion time and the genomics of host race formation in Rhagoletis pomonella fruit flies

Taxa harboring high levels of standing variation may be more likely to adapt to rapid environmental shifts and experience ecological speciation. Here, we characterize geographic and host‐related differentiation for 10,241 single nucleotide polymorphisms in Rhagoletis pomonella fruit flies to infer whether standing genetic variation in adult eclosion time in the ancestral hawthorn (Crataegus spp.)‐infesting host race, as opposed to new mutations, contributed substantially to its recent shift to earlier fruiting apple (Malus domestica). Allele frequency differences associated with early vs. late eclosion time within each host race were significantly related to geographic genetic variation and host race differentiation across four sites, arrayed from north to south along a 430‐km transect, where the host races co‐occur in sympatry in the Midwest United States. Host fruiting phenology is clinal, with both apple and hawthorn trees fruiting earlier in the North and later in the South. Thus, we expected alleles associated with earlier eclosion to be at higher frequencies in northern populations. This pattern was observed in the hawthorn race across all four populations; however, allele frequency patterns in the apple race were more complex. Despite the generally earlier eclosion timing of apple flies and corresponding apple fruiting phenology, alleles on chromosomes 2 and 3 associated with earlier emergence were paradoxically at lower frequency in the apple than hawthorn host race across all four sympatric sites. However, loci on chromosome 1 did show higher frequencies of early eclosion‐associated alleles in the apple than hawthorn host race at the two southern sites, potentially accounting for their earlier eclosion phenotype. Thus, although extensive clinal genetic variation in the ancestral hawthorn race exists and contributed to the host shift to apple, further study is needed to resolve details of how this standing variation was selected to generate earlier eclosing apple fly populations in the North.     

Refugial races and postglacial colonization history of the freshwater amphipod Gammarus lacustris in Northern Europe

The systematic structure and postglacial population history of the freshwater amphipod Gammarus lacustris were explored in an allozyme survey of 65 populations across Northern Europe. A strong multilocus pattern of differentiation discriminated populations of the north‐east (north‐eastern Norway, northern Finland) from those in the west and the south (southern and central Scandinavia, Denmark, Poland). This principal division is attributed to postglacial colonization of the area by two main refugial races or lineages, one from the east (Russia), the other from the south (north‐western European continent). The strongly diverged Eastern and Western races (Nei's D= 0.3, from 22 loci) now meet in a secondary contact zone across a narrow sector of northernmost Norway. Genetic population compositions in this zone vary in a mosaic pattern, and show no evidence of reproductive incompatibility. Similar contacts of eastern and western lineages, far older than the latest glaciation, are now known from a number of taxa and they constitute a general pattern in Fennoscandian phylogeography. Within the Western Gammarus race, the populations through coastal north‐western Norway are further distinguished from those in southern Scandinavia and Denmark by a set of unique alleles at high frequencies (D = 0.12). This suggests an independent early colonization of the coastal region by another distinct stock, either along an early deglaciated coastal corridor from the south‐west, or directly from the ice‐free continental shelf off the Norwegian coast – a hypothesis that has also previously been presented for G. lacustris, and parallels controversial suggestions of local refugia for other taxa in Scandinavia. The coastal population type only later could come into contact with Gammarus invading over the mountains from the south; these two population types now smoothly intergrade. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 79, 523–542.     

Frequency of hybridization between Ostrinia nubilalis E‐and Z‐pheromone races in regions of sympatry within the United States

Female European corn borer, Ostrinia nubilalis, produce and males respond to sex pheromone blends with either E‐ or Z‐Δ11‐tetradecenyl acetate as the major component. E‐ and Z‐race populations are sympatric in the Eastern United States, Southeastern Canada, and the Mediterranean region of Europe. The E‐ and Z‐pheromone races of O. nubilalis are models for incipient species formation, but hybridization frequencies within natural populations remain obscure due to lack of a high‐throughput phenotyping method. Lassance et al. previously identified a pheromone gland‐expressed fatty‐acyl reductase gene (pgfar) that controls the ratio of Δ11‐tetradecenyl acetate stereoisomers. We identified three single nucleotide polymorphism (SNP) markers within pgfar that are differentially fixed between E‐ and Z‐race females, and that are ≥98.2% correlated with female pheromone ratios measured by gas chromatography. Genotypic data from locations in the United States demonstrated that pgfar‐z alleles were fixed within historically allopatric Z‐pheromone race populations in the Midwest, and that hybrid frequency ranged from 0.00 to 0.42 within 11 sympatric sites where the two races co‐occur in the Eastern United States (mean hybridization frequency or heterozygosity (H_O) = 0.226 ± 0.279). Estimates of hybridization between the E‐ and Z‐races are important for understanding the dynamics involved in maintaining race integrity, and are consistent with previous estimates of low levels of genetic divergence between E‐ and Z‐races and the presence of weak prezygotic mating barriers.     

A second case of genetic host races in Rhagoletis? A population genetic comparison of sympatric host populations in the European cherry fruit fly,Rhagoletis cerasi

Despite an increasing acceptance in the biological community for sympatric speciation as a mode of species formation, well documented examples of sympatrically evolved ‘incipient species’ remain rare. The sympatric host races of apple maggot, Rhagoletis pomonella (Walsh), represent one of the most prominent case studies for sympatric speciation via a host shift. The European cherry fruit fly, R. cerasi (L.), shows strong ecological similarities to R. pomonella: (1) infestation of two different host plants, Lonicera xylosteum L. and Prunus avium L., and (2) divergent phenological and behavioral adaptations of flies on different hosts. The population genetic study presented here addresses whether the host associated populations of R. cerasi also represent genetically differentiated true host races. Out of a total of 29 allozyme loci examined, six were polymorphic and used to analyze six sympatric pairs of R. cerasi populations on Lonicera and Prunus from Switzerland and Germany. A direct comparison of allele frequencies between sympatric sites showed no pattern indicative of host races in R. cerasi. However, the hierarchical F‐statistic for one locus, mannose 6‐phosphate isomerase (Mpi), showed significant population differentiation that was in accordance with host race differentiation. Mpi is one of several loci that are also diagnostic for host race differentiation in R. pomonella. Results from Mpi suggest the formation of sympatric host races in R. cerasi, but additional polymorphic markers are necessary.     

Breeding systems and hybridization inPetrorhagia sect.Kohlrauschia (Caryophyllaceae)

In the diploid speciesP. prolifera, the evolution of autogamy has resulted in the presence of distinct selfing and outcrossing races. The change in breeding system toward autogamy is associated with a reduction in the size of floral features and a lower pollen production. The outbreeding populations have only been found in Greece, while the selfing populations have colonized much of Europe. The two races appear to be reproductively isolated from one another. —Petrorhagia sect.Kohlrauschia contains four species.P. glumacea is outbreeding and shows unilateral incompatibility with the small-flowered race ofP. prolifera and interfertility with the large-flowered race; althoughP. glumacea is sympatric with the large-flowered race, they do not appear to hybridize in the wild. Amongst the allopatric species, internal breeding barriers are also found. These may take the form of hybrid sterility or seed incompatibility.P. velutina is autogamous and reproductively isolated from all the other species of the section regardless of whether they are sympatric or allopatric. It is suggested that the evolution of autogamy has been an important factor in the spread ofP. velutina, the tetraploidP. nanteuilii, and the small-flowered race ofP. prolifera in Europe.     

Genetic control of durable resistance to yellow rust (Puccinia striiformis) in the wheat cultivar Hybride de Bersée

The wheat cultivar Hybride de Bersee is described as possessing durable resistance to yellow rust. Some races of Puccinia striiformis can infect it at the seedling stage but not severely in the field. Using euploid and aneuploid stocks of Bersee it was shown that a large part of this resistance was controlled by chromosome 5BS-7BS, the effect being detectable in seedlings and at later growth stages. Measurements of spore production from infected seedlings showed that the line ditelosomic for the 5BS arm of the chromosome was more resistant than the line ditelosomic for the 7BS arm under environments favouring high spore production, but more susceptible under environments favouring low spore production. Thus both arms of the chromosome could contribute to resistance in appropriate environments. Chromosome 5BS-7BS did not carry the factors controlling the race-specific resistance to race 37 E132 of P. striiformis. The possibility of exploiting the resistance carried by chromosome 5BS-7BS in wheat breeding programmes is discussed.     

Genetic Polymorphism and Evolution in Parthenogenetic Animals. II. Diploid and Polyploid SOLENOBIA TRIQUETRELLA (Lepidoptera: Psychidae)

Genic polymorphism at sixteen enzyme loci of four different chromosomal races of Solenobia triquetrella (bisexual, two diploid parthenogenetic races and tetraploid parthenogenetic) has been studied by starch gel electrophoresis. Isolated small diploid bisexual populations have rather uniform allele frequencies at all loci which we have studied. Diploid and tetraploid parthenogenetic individuals of this species are in general as heterozygous as bisexual ones. All parthenogenetic local populations are different from each other in the Alps. These parthenogenetic genotypes cannot be derived from a common ancestor through single mutations but rather bear evidence for a polyphyletic origin of parthenogenesis in Solenobia triquetrella. In the marginal distribution areas of the species in northern Europe single genotypes are spread over far larger areas than in the mountain regions of central Europe. This may be due to the old origin of parthenogenesis and polyploidy in northern Europe. No new parthenogenetic and polyploid strains have lately arisen in the regions outside of the Alps.     

A taxonomical re-evaluation of the Valais chromosome race of the common shrew<Emphasis Type="Italic">Sorex araneus</Emphasis> (Insectivora: Soricidae)

The common shrewSorex araneus Linnaeus, 1758 is subject to intense chromosomal polymorphism. About 65 chromosome races are presently known. One of these chromosome races (the Valais race) is karyologically, morphologically, biochemically, and genetically clearly distinct from all other chromosome races of the species. Recent studies of hybrid zones between the Valais race and other chromosome races in the Swiss and French Alps add further strong evidence for the specific taxonomic status of the Valais race. Chromosomes and diagnostic protein markers reveal sharp frequency clines and strong heterozygote deficits. In one hybrid zone, the maintenance of the strong genetic differentiation of the hybridizing taxa was confirmed by a study with autosomal microsatellites indicating minimal gene flow. A microsatellite marker on the Y-chromosome showed complete absence of male mediated gene flow suggesting hybrid male sterility. To clarify the taxonomic status of this taxon, additional analyses were conducted. A morphometric analysis of the mandible indicated the Valais race is morphologically as distinct from neighbouring chromosome races ofS. araneus as from other relatedSorex species. In a phylogeny based on complete mitochondrial DNA cytochromeb gene sequences, the Valais race clearly appears as the sister taxon to all other races ofS. araneus. Therefore, the chromosome race Valais ofS. araneus herein is elevated to specific status and the nameSorex antinorii Bonaparte, 1840 is applied.     

EXTREME KARYOTYPIC VARIATION IN A MUS MUSCULUS DOMESTICUS HYBRID ZONE: THE TOBACCO MOUSE STORY REVISITED

The Robertsonian fusion is a common chromosomal mutation among mammal species and is especially prevalent in the West European house mouse, Mus musculus domesticus. More than 40 races of the house mouse exist in Europe, including the famous “tobacco mouse” (Poschiavo race) of Val Poschiavo, Switzerland. Documented here is the discovery of an extreme case of karyotypic variation in the neighboring Upper Valtellina, Italy. In a 20-km stretch of the valley, 32 karyotypes were observed, including five chromosomal races and 27 hybrid types. One previously unknown race is reported, the “Mid Valtellina” race, with a diploid number of 2n = 24 and the Robertsonian fusions Rb(1.3), Rb(4.6), Rb(5.15), Rb(7.18), Rb(8.12), Rb(9.14), Rb(11.13), and Rb(16.17). The Poschiavo race (2n = 26), Upper Valtellina race (2n = 24), Lower Valtellina race (2n = 22) and all-acrocentric race (2n = 40) were also present. The races form a patchy distribution, which we term a “mottled hybrid zone.” Geographical position, isolation, extinction, recolonization, and selection against hybrids are all believed to be instrumental in the origin and evolution of this complex system. Previous studies of house mice from Upper Valtellina indicated that two of the races in the valley (the Upper Valtellina and Poschiavo races) may have speciated in the village of Migiondo. We discuss the possibility that there may have been a reinforcement event in this village.     

Molecular characterization of resistance to Heterodera glycines in soybean PI 438489B

Soybean (Glycine max L. Merr.) plant introduction (PI) 438489B is a newly found germplasm source that has resistance to multiple soybean cyst nematode (Heterodera glycines Ichinohe, SCN) races. We studied the inheritance of resistance to SCN races 1, 2, 3, 5 and 14 in PI 438489B using F₂ and F_2:3 families, which were generated by crossing to the susceptible cultivar ’Hamilton.’ The objectives of this study were to investigate the inheritance for resistance to SCN races in PI 438489B, to find molecular markers associated with resistances, and to study the allelic relationships among resistance loci for different SCN races. The results showed that the responses to SCN races were approximately normally distributed with large environmental effects, and were also highly correlated, which implied that genes giving resistance to different races were similar. The narrow-sense heritabilities of resistance to all five SCN races ranged from 0.55 to 0.88. Fifty one restriction fragment length polymorphism (RFLP) markers and 64 simple sequence repeat (SSR) markers were found to be polymorphic in the F₂ population. Quantitative trait loci (QTLs) associated with resistance to SCN races were anchored on soybean linkage groups (LGs) A1, A2, B1, B2, C1, C2, D1a, E and G. These QTLs explained 47.3%, 45.8%, 51.5%, 34.5% and 37.2% of the total phenotypic variances, respectively, for each race we investigated. Some QTLs for different races encompassed the same region of flanking markers; therefore, QTLs for multiple races may be linked or pleiotropic effects may be involved. Some loci provided resistance in a race-specific manner. Resistance to SCN race 14 had a different pattern compared to other races. Our results indicated that resistance to race 14 did not include loci on LGs A2 and G. These flanking markers associated with QTLs could be used to select for resistance to multiple SCN races in soybean breeding programs. Received: 25 March 2000 / Accepted: 4 August 2000