Correlated extinctions, colonizations and population fluctuations in a highly connected ringlet butterfly metapopulation

 The persistence of metapopulations is likely to be highly dependent on whether population dynamics are correlated among habitat patches as a result of migration between patches and spatially-correlated environmental stochasticity (weather effects). We examined whether population dynamics of the ringlet butterfly, Aphantopus hyperantus, were synchronous in an area of approximately 0.5 km², with respect to extinction, colonization and population fluctuations. Monks Wood Butterfly Monitoring Scheme transect count data from 1973 to 1995, revealed (A) a major environmental perturbation, the drought of 1976, which caused synchronized extinctions of A. hyperantus in subsequent years, (B) synchronized recolonization in years following the large number of apparent extinctions, and (C) population changes by A. hyperantus were highly correlated in many of the 14 sections of the transect, presumably reflecting similar responses to environmental stochasticity, and the exchange of individuals among sections. However, extinction and population synchrony depended on habitat type. Following the 1976 drought, A. hyperantus apparently became extinct from the most open and most shady habitats it occupied, with some persistence in habitats of intermediate shading, thus showing retraction to core populations in central parts of an environmental gradient, albeit with an average shift to relatively open habitat. Populations at extreme ends of the environmental gradient occupied by A. hyperantus fluctuated least synchronously, suggesting a potential buffering effect of habitat heterogeneity, but this was not crucial to survival after the 1976 drought. Thus, not all habitats are equally important to persistence. Correlated temporal dynamics, variation in habitat quality and the interaction between habitat quality and temporal environmental stochasticity are important determinants of metapopulation persistence and should be incorporated in metapopulation models. Received: 26 April 1996 / Accepted: 17 July 1996     

The effect of an agro-pasture landscape on diversity and migration patterns of frugivorous butterflies in Chiapas, Mexico

Few relevant data are available to analyze how landscape structure and composition affect the abundance and movement patterns of tropical insects. Using mark-release recapture experiments we examine the effect of an agro-pasture matrix on changes in diversity and migration of frugivorous butterflies in a tropical fragmented landscape in southern Mexico. In total, 53 frugivorous butterfly species were recorded in the entire landscape. Butterfly species composition was much more similar between sites than plant composition. A total of 3,501 individuals belonging to 41 butterfly species were captured, out of which 23 species (56%) were recaptured at least once. A large fraction of individuals was recaptured at the site of release (91%). We failed to find a significant relationship between the proportion occupied by the matrix and rates of residence, emigration, and immigration. Our results suggest that matrix quality in this and other traditionally managed agro-pasture landscapes plays a key role in both keeping important levels of biodiversity and maintaining constant movements of butterflies between otherwise isolated habitat patches.     

Comparative mapping of QTLs for agronomic traits of rice across environments by using a doubled-haploid population

We report here the RFLP mapping of quantitative trait loci (QTLs) which affect some important agronomic traits in cultivated rice. An anther culture-derived doubled-haploid (DH) population was established from a cross between indica and japonica rice varieties. A molecular linkage map comprising 137 markers was constructed based on this population which covered the rice genome at intervals of 14.8 cM on average. The linkage map was used to locate QTLs for such important agronomic traits as heading date, plant height, number of spikelets per panicle, number of grains per panicle, 1 000-grain weight and the percentage of seed set, by interval mapping. Evidence of genotype-by-environment interaction was found by comparing QTL maps of the same population grown in three diverse environments. A total of 22 QTLs for six agronomic traits was detected which were significant in at least one environment, but only seven were significant in all three environments; seven were significant in two environments and eight could only be detected in a single environment. However, QTLs-by-environment interaction was trait dependent. QTLs for spikelets and grains per panicle were common across environments while traits like heading date and plant height were more sensitive to environment. Received: 22 February 1996 / Accepted: 10 May 1996     

Relationships among climate,latitude and migration: Australian butterflies are not temperate-zone birds

We examined the distribution of butterflies over the mostly arid and semi-arid continent of Australia and analyzed the proportion of migrant species and species diversity with respect to an array of climatic and geographic variables. On a continent-wide scale, latitude explained virtually no variance in either proportion of migrants (r ²=0.01) or species diversity (r ²=0.03) in Australian butterflies. These results are in marked contrast to those for temperate-zone birds from three continents where latitude explained between 82 and 98% of the variance in frequency of migrants and also accounted for much of the variance in bird species diversity. In eastern Australia where rainfall regimes are similar to those in temperate Europe and North and South America, latitude explains 78% of the variance in frequency of butterfly migrants. In both eastern and central Australia, latitude also accounts for relatively high proportions of the variance in species diversity. Rainfall patterns and especially soil moisture are negatively associated with migration frequency in Australian butterfly faunas, both alone and in combination with other climate variables. Where moisture levels are relatively high, as in eastern Australia, measures of temperature are associated with migration frequency, a result consistent with findings for temperate-zone birds, suggesting latitude is a surrogate for temperature. The ultimate causes of migration in temperate-zone birds and Australian butterflies are the uneven temporal, and in Australia also spatial, distribution of resources. Uneven distribution is brought about primarily by temperature in temperate regions and by erratic rainfall over much of arid Australia. As a key determinant of productivity, especially in the tropics and subtropics, aridity is likely to be an important determinant of the global distributions of migrants. Received: 14 July 1999 / Accepted: 12 January 2000     

The diffusion model for migration and selection in a plant population

 The diffusion approximation is derived for migration and selection at a multiallelic locus in a partially selfing plant population subdivided into a lattice of colonies. Generations are discrete and nonoverlapping; both pollen and seeds disperse. In the diffusion limit, the genotypic frequencies at each point are those determined at equilibrium by the local rate of selfing and allelic frequencies. If the drift and diffusion coefficients are taken as the appropriate linear combination of the corresponding coefficients for pollen and seeds, then the migration terms in the partial differential equation for the allelic frequencies have the standard form for a monoecious animal population. The selection term describes selection on the local genotypic frequencies. The boundary conditions and the unidimensional transition conditions for a geographical barrier and for coincident discontinuities in the carrying capacity and migration rate have the standard form. In the diallelic case, reparametrization renders the entire theory of clines and of the wave of advance of favorable alleles directly applicable to plant populations. Received 30 August 1995; received in revised form 23 February 1996     

Linking individual foraging behavior and population spatial distribution in patchy environments: a field example with Mediterranean blue tits

 In spite of recent theoretical interest, few field studies have addressed the links between individual behavioral decisions and population distribution. This work analyzes the foraging behavior of individuals and the spatial distribution of a population of blue tits (Parus caeruleus) just before the main breeding season, when blue tit foraging was not affected by central-place or flocking behaviors. The study was carried out in open holm oak Quercus ilex woodlands (dehesas) that are patchy for canopy-foraging birds because of the scattered arrangement of trees. Residence time on each tree was not correlated either with previous flight time or with prey abundance in trees. Flight distances between trees were larger than average distances estimated in random samples of holm oaks taken close to foraging birds. Trees were not selected by birds on the basis of their expected energy costs and rewards. Bird abundance was not related to food availability in trees or to tree size across dehesas. However, bird abundance was strongly correlated with tree density and with the availability of tree holes for nesting, to the extent that the proportion of tits matched the proportions of both tree abundance and hole abundance across study plots. Overall, neither the behavior of individuals nor the distribution of the population of blue tits corresponded with food resources, which appeared superabundant; instead, tits appeared to behave and be distributed according to the distribution of structural resources such as trees and tree holes for nesting. Received: 18 August 1996 / Accepted: 14 March 1997     

Mapping of two genes for resistance to clubroot (Plasmodiophora brassicae) in a population of doubled haploid lines of Brassica oleracea by means of RFLP and AFLP markers

A genetic map covering 615 cM in 12 linkage groups was assembled based on 92 RFLP and AFLP markers segregating in a population of 107 doubled haploid lines (DH lines) of Brassica oleracea. The DH-line population was obtained through microspore culture from the of two homozygous parents: DH-line Bi derived from the cabbage landrace Bindsachsener, and DH-line Gr from broccoli cv ‘Greenia’. Sixty-five percent of the loci, and in some cases complete linkage groups, displayed distorted segregation ratios, a frequency much higher than that observed in populations of the same species. DH-line Bi was resistant to clubroot, which is caused by a Dutch field isolate of Plasmodiophora brassicae. Resistance in the DH-line population was determined in two ways: by assigning symptom grades to each plant, and by measuring the fresh weights of the healthy and affected parts of the root system of each plant. Using a multiple QTL mapping approach to analyze the fresh weight data, we found two loci for clubroot resistance; these were designated pb-3 and pb-4. The additive effects of these loci were responsible for 68% of the difference between the parents and for 60% of the genetic variance among DH-line means. Also, indications for the presence of two additional, minor QTLs were found. Analysis of symptom grades revealed the two QTLs pb-3 and pb-4, as well as one of the two minor QTLs indicated by analysis of the fresh weight data. Received: 29 April 1996 / Accepted: 10 May 1996     

On the role of body size in a tri-trophic metapopulation model

 A particular tri-trophic (resource, prey, predator) metapopulation model with dispersal of preys and predators is considered in this paper. The analysis is carried out numerically, by finding the bifurcations of the equilibria and of the limit cycles with respect to prey and predator body sizes. Two routes to chaos are identified. One is characterized by an intriguing cascade of flip and tangent bifurcations of limit cycles, while the other corresponds to the crisis of a strange attractor. The results are summarized by partitioning the space of body sizes in eight subregions, each one of which is associated to a different asymptotic behavior of the system. Emphasis is put on the possibility of having different modes of coexistence (stationary, cyclic, and chaotic) and/or extinction of the predator population. Received 1 August 1995; received in revised form 8 January     

The effects of plant dispersion and prey density on parasitism rates in a naturally patchy habitat

Despite extensive research on parasitoid-prey interactions and especially the effects of heterogeneity in parasitism on stability, sources of heterogeneity other than prey density have been little investigated. This research examines parasitism rates by three parasitoid species in relationship to prey density and habitat spatial pattern. The herbivore Itame andersoni (Geometridae) inhabits a subdivided habitat created by patches of its host plant, Dryas drummondii, in the Wrangell Mountains of Alaska. Dryas colonizes glacial moraines and spreads clonally to form distinct patches. Habitat subdivision occurs both on the patch scale and on the larger spatial scale of sites due to patchy successional patterns. Itame is attacked by three parasitoids: an ichneumonid wasp (Campoletis sp.), a braconid wasp (Aleiodes n. sp.), and the tachinid fly (Phyrxe pecosensis). I performed a large survey study at five distinct sites and censused Itame density and parasitism rates in 206 plant patches for 1–3 years. Parasitism rates varied with both plant patch size and isolation and also between sites, and the highest rates of overall parasitism were in the smallest patches. However, the effects of both small- and large-scale heterogeneity on parasitism differed for the three parasitoid species. There was weak evidence that Itame density was positively correlated with parasitism for the braconid and tachinid at the patch scale, but density effects differed for different patch sizes, patch isolations, and sites. At the site scale, there was no evidence of positive, but some indication of negative density-dependent parasitism. These patterns do not appear to be driven by negative interactions between the three parasitoid species, but reflect, rather, individual differences in habitat use and response to prey density. Finally, there was no evidence that parasitism strongly impacted the population dynamics of Itame. These results demonstrate the importance of considering habitat pattern when examining spatial heterogeneity of parasitism and the impacts of parasitoids. Received: 3 June 1999 / Accepted: 4 October 1999     

Flow cytometric analysis of the chromosomes and stability of a wheat cell-culture line

A rapidly growing, long-term suspension culture derived from Triticum aestivum L. (wheat) was synchronized using hydroxyurea and colchicine, and a chromosome suspension with chromosomes was made. After staining with the DNA-specific fluorochromes Hoechst 33258 and Chromomycin univariate and bivariate flow-cytometry histograms showed 15 clearly resolved peaks corresponding to individual chromosome types or groups of chromosomes with similar DNA contents. The flow karyotype was closely similar to a histogram of DNA content measurements of Feulgen-stained chromosomes made by microdensitometry. We were able to show the stability of the flow karyotype of the cell line over a year, while a parallel subculture had a slightly different, stable, karyotype following different growth conditions. The data indicate that flow cytometric analysis of plant karyotypes enables accurate, statistically precise chromosome classification and karyotyping of cereals. There was little overlap between individual flow-histogram peaks, so the method is useful for flow sorting and the construction of chromosome specific-recombinant DNA libraries. Using bivariate analysis, the AT:GC ratio of all the chromosomes was remarkably similar, in striking contrast to mammalian flow karyotypes. We speculate about a fundamental difference in organization and homogenization of DNA sequences between chromosomes within mammalian and plant genomes. Received: 24 April 1996 / Accepted: 24 May 1996     

The scale of resource specialization and the distribution and abundance of lycaenid butterflies

Numerous hypotheses have been proposed for the commonly observed, positive relationship between local abundance and geographic distribution in groups of closely related species. Here I consider how hostplant specialization and abundance affect the relative abundance and distribution of lycaenid butterflies (Lepidoptera: Lycaenidae). I first discuss three components of specialization: local specialization, turnover of specialization across a species’ range, and the minimum number of resources (or habitats) required by a species. Within this framework, I then consider one dimension of a lycaenid species’ niche, larval hostplant specialization. In a subalpine region of Colorado, I surveyed 11 lycaenid species and their hostplants at 17 sites. I compare this local information to continental hostplant use and large-scale distributions of the lycaenids and their hostplants. Local abundance of a lycaenid species is positively correlated with its local distribution (the number of sites occupied), but not with its regional or continental distribution. Neither local specialization (the number of hostplants used within one habitat) nor continental specialization (the number of hostplants used across many habitats) is correlated with local lycaenid abundance. Continental specialization is positively correlated with a species’ continental distribution, however. Finally, while generalist butterflies tend to have more hostplant available to them, differences in resource availability do not explain the differences in butterfly abundance. Although local abundance is correlated only with local distribution, I suggest that abundance-distribution relationships might emerge at regional and continental scales if local abundance were averaged across many habitat types. Consideration of the scale of a species’ resource specialization (within or among habitats) appears to be key to understanding the relationships between resource specialization, resource availability, and a species’ abundance and distribution. Received: 1 September 1999 / Accepted: 12 December 1999     

Choice of optimal oviposition sites by Hoplobatrachus occipitalis (Anura: Ranidae) in an unpredictable and patchy environment

 The rock pools on the river bank of the Comoé National Park (West Africa) provide a very diverse and unpredictable environment for anuran larval development. Because rock pools differ considerably in biotic and abiotic parameters, it should be adaptive for reproducing anurans to choose the most suitable oviposition sites. During the beginning of each rainy season (March to May), from 1991 to 1995, we investigated the allocation of breeding sites by Hoplobatrachus occipitalis, counted the number of eggs, and measured several biotic and abiotic factors that might influence the choice of a spawning site. The probability of predation by conspecific cannibalistic tadpoles and the water-holding capacity (WHC) of pools were the best predictor of number of eggs laid. We experimentally investigated the influence of these two parameters on egg-laying and showed that adults potentially can assess the presence, density, and size of tadpoles in pools by chemical cues. Likewise, manipulation of the water-holding capacity caused a rapid change in egg-laying behavior. To assess the risk of desiccation, frogs have to visit familiar pools repeatedly to monitor the decrease in volume over time and thus gain information about the pools’ water-holding capacity. Received: 22 April 1996 / Accepted: 17 July 1996     

Highly polymorphic microsatellites of rice consist of AT repeats, and a classification of closely related cultivars with these microsatellite loci

Microsatellites consisting of AT repeats are highly polymorphic in rice genomes and can be used to distinguish between even closely related japonica cultivars in Japan. Polymorphisms of 20 microsatellite loci were determined using 59 japonica cultivars, including both domestic and modern Japanese cultivars. Although the polymorphisms of these 20 microsatellite loci indicated that the Japanese cultivars were genetically quite similar, microsatellites consisting of AT repeats showed high gene diversity even among such closely related cultivars. Combinations of these hypervariable microsatellites can be employed to classify individual cultivars, since the microsatellites were stable within each cultivar. An identification system based on these highly polymorphic microsatellites could be used to maintain the purity of rice seeds by eliminating contamination. A parentage diagnosis using 17 polymorphic microsatellite loci clearly demonstrated that plants which carried desired chromosome regions had been selected in breeding programs. Thus, these hypervariable microsatellites consisting of AT repeats should promote the selection of plants which carry desired chromosomes from genetically similar parents. Backcrossing could also help to eliminate unnecessary chromosome regions with microsatellite polymorphisms at an early stage in breeding programs. Received: 8 July 1996 / Accepted: 12 July 1996     

The effects of two parasitoids on the life history and metapopulation structure of the intertidal snail Littoraria filosa in different-sized patches of mangrove forest

We describe differences in life history of the intertidal arboreal snail, Littoraria filosa, among patches of mangroves ranging in size from isolated trees to large stands several square kilometres in area. Recruitment of L. filosa occurred from mid spring (October) to early winter (June), recruits grew rapidly and copulating adults were found during the following September–April. Populations within large patches of forest were annuals; all or most individuals died between October–January (spring–midsummer). In contrast, those in smaller peripheral patches were more likely to survive over the summer but survival differed among patches and years. These differences in life history were caused by a parasitoid fly (genus Sarcophaga) that attacked L. filosa 10 mm and longer and was present in all large patches, but absent from, or rare, in smaller peripheral patches. Experimental introductions to isolated trees confirmed that the fly could kill L. filosa. Another sarcophagid parasitoid that attacked L. filosa from 4 to less than 10 mm long was also found in every patch. The combined effects of these parasitoids appear to determine the metapopulation structure of L. filosa. Most adults in large patches were killed by the larger fly during early summer. Summer recruits were often killed by the smaller fly within a month of settlement and when this happened effective recruitment of L. filosa was reduced to autumn. The planktotrophic larval stage of L. filosa lasts less than 1 month, so the source of autumn recruits to all patches must have been adults that survived the early summer, most of which were in small patches or on isolated trees. Consequently these ”peripheral sources” are likely to be important for persistence of the metapopulation of L. filosa. The results of this study demonstrate that metapopulation structure may be determined by complex interactions and that common models cannot be assumed to apply in all habitats. Received: 15 September 1999 / Accepted: 31 January 2000     

Characterization of spelt (Triticum spelta L.) forms by gel electrophoretic analyses of seed storage proteins. I. The gliadins

A comparison betweeen the electropherograms of the spelt and wheat cultivars showed specific differences in the gliadin band patterns which provided the possibility of a clear classification into spelt or wheat. A special nomenclature was developed to be able to improve the presentation of the gliadin band pattern of spelt, which is different from that of wheat. This nomenclature, however, has not yet been applied to other cereals. The gliadin band patterns were presented in a schematic form. As a parameter for comparison, idealized band patterns of both wheat and spelt were developed by comparing the proportions of the bands of all available types. When comparing the gliadin band patterns of the spelt cross-breeds with their corresponding parental generations, it was noted that the same parental bands were not always transmitted and that the cross-breeds showed differences in the intensity, mobility, occurrence, and the splitting of single bands. In general it can be said that the band pattern of the daughter generation – even in the examined and generations – is more similar to the band pattern of the mother than to that of the father, which proves a maternal effect. Received: 29 June 1996 / Accepted: 12 July 1996     

The effect of migration on metapopulation stability is qualitatively unaffected by demographic and spatial heterogeneity

Coupled map lattices (CMLs), using two coupled logistic equations, have been extensively used to model the dynamics of two-patch ecological systems. Such studies have revealed that migration rate plays an important role in determining the dynamics of the system, particularly when the two maps differ in their intrinsic growth rate parameter, r. However, under more realistic assumptions, a metapopulation can be expected to consist of more than two subpopulations, each with its own demographic parameters, which will in part be a function of the environment of that patch. The role of the spatial arrangement of heterogeneous (i.e. with different r values) subpopulations in shaping the dynamics of such a metapopulation has rarely been investigated. Here, we study the effect of demographic and spatial heterogeneity on the stability of one- and two-dimensional systems of 64 coupled Ricker maps with different r values, under periodic and absorbing boundary conditions. We show that the effects of migration rate on metapopulation stability do not depend upon either the precise spatial arrangement of the subpopulations in the lattice, or on the presence of a moderate proportion of vacant (uninhabitable) patches in the lattice. The results, thus, suggest that metapopulation models are robust to variation in spatial arrangement of patch quality and, hence, of demographic parameters. We also show that for any given arrangement of the patches, maximum stability of the metapopulation occurs when the migration levels are intermediate, a result that agrees well with previous studies on two-map CML systems.     

The mapping of phytochrome genes and photomorphogenic mutants of tomato

The map positions of five previously described phytochrome genes have been determined in tomato (Lycopersicon esculentum Mill.) The position of the yg-2 gene on chromosome 12 has been confirmed and the classical map revised. The position of the phytochrome A (phy A)-deficient fri mutants has been refined by revising the classical map of chromosome 10. The position of the PhyA gene is indistinguishable from that of the fri locus. The putative phyB1-deficient tri mutants were mapped by classical and RFLP analysis to chromosome 1. The PhyB1 gene, as predicted, was located at the same position. Several mutants with the high pigment (hp) phenotype, which exaggerates phytochrome responses, have been reported. Allelism tests confirmed that the hp-2 mutant is not allelic to other previously described hp (proposed here to be called hp-1) mutants and a second stronger hp-2 allele (hp-2 ^j ) was identified. The hp-2 gene was mapped to the classical, as well as the RFLP, map of chromosome 1. Received: 24 May 1996 / Accepted: 14 June 1996     

Genetic elimination of a starch granule protein, SGP-1, of wheat generates an altered starch with apparent high amylose

A starch granule protein, SGP-1, is a starch synthase bound to starch granules in wheat endosperm. A wheat lacking SGP-1 was produced by crossing three variants each deficient in one of three SGP-1 classes, namely SGP-A1, -B1 or -D1. This deficient wheat (SGP–1 null wheat) showed some alterations in endosperm starch, meaning that SGP-1 is involved in starch synthesis. Electrophoretic experiments revealed that the levels of two starch granule proteins, SGP-2 and -3, decreased considerably in the SGP-1 null wheat though that of the waxy protein (granule-bound starch syn- thase I) did not. The A-type starch granules were deformed. Apparent high amylose level (30.8–37.4%) was indicated by colorimetric measurement, amperometric titration, and the concanavalin A method. The altered structure of amylopectin was detected by both high- performance size-exclusion chromatography and high-performance anion exchange chromatography. Levels of amylopectin chains with degrees of polymerization (DP) 6–10 increased, while DP 11–25 chains decreased. A low starch crystallinity was shown by both X-ray diffraction and differential scanning calorimetry (DSC) analyses because major peaks were absent. Abnormal crystallinity was also suggested by the lack of a polarized cross in SGP-1 null starch. The above results suggest that SGP-1 is responsible for amylopectin synthesis. Since the SGP-1 null wheat produced novel starch which has not been described before, it can be used to expand variation in wheat starch. Received: 30 April 1999 / Accepted: 9 November 1999     

Analytical solutions of a minimal model of species migration in a bounded domain

 A minimal model of species migration is presented which takes the form of a parabolic equation with boundary conditions and initial data. Solutions to the differential problem are obtained that can be used to describe the small- and large-time evolution of a species distribution within a bounded domain. These expressions are compared with the results of numerical simulations and are found to be satisfactory within appropriate temporal regimes. The solutions presented can be used to describe existing observations of nematode distributions, can be used as the basis for further work on nematode migration, and may also be interpreted more generally. Received: 15 August 1999     

Shading reduces exploitation of soil nitrate and phosphate by Agropyron desertorum and Artemisia tridentata from soils with patchy and uniform nutrient distributions

 Shading may both lessen the demand for soil nutrients and also the energy supply for nutrient acquisition. Since root foraging for nutrients in patchy environments can be energy-costly, especially for an immobile nutrient such as phosphate (P), the effects of shading may be most expected in heterogeneous soils. Plant acquisition of nitrate (N) and phosphate from soils with patchy and uniform nutrient distributions was determined in a field study under open sunlight and with shading for two common perennial Great Basin shrub steppe species, Agropyron desertorum and Artemisia tridentata. Partial shading in a pattern which can occur in shrub steppe vegetation significantly decreased plant N and P acquisition from soils both in the patchy and the uniform nutrient treatments. Artemisia was more affected by the shading than was Agropyron. Exploitation of the rather immobile P ion by both species was reduced to a much greater degree by the shading in the patchy distribution treatment than in the uniform nutrient treatment. As expected, plant acquisition of the more mobile N varied little with nutrient distribution treatment for both species and the depression of N acquisition by shading was the same in both nutrient distributions. The effects of shading appeared to have had its primary influence on different components of root foraging in the two species, especially in the nutrient-rich patches. For Agropyron shading primarily affected root proliferation, as indicated by reduced root density in patches. For Artemisia, shading most influenced root physiological uptake capacity and this was most pronounced in the nutrient-rich patches. While aboveground competition for light may generally reduce nutrient acquisition, the effects appear to be most pronounced if root systems of these steppe species are foraging for nutrients such as P in spatially heterogeneous soils. Received: 29 February 1996 / Accepted: 16 July 1996