Foraging and community consequences of seed size for coexisting Negev Desert granivores

We examined the effects of seed size on patch use and diet selection for three co-existing Negev Desert granivores: Allenby's gerbil ( Gerbillus allenbyi ), greater Egyptian sand gerbil ( Gerbillus pyramidum ), and crested lark ( Galerida cristata ). We manipulated size and spatial distribution of seeds in experimental food patches and quantified foraging behavior by measuring giving-up densities (GUDs: the amount of food remaining in a resource patch following exploitation by a forager). In one experiment, we presented small (<1.4 mm in diameter cracked wheat), medium (2.0–3.3 mm), and large (>3.4 mm) seeds in separate trays; in a second, we presented small and medium seeds separately and mixed together. Gerbils had a higher handling time efficiency on smaller seeds, but a much higher encounter probability on larger seeds (20 times higher on large than medium seeds, and 2–5 times higher on medium than small seeds). This led gerbils to have significantly lower GUDs on larger seeds than smaller seeds and to harvest a higher proportion of the larger seeds. When presented with rich and poor patches, G. allenbyi tended to equalize GUDs in both patches, indicating a quitting harvest rate rule for patch exploitation. In contrast, larks appeared to use a fixed time rule for patch exploitation. For larks, seed size did not influence encounter probabilities, and they showed no seed-size selectivity. Still, larks had higher handling efficiencies on smaller than larger seeds, and consequently had a significantly lower GUD on small than medium seeds. Despite large differences between the gerbils and larks in their foraging, our results do not support species coexistence via seed-size partitioning: the larks had much higher GUDs than the gerbils on all seed sizes. Nonetheless, seed size, seed abundance, seed distribution and the animal's patch use behavior all played major roles in determining gerbils' and larks' diet selectivities and GUDs.     

Allelopathic effects of three plant invaders on germination of native species: a field study

The ability of some invasive plant species to produce biochemical compounds toxic to native species, called allelopathy, is thought to be one of the reasons for their success when introduced to a novel range, an idea known as the Novel Weapons Hypothesis. However, support for this hypothesis mainly comes from bioassays and experiments conducted under controlled environments, whereas field evidence is rare. In a field experiment, we investigated whether three plant species invasive in Europe, Solidago gigantea, Impatiens glandulifera and Erigeron annuus, inhibit the germination of native species through allelopathy more than an adjacent native plant community. At three sites for each invasive species, we compared the germination of native species that were sown on invaded and non-invaded plots. Half of these plots were amended with activated carbon to reduce the influence of potential allelopathic compounds. The germination of sown seeds and of seeds from the seedbank was monitored over a period of 9 weeks. Activated carbon generally enhanced seed germination. This effect was equally pronounced in invaded and adjacent non-invaded plots, indicating that invasive species do not suppress germination more than a native plant community. In addition, more seeds germinated from the seedbank on invaded than on non-invaded soil, probably due to previous suppression of germination by the invasive species. Our field study does not provide evidence for the Novel Weapons Hypothesis with respect to the germination success of natives. Instead, our results suggest that if invasive species release allelopathic compounds that suppress germination, they do so to a similar degree as the native plant community.     

An Experimental Study of Competition Between Seed-eating Desert Rodents and Ants

Reciprocal increases in rodent and ant densities on 0.1 ha plotsfrom which the other taxon had been excluded demonstrate thatthese distantly related desert granivores compete for seeds.Relative to unmanipulated control plots, numbers of ant coloniesincreased 71% on plots where rodents were excluded; rodentsincreased 20% in numbers of individuals and 29% in biomass inthe absence of ants. Comparisons of seed levels in the soiland of annual plant densities on experimental and control plotsprovide evidence that the rodent and ant populations are limitedby and compete for food. Greater numbers of seeds and annualsoccurred on plots where rodents and ants had been excluded thanon plots where both taxa were present. Particular species ofannuals were reduced in density by foraging of rodents. Antsincreased species diversity by differentially harvesting seedsof the most common species. Results of these and other recentstudies suggest that competition among distantly related organismsplays a major role in the organization of ecological communities.     

Contingent fire effects on granivore removal of exotic woody plant seeds in longleaf pine savannas

Prescribed fire is increasingly used to inhibit woody encroachment into fire-dependent ecosystems, yet its effects on other processes influencing invasion are poorly understood. We investigated how fire influences exotic woody invasion through its effects on granivore activity, and whether these effects depend on the habitat in which seed predation takes place. We quantified seed removal for four species of exotic woody plants (Albizia julibrissin, Elaeagnus umbellata, Melia azedarach and Triadica sebifera) in 17 sites in longleaf pine savanna that varied in time since fire (one or three growing seasons post-fire) in the sandhills region of North Carolina, USA. Within each site, we established paired plots in upland and upland-wetland ecotone communities and presented seeds in depots that allowed either arthropod, or arthropod and small vertebrate access. We found that differences in seed removal with time since fire were contingent on habitat and granivore community. In ecotones, three of four species had higher proportions of seeds removed from plots that were three growing seasons post-fire than plots one growing season post-fire, whereas only T. sebifera showed this pattern in upland habitat. Allowing vertebrate granivores access to seeds enhanced seed removal, and this effect was strongest in ecotone habitat. While granivores removed seeds of all four plant species, removal of E. umbellata was significantly higher than that of the other species, suggesting that granivores exhibited seed selection. These findings suggest that ecotone habitats in this system experience greater seed removal than upland habitats, particularly as time since fire increases, and differences are mainly due to the activity of vertebrate granivores. Such differences in seed removal, together with seed selection, may contribute to variation in exotic woody invasion of longleaf pine savannas.     

Taxonomic status of the Forest Buzzard Buteo oreophilus trizonatus

The foraging efficiencies of four sympatric southern African seed-eating birds, namely Bronze Mannikin Spermestes cucullatus, Cape Sparrow Passer melanurus, Southern Red Bishop Euplectes orix and Thick-billed Weaver Amblyospiza albifrons, and a domesticated species, the Bengalese Finch Lonchura domestica, were measured and compared using giving-up densities (the amount of food remaining following patch exploitation) in experimental food patches. Foraging efficiency was quantified using giving-up densities by offering individual birds equal foraging opportunities. A low giving-up density displays the ability of a forager to profitably harvest food at low resource densities and to gainfully exploit the foraging opportunities overlooked by a less efficient forager. Ten individuals of each of the five species were allowed to forage on six different seed types. Thick-billed Weavers had significantly lower giving-up densities for all seed types except the smallest, namely red manna. Bronze Mannikins showed the converse trend, foraging most efficiently on the smallest seeds. The results of the present study revealed that Thick-billed Weavers were the most efficient foragers (i.e. had the lowest giving-up densities on seeds in feeding trays).     

What do mice select for in seeds?

Summary Knowledge of the basis upon which granivores select seeds is crucial to the understanding of granivory. In this study the preferences of three rodent granivores among seeds of 11 plants from the semi-arid Karoo of South Africa were estimated, and related to the physical and chemical attributes of the seeds. Seed weights and calorific, moisture, protein, polyphenol, ash, lipid and silica contents were estimated and cell contents, soluble ash and soluble carbohydrate contents were derived from these values. These attributes were determined for both the intact seeds and the portion of the seed that is ingested by the mice. The efficiency with which mice ingested the seeds (in terms of time and mass) was recorded. All three mouse species ranked the seeds similarly, and the two species for which handling efficiency was measured did not differ in this regard. Preference hierachies were highly correlated with the rate of energy intake, as predicted by optimal foraging theory. There was no correlation between rodent preferences and the gross energy content of the seeds, emphasising the importance of measuring relevant parameters. The energy yield of the seeds calculated here, in conjunction with rodent population energy requirements and dietary data, may be used to estimate potential granivore impact on the seed production of the plant community.     

Foraging,food selection and worker size in the seed harvesting ant <Emphasis Type="Italic">Messor bouvieri</Emphasis>

Interactions among granivores and seeds depend on the foraging behaviour and morphology of the granivores and on the attributes and availability of seeds. We investigated seed selection by the seed harvesting ant Messor bouvieri in three adjacent plant communities in Spain by relating the harvested seeds to those in the seed rain. Preference was positively correlated with seed size and abundance which accounted for 43% and 20% of the variance respectively. Contrary to predictions of central place foraging theory, the size of seeds harvested did not increase with distance from the nest. Inclusion of a less-preferred item in the diet was more strongly related to the abundance of more-preferred items (60% of the variance) than the abundance of the less-preferred item (14% of the variance). Worker size accounted for 20–30% of the variance in the size of harvested seeds, although small workers did not appear to be constrained by load size for the range of seeds available. The body size of ants was significantly larger in the community with the greatest proportion of large seeds, although this was not due to their ability to carry larger loads or due to the greater force required to crush these seeds. The strong preference of M. bouvieri for large seeds may have important consequences for the plant communities in which they forage. Received: 30 November 1999 / Accepted: 14 April 2000     

Effects of body size,seed density,and soil characteristics on rates of seed harvest by heteromyid rodents

Summary Although it is well established that coexisting heteromyid rodent species forage in different microhabitats, we do not yet know the basis for divergent microhabitat choice. One possibility is that seed harvest rates differ among microhabitats, and each species forages where it can extract seeds most efficiently. Microhabitats vary in several factors that could affect heteromyid foraging efficiency, including seed density, soil organic content and particle size distribution. We have explored the effect of each of these variables on harvest rates of several species feeding from petri dishes containing known densities of millet seeds embedded in soil of known particle size and density. Results indicate that the number of seeds harvested per second increases uniformly with seed density and soil density and decreases with soil particle size. Body size affects these relationships: larger animals have higher harvest rates for a given set of conditions and experience a greater relative change in harvest rate for a give change in conditions. This implies that heteromyids can be expected to exhibit species-specific microhabitat preferences while foraging in nature.     

Competition between birds and mammals: A comparison of giving-up densities between crested larks and gerbils

We combined the concept of mechanisms of co-existence with the approach of giving-up densities to study inter-taxon competition between seed-eating birds and mammals. We measured feeding behaviour in food patches to define and study the guild of seed-eating vertebrates occupying sandy habitats at Bir Asluj, Negev Desert, Israel. Despite a large number of putatively granivorous rodents and birds at the site, two gerbil species (Allenbys gerbil, Gerbillus allenbyi, and the greater Egyptian gerbil, G. pyramidum) dominated nocturnal foraging, and a single bird species (crested lark, Galerida cristata) contributed all of the daytime foraging. We used giving-up densities to quantify foraging behaviour and foraging efficiencies. A low giving-up density demonstrates the ability of a forager to profitably harvest food at low abundances and to profitably utilize the foraging opportunities left behind by the less efficient forager. Gerbils had lower giving-up densities in the bush than open microhabitat, and lower giving-up densities in the semi-stabilized than stabilized sand habitats. Crested larks showed the opposite: lower giving-up densities in the open than bush, and on the stabilized than semi-stabilized sand habitats. Despite these patterns, gerbils had substantially lower giving-up densities than crested larks in both microhabitats, all sand habitats, and during each month. Several mechanisms may permit the crested lark to co-exist with the gerbils. Larks may be cream skimmers on the high spatial and temporal variability in seed abundances. Larks may rely on insects, fruit or smaller seeds. Or, larks may rely on adjacent rocky habitats.     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997     

Seed and microsite limitation in a late-successional old field: the effects of water, adults, litter, and small mammals on seeds and seedlings

Many plant communities are recruitment limited, which may occur because there are either too few seeds to fill available microsites, too few available microsites, or both. In a recruitment-limited, Minnesota, USA old field, we tested among these alternatives in a three-phase study. In phase 1, we estimated the production of late-successional forb and C4 grass seeds. In phase 2, we experimentally modified field establishment conditions with rainfall amendment, adult plant thinning, litter reduction, and small mammal exclusion. We then measured recruitment. On average, of the nearly 2,600 seeds produced m^?2, only 164 seeds m^?2 were present and living after overwintering, as measured by field and greenhouse germination. Furthermore, on average, only 9 of those 164 seeds m^?2 germinated in the field, even under the relaxed establishment conditions of our four experimental treatments. Although adults of C4 grasses dominate the field, surprisingly few C4 grass seedlings germinated. To understand why, in phase 3, we added seed of the three dominant C4 grasses into the same plots the following year. Their ability to recruit into control plots compared with treated plots was relatively lower than for the ambiently recruited forbs from phase 2, suggesting that the competitively dominant C4 grasses have greater difficulty establishing in the extant community. Of the seeds that did germinate in the field in phases 2 and 3, all four experimental treatments significantly improved at least one stage of establishment, with the rainfall amendment having the greatest overall effect across species. In total, our results suggest that seed limitation was exacerbated by microsite limitation via multiple mechanisms.     

Effects of harvester ants on plant species distribution and abundance in a serpentine grassland

Seed harvesting ants can have important effects on the composition and structure of plant communities. We investigated two effects of Messor andrei, the black seed-harvesting ant, on a serpentine grassland plant community in northern California. First, to determine if selective seed predation by ants affects plant community composition, we excluded harvester ants from 1-mediameter circular plots of grassland. Abundances of all species on these plots and on control plots were measured before and after exclosure. Second, to determine if M. andrei nest mounds affect plant community composition, we compared plant species abundances on and off nest mounds. M. andrei deposit large amounts of organic matter on their nest mounds over a foraging season, so mounds may alter the edaphic environment. The exclusion of seed-harvesting activity did not cause changes in the plant community. Nest mounds had a strong effect on plant communities: there were many more grasses and fewer forbs on ant mounds, although at least one forb, Lepidium nitidum, produced twice as many seeds when it grew on nest mounds. We found that nest mounds formed islands of higher-temperature soil in the serpentine grassland. Received: 31 March 1997 / Accepted: 6 May 1997     

Effects of grazing on soil seed bank dynamics: An approach with functional groups

Abstract. The relationship between intensity and timing of cattle grazing on changes in the size and composition of the soil seed bank were investigated in a 3‐yr study in a Mediterranean grassland in northeastern Israel. Treatments included manipulations of stocking rates and of grazing regimes, in a factorial design. The retrieved soil seed bank community was rich in species, with 133 species accounting for 80% of the 166 species recorded at the site. Within the seed bank, 89% of the species were annuals. Seed bank dynamics was analysed in terms of plant functional groups and germination strategies. In terms of total seed bank density and including all functional groups, 42% of the seeds present in the soil did not germinate under watering conditions. The dormancy level differed greatly among functional groups. The seed bank of annual legumes, crucifers, annual thistles and annual forbs had a large fraction of non‐germinated seeds and characterized areas grazed early in the growing season under high and very high grazing intensity. These functional groups were considered to have a higher potential for persistent seed banks production. In contrast, short and tall annual grasses and tall perennial grasses, that were dominant in ungrazed or moderately grazed paddocks, generally had seed banks with a very small fraction of non‐germinated seeds. Seed bank densities varied widely between grazing treatments and years. Under continuous grazing, heavy grazing pressure reduced seed bank densities of grasses and crucifers in comparison to moderate grazing. The greatest reduction on the seed bank densities resulted from heavy grazing concentrated during the seed‐set stages.     

The strength of seeds and their destruction by granivorous insects

The influence of seed structure and strength on their destruction by granivores is central to understanding the dynamics of granivore-plant interactions. For up to nine seed species, the effects of seed size (cm³), mass (mg), density (mg/cm³) and coat strength (MPa) on the damage inflicted by three post-dispersal granivores (Harpalus pensylvanicus, Anisodactylus sanctaecrucis, and Gryllus pennsylvanicus) were evaluated. Seed destruction rates by G. pennsylvanicus were statistically unrelated to the size and toughness of the seeds. Seed densities significantly affected their destruction by A. sanctaecrucis and H. pensylvanicus, as did seed size, mass, and strength in H. pensylvanicus under choice conditions. The carabid beetles destroyed more of the small, denser seeds with stronger seed coats. The results show that different granivores are able to distinguish the structural strength and physical density of seeds as well as seed size. The relative ability of granivores to detect these seed characteristics offers a way in which diverse communities of post-dispersal insect granivores can persist within a single habitat. The authors redefine how the strength of biological structures should be evaluated in ecological studies, using guidelines commonplace in the field of engineering. Handling Editor: Stanislav Gorb The U.S. Government’s right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged     

Small vertebrate granivores reduce seedling emergence in native tallgrass prairie restoration

High seed cost and low rates of establishment make tallgrass prairie restorations challenging and expensive endeavors. Typical seedling emergence rates in prairie restorations are approximately 10% and the causes of seed mortality are poorly understood. In this study, we examined the impact of small vertebrate granivores on prairie restoration by comparison of seedling emergence in open (sham) versus closed exclosures at three newly restored sites. To assess other causes of seed loss, we also tracked seed fates at one prairie restoration site. We coated seeds of four prairie species with fluorescent dye, placed them under closed exclosures, and monitored their fate (emerging seedling, partially germinated, nongerminated/viable, and nongerminated/nonviable) over a 5‐month period. On average, 9.6 more seedlings/m² emerged in the closed than the opened exclosures, suggesting that small vertebrate granivores reduce seedling emergence in prairie restoration. Granivores influenced the composition of the emerging community but did not preferentially consume large‐seeded species. In the seed‐tracking experiment, we found that greater than 70% of seeds were lost within 30 days of sowing, that seed recovery and viability both decreased with time in soil, and that seed fates differed between species. Collectively, our results indicate that small vertebrate granivores are an important cause of seed loss in prairie restoration, but unidentified belowground (e.g. fungal decomposition, invertebrate predation) and environmental (wind, rain) factors account for a greater proportion of total seed loss. Until these causes of seed loss are better understood, high seed costs will persist and continue to impede prairie restoration.     

Spatial organization of a desert rodent community: food addition and species removal

Summary From 1977 through 1983 we conducted experiments on a desert rodent community where supplemental seeds were added or certain rodent species and ants were removed from 0.25-ha fenced plots in a Chihuahuan Desert site in southeastern Arizona, USA. In this paper we examine the patterns of microhabitat use relative to vegetative cover by 11 rodent species. The results show that: i) removal of the largest seed-eating species, Dipodomys spectabilis, produced the most pervasive and dramatic shifts in microhabitat use by the remaining rodent species; ii) adding seeds or removing ants had little effect on the spatial use of microhabitats by rodents in this community; and iii) non-granivores were just as likely as granivores to shift microhabitat use when other granivores were removed. We believe these results indicate that both food and foraging microsites are limited but the relegation of subdominant species to lesspreferred microhabitats by the large Dipodomys spectabilis is the major factor underlying the spatial organization of this community. Results also demonstrate that strong interactions among species increase the probability that pathways of indirect interactions through intermediary species are important; these complex linkages may include species that overlap little in food preferences.     

Teasing Apart the Effects of Seed Size and Energy Content on Rodent Scatter-Hoarding Behavior

Scatter-hoarding rodents are known to play a crucial role in the seed dispersal of many plant species. Numerous studies have indicated that both seed size and the energy content of seeds can affect rodent foraging behavior. However, seed size is usually associated with energy content per seed, making it difficult to isolate how seed size and energy affect rodent foraging preferences. This study used 99 treatments of artificial seeds (11 seed sizes×9 levels of energy content) to tease apart the effect of seed size and energy content on rodent seed-caching behavior. Both seed traits showed significant effects, but their details depended on the stage of the rodent foraging process. Seeds with higher energy content were harvested more rapidly while seed size only had a modest effect on harvest rate. However, after harvesting, seed size showed a much stronger effect on rodent foraging behavior. Rodents’ choice of which seeds to remove and cache, as well as seed dispersal distance, seemed to reflect an optimal seed size. Our findings could be adapted in future studies to gain a better understanding of scatter-hoarding rodent foraging behavior, and the co-evolutionary dynamics between plant seed production and seed dispersers.     

Driven to distraction: detecting the hidden costs of flea parasitism through foraging behaviour in gerbils

Gerbilline rodents such as Allenby's gerbils (Gerbillus andersoni allenbyi), when parasitized by fleas such as Synosternus cleopatrae pyramidis, devote long hours of grooming to remove the ectoparasites. Yet no detrimental energetic or immunological effects of the ectoparasites have been found in adult Allenby's gerbil. Why should gerbils go to such trouble? We tested for the various ways that fleas can negatively affect gerbils by manipulating flea infestation on gerbils and the presence of a fox. We demonstrate that gerbils responded to fleas by leaving resource patches at higher giving-up densities. Furthermore, they stayed in those resource patches less time and left them at higher quitting harvest rates so long as a fox was also present. When flea-ridden, gerbils also abandoned using vigilance to manage risk and relied mainly on time allocation. Thus, having fleas imposed a foraging cost similar in nature to that arising from the risk of predation from foxes and may be even larger in magnitude. More than that, the presence of fleas acted as a magnifier of foraging costs, especially those arising from the risk of predation. The fleas reduced the gerbils' foraging aptitude and altered how they went about managing risk of predation. We hypothesize that fleas reduce the attention that gerbils otherwise have for foraging and predator detection. We suggest that this is the major cost of ectoparasitism.     

Native granivores reduce the establishment of native grasses but not invasive <Emphasis Type="Italic">Bromus tectorum</Emphasis>

Seed predation can structure plant communities by imposing strong population controls on some species but not others. In this context, studies from various ecosystems report that native granivores selectively forage for seeds from native species over seeds from exotic invaders, which could disproportionately favor the establishment of invaders and facilitate their dominance in communities. However, few studies have connected selective foraging for native seeds to differential patterns of establishment among native and invasive species. Thus, the extent to which preferential foraging for native seeds favors the establishment of invasive plants is unclear. Here, we used experimental seed additions and exclosure treatments at five field sites distributed across?≈?80,000 km² of the Great Basin Desert, USA to compare the effects of rodent foraging on the establishment of less-preferred cheatgrass (Bromus tectorum—an annual species native to Eurasia that is exotic and highly invasive across the Great Basin) and four species of more-preferred native grasses that commonly co-occur with cheatgrass. Rodent foraging reduced the establishment of each native species by at least 80% but had no effect on the establishment of cheatgrass, and this finding was consistent across study sites. Our results suggest that selective foraging for native species may favor the establishment of cheatgrass over native grasses, potentially exacerbating one of the most extensive plant invasions in North America.     

Effects of plant competition,seed predation,and nutrient limitation on seedling survivorship of spotted knapweed (<Emphasis Type="Italic">Centaurea stoebe</Emphasis>)

We measured seed germination and seedling survivorship of spotted knapweed, Centaurea stoebe, in a series of laboratory and field experiments to evaluate the efficacy of seed limitation as a management focus. This work was initiated 6 years after introduction of several biological control agents. The soil seed bank of the site used in this study contained a mean density of 5,848 seeds/m² (ranging from 0 to 16,364 seeds/m²), and 92% of the seeds isolated from soils were shriveled, discolored, and/or partially decayed. Additionally, none of the intact seeds germinated, suggesting that the viable seed bank at our field study site has been exhausted. Centaurea stoebe seeds were planted into pots under a range of soil nitrogen (N) availability, with half of the pots containing a single density of previously established seedlings of a native cool-season grass, slender wheatgrass (Elymus trachycaulus). A watering regime mimicking local precipitation was applied. Spotted knapweed exhibited large biomass responses to N addition, but the presence of grasses suppressed the ability to exploit this N. Surprisingly, low soil N conditions improved knapweed survivorship in the presence of grasses. Nevertheless, recruitment and biomass were still far below the levels reached in the absence of competition. To evaluate the effect of density on successful recruitment, Centaurea stoebe seed was introduced into a meadow at three densities matching reduced levels of seed production under the constraints of seed predators. These densities were sown with or without a seed mixture of native species, into an existing plant community lacking C. stoebe, and seedling recruitment was recorded over 2.5 years. Across all plots and densities sown (568–2,272 seeds m⁻² year⁻¹), seedling recruitment was less than 1%. The invasion potential of spotted knapweed was greatly diminished when realistic levels of plant competition and biological control limit seed production. We therefore conclude that a combination of seed limitation and shortage of ‘safe sites’ within undisturbed vegetation can limit densities of C. stoebe.