Dispersal and establishment filters influence the assembly of restored prairie plant communities

Community assembly filters, which in theory determine the suite of species that arrive at and establish in a community, have tremendous conceptual relevance to restoration. However, the concept has remained largely theoretical, with a paucity of empirical tests. As such, the applicability of assembly filters theory to ecological restoration remains incompletely known. We tested the relative strengths of dispersal and establishment filters by comparing the plant species composition, measured by species' presence/absence, in 29 restored prairies with the seed mixes used to restore each prairie. We found that both establishment and dispersal filters limited prairie similarity to the seed mix. Sown species responded differentially to filters, with a few species limited only by dispersal (seed density), many others limited only by establishment conditions (i.e. organic matter and sand content of soils, land use history, and fire frequency), and others limited by both dispersal and establishment filters. A few species, typically those sown most often, were not restricted by dispersal or establishment filters, likely because they were sown in high enough densities and all sites had suitable environmental conditions. Finally, one group of species established poorly, but we could not attribute this to either dispersal or establishment filters. This information can help land managers select species likely to establish in restorations when sown at sufficient densities. These results illustrate that dispersal and establishment filters limit the establishment of species in restored communities and these filters are species‐dependent. Identifying the most limiting filter(s) for species will inform strategies to increase their establishment success.     

Independent effects of arbuscular mycorrhiza and earthworms on plant diversity and newcomer plant establishment

Questions: How do arbuscular mycorrhiza and earthworms affect the structure and diversity of a ruderal plant community? Is the establishment success of newcomer plants enhanced by these soil organisms and their interactions? Methods: We grew a native ruderal plant community composed of different functional groups (grasses, legumes and forbs) in the presence and absence of arbuscular mycorrhizal fungi (AMF) and endogeic earthworms in mesocosms. We introduced seeds of five, mainly exotic, plant species from the same functional groups after a disturbance simulating mowing. The effects of the soil organisms on the native ruderal plant community and seedling establishment of the newcomer plants were assessed. Results: After disturbance, the total above‐ground regrowth of the native plant community was not affected by the soil organisms. However, AMF increased plant diversity and shoot biomass of forbs, but decreased shoot biomass of grasses of the native plant community. Earthworms led to a reduction in total root biomass. Establishment of the introduced newcomer plants increased in the presence of AMF and earthworms. Especially, seedling establishment of the introduced non‐native legume Lupinus polyphyllus and the native forb Plantago lanceolata was promoted in the presence of AMF and earthworms, respectively. The endogeic earthworms gained more weight in the presence of AMF and led to increased extraradical AMF hyphal length in soil. However, earthworms did not seem to modify the effect of AMF on the plant community. Conclusion: The present study shows the importance of mutualistic soil organisms in mediating the establishment success of newcomer plants in a native plant community. Mutualistic soil organisms lead to changes in the structure and diversity of the native plant community and might promote newcomer plants, including exotic species.     

Effects of ungulates and prairie dogs on seed banks and vegetation in a North American mixed-grass prairie

The relationship between vegetation cover and soil seed banks was studied in five different ungulate herbivore-prairie dog treatment combinations at three northern mixed-grass prairie sites in Badlands National Park, South Dakota. There were distinct differences in both the seed bank composition and the aboveground vegetation between the off-prairie dog colony treatments and the on-colony treatments. The three on-colony treatments were similar to each other at all three sites with vegetation dominated by the forbs Dyssodia papposa, Hedeoma spp., Sphaeralcea coccinea, Conyza canadensis, and Plantago patagonica and seed banks dominated by the forbs Verbena bracteata and Dyssodia papposa. The two off-colony treatments were also similar to each other at all three sites. Vegetation at these sites was dominated by the grasses Pascopyrum smithii, Bromus tectorum and Bouteloua gracilis and the seed banks were dominated by several grasses including Bromus tectorum, Monroa squarrosa, Panicum capillare, Sporobolus cryptandra and Stipa viridula. A total of 146 seedlings representing 21 species germinated and emerged from off-colony treatments while 3069 seedlings comprising 33 species germinated from on-colony treatments. Fifteen of the forty species found in soil seed banks were not present in the vegetation, and 57 of the 82 species represented in the vegetation were not found in the seed banks. Few dominant species typical of mixed-grass prairie vegetation germinated and emerged from seed banks collected from prairie dog colony treatments suggesting that removal of prairie dogs will not result in the rapid reestablishment of representative mixed-grass prairie unless steps are taken to restore the soil seed bank.     

Invasion by Solidago species has limited impacts on soil seed bank communities

Increasing attention in invasion biology is being paid to measuring and understanding the impacts of invasive species. For plant invasions, however, the impact of invasion on soil seed bank communities has been under-studied. At six sites in southern Germany, we investigated whether areas invaded by Solidago gigantea and Solidago canadensis experienced a reduction in seed bank species richness, size and diversity, and a change in species composition compared to adjacent uninvaded areas. We found no overall effect of invasion on seed bank size, or on species richness and diversity. Seed bank size significantly decreased from 0–5 cm to 5–10 cm depth in both invaded and uninvaded areas. A significant amount of variation in species composition was explained by invasion, but it was only one-tenth of that explained solely by site effects. Our study suggests that invasion by Solidago species may not have the same impacts on the soil seed banks of native species as other invasive perennial forbs that have so far been studied.     

Native-species seed additions do not shift restored prairie plant communities from exotic to native states

Exotic- and native-dominated communities can exist as alternate states in landscapes, but whether exotic-dominated states are persistent in the face of propagule pressure from native species is not well known. Here, we asked whether adding native seeds to low diversity, exotic-dominated patches would shift them to a more diverse, native state by using a long-term experiment with tallgrass prairie species in Iowa, USA. Previous work established that community assembly history led to alternate exotic or native states of perennial species. We added native seeds to plots in the spring after removing aboveground biomass with fire. We found that an experimental seed addition did not cause a shift from exotic to native states. Plots seeded eight years earlier in spring and without a priority effect continued to have the highest abundance and diversity of native species and lowest proportion of exotics. Our results suggest that exotic-dominated states in restorations can persist in the face of native species propagule pressure. Thus, assembly history can play a strong role in generating and maintaining alternate states over long time frames that are relevant to restoration. New restoration projects in exotic-dominated landscapes should maximize effort toward establishing native species during initial stages of restoration.     

Assessing the impact of plant invasions on soil seed bank communities: use of univariate and multivariate statistical approaches

Questions: Are soil seed banks affected by invasions of alien plants? How can we rigorously assess alterations in seed bank communities associated with invasive species and account for the high spatial variability of seed bank data? How do multivariate approaches compare with more traditional approaches based on analysis of variance? Location: Three riparian sites, Ireland. Methods: A protocol based on a combination of multivariate techniques was used to characterize soil seed bank communities associated with the herbaceous invasive species Heracleum mantegazzianum in May and October. Permutational multivariate analysis of variance (PERMANOVA) was used to test the effects of the factors “invasion”, “site”, “plot” and “depth” on the soil seed bank, while multivariate analysis of dispersion (PERMDISP) provided a measure of the variability of seed bank data at different spatial scales. Similarity percentages analysis (SIMPER) was used to identify the species that contributed most to the differences between invaded and uninvaded communities. A comparison between the results of PERMANOVA and ANOVA analyses was also made. Results: The composition of seed bank communities invaded by H. mantegazzianum differed significantly from that of uninvaded seed banks. Invaded seed banks were less diverse and had reduced abundance, and were dominated by only a few species, such as Urtica dioica and Juncus effusus. Such patterns were recorded at each of three depth categories, indicating that invasive plants can affect both the transient and the more persistent component of the soil seed bank. Seed bank variability was significantly higher within uninvaded areas, supporting the notion that invasions tend to lead to more homogeneous communities. Conclusion: The analytical protocol used in this study was effective in quantifying the effect of plant invasions, at different spatial scales, providing a statistically robust analysis of alterations in soil seed bank communities. Compared to ANOVA, this protocol provided more biological information and was more appropriate for analysis of the data. This approach is therefore recommended in soil seed bank and invasion ecological studies.     

Responses of plant and bird communities to prescribed burning in tallgrass prairies

Historic losses and fragmentation of tallgrass prairie habitat to agriculture and urban development have led to declines in diversity and abundance of plants and birds associated with such habitat. Prescribed burning is a management strategy that has potential for restoring and rejuvenating prairies in fragmented landscapes, and through such restoration, might create habitat for birds dependent upon prairies. To provide improved data for management decision-making regarding the use of prescribed fire in tallgrass prairies, we compared responses of plant and bird communities on five burned and five unburned tallgrass prairie fragments at the DeSoto National Wildlife Refuge, Iowa, USA, from 1995 to 1997. Overall species richness and diversity were unaffected by burning, but individual species of plants and birds were affected by year-treatment interactions, including northern bobwhite (Colinus virginianus) and ring-necked pheasant (Phasianus colchicus), which showed time-delayed increases in density on burned sites. Analyses of species/area relationships indicated that, collectively, many small sites did make significant contributions to plant biodiversity at landscape levels, supporting the overall conservation value of prairie fragments. In contrast, most birds species were present on larger sites. Thus, higher biodiversity in bird communities which contain area-sensitive species might require larger sites able to support larger, more stable populations, greater habitat heterogeneity, and greater opportunity for niche separation.     

The effects of irrigation effluent on a Western Canadian prairie river

Field studies were conducted during the peak irrigation period(July–August) at the mouth, 50 m upstream and 50 m downstreamoffour irrigation return flows draining a 50 km stretch of theOldmanRiver in the short grass prairie zone of southern Alberta.Watersamples were analyzed weekly. Physico-chemical variableschangesonly slightly at each site but salinity and conductivity weresignificantly higher at the downstream site (Expanse Coulee)andall sites showed an increase over 1978 data. Conductivityalong theleft bank of the river was measured at 50 m intervalsdownstream ofExpanse Coulee until, after 1.5 km, it declined to theupstreamlevel which in the laboratory was shown to require a 660%dilutionof the effluent. In field experiments conducted at ExpanseCouleeit was shown that macroinvertebrate biomass was significantlylowerdownstream of the mouth and significantly higher upstream.Chlorophyll-a was not significantly different betweenthemouth and upstream with both significantly higher thandownstreamwhere it was not detectable. Inorganic carbon assimilation byepilithic algae and heterotrophic potential of the bacteriawereboth significantly higher at the mouth and upstream andsignificantly lower downstream.Organic matter in the sediment was not significantly differentupstream or downstream of the mouth but there was a higherpercentage of large particles (125–250 m) at the mouthanddownstream of EC a higher percentage of medium(60–125 m) andsmall (0–60 m) particles were found 50 m upstream. It isconcluded that at present differences in invertebrate biomass,chlorophyll-a, inorganic carbon assimilation byepilithicalgae and heterotrophic potential of epilithic bacteria atExpanseCoulee are not directly attributable to increase in salinitybutare more probably the result of increased suspended solids andsaltation.     

Influence of the invasive shrub Nicotiana glauca Graham on the plant seed bank in various locations in Taif region,western of Saudi Arabia

Invasive species have been considered as one of the most serious threats to the biodiversity of various ecosystems, particularly in arid regions. The present study aimed to assess the influence of the invasive shrub Nicotiana glauca on the biodiversity of different habitats in Taif region, Saudi Arabia as well as to determine the highest habitat with seed bank of N. glauca. Soil samples were collected from three locations (Alwaht, Ash-shafa, and Ar Ruddaf), invaded with N. glauca, and analyzed for the soil seed bank. A soil seed experiment was designed in a greenhouse, whereby emerged plant seedlings were left to grow for three months and identified as well as the species density and biodiversity were assessed under and outside the canopy of N. glauca. Also, the floristic composition, life forms, and chorotype spectra of the plant species of the seed bank were analyzed. A total of 42 species, belonging to 23 families, were recorded in the soil seed bank. Asteraceae, Poaceae, and Cyperaceae were the major families (42.9%). The life form spectra of the recorded species were dominated by Therophytes (59.5%). Chorotype spectra analysis revealed that Mediterranean, Saharo-Arabian, and Irano-Turanian were the most represented elements. The species richness and evenness were higher outside the canopy, which indicates a negative effect of the invasive shrub N. glauca on the plant biodiversity in the study area, particularly in Ar Ruddaf location. This could be attributed to the competition or allelopathic effect of N. glauca. In contrast, the density of N. glauca seeds was higher under the canopy compared to outside. The soil nutrients and moisture under the canopy were higher than outside canopy. The present study provides a deeper understanding of the most susceptible habitats or communities to the invasion by N. glauca and thereby open the challenge toward control of this noxious plant and vegetation restoration.     

Incomplete recovery of plant diversity in restored prairie wetlands on agricultural landscapes

Restoration efforts are being implemented globally to mitigate the degradation and loss of wetland habitat; however, the rate and success of wetland vegetation recovery post‐restoration is highly variable across wetland classes and geographies. Here, we measured the recovery of plant diversity along a chronosequence of restored temporary and seasonal prairie wetlands ranging from 0 to 23 years since restoration, including drained and natural wetlands embedded in agricultural and natural reserve landscapes in central Alberta, Canada. We assessed plant diversity using the following structural indicators: percent cover of hydrophytes, native and non‐native species, species richness, and community composition. Our findings indicate that plant diversity recovered to resemble reference wetlands in agricultural landscapes within 3–5 years of restoration; however, restored wetlands maintained significantly lower species richness and a distinct community composition compared to reference wetlands located within natural reserves. Early establishment of non‐native species during recovery, dispersal limitation, and depauperated native seed bank were probable barriers to complete recovery. Determining the success of vegetation recovery provides important knowledge that can be used to improve restoration strategies, especially considering projected future changes in land use and climate.     

Interannual variation in precipitation and other planting conditions impacts seedling establishment in sown plant communities

Ecological restoration can reverse biodiversity loss worldwide, yet restoration goals and outcomes vary widely, which limits this potential. Divergent restoration outcomes may stem from variation in conditions at the outset of restoration, but empirical evidence is lacking and typically confounded with site differences. Additionally, precipitation is usually cited as the source of this variation, although a wide range of conditions can vary annually. We tested for effects of planting year on seedling establishment by installing identical restorations in three different years. Within those years, we manipulated rainfall with rain‐out shelters to disentangle the effects of precipitation from other annually variable conditions. Additionally, we tested whether increasing seed mix richness buffers against adverse planting conditions. For the first growing season after planting, we followed emergence and survival of sown prairie species and nonsown weed species to determine how planting year conditions influence an establishing plant community, if at all. We found that seedling establishment differed across planting years and precipitation treatments, and that varying emergence patterns by species led to differences in the composition of the first‐year community. We also found significant variation in sown species establishment across years when precipitation was held constant, illustrating the previously overlooked role of nonprecipitation drivers on planting year effects. Higher seed mix richness did not consistently improve establishment of sown species under different planting conditions. This research provides important experimental evidence for effects of interannual variation in planting conditions on first‐year establishment. Future work will examine how these initial changes affect longer‐term assembly dynamics.     

Effect of grazing removal on aboveground vegetation and soil seed bank composition in sub-alpine grasslands of northern Iran

Background: Knowledge about vegetation and soil seed bank composition and the processes that contribute to vegetation recovery after the removal of heavy grazing is lacking in sub-alpine ecosystems.

Aims: In order to assess the effects of large herbivores on above-ground vegetation (AGV) and soil seed bank (SSB) characteristics, intensively sheep-grazed areas were compared to adjacent areas where grazing had been removed 10 years previously in a sub-alpine grassland of northern Iran.

Methods: A total of 40 4-m² (2 m × 2 m) plots were established in each treatment, and soil samples were collected from each plot within a depth of 10 cm. Plant species composition was determined for each plot during the flowering stage of herbaceous species in June 2011. The seedling emergence method was used to estimate SSB composition.

Results: A total of 45 species (23 annuals and 22 perennials) emerged from the soil samples of the grazed area, while the number of species emerged from the soil samples of the previously grazing area was 76 (37 annuals and 39 perennials). The removal of grazing led to a significant increase in species richness and seed density in the SSB. Species turnover of AGV was higher, and that of the SSB was similar for grazed areas compared with areas that were formerly grazed. Detrended correspondence analysis ordination of AGV composition showed a clearly separate structure between grazed plots and plots where grazing has been removed, while the segregation was less clear for SSB composition.

Conclusions: We concluded that restoration of locally degraded sites cannot rely on the SSB when grazing is stopped simultaneously and unvegetated gaps are colonised by vegetative growth rather than by seed.     

The effects of fire frequency and grazing on tallgrass prairie productivity and plant composition are mediated through bud bank demography

Periodic fire, grazing, and a variable climate are considered the most important drivers of tallgrass prairie ecosystems, having large impacts on the component species and on ecosystem structure and function. We used long-term experiments at Konza Prairie Biological Station to explore the underlying demographic mechanisms responsible for tallgrass prairie responses to two key ecological drivers: fire and grazing. Our data indicate that belowground bud banks (populations of meristems associated with rhizomes or other perennating organs) mediate tallgrass prairie plant response. Fire and grazing altered rates of belowground bud natality, tiller emergence from the bud bank, and both short-term (fire cycle) and long-term (>15 year) changes in bud bank density. Annual burning increased grass bud banks by 25% and decreased forb bud banks by 125% compared to burning every 4 years. Grazing increased the rate of emergence from the grass bud bank resulting in increased grass stem densities while decreasing grass bud banks compared to ungrazed prairie. By contrast, grazing increased both bud and stem density of forbs in annually burned prairie but grazing had no effect on forb bud or stem density in the 4-year burn frequency treatment. Lastly, the size of the reserve grass bud bank is an excellent predictor of long-term ANPP in tallgrass prairie and also of short-term interannual variation in ANPP associated with fire cycles, supporting our hypothesis that ANPP is strongly regulated by belowground demographic processes. Meristem limitation due to management practices such as different fire frequencies or grazing regimes may constrain tallgrass prairie responses to interannual changes in resource availability. An important consequence is that grasslands with a large bud bank may be the most responsive to future climatic change or other global change phenomena such as nutrient enrichment, and may be most resistant to exotic species invasions.     

Fungal endophyte infection increases tall fescue's survival,growth, and flowering in a reconstructed prairie

Grasslands in North America are increasingly threatened by land conversion and ecological degradation, prompting restoration efforts to increase native plant species diversity and improve wildlife habitat. A major challenge is the removal and management of nonnative invasive species such as tall fescue (Schedonorus arundinaceus), which has a symbiotic association with a fungal endophyte (Epichloë coenophiala) that modifies its ecological interactions. Using transplanted clumps of the cultivar Kentucky‐31, we tested the effects of endophyte infection on tall fescue's survival and performance (tiller production, flowering, and basal area) for 5 years in a central Kentucky reconstructed prairie. We predicted that endophyte infected (E+) clumps would have increased performance compared to endophyte‐free (E?) clumps. Overall, E+ clumps had greater survival, tiller production, flowering tiller production, and basal area, but not reproductive effort (proportion of tillers flowering) as compared to E? clumps. However, survival and trends in tiller number and basal area over the 5‐year period suggested experimental tall fescue populations were in decline in the reconstructed prairie, although the E? population declined more rapidly. Our study provides evidence that endophyte infection improved tall fescue's growth and survival in a postreconstruction plant community, at least in the early years following reconstruction, and may increase the invasive potential of this nonnative species in prairie restorations.     

科尔沁沙地植被恢复演替进程中群落土壤种子库研究

研究了科尔沁沙地植被恢复演替进程中各群落土壤种子库种类组成、密度以及物种多样性等特征。主要结论是:①科尔沁沙地植被退化后的恢复演替是一种次生的中途恢复演替。演替各阶段土壤种子库密度为流动沙丘<半流动沙丘<固定沙丘,从流动到半流动沙丘阶段,土壤种子库密度平均增加了709%;从半流动到固定沙丘阶段,土壤种子库密度平均增加了393%。从流动到半流动沙丘阶段是土壤种子库密度的快速增长期。②植被恢复演替过程中,土壤种子库组成均以1年生草本植物为主(优势度为60.40%~91.83%),到演替中后期阶段,多年生草本植物的种类有所增加,但其所占比例仍很小。③演替各阶段群落种子库物种多样性指数分别为0.6616、0.7736、0.7281、1.0939、1.0648和0.9682,可见种子库物种多样性最高的群落并非是演替历史最大的群落。④恢复演替系列各阶段土壤种子库间的相似性系数都较大,在0.368~1.000范围之内,任一群落总是与其下一阶段最邻近的群落具有最高的相似性系数。     

Effects of mycorrhizae on plant growth and dynamics in experimental tall grass prairie microcosms

Experimental microcosms (40 X 52 X 32 cm) containing an assemblage of eight tallgrass prairie grass and forb species in native prairie soil were maintained under mycorrhizal (untreated control) or mycorrhizal-suppressed (fungicide-treated) conditions to examine plant growth, demographic, and community responses to mycorrhizal symbiosis. The fungicide benomyl successfully reduced mycorrhizal root colonization in the fungicide-treated microcosms to only 6.4% (an 83% reduction relative to mycorrhizal controls). Suppression of mycorrhizas resulted in a 31% reduction in total net aboveground plant production and changes in the relative production of C4 and C3 plants. The C4 tallgrasses Andropogon gerardi and Sorghastrum nutans produced less plant biomass in the fungicide-treated microcosms, and had a greater ratio of reproductive to vegetative biomass. Cool-season C3 grasses, Koeleria pyramidata and Poa pratensis accumulated more biomass and were a significantly greater proportion of total community biomass in mycorrhizal-suppressed microcosms. Forbs showed variable responses to mycorrhizal suppression. The two legumes Amorpha canescens and Dalea purpurea had significantly lower survivorship in the fungicide-treated microcosms, relative to the controls. The results confirm the high mycorrhizal dependency and growth responsiveness of dominant prairie grasses, and indicate that differential growth and demographic responses to mycorrhizal colonization among species may significantly affect plant productivity and species relative abundances in tallgrass prairie.     

Temporal effects of agri-environment schemes on ditch bank plant species

Many of the agri-environment schemes (AES) implemented in the Western Peat District of the Netherlands have as their objective the conservation of the diversity of ditch bank plants. We investigated the effects of AES on ditch bank species in this area, using a dataset collected by 377 farmers who managed and monitored ditch banks during a 10-year period. We found that species richness has increased minimally over the last 10 years in ditch banks. Yet, we found no differences in increases in time between ditch banks with and without AES. In both ditch bank types plant species composition changed to species with higher nitrogen tolerance. Furthermore, species that disperse over long distances by water increased, whereas species with no capacity to disperse over long distances declined in both ditch bank types. This indicates that changes in vegetation composition in ditch banks are affected by other factors than AES.     

Legacies of afforestation on soil nematode community composition,structure, and diversity in a northern Canadian prairie

Plant and Soil - We examined how legacies of afforestation affect soil food webs using the composition, structure, and diversity of soil nematode communities along a prairie restoration...     

Species richness and evenness respond in a different manner to propagule density in developing prairie microcosm communities

Diversity has two basic components: richness, or number of species in a given area, and evenness, or how relative abundance or biomass is distributed among species. Previously, we found that richness and evenness can be negatively related across plant communities and that evenness can account for more variation in Shannon’s diversity index (H′) than richness, which suggests that relationships among diversity components can be complex. Non-positive relationships between evenness and richness could arise due to the effects of migration rate or local species interactions, and relationships could vary depending on how these two processes structure local communities. Here we test whether diversity components are equally or differentially affected over time by changes in seed density (and associated effects on established plant density and competition) in greenhouse communities during the very early stages of community establishment. In our greenhouse experiment, we seeded prairie microcosms filled with bare field soil at three densities with draws from a mix of 22 grass and forb species to test if increased competition intensity or seedling density would affect the relationships among diversity components during early community establishment. Increased seed density treatments caused diversity components to respond in a different manner and to have different relationships with time. Richness increased linearly with seed density early in the experiment when seedling emergence was high, but was unrelated to density later in the experiment. Evenness decreased log-linearly with seed densities on all sampling dates due to a greater dominance by Rudbeckia hirta with higher densities. Early in the experiment, diversity indices weakly reflected differences in richness, but later, after the competitive effects of Rudbeckia hirta became more intense, diversity indices more strongly reflected differences in evenness. This suggests that species evenness and diversity indices do not always positively covary with richness. Based on these results, we suggest that evenness and richness can be influenced by different processes, with richness being more influenced by the number of emerging seedlings and evenness more by species interactions like competition. These results suggest that both diversity components should be measured in plant diversity studies whenever it is possible.