Linking individual response to biotic interactions with community structure: a trait-based framework

1.  Due to species-specificity of the outcomes of biotic interactions, it is difficult to generalize from observed biotic interactions at the individual plant level to the effect of those interactions at the community level. To evaluate the importance of biotic interactions in shaping plant communities, it is necessary to understand how the outcomes of the complex interactions observed at the individual level can influence community structure.
2.  Here, we propose a trait-based framework that identifies and organises mechanisms affecting community structure (here described with relative abundances of plant functional traits – i.e. the distribution of trait values at the community level). We applied our approach to a single leaf trait, specific leaf area (SLA), to link individual responses to plant interactions with community structure (SLA distribution observed at the community level) and to test whether biotic interactions can predict the functional composition of subalpine grasslands. We evaluated the generality of our model through a cross-validation with a set of eight subalpine grasslands independent from the four fields used to build the model.
3.  We found that competition and facilitation were able to explain the functional composition of subalpine grasslands, and the relevant fitness components (survival or growth) explaining this link changed depending on the limiting resources. When soil water availability was limiting, positive plant-plant interactions acting on survival were able to explain community structure. In contrast, when no water limitation was observed competition acting on individual growth was the main driver of community structure.
4.  Our framework enables evaluation of the consequences of biotic interactions observed at individual level on community structure, thereby indicating when and where different types of plant-plant interactions are important.     

Little fox on the prairie: genetic structure and diversity throughout the distribution of a grassland carnivore in the United States

The Great Plains region is fragmented by natural and anthropogenic sources, yet the interaction between habitat fragmentation and genetic structure in this region has received limited study. Swift fox (Vulpes velox) are closely associated with short and mixed grass ecosystems, providing an opportunity to study patterns of gene flow, diversity and genetic structure in this area. We collected 589 samples throughout the species’ distribution in the United States and analyzed these samples using 15 microsatellite loci and a 250 base pair sequence of the mitochondrial DNA control region. We detected three levels of spatial genetic structure using microsatellite markers and identified six mitochondrial haplotypes, five of which showed spatial clustering. Differentiation between groups was significant while genetic diversity within groups was generally high. Anthropogenic influences, particularly agriculture, appear to reducing gene flow, especially in the central portion of the species’ range. Conservation measures should be taken to remediate these impacts and to maintain future gene flow in light of expected agricultural expansion in the Great Plains. Potential evolutionary significant units are identified, although further investigation using ecological indicators and adaptive loci is recommended to characterize the adaptive distinctiveness of swift fox populations.     

Black-tailed prairie dogs and the structure of avian communities on the shortgrass plains

We tested the hypothesis that black-tailed prairie dogs (Cynomys ludovicianus) influence avian community structure on the shortgrass prairie. We surveyed 36 prairie dog towns and 36 paired sites without prairie dogs during summer and fall of 1997, 1998, and 1999 in the Oklahoma Panhandle. Our surveys totaled 9,040 individual observations for 73 avian species. Significantly distinct avian communities were present on prairie dog towns when compared to sites within four different macrohabitats of the surrounding landscape: open rangeland, scrub/sandsage (Artemisia filifolia) habitats, Conservation Reserve Program (CRP) plots, and fallow crop fields. Relative densities of all bird species combined was higher on prairie dog towns versus paired sites in summer and fall. Mean species richness of birds was significantly higher on prairie dog towns than paired sites during summer, but there were no significant differences in fall. Open rangeland had the highest mean species richness in fall. Assemblages of avian communities differed significantly between prairie dog towns and the four macrohabitat types during summer. Burrowing owls (Athene cunicularia), killdeer (Charadrius vociferous), horned larks (Eremophila alpestris), and meadowlarks (Sturnella spp.) were positively and significantly associated with prairie dog towns during summer, while horned larks and ferruginous hawks (Buteo regalis) were significantly associated with prairie dog towns during fall. Even in their current remnant state, black-tailed prairie dogs continue to play a significant role in the assembly of ecological communities across the Great Plains. Conservation of prairie dogs goes well beyond a single species, and is an important strategy for the preservation of the prairie ecosystem as a whole.     

Burrowing activities of kangaroo rats and patterns in plant species dominance at a shortgrass steppe-desert grassland ecotone

Abstract. Our objective was to evaluate the effects of burrowing activities by banner-tail kangaroo rats (Dipodomys spectabilis Merriam) on plant community structure and species dominance for two patch types at the ecotone between shortgrass steppe and desert grassland in New Mexico, USA. 10 mounds produced by kangaroo rats were selected in patches dominated by Bouteloua gracilis (the dominant in shortgrass steppe communities) and 10 mounds were selected in patches dominated by B. eriopoda (the dominant in Chihuahuan desert grasslands). Plant cover and density by species were sampled from three locations associated with each mound: the mound proper, the edge of the mound in the transition area, and the off-mound vegetation. Similar cover of B. eriopoda for the edges of mounds in both patch types indicates the ability of this species to respond to animal disturbances regardless of the amount of cover in the surrounding undisturbed vegetation. By contrast, cover of B. gracilis was low for all mounds and mound edges in patches dominated by this species. Much higher cover of B. eriopoda on mound edges compared to the undisturbed vegetation in B. gracilis-dominated patches indicates that kangaroo rats have important positive effects on this species. Lower cover of perennial grasses and higher cover of forbs, shrubs, and succulents on the edges of mounds in B. eriopoda-dominated patches compared to patches dominated by B. gracilis indicate the importance of surrounding vegetation to plant responses on disturbed areas. Our results show that kangaroo rats have important effects on both species dominance and composition for different patch types, and may provide a mechanism for small-scale dominance patterns at an ecotone; thus providing further support for their role as keystone species in desert grasslands.     

Positive interactions between desert granivores: localized facilitation of harvester ants by kangaroo rats

Facilitation, when one species enhances the environment or performance of another species, can be highly localized in space. While facilitation in plant communities has been intensely studied, the role of facilitation in shaping animal communities is less well understood. In the Chihuahuan Desert, both kangaroo rats and harvester ants depend on the abundant seeds of annual plants. Kangaroo rats, however, are hypothesized to facilitate harvester ants through soil disturbance and selective seed predation rather than competing with them. I used a spatially explicit approach to examine whether a positive or negative interaction exists between banner-tailed kangaroo rat (Dipodomys spectabilis) mounds and rough harvester ant (Pogonomyrmex rugosus) colonies. The presence of a scale-dependent interaction between mounds and colonies was tested by comparing fitted spatial point process models with and without interspecific effects. Also, the effect of proximity to a mound on colony mortality and spatial patterns of surviving colonies was examined. The spatial pattern of kangaroo rat mounds and harvester ant colonies was consistent with a positive interspecific interaction at small scales (<10 m). Mortality risk of vulnerable, recently founded harvester ant colonies was lower when located close to a kangaroo rat mound and proximity to a mound partly predicted the spatial pattern of surviving colonies. My findings support localized facilitation of harvester ants by kangaroo rats, likely mediated through ecosystem engineering and foraging effects on plant cover and composition. The scale-dependent effect of kangaroo rats on abiotic and biotic factors appears to result in greater founding and survivorship of young colonies near mounds. These results suggest that soil disturbance and foraging by rodents can have subtle impacts on the distribution and demography of other species.     

Nitrogen-fixing bacteria, arbuscular mycorrhizal fungi, and the productivity and structure of prairie grassland communities

Due to their complementary roles in meeting plant nutritional needs, arbuscular mycorrhizal fungi (AMF) and nitrogen-fixing bacteria (N₂-fixers) may have synergistic effects on plant communities. Using greenhouse microcosms, we tested the effects of AMF, N₂-fixers (symbiotic: rhizobia, and associative: Azospirillum brasilense), and their potential interactions on the productivity, diversity, and species composition of diverse tallgrass prairie communities and on the productivity of Panicum virgatum in monoculture. Our results demonstrate the importance of AMF and N₂-fixers as drivers of plant community structure and function. In the communities, we found a positive effect of AMF on diversity and productivity, but a negative effect of N₂-fixers on productivity. Both AMF and N₂-fixers affected relative abundances of species. AMF shifted the communities from dominance by Elymus canadensis to Sorghastrum nutans, and seven other species increased in abundance with AMF, accounting for the increased diversity. N₂-fixers led to increases in Astragalus canadensis and Desmanthus illinoense, two legumes that likely benefited from the presence of the appropriate rhizobia symbionts. Sorghastrum nutans declined 44?% in the presence of N₂-fixers, with the most likely explanation being increased competition from legumes. Panicum monocultures were more productive with AMF, but showed no response to N₂-fixers, although inference was constrained by low Azospirillum treatment effectivity. We did not find interactions between AMF and N₂-fixers in communities or Panicum monocultures, indicating that short-term effects of these microbial functional groups are additive.     

Response of lizard community structure to desert grassland restoration mediated by a keystone rodent

Many grasslands in the Chihuahuan Desert have transformed to shrublands dominated by creosotebush (Larrea tridentata). Grassland restoration efforts have been directed at controlling creosotebush by applying herbicide over large spatial scales. However, we have a limited understanding of how landscape-scale restoration affects biodiversity. We examined whether restoration treatments in southern New Mexico, USA have influenced the community structure of lizards, which are sensitive to shrub encroachment. We compared lizard community structure on 21 areas treated with herbicide from 7 to 29 years ago with paired untreated areas that were dominated by shrubs and matched by geomorphology, soils, and elevation. To examine mechanisms underlying responses to restoration, we tested whether the abundance of a grassland specialist, Aspidoscelis uniparens, depended on time since treatment, treatment area and isolation, and local habitat quality. Because lizards use rodent burrows as habitat, we tested whether community structure and A. uniparens abundance depended on the abundance of the keystone rodent, Dipodomys spectabilis. Treated areas had reduced shrub cover and increased grass cover compared to untreated areas. Lizard community composition differed strongly between areas, with four species responding to treatments. Divergence in community composition between treated–untreated pairs was greatest for old treatments (≥22 years), and community composition was influenced by D. spectabilis. In particular, the abundance of A. uniparens was greatest on old treatments with a high density of D. spectabilis. Overall, our results demonstrate lizard community structure responds to grassland restoration efforts, and keystone species can shape restoration outcomes. Reestablishment of keystone species may be a critical constraint on the recovery of animal biodiversity after habitat restoration.     

Promotion of exotic weed establishment by endangered giant kangaroo rats (Dipodomys ingens) in a California grassland

Giant kangaroo rats (Dipodomys ingens) continually modify their burrow precincts by digging tunnels, clipping plants, and other activities. In the valley grasslands of the Carrizo Plain Natural Area (San Luis Obispo County, California), this chronic disturbance to soil and vegetation promoted the establishment of exotic ruderal and early successional plant species. Erodium cicutarium, Bromus madritensis ssp. rubens, and other Mediterranean annuals were found to constitute a very large proportion of the vegetation on giant kangaroo rat precincts. When vegetation on precincts was compared with the vegetation in less disturbed intermediate areas located between precincts, species richness, cover and frequency of exotic plants were significantly greater on precincts. The reverse was found for native species. In addition, exotic species encountered in this study had significantly larger seeds than did native species, suggesting that these granivorous kangaroo rats preferentially cache large weed seeds on their precincts. Since the kangaroo rats depend on exotic plants for food and the exotic plants depend upon the kangaroo rats to disturb their habitat continually, the weed-kangaroo rat relationship is mutualistic. This strong relationship may also inhibit population growth of native grassland plants which occupy disturbed habitats but have difficulty competing with exotic weeds for resources. From a conservation perspective, this mutualism presents an intractable management dilemma. Restoration of valley grasslands where endangered giant kangaroo rats occur, to conditions where native species dominate, may be impossible.     

Phenological influences on the albedo of prairie grassland and crop fields

 The albedo of land surfaces is strongly affected by transient surface conditions. For a vegetated surface, albedo can change with soil moisture, fractional canopy cover, and plant phenology as well as solar zenith angle (θ_s). In this study, the relationships between albedo and plant phenology in prairie grassland and agricultural crops were examined by removing the effect of variations in θ_s. Albedos were evaluated at a constant θ_s, which was assigned to be 20° in this study. For days with a minimum θ_s larger than 20°, a polynomial function that relates albedos to θ_s was derived and applied to estimate the albedos at 20°. After the removal of θ_s variations, the observed albedos of grassland at the Konza Prairie showed a linearly decreasing trend from spring to winter due to prairie phenology. The observed albedo of a maize field showed a clear increase from seedling to peak green stage, except when drought caused a decrease in near-infrared reflectance. The observed albedos of a winter wheat field showed an uninterrupted decrease from peak green to harvesting stage. Received: 30 May 1998 / Accepted 15 July1998     

Partitioning the effects of an ecosystem engineer: kangaroo rats control community structure via multiple pathways

1.?Ecosystem engineers impact communities by altering habitat conditions, but they can also have strong effects through consumptive, competitive and other non-engineering pathways. 2.?Engineering effects can lead to fundamentally different community dynamics than non-engineering effects, but the relative strengths of these interactions are seldom quantified. 3.?We combined structural equation modelling and exclosure experiments to partition the effects of a keystone engineer, the giant kangaroo rat (Dipodomys ingens), on plants, invertebrates and vertebrates in a semi-arid California grassland. 4.?We separated the effects of burrow creation from kangaroo rat density and found that kangaroo rats increased the diversity and abundance of other species via both engineering and non-engineering pathways. 5.?Engineering was the primary factor structuring plant and small mammal communities, whereas non-engineering effects structured invertebrate communities and increased lizard abundance. 6.?These results highlight the importance of the non-engineering effects of ecosystem engineers and shed new light on the multiple pathways by which strong-interactors shape communities.     

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.     

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...     

Ecosystem engineering varies spatially: a test of the vegetation modification paradigm for prairie dogs

Colonial, burrowing herbivores can be engineers of grassland and shrubland ecosystems worldwide. Spatial variation in landscapes suggests caution when extrapolating single‐place studies of single species, but lack of data and the need to generalize often leads to ‘model system’ thinking and application of results beyond appropriate statistical inference. Generalizations about the engineering effects of prairie dogs (Cynomys sp.) developed largely from intensive study at a single complex of black‐tailed prairie dogs C. ludovicianus in northern mixed prairie, but have been extrapolated to other ecoregions and prairie dog species in North America, and other colonial, burrowing herbivores. We tested the paradigm that prairie dogs decrease vegetation volume and the cover of grasses and tall shrubs, and increase bare ground and forb cover. We sampled vegetation on and off 279 colonies at 13 complexes of 3 prairie dog species widely distributed across 5 ecoregions in North America. The paradigm was generally supported at 7 black‐tailed prairie dog complexes in northern mixed prairie, where vegetation volume, grass cover, and tall shrub cover were lower, and bare ground and forb cover were higher, on colonies than at paired off‐colony sites. Outside the northern mixed prairie, all 3 prairie dog species consistently reduced vegetation volume, but their effects on cover of plant functional groups varied with prairie dog species and the grazing tolerance of dominant perennial grasses. White‐tailed prairie dogs C. leucurus in sagebrush steppe did not reduce shrub cover, whereas black‐tailed prairie dogs suppressed shrub cover at all complexes with tall shrubs in the surrounding habitat matrix. Black‐tailed prairie dogs in shortgrass steppe and Gunnison's prairie dogs C. gunnisoni in Colorado Plateau grassland both had relatively minor effects on grass cover, which may reflect the dominance of grazing‐tolerant shortgrasses at both complexes. Variation in modification of vegetation structure may be understood in terms of the responses of different dominant perennial grasses to intense defoliation and differences in foraging behavior among prairie dog species. Spatial variation in the engineering role of prairie dogs suggests spatial variation in their keystone role, and spatial variation in the roles of other ecosystem engineers. Thus, ecosystem engineering can have a spatial component not evident from single‐place studies.     

The effects of individual interactions and habitat preferences on spatial structure in a grassland bird community

Spatial structure and the distribution of individuals within a community might be influenced by several factors such as habitat heterogeneity and local interactions among individuals of the same and different species. We investigated the spatial distributions of eight bird species in a grassland community during the breeding season and examined whether the spatial distributions of individuals were influenced by interactions among neighboring individuals or different habitat preferences of different bird species.
In order to identify the effects of the interactions among neighboring individuals and habitat preference, we developed a randomization test in which species identifications were randomly allocated to the observed individual positions within areas with the same vegetation structure. The randomization test indicated that individuals tend to have territories near the territories of individuals of the same species or of a particular species more frequently (or less frequently) than those expected from random distributions of individuals.
Among these associations, only one case was explained by individual interactions, and 19 cases were explained by habitat preference.
The results suggest that both individual interactions and habitat preference affected the spatial distributions of individuals and possibly influence the species compositions and diversity in grassland bird communities.     

Earthworm-mycorrhiza interactions can affect the diversity, structure and functioning of establishing model grassland communities

Both earthworms and arbuscular mycorrhizal fungi (AMF) are important ecosystem engineers co-occurring in temperate grasslands. However, their combined impacts during grassland establishment are poorly understood and have never been studied. We used large mesocosms to study the effects of different functional groups of earthworms (i.e., vertically burrowing anecics vs. horizontally burrowing endogeics) and a mix of four AMF taxa on the establishment, diversity and productivity of plant communities after a simulated seed rain of 18 grassland species comprising grasses, non-leguminous forbs and legumes. Moreover, effects of earthworms and/or AMF on water infiltration and leaching of ammonium, nitrate and phosphate were determined after a simulated extreme rainfall event (40 l m(-2)). AMF colonisation of all three plant functional groups was altered by earthworms. Seedling emergence and diversity was reduced by anecic earthworms, however only when AMF were present. Plant density was decreased in AMF-free mesocosms when both anecic and endogeic earthworms were active; with AMF also anecics reduced plant density. Plant shoot and root biomass was only affected by earthworms in AMF-free mesocosms: shoot biomass increased due to the activity of either anecics or endogeics; root biomass increased only when anecics were active. Water infiltration increased when earthworms were present in the mesocosms but remained unaffected by AMF. Ammonium leaching was increased only when anecics or a mixed earthworm community was active but was unaffected by AMF; nitrate and phosphate leaching was neither affected by earthworms nor AMF. Ammonium leaching decreased with increasing plant density, nitrate leaching decreased with increasing plant diversity and density. In order to understand the underlying processes of these interactions further investigations possibly under field conditions using more diverse belowground communities are required. Nevertheless, this study demonstrates that belowground-aboveground linkages involving earthworms and AMF are important mediators of the diversity, structure and functioning of plant communities.