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Rangelands provide numerous ecosystem services, including forage for livestock grazing. Effective grazing management requires measuring forage availability, which influences the level of grazing that can be sustained while maintaining healthy ecological conditions. However, spatiotemporal variability makes such determinations of forage quantity difficult, potentially hindering optimal management. These determinations are especially difficult across large, remote areas. To address this, we developed an approach using data from a one-time sampling of vegetation throughout the Uintah and Ouray Reservation in northeast Utah. By associating these data in a random forest model with environmental and climatic covariates that vary annually, we produced yearly predictions of forage availability on a pixel-by-pixel basis for the Reservation and surroundings from 1984 to 2018. This method addresses and quantifies the spatiotemporal variability of available forage. The model confirms that forage availability is highly variable throughout the area. On average, forage availability in Reservation management units declined as much as 32% below median availability in some years and increased 33% above median availability in others. Moreover, some management units experienced large increases in favorable years but less significant declines in unfavorable years, while the opposite was true in others. In comparison to determining a single “typical” forage availability of management units, recognizing this inherent variability and how it differs between units provides a fuller picture of the range of possible forage availability. This can improve grazing management by revealing how forage quantities vary from year to year and may help avoid forage overutilization during unfavorable years such as drought. The model can continue to be used into the future to monitor vegetation trends, though with ongoing climate and vegetation changes periodic recalibration may be necessary. In addition, the modeling method may be applicable to other similar study systems.

We would like to thank Irene Martín Brull, Josep A. Ribera i Torró and Nuria García Díaz for their help in the field. This study was funded by the Spanish Ministry of Economy and Finance (grant CGL2016-79950- R) and the Junta de Andalucia I+D+i program (grant P18-RTJ- 4704)., All data are available from the Dryad Digital Repository https://doi. org/10.5061/dryad.mcvdn ck29 (Rubio de Casas et al., 2021)., The peer review history for this article is available at https://publons.com/publon/10.1111/1365-2745.13831., 1. Cultivated plant species often naturalize and enter wild communities in a process known as feralization. To successfully feralize, crops must overcome ecological barriers and may undergo selection on certain traits, diverging phenotypically and genetically from their crop ancestors. In spite of the agronomic and ecological relevance of crop feralization, the eco-evolutionary dynamics driving it remain understudied. 2. In this paper, we evaluated phenotypic and genotypic differentiation in fruit and seed traits during the naturalization of the almond tree (Prunus dulcis (Mill.) D.A. Webb) in SE Iberia and evaluated the potential role of natural selection in this process. To do so, we investigated the patterns of genetic divergence between cultivated and feral populations using functional (the cyanogenesis Sk gene) and neutral (17 SSR loci) markers and analysed morphological and biochemical traits in kernels of 342 individuals from 15 cultivated and 24 feral populations. 3. We detected very little genetic differentiation in neutral markers between cultivated and feral populations. The majority of the observed genetic variation was due to differences within each type. Conversely, the recessive allele sk responsible for seed toxicity was significantly more frequent in feral populations. Phenotypic differentiation between cultivated and naturalized almond populations was also significant. Feral almond kernels were smaller and lighter, had denser and more resistant shells (endocarps) and more toxic seeds. Selection analyses indicated that these genetic and phenotypic patterns might be driven by directional selection on fruit and seed traits, potentially linked to defence against predation. 4. Synthesis. Our findings indicate that almond naturalization is consistent with strong directional selection on fruits and seeds, leading to smaller and more toxic seeds encased in harder endocarps. Accordingly, we propose that feralization of this crop is, at least to some degree, driven by adaptive evolution of dispersal and recruitment traits., Secretaría de Estado de Investigación, Desarrollo e Innovación, Grant/Award Number: CGL2016-79950- R, Junta de Andalucia I+D+i program (grant P18-RTJ- 4704)

Mutualism effectiveness, the contribution of an interacting organism to its partner's fitness, is defined as the number of immediate outcomes of the interactions (quantity component) multiplied by the probability that an immediate outcome results in a new individual (quality component). These components form a two-dimensional effectiveness landscape with each species' location determined by its values of quantity (x-axis) and quality (y-axis). We propose that the evolutionary history of mutualistic interactions leaves a footprint that can be identified by three properties of the spatial structure of effectiveness values: dispersion of effectiveness values, relative contribution of each component to the effectiveness values and correlation between effectiveness components. We illustrate this approach using a large dataset on synzoochory, seed dispersal by seed-caching animals. The synzoochory landscape was clumped, with effectiveness determined primarily by the quality component, and with quantity and quality positively correlated. We suggest this type of landscape structure is common in generalised coevolved mutualisms, where multiple functionally equivalent, high-quality partners exert similarly strong selection. Presumably, only those organisms located in high-quality regions will impact the evolution of their partner. Exploring properties of effectiveness landscapes in other mutualisms will provide new insight into the evolutionary and ecological consequences of mutualisms.

Although dispersal is generally viewed as a crucial determinant for the fitness of any organism, our understanding of its role in the persistence and spread of plant populations remains incomplete. Generalizing and predicting dispersal processes are challenging due to context dependence of seed dispersal, environmental heterogeneity and interdependent processes occurring over multiple spatial and temporal scales. Current population models often use simple phenomenological descriptions of dispersal processes, limiting their ability to examine the role of population persistence and spread, especially under global change. To move seed dispersal ecology forward, we need to evaluate the impact of any single seed dispersal event within the full spatial and temporal context of a plant’s life history and environmental variability that ultimately influences a population’s ability to persist and spread. In this perspective, we provide guidance on integrating empirical and theoretical approaches that account for the context dependency of seed dispersal to improve our ability to generalize and predict the consequences of dispersal, and its anthropogenic alteration, across systems. We synthesize suitable theoretical frameworks for this work and discuss concepts, approaches and available data from diverse subdisciplines to help operationalize concepts, highlight recent breakthroughs across research areas and discuss ongoing challenges and open questions. We address knowledge gaps in the movement ecology of seeds and the integration of dispersal and demography that could benefit from such a synthesis. With an interdisciplinary perspective, we will be able to better understand how global change will impact seed dispersal processes, and potential cascading effects on plant population persistence, spread and biodiversity., Although dispersal is generally viewed as a crucial determinant for the fitness of any organism, our understanding of its role in the persistence and spread of plant populations remains incomplete. In this perspective, we provide guidance on integrating empirical and theoretical approaches that account for the context dependency of seed dispersal to improve our ability to generalize and predict the consequences of dispersal, and its anthropogenic alteration, across systems. With an interdisciplinary perspective, we will be able to better understand how global change will impact seed dispersal processes, and potential cascading effects on plant population persistence, spread and biodiversity.

Roads associated with energy development have fragmented much of the Uinta Basin, the Colorado Plateau in general, and other areas of western North America. Beyond reducing available habitat, spreading exotic species, and creating barriers to dispersal, unpaved roads also increase dust loads on plants and pollinators, which may reduce plant growth and reproduction. We studied the effects of an unpaved road on reproduction of an endangered Utah endemic shrub. We measured the size and reproductive output of 156 plants and the dust deposition in plots at increasing distances from the road. We also hand outcrossed 240 flowers from 80 plants to help determine if any reduced reproduction was due to pre- or postpollination mechanisms. Additionally, we experimentally dusted 3 leaves on 30 plants (n = 90) and measured stomatal conductance pre-dust and post-dust. We also dusted 3 flowers on 10 plants (n = 30) prior to hand pollination and measured fruit set. Generalized linear mixed models were used to examine the relationship between reproduction and dust deposition. When controlling for plant size and distance from the road, fruit set was negatively correlated with increasing levels of dust deposition (F1, 15 = 5.26, P = 0.036). The number of seeds per plant, mean plant seed weight, and the proportion of hand-pollinated flowers that set fruit were also negatively correlated with dust, although not significantly. Dusting significantly reduced stomatal conductance (F1, 58 = 87.56, P < 0.001). Eighty percent of hand pollinated flowers (24 of 30) set fruit after dusting. These results demonstrate that road dust reduces H. suffrutescens reproduction, although the mechanisms are not clear. Although dust negatively affected physiological processes, hand-pollination results suggest that dust might be disrupting pollination. This study documents the effects of road dust on the reproduction of an endangered shrub in Utah's Uinta Basin and highlights the need for further research into the effects of roads and dust on nearby plants.

There is growing realization that intraspecific variation in seed dispersal can have important ecological and evolutionary consequences. However, we do not have a good understanding of the drivers or causes of intraspecific variation in dispersal, how strong an effect these drivers have, and how widespread they are across dispersal modes. As a first step to developing a better understanding, we present a broad, but not exhaustive, review of what is known about the drivers of intraspecific variation in seed dispersal, and what remains uncertain. We start by decomposing ‘drivers of intraspecific variation in seed dispersal’ into intrinsic drivers (i.e. variation in traits of individual plants) and extrinsic drivers (i.e. variation in ecological context). For intrinsic traits, we further decompose intraspecific variation into variation among individuals and variation of trait values within individuals. We then review our understanding of the major intrinsic and extrinsic drivers of intraspecific variation in seed dispersal, with an emphasis on variation among individuals. Crop size is the best-supported and best-understood intrinsic driver of variation across dispersal modes; overall, more seeds are dispersed as more seeds are produced, even in cases where per seed dispersal rates decline. Fruit/seed size is the second most widely studied intrinsic driver, and is also relevant to a broad range of seed dispersal modes. Remaining intrinsic drivers are poorly understood, and range from effects that are probably widespread, such as plant height, to drivers that are most likely sporadic, such as fruit or seed colour polymorphism. Primary extrinsic drivers of variation in seed dispersal include local environmental conditions and habitat structure. Finally, we present a selection of outstanding questions as a starting point to advance our understanding of individual variation in seed dispersal., Intraspecific variation in the quantity and quality of seed dispersal has important ecological and evolutionary consequences, yet it is generally ignored in favour of simply using population mean values. In a broad but not exhaustive review we show that drivers of intraspecific variation in seed dispersal are diverse and pervasive. These include both intrinsic (i.e. variation in traits of individual plants) and extrinsic (i.e. variation in ecological context) drivers. Further, these drivers appear to interact frequently in complex ways. Current theory does not include or account for these complex and interacting drivers.

As the single opportunity for plants to move, seed dispersal has an important impact on plant fitness, species distributions and patterns of biodiversity. However, models that predict dynamics such as risk of extinction, range shifts and biodiversity loss tend to rely on the mean value of parameters and rarely incorporate realistic dispersal mechanisms. By focusing on the mean population value, variation among individuals or variability caused by complex spatial and temporal dynamics is ignored. This calls for increased efforts to understand individual variation in dispersal and integrate it more explicitly into population and community models involving dispersal. However, the sources, magnitude and outcomes of intraspecific variation in dispersal are poorly characterized, limiting our understanding of the role of dispersal in mediating the dynamics of communities and their response to global change. In this manuscript, we synthesize recent research that examines the sources of individual variation in dispersal and emphasize its implications for plant fitness, populations and communities. We argue that this intraspecific variation in seed dispersal does not simply add noise to systems, but, in fact, alters dispersal processes and patterns with consequences for demography, communities, evolution and response to anthropogenic changes. We conclude with recommendations for moving this field of research forward., Seed dispersal—the single opportunity for plants to move—is important for plant fitness, species distributions and patterns of biodiversity. Models that predict extinction risk of species, range shifts and biodiversity tend to rely on average dispersal distances. However, we know that seed dispersal is highly variable even within a single species (e.g. some seeds go very far and some barely move away from their parent plant, some seeds end up in great quality habitats and some end up on roads). This paper looks at the consequences of this variation in seed dispersal for plants and their ability to respond to future global change.

Synzoochory is the dispersal of seeds by seed-caching animals. The animal partner in this interaction plays a dual role, acting both as seed disperser and seed predator. We propose that this duality gives to synzoochory two distinctive features that have crucial ecological and evolutionary consequences. First, because plants attract animals that have not only positive (seed dispersal) but also negative (seed predation) impacts on their fitness, the evolution of adaptations to synzoochory is strongly constrained. Consequently, it is not easy to identify traits that define a synzoochorous dispersal syndrome. The absence of clear adaptations entails the extra difficulty of identifying synzoochorous plants by relying on dispersal traits, limiting our ability to explore the full geographic, taxonomic and phylogenetic extent of synzoochory. Second, the positive and negative outcomes of interactions with synzoochorous animals are expressed simultaneously. Consequently, synzoochorous interactions are not exclusively mutualistic or antagonistic, but are located at some point along a mutualism-antagonism continuum. What makes synzoochory interesting and unique is that the position of each partner along the continuum can be evaluated for every plant—animal interaction, and thus the continuum can be precisely described by assessing the relative frequency of positive and negative interaction events in each pairwise interaction. Herein we explore these two main features of synzoochory with a comprehensive quantitative survey of published studies on synzoochory. Synzoochory has been recorded for at least 1339 plant species differing in life forms, from annual and short-lived herbs to long-lived trees, belonging to 641 genera and 157 families widely distributed across the globe and across the seed plant phylogeny. Over 30 animal families belonging to five disparate taxonomic groups (rodents, marsupials, birds, insects, and land crabs) potentially act as synzoochorous dispersers. Although synzoochory appears to be fundamentally a secondary dispersal mode, many abundant and dominant trees are primarily synzoochorous. In addition, we found evidence of the existence of diplosynzoochory (caching animals acting both as primary and secondary dispersers of the same individual seed), mostly in nut-bearing trees. Finally, we found that synzoochorous interactions are widely spread across the mutualism—antagonism continuum. Nevertheless, there were some differences among disperser species and functional groups. Corvids and some rodents (cricetids, nesomyids, sciurids) were located in the positive-effects region of the continuum and presumably behave mostly as dispersers, whereas land crabs and insects were located in the negative-effects extreme and behave mostly as seed predators. Our review demonstrates that synzoochory is not an anecdotal ecological interaction. Rather, it is pivotal to the functioning of many ecosystems where the natural regeneration of keystone plant species depends on the activity of granivorous animals that play a dual role. This distinctive interaction should not be ignored if we wish to have an accurate understanding of the functioning of natural systems., J.M.G. and P.J. were supported by CYTED program (Red Temática 418RT0555). E.W.S. is supported by the Ecology Center and the Utah Agricultural Experiment Station (UAES), Utah State University. P.J. is supported by grant CGL2017‐82847‐P from the Spanish Ministry of Science, Innovation, and Universities (AEI).

When climatic or environmental conditions change, plant populations must either adapt to these new conditions, or track their niche via seed dispersal. Adaptation of plants to different abiotic environments has mostly been discussed with respect to physiological and demographic parameters that allow local persistence. However, rapid modifications in response to changing environmental conditions can also affect seed dispersal, both via plant traits and via their dispersal agents. Studying such changes empirically is challenging, due to the high variability in dispersal success, resulting from environmental heterogeneity, and substantial phenotypic variability of dispersal-related traits of seeds and their dispersers. The exact mechanisms that drive rapid changes are often not well understood, but the ecological implications of these processes are essential determinants of dispersal success, and deserve more attention from ecologists, especially in the context of adaptation to global change. We outline the evidence for rapid changes in seed dispersal traits by discussing variability due to plasticity or genetics broadly, and describe the specific traits and biological systems in which variability in dispersal is being studied, before discussing some of the potential underlying mechanisms. We then address future research needs and propose a simulation model that incorporates phenotypic plasticity in seed dispersal. We close with a call to action and encourage ecologists and biologist to embrace the challenge of better understanding rapid changes in seed dispersal and their consequences for the reaction of plant populations to global change., Global ecological change is causing plant populations to either adapt or move in response to new environmental conditions. In many species it is thought that seed dispersal may not allow populations to move at a rate that tracks the shifting climate. However, phenotypic plasticity and rapid genetic changes in traits associated with seed dispersal may modify the response capabilities of plants and allow them to responds more quickly. We explore the evidence for rapid modification of seed dispersal traits and propose a path forward to better understand the ways in which seed dispersal may buffer the effects of global ecological change.

Invasion and dominance of exotic grasses and increased fire frequency threaten native ecosystems worldwide. In the Great Basin region of the western United States, woody and herbaceous fuel treatments are implemented to decrease the effects of wildfire and increase sagebrush (Artemisia spp.) ecosystem resilience to disturbance and resistance to exotic annual grasses. High cover of the exotic annual cheatgrass (Bromus tectorum) after treatments increases fine fuels, which in turn increases the risk of passing over a biotic threshold to a state of increased wildfire frequency and conversion to cheatgrass dominance. Sagebrush ecosystem resilience to wildfire and resistance to cheatgrass depend on climatic conditions and abundance of perennial herbaceous species that compete with cheatgrass. In this study, we used longer‐term data to evaluate the relationships among soil climate conditions, perennial herbaceous cover, and cheatgrass cover following fuel management treatments across the environmental gradients that characterize sagebrush ecosystems in the Great Basin. We examined the effects of woody and herbaceous fuel treatments on soil temperature, soil water availability (13–30 and 50 cm depths), and native and exotic plant cover on six sagebrush sites lacking piñon (Pinus spp.) or juniper (Juniperus spp.) tree expansion and 11 sagebrush sites with tree expansion. Both prescribed fire and mechanical treatments increased soil water availability on woodland sites and perennial herbaceous cover on some woodland and sagebrush sites. Prescribed fire also slightly increased soil temperatures and especially increased cheatgrass cover compared to no treatment and mechanical treatments on most sites. Non‐metric dimensional scaling ordination and decision tree partition analysis indicated that sites with warmer late springs and warmer and wetter falls had higher cover of cheatgrass. Sites with wetter winters and early springs (March–April) had higher cover of perennial herbs. Our findings suggest that site resistance to cheatgrass after fire and fuel control treatments decreases with a warmer and drier climate. This emphasizes the need for management actions to maintain and enhance perennial herb cover, such as implementing appropriate grazing management, and revegetating sites that have low abundance of perennial herbs in conjunction with fuel control treatments.

We investigated fruit set, seed set, and germination requirements of shrubby reed-mustard (Hesperidanthus suffrutescens), an endangered endemic shrub in the Uinta Basin of eastern Utah, U.S.A. To determine the degree of self-compatibility, 120 plants received four pollination treatments each for 2 y. Treatments included autogamy, geitonogamy, xenogamy, and an open control. Autogamy and geitonogamy produced substantially and significantly fewer fruits, seeds, and seeds per fruit than xenogamy, suggesting little self-compatibility. Additionally, the maternal reproductive success of outcrossed flowers was significantly greater than open control flowers suggesting pollen limitation, perhaps due to reduced pollinator abundance or activity. One study population showed significantly reduced reproduction during the second year, perhaps resulting from impacts of anthropogenic disturbance. Although both selfing treatments produced significantly fewer seeds, there were no significant differences in germinat...

The Sagebrush Steppe Treatment Evaluation Project (SageSTEP) is an integrated long-term study that evaluates ecological effects of alternative treatments designed to reduce woody fuels and to stimulate the herbaceous understory of sagebrush steppe communities of the Intermountain West. This synopsis summarizes results through 3 yr posttreatment. Woody vegetation reduction by prescribed fire, mechanical treatments, or herbicides initiated a cascade of effects, beginning with increased availability of nitrogen and soil water, followed by increased growth of herbaceous vegetation. Response of butterflies and magnitudes of runoff and erosion closely followed herbaceous vegetation recovery. Effects on shrubs, biological soil crust, tree cover, surface woody fuel loads, and sagebrush-obligate bird communities will take longer to be fully expressed. In the short term, cool wet sites were more resilient than warm dry sites, and resistance was mostly dependent on pretreatment herbaceous cover. At least 10 yr of posttreatment time will likely be necessary to determine outcomes for most sites. Mechanical treatments did not serve as surrogates for prescribed fire in how each influenced the fuel bed, the soil, erosion, and sage-obligate bird communities. Woody vegetation reduction by any means resulted in increased availability of soil water, higher herbaceous cover, and greater butterfly numbers. We identified several trade-offs (desirable outcomes for some variables, undesirable for others), involving most components of the study system. Trade-offs are inevitable when managing complex natural systems, and they underline the importance of asking questions about the whole system when developing management objectives. Substantial spatial and temporal heterogeneity in sagebrush steppe ecosystems emphasizes the point that there will rarely be a “recipe” for choosing management actions on any specific area. Use of a consistent evaluation process linked to monitoring may be the best chance managers have for arresting woodland expansion and cheatgrass invasion that may accelerate in a future warming climate.

Despite the critical need to improve degraded herbaceous understory conditions in many semiarid ecosystems, the influence of soil properties on seedling emergence of species seeded in shrubland plant communities is largely unexplored. We evaluated emergence patterns of 6 restoration species in soils from wyomingensis (i.e., Wyoming big sagebrush, Artemisia tridentata ssp. wyomingensis [Beetle & A. Young] S.L. Welsh) and vaseyana (i.e., mountain big sagebrush, A. t. ssp. vaseyana [Rydb.] Beetle) plant communities that differed in soil texture, soil organic matter content, and soil water-holding capacity. We conducted 2 separate experiments that regularly wetted soils to standardized soil water potentials (i.e., field capacity; −0.03 MPa) and allowed differences in evaporation to create distinct wet-dry watering pattern cycles over a 26–29 d period. Our objectives were to compare soil attributes of wyomingensis and vaseyana soils, evaluate whether emergence patterns of restoration species vary within these soils, and determine how these patterns are altered by soil water-content levels. We found differences in soil texture and organic matter between soils and thus soil water-holding capacity: finer-textured vaseyana soils held roughly twofold more water than coarse-textured wyomingensis soils. Seeds in vaseyana soils were exposed to fewer wet-dry cycles compared to wyomingensis soils because of the greater capacity of vaseyana soils to retain water. Restoration species also collectively exhibited greater emergence in vaseyana soils than in wyomingensis soils, yet emergence patterns were vastly different among species, and differences between soils became more pronounced under low soil water for only 2 species. We conclude that the manner in which soils and water uniquely influenced emergence patterns provides new insights into species suitability for restoration sites and how inherent soil differences may constrain seeding success.

There is increasing recognition that both competition and facilitation are important drivers of plant community dynamics in arid and semi-arid environments. Decades of research have provided a litany of examples of the potential for shrubs as nurse plants for establishment of desirable species, especially in water-limited environments. However, interactions with the existing understory community may alter the outcome of interactions between shrubs and understory plants. A manipulative experiment was conducted to disentangle interactions between a native forb species (Penstemon palmeri A. Gray), a native shrub (Artemisia tridentata Nutt.), and a diverse understory of exotic and native forbs and grasses in a semi-arid shrubland of Northern Utah, USA. Seedlings of P. palmeri were transplanted in a factorial design: (1) beneath shrub canopies or into their interspaces and (2) with understory interactions retained or removed. Transplant survival was tracked for roughly 1 year. Shrubs appeared to facilitate P. palmeri survival while interactions with the existing understory community were equivalently negative, leading to overall neutral interactions. Further, positive shrub interactions and negative understory interactions appeared to operate independently and simultaneously. While the debate over the importance of facilitation and competition in driving plant community dynamics continues, our observations strongly suggest that both have considerable effects on plant establishment in A. tridentata communities. Furthermore, our results inform the conservation and restoration of P. palmeri populations, and suggest the utility of nurse shrubs and/or understory thinning as strategies for increasing the diversity of desirable species in the arid and semi-arid western United States shrublands.

Plant spatial patterns critically influence community dynamics, including plant interactions, resource distribution, and community invasibility. Research suggests that resistance of western US plant communities to further invasion by the exotic annual grass Bromus tectorum may be linked to the positions of, and spacing between, perennial plants. In particular, gaps between aggregated clusters of perennial plants may facilitate B. tectorum invasion by providing safe sites for seed germination and establishment. We tested the effects of random, regular, and aggregated bunchgrass patterns, manipulated at both community (plot) and neighborhood scales, on B. tectorum biomass and spikelet production after experimental seed addition. We found strong evidence of treatment effects on both biomass and spikelets, which varied between treatments by approximately 2.5-fold. Mean biomass and spikelet counts were lowest in plots in which bunchgrasses were aggregated at both community and neighborhood scales, likely due to the increased competition. Although not statistically distinguishable from most other treatments, B. tectorum biomass and spikelet counts were highest in plots with bunchgrass patterns that were random at the community scale and aggregated at the neighborhood scale. These plots were characterized by relatively large gaps between bunchgrass clusters, suggesting that B. tectorum may exploit gaps between aggregated perennial plants. Our results support the emerging hypothesis that community resistance to B. tectorum invasion could be increased through manipulation of perennial vegetation to reduce basal gap size and connectivity.

Summary Rodents frequently forage in a density-dependent manner, increasing harvesting in patches with greater seed densities. Although seldom considered, seed harvesting may also depend on the species identities of other individuals in the seed neighbourhood. When the seed harvest of a focal species increases in association with another seed species, the focal species suffers from Associational Susceptibility. In contrast, if seeds of the focal species are harvested less when in association with a second species, the focal species benefits from Associational Resistance. To evaluate density dependence and associational effects among seeds in mixtures, we conducted seed removal experiments using a completely additive design patterned after a two-species competition experiment using seeds of either Achnatherum hymenoides (Indian ricegrass), Leymus cinereus (basin wildrye) or Pseudoroegneria spicata (bluebunch wheatgrass), all native perennial grasses, combined with seeds of Bromus tectorum (cheatgrass), a non-native annual grass. The experiment involved placing five fixed quantities of the native seeds mixed with five fixed quantities of B. tectorum seeds in a factorial design, resulting in 35 seed mixture combinations. The seed-eating rodent community at our study sites, in order of abundance, is composed of Peromyscus maniculatus (North American deer mouse), Dipodomys ordii (Ord's kangaroo rat) and Perognathus parvus (Great Basin pocket mouse). Native seed harvesting was density dependent, with a greater proportion of seeds being harvested as density increased. In the mixed density model, the presence of B. tectorum did not affect harvest of any of the native species' seeds when analysed individually. However, when all three native species were analysed together, increasing quantities of B. tectorum resulted in reduced harvest of native seeds, demonstrating weak but significant Associational Resistance. In contrast, harvest of B. tectorum seeds increased when in combination with any of the native seed species individually, indicating relatively strong Associational Susceptibility. These results demonstrate that seed harvest is determined not just by seed density, but also by the local seed neighbourhood and suggest that associational effects between native seeds and B. tectorum can occur in field conditions. The ecological implications of seed selection and associational effects on plant populations in natural and managed systems are also discussed.

Sagebrush steppe ecosystems in the United States currently (2016) occur on only about one-half of their historical land area because of changes in land use, urban growth, and degradation of land, including invasions of non-native plants. The existence of many animal species depends on the existence of sagebrush steppe habitat. The greater sage-grouse (Centrocercus urophasianus) depends on large landscapes of intact habitat of sagebrush and perennial grasses for their existence. In addition, other sagebrush-obligate animals have similar requirements and restoration of landscapes for greater sage-grouse also will benefit these animals. Once sagebrush lands are degraded, they may require restoration actions to make those lands viable habitat for supporting sagebrush-obligate animals, livestock, and wild horses, and to provide ecosystem services for humans now and for future generations. When a decision is made on where restoration treatments should be applied, there are a number of site-specific decisions managers face before selecting the appropriate type of restoration. This site-level decision tool for restoration of sagebrush steppe ecosystems is organized in nine steps. ●Step 1 describes the process of defining site-level restoration objectives. ●Step 2 describes the ecological site characteristics of the restoration site. This covers soil chemistry and texture, soil moisture and temperature regimes, and the vegetation communities the site is capable of supporting. ●Step 3 compares the current vegetation to the plant communities associated with the site State and Transition models. ●Step 4 takes the manager through the process of current land uses and past disturbances that may influence restoration success. ●Step 5 is a brief discussion of how weather before and after treatments may impact restoration success. ●Step 6 addresses restoration treatment types and their potential positive and negative impacts on the ecosystem and on habitats, especially for greater sage-grouse. We discuss when passive restoration options may be sufficient and when active restoration may be necessary to achieve restoration objectives. ●Step 7 addresses decisions regarding post-restoration livestock grazing management. ●Step 8 addresses monitoring of the restoration; we discuss important aspects associated with implementation monitoring as well as effectiveness monitoring. ●Step 9 takes the information learned from monitoring to determine how restoration actions in the future might be adapted to improve restoration success.

A core interest in studies of mutualistic interactions is the ‘effectiveness’ of mutualists in providing benefits to their partners. In plant-animal mutualisms it is widely accepted that the total effect of a mutualist on its partner is estimated as (1) a ‘quantity’ component multiplied by (2) a ‘quality’ component, although the meanings of ‘effectiveness,’ ‘quantity,’ and ‘quality’ and which terms are applied to these metrics vary greatly across studies. In addition, a similar quantity × quality = total effect approach has not been applied to other types of mutualisms, although it could be informative. Lastly, when a total effect approach has been applied, it has invariably been from a phytocentric perspective, focussing on the effects of animal mutualists on their plant partner. This lack of a common framework of ‘effectiveness’ of mutualistic interactions limits generalisation and the development of a broader understanding of the ecology and evolution of mutualisms. In this paper, we propose a general framework and demonstrate its utility by applying it to both partners in five different types of mutualisms: pollination, seed dispersal, plant protection, rhizobial, and mycorrhizal mutualisms. We then briefly discuss the flexibility of the framework, potential limitations, and relationship to other approaches.

Granivore foraging decisions affect consumer success and determine the quantity and spatial pattern of seed survival. These decisions are influenced by environmental variation at spatial scales ranging from landscapes to local foraging patches. In a field experiment, the effects of seed patch variation across three spatial scales on seed removal by western harvester ants Pogonomyrmex occidentalis were evaluated. At the largest scale we assessed harvesting in different plant communities, at the intermediate scale we assessed harvesting at different distances from ant mounds, and at the smallest scale we assessed the effects of interactions among seed species in local seed neighborhoods on seed harvesting (i.e. resource–consumer interface). Selected seed species were presented alone (monospecific treatment) and in mixture with Bromus tectorum (cheatgrass; mixture treatment) at four distances from P. occidentalis mounds in adjacent intact sagebrush and non-native cheatgrass-dominated communities in the Great Basin, Utah, USA. Seed species differed in harvest, with B. tectorum being least preferred. Large and intermediate scale variation influenced harvest. More seeds were harvested in sagebrush than in cheatgrass-dominated communities (largest scale), and the quantity of seed harvested varied with distance from mounds (intermediate-scale), although the form of the distance effect differed between plant communities. At the smallest scale, seed neighborhood affected harvest, but the patterns differed among seed species considered. Ants harvested fewer seeds from mixed-seed neighborhoods than from monospecific neighborhoods, suggesting context dependence and potential associational resistance. Further, the effects of plant community and distance from mound on seed harvest in mixtures differed from their effects in monospecific treatments. Beyond the local seed neighborhood, selection of seed resources is better understood by simultaneously evaluating removal at multiple scales. Associational effects provide a useful theoretical basis for better understanding harvester ant foraging decisions. These results demonstrate the importance of ecological context for seed removal, which has implications for seed pools, plant populations and communities.

Ex situ conservation of rare plant species requires an understanding of germination requirements. We report the first successful germination trials for Maguire primrose (Primula cusickiana var. maguirei[L.O. Williams] N.H. Holmgren & S. Kelso), a threatened perennial herb narrowly restricted to a 20-km canyon in northern Utah, USA. Seeds collected from different sites (upper or lower canyon populations) and flower morphs (pin or thrum) that had been stored for either 1 or 2 years were cold, moist stratified in darkness at 1–2°C for 270 days. Independent of treatment, time to first germination was approximately 75 days, time to 50% germination (T50) was approximately 165 days, and total germination was 55.8%. Higher mean germination was observed in seeds stored for 2 years compared with seeds stored for 1 year, as well as in seeds from pin flowers compared with thrum flowers. There was not a significant effect of site on mean germination, but there was a significant storage time × site interaction. Seeds stored for 2 years also had a lower mean T50 versus seeds stored for 1 year. In addition, seeds from upper canyon populations had a higher mean T50 than those from lower canyon populations. Our results suggest that short-term storage should not significantly decrease seed viability, and that both flower morph and canyon location may influence germination. Evidence suggests that germination patterns are adaptive. Our results contribute to our understanding of the germination biology of P. cusickiana var. maguirei and inform future conservation efforts for this threatened species.

Community-structuring processes continue to be of great interest to plant ecologists, and plant spatial patterns have been linked to processes including disturbance, dispersal, environmental heterogeneity, and plant interactions. Under the assumption that the analysis of the spatial structure of plant communities can help to elucidate the type and importance of the predominant community-structuring processes, many studies have analyzed point pattern data on various plant species. A variety of methods have been devised to acquire point pattern data for individual plants, however, the classic tradeoff between the speed of acquisition and the precision of spatial data has meant that large and precise datasets on plant locations are difficult to obtain. The primary goal of this study was to develop a GPS-based methodology for the rapid collection of precise spatial data on plant locations in a semi-arid shrubland in the Great Basin, USA. The secondary goal was to demonstrate a potential application of this approach by using recently developed univariate and bivariate spatial statistics to test for aggregation within the shrub community, as observed in other semi-arid shrublands. We efficiently mapped 2,358 individuals of five shrub species with a spatial error of ≤0.02 m, and found strong statistical evidence of fine-scale aggregation (1) independent of species, (2) within species, and (3) between two species pairs. Our approach is useful for rapidly collecting precise point pattern data in plant communities, and has other applications related to population modeling, GIS analysis, and conservation.

Growth in seed dispersal studies has been fast-paced since the seed disperser effec- tiveness (SDE) framework was developed 17 yr ago. Thus, the time is ripe to revisit the framework in light of accumulated new insight. Here, we first present an over- view of the framework, how it has been applied, and what we know and do not know. We then introduce the SDE landscape as the two-dimensional representa- tion of the possible combinations of the quantity and the quality of dispersal and with elevational contours representing isoclines of SDE. We discuss the structure of disperser assemblages on such landscapes. Following this we discuss recent advances and ideas in seed dispersal in the context of their impacts on SDE. Finally, we highlight a number of emerging issues that provide insight into SDE. Overall, the SDE framework successfully captures the complexities of seed dispersal. We advocate an expanded use of the term dispersal encompassing the multiple recruit- ment stages from fruit to adult. While this entails difficulties in estimating SDE, it is a necessary expansion if we are to understand the central relevance of seed dispersal in plant ecology and evolution.

Aim The exotic annual cheatgrass (Bromus tectorum) is fast replacing sagebrush (Artemisia tridentata) communities throughout the Great Basin Desert and nearby regions in the Western United States, impacting native plant communities and altering fire regimes, which contributes to the long-term persistence of this weedy species. The effect of this conversion on native faunal communities remains largely unexamined. We assess the impact of conversion from native perennial to exotic annual plant communities on desert rodent communities. Location Wyoming big sagebrush shrublands and nearby sites previously converted to cheatgrass-dominated annual grasslands in the Great Basin Desert, Utah, USA. Methods At two sites in Tooele County, Utah, USA, we investigated with Sherman live trapping whether intact sagebrush vegetation and nearby converted Bromus tectorum-dominated vegetation differed in rodent abundance, diversity and community composition. Results Rodent abundance and species richness were considerably greater in sagebrush plots than in cheatgrass-dominated plots. Nine species were captured in sagebrush plots; five of these were also trapped in cheatgrass plots, all at lower abundances than in the sagebrush. In contrast, cheatgrass-dominated plots had no species that were not found in sagebrush. In addition, the site that had been converted to cheatgrass longer had lower abundances of rodents than the site more recently converted to cheatgrass-dominated plots. Despite large differences in abundances and species richness, Simpson’s D diversity and Shannon-Wiener diversity and Brillouin evenness indices did not differ between sagebrush and cheatgrass-dominated plots. Main conclusions This survey of rodent communities in native sagebrush and in converted cheatgrass-dominated vegetation suggests that the abundances and community composition of rodents may be shifting, potentially at the larger spatial scale of the entire Great Basin, where cheatgrass continues to invade and dominate more landscape at a rapid rate.

Two varieties of sterile annual hybrid (SAH) cereal grasses (Triticum aestivum×Elytrigia elongata and Triticum sp. ×Secale sp.) have been developed as tools for restoration to circumvent problems associated with the use of exotic perennial grasses in wildlands. Recent research has shown that both varieties of SAH grasses performed better than native species when sown into Cheatgrass (Bromus tectorum)-dominated rangelands. The primary goal of this study was to see whether early emergence was a potential mechanism contributing to the success of SAH grasses. Second, we investigated whether the SAH grasses were indeed sterile. Seeds of both SAH varieties were germinated in growth chambers along with B. tectorum and four perennial grasses. Seeds were exposed to three moisture levels and four temperature regimes. In a separate experiment, seed from 10 individuals of each SAH variety grown in a greenhouse were tested for sterility. Triticum aestivum×E. elongata consistently emerged fastest under all moisture and temperature levels, followed by Triticum sp. ×Secale sp., then B. tectorum, and, finally, the four perennial grass species. All interactions also were significant; these interactions modified without negating the general patterns revealed in the main effects. Seed production in the sterility trial for both SAH varieties was highly variable among individuals and all seed that were produced germinated. The ability of SAH grasses to maintain rapid emergence under adverse conditions suggests that they may be a useful tool in restoration; however, their lack of true sterility may contradict the goals of land managers.

Biological invasions are one of the greatest threats to native species in natural ecological systems. One of the most successful invasive species is Bromus tectorum L. (cheatgrass), which is having marked impacts on native plant communities and ecosystem processes. However, we know little about the effects of this invasion on native animal species in the Intermountain West. Because ants have been used to detect ecological change associated with anthropogenic land use, they seem well suited for a preliminary evaluation of the consequences of cheatgrass-driven habitat conversion. In our study, we used pitfall traps to assess ant community assemblages in intact sagebrush and nearby cheatgrass-dominated vegetation. Ant abundance was about 10-fold greater in cheatgrass-dominated plots than in sagebrush plots. We also noted differences in diversity and evenness between habitat types at both the species and the functional-group levels of organization. At the species level, Shannon's diversity index was...

The factors that determine plant community resistance to exotic annual Bromus species (Bromus hereafter) are diverse and context specific. They are influenced by the environmental characteristics and attributes of the community, the traits of Bromus species, and the direct and indirect interactions of Bromus with the plant community. Environmental factors, in particular ambient and soil temperatures, have significant effects on the ability of Bromus to establish and spread. Seasonality of precipitation relative to temperature influences plant community resistance to Bromus through effects on soil water storage, timing of water and nutrient availability, and dominant plant life forms. Differences among plant communities in how well soil resource use by the plant community matches resource supply rates can influence the magnitude of resource fluctuations due to either climate or disturbance and thus the opportunities for invasion. The spatial and temporal patterns of resource availability and acquisition of growth resources by Bromus versus native species strongly influence resistance to invasion. Traits of Bromus that confer a “priority advantage” for resource use in many communities include early-season germination and high growth and reproductive rates. Resistance to Bromus can be overwhelmed by high propagule supply, low innate seed dormancy, and large, if short-lived, seed banks. Biological crusts can inhibit germination and establishment of invasive annual plants, including several annual Bromus species, but are effective only in the absence of disturbance. Herbivores can have negative direct effects on Bromus, but positive indirect effects through decreases in competitors. Management strategies can be improved through increased understanding of community resistance to exotic annual Bromus species.

In this study we assessed the effectiveness of rodents as dispersers of Quercus ilex in a patchy landscape in southeastern Spain. We experimentally followed the fates of 3,200 marked and weighed acorns from dispersal through the time of seedling emergence over three years. Rodents handled about 99% of acorns, and dispersed 67% and cached 7.4% of the dispersed acorns. Most caches were recovered and consumed, and only 1.3% of the original experimental acorns were found alive in caches the following spring. Dispersal distances were short (mean = 356.2 cm, median = 157 cm) and strongly right-skewed. Heavier acorns were dispersed further and were more likely to be cached and survive than lighter acorns. All caches were in litter or soil, and each contained a single acorn. Rodents moved acorns nonrandomly, mostly to oaks and pines. Most surviving acorns were either in oaks, a poor microhabitat for oak recruitment, or shrubs, a suitable microhabitat for oak recruitment. Our results suggest that rodents, by burying a relatively high proportion of acorns singly in shrubs and pines, act as moderately effective dispersers of Q. ilex. Nonetheless, this dispersal comes at a very heavy cost.

Sagebrush steppe ecosystems in the United States currently occur on only about one-half of their historical land area because of changes in land use, urban growth, and degradation of land, including invasions of non-native plants. The existence of many animal species depends on the existence of sagebrush steppe habitat. The greater sage-grouse (Centrocercus urophasianus) is a landscape-dependent bird that requires intact habitat and combinations of sagebrush and perennial grasses to exist. In addition, other sagebrush-obligate animals also have similar requirements and restoration of landscapes for greater sage-grouse also will benefit these animals. Once sagebrush lands are degraded, they may require restoration actions to make those lands viable habitat for supporting sagebrush-obligate animals. This restoration handbook is the first in a three-part series on restoration of sagebrush ecosystems. In Part 1, we discuss concepts surrounding landscape and restoration ecology of sagebrush ecosystems and greater sage-grouse that habitat managers and restoration practitioners need to know to make informed decisions regarding where and how to restore specific areas. We will describe the plant dynamics of sagebrush steppe ecosystems and their responses to major disturbances, fire, and defoliation. We will introduce the concepts of ecosystem resilience to disturbances and resistance to invasions of annual grasses within sagebrush steppe. An introduction to soils and ecological site information will provide insights into the specific plants that can be restored in a location. Soil temperature and moisture regimes are described as a tool for determining resilience and resistance and the potential for various restoration actions. Greater sage-grouse are considered landscape birds that require large areas of intact sagebrush steppe; therefore, we describe concepts of landscape ecology that aid our decisions regarding habitat restoration. We provide a brief overview of restoration techniques for sage-grouse habitat restoration. We conclude with a description of the critical nature of monitoring for adaptive management of sagebrush steppe restoration at landscape- and project-specific levels.

The alien grass Bromus tectorum dominates stable annual-plant communities that have replaced native shrub-perennial grass communities over much of the semi-arid western United States. We conducted field competition experiments between B. tectorum and a native grass, Elymus elymoides, on two sites to determine the effects of B. tectorum competition on perennial grasses, and the role of B. tectorum competition in the stability of B. tectorum-dominated communities. B. tectorum competition acting on seedling-stage E. elymoides plants greatly reduced first-year relative growth rates and biomass which, in turn, reduced second-year survival, biomass, and flowering. However, B. tectorum competition acting on older E. elymoides plants had much less effect, which may help to explain why intact perennial-plant communities are resistant to B. tectorum invasion. At the drier site, direct effects of B. tectorum competition were less, but competition and drier habitat combined to produce greater E. elymoides mortality.

Agricultural land abandonment is rampant in present day Europe. A major consequence of this phenomenon is the re-colonization of these areas by the original vegetation. However, some agricultural, exotic species are able to naturalize and colonize these abandoned lands. In this study we explore the ability of almonds (Prunus dulcis D.A. Webb.) to establish in abandoned croplands in semi-arid areas of SE Iberian Peninsula. Domesticated during the early Holocene in SW Asia and the Eastern Mediterranean, the almond has spread as a crop all over the world. We established three plots adjacent to almond orchards on land that was abandoned and reforested with Aleppo pine (Pinus halepensis Mill.) and Holm oak (Quercus ilex L.) about 20 years ago. We recorded the abundance of almond seeds, seedlings, juveniles, pre-reproductives and adults in these plots, and determined their recruitment microhabitat. We found natural regeneration in all three plots, with almond density higher than 150 individuals/ha in each plot. About half of the almonds in the plots were juveniles, although 7% were one-year seedlings and 3% were adult trees. This suggests that colonization of abandoned lands is an ongoing gradual process. Most naturally established almonds were under tree cover, although this was not the most abundant microhabitat, avoiding the negative impact of browsing by Red Deer (Cervus elaphus) and domestic sheep and goats. In addition, the mean distance from the nearest seed source was 77 m. These findings suggest the action of mobile almond dispersers. Several Corvid species including the Carrion Crow (Corvus corone) and the Magpie (Pica pica) have been observed consuming and moving almonds in the study area. Our study has demonstrated that almonds are colonizing abandoned lands mostly as a consequence of effective seed dispersal by some animals. We show that almonds have the potential to become fully naturalized in the near future in many semi-arid areas of the Western Mediterranean.

Sagebrush steppe ecosystems in the United States currently (2015) occur on only about one-half of their historical land area because of changes in land use, urban growth, and degradation of land, including invasions of non-native plants. The existence of many animal species depends on the existence of sagebrush steppe habitat. The greater sage-grouse (Centrocercus urophasianus) is a landscape-dependent bird that requires intact habitat and combinations of sagebrush and perennial grasses to exist. In addition, other sagebrush-obligate animals also have similar requirements and restoration of landscapes for greater sage-grouse also will benefit these animals. Once sagebrush lands are degraded, they may require restoration actions to make those lands viable habitat for supporting sagebrush-obligate animals. Land managers do not have resources to restore all locations because of the extent of the restoration need and because some land uses are not likely to change, therefore, restoration decisions made at the landscape to regional scale may improve the effectiveness of restoration to achieve landscape and local restoration objectives. We present a landscape restoration decision tool intended to assist decision makers in determining landscape objectives, to identify and prioritize landscape areas where sites for priority restoration projects might be located, and to aid in ultimately selecting restoration sites guided by criteria used to define the landscape objectives. The landscape restoration decision tool is structured in five sections that should be addressed sequentially. Each section has a primary question or statement followed by related questions and statements to assist the user in addressing the primary question or statement. This handbook will guide decision makers through the important process steps of identifying appropriate questions, gathering appropriate data, developing landscape objectives, and prioritizing landscape patches where potential sites for restoration projects may be located. Once potential sites are selected, land managers can move to the site-specific decision tool to guide restoration decisions at the site level.

We assessed the effects of the litter layer, soil surface conditions, and microhabitat on emergence of the tree Cercocarpus ledifolius , and how litter affected establishment and survival of first year seedlings of this species. Natural seedling emergence was similar in three microhabitats: open interspaces (0.12kg litter m −2 ), beneath Artemisia tridentata canopies (0.27kg litter m −2 ), and beneath C. ledifolius canopies (2.5kg litter m −2 ). However, because seed dispersal is highest under trees, and lowest in the open, seedling emergence seemed to vary and have highest values in open interspaces. Emergence rates did not vary between soil surface treatments, but were lower in experimental litter treatments than in treatments, without litter, independent of microhabitat. In contrast, litter beneath trees favored subsequent seedling establishment and survival. This study shows that the role of the litter layer varies during the first year of the life of this species, with litter inhibiting initial emergence but enhancing early seedling survival.

Current paradigm suggests that spatial and temporal competition for resources limit an exotic invader, cheatgrass (Bromus tectorum L.), which once established, alters fire regimes and can result in annual grass dominance in sagebrush steppe. Prescribed fire and fire surrogate treatments (mowing, tebuthiuron, and imazapic) are used to reduce woody fuels and increase resistance to exotic annuals, but may alter resource availability and inadvertently favor invasive species. We used four study sites within the Sagebrush Steppe Treatment Evaluation Project (SageSTEP) to evaluate 1) how vegetation and soil resources were affected by treatment, and 2) how soil resources influenced native herbaceous perennial and exotic annual grass cover before and following treatment. Treatments increased resin exchangeable NH4 þ ,N O3 � ,H 2PO4 � , and K þ , with the largest increases caused by prescribed fire and prolonged by application of imazapic. Burning with imazapic application also increased the number of wet growing degree days. Tebuthiuron and imazapic reduced exotic annual grass cover, but imazapic also reduced herbaceous perennial cover when used with prescribed fire. Native perennial herbaceous species cover was higher where mean annual precipitation and soil water resources were relatively high. Exotic annual grass cover was higher where resin exchangeable H2PO4 � was high and gaps between perennial plants were large. Prescribed fire, mowing, and tebuthiuron were successful at increasing perennial herbaceous cover, but the results were often ephemeral and inconsistent among sites. Locations with sandy soil, low mean annual precipitation, or low soil water holding capacity were more likely to experience increased exotic annual grass cover after treatment, and treatments that result in slow release of resources are needed on these sites. This is one of few studies that correlate abiotic variables to native and exotic species cover across a broad geographic setting, and that demonstrates how soil resources potentially influence the outcome of management treatments.

Utah juniper (Juniperus osteosperma) is one of many plant species that produce large numbers of fruits containing parthenocarpic or otherwise empty or inviable seeds. We tested the hypothesis that production of empty fruits in this species results in reduced levels of predation on fertile seeds. In a population in west-central Utah, we estimated the proportion of fruits with filled seeds in trees suffering high levels of fruit destruction by the seed-eating bird Parus inornatus and in neighbouring trees similar in crown and fruit-crop size but suffering negligible predation. We found that the heavily attacked trees had higher proportions of filled seeds. Thus, juniper may benefit from producing fruits that contain no offspring. This is the first study to demonstrate that empty seeds may reduce predation by vertebrate seed eaters and the first to demonstrate discrimination based on seed filling at the level of whole plants.

Natural formation of treefall gaps plays an integral role in the ecological and evolutionary dynamics of many tropical forests, affecting the spatiotemporal distribution of plants and the animals that interact with them. This study examines the impact of treefall gaps on the spatial and temporal patchiness of ant assemblages in a moist lowland forest in Panama. Using pitfall traps and honey baits, we compared ant assemblages in five 1 to 2-year-old treefall gaps (ca 100 m2) and five adjacent plots (ca 100 m2) in undisturbed forest understory at three different times of year (late wet season, late dry season, and early wet season). We found little evidence that ant assemblages respond dramatically to the formation of treefall gaps and the differences in habitat qualities they produce. Ant abundance, species richness, species composition, and rates of resource discovery did not differ between gaps and forest understory. However, we did find significant differences in numerical abundance related to forest stratum (ground vs vegetation) and resource type in pitfall traps (oil-cockroach vs honey), and significant differences in ant species richness and rates of resource discovery across seasons. While habitat effects by themselves were never statistically significant, habitat and seasonal differences in species richness interacted significantly to produce complex, season-dependent differences among gap and forest habitats. These results suggest that the formation of natural treefall gaps has less of an effect on Neotropical ant assemblages compared to other groups of organisms (e.g., plants, birds) or other causes of patchiness (e.g., ant mosaics, moisture availability, army ant predation). The results of our study also have important implications for the underlying causes of habitat differences in the distribution of ant-defended plants.

In this study, we explore the reproductive biology of Cercocarpus ledifolius (Rosaceae), a species showing extremely limited recruitment throughout its range in the Intermountain West, USA. We examined (1) self-compatibility, (2)the potential for wind pollination, (3) pollen limitation of seed production, and (4)loss of developing seeds. Through hand-pollination experiments, we determined that C. ledifolius flowers are highly self-compatible. Abundant seeds were produced from flowers pollinated with pollen from the same plant, and the level of seed production did not differ from control outcrossed treatments. Furthermore, we measured substantial airborne pollen loads within C. ledifolius stands, indicating transport of pollen via wind and a strong potential for wind pollination, despite apparent adaptations for insect pollination. Pollen did not limit seed production; branches on which all flowers were handpollinated with outcrossed pollen did not have greater seed production than control branches open only to natural pollination. Lastly, virtually all initiated seeds developed to maturity, indicating low seed abortion rates and minimal loss to pre-dispersal seed predators. Natural levels of fruit production were at or greater than mean fruit production for woody perennials. In summary, we found no evidence that the quantity of pollen, the source of pollen (self vs. outcrossed), or developing seed loss is seriously limiting recruitment of this species.

The ecological forces determining where within a landscape plants recruit and grow has been termed proximal habitat choice. Habitat choice is imposed first by a heterogeneous pattern of seed dispersal across the patches that make up the landscape and second by environmental variation that favors plant survival in some patches more than in others. Seedseedling conflicts can occur during both of these phases of habitat choice if conditions or traits that are favorable for seeds are unfavorable for seedlings. During the dispersal phase, smaller seeds may have a greater probability of dispersal than larger seeds, and thus a greater probability of escape from predation, but they contain fewer reserves for support of the establishing seedling. After dispersal, environmental characteristics of a given patch type that lead to disproportionately high seed survival may lead to disproportionately low seedling survival. Considering three hypothetical landscapes, each composed of five patch types, I demonstrate that seed-seedling conflicts can have a major impact on both the overall quantity of recruitment at the landscape level and on the distribution of recruitment among patches. Available empirical evidence suggests these conflicts may be widespread in natural systems. Given their potential importance and extent, seed-seedling conflicts may play a previously unrecognized role in habitat choice.

Seed dispersal determines the conditions that seeds, seedlings, and saplings confront, and thus potentially provides a unifying theme for plant population ecology. This potential is seldom realized...