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Negative effects of an allelopathic invader on AM fungal plant species drive community‐level responses

The mechanisms causing invasive species impact are rarely empirically tested, limiting our ability to understand and predict subsequent changes in invaded plant communities. Invader disruption of native mutualistic interactions is a mechanism expected to have negative effects on native plant species. Specifically, disruption of native plant‐fungal mutualisms may provide non‐mycorrhizal plant invaders an advantage over mycorrhizal native plants. Invasive Alliaria petiolata (garlic mustard) produces secondary chemicals toxic to soil microorganisms including mycorrhizal fungi, and is known to induce physiological stress and reduce population growth rates of native forest understory plant species. Here, we report on a 11‐yr manipulative field experiment in replicated forest plots testing if the effects of removal of garlic mustard on the plant community support the mutualism disruption hypothesis within the entire understory herbaceous community. We compare community responses for two functional groups: the mycorrhizal vs. the non‐mycorrhizal plant communities. Our results show that garlic mustard weeding alters the community composition, decreases community evenness, and increases the abundance of understory herbs that associate with mycorrhizal fungi. Conversely, garlic mustard has no significant effects on the non‐mycorrhizal plant community. Consistent with the mutualism disruption hypothesis, our results demonstrate that allelochemical producing invaders modify the plant community by disproportionately impacting mycorrhizal plant species. We also demonstrate the importance of incorporating causal mechanisms of biological invasion to elucidate patterns and predict community‐level responses.

Ecology↗

Soil properties and perceived disturbance of grasslands subjected to mechanized military training: Evaluation of an index

Mechanized maneuver training impacts the landscape by creating depressions, compacting soils, producing bare ground areas, transporting seeds of invasive plants, and crushing vegetation. We measured 3 physical, 13 chemical, and 2 biological soil properties and used a disturbance index (DI) based on perceptions of soil conditions on a military installation to assess the condition of 100 ?? 100 m plots (1 ha): 10 in 2002 and 10 in 2004. Potential DI scores range from 0 (no appreciable evidence of disturbance) to 1 (>95 per cent of the plot disturbed). Bulk density, porosity (%), and water content (%) - all at 5-1-10-0cm depth, and nematode family richness (NFR) were significantly, negatively correlated (Spearman coefficients, rs) with the DI of both years. The strong negative correlation (rs., = -0.69 in 2002, -0.79 in 2004) of NFR with the DI appears to reflect the status of nematode diversity and, therefore, may serve as a useful, inexpensive approach to rapidly assessing grasslands subjected to mechanized military training. Copyright ?? 2007 John Wiley & Sons, Ltd.

Land Degradation and Development↗

Introduction: Exotic annual Bromus in the western USA

The spread and impacts of exotic species are unambiguous, global threats to many ecosystems. A prominent example is the suite of annual grasses in the Bromus genus ( Bromus hereafter) that originate from Europe and Eurasia but have invaded or are invading large areas of the Western USA. This book brings a diverse, multidisciplinary group of authors together to synthesize current knowledge, research needs, and management implications for Bromus . Exotic plant invasions are multifaceted problems, and understanding and managing them requires the biological, ecological, sociological, and economic perspectives that are integrated in this book. Knowing how well information from one geographic or environmental setting can transfer to another is a key need for broadly distributed Bromus species especially given ongoing climate change. Thus, the chapters in the book compare and contrast invasibility of different ecoregions and invasiveness of different Bromus species. A universal theme is managing for ecosystems that are resilient to disturbance and resistant to invasion which will be essential for adaptation to the human-caused problem of Bromus in the Western USA.

Book chapter↗

Fire and grazing influence site resistance to Bromus tectorum through their effects on shrub, bunchgrass and biocrust communities in the Great Basin (USA)

Shrubs, bunchgrasses and biological soil crusts (biocrusts) are believed to contribute to site resistance to plant invasions in the presence of cattle grazing. Although fire is a concomitant disturbance with grazing, little is known regarding their combined impacts on invasion resistance. We are the first to date to test the idea that biotic communities mediate the effects of disturbance on site resistance. We assessed cover of Bromus tectorum , shrubs, native bunchgrasses, lichens and mosses in 99 burned and unburned plots located on similar soils where fires occurred between 12 and 23 years before sampling. Structural equation modeling was used to test hypothesized relationships between environmental and disturbance characteristics, the biotic community and resistance to B. tectorum cover. Characteristics of fire and grazing did not directly relate to cover of B. tectorum. Relationships were mediated through shrub, bunchgrass and biocrust communities. Increased site resistance following fire was associated with higher bunchgrass cover and recovery of bunchgrasses and mosses with time since fire. Evidence of grazing was more pronounced on burned sites and was positively correlated with the cover of B. tectorum , indicating an interaction between fire and grazing that decreases site resistance. Lichen cover showed a weak, negative relationship with cover of B. tectorum . Fire reduced near-term site resistance to B. tectorum on actively grazed rangelands. Independent of fire, grazing impacts resulted in reduced site resistance to B. tectorum , suggesting that grazing management that enhances plant and biocrust communities will also enhance site resistance.

Great Basin↗

Diel biogeochemical processes in terrestrial waters

Many biogeochemical processes in rivers and lakes respond to the solar photocycle and produce persistent patterns of measureable phenomena that exhibit a day–night, or 24-h, cycle. Despite a large body of recent literature, the mechanisms responsible for these diel fluctuations are widely debated, with a growing consensus that combinations of physical, chemical, and biological processes are involved. These processes include streamflow variation, photosynthesis and respiration, plant assimilation, and reactions involving photochemistry, adsorption and desorption, and mineral precipitation and dissolution. Diel changes in streamflow and water properties such as temperature, pH, and dissolved oxygen concentration have been widely recognized, and recently, diel studies have focused more widely by considering other constituents such as dissolved and particulate trace metals, metalloids, rare earth elements, mercury, organic matter, dissolved inorganic carbon (DIC), and nutrients. The details of many diel processes are being studied using stable isotopes, which also can exhibit diel cycles in response to microbial metabolism, photosynthesis and respiration, or changes in phase, speciation, or redox state. In addition, secondary effects that diel cycles might have, for example, on biota or in the hyporheic zone are beginning to be considered. This special issue is composed primarily of papers presented at the topical session “Diurnal Biogeochemical Processes in Rivers, Lakes, and Shallow Groundwater” held at the annual meeting of the Geological Society of America in October 2009 in Portland, Oregon. This session was organized because many of the growing number of diel studies have addressed just a small part of the full range of diel cycling phenomena found in rivers and lakes. This limited focus is understandable because (1) fundamental aspects of many diel processes are poorly understood and require detailed study, (2) the interests and expertise of individual scientists typically do not encompass the wide diversity and range of processes that produce diel cycles, and (3) the logistics of making field measurements for 24-h periods has limited recognition and understanding of these important cycles. Thus, the topical session brought together hydrologists, biologists, geochemists, and ecologists to discuss field studies, laboratory experiments, theoretical modeling, and measurement techniques related to diel cycling. Hopefully with the cross-disciplinary synergy developed at the session as well as by this special issue, a more comprehensive understanding of the interrelationships between the diel processes will be developed. Needless to say, understanding diel processes is critical for regulatory agencies and the greater scientific community. And perhaps more importantly, expanded knowledge of biogeochemical cycling may lead to better predictions of how aquatic ecosystems might react to changing conditions of contaminant loading, eutrophication, climate change, drought, industrialization, development, and other variables.

Chemical Geology↗

Beyond theories of plant invasions: Lessons from natural landscapes

There are a growing number of contrasting theories about plant invasions, but most are only weakly supported by small-scale field experiments, observational studies, and mathematical models. Among the most contentious theories is that species-rich habitats should be less vulnerable to plant invasion than species-poor sites, stemming from earlier theories that competition is a major force in structuring plant communities. Early ecologists such as Charles Darwin (1859) and Charles Elton (1958) suggested that a lack of intense interspecific competition on islands made these low-diversity habitats vulnerable to invasion. Small-scale field experiments have supported and contradicted this theory, as have various mathematical models. In contrast, many large-scale observational studies and detailed vegetation surveys in continental areas often report that species-rich areas are more heavily invaded than species-poor areas, but there are exceptions here as well. In this article, I show how these seemingly contrasting patterns converge once appropriate spatial and temporal scales are considered in complex natural environments. I suggest ways in which small-scale experiments, mathematical models, and large- scale observational studies can be improved and better integrated to advance a theoretically based understanding of plant invasions.

Comments on Theoretical Biology↗

Lead in the environment

Anthropogenic uses of lead have probably altered its availability and environmental distribution more than any other toxic element. Consequently, lead concentrations in many living organisms may be approaching thresholds of toxicity for the adverse effects of lead. Such thresholds are difficult to define, as they vary with the chemical and physical form of lead, exposure regime, other elements present and also vary both within and between species. The technological capability to accurately quantify low lead concentrations has increased over the last decade, and physiological and behavioral effects have been measured in wildlife with tissue lead concentrations below those previously considered safe for humans.s.236 Consequently. lead criteria for the protection of wildlife and human health are frequently under review, and 'thresholds' of lead toxicity are being reconsidered. Proposed lead criteria for the protection of natural resources have been reviewed by Eisler. Uptake of lead by plants is limited by its generally low availability in soils and sediments, and toxicity may be limited by storage mechanisms and its apparently limited translocation within most plants. Lead does not generally accumulate within the foliar parts of plants, which limits its transfer to higher trophic levels. Although lead may concentrate in plant and animal tissues, no evidence of biomagnification exists. Acid deposition onto surface waters and soils with low buffering capacity may influence the availability of lead for uptake by plants and animals, and this may merit investigation at susceptible sites. The biological significance of chronic low-level lead exposure to wildlife is sometimes difficult to quantify. Animals living in urban environments or near point sources of lead emission are inevitably subject to greater exposure to lead and enhanced risk of lead poisoning. Increasingly strict controls on lead emissions in many countries have reduced exposure to lead from some sources. and the .reduction of lead in gasoline has resulted in lower tissue lead concentrations in humans and wildlife from many, particularly urban, locations. However, it has been suggested that increasing use of organic lead compounds as catalysts for the production of plastics and as wood preservatives and biocides could adversely affect wildlife. The most significant source of direct wildlife mortality from lead is spent gunshot and fishing sinkers. Elevated mortality from shot ingestion in avian species resulted in the introduction of nontoxic (steel) shot zones along certain flyways in the United States in the mid-1970s and a total ban on the use of lead for waterfowl and coot hunting nationwide by 1992. Several other countries are now following suit and have either banned or are in the process of restricting the use of lead shot for waterfowl hunting. In the United States it has been estimated that since the 1986 hunting season. when the use of nontoxic shot became widespread. over 6 million ducks have not been lost to lead poisoning. Raptors, especially eagles, have also apparently benefited. although lead poisoning from ingestion of bullet fragments remains a problem for the critically threatened California condor. Quantifying reductions in lead mortality rates would be difficult since eagle populations throughout North America are rapidly recovering from other anthropogenic perturbations, especially organochlorine pesticides.

Book chapter↗

Control of invasive weeds with prescribed burning

Prescribed burning has primarily been used as a tool for the control of invasive late-season annual broadleaf and grass species, particularly yellow starthistle, medusahead, barb goatgrass, and several bromes. However, timely burning of a few invasive biennial broadleaves (e.g., sweetclover and garlic mustard), perennial grasses (e.g., bluegrasses and smooth brome), and woody species (e.g., brooms and Chinese tallow tree) also has been successful. In many cases, the effectiveness of prescribed burning can be enhanced when incorporated into an integrated vegetation management program. Although there are some excellent examples of successful use of prescribed burning for the control of invasive species, a limited number of species have been evaluated. In addition, few studies have measured the impact of prescribed burning on the long-term changes in plant communities, impacts to endangered plant species, effects on wildlife and insect populations, and alterations in soil biology, including nutrition, mycorrhizae, and hydrology. In this review, we evaluate the current state of knowledge on prescribed burning as a tool for invasive weed management.

Weed Technology↗

Invasive Species Science Branch: research and management tools for controlling invasive species

Invasive, nonnative species of plants, animals, and disease organisms adversely affect the ecosystems they enter. Like “biological wildfires,” they can quickly spread and affect nearly all terrestrial and aquatic ecosystems. Invasive species have become one of the greatest environmental challenges of the 21st century in economic, environmental, and human health costs, with an estimated effect in the United States of more than $120 billion per year. Managers of the Department of the Interior and other public and private lands often rank invasive species as their top resource management problem. The Invasive Species Science Branch of the Fort Collins Science Center provides research and technical assistance relating to management concerns for invasive species, including understanding how these species are introduced, identifying areas vulnerable to invasion, forecasting invasions, and developing control methods. To disseminate this information, branch scientists are developing platforms to share invasive species information with DOI cooperators, other agency partners, and the public. From these and other data, branch scientists are constructing models to understand and predict invasive species distributions for more effective management. The branch also has extensive herpetological and population biology expertise that is applied to harmful reptile invaders such as the Brown Treesnake on Guam and Burmese Python in Florida.

Fact Sheet↗

High severity fire in forests of the southwest: Conservation implications. Progress Report August 2005

The occurrence of large, severe fires in southwestern ponderosa pine (Pinus ponderosa) forests has resulted in concern that these forests may not persist under such an extreme disturbance regime. In our research, we are examining the outcomes of high-severity fire in ponderosa pine forests and their neighboring communities across an elevational gradient. One goal of our work is to contribute to understanding the resiliency of these systems, but we also want to investigate the conservation values intrinsic to the diverse communities that represent alternative successional trajectories after severe fire. One assumption of our research is that the spatial pattern of a disturbance becomes increasingly important when the disturbance is large and biological legacies are few and sparse. We ask, therefore, what spectrum of plant communities results from high severity fire, and what is their relationship to spatial patterns of severity mapped in early post fire timeframes? Also, do spatial patterns of older burns (1950–80) differ from recent burns (1998–present) in ways that make us expect successional changes years from now to differ from those we observed at our older burn field sites? Here, we describe the first stages of our work in mapping burn severity at old and new burns as well as the work we have recently completed at our two field sites. The report is organized under our two main objectives with the purpose of summarizing the steps we have taken in working toward these objectives, as well as changes we have made in methodologies since the original study plan. We present some general observations and plans for the next steps in data analysis and product generation. This report, the study plan, a photograph gallery, slide presentations, and our contact information are available on the project Web site http://www.umass.edu/landeco/research/swfire/swfire.html .

Open-File Report↗

Klamath Mountains Ecoregion: Chapter 13 in Status and trends of land change in the Western United States--1973 to 2000

The Klamath Mountains Ecoregion covers approximately 47,791 km 2 (18,452 mi 2 ) of the Klamath and Siskiyou Mountains of northern California and southern Oregon (fig. 1) (Omernik, 1987; U.S. Environmental Protection Agency, 1997). The ecoregion is flanked by the Coast Range Ecoregion to the west, the Southern and Central California Chaparral and Oak Woodlands Ecoregion to the south, the Cascades and the Eastern Cascades Slopes and Foothills Ecoregions to the east, and the Willamette Valley Ecoregion to the north. The mild Mediterranean climate of the ecoregion is characterized by hot, dry summers and wet winters; the amount of winter moisture varies within the ecoregion, decreasing from west to east. The Klamath–Siskiyou Mountains region is widely recognized as an important biodiversity hotspot (Whittaker, 1960; Kruckeberg, 1984; Wagner, 1997; DellaSala and others, 1999), containing more than 3,500 plant species, more than 200 of which are endemic (Sawyer, 2007). A biological assessment by DellaSala and others (1999) ranked the Klamath–Siskiyou Mountains region as the fifth richest coniferous forest in terms of species diversity. In addition, the International Union for the Conservation of Nature considers the region an area of notable botanical importance (Wagner, 1997). Twenty-nine different species of conifers can be found in the Klamath Mountains Ecoregion (Sawyer, 1996).

California;Oregon↗

Prior knowledge-based approach for associating contaminants with biological effects: A case study in the St. Croix River basin, MN, WI, USA

Evaluating potential adverse effects of complex chemical mixtures in the environment is challenging. One way to address that challenge is through more integrated analysis of chemical monitoring and biological effects data. In the present study, water samples from five locations near two municipal wastewater treatment plants in the St. Croix River basin, on the border of MN and WI, USA, were analyzed for 127 organic contaminants. Known chemical-gene interactions were used to develop site-specific knowledge assembly models (KAMs) and formulate hypotheses concerning possible biological effects associated with chemicals detected in water samples from each location. Additionally, hepatic gene expression data were collected for fathead minnows ( Pimephales promelas ) exposed in situ , for 12 d, at each location. Expression data from oligonucleotide microarrays were analyzed to identify functional annotation terms enriched among the differentially-expressed probes. The general nature of many of the terms made hypothesis formulation on the basis of the transcriptome-level response alone difficult. However, integrated analysis of the transcriptome data in the context of the site-specific KAMs allowed for evaluation of the likelihood of specific chemicals contributing to observed biological responses. Thirteen chemicals (atrazine, carbamazepine, metformin, thiabendazole, diazepam, cholesterol, p-cresol, phenytoin, omeprazole, ethyromycin, 17β-estradiol, cimetidine, and estrone), for which there was statistically significant concordance between occurrence at a site and expected biological response as represented in the KAM, were identified. While not definitive, the approach provides a line of evidence for evaluating potential cause-effect relationships between components of a complex mixture of contaminants and biological effects data, which can inform subsequent monitoring and investigation.

Minnesota, Wisconsin↗

Biocrust and seed placement influence emergence rates of perennial grass Elymus elymoides across five North American deserts

1. Dryland vascular plant emergence is often limited by water availability especially with projected climate and precipitation changes. Biological soil crusts (biocrusts) can promote water retention and nutrient availability that benefit germination, and emergence yet can also act as a surface barrier preventing critical seed soil contact and hindering emergence. Alongside these factors, dryland fire frequency has increased with the inclusion of invasive species and vegetation structural changes. With enhanced continuous fine fuel distribution there is a high potential to disrupt biocrust-plant interactions and influence associated management actions. 2. This study explores the dynamic relationship between biocrusts and fire-related heating effects on seedling emergence across five North American deserts: the Chihuahuan Desert, Colorado Plateau, Great Basin, Mojave Desert and Sonoran Desert. We conducted a greenhouse-based seedling emergence experiment using Elymus elymoides (bottlebrush squirreltail), a common perennial grass, with biocrust and bare soil mesocosms in which half were heated to mimic the effects of wildfire temperature. 3. The variables that had the greatest influence on germination rate and germination timing were the presence of biocrust and seed placement (on top of vs within the biocrust/soil matrix). Emergence rate was greatest atop bare soil followed by seeds inserted into biocrust. Emergence timing was faster with biocrust present, but only when seeds were inserted into it. Desert origin of biocrust and soil collection also influenced germination where the probability of any one seed emerging was highest in the Chihuahuan and Mojave desert sites relative to the Sonoran desert site which showed the lowest probability. Heating had mixed effects whereby it increased the likelihood of emergence but did not affect the overall rate or timing. 4. Synthesis and applications . This study underscores the importance of healthy and impaired biocrusts in dryland systems and suggests ways in which the combination of biocrust and seed placement can influence plant establishment, in addition to providing insight into seeding strategies for managers and restoration practitioners working in dryland sites.

Arizona, California, Colorado, Nevada, New Mexico↗

An adaptive decision framework for the conservation of a threatened plant

Mead's milkweed Asclepias meadii , a long-lived perennial herb of tallgrass prairie and glade communities of the central United States, is a species designated as threatened under the U.S. Endangered Species Act. Challenges to its successful management include the facts that much about its life history is unknown, its age at reproductive maturity is very advanced, certain life stages are practically unobservable, its productivity is responsive to unpredictable environmental events, and most of the known populations occur on private lands unprotected by any legal conservation instrument. One critical source of biological uncertainty is the degree to which fire promotes growth and reproductive response in the plant. To aid in its management, we developed a prototype population-level state-dependent decision-making framework that explicitly accounts for this uncertainty and for uncertainties related to stochastic environmental effects and vital rates. To parameterize the decision model, we used estimates found in the literature, and we analyzed data from a long-term monitoring program where fates of individual plants were observed through time. We demonstrate that different optimal courses of action are followed according to how one believes that fire influences reproductive response, and we show that the action taken for certain population states is informative for resolving uncertainty about competing beliefs regarding the effect of fire. We advocate the use of a model-predictive approach for the management of rare populations, particularly when management uncertainty is profound. Over time, an adaptive management approach should reduce uncertainty and improve management performance as predictions of management outcome generated under competing models are continually informed and updated by monitoring data.

Journal of Fish and Wildlife Management↗

Small mammals as indicators of short-term and long-term disturbance in mixed prairie

Disturbance by military maneuvers over short and long time scales may have differential effects on grassland communities. We assessed small mammals as indicators of disturbance by military maneuvers in a mixed prairie in southern Oklahoma USA. We examined sites on two soil series, Foard and Lawton, across a gradient of disturbance intensity. A MANOVA showed that abundance of small mammals was associated (p = 0.03) with short-term (cover of vehicle tracks) disturbance but was not associated (p = 0.12) with long-term (loss of soil organic carbon, SOC) disturbance intensity. At the individual species level, Sigmodon hispidus (cotton rat) and Peromyscus maniculatus (deer mouse) occurred across all levels of disturbance and in both soil types. Only P. maniculatus abundance changed (p < 0.01) with short-term disturbance and increased by about one individual per 5% of additional track-cover. Abundance of P. maniculatus also increased (p = 0.04) by about three individuals per 1% increase in soil carbon. Chaetodipus hispidus (hispid pocket mouse) and Reithrodontomys fulvescens (fulvous harvest mouse) only occurred in single soil types limiting their potential as more general indicators. Abundance of P. maniculatus was positively related to shifts in plant species composition and likely reflected changes in vegetation structure (i.e. litter depth) and forage availability resulting from disturbance. Peromyscus maniculatus may be a useful biological indicator of ecosystem change because it responded predictably to both long-term and short-term disturbance and, when coupled with soil, plant, and disturbance history variables, can reveal land condition trends. ?? Springer Science+Business Media B.V. 2007.

Environmental Monitoring and Assessment↗

Biodiversity influences plant productivity through niche–efficiency

The loss of biodiversity is threatening ecosystem productivity and services worldwide, spurring efforts to quantify its effects on the functioning of natural ecosystems. Previous research has focused on the positive role of biodiversity on resource acquisition (i.e., niche complementarity), but a lack of study on resource utilization efficiency, a link between resource and productivity, has rendered it difficult to quantify the biodiversity–ecosystem functioning relationship. Here we demonstrate that biodiversity loss reduces plant productivity, other things held constant, through theory, empirical evidence, and simulations under gradually relaxed assumptions. We developed a theoretical model named niche–efficiency to integrate niche complementarity and a heretofore-ignored mechanism of diminishing marginal productivity in quantifying the effects of biodiversity loss on plant productivity. Based on niche–efficiency, we created a relative productivity metric and a productivity impact index (PII) to assist in biological conservation and resource management. Relative productivity provides a standardized measure of the influence of biodiversity on individual productivity, and PII is a functionally based taxonomic index to assess individual species’ inherent value in maintaining current ecosystem productivity. Empirical evidence from the Alaska boreal forest suggests that every 1% reduction in overall plant diversity could render an average of 0.23% decline in individual tree productivity. Out of the 283 plant species of the region, we found that large woody plants generally have greater PII values than other species. This theoretical model would facilitate the integration of biological conservation in the international campaign against several pressing global issues involving energy use, climate change, and poverty.

Proceedings of the National Academy of Sciences of↗

Developing terrestrial, multi-taxon indices of biological integrity: An example from coastal sage scrub

We screened 351 species or genera for their response to disturbance in coastal sage scrub (CSS) to develop a 15-metric, 5-taxon Index of Biological Integrity (IBI). We collected data on ants, birds, herpetofauna, small mammals, and plants for two years on 46 sites established across a gradient of disturbance in three reserves. The gradient spanned relatively intact CSS with thick stands of shrubs, to former CSS stands type-converted to exotic grasses. ANOVAs and clustering analyses indicated the IBI could distinguish four levels of disturbance in CSS. General measures of community structure, such as richness, did not show changes across the gradient for most taxa, and responses of taxa across the gradient were varied and rarely correlated. However, turnover in species or genera across the gradient was common across all taxa as shrub-obligate life forms were replaced by those favoring grassy or disturbed habitats. Our data indicate index-based approaches based on data collected across disturbance gradients may outperform more traditional community level metrics when responses to anthropogenic influences are complex and vary across species. ?? 2007 Elsevier Ltd. All rights reserved.

Biological Conservation↗

Emergence of ratio-dependent and predator-dependent functional responses for pollination mutualism and seed parasitism

Prey ( N ) dependence [ g ( N )], predator ( P ) dependence [ g ( P ) or g ( N , P )], and ratio dependence [ f ( P / N )] are often seen as contrasting forms of the predator's functional response describing predator consumption rates on prey resources in predator&ndash;prey and parasitoid&ndash;host interactions. Analogously, prey-, predator-, and ratio-dependent functional responses are apparently alternative functional responses for other types of consumer&ndash;resource interactions. These include, for example, the fraction of flowers pollinated or seeds parasitized in pollination (pre-dispersal) seed-parasitism mutualisms, such as those between fig wasps and fig trees or yucca moths and yucca plants. Here we examine the appropriate functional responses for how the fraction of flowers pollinated and seeds parasitized vary with the density of pollinators (predator dependence) or the ratio of pollinator and flower densities (ratio dependence). We show that both types of functional responses can emerge from minor, but biologically important variations on a single model. An individual-based model was first used to describe plant&ndash;pollinator interactions. Conditional upon on whether the number of flowers visited by the pollinator was limited by factors other than search time (e.g., by the number of eggs it had to lay, if it was also a seed parasite), and on whether the pollinator could directly find flowers on a plant, or had to search, the simulation results lead to either a predator-dependent or a ratio-dependent functional response. An analytic model was then used to show mathematically how these two cases can arise.

Ecological Modelling↗