A comparison of population viability measures.

The viability of populations can be quantified with several measures, such as the probability of extinction, the mean time to extinction, or the population size. While conservation management decisions can be based on these measures, it has not yet been explored systematically if different viability measures rank species and scenarios similarly and if one viability measure can be converted into another to compare studies. To address this challenge, we conducted a quantitative comparison of eight viability measures based on the simulated population dynamics of more than 4500 virtual species. We compared (a) the ranking of scenarios based on different viability measures, (b) assessed direct correlations between the measures, and (c) explored if parameters in the simulation models can alter the relationship between pairs of viability measures. We found that viability measures ranked species similarly. Despite this, direct correlations between the different measures were often weak and could not be generalized. This can be explained by the loss of information due to the aggregation of raw data into a single number, the effect of model parameters on the relationship between viability measures, and because distributions, such as the probability of extinction over time, cannot be ranked objectively. Similar scenario rankings by different viability measures show that the choice of the viability metric does in many cases not alter which population is regarded more viable or which management option is the best. However, the more two scenarios or populations differ, the more likely it becomes that different measures produce different rankings. We thus recommend that PVA studies publish raw simulation data, which not only describes all risks and opportunities to the reader but also facilitates meta-analyses of PVA studies.

EFForTS-LGraf: A landscape generator for creating smallholder-driven land-use mosaics.

Spatially-explicit simulation models are commonly used to study complex ecological and socio-economic research questions. Often these models depend on detailed input data, such as initial land-cover maps to set up model simulations. Here we present the landscape generator EFFortS-LGraf that provides artificially-generated land-use maps of agricultural landscapes shaped by small-scale farms. EFForTS-LGraf is a process-based landscape generator that explicitly incorporates the human dimension of land-use change. The model generates roads and villages that consist of smallholder farming households. These smallholders use different establishment strategies to create fields in their close vicinity. Crop types are distributed to these fields based on crop fractions and specialization levels. EFForTS-LGraf model parameters such as household area or field size frequency distributions can be derived from household surveys or geospatial data. This can be an advantage over the abstract parameters of neutral landscape generators. We tested the model using oil palm and rubber farming in Indonesia as a case study and validated the artificially-generated maps against classified satellite images. Our results show that EFForTS-LGraf is able to generate realistic land-cover maps with properties that lie within the boundaries of landscapes from classified satellite images. An applied simulation experiment on landscape-level effects of increasing household area and crop specialization revealed that larger households with higher specialization levels led to spatially more homogeneous and less scattered crop type distributions and reduced edge area proportion. Thus, EFForTS-LGraf can be applied both to generate maps as inputs for simulation modelling and as a stand-alone tool for specific landscape-scale analyses in the context of ecological-economic studies of smallholder farming systems.

The role of biotic factors during plant establishment in novel communities assessed with an agent-based simulation model.

BACKGROUND: Establishment success of non-native species is not only influenced by environmental conditions, but also by interactions with local competitors and enemies. The magnitude of these biotic interactions is mediated by species traits that reflect competitive strength or defence mechanisms. Our aim was to investigate the importance of species traits for successful establishment of non-native species in a native community exhibiting biotic resistance in the form of competition and herbivory. METHODS: We developed a trait-based, individual-based simulation model tracking the survival of non-native plants in a native community. In the model, non-native plants are characterized by high or low values of competition and defence traits. Model scenarios included variation of initial number of non-natives, intensity of competitive interaction, density of herbivores and density as well as mixture of the native community. RESULTS: Traits related to competition had a much greater impact on survival of non-native species than traits related to defence. Survival rates of strong competitors never fell below 50% while survival of weak competitors averaged at about 10%. Weak competitors were also much more susceptible to competitive pressures such as community density, composition and competition intensity. Strong competitors responded negatively to changes in competition intensity, but hardly to composition or density of the native community. High initial numbers of non-native individuals decreased survival rate of strong competitors, but increased the survival rate of weak competitors. Survival under herbivore attack was only slightly higher for plants with high defensive ability than for those with low defensive ability. Surprisingly, though, herbivory increased survival of species classified as weak competitors. DISCUSSION: High survival rates of strong non-native competitors relate to a higher probability of successful establishment than for weak competitors. However, the reduced survival of strong competitors at high initial numbers indicates a self-thinning effect, probably mediated by a strongly competitive milieu. For weak competitors, our model emphasizes positive effects of high propagule pressure known from field studies. General effects of herbivory or defence abilities on survival were not supported by our model. However, the positive effect of herbivory on survival of weak competitors indicated side effects of herbivory, such as weakening resident competitors. This might play an important role for establishment of non-natives in a new community.

Land-use change in oil palm dominated tropical landscapes-An agent-based model to explore ecological and socio-economic trade-offs.

Land-use changes have dramatically transformed tropical landscapes. We describe an ecological-economic land-use change model as an integrated, exploratory tool used to analyze how tropical land-use change affects ecological and socio-economic functions. The model analysis seeks to determine what kind of landscape mosaic can improve the ensemble of ecosystem functioning, biodiversity, and economic benefit based on the synergies and trade-offs that we have to account for. More specifically, (1) how do specific ecosystem functions, such as carbon storage, and economic functions, such as household consumption, relate to each other? (2) How do external factors, such as the output prices of crops, affect these relationships? (3) How do these relationships change when production inefficiency differs between smallholder farmers and learning is incorporated? We initialize the ecological-economic model with artificially generated land-use maps parameterized to our study region. The economic sub-model simulates smallholder land-use management decisions based on a profit maximization assumption. Each household determines factor inputs for all household fields and decides on land-use change based on available wealth. The ecological sub-model includes a simple account of carbon sequestration in above-ground and below-ground vegetation. We demonstrate model capabilities with results on household consumption and carbon sequestration from different output price and farming efficiency scenarios. The overall results reveal complex interactions between the economic and ecological spheres. For instance, model scenarios with heterogeneous crop-specific household productivity reveal a comparatively high inertia of land-use change. Our model analysis even shows such an increased temporal stability in landscape composition and carbon stocks of the agricultural area under dynamic price trends. These findings underline the utility of ecological-economic models, such as ours, to act as exploratory tools which can advance our understanding of the mechanisms underlying the trade-offs and synergies of ecological and economic functions in tropical landscapes.

A review of the ecosystem functions in oil palm plantations, using forests as a reference system.

Oil palm plantations have expanded rapidly in recent decades. This large-scale land-use change has had great ecological, economic, and social impacts on both the areas converted to oil palm and their surroundings. However, research on the impacts of oil palm cultivation is scattered and patchy, and no clear overview exists. We address this gap through a systematic and comprehensive literature review of all ecosystem functions in oil palm plantations, including several (genetic, medicinal and ornamental resources, information functions) not included in previous systematic reviews. We compare ecosystem functions in oil palm plantations to those in forests, as the conversion of forest to oil palm is prevalent in the tropics. We find that oil palm plantations generally have reduced ecosystem functioning compared to forests: 11 out of 14 ecosystem functions show a net decrease in level of function. Some functions show decreases with potentially irreversible global impacts (e.g. reductions in gas and climate regulation, habitat and nursery functions, genetic resources, medicinal resources, and information functions). The most serious impacts occur when forest is cleared to establish new plantations, and immediately afterwards, especially on peat soils. To variable degrees, specific plantation management measures can prevent or reduce losses of some ecosystem functions (e.g. avoid illegal land clearing via fire, avoid draining of peat, use of integrated pest management, use of cover crops, mulch, and compost) and we highlight synergistic mitigation measures that can improve multiple ecosystem functions simultaneously. The only ecosystem function which increases in oil palm plantations is, unsurprisingly, the production of marketable goods. Our review highlights numerous research gaps. In particular, there are significant gaps with respect to socio-cultural information functions. Further, there is a need for more empirical data on the importance of spatial and temporal scales, such as differences among plantations in different environments, of different sizes, and of different ages, as our review has identified examples where ecosystem functions vary spatially and temporally. Finally, more research is needed on developing management practices that can offset the losses of ecosystem functions. Our findings should stimulate research to address the identified gaps, and provide a foundation for more systematic research and discussion on ways to minimize the negative impacts and maximize the positive impacts of oil palm cultivation.

Root-Lesion Nematodes Suppress Cabbage Aphid Population Development by Reducing Aphid Daily Reproduction.

Empirical studies have shown that belowground feeding herbivores can affect the performance of aboveground herbivores in different ways. Often the critical life-history parameters underlying the observed performance effects remain unexplored. In order to better understand the cause for the observed effects on aboveground herbivores, these ecological mechanisms must be better understood. In this study we combined empirical experiments with a modeling approach to analyze the effect of two root feeding endoparasitic nematodes with different feeding strategies on the population growth of the aboveground feeding specialist aphid Brevicoryne brassicae on Brassica nigra. The aim was to test whether emerging differences in life history characteristics (days until reproduction, daily reproduction) would be sufficient to explain observed differences in aphid population development on plants with and without two species of nematodes. Aphid numbers were lower on plants with Pratylenchus penetrans in comparison to aphid numbers on plants with Meloidogyne spp. A dedicated experiment showed that aphid daily reproduction was lower on plants with P. penetrans (3.08 offspring female(-1) day(-1)) in comparison to both uninfested plants and plants with Meloidogyne spp. (3.50 offspring female(-1) day(-1)). The species-specific reduction of aphid reproduction appeared independent of changes in amino acids, soluble sugars or the glucosinolate sinigrin in the phloem. An individual-based model revealed that relatively small differences in reproduction rate per female were sufficient to yield a similar difference in aphid populations as was found in the empirical experiments.

Modeling microbial growth and dynamics.

Modeling has become an important tool for widening our understanding of microbial growth in the context of applied microbiology and related to such processes as safe food production, wastewater treatment, bioremediation, or microbe-mediated mining. Various modeling techniques, such as primary, secondary and tertiary mathematical models, phenomenological models, mechanistic or kinetic models, reactive transport models, Bayesian network models, artificial neural networks, as well as agent-, individual-, and particle-based models have been applied to model microbial growth and activity in many applied fields. In this mini-review, we summarize the basic concepts of these models using examples and applications from food safety and wastewater treatment systems. We further review recent developments in other applied fields focusing on models that explicitly include spatial relationships. Using these examples, we point out the conceptual similarities across fields of application and encourage the combined use of different modeling techniques in hybrid models as well as their cross-disciplinary exchange. For instance, pattern-oriented modeling has its origin in ecology but may be employed to parameterize microbial growth models when experimental data are scarce. Models could also be used as virtual laboratories to optimize experimental design analogous to the virtual ecologist approach. Future microbial growth models will likely become more complex to benefit from the rich toolbox that is now available to microbial growth modelers.

Spatial scales of interactions among bacteria and between bacteria and the leaf surface.

Microbial life on plant leaves is characterized by a multitude of interactions between leaf colonizers and their environment. While the existence of many of these interactions has been confirmed, their spatial scale or reach often remained unknown. In this study, we applied spatial point pattern analysis to 244 distribution patterns of Pantoea agglomerans and Pseudomonas syringae on bean leaves. The results showed that bacterial colonizers of leaves interact with their environment at different spatial scales. Interactions among bacteria were often confined to small spatial scales up to 5-20 µm, compared to interactions between bacteria and leaf surface structures such as trichomes which could be observed in excess of 100 µm. Spatial point-pattern analyses prove a comprehensive tool to determine the different spatial scales of bacterial interactions on plant leaves and will help microbiologists to better understand the interplay between these interactions.

Adaptive and selective seed abortion reveals complex conditional decision making in plants.

Behavior is traditionally attributed to animals only. Recently, evidence for plant behavior is accumulating, mostly from plant physiological studies. Here, we provide ecological evidence for complex plant behavior in the form of seed abortion decisions conditional on internal and external cues. We analyzed seed abortion patterns of barberry plants exposed to seed parasitism and different environmental conditions. Without abortion, parasite infestation of seeds can lead to loss of all seeds in a fruit. We statistically tested a series of null models with Monte Carlo simulations to establish selectivity and adaptiveness of the observed seed abortion patterns. Seed abortion was more frequent in parasitized fruits and fruits from dry habitats. Surprisingly, seed abortion occurred with significantly greater probability if there was a second intact seed in the fruit. This strategy provides a fitness benefit if abortion can prevent a sibling seed from coinfestation and if nonabortion of an infested but surviving single seed saves resources invested in the fruit coat. Ecological evidence for complex decision making in plants thus includes a structural memory (the second seed), simple reasoning (integration of inner and outer conditions), conditional behavior (abortion), and anticipation of future risks (seed predation).

Heterodera schachtii nematodes interfere with aphid-plant relations on Brassica oleracea.

Aboveground and belowground herbivore species modify plant defense responses differently. Simultaneous attack can lead to non-additive effects on primary and secondary metabolite composition in roots and shoots. We previously found that aphid (Brevicoryne brassicae) population growth on Brassica oleracea was reduced on plants that were infested with nematodes (Heterodera schachtii) prior (4 weeks) to aphid infestation. Here, we examined how infection with root-feeding nematodes affected primary and secondary metabolites in the host plant and whether this could explain the increase in aphid doubling time from 3.8 to 6.7 days. We hypothesized that the effects of herbivores on plant metabolites would depend on the presence of the other herbivore and that nematode-induced changes in primary metabolites would correlate with reduced aphid performance. Total glucosinolate concentration in the leaves was not affected by nematode presence, but the composition of glucosinolates shifted, as gluconapin concentrations were reduced, while gluconapoleiferin concentrations increased in plants exposed to nematodes. Aphid presence increased 4-methoxyglucobrassicin concentrations in leaves, which correlated positively with the number of aphids per plant. Nematodes decreased amino acid and sugar concentrations in the phloem. Aphid population doubling time correlated negatively with amino acids and glucosinolate levels in leaves, whereas these correlations were non-significant when nematodes were present. In conclusion, the effects of an herbivore on plant metabolites were independent of the presence of another herbivore. Nematode presence reduced aphid population growth and disturbed feeding relations between plants and aphids.

Testing the paradox of enrichment along a land use gradient in a multitrophic aboveground and belowground community.

In the light of ongoing land use changes, it is important to understand how multitrophic communities perform at different land use intensities. The paradox of enrichment predicts that fertilization leads to destabilization and extinction of predator-prey systems. We tested this prediction for a land use intensity gradient from natural to highly fertilized agricultural ecosystems. We included multiple aboveground and belowground trophic levels and land use-dependent searching efficiencies of insects. To overcome logistic constraints of field experiments, we used a successfully validated simulation model to investigate plant responses to removal of herbivores and their enemies. Consistent with our predictions, instability measured by herbivore-induced plant mortality increased with increasing land use intensity. Simultaneously, the balance between herbivores and natural enemies turned increasingly towards herbivore dominance and natural enemy failure. Under natural conditions, there were more frequently significant effects of belowground herbivores and their natural enemies on plant performance, whereas there were more aboveground effects in agroecosystems. This result was partly due to the "boom-bust" behavior of the shoot herbivore population. Plant responses to herbivore or natural enemy removal were much more abrupt than the imposed smooth land use intensity gradient. This may be due to the presence of multiple trophic levels aboveground and belowground. Our model suggests that destabilization and extinction are more likely to occur in agroecosystems than in natural communities, but the shape of the relationship is nonlinear under the influence of multiple trophic interactions.

Microbiology of the phyllosphere: a playground for testing ecological concepts.

Many concepts and theories in ecology are highly debated, because it is often difficult to design decisive tests with sufficient replicates. Examples include biodiversity theories, succession concepts, invasion theories, coexistence theories, and concepts of life history strategies. Microbiological tests of ecological concepts are rapidly accumulating, but have yet to tap into their full potential to complement traditional macroecological theories. Taking the example of microbial communities on leaf surfaces (i.e. the phyllosphere), we show that most explorations of ecological concepts in this field of microbiology focus on autecology and population ecology, while community ecology remains understudied. Notable exceptions are first tests of the island biogeography theory and of biodiversity theories. Here, the phyllosphere provides the unique opportunity to set up replicated experiments, potentially moving fields such as biogeography, macroecology, and landscape ecology beyond theoretical and observational evidence. Future approaches should take advantage of the great range of spatial scales offered by the leaf surface by iteratively linking laboratory experiments with spatial simulation models.

Reduction of rare soil microbes modifies plant-herbivore interactions.

Rare species are assumed to have little impact on community interactions and ecosystem processes. However, very few studies have actually attempted to quantify the role of rare species in ecosystems. Here we compare effects of soil community assemblages on plant-herbivore interactions and show that reduction of rare soil microbes increases both plant biomass and plant nutritional quality. Two crop plant species growing in soil where rare microbes were reduced, had tissues of higher nutritional quality, which theoretically makes them more susceptible to pest organisms such as shoot-feeding aphids and root-feeding nematodes. Reduction of rare microbes increased aphid body size in the absence of nematodes; nematodes always reduced aphid body size independent of the soil microbial community. This study is the first to show that rare soil microbes are not redundant but may play a role in crop protection by enhancing aboveground and belowground plant defence. It remains to be tested whether these are direct effects of rare soil microbes on plants and herbivores, or indirect effects via shifts in the microbial soil community assemblages.