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Long-term impacts of livestock herbivory on herbaceous and woody vegetation in semiarid savannas
(2000)
Simulated long-term vegetation response to alternative stocking strategies in savanna rangelands
(2000)
Ecological buffering mechanisms in savannas : a unifying theory of long-term tree-grass coexistence
(2000)
Environmental heterogeneity is a major determinant of plant population dynamics. In semi-arid Kalahari savannas, heterogeneity is created by savanna structure, i.e. by the spatial arrangement and temporal dynamics of woody plant and open grassland microsites. We formulate a conceptual model describing the effects of savanna dynamics on the population dynamics of the animal-dispersed shrub Grewia flava. From empirical results we derive model rules describing effects of savanna structure on several processes in Grewia's life cycle. By formulating the model, we summarise existing information on Grewia demography and identify gaps in this knowledge. Despite a number of such gaps, the model can be used to make certain quantitative predictions. As an example, we apply the model to investigate the role of seed dispersal in Grewia encroachment on rangelands. Model results show that cattle promote encroachment by depositing substantial numbers of seeds in open areas, where Grewia is otherwise dispersal-limited. Finally, we draw some general conclusions about Grewia's life history and population dynamics. Under natural conditions, concentrated seed deposition under woody plants appears to be a key process causing the observed association between Grewia and other woody plants. Furthermore, low rates of recruitment and high adult survival result in slow-motion dynamics of Grewia populations. As a consequence, Grewia populations interact with savanna dynamics on long temporal and short to intermediate spatial scales.
Structural and animal species diversity in arid and semi-arid savannas of the southern Kalahari
(2004)
The small fox tapeworm (Echinococcus multilocularis) shows a heterogeneous spatial distribution in the intermediate host (Microtus arvalis). To identify the ecological processes responsible for this heterogeneity, we developed a spatially explicit simulation model. The model combines individual-based (foxes, Vulpes vulpes) and grid- based (voles) techniques to simulate the infections in both intermediate and definite host. If host populations are homogeneously mixed, the model reproduces field data for parasite prevalence only for a limited number of parameter combinations. As ecological parameters inevitably vary to a certain degree, we discarded the homogeneous mixing model as insufficient to gain insight into the ecology of the fox tapeworm cycle. We analysed five different model scenarios, each focussing on an ecological process that might be responsible for the heterogeneous spatial distribution of E multilocularis in the intermediate host. Field studies revealed that the prevalence ratio between intermediate and definite host remains stable over a wide range of ecological conditions. Thus, by varying the parameters in simulation experiments, we used the robustness of the agreement between field data and model output as quality criterion for the five scenarios. Only one of the five scenarios was found to reproduce the prevalence ratio over a sufficient range of parameter combinations. In the accentuated scenario most tapeworm eggs die due to bad environmental conditions before they cause infections in the intermediate host. This scenario is supported by the known sensitivity of tapeworm eggs to high temperatures and dry conditions. The identified process is likely to lead to a heterogeneous availability of infective eggs and thus to a clumped distribution of infected intermediate hosts. In conclusion, areas with humid conditions and low temperatures must be pointed out as high risk areas for human exposure to E. multilocularis eggs as well. (C) 2004 on behalf of Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved
There is concern about the lack of recruitment of Acacia trees in the Negev desert of Israel. We have developed three models to estimate the frequency of recruitment necessary for long-term population survival (i.e. positive average population growth for 1,000 years and <10% probability of extinction). Two models assume purely episodic recruitment based on the general notion that recruitment in and environments is highly episodic. They differ in that the deterministic model investigates average dynamics while the stochastic model does not. Studies indicating that recruitment episodes in and environments have been overemphasized motivated the development of the third model. This semi-stochastic model simulates a mixture of continuous and episodic recruitment. Model analysis was done analytically for the deterministic model and via running model simulations for the stochastic and semi-stochastic models. The deterministic and stochastic models predict that, on average, 2.2 and 3.7 recruitment events per century, respectively, are necessary to sustain the population. According to the semi-stochastic model, 1.6 large recruitment events per century and an annual probability of 50% that a small recruitment event occurs are needed. A consequence of purely episodic recruitment is that all recruitment episodes produce extremely large numbers of recruits (i.e. at odds with field observations), an evaluation that holds even when considering that rare events must be large. Thus, the semi- stochastic model appears to be the most realistic model. Comparing the prediction of the semi-stochastic model to field observations in the Negev desert shows that the absence of observations of extremely large recruitment events is no reason for concern. However, the almost complete absence of small recruitment events is a serious reason for concern. The lack of recruitment may be due to decreased densities of large mammalian herbivores and might be further exacerbated by possible changes in climate, both in terms of average precipitation and the temporal distribution of rain
In ecology much attention has been paid towards seed dispersal of fleshy-fruited plants, however, knowledge is lacking about the Iona-term demographic consequences of variation in dispersal distance and fruit removal rate, particularly given the natural variability of the environment the organism lives in. In this study we used a spatially explicit, two-level stochastic computer model to simulate population dynamics of a fleshy-fruited shrub living in the sub-canopy of solitary savanna trees. On the landscape level we implemented three realistic scenarios of savanna landscape dynamics for a period of 500 years with equal inter-annual mean of environmental variables. The first scenario is representative of a relatively constant environment with normal variability in precipitation, constant tree density and random tree recruitment pattern. The second and third scenarios represent positive auto-correlated, cyclic patterns with alternating phases of tree cover increase and decrease corresponding with favorable and unfavorable rain phases. Our simulation experiments show that when fruit removal rate is extremely low, population persistence is enhanced under relatively constant rain conditions, while alternating rain phases of the cyclic scenarios lead to a significant population decrease. This result confirms previous findings that periodically fluctuating environments may increase local extinction risk. However, when dispersal distance is a limiting factor (whilst removal rate was sufficiently high), tree clumps typically forming in wet phases of both cyclic scenarios compensated for the negative effect of low dispersal distances, while the constant scenario with random tree pattern and larger inter-tree distances resulted in a significant population decline. (C) 2003 Elsevier B.V. All rights reserved
Shrub encroachment, i.e. the increase in woody plant cover, is a major concern for livestock farming in southern Kalahari savannas. We developed a grid-based computer model simulating the population dynamics of Grewia flava, a common, fleshy-fruited encroaching shrub. In the absence of large herbivores, seeds of Grewia are largely deposited in the sub-canopy of Acacia erioloba. Cattle negate this dispersal limitation by browsing on the foliage of Grewia and dispersing seeds into the grassland matrix. In this study we first show that model predictions of Grewia cover dynamics are realistic by comparing model output with shrub cover estimates obtained from a time series of aerial photographs. Subsequently, we apply a realistic range of intensity of cattle-induced seed dispersal combined with potential precipitation and fire scenarios. Based on the simulation results we suggest that cattle may facilitate shrub encroachment of Grewia. The results show that the severity of shrub encroachment is governed by the intensity of seed dispersal. For a high seed dispersal intensity without fire (equivalent to a high stocking rate) the model predicts 56% shrub cover and 85% cell cover after 100 yr. With fire both recruitment and shrub cover are reduced, which may, under moderate intensities, prevent shrub encroachment. Climate change scenarios with two-fold higher frequencies of drought and wet years intensified shrub encroachment rates, although long-term mean of precipitation remained constant. As a management recommendation we suggest that shrub encroachment on rangelands may be counteracted by frequent fires and controlling cattle movements to areas with a high proportion of fruiting Grewia shrubs
In a selected literature survey we reviewed studies on the habitat heterogeneity-animal species diversity relationship and evaluated whether there are uncertainties and biases in its empirical support. We reviewed 85 publications for the period 1960-2003. We screened each publication for terms that were used to define habitat heterogeneity, the animal species group and ecosystem studied, the definition of the structural variable, the measurement of vegetation structure and the temporal and spatial scale of the study. The majority of studies found a positive correlation between habitat heterogeneity/diversity and animal species diversity. However, empirical support for this relationship is drastically biased towards studies of vertebrates and habitats under anthropogenic influence. In this paper we show that ecological effects of habitat heterogeneity may vary considerably between species groups depending on whether structural attributes are perceived as heterogeneity or fragmentation. Possible effects may also vary relative to the structural variable measured. Based upon this, we introduce a classification framework that may be used for across-studies comparisons. Moreover, the effect of habitat heterogeneity for one species group may differ in relation to the spatial scale. In several studies, however, different species groups are closely linked to 'keystone structures' that determine animal species diversity by their presence. Detecting crucial keystone structures of the vegetation has profound implications for nature conservation and biodiversity management.
The predicted climate change causes deep concerns on the effects of increasing temperatures and changing precipitation patterns on species viability and, in turn, on biodiversity. Models of Population Viability Analysis (PVA) provide a powerful tool to assess the risk of species extinction. However, most PVA models do not take into account the potential effects of behavioural adaptations. Organisms might adapt to new environmental situations and thereby mitigate negative effects of climate change. To demonstrate such mitigation effects, we use an existing PVA model describing a population of the tawny eagle (Aquila rapax) in the southern Kalahari. This model does not include behavioural adaptations. We develop a new model by assuming that the birds enlarge their average territory size to compensate for lower amounts of precipitation. Here, we found the predicted increase in risk of extinction due to climate change to be much lower than in the original model. However, this "buffering" of climate change by behavioural adaptation is not very effective in coping with increasing interannual variances. We refer to further examples of ecological "buffering mechanisms" from the literature and argue that possible buffering mechanisms should be given due consideration when the effects of climate change on biodiversity are to be predicted. (c) 2004 Elsevier B.V. All rights reserved
Agent-based complex systems are dynamic networks of many interacting agents; examples include ecosystems, financial markets, and cities. The search for general principles underlying the internal organization of such systems often uses bottom-up simulation models such as cellular automata and agent-based models. No general framework for designing, testing, and analyzing bottom-up models has yet been established, but recent advances in ecological modeling have come together in a general strategy we call pattern-oriented modeling. This strategy provides a unifying framework for decoding the internal organization of agent-based complex systems and may lead toward unifying algorithmic theories of the relation between adaptive behavior and system complexity
Question: Is there a relationship between size and death in the Iona-lived, deep-rooted tree, Acacia erioloba, in a semi-arid savanna? What is the size-class distribution of A. erioloba mortality? Does the mortality distribution differ from total tree size distribution? Does A. erioloba mortality distribution match the mortality distributions recorded thus far in other environments? Location: Dronfield Ranch, near Kimberley, Kalahari, South Africa. Methods: A combination of aerial photographs and a satellite image covering 61 year was used to provide long-term spatial data on mortality. We used aerial photographs of the study area from 1940, 1964, 1984, 1993 and a satellite image from 2001 to follow three plots covering 510 ha. We were able to identify and individually follow ca. 3000 individual trees from 1940 till 2001. Results: The total number of trees increased over time. No relationship between total number of trees and mean tree size was detected. There were no trends over time in total number of deaths per plot or in size distributions of dead trees. Kolmogorov-Smirnov tests showed no differences in size class distributions for living trees through time. The size distribution of dead trees was significantly different from the size distribution of all trees present on the plots. Overall, the number of dead trees was low in small size classes, reached a peak value when canopy area was 20 - 30 m(2), and declined in lamer size-classes. Mortality as a ratio of dead vs. total trees peaked at intermediate canopy sizes too. Conclusion: A. erioloba mortality was size-dependent, peaking at intermediate sizes. The mortality distribution differs from all other tree mortality distributions recorded thus far. We suggest that a possible mechanism for this unusual mortality distribution is intraspecific competition for water in this semi-arid environment.
In semiarid savannas of Southern Africa current land use practices and climate change may lead to substantial changes of vegetation structure in the near future, however uncertainty remains about the potential consequences and the magnitude of change. In this paper we study the impact of climate change, cattle grazing, and wood cutting on shrub cover dynamics in savannas of the southern Kalahari. We use an established savanna ecosystem model to simulate landscape dynamics in terms of rainfall, fire and distribution of the dominant tree Acacia erioloba. We then incorporate these data into a spatial population model of the common, fleshy-fruited shrub Grewia flava and investigate shrub cover dynamics for a period of 100 years. Depending on the intensity of commercial wood cutting practices tree removal of A. erioloba led to a strong decline of the G. flava population, as shrub recruitment is concentrated in tree sub-canopies due to bird-mediated seed dispersal. Under climate change shrub cover slightly decreased with decreasing precipitation and was unchanged with increase in precipitation variability. Contrarily, grazing by cattle strongly increased shrub cover and facilitated shrub encroachment because of cattle-induced distribution of G. flava seeds into the matrix vegetation. Knowledge of the latter process is particularly important because shrub invasion is a major concern for conservation and savanna rangeland management as a result of its adverse effects on livestock carrying capacity and biodiversity
Predicting the breeding success of large raptors in arid southern Africa : a first assessment
(2006)
Raptors are often priorities for conservation efforts and breeding success is a target measure for assessing their conservation status. The breeding success of large raptors in and southern Africa is thought to be higher in years of high rainfall. While this correlation has been found in several studies, it has not yet been shown for data from a wider geographical area. In conservation research, it is important to explore the differences between spatially- separated populations to estimate and to compare their conservation status, and to deduce specific management strategies. Using a theoretical approach, we develop a simplistic model to explain the breeding success-rainfall relationship in large African raptors at larger spatial scales. Secondly, we validate this model and we show that the inclusion of field data leads to consistent predictions. In particular, we recommend that the average size of the 'effective territory' should be included in the relationship between annual rainfall and breeding success of raptors in and southern Africa. Accordingly, we suggest that breeding success is a function of precipitation and inter- nest distance. We present a new measure of territory quality depending on rainfall and territory size. We suggest that our model provides a useful first approach to assess breeding success in large raptors of and southern Africa. However, we strongly emphasise the need to gather more data to further verify our model. A general problem in conservation research is to compare the status of populations assessed in different study areas under changing environmental conditions. Our simplistic approach indicates that this problem can be overcome by using a weighted evaluation of a target measure (i.e. breeding success), taking regional differences into account
Environmental gradients represent an ideal framework for studying adaptive variation in the life history of plant species. However, on very steep gradients, largely contrasting conditions at the two gradient ends often limit the distribution of the same species across the whole range of environmental conditions. Here, we study phenotypic variation in a winter annual crucifer Biscutella didyma persisting along a steep gradient of increasing rainfall in Israel. In particular, we explored whether the life history at the arid end of the gradient indicates adaptations to drought and unpredictable conditions, while adaptations to the highly competitive environment prevail at the mesic Mediterranean end. We examined several morphological and reproductive traits in four natural populations and in populations cultivated in standard common environment. Plants from arid environments were faster in phenological development, more branched in architecture and tended to maximize reproduction, while the Mediterranean plants invested mainly in vertical vegetative growth. Differences between cultivation and field in diaspore production were very large for arid populations as opposed to Mediterranean ones, indicating a larger potential to increase reproduction under favorable conditions. Our overall findings indicate two strongly opposing selective forces at the two extremes of the aridity gradient, which result in contrasting strategies within the studied annual plant species
In most stochastic models addressing the persistence of small populations, environmental noise is included by imposing a synchronized effect of the environment on all individuals. However, buffer mechanisms are likely to exist that may counteract this synchronization to some degree. We have studied whether the flexibility in the mating system, which has been observed in some bird species, is a potential mechanism counteracting the synchronization of environmental fluctuations. Our study organism is the lesser spotted woodpecker Picoides minor (Linnaeus), a generally monogamous species. However, facultative polyandry, where one female mates with two males with separate nests, was observed in years with male-biased sex ratio. We constructed an individual-based model from data and observations of a population in Taunus, Germany. We tested the impact of three behavioural scenarios on population persistence: (1) strict monogamy; (2) polyandry without costs; and (3) polyandry assuming costs in terms of lower survival and reproductive success for secondary males. We assumed that polyandry occurs only in years with male-biased sex ratio and only for females with favourable breeding conditions. Even low rates of polyandry had a strong positive effect on population persistence. The increase of persistence with carrying capacity was slower in the monogamous scenario, indicating strong environmental noise. In the polyandrous scenarios, the increase of persistence was stronger, indicating a buffer mechanism. In the polyandrous scenarios, populations had a higher mean population size, a lower variation in number of individuals, and recovered faster after a population breakdown. Presuming a realistic polyandry rate and costs for polyandry, there was still a strong effect of polyandry on persistence. The results show that polyandry and in general flexibility in mating systems is a buffer mechanism that can significantly reduce the impact of environmental and demographic noise in small populations. Consequently, we suggest that even behaviour that seems to be exceptional should be considered explicitly when predicting the persistence of populations
Use of large Acacia trees by the cavity dwelling Black-tailed Tree Rat in the southern Kalahari
(2006)
Recent extensive harvesting of large, often dead Acacia trees in and savanna of southern Africa is cause for concern about the conservation status of the arid savanna and its animal community. We mapped vegetation and nests of the Black-tailed Tree Rat Thallomy's nigricauda to assess the extent to which the rats depend on particular tree species and on the existence of dead, standing trees. The study was conducted in continuous Acacia woodland on the southern and eastern edge of the Kalahari, South Africa. Trees in which there were tree rat nests were compared with trees of similar size and vigour to identify the characteristics of nest sites. Spatial analysis of tree rat distribution was conducted using Ripley's-L function. We found that T nigricauda was able to utilize all available tree species, as long as trees were large and old enough so that cavities were existing inside the stem. The spatial distribution of nest trees did not show clumping at the investigated scale, and we therefore reject the notion of the rats forming colonies when inhabiting continuous woodlands. The selection of a particular tree as a nest site was furthermore depending on the close proximity of the major food plant, Acacia mellifera. This may limit the choice of suitable nest sites. since A. mellifera was less likely to grow within a vegetation patch containing a large trees than in patches without large trees.
Small livestock is an important resource for rural human populations in dry climates. How strongly will climate change affect the capacity of the rangeland? We used hierarchical modelling to scale quantitatively the growth of shrubs and annual plants, the main food of sheep and goats, to the landscape extent in the eastern Mediterranean region. Without grazing, productivity increased in a sigmoid way with mean annual precipitation. Grazing reduced productivity more strongly the drier the landscape. At a point just under the stocking capacity of the vegetation, productivity declined precipitously with more intense grazing due to a lack of seed production of annuals. We repeated simulations with precipitation patterns projected by two contrasting IPCC scenarios. Compared to results based on historic patterns, productivity and stocking capacity did not differ in most cases. Thus, grazing intensity remains the stronger impact on landscape productivity in this dry region even in the future.
Modelling and empirical studies have shown that input from the regional seed pool is essential to maintain local species diversity. However, most of these studies have concentrated on simplified, if not neutral, model systems, and focus on a limited subset of species or on aggregated measures of diversity only (e.g., species richness or Shannon diversity). Thus they ignore more complex species interactions and important differences between species. To gain a better understanding of how seed immigration affects community structure at the local scale in real communities we conducted computer simulation experiments based on plant functional types (PFTs) for a species-rich, fire-prone Mediterranean-type shrubland in Western Australia. We developed a spatially explicit simulation model to explore the community dynamics of 38 PFTs, defined by seven traits - regeneration mode, seed production, seed size, maximum crown diameter, drought tolerance, dispersal mode and seed bank type - representing 78 woody species. Model parameterisation is based on published and unpublished data on the population dynamics of shrub species collected over 18 years. Simulation experiments are based on two contrasting seed immigration scenarios: (1) the 'equal seed input number' scenario, where the number of immigrant seeds is the same for all PFTs, and (2) the 'equal seed input mass' scenario, where the cumulative mass of migrating seeds is the same for all PFTs. Both scenarios were systematically tested and compared for different overall seed input values. Without immigration the local community drifts towards a state with only 13 coexisting PFTs. With increasing immigration rates in terms of overall mass of seeds the simulated number of coexisting PFTs and Shannon diversity quickly approaches values observed in the field. The equal seed mass scenario resulted in a more diverse community than did the seed number scenario. The model successfully approximates the frequency distributions (relative densities) of all individual plant traits except seed size for scenarios associated with equal seed input mass and high immigration rate. However, no scenario satisfactorily approximated the frequency distribution for all traits in combination. Our results show that regional seed input can explain the more aggregated measures of local community structure, and some, but not all, aspects of community composition. This points to the possible importance of other (untested) processes and traits (e.g., dispersal vectors) operating at the local scale. Our modelling framework can readily allow new factors to be systematically investigated, which is a major advantage compared to previous simulation studies, as it allows us to find structurally realistic models, which can address questions pertinent to ecological theory and to conservation management.
Plant population modelling has been around since the 1970s, providing a valuable approach to understanding plant ecology from a mechanistic standpoint. It is surprising then that this area of research has not grown in prominence with respect to other approaches employed in modelling plant systems. In this review, we provide an analysis of the development and role of modelling in the field of plant population biology through an exploration of where it has been, where it is now and, in our opinion, where it should be headed. We focus, in particular, on the role plant population modelling could play in ecological forecasting, an urgent need given current rates of regional and global environmental change. We suggest that a critical element limiting the current application of plant population modelling in environmental research is the trade-off between the necessary resolution and detail required to accurately characterize ecological dynamics pitted against the goal of generality, particularly at broad spatial scales. In addition to suggestions how to overcome the current shortcoming of data on the process-level we discuss two emerging strategies that may offer a way to overcome the described limitation: (1) application of a modern approach to spatial scaling from local processes to broader levels of interaction and (2) plant functional-type modelling. Finally we outline what we believe to be needed in developing these approaches towards a 'science of forecasting'.
Dry lands are exposed to a highly variable environment and face a high risk of degradation. The effects of climate change are likely to increase this risk; thus a profound knowledge of the system dynamics is crucial for evaluating management options. This applies particularly for the interactions between water and vegetation, which exhibit strong feedbacks. To evaluate these feedbacks and the effects of climate change on soil moisture dynamics, we developed a generic, process-based, spatially explicit soil moisture model of two soil layers, which can be coupled with vegetation models. A time scale relevant for ecological processes can be simulated without difficulty, and the model avoids complex parameterization with data that are unavailable for most regions of the world. We applied the model to four sites in Israel along a precipitation and soil type gradient and assessed the effects of climate change by comparing possible climatic changes with present climate conditions. The results show that in addition to temperature, the total amount of precipitation and its intra-annual variability are an important driver of soil moisture patterns. This indicates that particularly with regard to climate change, the approach of many ecological models that simulate water dynamics on an annual base is far too simple to make reliable predictions. Thus, the introduced model can serve as a valuable tool to improve present ecological models of dry lands because of its focus on the applicability and transferability.
In common garden experiments, a number of genotypes are raised in a common environment in order to quantify the genetic component of phenotypic variation. Common gardens are thus ideally suited for disentangling how genetic and environmental factors contribute to the success of invasive species in their new non-native range. Although common garden experiments are increasingly employed in the study of invasive species, there has been little discussion about how these experiments should be designed for greatest utility. We argue that this has delayed progress in developing a general theory of invasion biology. We suggest a minimum optimal design (MOD) for common garden studies that target the ecological and evolutionary processes leading to phenotypic differentiation between native and invasive ranges. This involves four elements: (A) multiple, strategically sited garden locations, involving at the very least four gardens (2 in the native range and 2 in the invaded range); (B) careful consideration of the genetic design of the experiment; (C) standardization of experimental protocols across all gardens; and (D) care to ensure the biosafety of the experiment. Our understanding of the evolutionary ecology of biological invasions will be greatly enhanced by common garden studies, if and only if they are designed in a more systematic fashion, incorporating at the very least the MOD suggested here.
We investigated sex-specific parental care behaviour of lesser spotted woodpeckers Picoides minor in the low mountain range Taunus, Germany. Observed parental care included incubation, nest sanitation as well as brooding and feeding of nestlings. Contributions of the two sexes to parental care changed in progress of the breeding period. During incubation and the first half of the nestling period, parental care was divided equally between partners. However, in the late nestling stage, we found males to feed their nestlings irrespective of brood size while females considerably decreased feeding rate with the number of nestlings. This behaviour culminated in desertion of small broods by females shortly before fledging. The fact that even deserted nests were successful indicates that males were able to compensate for the females' absence. Interestingly, the mating of one female with two males with separate nests could be found in the population, which confirms earlier findings of polyandry in the lesser spotted woodpecker. We conclude that biparental care is not essential in the later stage and one partner can reduce effort and thus costs of parental care, at least in small broods where the mate is able to compensate for that behaviour. Reduced care and desertion appears only in females, which might be caused by a combination of two traits: First, females might suffer higher costs of investment in terms of mortality and secondly, male-biased sex ratio in the population generally leads to higher mating probabilities for females in the following breeding season. The occurrence of polyandry seems to be a result of these conditions.
We studied the effects of overgrazing on the foraging behaviour of the lizard Pedioplanis l. lineoocellata (Spotted Sand Lizard), a sit-and-wait forager, in habitats of differing vegetation states to determine the effects of habitat degradation on this species. At high grazing intensity where vegetation cover and diversity is low, the lizard P. lineoocellata moves more frequently, spends more time moving and covers larger distances than in habitats where vegetation cover and diversity is high. These behavioural changes in movement patterns can be explained by less abundant prey in habitats with low vegetation cover and diversity. Although morphology, phylogeny and physiology of P. lineoocellata should constrain the change in foraging behaviour, the species has modified its foraging strategy from sit- and-wait to actively foraging. We assume that this behavioural flexibility of P. lineoocellata is a buffer mechanism enabling the species to use and survive in degraded (unfavourable) habitats.
Tree size and herbivory determine below-canopy grass quality and species composition in savannahs
(2009)
Large single-standing trees are rapidly declining in savannahs, ecosystems supporting a high diversity of large herbivorous mammals. Savannah trees are important as they support both a unique flora and fauna. The herbaceous layer in particular responds to the structural and functional properties of a tree. As shrubland expands stem thickening occurs and large trees are replaced by smaller trees. Here we examine whether small trees are as effective in providing advantages for grasses growing beneath their crowns as large trees are. The role of herbivory in this positive tree- grass interaction is also investigated. We assessed soil and grass nutrient content, structural properties, and herbaceous species composition beneath trees of three size classes and under two grazing regimes in a South African savannah. We found that grass leaf content (N and P) beneath the crowns of particularly large (ca. 3.5 m) and very large trees (ca. 9 m) was as much as 40% greater than the same grass species not growing under a tree canopy, whereas nutrient contents of grasses did not differ beneath small trees (< 2.3 m). Moderate herbivory enhanced these effects slightly. Grass species composition differed beneath and beyond the tree canopy but not between tree size classes. As large trees significantly improve the grass nutrient quality for grazers in contrast to smaller trees, the decline of the former should be halted. The presence of trees further increases grass species diversity and patchiness by favouring shade- tolerant species. Both grazing wildlife and livestock will benefit from the presence of large trees because of their structural and functional importance for savannahs.
Shrub encroachment linked to heavy grazing has dramatically changed savanna landscapes, and is a major form of rangeland degradation. Our understanding of how shrub encroachment affects arthropod communities is poor, however. Here, we investigate the effects of shrub encroachment on abundance and diversity of ground-dwelling (wingless) arthropods at varying levels of shrub cover in the southern Kalahari. We also ascertain if invertebrate assemblage composition changes with habitat structure and identify which aspects of habitat structure (e.g., grass cover, herbaceous plant cover, shrub density) correlate most strongly with these changes. Ant, scorpion and dung beetle abundance increased with shrub cover, whereas grasshoppers and solifuges declined. Spider and beetle abundance exhibited hump-shaped relationships with shrub cover. RTU richness within orders either mirrored abundances, or exhibited no trend. Shrub density was the habitat component most correlated with similarities between invertebrate assemblages. Ground-dwelling arthropods showed clear shifts in species assemblage composition at a similarity level of 65% according to shrub density. Changes in indicator species showed that within the Tenebrionidae (darkling beetles), certain species respond positively to shrub thickening, replacing other species within the Family. Small-bodied, wingless Scarabaeidae (dung beetles) tended to increase with increased shrub density and three species emerged as significant indicators of more thickened habitats, although this might be a response to greater dung availability, rather than habitat structure itself. We conclude that because ground- dwelling invertebrates showed such clear responses in species assemblage composition, they present excellent candidates for use as indicator species in further studies into bush encroachment.
SDM performance varied for different range dynamics. Prediction accuracies decreased when abrupt range shifts occurred as species were outpaced by the rate of climate change, and increased again when a new equilibrium situation was realised. When ranges contracted, prediction accuracies increased as the absences were predicted well. Far- dispersing species were faster in tracking climate change, and were predicted more accurately by SDMs than short- dispersing species. BRTs mostly outperformed GLMs. The presence of a predator, and the inclusion of its incidence as an environmental predictor, made BRTs and GLMs perform similarly. Results are discussed in light of other studies dealing with effects of ecological traits and processes on SDM performance. Perspectives are given on further advancements of SDMs and for possible interfaces with more mechanistic approaches in order to improve predictions under environmental change.
Aim This study aims to link demographic traits and post-glacial recolonization processes with genetic traits in Himantoglossum hircinum (L.) Spreng (Orchidaceae), and to test the implications of the central-marginal concept (CMC) in Europe. Location Twenty sites covering the entire European distribution range of this species. Methods We employed amplified fragment length polymorphism (AFLP) markers and performed a plastid microsatellite survey to assess genetic variation in 20 populations of H. hircinum located along central-marginal gradients. We measured demographic traits to assess population fitness along geographical gradients and to test for correlations between demographic traits and genetic diversity. We used genetic diversity indices and analyses of molecular variance (AMOVA) to test hypotheses of reduced genetic diversity and increased genetic differentiation and isolation from central to peripheral sites. We used Bayesian simulations to analyse genetic relationships among populations. Results Genetic diversity decreased significantly with increasing latitudinal and longitudinal distance from the distribution centre when excluding outlying populations. The AMOVA revealed significant genetic differentiation among populations (F-ST = 0.146) and an increase in genetic differentiation from the centre of the geographical range to the margins (except for the Atlantic group). Population fitness, expressed as the ratio N-R/N, decreased significantly with increasing latitudinal distance from the distribution centre. Flower production was lower in most eastern peripheral sites. The geographical distribution of microsatellite haplotypes suggests post-glacial range expansion along three major migratory pathways, as also supported by individual membership fractions in six ancestral genetic clusters (C1-C6). No correlations between genetic diversity (e.g. diversity indices, haplotype frequency) and population demographic traits were detected. Main conclusions Reduced genetic diversity and haplotype frequency in H. hircinum at marginal sites reflect historical range expansions. Spatial variation in demographic traits could not explain genetic diversity patterns. For those sites that did not fit into the CMC, the genetic pattern is probably masked by other factors directly affecting either demography or population genetic structure. These include post-glacial recolonization patterns and changes in habitat suitability due to climate change at the northern periphery. Our findings emphasize the importance of distinguishing historical effects from those caused by geographical variation in population demography of species when studying evolutionary and ecological processes at the range margins under global change.
Grazing is known as one of the key factors for diversity and community composition in grassland ecosystems, but the response of plant communities towards grazing varies remarkably between sites with different environmental conditions. It is generally accepted that grazing increases plant diversity in productive environments, while it tends to reduce diversity in unproductive habitats (grazing reversal hypothesis). Despite empirical evidence for this pattern the mechanistic link between modes of plant-plant competition and grazing response at the community level still remains poorly understood. Root-competition in particular has rarely been included in theoretical studies, although it has been hypothesized that variations in productivity and grazing regime can alter the relative importance of shoot- and root-competition. We therefore developed an individual-based model based on plant functional traits to investigate the response of a grassland community towards grazing. Models of different complexity, either incorporating only shoot competition or with distinct shoot- and root-competition, were used to study the interactive effects of grazing, resource availability, and the mode of competition (size-symmetric or asymmetric). The pattern predicted by the grazing reversal hypothesis (GRH) can only be explained by our model if shoot- and root-competition are explicitly considered and if size asymmetry of above- and symmetry of below-ground competition is assumed. For this scenario, the model additionally reproduced empirically observed plant trait responses: erect and large plant functional types (PFTs) dominated without grazing, while frequent grazing favoured small PFTs with a rosette growth form. We conclude that interactions between shoot- and root-competition and size symmetry/asymmetry of plant-plant interactions are crucial in order to understand grazing response under different habitat productivities. Our results suggest that future empirical trait surveys in grassland communities should include root traits, which have been largely ignored in previous studies, in order to improve predictions of plants" responses to grazing.
The interaction between ecological and hydrological processes is particularly important in arid and semi-arid regions. Often the interaction between these processes is not completely understood and they are studied separately. We developed a grid-based computer model simulating the dynamics of the four most common vegetation types (perennial grass, annuals, dwarf shrubs and shrubs) and related hydrological processes in the region studied. Eco-hydrological interactions gain importance in rangelands with increasing slope, where vegetation cover obstructs run-off and decreases evaporation from the soil. Overgrazing can influence these positive feedback mechanisms. In this study, we first show that model predictions of cover and productivity of the vegetation types are realistic by comparing them with estimates obtained from field surveys. Then, we apply a realistic range in slope angle combined with two land use regimes (light versus heavy grazing intensity). Our simulation results reveal that hydrological processes and associated productivity are strongly affected by slope, whereas the magnitude of this impact depends on overgrazing. Under low stocking rates, undisturbed vegetation is maintained and run-off and evaporation remain low on flat plains and gentle slope. On steep slopes, run-off and evaporation become larger, while water retention potential decreases, which leads to reduced productivity. Overgrazing, however, reduces vegetation cover and biomass production and the landscape"s ability to conserve water decreases even on flat plains and gentle slopes. Generally, the abundance of perennial grasses and shrubs decreases with increasing slope and grazing. Dominance is shifted towards shrubs and annuals. As a management recommendation we suggest that different vegetation growth forms should not only be regarded as forage producers but also as regulators of ecosystem functioning. Particularly on sloping range lands, a high percentage of cover by perennial vegetation insures that water is retained in the system.
Scaling up ecohydrological processes : role of surface water flow in water-limited landscapes
(2009)
In this study, we present a stochastic landscape modeling approach that has the power to transfer and integrate existing information on vegetation dynamics and hydrological processes from the small scale to the landscape scale. To include microscale processes like ecohydrological feedback mechanisms and spatial exchange like surface water flow, we derive transition probabilities from a fine-scale simulation model. We applied two versions of the landscape model, one that includes and one that disregards spatial exchange of water to the situation of a sustainably used research farm and communally used and degraded rangeland in semiarid Namibia. Our simulation experiments show that including spatial exchange of overland flow among vegetation patches into our model is a precondition to reproduce vegetation dynamics, composition, and productivity, as well as hydrological processes at the landscape scale. In the model version that includes spatial exchange of water, biomass production at light grazing intensities increases 2.24-fold compared to the model without overland flow. In contrast, overgrazing destabilizes positive feedbacks through vegetation and hydrology and decreases the number of hydrological sinks in the model with overland flow. The buffer capacity of these hydrological sinks disappears and runoff increases. Here, both models predicted runoff losses from the system and artificial droughts occurring even in years with good precipitation. Overall, our study reveals that a thorough understanding of overland flow is an important precondition for improving the management of semiarid and arid rangelands with distinct topography.
The need to implement sustainable resource management regimes for semi-arid and arid rangelands is acute as non- adapted grazing strategies lead to irreversible environmental problems such as desertification and associated loss of economic support to society. In these sensitive ecosystems, traditional sectoral, disciplinary approaches will not work to attain sustainability: achieving a collective vision of how to attain sustainability requires interactive efforts among disciplines in a more integrated approach. Therefore, we developed an integrated ecological-economic approach that consists of an ecological and an economic module and combines relevant processes on either level. Parameters for both modules are adjusted for an arid dwarf shrub savannah in southern Namibia. The economic module is used to analyse decisions of different virtual farmer types on annual stocking rates depending on their knowledge how the ecosystem works and climatic conditions. We used a dynamic linear optimisation model to simulate farm economics and livestock dynamics. The ecological module is used to simulate the impact of the farmers' land-use decision, derived by the economic module, on ecosystem dynamics and resulting carrying capacity of the system for livestock. Vegetation dynamics, based on the concept of State-and-transition models, and forage productivity for both modules is derived by a small- scale and spatially explicit vegetation model. This mechanistic approach guarantees that data collected and processes estimated at smaller scales are included in our application. Simulation results of the ecological module were successfully compared to simulation results of the optimisation model for a time series of 30 years. We revealed that sustainable management of semi-arid and arid rangelands relies strongly on rangeland managers' understanding of ecological processes. Furthermore, our simulation results demonstrate that the projected lower annual rainfall due to climate change adds an additional layer of risk to these ecosystems that are already prone to land degradation.
The long-term persistence of populations and species depends on the successful recruitment of individuals. The generative recruitment of plants may be limited by a lack of suitable germination and establishment conditions. Establishment limitation may especially be caused by the competitive effect of surrounding dense vegetation, which is believed to restrict the recruitment success of many plant species to small open patches ('safe sites'). We conducted experiments to clarify the roles of germination and seedling establishment as limiting processes in the recruitment of Juncus atratus Krock., a rare and threatened herbaceous perennial river corridor plant in Central Europe. Light intensity had a positive effect on germination. However, some seedlings emerged even in total darkness and the germination rate at 1% light intensity was more than half of that at 60% light intensity. Seedling establishment in the field after 10 weeks was 30% on bare ground, but it was close to zero in grassland. Establishment in the growth chamber after 8 weeks was close to 75% for seedlings that germinated underwater, but only about 35% for seedlings that germinated afloat. Furthermore, establishment decreased with flooding duration on bare ground, but increased with flooding duration in grassland. These data indicate that establishment, rather than germination, is a critical life stage in Central European populations off. atratus. They furthermore indicate that the competition of surrounding vegetation for water limits seedling establishment under field conditions without flooding, largely restricting establishment success to bare ground habitats. In contrast, grassland is more suitable for the recruitment off. atratus than bare ground under prolonged flooding. Grassland may facilitate the establishment off. atratus seedlings during long- lasting floods by supplying oxygen to the soil through aerenchyma. The shift from competition to facilitation in grassland occurred after 30 days of flooding, i.e. within the ontogeny of individual plants. The specific recruitment requirements off. arrows may be a main cause of its rarity in modern Central Europe. In order to prevent regional extinction off. atratus, we suggest maintaining or re-establishing natural hydrodynamics in the species' habitats.
Question: The majority of studies investigating the impact of climate change on local plant communities ignores changes in regional processes, such as immigration from the regional seed pool. Here we explore: (i) the potential impact of climate change on composition of the regional seed pool, (ii) the influence of changes in climate and in the regional seed pool on local community structure, and (iii) the combinations of life history traits, i.e. plant functional types (PFTs), that are most affected by environmental changes. Location: Fire-prone, Mediterranean-type shrublands in southwestern Australia. Methods: Spatially explicit simulation experiments were conducted at the population level under different rainfall and fire regime scenarios to determine the effect of environmental change on the regional seed pool for 38 PFTs. The effects of environmental and seed immigration changes on local community dynamics were then derived from community-level experiments. Classification tree analyses were used to investigate PFT- specific vulnerabilities to climate change. Results: The classification tree analyses revealed that responses of PFTs to climate change are determined by specific trait characteristics. PFT-specific seed production and community patterns responded in a complex manner to climate change. For example, an increase in annual rainfall caused an increase in numbers of dispersed seeds for some PFTs, but decreased PFT diversity in the community. Conversely, a simulated decrease in rainfall reduced the number of dispersed seeds and diversity of PFTs. Conclusions: PFT interactions and regional processes must be considered when assessing how local community structure will be affected by environmental change.
Conservation actions need to account for and be adapted to address changes that will occur under global climate change. The identification of stresses on biological diversity (as defined in the Convention on Biological Diversity) is key in the process of adaptive conservation management. We considered any impact of climate change on biological diversity a stress because such an effect represents a change (negative or positive) in key ecological attributes of an ecosystem or parts of it. We applied a systemic approach and a hierarchical framework in a comprehensive classification of stresses to biological diversity that are caused directly by global climate change. Through analyses of 20 conservation sites in 7 countries and a review of the literature, we identified climate-change-induced stresses. We grouped the identified stresses according to 3 levels of biological diversity: stresses that affect individuals and populations, stresses that affect biological communities, and stresses that affect ecosystem structure and function. For each stress category, we differentiated 3 hierarchical levels of stress: stress class (thematic grouping with the coarsest resolution, 8); general stresses (thematic groups of specific stresses, 21); and specific stresses (most detailed definition of stresses, 90). We also compiled an overview of effects of climate change on ecosystem services using the categories of the Millennium Ecosystem Assessment and 2 additional categories. Our classification may be used to identify key climate-change-related stresses to biological diversity and may assist in the development of appropriate conservation strategies. The classification is in list format, but it accounts for relations among climate-change-induced stresses.
Low-dimensional trade-offs fail to explain richness and structure in species-rich plant communities
(2011)
Mathematical models and ecological theory suggest that low-dimensional life history trade-offs (i.e. negative correlation between two life history traits such as competition vs. colonisation) may potentially explain the maintenance of species diversity and community structure. In the absence of trade-offs, we would expect communities to be dominated by 'super-types' characterised by mainly positive trait expressions. However, it has proven difficult to find strong empirical evidence for such trade-offs in species-rich communities. We developed a spatially explicit, rule-based and individual-based stochastic model to explore the importance of low-dimensional trade-offs. This model simulates the community dynamics of 288 virtual plant functional types (PFTs), each of which is described by seven life history traits. We consider trait combinations that fit into the trade-off concept, as well as super-types with little or no energy constraints or resource limitations, and weak PFTs, which do not exploit resources efficiently. The model is parameterised using data from a fire-prone, species-rich Mediterranean-type shrubland in southwestern Australia. We performed an exclusion experiment, where we sequentially removed the strongest PFT in the simulation and studied the remaining communities. We analysed the impact of traits on performance of PFTs in the exclusion experiment with standard and boosted regression trees. Regression tree analysis of the simulation results showed that the trade-off concept is necessary for PFT viability in the case of weak trait expression combinations such as low seed production or small seeds. However, species richness and diversity can be high despite the presence of super-types. Furthermore, the exclusion of super-types does not necessarily lead to a large increase in PFT richness and diversity. We conclude that low-dimensional trade-offs do not provide explanations for multi-species co-existence contrary to the prediction of many conceptual models.
Scale-dependent determinants of plant species richness in a semi-arid fragmented agro-ecosystem
(2011)
Aims: (1) Understanding how the relationship between species richness and its determinants depends on the interaction between scales at which the response and explanatory variables are measured. (2) Quantifying the relative contributions of local, intermediate and large-scale determinants of species richness in a fragmented agro-ecosystem. (3) Testing the hypothesis that the relative contribution of these determinants varies with the grain size at which species richness is measured.
Location: A fragmented agro-ecosystem in the Southern Judea Lowland, Israel, within a desert-Mediterranean transition zone.
Methods: Plant species richness was estimated using hierarchical nested sampling in 81 plots, positioned in 38 natural vegetation patches within an agricultural matrix (mainly wheat fields) among three land units along a sharp precipitation gradient. Explanatory variables included position along that gradient, patch area, patch isolation, habitat heterogeneity and overall plant density. We used general linear models and hierarchical partitioning of variance to test and quantify the effect of each explanatory variable on species richness at four grain sizes (0.0625, 1, 25 and 225m(2)).
Results: Species richness was mainly affected by position along a precipitation gradient and overall plant density, and to a lesser extent by habitat heterogeneity. It was also significantly affected by patch area and patch isolation, but only for small grain sizes. The contribution of each explanatory variable to explained variance in species richness varied with grain size, i.e. scale-dependent. The influence of geographic position and habitat heterogeneity on species richness increased with grain size, while the influence of plant density decreased with grain size.
Main conclusions: Species richness is determined by the combined effect of several scale-dependent determinants. Ability to detect an effect and effect size of each determinant varies with the scale (grain size) at which it is measured. The combination of a multi-factorial approach and multi-scale sampling reveals that conclusions drawn from studies that ignore these dimensions are restricted and potentially misleading.
Morphological plasticity is a striking characteristic of plants in natural communities. In the context of foraging behavior particularly, root plasticity has been documented for numerous species. Root plasticity is known to mitigate competitive interactions by reducing the overlap of the individuals' rhizospheres. But despite its obvious effect on resource acquisition, plasticity has been generally neglected in previous empirical and theoretical studies estimating interaction intensity among plants. In this study, we developed a semi-mechanistic model that addresses this shortcoming by introducing the idea of compensatory growth into the classical-zone-of influence (ZOI) and field-of-neighborhood (FON) approaches. The model parameters describing the belowground plastic sphere of influence (PSI) were parameterized using data from an accompanying field experiment. Measurements of the uptake of a stable nutrient analogue at distinct distances to the neighboring plants showed that the study species responded plastically to belowground competition by avoiding overlap of individuals' rhizospheres. An unexpected finding was that the sphere of influence of the study species Bromus hordeaceus could be best described by a unimodal function of distance to the plant's center and not with a continuously decreasing function as commonly assumed. We employed the parameterized model to investigate the interplay between plasticity and two other important factors determining the intensity of competitive interactions: overall plant density and the distribution of individuals in space. The simulation results confirm that the reduction of competition intensity due to morphological plasticity strongly depends on the spatial structure of the competitive environment. We advocate the use of semi-mechanistic simulations that explicitly consider morphological plasticity to improve our mechanistic understanding of plant interactions.
Understanding and predicting the composition and spatial structure of communities is a central challenge in ecology. An important structural property of animal communities is the distribution of individual home ranges. Home range formation is controlled by resource heterogeneity, the physiology and behaviour of individual animals, and their intra- and interspecific interactions. However, a quantitative mechanistic understanding of how home range formation influences community composition is still lacking. To explore the link between home range formation and community composition in heterogeneous landscapes we combine allometric relationships for physiological properties with an algorithm that selects optimal home ranges given locomotion costs, resource depletion and competition in a spatially-explicit individual-based modelling framework. From a spatial distribution of resources and an input distribution of animal body mass, our model predicts the size and location of individual home ranges as well as the individual size distribution (ISD) in an animal community. For a broad range of body mass input distributions, including empirical body mass distributions of North American and Australian mammals, our model predictions agree with independent data on the body mass scaling of home range size and individual abundance in terrestrial mammals. Model predictions are also robust against variation in habitat productivity and landscape heterogeneity. The combination of allometric relationships for locomotion costs and resource needs with resource competition in an optimal foraging framework enables us to scale from individual properties to the structure of animal communities in heterogeneous landscapes. The proposed spatially-explicit modelling concept not only allows for detailed investigation of landscape effects on animal communities, but also provides novel insights into the mechanisms by which resource competition in space shapes animal communities.
Habitat loss poses a severe threat to biodiversity. While many studies yield valuable information on how specific species cope with such environmental modification, the mechanistic understanding of how interacting species or whole communities are affected by habitat loss is still poor. Individual movement plays a crucial role for the space use characteristics of species, since it determines how individuals perceive and use their heterogeneous environment. At the community level, it is therefore essential to include individual movement and how it is influenced by resource sharing into the investigation of consequences of habitat loss. To elucidate the effects of foraging movement on communities in face of habitat loss, we here apply a recently published spatially-explicit and individual-based model of home range formation. This approach allows predicting the individual size distribution (ISD) of mammal communities in simulation landscapes that vary in the amount of suitable habitat. We apply three fundamentally different foraging movement approaches (central place forager (CPF), patrolling forager (PF) and body mass dependent nomadic forager (BNF)). Results show that the efficiency of the different foraging strategies depends on body mass, which again affects community structure in face of habitat loss. CPF is only efficient for small animals, and therefore yields steep ISD exponents on which habitat loss has little effect (due to a movement limitation of body mass). PF and particularly BNF are more efficient for larger animals, resulting in less steep ISDs with higher mass maxima, both showing a threshold behaviour with regard to loss of suitable habitat. These findings represent a new way of explaining observed extinction thresholds, and therefore indicate the importance of individual space use characterized by physiology and behaviour, i.e. foraging movement, for communities and their response to habitat loss. Findings also indicate the necessity to incorporate the crucial role of movement into future conservation efforts of terrestrial communities.
Disturbances are characteristic for many ecosystems. However, we still lack generalizations concerning their role in shaping communities, particularly when disturbances co-occur. To study such effects, we used a novel modeling approach that is unrestricted by a priori tradeoffs among specific plant traits, except for those generated by allocation principles. Thus, trait combinations were emergent properties associated with biotic and abiotic constraints. Specifically, we asked which traits dominate under specific disturbance regimes, whether single and combined disturbance regimes promote similar trait tradeoffs and how complex disturbance regimes affect species richness and functional diversity. Overall, disturbances' temporal properties governed the outcome of combined disturbances and were a stronger assortative force than spatial disturbance properties: low temporal predictability decreased seed-dispersability and dormancy, but increased competitive ability and disturbance tolerance. Evidence for tradeoffs between different colonization modes and between dormancy and disturbance tolerance were found, while surprisingly, the widely accepted colonization-competition tradeoff was not generated. Diversity was highest at intermediate disturbance intensity, but decreased monotonically with increasing unpredictability. In accordance with our results, future models should avoid restrictive assumptions about tradeoffs to generate robust and more general predictions about the role of disturbances for community dynamics.
The response of species diversity to dispersal capability is inherently scale-dependent: increasing dispersal capability is expected to increase diversity at the local scale, while decreasing diversity at the metacommunity scale. However, these expectations are based on model formulations that neglect dispersal limitation and species segregation at the local scale. We developed a unifying framework of dispersaldiversity relationships and tested the generality of these expectations. For this purpose we used a spatially-explicit neutral model with various combinations of survey area (local scale) and landscape size (metacommunity scale). Simulations were conducted using landscapes of finite and of conceptually infinite size. We analyzed the scale-dependence of dispersal-diversity relationships for exponentially-bounded versus fat-tailed dispersal kernels, several levels of speciation rate and contrasting assumptions on recruitment at short dispersal distances. We found that the ratio of survey area to landscape size is a major determinant of dispersaldiversity relationships. With increasing survey-to-landscape area ratio the dispersaldiversity relationship switches from monotonically increasing through a U-shaped pattern (with a local minimum) to a monotonically decreasing pattern. Therefore, we provide a continuous set of dispersaldiversity relationships, which contains the response shapes reported previously as extreme cases. We suggest the mean dispersal distance with the minimum of species diversity (minimizing dispersal distance) for a certain scenario as a key characteristic of dispersaldiversity relationships. We show that not only increasing mean dispersal distances, but also increasing variances of dispersal can enhance diversity at the local scale, given a diverse species pool at the metacommunity scale. In conclusion, the response of diversity to variations of dispersal capability at spatial scales of interest, e.g. conservation areas, can differ more widely than expected previously. Therefore, land use and conservation activities, which manipulate dispersal capability, need to consider the landscape context and potential species pools carefully.
1. The complex, nonlinear response of dryland systems to grazing and climatic variations is a challenge to management of these lands. Predicted climatic changes will impact the desertification of drylands under domestic livestock production. Consequently, there is an urgent need to understand the response of drylands to grazing under climate change. 2. We enhanced and parameterized an ecohydrological savanna model to assess the impacts of a range of climate change scenarios on the response of a semi-arid African savanna to grazing. We focused on the effects of temperature and CO2 level increase in combination with changes in inter- and intra-annual precipitation patterns on the long-term dynamics of three major plant functional types. 3. We found that the capacity of the savanna to sustain livestock grazing was strongly influenced by climate change. Increased mean annual precipitation and changes in intra-annual precipitation pattern have the potential to slightly increase carrying capacities of the system. In contrast, decreased precipitation, higher interannual variation and temperature increase are leading to a severe decline of carrying capacities owing to losses of the perennial grass biomass. 4. Semi-arid rangelands will be at lower risk of shrub encroachment and encroachment will be less intense under future climatic conditions. This finding holds in spite of elevated levels of atmospheric CO2 and irrespective of changes in precipitation pattern, because of the drought sensitivity of germination and establishment of encroaching species. 5. Synthesis and applications. Changes in livestock carrying capacities, both positive and negative, mainly depend on the highly uncertain future rainfall conditions. However, independent of the specific changes, shrub encroachment becomes less likely and in many cases less severe. Thus, managers of semi-arid rangelands should shift their focus from woody vegetation towards perennial grass species as indicators for rangeland degradation. Furthermore, the resulting reduced competition from woody vegetation has the potential to facilitate ecosystem restoration measures such as re-introduction of desirable plant species that are only little promising or infeasible under current climatic conditions. On a global scale, the reductions in standing biomass resulting from altered degradation dynamics of semi-arid rangelands can have negative impacts on carbon sequestration.
Fragmentation and loss of habitat are major threats to animal communities and are therefore important to conservation. Due to the complexity of the interplay of spatial effects and community processes, our mechanistic understanding of how communities respond to such landscape changes is still poor. Modelling studies have mostly focused on elucidating the principles of community response to fragmentation and habitat loss at relatively large spatial and temporal scales relevant to metacommunity dynamics. Yet, it has been shown that also small scale processes, like foraging behaviour, space use by individuals and local resource competition are also important factors. However, most studies that consider these smaller scales are designed for single species and are characterized by high model complexity. Hence, they are not easily applicable to ecological communities of interacting individuals. To fill this gap, we apply an allometric model of individual home range formation to investigate the effects of habitat loss and fragmentation on mammal and bird communities, and, in this context, to investigate the role of interspecific competition and individual space use. Results show a similar response of both taxa to habitat loss. Community composition is shifted towards higher frequency of relatively small animals. The exponent and the 95%-quantile of the individual size distribution (ISD, described as a power law distribution) of the emerging communities show threshold behaviour with decreasing habitat area. Fragmentation per se has a similar and strong effect on mammals, but not on birds. The ISDs of bird communities were insensitive to fragmentation at the small scales considered here. These patterns can be explained by competitive release taking place in interacting animal communities, with the exception of bird's buffering response to fragmentation, presumably by adjusting the size of their home ranges. These results reflect consequences of higher mobility of birds compared to mammals of the same size and the importance of considering competitive interaction, particularly for mammal communities, in response to landscape fragmentation. Our allometric approach enables scaling up from individual physiology and foraging behaviour to terrestrial communities, and disentangling the role of individual space use and interspecific competition in controlling the response of mammal and bird communities to landscape changes.
Disturbances' role in shaping communities is well documented but highly disputed. We suggest replacing the overused two-trait trade-off approach with a functional group scheme, constructed from combinations of four key traits that represent four classes of species' responses to disturbances. Using model results and field observations from sites affected by two highly different disturbances, we demonstrated that popular dichotomous trade-offs are not sufficient to explain community dynamics, even if some emerge under certain conditions. Without disturbances, competition was only sufficient to predict species survival but not relative success, which required some escape mechanism (e.g., long-term dormancy). With highly predictable and large-scale disturbances, successful species showed a combination of high individual tolerance to disturbance and, more surprisingly, high competitive ability. When disturbances were less predictable, high individual tolerance and long-term seed dormancy were favored, due to higher environmental uncertainty. Our study demonstrates that theories relying on a small number of predefined trade-offs among traits (e.g., competition-colonization trade-off) may lead to unrealistic results. We suggest that the understanding of disturbance-community relationships can be significantly improved by employing sets of relevant trait assemblies instead of the currently common approach in which trade-offs are assumed in advance.
Aim To assess how habitat loss and climate change interact in affecting the range dynamics of species and to quantify how predicted range dynamics depend on demographic properties of species and the severity of environmental change. Location South African Cape Floristic Region. Methods We use data-driven demographic models to assess the impacts of past habitat loss and future climate change on range size, range filing and abundances of eight species of woody plants (Proteaceae). The species-specific models employ a hybrid approach that simulates population dynamics and long-distance dispersal on top of expected spatio-temporal dynamics of suitable habitat. Results Climate change was mainly predicted to reduce range size and range filling (because of a combination of strong habitat shifts with low migration ability). In contrast, habitat loss mostly decreased mean local abundance. For most species and response measures, the combination of habitat loss and climate change had the most severe effect. Yet, this combined effect was mostly smaller than expected from adding or multiplying effects of the individual environmental drivers. This seems to be because climate change shifts suitable habitats to regions less affected by habitat loss. Interspecific variation in range size responses depended mostly on the severity of environmental change, whereas responses in range filling and local abundance depended mostly on demographic properties of species. While most surviving populations concentrated in areas that remain climatically suitable, refugia for multiple species were overestimated by simply overlying habitat models and ignoring demography. Main conclusions Demographic models of range dynamics can simultaneously predict the response of range size, abundance and range filling to multiple drivers of environmental change. Demographic knowledge is particularly needed to predict abundance responses and to identify areas that can serve as biodiversity refugia under climate change. These findings highlight the need for data-driven, demographic assessments in conservation biogeography.
Semi-natural grasslands, biodiversity hotspots in Central-Europe, suffer from the cessation of traditional land-use. Amount and intensity of these changes challenge current monitoring frameworks typically based on classic indicators such as selected target species or diversity indices. Indicators based on plant functional traits provide an interesting extension since they reflect ecological strategies at individual and ecological processes at community levels. They typically show convergent responses to gradients of land-use intensity over scales and regions, are more directly related to environmental drivers than diversity components themselves and enable detecting directional changes in whole community dynamics. However, probably due to their labor- and cost intensive assessment in the field, they have been rarely applied as indicators so far.
Here we suggest overcoming these limitations by calculating indicators with plant traits derived from online accessible databases. Aiming to provide a minimal trait set to monitor effects of land-use intensification on plant diversity we investigated relationships between 12 community mean traits, 2 diversity indices and 6 predictors of land-use intensity within grassland communities of 3 different regions in Germany (part of the German 'Biodiversity Exploratory' research network). By standardization of traits and diversity measures, use of null models and linear mixed models we confirmed (i) strong links between functional community composition and plant diversity, (ii) that traits are closely related to land-use intensity, and (iii) that functional indicators are equally, or even more sensitive to land-use intensity than traditional diversity indices. The deduced trait set consisted of 5 traits, i.e., specific leaf area (SLA), leaf dry matter content (LDMC), seed release height, leaf distribution, and onset of flowering. These database derived traits enable the early detection of changes in community structure indicative for future diversity loss. As an addition to current monitoring measures they allow to better link environmental drivers to processes controlling community dynamics.
Understanding the regional dynamics of plant communities is crucial for predicting the response of plant diversity to habitat fragmentation. However, for fragmented landscapes the importance of regional processes, such as seed dispersal among isolated habitat patches, has been controversially debated. Due to the stochasticity and rarity of among-patch dispersal and colonization events, we still lack a quantitative understanding of the consequences of these processes at the landscape-scale. In this study, we used extensive field data from a fragmented, semi-arid landscape in Israel to parameterize a multi-species incidence-function model. This model simulates species occupancy pattern based on patch areas and habitat configuration and explicitly considers the locations and the shapes of habitat patches for the derivation of patch connectivity. We implemented an approximate Bayesian computation approach for parameter inference and uncertainty assessment. We tested which of the three types of regional dynamics - the metacommunity, the mainland-island, or the island communities type - best represents the community dynamics in the study area and applied the simulation model to estimate the extinction debt in the investigated landscape. We found that the regional dynamics in the patch-matrix study landscape is best represented as a system of highly isolated island' communities with low rates of propagule exchange among habitat patches and consequently low colonization rates in local communities. Accordingly, the extinction rates in the local communities are the main drivers of community dynamics. Our findings indicate that the landscape carries a significant extinction debt and in model projections 33-60% of all species went extinct within 1000 yr. Our study demonstrates that the combination of dynamic simulation models with field data provides a promising approach for understanding regional community dynamics and for projecting community responses to habitat fragmentation. The approach bears the potential for efficient tests of conservation activities aimed at mitigating future losses of biodiversity.
Quantifying the association of plant functional traits to environmental gradients is a promising approach for understanding and projecting community responses to land use and climatic changes. Although habitat fragmentation and climate are expected to affect plant communities interactively, there is a lack of empirical studies addressing trait associations to fragmentation in different climatic regimes.
In this study, we analyse data on the key functional traits: specific leaf area (SLA), plant height, seed mass and seed number. First, we assess the evidence for the community assembly mechanisms habitat filtering and competition at different spatial scales, using several null-models and a comprehensive set of community-level trait convergence and divergence indices. Second, we analyse the association of community-mean traits with patch area and connectivity along a south-north productivity gradient.
We found clear evidence for trait convergence due to habitat filtering. In contrast, the evidence for trait divergence due to competition fundamentally depended on the null-model used. When the null-model controlled for habitat filtering, there was only evidence for trait divergence at the smallest sampling scale (0.25 m x 0.25 m). All traits varied significantly along the S-N productivity gradient. While plant height and SLA were consistently associated with fragmentation, the association of seed mass and seed number with fragmentation changed along the S-N gradient.
Our findings indicate trait convergence due to drought stress in the arid sites and due to higher productivity in the mesic sites. The association of plant traits to fragmentation is likely driven by increased colonization ability in small and/or isolated patches (plant height, seed number) or increased persistence ability in isolated patches (seed mass).
Our study provides the first empirical test of trait associations with fragmentation along a productivity gradient. We conclude that it is crucial to study the interactive effects of different ecological drivers on plant functional traits.
2. We present a hierarchical model that integrates observations from multiple sources to estimate spatio-temporal abundance trends. The model links annual population densities on a spatial grid to both long-term count data and to opportunistic occurrence records from a citizen science programme. Specific observation models for both data types explicitly account for differences in data structure and quality.
3. We test this novel method in a virtual study with simulated data and apply it to the estimation of abundance dynamics across the range of a butterfly species (Pyronia tithonus) in Great Britain between 1985 and 2004. The application to simulated and real data demonstrates how the hierarchical model structure accommodates various sources of uncertainty which occur at different stages of the link between observational data and the modelled abundance, thereby it accounts for these uncertainties in the inference of abundance variations.
4. We show that by using hierarchical observation models that integrate different types of commonly available data sources, we can improve the estimates of variation in species abundances across space and time. This will improve our ability to detect regional trends and can also enhance the empirical basis for understanding range dynamics.
Patterns of past vegetation changes over time and space can help facilitate better understanding of the interactions among climate, ecosystem, and human impact. Biome changes in China over the last 22,000 yr (calibrated radiocarbon date, a BP) were numerically reconstructed by using a standard approach of pollen-plant functional type-biome assignment (biomization). The biomization procedure involves pollen data from 2434 surface sites and 228 fossil sites with a high quality of pollen count and C-14 dating, 51 natural and three anthropogenic plant functional types (PFTs), as well as 19 natural and one anthropogenic biome. Surface pollen-based reconstruction of modern natural biome patterns is in good agreement (74.4%) with actual vegetation distribution in China. However, modem large-scale anthropogenic biome reconstruction has not been successful based on the current setup of three anthropogenic PFTs (plantation, secondary, and disturbed PFT) because of the limitation of non-species level pollen identification and the difficulty in the clear assignment of disturbed PFTs. The non-anthropogenic biome distributions of 44 time slices at 500-year intervals show large-scale discrepant and changed vegetation patterns from the last glacial maximum (LGM) to the Holocene throughout China. From 22 ka BP to 19 ka BP, temperate grassland, xerophytic shrubland, and desert dominated northern China, whereas cold or cool forests flourished in central China. Warm-temperate evergreen forests were restricted to far southern China, and tropical forests were absent During 18.5 ka BP to 12 ka BP, cold, cool, and dry biomes extended to some parts of northern, westem, and eastern China. Warm-temperate evergreen and mixed forests gradually expanded to occupy the whole of southern China. A slight northward shift of forest biomes occurred from 15 ka BP to 12 lea BP. During 11.5 ka BP to 9 ka BP, temperate grassland and shrubland gradually stretched to northern and western China. Cold and cool forests widely expanded into northern and central China, as well as in the northern margin of South China along with temperate deciduous forest. Since the early mid-Holocene (approximately 8.5 ka BP to 5.5 ka BP), all forest biomes shifted northward at the expense of herbaceous and shrubby biomes. Simultaneously, cold and cool forest biomes occupied the marginal areas of the Tibetan Plateau and the high mountains in western China. During the middle to late Holocene, from 5 ka to the present, temperate grassland and xerophytic shrubland expanded to the south and east, whereas temperate deciduous forests slightly shifted southward. After 3 lea BP, forest biomes were absent in western China and on the Tibetan plateau surface. Dramatic biome shifts from the LGM to the Holocene were observed in the forest-grassland ecotone and transitional zones between temperate and subtropical climates, between subtropical and tropical regions, and in the mountainous margins of the eastern Tibetan Plateau. Evidence showed more human disturbances during the late Holocene. More pollen records and historical documents are therefore further needed to understand fully the human disturbance-induced large-scale forest changes. In addition, more classifications of anthropogenic biome or land cover, more distinct assignment of pollen taxa to anthropogenic PFTs, and more effective numerical and/or mechanistic techniques in building large-scale human disturbances are required. (C) 2014 Elsevier B.V. All rights reserved.
Identifying drivers of species diversity is a major challenge in understanding and predicting the dynamics of species-rich semi-natural grasslands. In particular in temperate grasslands changes in land use and its consequences, i.e. increasing fragmentation, the on-going loss of habitat and the declining importance of regional processes such as seed dispersal by livestock, are considered key drivers of the diversity loss witnessed within the last decades.
Savanna rangelands worldwide are threatened by shrub encroachment, i.e. the increase of woody plant species at the cost of perennial grasses, causing a strong decline in the productivity of domestic livestock production. Although recent studies indicate that fire might be of great importance for semi-arid and arid savanna dynamics, it is largely not applied in the management of semi-arid rangelands especially with regard to woody plant control. We used the eco-hydrological savanna model EcoHyD to simulate the effects of different fire management strategies on semi-arid savanna vegetation and to assess their longterm suitability for semi-arid rangeland management. Simulation results show that prescribed fires, timed to kill tree seedlings prevented shrub encroachment for a broad range of livestock densities while the possible maximum long-term cattle densities on the simulated semi-arid rangeland in Namibia increased by more than 30%. However, when grazing intensity was too high, fire management failed in preventing shrub encroachment.
Our findings indicate that with regard to fire management a clear distinction between mesic and more arid savannas is necessary: While the frequency of fires is of relevance for mesic savannas, we recommend a fire management focussing on the timing of fire for semi-arid and arid savannas. (C) 2014 Elsevier Ltd. All rights reserved.
Our current understanding regarding the functioning of the savanna ecosystem describes savannas as either competition- or disturbance-dependent. Within this generalized view, the role and importance of facilitation have been mostly neglected. This study presents a mathematical model of savannas with coupled soil moisture-vegetation dynamics, which includes interspecific competition and environmental disturbance. We find that there exist environmental and climatic conditions where grass facilitation toward trees plays an important role in supporting tree cover and by extension preserving the savanna biome. We, therefore, argue that our theoretical results in combination with the first empirical studies on the subject should stimulate further research into the role of facilitation in the savanna ecosystem, particularly when analyzing the impact of past and projected climatic changes on it. (C) 2015 Elsevier B.V. All rights reserved.
Small scale distribution of insect root herbivores may promote plant species diversity by creating patches of different herbivore pressure. However, determinants of small scale distribution of insect root herbivores, and impact of land use intensity on their small scale distribution are largely unknown. We sampled insect root herbivores and measured vegetation parameters and soil water content along transects in grasslands of different management intensity in three regions in Germany. We calculated community-weighted mean plant traits to test whether the functional plant community composition determines the small scale distribution of insect root herbivores. To analyze spatial patterns in plant species and trait composition and insect root herbivore abundance we computed Mantel correlograms. Insect root herbivores mainly comprised click beetle (Coleoptera, Elateridae) larvae (43%) in the investigated grasslands. Total insect root herbivore numbers were positively related to community-weighted mean traits indicating high plant growth rates and biomass (specific leaf area, reproductive-and vegetative plant height), and negatively related to plant traits indicating poor tissue quality (leaf C/N ratio). Generalist Elaterid larvae, when analyzed independently, were also positively related to high plant growth rates and furthermore to root dry mass, but were not related to tissue quality. Insect root herbivore numbers were not related to plant cover, plant species richness and soil water content. Plant species composition and to a lesser extent plant trait composition displayed spatial autocorrelation, which was not influenced by land use intensity. Insect root herbivore abundance was not spatially autocorrelated. We conclude that in semi-natural grasslands with a high share of generalist insect root herbivores, insect root herbivores affiliate with large, fast growing plants, presumably because of availability of high quantities of food. Affiliation of insect root herbivores with large, fast growing plants may counteract dominance of those species, thus promoting plant diversity.
Density regulation influences population dynamics through its effects on demographic rates and consequently constitutes a key mechanism explaining the response of organisms to environmental changes. Yet, it is difficult to establish the exact form of density dependence from empirical data. Here, we developed an individual-based model to explore how resource limitation and behavioural processes determine the spatial structure of white stork Ciconia ciconia populations and regulate reproductive rates. We found that the form of density dependence differed considerably between landscapes with the same overall resource availability and between home range selection strategies, highlighting the importance of fine-scale resource distribution in interaction with behaviour. In accordance with theories of density dependence, breeding output generally decreased with density but this effect was highly variable and strongly affected by optimal foraging strategy, resource detection probability and colonial behaviour. Moreover, our results uncovered an overlooked consequence of density dependence by showing that high early nestling mortality in storks, assumed to be the outcome of harsh weather, may actually result from density dependent effects on food provision. Our findings emphasize that accounting for interactive effects of individual behaviour and local environmental factors is crucial for understanding density-dependent processes within spatially structured populations. Enhanced understanding of the ways animal populations are regulated in general, and how habitat conditions and behaviour may dictate spatial population structure and demographic rates is critically needed for predicting the dynamics of populations, communities and ecosystems under changing environmental conditions.
Conservation actions need to account for global climate change and adapt to it. The body of the literature on adaptation options is growing rapidly, but their feasibility and current state of implementation are rarely assessed. We discussed the practicability of adaptation options with conservation managers analysing three fields of action: reducing the vulnerability of conservation management, reducing the vulnerability of conservation targets (i.e. biodiversity) and climate change mitigation. For all options, feasibility, current state of implementation and existing obstacles to implementation were analysed, using the Federal State of Brandenburg, Germany, as a case study. Practitioners considered a large number of options useful, most of which have already been implemented at least in part. Those options considered broadly implemented resemble mainly conventional measures of conservation without direct relation to climate change. Managers are facing several obstacles for adapting to climate change, including political reluctance to change, financial and staff shortages in conservation administrations and conflictive EU funding schemes in agriculture. A certain reluctance to act, due to the high degree of uncertainty with regard to climate change scenarios and impacts, is widespread. A lack of knowledge of appropriate methods such as adaptive management often inhibits the implementation of adaptation options in the field of planning and management. Based on the findings for Brandenburg, we generally conclude that it is necessary to focus in particular on options that help to reduce vulnerability of conservation management itself, i.e. those that enhance management effectiveness. For instance, adaptive and proactive risk management can be applied as a no-regrets option, independently from specific climate change scenarios or impacts, strengthening action under uncertainty.
Changes in land-use are supposed to be among the severest prospective threats to plant diversity worldwide. In semi-natural temperate grasslands, the cessation of traditional land use like livestock grazing is considered to be one of the most important drivers of the diversity loss witnessed within the last decades. Despite of the enormous number of studies on successional pathways following grazing abandonment there is no general pattern of how grassland communities are affected in terms of diversity, trait composition and pace of succession. To gain a comprehensive picture is difficult given the heterogeneity of environments and the time and effort needed for long-term investigations. We here use a proven individual- and trait-based grassland community model to analyze short- and long-term consequences of grazing abandonment under different assumptions of resource availability, pre-abandonment grazing intensity and regional isolation of communities.
Grazing abandonment led to a decrease of plant functional type (PFT) diversity in all but two scenarios in the long-term. In short-term we also found an increase or no change in Shannon diversity for several scenarios. With grazing abandonment we overall found an increase in maximum plant mass, clonal integration and longer lateral spread, a decrease in rosette plant types and in stress tolerant plants, as well as an increase in grazing tolerant and a decrease in grazing avoiding plant types. Observed changes were highly dependent on the regional configuration of communities, prevalent resource conditions and land use intensity before abandonment. While long-term changes took around 10-20 years in resource rich conditions, new equilibria established in resource poor conditions only after 30-40 years.
Our results confirm the potential threats caused by recent land-use changes and the assumption that oligotrophic communities are more resistant than mesotrophic communities also for long-term abandonment. Moreover, results revealed that species-rich systems are not per se more resistant than species-poor grasslands. (C) 2015 Elsevier B.V. All rights reserved.
Fertilization causes species loss and species dominance changes in plant communities worldwide. However, it still remains unclear how fertilization acts upon species functional traits, e.g. seed mass. Seed mass is a key trait of the regeneration strategy of plants, which influences a range of processes during the seedling establishment phase. Fertilization may select upon seed mass, either directly by increased nutrient availability or indirectly by increased competition. Since previous research has mainly analyzed the indirect effects of fertilization, we disentangled the direct and indirect effects to examine how nutrient availability and competition influence the seed mass relationships on four key components during seedling establishment: seedling emergence, time of seedling emergence, seedling survival and seedling growth. We conducted a common garden experiment with 22 dry grassland species with a two-way full factorial design that simulated additional nutrient supply and increased competition. While we found no evidence that fertilization either directly by additional nutrient supply or indirectly by increased competition alters the relationship between seed mass and (time of) seedling emergence, we revealed that large seed mass is beneficial under nutrient-poor conditions (seedlings have greater chances of survival, particularly in nutrient-poor soils) as well as under competition (large-seeded species produced larger seedlings, which suffered less from competition than small-seeded species). Based on these findings, we argue that both factors, i.e. nutrient availability and competition intensity, ought to be considered to understand how fertilization influences seedling establishment and species composition with respect to seed mass in natural communities. We propose a simple conceptual model, in which seed mass in natural communities is determined by competition intensity and nutrient availability. Here, we hypothesize that seed mass shows a U-shaped pattern along gradients of soil fertility, which may explain the contrasting soil fertility-seed mass relationships found in the recent literature.
East Africa hosts a striking diversity of terrestrial ecosystems, which vary both in space and time due to complex regional topography and a dynamic climate. The structure and functioning of these ecosystems under this environmental setting can be studied with dynamic vegetation models (DVMs) in a spatially explicit way. Yet, regional applications of DVMs to East Africa are rare and a comprehensive validation of such applications is missing. Here, we simulated the present-day and mid-Holocene vegetation of East Africa with the DVM, LPJ-GUESS and we conducted an exhaustive comparison of model outputs with maps of potential modern vegetation distribution, and with pollen records of local change through time. Overall, the model was able to reproduce the observed spatial patterns of East African vegetation. To see whether running the model at higher spatial resolutions (10′ × 10′) contribute to resolve the vegetation distribution better and have a better comparison scale with the observational data (i.e. pollen data), we run the model with coarser spatial resolution (0.5° × 0.5°) for the present-day as well. Both the area- and point-wise comparison showed that a higher spatial resolution allows to better describe spatial vegetation changes induced by the complex topography of East Africa. Our analysis of the difference between modelled mid-Holocene and modern-day vegetation showed that whether a biome shifts to another is best explained by both the amount of change in precipitation it experiences and the amount of precipitation it received originally. We also confirmed that tropical forest biomes were more sensitive to a decrease in precipitation compared to woodland and savanna biomes and that Holocene vegetation changes in East Africa were driven not only by changes in annual precipitation but also by changes in its seasonality.
QuestionThe empirical evidence of root herbivory effects on plant community composition and co-existence is contradictory. This originates from difficulties connected to below-ground research and confinement of experimental studies to a small range of environmental conditions. Here we suggest coupling experimental data with an individual-based model to overcome the limitations inherent in either approach. To demonstrate this, we investigated the consequences of root herbivory, as experimentally observed on individual plants, on plant competition and co-existence in a population and community context under different root herbivory intensities (RHI), fluctuating and constant root herbivore activity and grazing along a resource gradient. LocationBerlin, Germany, glasshouse; Potsdam, Germany, high performance cluster computer. MethodsThe well-established community model IBC-Grass was adapted to allow for a flexible species parameterization and to include annual species. Experimentally observed root herbivory effects on performance of eight common grassland plant species were incorporated into the model by altering plant growth rates. We then determined root herbivore effects on plant populations, competitive hierarchy and consequences for co-existence and community diversity. ResultsRoot herbivory reduced individual biomass, but temporal fluctuation allowed for compensation of herbivore effects. Reducing resource availability strongly shifted competitive hierarchies, with, however, more similar hierarchies along the gradient under root herbivory, pointing to reduced ecological species differences. Consequently, negative effects on co-existence and diversity prevailed, with the exception of a few positive effects on co-existence of selected species pairs. Temporal fluctuation alleviated but did not remove negative root herbivore effects, despite of the stronger influence of intra- compared to interspecific competition. Grazing in general augmented co-existence. Most interestingly, grazing interacted with RHI and resource availability by promoting positive effects of root herbivory. ConclusionsThrough integrating experimental data on the scale of individual plants with a simulation model we verified that root herbivory could affect plant competition with consequences for species co-existence. Our approach demonstrates the benefit that accrues when empirical and modelling approaches are brought more closely together, and that gathering data on distinct processes and under specific conditions, combined with appropriate models, can be used to answer challenging research questions in a more general way.
Although many birds are socially monogamous, most (>75%) studied species are not strictly genetically monogamous, especially under high breeding density. We used molecular tools to reevaluate the reproductive strategy of the socially monogamous white stork (Ciconia ciconia) and examined local density effects. DNA samples of nestlings (Germany, Spain) were genotyped and assigned relationships using a two-program maximum likelihood classification. Relationships were successfully classified in 79.2% of German (n = 120) and 84.8% of Spanish (n = 59) nests. For each population respectively, 76.8% (n = 73) and 66.0% (n = 33) of nests contained only full-siblings, 10.5% (n = 10) and 18.0% (n = 9) had half-siblings (at least one nestling with a different parent), 3.2% (n = 3) and 10.0% (n = 5) had unrelated nestlings (at least two nestlings, each with different parents), and 9.5% (n = 9) and 6.0% (n = 3) had “not full-siblings” (could not differentiate between latter two cases). These deviations from strict monogamy place the white stork in the 59th percentile for extra-pair paternity among studied bird species. Although high breeding density generally increases extra-pair paternity, we found no significant association with this species’ mating strategies. Thus although genetic monogamy is indeed prominent in the white stork, extra-pair paternity is fairly common compared to other bird species and cannot be explained by breeding density.
The abandonment of military areas leads to succession processes affecting valuable open-land habitats and is considered to be a major threat for European butterflies. We assessed the ability of hyper spectral remote sensing data to spatially predict the occurrence of one of the most endangered butterfly species (Hipparchia statilinus) in Brandenburg (Germany) on the basis of habitat characteristics at a former military training area. Presence-absence data were sampled on a total area of 36 km(2), and N = 65 adult individuals of Hipparchia statilinus could be detected. The floristic composition within the study area was modeled in a three-dimensional ordination space. Occurrence probabilities for the target species were predicted as niches between ordinated floristic gradients by using Regression Kriging of Indicators. Habitat variance could be explained by up to 81 % with spectral variables at a spatial resolution of 2 x 2 m by transferring PLSR models to imagery. Ordinated ecological niche of Hipparchia statilinus was tested against environmental predictor variables. N = 6 variables could be detected to be significantly correlated with habitat preferences of Hipparchia statilinus. They show that Hipparchia statilinus can serve as a valuable indicator for the evaluation of the conservation status of Natura 2000 habitat type 2330 (inland dunes with open Corynephorus and Agrostis grasslands) protected by the Habitat Directive (Council Directive 92/43/EEC). The authors of this approach, conducted in August 2013 at Doberitzer Heide Germany, aim to increase the value of remote sensing as an important tool for questions of biodiversity research and conservation.
1. Migration conveys an immense challenge, especially for juvenile birds coping with enduring and risky journeys shortly after fledging. Accordingly, juveniles exhibit considerably lower survival rates compared to adults, particularly during migration. Juvenile white storks (Ciconia ciconia), which are known to rely on adults during their first fall migration presumably for navigational purposes, also display much lower annual survival than adults.
2. Using detailed GPS and body acceleration data, we examined the patterns and potential causes of age-related differences in fall migration properties of white storks by comparing first-year juveniles and adults. We compared juvenile and adult parameters of movement, behaviour and energy expenditure (estimated from overall dynamic body acceleration) and placed this in the context of the juveniles’ lower survival rate.
3. Juveniles used flapping flight vs. soaring flight 23% more than adults and were estimated to expend 14% more energy during flight. Juveniles did not compensate for their higher flight costs by increased refuelling or resting during migration. When juveniles and adults migrated together in the same flock, the juvenile flew mostly behind the adult and was left behind when they separated. Juveniles showed greater improvement in flight efficiency throughout migration compared to adults which appears crucial because juveniles exhibiting higher flight costs suffered increased mortality.
4. Our findings demonstrate the conflict between the juveniles’ inferior flight skills and their urge to keep up with mixed adult–juvenile flocks. We suggest that increased flight costs are an important proximate cause of juvenile mortality in white storks and likely in other soaring migrants and that natural selection is operating on juvenile variation in flight efficiency.
Climate change and land use management practices are major drivers of biodiversity in terrestrial ecosystems. To understand and predict resulting changes in community structures, individual-based and spatially explicit population models are a useful tool but require detailed data sets for each species. More generic approaches are thus needed. Here we present a trait-based functional type approach to model savanna birds. The aim of our model is to explore the response of different bird functional types to modifications in habitat structure. The functional types are characterized by different traits, in particular body mass, which is related to life-history traits (reproduction and mortality) and spatial scales (home range area and dispersal ability), as well as the use of vegetation structures for foraging and nesting, which is related to habitat quality and suitability. We tested the performance of the functional types in artificial landscapes varying in shrub:grass ratio and clumping intensity of shrub patches. We found that an increase in shrub encroachment and a decrease in habitat quality caused by land use mismanagement and climate change endangered all simulated bird functional types. The strength of this effect was related to the preferred habitat. Furthermore, larger-bodied insectivores and omnivores were more prone to extinction due to shrub encroachment compared to small-bodied species. Insectivorous and omnivorous birds were more sensitive to clumping intensity of shrubs whereas herbivorous and carnivorous birds were most affected by a decreasing amount of grass cover. From an applied point of view, our findings emphasize that policies such as woody plant removal and a reduction in livestock stocking rates to prevent shrub encroachment should prioritize the enlargement of existing grassland patches. Overall, our results show that the combination of an individual-based modelling approach with carefully defined functional types can provide a powerful tool for exploring biodiversity responses to environmental changes. Furthermore, the increasing accumulation of worldwide data sets on species’ core and soft traits (surrogates to determine core traits indirectly) on one side and the refinement of conceptual frameworks for animal functional types on the other side will further improve functional type approaches which consider the sensitivities of multiple species to climate change, habitat loss, and fragmentation.
The El Nino-Southern Oscillation (ENSO) is the main driver of the interannual variability in eastern African rainfall, with a significant impact on vegetation and agriculture and dire consequences for food and social security. In this study, we identify and quantify the ENSO contribution to the eastern African rainfall variability to forecast future eastern African vegetation response to rainfall variability related to a predicted intensified ENSO. To differentiate the vegetation variability due to ENSO, we removed the ENSO signal from the climate data using empirical orthogonal teleconnection (EOT) analysis. Then, we simulated the ecosystem carbon and water fluxes under the historical climate without components related to ENSO teleconnections. We found ENSO-driven patterns in vegetation response and confirmed that EOT analysis can successfully produce coupled tropical Pacific sea surface temperature-eastern African rainfall teleconnection from observed datasets. We further simulated eastern African vegetation response under future climate change as it is projected by climate models and under future climate change combined with a predicted increased ENSO intensity. Our EOT analysis highlights that climate simulations are still not good at capturing rainfall variability due to ENSO, and as we show here the future vegetation would be different from what is simulated under these climate model outputs lacking accurate ENSO contribution. We simulated considerable differences in eastern African vegetation growth under the influence of an intensified ENSO regime which will bring further environmental stress to a region with a reduced capacity to adapt effects of global climate change and food security.
Natural grassland communities are threatened by a variety of factors, such as climate change and increasing land use by mankind. The use of plant protection products (synthetic or organic) is mandatory in agricultural food production. To avoid adverse effects on natural grasslands within agricultural areas, synthetic plant protection products are strictly regulated in Europe. However, effects of herbicides on non-target terrestrial plants are primarily studied on the level of individual plants neglecting interactions between species. In our study, we aim to extrapolate individual-level effects to the population and community level by adapting an existing spatio-temporal, individual-based plant community model (IBC-grass). We analyse the effects of herbicide exposure for three different grassland communities: 1) representative field boundary community, 2) Calthion grassland community, and 3) Arrhenatheretalia grassland community. Our simulations show that herbicide depositions can have effects on non-target plant communities resulting from direct and indirect effects on population level. The effect extent depends not only on the distance to the field, but also on the specific plant community, its disturbance regime (cutting frequency, trampling and grazing intensity) and resource level. Mechanistic modelling approaches such as IBC-grass present a promising novel approach in transferring and extrapolating standardized pot experiments to community level and thereby bridging the gap between ecotoxicological testing (e.g. in the greenhouse) and protection goals referring to real world conditions.
The El Nino-Southern Oscillation (ENSO) is the main driver of the interannual variability in eastern African rainfall, with a significant impact on vegetation and agriculture and dire consequences for food and social security. In this study, we identify and quantify the ENSO contribution to the eastern African rainfall variability to forecast future eastern African vegetation response to rainfall variability related to a predicted intensified ENSO. To differentiate the vegetation variability due to ENSO, we removed the ENSO signal from the climate data using empirical orthogonal teleconnection (EOT) analysis. Then, we simulated the ecosystem carbon and water fluxes under the historical climate without components related to ENSO teleconnections. We found ENSO-driven patterns in vegetation response and confirmed that EOT analysis can successfully produce coupled tropical Pacific sea surface temperature-eastern African rainfall teleconnection from observed datasets. We further simulated eastern African vegetation response under future climate change as it is projected by climate models and under future climate change combined with a predicted increased ENSO intensity. Our EOT analysis highlights that climate simulations are still not good at capturing rainfall variability due to ENSO, and as we show here the future vegetation would be different from what is simulated under these climate model outputs lacking accurate ENSO contribution. We simulated considerable differences in eastern African vegetation growth under the influence of an intensified ENSO regime which will bring further environmental stress to a region with a reduced capacity to adapt effects of global climate change and food security.
Protected areas are arguably the most important instrument of biodiversity conservation. To keep them fit under climate change, their management needs to be adapted to address related direct and indirect changes. In our study we focus on the adaptation of conservation management planning, evaluating management plans of 60 protected areas throughout Germany with regard to their climate change-robustness. First, climate change-robust conservation management was defined using 11 principles and 44 criteria, which followed an approach similar to sustainability standards. We then evaluated the performance of individual management plans concerning the climate change-robustness framework. We found that climate change-robustness of protected areas hardly exceeded 50 percent of the potential performance, with most plans ranking in the lower quarter. Most Natura 2000 protected areas, established under conservation legislation of the European Union, belong to the sites with especially poor performance, with lower values in smaller areas. In general, the individual principles showed very different rates of accordance with our principles, but similarly low intensity. Principles with generally higher performance values included holistic knowledge management, public accountability and acceptance as well as systemic and strategic coherence. Deficiencies were connected to dealing with the future and uncertainty. Lastly, we recommended the presented principles and criteria as essential guideposts that can be used as a checklist for working towards more climate change-robust planning.
Intraspecific trait variability plays an important role in species adaptation to climate change. However, it still remains unclear how plants in semi-arid environments respond to increasing aridity. We investigated the intraspecific trait variability of two common Mediterranean annuals (Geropogon hybridus and Crupina crupinastrum) with similar habitat preferences. They were studied along a steep precipitation gradient in Israel similar to the maximum predicted precipitation changes in the eastern Mediterranean basin (i.e. -30% until 2100). We expected a shift from competitive ability to stress tolerance with decreasing precipitation and tested this expectation by measuring key functional traits (canopy and seed release height, specific leaf area, N-and P-leaf content, seed mass). Further, we evaluated generative bet-hedging strategies by different seed traits. Both species showed different responses along the precipitation gradient. C. crupinastrum exhibited only decreased plant height toward saridity, while G. hybridus showed strong trends of generative adaptation to aridity. Different seed trait indices suggest increased bet-hedging of G. hybridus in arid environments. However, no clear trends along the precipitation gradient were observed in leaf traits (specific leaf area and leaf N-/P-content) in both species. Moreover, variance decomposition revealed that most of the observed trait variation (>> 50%) is found within populations. The findings of our study suggest that responses to increased aridity are highly species-specific and local environmental factors may have a stronger effect on intraspecific trait variation than shifts in annual precipitation. We therefore argue that trait-based analyses should focus on precipitation gradients that are comparable to predicted precipitation changes and compare precipitation effects to effects of local environmental factors. (C) 2017 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.
Spatial environmental heterogeneity is considered a fundamental factor for the maintenance of plant species richness. However, it still remains unclear whether heterogeneity may also facilitate coexistence at fine grain sizes or whether other processes, like mass effects and source sink dynamics due to dispersal, control species composition and diversity at these scales. In this study, we used two complimentary analyses to identify the role of heterogeneity within 15 m x 15 m plots for the coexistence of species-rich annual communities in a semi-arid environment along a steep precipitation gradient. Specifically, we: (a) analyzed the effect of environmental heterogeneity on species, functional and phylogenetic diversity within microsites (alpha diversity, 0.06 m(2) and 1 m(2)), across microsites (beta diversity), and diversity at the entire plot (gamma diversity); (b) further we used two null models to detect non-random trait and phylogenetic patterns in order to infer assembly processes, i.e. whether co-occurring species tend to share similar traits (trait convergence) or dissimilar traits (trait divergence). In general, our results showed that heterogeneity had a positive effect on community diversity. Specifically, for alpha diversity, the effect was significant for functional diversity, and not significant for either species or phylogenetic diversities. For beta diversity, all three measures of community diversity (species, functional, and phylogenetic) increased significantly, as they also did for gamma diversity, where functional measures were again stronger than for species or phylogenetic measures. In addition, the null model approach consistently detected trait convergence, indicating that species with similar traits tended to co-occur and had high abundances in a given microsite. While null model analysis across the phylogeny partly supported these trait findings, showing phylogenetic underdispersion at the 1m(2) grain size, surprisingly when species abundances in microsites were analyzed they were more evenly distributed across the phylogenetic tress than expected (phylogenetic overdispersion). In conclusion, our results provide compelling support that environmental heterogeneity at a relatively fine scale is an important factor for species co-existence as it positively affects diversity as well as influences species assembly. Our study underlines the need for trait-based approaches conducted at fine grain sizes in order to better understand species coexistence and community assembly. (C) 2017 Elsevier GmbH. All rights reserved.
Periodic environments determine the life cycle of many animals across the globe and the timing of important life history events, such as reproduction and migration. These adaptive behavioural strategies are complex and can only be fully understood (and predicted) within the framework of natural selection in which species adopt evolutionary stable strategies. We present sOAR, a powerful and user-friendly implementation of the well-established framework of optimal annual routine modelling. It allows determining optimal animal life history strategies under cyclic environmental conditions using stochastic dynamic programming. It further includes the simulation of population dynamics under the optimal strategy. sOAR provides an important tool for theoretical studies on the behavioural and evolutionary ecology of animals. It is especially suited for studying bird migration. In particular, we integrated options to differentiate between costs of active and passive flight into the optimal annual routine modelling framework, as well as options to consider periodic wind conditions affecting flight energetics. We provide an illustrative example of sOAR where food supply in the wintering habitat of migratory birds significantly alters the optimal timing of migration. sOAR helps improving our understanding of how complex behaviours evolve and how behavioural decisions are constrained by internal and external factors experienced by the animal. Such knowledge is crucial for anticipating potential species’ response to global environmental change.