Showing papers in "Oikos in 2003"

Variation in trophic shift for stable isotope ratios of carbon, nitrogen, and sulfur

Abstract: Use of stable isotope ratios to trace pathways of organic matter among consumers requires knowledge of the isotopic shift between diet and consumer. Variation in trophic shift among consumers can be substantial. For data from the published literature and supplementary original data (excluding fluid-feeding consumers), the mean isotopic shift for C was + 0.5 + 0.13%o rather than 0.0%o, as commonly assumed. The shift for C was higher for consumers analyzed as muscle (+ 1.3 + 0.30%o) than for consumers analyzed whole (+ 0.3 +0.14%o). Among consumers analyzed whole, the trophic shift for C was lower for consumers acidified prior to analysis (-0.2 + 0.21%o) than for unacidified samples ( +0.5 + 0.17%o). For N, trophic shift was lower for consumers raised on invertebrate diets (+ 1.4 + 0.21%o) than for consumers raised on other high-protein diets (+3.3 +0.26%o) and was intermediate for consumers raised on plant and algal diets (+2.2 +0.30%o). The trophic shift for S differed between high-protein (+ 2.0 + 0.65%o) and low-protein diets (-0.5 + 0.56%o). Thus, methods of analysis and dietary differences can affect trophic shift for consumers; the utility of stable isotope methods can be improved if this information is incorporated into studies of trophic relationships. Although few studies of stable isotope ratios have considered variation in the trophic shift, such variation is important because small errors in estimates of trophic shift can result in large errors in estimates of the contribution of sources to consumers or in estimates of trophic position.

Arguments for rejecting the sequential Bonferroni in ecological studies

Abstract: Interpretation of results that include multiple statistical tests has been an issue of great concern for some time in the ecological literature. The basic problem is that when multiple tests are undertaken, each at the same significance level ( ), the probability of achieving at least one significant result is greater than that significance level (Zaykin et al. 2002). Therefore, there is an increased probability of rejecting a null hypothesis when it would be inappropriate to do so. The typical solution to this problem has been lowering the values for the table (i.e. establishing a table-wide significance level) and therefore reducing the probability of a spurious result. Specifically, the most common procedure has been the application of the sequential Bonferroni adjustment (Holm 1979, Miller 1981, Rice 1989). Arguments in this essay address the problems of adjusting probability values for tables of multiple statistical tests, and more specifically argue for rejection of the sequential Bonferroni as a solution to this problem. Since the influential publication of Rice (1989), the sequential Bonferroni correction has become the primary method of addressing the problem of multiple statistical tests in ecological research. The sequential Bonferroni adjusts the table-wide p-value to keep it constant at 0.05, and subsequently reduces the probability of a spurious result. Although other methods exist for addressing tables of multiple statistical tests, the sequential Bonferroni has become the most commonly utilized process. However, this method has several flaws ranging from mathematical to logical to practical that argue for rejecting this method in ecological studies.

Long-distance migration: evolution and determinants

Abstract: Long distance migration has evolved in many organisms moving through different media and using various modes of locomotion and transport. Migration continues to evolve or become suppressed as shown by ongoing dynamic and rapid changes of migration patterns. This great evolutionary flexibility may seem surprising for such a complex attribute as migration. Even if migration in most cases has evolved basically as a strategy to maximise fitness in a seasonal environment, its occurrence and extent depend on a multitude of factors. We give a brief overview of different factors (e.g. physical, geographical, historical, ecological) likely to facilitate and/or constrain the evolution of long distance migration and discuss how they are likely to affect migration. The basic driving forces for migration are ecological and biogeographic factors like seasonality, spatiotemporal distributions of resources, habitats, predation and competition. The benefit of increased resource availability will be balanced by costs associated with the migratory process in terms of time (incl. losses of prior occupancy advantages), energy and mortality (incl. increased exposure to parasites). Furthermore, migration requires genetic instructions (allowing substantial room for learning in some of the traits) about timing, duration and distance of migration as well as about behavioural and physiological adaptations (fuelling, organ flexibility, locomotion, use of environmental transport etc) and control of orientation and navigation. To what degree these costs and requirements put constraints on migration often depends on body size according to different scaling relationships. From this expos it is clear that research on migration warrants a multitude of techniques and approaches for a complete as possible understanding of a very complex evolutionary syndrome. In addition, we also present examples of migratory distances in a variety of taxons. In recent years new techniques, especially satellite radio telemetry, provide new information of unprecedented accuracy about journeys of individual animals, allowing re-evaluation of migration, locomotion and navigation theories. (Less)

Mollusks as ecosystem engineers: the role of shell production in aquatic habitats

Abstract: Mollusk shells are abundant, persistent, ubiquitous physical structures in aquatic habitats. Using an ecosystem engineering perspective, we identify general roles of mollusk shell production in aquatic ecosystems. Shells are substrata for attachment of epibionts, provide refuges from predation, physical or physiological stress, and control transport of solutes and particles in the benthic environment. Changes in availability of these resources caused by shell production have important consequences for other organisms. Colonization of shelled habitat depends on individual shell traits and spatial arrangement of shells, which determine access of organisms to resources and the degree to which biotic or abiotic forces are modulated. Shell production will increase species richness at the landscape level if shells create resources that are not otherwise available and species are present that use these resources. Changes in the availability of resources caused by shells and the resulting effects on other organisms have both positive and negative feedbacks to these engineers. Positive feedbacks appear to be most frequently mediated by changes in resource availability, whereas negative feedbacks appear to be most frequently mediated by organisms. Given the diversity of species that depend upon resources controlled by shells and rapid changes in global shell production that are occurring due to human activities, we suggest that shell producers should not be neglected as a targets of conservation, restoration and habitat management.

Scavenging by vertebrates: behavioral, ecological, and evolutionary perspectives on an important energy transfer pathway in terrestrial ecosystems

Abstract: Carrion use by terrestrial vertebrates is much more prevalent than conventional theory implies, and, rather than a curiosity of animal behavior, is a key ecological process that must be accounted for. Human aversion to rotted substances and difficulties associated with identifying scavenged material in studies of food habits have contributed to the relative lack of information concerning scavenging behavior in vertebrates. Several lines of evidence, however, suggest that carrion resources are more extensively used by vertebrates than has been widely assumed: 1) a substantial number of animals die from causes other than predation and become available to scavengers, 2) a wide variety of vertebrate scavengers, rather than microbes or arthropods, consume most available carcasses, and 3) intense competition exists between vertebrate scavengers and decomposers, especially in warm climates. Although vultures are best adapted to use carrion, nearly all vertebrate predators are also scavengers to some extent. The costs and benefits associated with carrion use influences the evolution of scavenging behavior in vertebrates, resulting in a continuum of facultative scavengers that use carrion to varying degrees. The realized usage of carrion by a vertebrate species is influenced by the speed and efficiency with which it forages, its visual and olfactory abilities, and its capacity for detoxifying products of decomposition. A deeper understanding of carrion use by facultative scavengers will improve our knowledge of community and ecosystem processes, especially the flow of energy through food webs.

Effects of landscape context on herbivory and parasitism at different spatial scales

Abstract: Local community structure and interactions have been shown to depend partly on landscape context. In this paper we tested the hypothesis that the spatial scale experienced by an organism depends on its trophic level. We analyzed plant-herbivore and herbivore-parasitoid interactions in 15 agricultural landscapes differing in structural complexity using the rape pollen beetle (Meligethes aeneus), an important pest on oilseed rape (Brassica napus), and its parasitoids. In the very center of each landscape a patch of potted rape plants was placed in a grassy field margin strip for standardized measurement. Percent non-crop area of landscapes was negatively related to plant damage caused by herbivory and positively to the herbivores’ larval mortality resulting from parasitism. In a geographic scale analysis, we quantified the structure of the 15 landscapes for eight circular sectors ranging from 0.5 to 6 km diameter. Correlations between parasitism and non-crop areas as well as between herbivory and non-crop area were strongest at a scale of 1.5 km, thereby not supporting the view that higher trophic levels experience the world at a larger spatial scale. However, the predictive power of non-crop area changed only slightly for herbivory, but greatly with respect to parasitism as scales from 0.5 to 1.5 km and from 1.5 to 6 km diameter increased. Furthermore, the effect of non-crop area tended to be stronger in parasitism than herbivory suggesting a greater effect of changes in landscape context on parasitoids. This is in support of the general idea that higher trophic levels should be more susceptible to disturbance.

Methods for estimating long-distance dispersal

Abstract: Long-distance dispersal (LDD) includes events in which propagules arrive, but do not necessarily establish, at a site far removed from their origin. Although important in a variety of ecological contexts, the system-specific nature of LDD makes "far removed" difficult to quantify, partly, but not exclusively, because of inherent uncertainty typically involved with the highly stochastic LDD processes. We critically review the main methods employed in studies of dispersal, in order to facilitate the evaluation of their pertinence to specific aspects of LDD research. Using a novel classification framework we identify six main methodological groups: biogeographical: Eulerian and Lagrangian movement/redistributiona; short-term and long-term genetic analyses; and modeling. We briefly discuss the strengths and weaknesses of the most promising methods available for estimation of LDD. illustrating them with examples from current studies. The rarity of LDD events will continue to make collecting, analyzing, and interpreting the necessary data difficult, and a simple and comprehensive definition of LDD will remain elusive. However, considerable advances have been made in some methodological areas, such as miniaturization of tracking devices, elaboration of stable isotope and genetic analyses, and refinement of mechanistic models. Combinations of methods are increasingly used to provide improved insight on LDD from multiple angles. However, human activities substantially, increase the variety of long-distance transport avenues, making the estimation of LDD even more challenging.

Towards a functional resource-based concept for habitat: a butterfly biology viewpoint

Abstract: The habitat is the basic unit for developments in life history, population dynamics, landscape ecology and conservation of organisms. It is frequently treated as a particulate, invariant and homogeneous entity (a patch). Here we examine the implications of using this concept of habitat in butterfly biology. In doing so, we suggest the alternative approach of applying a functional resource-based concept of habitat. This recognises the fundamental requirements of organisms, consumables and utilities, the latter describing suitable environmental conditions as well as essential substrates. We argue that a resource-based concept is critical for butterfly conservation and call for the development of a resource database on butterfly biology.

Guilds or functional groups: does it matter?

Abstract: Although most researchers use the terms “guild” and “functional group” more or less synonymously, these two concepts bear different meanings. The guild concept refers primarily to the mechanisms of resource sharing by species in a competitive context whereas the functional groups concept is concerned with how a resource or any other ecological component is processed by different species to provide a specific ecosystem service or function. In many cases but not necessarily all, the two concepts are the two “faces” or “sides” of the same coin: the sharing by species of a similar resource is the guild facet (structural), while the ecosystem processes these species eventually perform through resource exploitation is the functional group facet. The two concepts differ in that competitive relationships within groups of species are not the focus of the functional group approach, exactly as processes or functions are not the focus of the guild approach. A group of species can be considered either as a guild or a functional group depending on the question addressed. Guild and functional group membership is independent of phylogenetic relationships but because species tend to share similar life history traits and adaptations through common evolutionary history, guild and functional group associates are often closely related. The concept of guild has had broader application in animal studies than in plant studies, whereas the reverse is true for the concept of functional group. Recent methodological advances to objectively partition species into guilds and functional groups, taking into consideration the most relevant characters or traits for delineating them, provide the means to construct an operational framework for making in situ and ex situ experiments that are urgently needed for a better understanding of the role of species in ecosystem functioning, especially in relation to global change concerns.

Using pattern-oriented modeling for revealing hidden information: a key for reconciling ecological theory and application

Abstract: We suggest that the conscious use of information that is “hidden” in distinct structures in nature itself and in data extracted from nature (=pattern) during the process of modeling (=pattern-oriented modeling) can substantially improve models in ecological application and conservation Observed patterns, such as time-series patterns and spatial patterns of presence/absence in habitat patches, contain a great deal of data on scales, site-history, parameters and processes Use of these data provides criteria for aggregating the biological information in the model, relates the model explicitly to the relevant scales of the system, facilitates the use of helpful techniques of indirect parameter estimation with independent data, and helps detect underlying ecological processes Additionally, pattern-oriented models produce comparative predictions that can be tested in the field We developed a step-by-step protocol for pattern-oriented modeling and illustrate the potential of this protocol by discussing three pattern-oriented population models: (1) a population viability analysis for brown bears (Ursusarctos) in northern Spain using time-series data on females with cubs of the year to adjust unknown model parameters; (2) a savanna model for detecting underlying ecological processes from spatial patterns of tree distribution; and (3) the incidence function model of metapopulation dynamics as an example of process integration and model generalization We conclude that using the pattern-oriented approach to its full potential will require a major paradigm shift in the strategies of modeling and data collection, and we argue that more emphasis must be placed on observing and documenting relevant patterns in addition to attempts to obtain direct estimates of model parameters

A review of long-distance movements by marine turtles, and the possible role of ocean currents

Abstract: Sea turtle movements often occur in open-sea unsheltered areas, and are therefore likely to be influenced by major oceanographic processes. Only recently has work started to examine the possible relationships of these movements with dynamic oceanic features, and consequently a clear picture of such interaction is only available in a few cases. Newborn sea turtles are thought to rely on oceanic currents to reach their pelagic nursery habitats. The actual extent and timing of these developmental migrations are known for only a few populations, but these movements probably last several years and range over thousands of km. Large juveniles that have been tracked during their pelagic stage were found to make long-distance movements, sometimes swimming against the prevailing currents. Older juveniles of most species leave the pelagic habitat to recruit to neritic developmental habitats. This is a very poorly documented phase of the sea turtle life-cycle, and the few available indications show that turtles may have to swim actively for enormous distances to counterbalance their previous drift with the current. The course and extent of adult postnesting migrations vary greatly among different turtle species, but two main patterns are evident. Some species, like green, hawksbill and loggerhead turtles, shuttle between the nesting beach and a specific feeding area used for the entire inter-reproductive period. In these cases, individuals swim, rather than drift, to complete their journeys, with possible advection due to currents sometimes helping them to quickly reach their target, but sometimes providing navigational challenges. Other species such as the olive ridley and the leatherback turtle, leave the coastal nesting areas to reach the pelagic environment where they forage, and perform wandering movements. Major oceanographic processes (such as main currents and eddies) have been recently shown to have a remarkable influence on leatherback movements, making it questionable whether these journeys are to be considered migrations or, rather, prolonged stays in vast feeding areas.

Current measures of nutrient resorption efficiency lead to a substantial underestimation of real resorption efficiency: facts and solutions.

Abstract: Nutrient resorption is an important process during leaf senescence, which helps plants to minimize nutrient losses. To quantify nutrient resorption, the parameter resorption efficiency is commonly used. This parameter describes the percentage of the nutrient pool withdrawn before leaf abscission. The nutrient pool is generally expressed on the basis of leaf mass or leaf area, assuming that these bases do not change during senescence. In this paper we firstly present a mathematical formula describing the effect of change in measurement basis on the difference between the real resorption efficiency (RRE) value and the measured resorption efficiency (MRE). This formula shows that even moderate senescence-related changes in a measurement basis can lead to considerable underestimation of RRE. Secondly, to estimate the general change in measurement basis we quantified leaf mass loss and leaf shrinkage during senescence from literature data. These data shows that mass loss percentages can be as high as 40%, and leaf shrinkage can be up to 20%. This level of change in basis seriously compromises the MRE when not corrected for. Using our formula and the reported average literature values of changes in leaf mass (21%) and leaf shrinkage (11%,) during senescence, we calculated that the average RRE for nitrogen and phosphorous of terrestrial plants is 6% (leaf area) to 10% (leaf mass) higher than the 50%, respectively 52% as reported by Aerts (1996). This implies that nutrient resorption from senescing leaves is even more important for nutrient retention in terrestrial plants than thought so far. We advocate that preselecting leaves and monitoring the measurement basis throughout the duration of the experiment should minimize the difference between MRE and RRE.

Can noise induce chaos

Abstract: An important component of the mathematical definition of chaos is sensitivity to initial conditions. Sensitivity to initial conditions is usually measured in a deterministic model by the dominant Lyapunov exponent (LE), with chaos indicated by a positive LE. The sensitivity measure has been extended to stochastic models; however, it is possible for the stochastic Lyapunov exponent (SLE) to be positive when the LE of the underlying deterministic model is negative, and vice versa. This occurs because the LE is a long-term average over the deterministic attractor while the SLE is the long-term average over the stationary probability distribution. The property of sensitivity to initial conditions, uniquely associated with chaotic dynamics in deterministic systems, is widespread in stochastic systems because of time spent near repelling invariant sets (such as unstable equilibria and unstable cycles). Such sensitivity is due to a mechanism fundamentally different from deterministic chaos. Positive SLE's should therefore not be viewed as a hallmark of chaos. We develop examples of ecological population models in which contradictory LE and SLE values lead to confusion about whether or not the population fluctuations are primarily the result of chaotic dynamics. We suggest that “chaos” should retain its deterministic definition in light of the origins and spirit of the topic in ecology. While a stochastic system cannot then strictly be chaotic, chaotic dynamics can be revealed in stochastic systems through the strong influence of underlying deterministic chaotic invariant sets.

Pay-offs and penalties of competing migratory schedules

Abstract: We relate variation in the timing of arrival by migrating birds breeding at northerly latitudes to individual differences in the prior accumulation of energy stores. Balancing starvation risks early in the season against the almost universal declining trend in reproductive prospects with advancing date is seen as an individual decision with fitness consequences. We review three studies implicating events at the staging sites or in winter in setting the individual migratory schedule. Climate change influences the timetable of a pied flycatcher (Ficedula hypoleuca) population breeding in The Netherlands and wintering in West Africa, followed since 1960. Mean air temperature in the period mid April-mid May (arrival and laying) increased and laying date advanced by 10 days. Still, in recent years most birds did not lay early enough to maximise fitness (determined by recruitment and parental survival) whereas many parents achieved this goal in 1980-1985. As the flycatchers have not started to arrive earlier, some ecological constraint further upstream is postulated (possibly the hurdle of the crossing of Sahara and Mediterranean). The ability to follow individual migrants provides a second avenue to assess the fitness implications of migratory schedules. Thus, brightly coloured male bar-tailed godwits (Limosa lapponica) captured in the Dutch Wadden Sea (the intermediate staging site linking a West African wintering area with breeding sites in arctic Russia) and traced with miniature radio-transmitters did not depart early. The 'best' males (with bright breeding plumage) were picked up by the listening stations in Sweden 650 km further along the migratory route ten days later than the paler individuals. If early arrival confers the competitive advantage of prior occupancy but increases mortality, the 'best' males may be able to afford arriving later and thus avoid some of the survival costs. Return rate of the 'bright' males to the staging site in later seasons was indeed higher than for the 'pale', early males. Intensive observation of pink-footed geese (Anser brachyrhynchus) fitted with coded neck-collars substantiate the tight relationship between energy stores (fat) accumulated up to final departure from the final staging site (Vesteralen, N. Norway) en route to the nesting grounds (Spitsbergen) and subsequent success. The breeding outcome of individual parents (accompanied by juveniles or not) could be related to observations of body condition before departure (visual 'abdominal profile index'). Recently, perceived conflicts with agriculture have resulted in widespread harassment by humans. The geese have: drastically shortened their stay on the Vesteralen, fail to achieve the body condition usual a decade ago and reproductive output has fallen. Although the geese are currently pioneering new staging sites, an adequate alternative has not materialised, underlining the critical role of the final take-off site.

The role of habitat selection behavior in population dynamics: source -sink systems and ecological traps

Abstract: Ecological traps, poor-quality habitat that nonetheless attract individuals, have been observed in both natural and human-altered settings. Until recently, ecological traps were considered a kind of source--sink system, but source-sink theory does not model maladaptive habitat choice, and therefore cannot accurately represent ecological traps or predict their population-level consequences. Although recent models of ecological traps addressed this problem, they used patch-based models containing only two habitats that were very different from one another, but were internally homogeneous. These sorts of patch models may not apply to many real populations, and using them for populations in landscapes with mosaic or gradient habitat structures may be misleading. I developed models that treat source -sink dynamics and ecological traps as special cases of a single process, in which the attractiveness and quality of the habitat are separate variables that can be either positively or negatively related, and in which habitat quality varies continuously throughout the landscape. As expected, sinks are less detrimental to populations than ecological traps, in which preferential use of poor habitat elevates extinction risk. Furthermore, ecological traps may be undetected, and may even appear to be sources, when population sizes are large, but may still prevent recovery in spite of the availability of high-quality habitat when populations drop below threshold levels. Conservation biologists do not routinely consider the possibility that apparent sinks are actually traps, but since traps should be associated with the rapidly changing and novel habitat characteristics primarily produced by human activities, ecological traps should be considered an important and potentially widespread conservation concern.

The distance dependence prediction of the Janzen-Connell hypothesis: a meta-analysis

Abstract: The Janzen-Connell hypothesis explains the maintenance of tropical diversity through the interacting effects of parent-centered dispersal patterns and distance- and density-dependent propagule survival. These effects were thought to support regular spacing of species within tropical forest, enhancing diversity. One of the predictions of the hypothesis is that seed and seedling survival should improve with increased parental distance. Although there are many independent tests of this hypothesis for individual species, there are few synthetic studies that have brought these data together to test its validity across species. This paper reports the results of a meta-analysis of the effect of distance on enhancing propagule survival, employing an odds-ratio effect size metric. We found no general support for the distance-dependent prediction of the hypothesis, and conclude that further testing to explore this hypothesis as a diversity-maintaining mechanism is unnecessary. However, we did find that distance from parent slightly reduces survivorship in the temperate zone, as contrasted with the tropics, and we saw stronger evidence in support of the hypothesis for seedlings than for seeds. The phenomenon of enhanced propagule survival with distance from the parent may be important for the population biology of particular species, but it is not a general phenomenon across communities, life history stages or life forms.

Interactions between above- and belowground insect herbivores as mediated by the plant defense system

Abstract: Plants are frequently attacked by both above- and belowground arthropod herbivores. Nevertheless, studies rarely consider root and shoot herbivory in conjunction. Here we provide evidence that the root-feeding insect Agriotes lineatus reduces the performance of the foliage feeding insect Spodoptera exigua on cotton plants. In a bioassay, S. exigua larvae were allowed to feed on either undamaged plants, or on plants that had previously been exposed to root herbivory, foliar herbivory, or a combination of both. Previous root herbivory reduced the relative growth rates as well as the food consumption of S. exigua by more than 50% in comparison to larvae feeding on the undamaged controls. We found no effects in the opposite direction, as aboveground herbivory by S. exigua did not affect the relative growth rates of root-feeding A. lineatus. Remarkably, neither did the treatment with foliar herbivory affect the food consumption and relative growth rate of S. exigua in the bioassay. However, this treatment did result in a significant change in the distribution of S. exigua feeding. Plants that had been pre-exposed to foliar herbivory suffered significantly less damage on their young terminal leaves. While plant growth and foliar nitrogen levels were not affected by any of the treatments, we did find significant differences between treatments with respect to the level and distribution of plant defensive chemicals (terpenoids). Exposure to root herbivores resulted in an increase in terpenoid levels in both roots as well as in mature and immature foliage. Foliar damage, on the other hand, resulted in high terpenoid levels in young, terminal leaves only. Our results show that root-feeding herbivores may change the level and distribution of plant defenses aboveground. Our data suggest that the reported interactions between below- and aboveground insect herbivores are mediated by induced changes in plant secondary chemistry.

Global warming and the export of dissolved organic carbon from boreal peatlands

Abstract: Peatlands occupy approximately 15% of boreal and sub-arctic regions, contain approximately one third of the world's soil carbon pool, and supply most of the dissolved organic carbon (DOC) entering boreal lakes and rivers and the Arctic Ocean. The high latitudes occupied by these peatlands are expected to see the greatest amount of climatic warming in the next several decades. In addition to increasing temperatures, climatic change could also affect the position of the water-table level and discharge from these peatlands. Changes in temperature, water tables, and discharge could affect delivery of DOC to downstream ecosystems where it exerts significant control over productivity, biogeochemical cycles, and attenuation of visible and UV radiation. We experimentally warmed and controlled water tables while measuring discharge in a factorial experiment in large mesocosms containing peat monoliths and intact plant communities from a bog and fen to determine the effects of climate change on DOC budgets. We show that the DOC budget is controlled largely by changes in discharge rather than by any effect of warming or position of the water-table level on DOC concentrations. Furthermore, we identify a critical discharge rate in bogs and fens for which the DOC budget switches from net export to net retention. We also demonstrate an exponential increase in trace gas CO 2 -C and CH 4 -C emissions coincident with increased retention of dissolved organic carbon from boreal peatlands.

Growing tall vs growing wide: tree architecture and allometry of Acacia karroo in forest, savanna, and arid environments

Abstract: In order to investigate how environmental factors other than light availability affect tree architecture, differences in branching architecture and allometry were analysed in populations of Acacia karroo Hein. from three different environments in South Africa: forests, savannas and arid-shrublands. Factors such as fire and herbivory have a large effect on tree life history in certain environments and are likely to have selected for trees that have different architectures from those of forest trees, whose major limitation is light assimilation. Significant differences were found in stem architecture and branching architecture between trees in each environment. Compared with forest trees, trees in savannas had an elongated growth form with small canopy and leaf areas, and tall, thin, unbranched trunks. Trees in arid areas showed opposite trends with wider canopies, and increased lateral branching. Savanna trees had significantly smaller spines than trees in other environments, and both forest and savanna trees showed delayed reproduction. These differences are probably related to a trade-off between an architecture geared towards rapid height-gain and one promoting lateral spread, and can be explained with reference to the different selective pressures in each environment. In forests, vertical and horizontal growth are both important. However, in savannas there is a great pressure for rapid vertical growth to escape fires, while in arid areas a defensive, lateral growth form is selected for. Savanna trees and arid karoo trees have evolved architectures that are more extreme vertically and laterally than the range of architectures displayed in a forest community.

Response of plant-pollinator communities to fire: changes in diversity, abundance and floral reward structure

Abstract: Globally, plant-pollinator communities are subject to a diverse array of perturbations and in many temperate and semi-arid systems fire is a dominant structuring force. We present a novel and highly integrated approach, which quantifies, in parallel, the response to fire of pollinator communities, floral communities and floral reward structure. Mt Carmel, Israel is a recognised bee-flower biodiversity hotspot, and using a chronosequence of habitats with differing post-fire ages, we follow the changes in plant-pollinator community organisation from immediately following a burn until full regeneration of vegetation. Initially, fire has a catastrophic effect on these communities, however, recovery is rapid with a peak in diversity of both flowers and bees in the first 2 years post-fire, followed by a steady decline over the next 50 years. The regeneration of floral communities is closely matched by that of their principal pollinators. At the community level we quantify, per unit area of habitat, key parameters of nectar and pollen forage known to be of importance in structuring pollinator communities. Nectar Volume, nectar water content, nectar concentration and the diversity of nectar foraging niches are all greatest immediately following fire with a steady decrease as regeneration proceeds. Temporal changes in energy availability for nectar, pollen, total energy (nectar + pollen) and relative importance of pollen to nectar energy show a similar general decline with site age, however, the pattern is less clear owing to the highly patchy distribution of floral resources. Changes in floral reward structure reflect the general shift from annuals (generally low-reward open access flowers) to perennials (mostly high-reward and restricted access flowers) as post-fire regeneration ensues. The impact of fire on floral communities and their associated rewards have clear implications for pollinator community structure and we discuss this and the role of other disturbance factors on these systems.

Mechanisms underlying shoal composition in the Trinidadian guppy, Poecilia reticulata

Abstract: Free-ranging groups are frequently assorted by phenotypic characters. However, very little is known about the underlying processes that determine this structuring. In this study, we investigate the mechanisms underlying the phenotypic composition of shoals of guppies (Poecilia reticulata) in a high-predation stream in Trinidad's Northern Mountain Range. We collected 57 entire wild shoals, which were strongly assorted by body length. Shoal encounters staged within an experimental arena showed shoal fission (but not fusion) events to be an important mechanism in generating phenotypic assortment. In the wild, fission and fusion between guppy shoals occurred extremely frequently and thus are unlikely to constrain the opportunities for shoal assortment. However, fission and fusion processes occur under the restrictions imposed by the distribution of individuals within the environment. We observed size specific segregation within the habitat in three dimensions, providing a passive mechanism that contributes to the maintenance of the observed homogeneity of group composition. Furthermore sex differences were found in social behaviour. Individual male guppies switched between shoals more frequently than females and left a shoal more often than females. We argue that shoal composition is determined by habitat segregation on a medium spatial scale and by fission/fusion processes on a small spatial scale (with sex-specific shoal dynamics adding a additional layer of complexity).

Morphology dependent foraging efficiency in perch: a trade-off for ecological specialization?

Abstract: The work in this thesis deals with the ecology and evolution of adaptive individual variation. Ecologists have long used niche theory to describe the ecology of a species as a whole, treating conspecific individuals as ecological equivalent. During recent years, research about individual variation in diet and morphology has gained interest in adaptive radiations and ecological speciation. Such variation among individual niche use may have important conservation implications as well as ecological and evolutionary implications. However, up to date we know very little about the extension of this phenomenon in natural populations and the mechanisms behind it.The results in this thesis show that the extension of individual diet specialization is widely spread throughout the animal kingdom. The variation in diet is mainly correlated to morphological variation but not always. Furthermore, this variation in diet and morphology among individuals could be both genetically determined and environmentally induced and it mainly comes from trade-offs in foraging efficiency between different prey types. The results from a number of studies of perch also show that individual perch differ in morphology and diet depending on habitat, where littoral perch has a deeper body compared to pelagic perch. This difference in morphology corresponds to functional expectations and is related to foraging efficiency trade-offs between foraging in the littoral and pelagic zone of a lake. The variation in morphology in perch is mainly due to phenotypic plasticity but there are also small genetic differences between the littoral and pelagic perch. Two separate studies show that both predation and competition may be important mechanism for the variation in morphology and diet in perch.In conclusion, the results in this thesis show that individual variation in diet and habitat choice is a common phenomenon with lots of ecological and evolutionary implications. However, there are many mechanisms involved in this phenomenon on which we are just about to start learning more about, and only further research in this area will give us the full insight.

The response of a three trophic level soil food web to the identity and diversity of plant species and functional groups

Abstract: Despite considerable recent interest in how biodiversity may influence ecosystem properties, the issue of how plant diversity and composition may affect multiple trophic levels in soil food webs remains essentially unexplored. We conducted a glasshouse experiment in which three plant species of each of three functional groups (grasses, N-fixing legumes and forbs) were grown in monoculture and in mixtures of three species (with the three species being in the same or different functional groups) and all nine species. Plant species identity had important effects on the biomasses or population densities of belowground primary consumers (microbial biomass, herbivorous nematodes) and two groups of secondary consumers (microbe-feeding nematodes and enchytraeids); the third consumer trophic level (predatory nematodes) was marginally not significantly affected at P=0.05. Plant species also influenced the relative importance of the bacterial-based and fungal-based energy channels for both the primary and secondary consumer trophic levels. Within-group diversity of only the soil microflora and herbivorous nematodes (both representing the basal consumer trophic level) were affected by plant species identity. However, community composition within all trophic groupings considered (herbivorous nematodes, microbes, microbe-feeding nematodes, predatory nematodes) was strongly influenced by what plant species were present. Despite the strong responses of the soil biota to plant species identity, there were few effects of plant species or functional group richness on any of the belowground response variables measured. Further, net primary productivity (NPP) was unaffected by plant diversity. Since some belowground response variables were correlated with NPP across treatments, it is suggested that belowground responses to plant diversity might become more apparent in situations when NPP itself responds to plant diversity. Our results point to plant species identity as having important multitrophic effects on soil food webs, both at the whole trophic group and within-group levels of resolution, and suggest that differences in plant traits across species may be important in driving the decomposer subsystem.

Plant species diversity, plant biomass and responses of the soil community on abandoned land across Europe: idiosyncracy or above-belowground time lags

Abstract: We examined the relationship between plant species diversity, productivity and the development of the soil community during early secondary succession on former arable land across Europe. We tested the hypothesis that increasing the initial plant species diversity enhances the biomass production and consequently stimulates soil microbial biomass and abundance of soil invertebrates. We performed five identical field experiments on abandoned arable land in five European countries (CZ, NL, SE, SP and UK) which allowed us to test our hypothesis in a range of climate, soil and other environmental factors that varied between the experimental sites. The initial plant diversity was altered by sowing seed mixtures of mid-successional grassland species with two or five grass species, one or five legumes and one or five forbs. The results of low and high sown diversity treatments were compared with plots that were naturally colonized by species present in the seed bank. In three out of the five field sites, there was no correlation between plant species number and plant biomass production, one site had a positive and the other a negative relation. Treatments with a high diversity seed mixture had a higher biomass than the naturally colonized plots. However, there was no significant difference between high and low sown diversity plots at four out of five sites. The three-year study did not give any evidence of a general bottom-up effect from increased plant biomass on biomass of bacteria, saprophytic fungi or abundance of microarthropods. The biomass of arbuscular mycorrhizal was negatively related to plant biomass. The abundance of nematodes increased after abandonment and was related to plant biomass at four sites. Our results support the hypothesis that plant species diversity may have idiosyncratic effects on soil communities, even though studies on a longer term could reveal time lags in the response to changes in composition and biomass production of plant communities.

Integrating methods that investigate how complementarity influences ecosystem functioning

Abstract: Separating the mechanisms that influence ecosystem functioning has been a goal of recent high profile experiments. Integrating the various experimental and analytical methods that attempt this goal across terrestrial and aquatic ecosystems, as well as careful definition of ‘complementarity’, produces novel insights and valuable lessons about new directions for research. (1) Experimental designs differ in temporal scale and whether standing stock or another ecosystem process was the response variable. (2) Mathematically identical variables in different designs have contrasting ecological interpretations. For example, different sets of ecological processes can contribute to different variables in different experimental designs. (3) The frequency of transgressive overyielding of standing stock (e.g. total above ground biomass) in polycultures implies little about the prevalence of transgressive overyielding in other ecosystem processes. (4) Measuring the contribution to ecosystem functioning of individual species, rather than just total ecosystem functioning of a polyculture, is not essential for estimating effects of complementarity. (5) Further research will profit from distinguishing standing stock from all other ecosystem functions. (6) None of the analytic methods can distinguish the effects of individual processes or mechanisms such as resource use differentiation, facilitation, or allelopathy, for which additional experimental treatments are required.

Spatial occurrence and colonisations in patch‐tracking metapopulations: local conditions versus dispersal

Abstract: We studied the relative importance of local variables and dispersal for the occurrence and colonisation of the epiphytic bryophytes Orthotricum speciosum (spore dispersed), and O. obtusifolium (spore and asexual gemmae) on aspen trees (‘patches’) in two forest landscapes (one old-growth and one fragmented) using multiple logistic regression. The relative importance of dispersal was quantified as the reduction of residual deviance for a connectivity variable. In modelling dispersal, we assumed that trees with low local abundance were recent colonisations, and that trees with high local abundance were diaspore sources for colonisation. The occurrence of O. speciosum in the fragmented landscape was most affected by shading, but also by connectivity, aspen diameter and vitality. In the old-growth landscape, connectivity was the single most important variable for recent colonisations, but its effect was lower than the sum of the effects of all local environmental variables. The occurrence of O. obtusifolium in the fragmented landscape was related to similar variables but the relative importance of these variables was different, and connectivity did not affect the probability of a recent colonisation in this species. We describe the epiphyte-tree system in the patch-tracking metapopulation model. In this model colonisations are distance dependent, but in contrast to the classical metapopulation model local extinctions are caused by deterministic patch destruction – once the epiphyte has colonised the tree it remains until the tree dies.

Interferences between Sphagnum and vascular plants: effects on plant community structure and peat formation

Abstract: The interference between vascular plants and peat mosses with respect to nitrogen and phosphorus was studied in a fertilization experiment and with respect to competition for light in a removal experiment in poor fens with either soligenous or topogenous hydrology using Narthecium ossifragum (L.) Huds. and three species of Sphagnum sect. Sphagnum as targets. Adding fertilizer either on the moss surface or below it confirmed the hypotheses of an asymmetric competition for nutrients, viz. that the Sphagnum mosses relied on the atmospheric supply while Narthecium depended on mineralization in the peat. The results of the removal experiments and the negatively correlated growth of Narthecium and Sphagnum mosses demonstrated a symmetric competition for light. The intensity of the competition for light increased as the availability of N and P increased. The nutrient resources in the total biomass decreased with decreasing standing crop of Narthecium. Only with a considerable amount of mineral nutrients in the biomass has Narthecium the capacity to grow ahead of Sphagnum, because the asymmetric competition for N and P gives Sphagnum the capacity to reduce the performance of vascular plants. The mosses are more efficient in their use of nutrients and produce a decay-resistant litter inducing low mineralization and increasing the peat accumulation rate, and that withdraws N and P from the rhizosphere. The Sphagnum mosses thus act as ecological engineers structuring the plant community and determining the carbon balance of the system. The development of ombrotrophic conditions through peat accumulation seems less probable on soligenous than on topogenous mires owing to the higher mineralization rate there supporting the growth of the vascular plants. Correspondingly, disturbances of the Sphagnum cover, such as through airborne pollutants, increase the productivity of the vascular plants and decrease the capacity for carbon accumulation.

Explaining productivity‐diversity relationships in plants

Abstract: Relationships between productivity and diversity in plant communities have been widely documented. Unimodal productivity-diversity relationships are most common along natural productivity gradients, and fertilization generally reduces diversity. Five distinct hypotheses invoke changes in competition to explain why diversity should decline from intermediate to high productivity. Because experiments measuring the effects of competition on diversity are rare, four of the five hypotheses have not been directly tested, but each hypothesis makes unique predictions that allow for indirect tests. The indirect evidence is often conflicting, and while none of the hypotheses can be rejected, only the dynamic equilibrium hypothesis is consistently supported. A new hypothesis, however, is supported by indirect evidence and may help to explain the variation in the shape of productivity-diversity relationships, as well as the most common patterns. Diversity may be high in environments that promote size symmetric competition, where soil resources limit growth and are homogeneously distributed within the soil volume explored by individual plants. Conversely, diversity may be low in environments that promote size asymmetric competition, where light is limiting, or where soil resources are limiting and are patchily distributed within rooting zones.

Travel schedules to the high arctic: barnacle geese trade‐off the timing of migration with accumulation of fat deposits

Abstract: On their way from the wintering area to the breeding grounds in Spitsbergen, barnacle geese Branta leucopsis stage on islands off the coast of Norway. The aim of this study was to describe when the geese migrate in relation to the body stores deposited and explore questions related to the concept of optimal migration schedules and on the possible mechanisms involved. We estimated fat stores by repeated assessments of the abdominal profile index of individually marked females throughout staging. Reproductive success was derived from observations of the same individuals later in the annual cycle. Females arriving late, or with low fat stores at arrival, achieved higher fat deposition rates, probably by spending more time foraging. But they were unable to match final fat scores of birds that arrived earlier or with larger fat stores. Reproductive success was correlated with the timing of migration and individuals departing at intermediate dates achieved highest success. The exact date of peak reproductive success depended on the size of fat stores accumulated, such that low-quality birds (depositing less fat) benefited most from an early departure to the breeding grounds. Observations in the breeding colonies showed that these birds did not initiate a nest earlier but they spent a longer time in Spitsbergen before settling. The length of stay in Norway was close to the prediction derived from an optimisation model relating spring events to eventual breeding success. Poorest performing birds stayed longer than expected, perhaps depositing more fat to avoid the risk of starvation. Two possible mechanisms of the timing of migration were contrasted and it seemed that the geese departed for migration as soon as they were unable to accumulate any more fat stores.

Regional and local patterns in plant species richness with respect to resource availability

Abstract: The hump-shaped relationship between plant species richness and productivity is a well-established and important paradigm. While plot-based species richness patterns on local scales have received much attention, little is currently known about species-based patterns on a regional scale. Using Ellenberg's indicator values for 1802 plant species in central Europe, we assess the patterns in regional species richness with respect to light, water, and mineral nutrient availability – three variables that strongly influence productivity. The results of this analysis are compared to those of published studies on smaller scales leading to the following conclusions: 1 On a regional scale in central Europe there is a hump-shaped relationship between soil nutrient supply and plant species richness within a given biome. 2 The peak in species richness for grasslands and wetlands occurs on nutrient-poor soils, while the peak for forests is on nutrient-rich soils. 3 Gradients in plant productivity controlled by different variables (i.e. water, nutrients, or disturbance) have dissimilar effects on plant species richness.