Flexible defense strategies: competition modifies investment in behavioral vs. morphological defenses

Competition is predicted to affect the expression of inducible defenses, but because costs of behavioral and morphological antipredator defenses differ along resource gradients, its effects on defenses may depend on the traits considered. We tested the predictions from different defense models in tadpoles of the common frog Rana temporaria, which exhibit both types of defenses. In an outdoor experiment, we exposed the tadpoles to nonlethal predators (Aeshna dragonfly larvae) and to a gradient of intraspecific competition. Morphological responses did not follow any of the expected patterns, since investment in defense was not affected by resource level. Instead, tail depth decreased in the absence of predators. Behavioral defenses followed a state-dependent model. Overall, the defense strategy of the tadpoles revealed a shift from morphological and behavioral defenses at low tadpole density to morphological defense only at high density. This difference probably reflects the different efficiency of the defenses. Hiding is an effective means of defense, but it is unsustainable when resources are scarce. Morphological responses become more important with increasing density to compensate for the increase in behavioral risk-taking. Our results indicate that competition can strongly affect reaction norms of inducible defenses and highlight the importance of integrating ecological parameters that affect the cost-benefit balance of phenotypic plasticity.     

Functional connectivity from a reef fish perspective: behavioral tactics for moving in a fragmented landscape

Functional connectivity, the degree to which the landscape facilitates or impedes movement, depends on how animals perceive costs and benefits associated with habitat features and integrate them into a movement path. There have been few studies on functional connectivity in marine organisms, despite its importance for the effectiveness of Marine Protected Areas. In this study, we asked how open sand and conspecific distribution affected functional connectivity of longfin damselfish (Stegastes diencaeus) on fringing reefs in Barbados. We translocated 102 individuals to sites varying in sand gap width and in configuration: Continuous (solid reef between release site and territory); Detour (sand along the direct path between release site and territory, but an alternative, continuous solid U-shaped reef path); and Patch (sand between release site and territory, but an alternative stepping stone path). We visually tracked and mapped every homing path. We found no evidence of a barrier to movement in the Continuous configuration, but sand was a partial barrier in Detour and Patch configurations. The probability of crossing the sand gap dropped below 50% when its width was > 1.85 m in Detour and > 3.90 m in Patch configuration. Damselfish avoiding large gaps took detours that approximated the route maximizing travel over reef, but they crossed more short sand gaps and fewer conspecific territories, suggesting avoidance of agonistic interactions. This study quantifies for the first time the size and steepness of a barrier to movement in a marine organism, and it provides evidence for effects of both landscape configuration and conspecific distribution on functional connectivity.     

Grassland invader responses to realistic changes in native species richness.

The importance of species richness for repelling exotic plant invasions varies from ecosystem to ecosystem. Thus, in order to prioritize conservation objectives, it is critical to identify those ecosystems where decreasing richness will most greatly magnify invasion risks. Our goal was to determine if invasion risks greatly increase in response to common reductions in grassland species richness. We imposed treatments that mimic management-induced reductions in grassland species richness (i.e., removal of shallow- and/or deep-rooted forbs and/or grasses and/or cryptogam layers). Then we introduced and monitored the performance of a notorious invasive species (i.e., Centaurea maculosa). We found that, on a per-gram-of-biomass basis, each resident plant group similarly suppressed invader growth. Hence, with respect to preventing C. maculosa invasions, maintaining overall productivity is probably more important than maintaining the productivity of particular plant groups or species. But at the sites we studied, all plant groups may be needed to maintain overall productivity because removing forbs decreased overall productivity in two of three years. Alternatively, removing forbs increased productivity in another year, and this led us to posit that removing forbs may inflate the temporal productivity variance as opposed to greatly affecting time-averaged productivity. In either case, overall productivity responses to single plant group removals were inconsistent and fairly modest, and only when all plant groups were removed did C. maculosa growth increase substantially over a no-removal treatment. As such, it seems that intense disturbances (e.g., prolonged drought, overgrazing) that deplete multiple plant groups may often be a prerequisite for C. maculosa invasion.     

Independent effects of fragmentation on forest songbirds: an organism-based approach.

The degree to which spatial patterns influence the dynamics and distribution of populations is a central question in ecology. This question is even more pressing in the context of rapid habitat loss and fragmentation, which threaten global biodiversity. However, the relative influence of habitat loss and landscape fragmentation, the spatial patterning of remaining habitat, remains unclear. If landscape pattern affects population size, managers may be able to design landscapes that mitigate habitat loss. We present the results of a mensurative experiment designed to test four habitat loss vs. fragmentation hypotheses. Unlike previous studies, we measured landscape structure using quantitative, spatially explicit habitat distribution models previously developed for two species: Blackburnian Warbler (Dendroica fusca) and Ovenbird (Seiurus aurocapilla). We used a stratified sampling design that reduced the confounding of habitat amount and fragmentation variables. Occurrence and reoccurrence of both species were strongly influenced by characteristics at scales greater than the individual territory, indicating little support for the random-sample hypothesis. However, the type and spatial extent of landscape influence differed. Both occurrence and reoccurrence of Blackburnian Warblers were influenced by the amount of poor-quality matrix at 300- and 2000-m spatial extents. The occurrence and reoccurrence of Ovenbirds depended on a landscape pattern variable, patch size, but only in cases when patches were isolated. These results support the hypothesis that landscape pattern is important for some species only when the amount of suitable habitat is low. Although theoretical models have predicted such an interaction between landscape fragmentation and composition, to our knowledge this is the first study to report empirical evidence of such nonlinear fragmentation effects. Defining landscapes quantitatively from an organism-based perspective may increase power to detect fragmentation effects, particularly in forest mosaics where boundaries between patches and matrix are ambiguous. Our results indicate that manipulating landscape pattern may reduce negative impacts of habitat loss for Ovenbird, but not Blackburnian Warbler. We emphasize that most variance in the occurrence of both species was explained by local scale or landscape composition variables rather than variables reflecting landscape pattern.     

Time allocation strategies in insect parasitoids: from ultimate predictions to proximate behavioral mechanisms

As most parasitoids are time limited, they usually die before they have laid all their eggs. In such cases, optimal foraging theory predicts that female parasitoids will adopt behavioral reproductive strategies enabling them to maximize progeny production per unit of time. One key situation in which parasitoid females must optimize their time budget is related to the fact that most of their hosts are distributed in discrete patches in the environment. In this review, I first present the results of basic theoretical models predicting female wasp search duration on a patch of hosts. I then compile and analyze all studies investigating the effect of different factors on parasitoid patch time allocation and patch-leaving decision rules. Different patch-leaving mechanisms that were proposed to explain the results obtained are discussed, along with statistical methods that should be used to estimate them from experimental data. Finally, ideas for future research are presented.     

Traplining by purple-throated carib hummingbirds: behavioral responses to competition and nectar availability

We examined the effects of nectar availability and competition on foraging preferences and revisit intervals of traplining female purple-throated caribs hummingbirds (Eulampis jugularis) to Heliconia patches shared by two individuals or visited solely by one individual. Birds at both shared and solitary patches preferred multiflowered to single-flowered inflorescences, but the magnitude of this preference depended on food availability and competition. During a year of low flower availability, females visited multiflowered inflorescences more frequently than single-flowered inflorescences only when nectar availability was experimentally enhanced; similarly, females at shared patches exhibited a significant preference for multiflowered inflorescences only after experimental increases in nectar availability. Experimental manipulations of nectar availability also had different effects on revisit intervals of birds at shared vs solitary patches. Birds at shared patches responded to patch-wide increases in nectar rewards by increasing the duration of their visit intervals, whereas birds at solitary patches did not. In contrast, birds at solitary patches responded to abrupt losses of nectar at flowers (simulating competition) by decreasing the duration of their visit intervals, whereas a bird at a shared patch did not alter its return interval. The contrasting results between shared vs solitary patches suggest that future studies of traplining behavior should incorporate levels of competition into their design.     

Influences of divergent behavioral strategies upon risk allocation in juvenile flatfishes

Animals balance feeding and anti-predator behaviors at various temporal scales. When risk is infrequent or brief, prey can postpone feeding in the short term and temporally allocate feeding behavior to less risky periods. If risk is frequent or lengthy, however, prey must eventually resume feeding to avoid fitness consequences. Species may exhibit different behavioral strategies, depending on the fitness tradeoffs that exist in their environment or across their life histories. North Pacific flatfishes that share juvenile rearing habitat exhibit a variety of responses to predation risk, but their response to risk frequency has not been examined. We observed the feeding and anti-predator behaviors of young-of-the-year English sole (Parophrys vetulus), northern rock sole (Lepidopsetta polyxystra), and Pacific halibut (Hippoglossus stenolepis)—three species that exhibit divergent anti-predator strategies—following exposure to three levels of predation risk: no risk, infrequent (two exposures/day), and frequent (five exposures/day). The English sole responded to the frequent risk treatment with higher feeding rates than during infrequent risk, following a pattern of behavioral response that is predicted by the risk allocation hypothesis; rock sole and halibut did not follow the predicted pattern, but this may be due to the limited range of treatments. Our observations of unique anti-predator strategies, along with differences in foraging and species-specific ecologies, suggest divergent trajectories of risk allocation for the three species.     

Water management strategies against toxic Microcystis blooms in the Dutch delta.

To prevent flooding of the Dutch delta, former estuaries have been impounded by the building of dams and sluices. Some of these water bodies, however, experience major ecological problems. One of the problem areas is the former Volkerak estuary that was turned into a freshwater lake in 1987. From the early 1990s onward, toxic Microcystis blooms dominate the phytoplankton of the lake every summer. Two management strategies have been suggested to suppress these harmful algal blooms: flushing the lake with fresh water or reintroducing saline water into the lake. This study aims at an advance assessment of these strategies through the development of a mechanistic model of the population dynamics of Microcystis. To calibrate the model, we monitored the benthic and pelagic Microcystis populations in the lake during two years. Field samples of Microcystis were incubated in the laboratory to estimate growth and mortality rates as functions of light, temperature, and salinity. Recruitment and sedimentation rates were measured in the lake, using traps, to quantify benthic-pelagic coupling of the Microcystis populations. The model predicts that flushing with fresh water will suppress Microcystis blooms when the current flushing rate is sufficiently increased. Furthermore, the inlet of saline water will suppress Microcystis blooms for salinities exceeding 14 g/L. Both management options are technically feasible. Our study illustrates that quantitative ecological knowledge can be a helpful tool guiding large-scale water management.     

景观规划与管理及其相关领域研究的新进展

景观规划与管理是景观生态学应用研究的热点,是以景观异质性和景观功能的定量研究为基础,以3S和模型模拟方法为技术支撑的综合性应用领域,它是沟通景观生态学理论研究和实践运用的桥梁。景观规划与管理和景观结构、景观功能与景观动态以及景观尺度、景观过程及景观格局等具有密切的联系,上述方面的相关理论与方法,共同支撑着景观规划与管理的客观实践。景观异质性是景观规划与管理的重要理论基础,景观功能是景观规划与管理的现实目标,景观生态研究方法是景观规划与景观管理的技术保障。20世纪90年代以来,随着学科的交叉与融合,景观规划与管理领域取得了一系列新进展,主要表现在理论、方法和应用方面的研究愈加广泛和深入,景观规划与管理方式也越来越智能化与人性化;景观规划管理的发展也在一定程度上推动了景观异质性、景观功能和景观研究方法的发展。目前,相关学科方法和技术的交叉融合成为研究景观规划与管理的重要趋势,3S和模型交叉融合促进景观规划与管理方法向着多元化、数字化和可视化发展。随着景观规划管理现实应用领域的拓展,景观异质性和景观功能在理论、方法、应用方面的研究进一步深化。     

Zooplankton responses to hypoxia: behavioral patterns and survival of three species of calanoid copepods

Seasonally recurrent and persistent hypoxic events in semi-enclosed coastal waters are characterized by bottom-water dissolved oxygen (d.o.) concentrations of < 2.0 ml l⁻¹. Shifts in the distribution patterns of zooplankters in association with these events have been documented, but the mechanisms responsible for these shifts have not been investigated. This study assessed interspecific differences in responses to hypoxia by several species of calanoid copepods common off Turkey Point, Florida, USA: Labidocera aestiva (Wheeler) (a summer/fall species), Acartia tonsa (Dana) (a ubiquitous year-round species), and Centropages hamatus (Lilljeborg) (a winter/spring species). Under conditions of moderate to severe hypoxia 24-h survival experiments were conducted for adults and nauplii of these species from August 1994 to October 1995. Experiments on adults used a flow-through system to maintain constant d.o. concentrations. Adults of A. tonsa showed no decline in survival with d.o. as low as 1.0 ml l⁻¹, sharp declines in survival at d.o. = 0.9 to 0.6 ml l⁻¹, and 100% mortality with d.o. = 0.5 ml l⁻¹. Adults of L. aestiva and C. hamatus were more sensitive to oxygen depletion: both species experienced significant decreases in survival for d.o. = 1.0 ml l⁻¹. Nauplii of L. aestiva and A. tonsa showed no significant mortality with d.o. = 1.1 to 1.5 ml␣l⁻¹ and d.o. = 0.24 to 0.5 ml l⁻¹, respectively. In addition, experiments investigating behavioral avoidance of moderate to severe hypoxia were carried out for adults of all three species. None of the three species effectively avoided either severely hypoxic (d.o. < 0.5 ml l⁻¹) or moderately hypoxic (d.o. ≈ 1.0 ml l⁻¹) bottom layers in stratified columns. These results suggest that in␣nearshore areas where development of zones of d.o. < 1.0 ml l⁻¹ may be sudden, widespread, or unpredictable, patterns of reduced copepod abundance in bottom waters may be due primarily to mortality rather than avoidance. Received: 31 August 1996 / Accepted: 24 September 1996     

我国海洋自然保护区面临的主要问题及管理策略

我国海洋自然保护区的建立,在保护海洋环境、物种资源,维护海洋生态平衡等方面起了重要作用;然而海洋自然保护区建设管理工作起步较晚、基础薄弱、保护区的管理正处于摸索阶段,还存在一些问题.文章提出了目前我国海洋自然保护区在管理体制、总体布局规划、经费投入、法律法规以及科研工作方面存在的主要问题,并根据这些问题提出相应解决的管理策略.     

Differences in olfactory sensitivity and behavioral responses among honey bees bred for hygienic behavior

Honey bees, Apis mellifera L., bred for hygienic behavior uncap and remove diseased and mite-infested brood. We hypothesized that within a colony bred for hygienic behavior, there would be differences in olfactory sensitivity among bees of the same age. We predicted that bees that initiate the behavior by perforating and uncapping brood would have greater olfactory sensitivity to the odor of the diseased brood, and would be better able to discriminate between odors of healthy and diseased brood, compared to bees that complete the behavior by removing the uncapped brood from the cells. Electroantennogram recordings of 15- to 21-day-old bees from three colonies demonstrated that bees collected while uncapping dead brood had significantly greater olfactory sensitivity to all concentrations of diseased brood odor compared to bees collected while removing brood. Proboscis-extension reflex discrimination conditioning demonstrated that 15- to 21-day-old bees collected while uncapping discriminated significantly better and generalized significantly less between the odors of diseased and healthy brood compared to bees collected while removing, when the odor of diseased brood was rewarded, but not when the odor of healthy brood was rewarded. Bees collected while uncapping brood that had been pierced with a pin had significantly less olfactory sensitivity than bees collected while uncapping freeze-killed brood, most likely because the pierced brood had greater stimulus intensity. Initiation of hygienic behavior depends on the olfactory sensitivity of the bee and stimulus intensity of the abnormal brood. Differential olfactory sensitivity and responsiveness among hygienic bees could lead to the apparent partitioning of the behavior into uncapping and removing components.Communicated by R.F.A. Moritz     

Denitrification and the nitrogen budget of a reservoir in an agricultural landscape.

Denitrification is an important process in aquatic sediments, but its role has not been assessed in the N mass balance of upper-Midwestern (USA) reservoirs that receive large agricultural riverine N inputs. We used a 4400-ha reservoir to determine the role of denitrification in the N mass balance and effectiveness in reducing downstream transport of NO(3-)N. Sediment denitrification was (1) measured monthly (March 2002-March 2003) at eight sites in the Lake Shelbyville reservoir in central Illinois using the acetylene inhibition, chloramphenicol technique, (2) scaled to the overall reservoir and compared to N not accounted for in a mass balance, and (3) estimated indirectly using long-term (1981-2003) mass balances of N in the reservoir. Denitrification rates in the reservoir were high during spring and early summer of 2002, when maximum NO(3-)N concentrations were measured (10-14 mg NO(3-)N/L). We estimated that denitrification for the year was between 2580 and 5150 Mg N. Missing N from the mass balance was 3004 Mg N, suggesting that sediment denitrification was the sink. Areal rates of sediment denitrification in the reservoir ranged from 62 to 225 g N x m(-2) x yr(-1), with rates a function of both denitrification intensity (microg N x g dry mass x h(-1)) and the overall mass of sediment present. From 1981 to 2003 the average NO(3-)N inlet flux was 8900 Mg N/yr. About 58% of the total NO(3-)N input was removed, and annual NO(3-)N removed as a percentage of inputs was significantly related to reservoir retention time (average = 0.36 yr for the 23 years, range = 0.21-0.84 yr). By scaling denitrification in Lake Shelbyville to other reservoirs in Illinois, we estimated a sink of 48900 Mg N/yr. When combined with estimated in-stream denitrification, 60900 Mg N/yr was estimated to be removed by sediment denitrification. This reduces riverine export from Illinois to the Gulf of Mexico, where the flux during the 1990s was about 244000 Mg N/yr, and illustrates the importance of reservoir denitrification as an N sink in Midwestern agricultural landscapes.     

Bee community shifts with landscape context in a tropical countryside.

The ongoing scientific controversy over a putative "global pollination crisis" underscores the lack of understanding of the response of bees (the most important taxon of pollinators) to ongoing global land-use changes. We studied the effects of distance to forest, tree management, and floral resources on bee communities in pastures (the dominant land-use type) in southern Costa Rica. Over two years, we sampled bees and floral resources in 21 pastures at three distance classes from a large (approximately 230-ha) forest patch and of three common types: open pasture; pasture with remnant trees; and pasture with live fences. We found no consistent differences in bee diversity or abundance with respect to pasture management or floral resources. Bee community composition, however, was strikingly different at forest edges as compared to deforested countryside only a few hundred meters from forest. At forest edges, native social stingless bees (Apidae: Meliponini) comprised approximately 50% of the individuals sampled, while the alien honeybee Apis mellifera made up only approximately 5%. Away from forests, meliponines dropped to approximately 20% of sampled bees, whereas Apis increased to approximately 45%. Meliponine bees were also more speciose at forest edge sites than at a distance from forest, their abundance decreased with continuous distance to the nearest forest patch, and their species richness was correlated with the proportion of forest cover surrounding sample sites at scales from 200 to 1200 m. Meliponines and Apis together comprise the eusocial bee fauna of the study area and are unique in quickly recruiting foragers to high-quality resources. The diverse assemblage of native meliponine bees covers a wide range of body sizes and flower foraging behavior not found in Apis, and populations of many bee species (including Apis), are known to fluctuate considerably from year to year. Thus, the forest-related changes in eusocial bee communities we found may have important implications for: (1) sustaining a diverse bee fauna in tropical countryside; (2) ensuring the effective pollination of a diverse native plant community; and (3) the efficiency and stability of agricultural pollination, particularly for short-time-scale, mass-flowering crops such as coffee.     

Integrating economic costs into the analysis of flexible conservation management strategies.

Flexible conservation management, where measures (e.g., mowing of meadows, removing invasive species) are selected in each decision period depending on the current state of the ecological system, is generally perceived as superior to fixed management, where the same measure is applied in each decision period independent of the current state of the system. In past comparisons of fixed and flexible conservation strategies the additional costs that arise only in flexible strategies have usually been ignored. In this paper, we present a framework to integrate costs of flexible management into the evaluation of flexible conservation strategies. Using the example of an endangered butterfly species we demonstrate that the costs of flexible management may reverse the rank order of flexible and fixed conservation strategies, such that fixed strategies may lead to better ecological results than flexible ones for the same financial budget.     

Conditioning uncertainty in ecological models: Assessing the impact of fire management strategies

A simple simulation model has been used to investigate whether large fires in Mediterranean regions are a result of extreme weather conditions or the cumulative effect of a policy of fire suppression over decades. The model reproduced the fire regime characteristics for a wide variety of regions of Mediterranean climate in California, France and Spain. The Generalised Likelihood Uncertainty Estimation (GLUE) methodology was used to assess the possibility of multiple model parameter sets being consistent with the available calibration data. The resulting set of behavioural models was used to assess uncertainty in the predictions. The results suggested that (1) for a given region, the total area burned is much the same whether suppression or prescribed fire policies are used or not; however fire suppression enhances fire intensity and prescribed burning reduces it; (2) the proportion of large fires can be reduced, but not eliminated, using prescribed fires, especially in areas which have the highest proportion of large fires.     

Grassland restoration with and without fire: evidence from a tree-removal experiment

Forest encroachment threatens the biological diversity of grasslands globally. Positive feedbacks can reinforce the process, affecting soils and ground vegetation, ultimately leading to replacement of grassland by forest species. We tested whether restoration treatments (tree removal, with or without fire) reversed effects of nearly two centuries of encroachment by Abies grandis and Pinus contorta into dry, montane meadows in the Cascade Range, Oregon, USA. In nine, 1-ha plots containing a patchy mosaic of meadow openings and forests of varying age (20 to > 140 yr), we compared three treatments affecting the ground vegetation: control (no trees removed), unburned (trees removed, slash burned in piles leaving 90% of the area unburned), and burned (trees removed, slash broadcast burned). We quantified changes over 3-4 years in soils, abundance and richness of species with differing habitat associations (meadow, forest, and ruderal), and recruitment of conifers. Except for a transient increase in available N (especially in burn scars), effects of burning on soils were minimal due, in part, to mixing by gophers. Tree removal greatly benefited meadow species at the expense of forest herbs. Cover and richness of meadow species increased by 47% and 38% of initial values in unburned plots, but changed minimally in burned plots. In contrast, cover and richness of forest herbs declined by 44% and 26% in unburned plots and by 79% and 58% in burned plots. Ruderal species and conifer seedlings were uncommon in both treatments. Although vegetation was consumed beneath burn piles, meadow species recovered significantly after three years. Long-term tree presence did not preclude recovery of meadow species; in fact, colonization was greater in older than in younger forests. In sum, temporal trends were positive for most indicators, suggesting strong potential for restoration. Contrary to conventional wisdom, tree removal without fire may be sufficient to shift the balance from forest to meadow species. In meadows characterized by historically infrequent fire, small-scale disturbances and competitive interactions may be more critical to ecosystem maintenance and restoration. Managers facing the worldwide phenomenon of tree invasion should critically evaluate the ecological vs. operational need for fire in ecosystem restoration.     

Disturbance and landscape dynamics in a changing world

Disturbance regimes are changing rapidly, and the consequences of such changes for ecosystems and linked social-ecological systems will be profound. This paper synthesizes current understanding of disturbance with an emphasis on fundamental contributions to contemporary landscape and ecosystem ecology, then identifies future research priorities. Studies of disturbance led to insights about heterogeneity, scale, and thresholds in space and time and catalyzed new paradigms in ecology. Because they create vegetation patterns, disturbances also establish spatial patterns of many ecosystem processes on the landscape. Drivers of global change will produce new spatial patterns, altered disturbance regimes, novel trajectories of change, and surprises. Future disturbances will continue to provide valuable opportunities for studying pattern-process interactions. Changing disturbance regimes will produce acute changes in ecosystems and ecosystem services over the short (years to decades) and long-term (centuries and beyond). Future research should address questions related to (1) disturbances as catalysts of rapid ecological change, (2) interactions among disturbances, (3) relationships between disturbance and society, especially the intersection of land use and disturbance, and (4) feedbacks from disturbance to other global drivers. Ecologists should make a renewed and concerted effort to understand and anticipate the causes and consequences of changing disturbance regimes.