Species, functional groups, and thresholds in ecological resilience

The cross-scale resilience model states that ecological resilience is generated in part from the distribution of functions within and across scales in a system. Resilience is a measure of a system's ability to remain organized around a particular set of mutually reinforcing processes and structures, known as a regime. We define scale as the geographic extent over which a process operates and the frequency with which a process occurs. Species can be categorized into functional groups that are a link between ecosystem processes and structures and ecological resilience. We applied the cross-scale resilience model to avian species in a grassland ecosystem. A species' morphology is shaped in part by its interaction with ecological structure and pattern, so animal body mass reflects the spatial and temporal distribution of resources. We used the log-transformed rank-ordered body masses of breeding birds associated with grasslands to identify aggregations and discontinuities in the distribution of those body masses. We assessed cross-scale resilience on the basis of 3 metrics: overall number of functional groups, number of functional groups within an aggregation, and the redundancy of functional groups across aggregations. We assessed how the loss of threatened species would affect cross-scale resilience by removing threatened species from the data set and recalculating values of the 3 metrics. We also determined whether more function was retained than expected after the loss of threatened species by comparing observed loss with simulated random loss in a Monte Carlo process. The observed distribution of function compared with the random simulated loss of function indicated that more functionality in the observed data set was retained than expected. On the basis of our results, we believe an ecosystem with a full complement of species can sustain considerable species losses without affecting the distribution of functions within and across aggregations, although ecological resilience is reduced. We propose that the mechanisms responsible for shaping discontinuous distributions of body mass and the nonrandom distribution of functions may also shape species losses such that local extinctions will be nonrandom with respect to the retention and distribution of functions and that the distribution of function within and across aggregations will be conserved despite extinctions.     

Space partitioning and interactions in an intertidal sand-burrowing amphipod guild

A nine-year study at a moderately exposed, fine sand intertidal beach at Long Sands, Maine, USA identified marked seasonal changes in the foci of sand-burrowing amphipod activities. These included seasonal population movements, degree of interspecific horizontal overlap, life stage segregation within and between species, and sand depth stratification. Behavioral interactions related to competition for available space are considered important in affecting the distribution and abundance of amphipod species, and community structure. Significant negative correlations were shown between the abundance and upper distributional limits of the smaller, lower shore Acanthohaustorius millsi and the abundance of the larger, higher shore species Haustorius canadensis. Tolerance limits to a variety of prevailing physical factors alone, were not considered a serious obstacle to occupation of higher intertidal level sands by A. millsi. Summer brooding females of both species occupied similar 5 to 10 cm sand depths, but were strongly segregated horizontally. Laboratory coexistence experiments during the reproductive period showed increased mortality of adults and negligible reproductive output of A. millsi when in combination with H. canadensis compared to controls where the former species was alone. A third species, Amphiporeia virginiana, occupied primarily very shallow sand (0 to 2.5 cm), and performed seasonal movements opposite in direction to A. millsi and H. canadensis. The presence, amount, and refinement of biological interactions across the wave exposure gradient requires further study. Until then, we consider as inappropriate the blanket designation of all types of intertidal sand communities as physically controlled.     

Multivariate trade-offs, succession, and phenological differentiation in a guild of colonial invertebrates

For competing species limited by one or few resources, diversity is thought to be maintained by trade-offs that allow niche differentiation without resource partitioning. However, few studies have quantified multiple key traits for each species in a guild and shown that trade-offs among these traits apply across the guild. Here we document strong bivariate and multivariate relationships among growth rate, fecundity, longevity, and overgrowth ability for six co-occurring colonial invertebrates. We find that all four of these traits are constrained to a single "fast-slow" niche axis that mechanistically relates life history variation to a colonization-competition trade-off. The location of species on this axis strongly predicts the timing of their peak abundance during succession. We also find that species closer to each other on the fast-slow axis are more likely to differ in reproductive phenology, suggesting a secondary dimension of niche differentiation for otherwise similar species.     

Lethal and nonlethal predator effects on an herbivore guild mediated by system productivity

Indirect effects propagated through intervening species in a food web have important effects on community properties. Traditionally, these indirect effects have been conceptualized as mediated through density changes of the intervening species, but it is becoming increasingly apparent that those mediated through trait (phenotypic) responses also can be very important. Because density- and trait-mediated indirect effects have different properties, it is critical that we understand the mechanisms of transmission in order to predict how they will interact, and when or where they will be important. In this study, we examined the mechanisms and consequences of the lethal (density-mediated) and nonlethal (trait-mediated) effects of a larval odonate predator on a guild of four herbivore species (a larval anuran and three species of snails) and their resources. We also manipulated system productivity in order to explore the effects of environmental context on the transmission of these two types of indirect effects. We show that trait-mediated effects arising from the predator can be very strong relative to density-mediated effects on both the competing herbivores and the species composition and production of their resources. A number of these indirect effects are shown to be contingent on productivity of the system. We further present evidence that trait- and density-mediated indirect effects originating from a predator may be transmitted independently through different routes in a food web, particularly when spatial responses of the transmitting prey are involved. Finally, effects on prey growth due to trait responses to the predator varied from negative to positive in predictable ways as a function of time and indirect effects on the larger food web. These results indicate the important role that trait-mediated indirect effects can play in trophic cascades and keystone predator interactions, and we discuss how the mechanisms involved can be incorporated in theory.     

Transvalued Species in an African Forest

Abstract: We combined ethnographic investigations with repeated ecological transect surveys in the Dzanga‐Sangha Dense Forest Reserve (RDS), Central African Republic, to elucidate consequences of intensifying mixed use of forests. We devised a framework for transvaluation of wildlife species, which means the valuing of species on the basis of their ecological, economic, and symbolic roles in human lives. We measured responses to hunting, tourism, and conservation of two transvalued species in RDS: elephants (Loxodonta cyclotis) and gorillas (Gorilla gorilla). Our methods included collecting data on encounter rates and habitat use on line transects. We recorded cross‐cultural variation in ideas about and interactions with these species during participant observation of hunting and tourism encounters and ethnographic interviews with hunters, conservation staff, researchers, and tourists. Ecologically, gorillas used human‐modified landscapes successfully, and elephants were more vulnerable than gorillas to hunting. Economically, tourism and encounters with elephants and gorillas generated revenues and other benefits for local participants. Symbolically, transvaluation of species seemed to undergird competing institutions of forest management that could prove unsustainable. Nevertheless, transvaluation may also offer alternatives to existing social hierarchies, thereby integrating local and transnational support for conservation measures. The study of transvaluation requires attention to transnational flows of ideas and resources because they influence transspecies interactions. Cross‐disciplinary in nature, transvalution of species addresses the political and economic challenges to conservation because it recognizes the varied human communities that shape the survival of wildlife in a given site. Transvaluation of species could foster more socially inclusive management and monitoring approaches attuned to competing economic demands, specific species behaviors, and human practices at local scales.     

Temporal variability in functional responses and prey selectivity of the pelagic mysid, Mysis mixta, in natural prey assemblages

Predation rates and prey selection of the pelagic mysid shrimp, Mysis mixta, were studied experimentally in the northern Baltic Sea in 1998 during their most intensive growth period, from June to October. Functional responses during 5 months were determined by providing the mysids with a natural zooplankton assemblage, diluted to several different concentrations. The results show that ingestion rate increased, along with mysid growth, from early summer to autumn and that saturation level was reached between 400 and 500 μg C l⁻¹. Ingestion rates increased with increasing prey concentration, and sigmoidal curves explained mostly the variation in ingestion rates (explanatory levels of 86–97%). Prey selection was evident in June, July and August, though weaker during the latter 2 months. Selection differed between the studied months but, generally, copepods were more positively selected than cladocerans. Rotifers were the main prey during June and July, when mysids were small, while larger mysids fed on copepods and cladocerans. Of the copepods, Eurytemora affinis was a truly selected species. This study shows that mysids feed on many zooplankton taxa and that they undergo ontogenetic diet shifts. Received: 19 July 2000 / Accepted: 19 October 2000     

Alloimmune responses between Acropora hemprichi conspecifics: nontransitive patterns of overgrowth and delayed cytotoxicity

Allogeneic interactions between Acropora hemprichi (Ehrenberg) colonies were followed in situ up to 18 mo between 1990 and 1992 in Eilat, Red Sea. We established all 21 possible branch pair combinations among seven large colonies in three to eight replicates each (n=92 pairs), in addition in 14 pairs of grafts within-colonies. All isografts resulted in fusion. Allogeneic interactions revealed a variety of effector mechanisms which involved the appearance of dark blue-purple color in the tissues of interacting branch tips (<1 cm), unilateral overgrowths (1.0 to 1.5 mm mo^-1 for 1 to 8 mo), rejections, healings, continuous overgrowths, reversals, repeated rejections and more. Direction of overgrowth was usually specific to a given combination of colonies. A hierarchial network of overgrowths was obtained by analyzing overgrowth directions of all pairwise combinations, which included 30 transitive and five nontransitive three-way combinations. These outcomes, similarly to responses of allogeneic encounters recorded from other coral species, show that colonies can respond selectively to different types of allogeneic challenges. It is suggested that corals may have the capacity to distinguish self from nonself by recognizing nonself attributes presented on conspecifics.     

川西北高寒草甸两种蚯蚓的互作及其对土壤养分的影响

以川西北高寒草甸中普遍存在的两种蚯蚓(微小双胸蚓Bimastus parvus和威廉腔蚓Metaphire guillemi)为研究对象,采用野外微宇宙实验方法比较单独接种和混合接种时两种蚯蚓的数量、分布和繁殖状况,以及牛粪分解率和土壤养分含量的变化,探讨不同蚯蚓种群间的相互作用及其对土壤养分的影响.结果发现:1)两种蚯蚓混合接种加快了牛粪分解,增加了下层(10~20 cm)土壤可溶性氮含量,但对牛粪分解速率和土壤可溶性氮含量均不存在交互作用;2)在混合接种处理中,微小双胸蚓的死亡比率显著高于威廉腔蚓的死亡比率;3)两种蚯蚓混合接种处理时,威廉腔蚓的繁殖明显受到抑制,而微小双胸蚓的幼蚓数量有所增加.研究表明,威廉腔蚓和微小双胸蚓共存能够发生竞争性相互作用,加快地面上的牛粪分解,增加土壤可溶性氮含量,从而可能提高高寒草甸的初级生产力.     

Ecological differences and coexistence in a guild of microparasites: Bartonella in wild rodents

The study of ecological differences among coexisting microparasites has been largely neglected, but it addresses important and unusual issues because there is no clear distinction in such cases between conventional (resource) and apparent competition. Here patterns in the population dynamics are examined for four species of Bartonella (bacterial parasites) coexisting in two wild rodent hosts, bank voles (Clethrionomys glareolus) and wood mice (Apodemus sylvaticus). Using generalized linear modeling and mixed effects models, we examine, for these four species, seasonal patterns and dependencies on host density (both direct and delayed) and, having accounted for these, any differences in prevalence between the two hosts. Whereas previous studies had failed to uncover species differences, here all four were different. Two, B. doshiae and B. taylorii, were more prevalent in wood mice, and one, B. birtlesii, was more prevalent in bank voles. B. birtlesii, B. grahamii, and B. taylorii peaked in prevalence in the fall, whereas B. doshiae peaked in spring. For B. birtlesii in bank voles, density dependence was direct, but for B. taylorii in wood mice density dependence was delayed. B. birtlesii prevalence in wood mice was related to bank vole density. The implications of these differences for species coexistence are discussed.     

Detecting general plant functional type responses in fragmented landscapes using spatially-explicit simulations

Habitat loss and increasing landscape fragmentation are known to be key forces driving the ongoing loss of plant species diversity. While the combined effects of increasing isolation and decreasing population size have been studied intensively; it is less understood how plant population performance in heterogeneous landscapes is affected by changes in fragmentation alone.     

Local interactions between predators and prey call into question commonly used functional responses

Commonly used functional response models (Holling’s type I and type II models) assume that the encounter rate of a predator increases linearly with prey density, provided that the predator is searching for prey. In other other words, aN (a is the baseline encounter rate and N is prey density) describes the encounter rate. This study examined whether the models are adequate when predators and prey interact locally by using a spatially explicit individual based model because local interactions affect the spatial distribution of predators and prey, which also affects the encounter rate. Predators were assumed to possess a spatial perception range that influenced their foraging behavior (e.g., if a prey is in the perception range, the predator moves towards the prey). The effect of antipredator behavior by prey was also examined. The results suggest that prey and predator densities as well as handling time affect the baseline rate (i.e., parameter a) as opposed to the common assumption that the parameter is constant. The nature of model deviations depended on both the antipredator behavior and the predators’ perception range. Understanding these deviations is important as they qualitatively affect community dynamics.     

Economic Science, Endangered Species, and Biodiversity Loss

Abstract: Although economic analysis can be used to argue for preservation of species and habitats, many natural assets represent inferior investments in society's asset portfolio. We demonstrate this for the case of ancient temperate rainforests and minke whales (  Balaenoptera acutorostrata ). For both rainforests and whales, we determined their value for harvest and balanced this against society's valuation of the preserved stock. For the market and nonmarket data available, we then determined how much rainforest and how many minke whales global society should keep in its asset portfolio. Although ecologists increasingly attempt to justify preservation of biological assets on economic grounds, we argue that this might be a dangerous approach to take. Ultimately, it may be necessary to reexamine the ethical foundations for conservation of nature and biodiversity, including the economist's use of utilitarianism. We suggest that the safe minimum standard approach may prove useful in practice.     

Regional Decline of an Iconic Amphibian Associated with Elevation,Land‐Use Change,and Invasive Species

Abstract: Ecological theory predicts that species with restricted geographic ranges will have the highest probability of extinction, but species with extensive distributions and high population densities can also exhibit widespread population losses. In the western United States populations of northern leopard frogs (Lithobates pipiens)—historically one of the most widespread frogs in North America—have declined dramatically in abundance and geographic distribution. To assess the status of leopard frogs in Colorado and evaluate causes of decline, we coupled statewide surveys of 196 historically occupied sites with intensive sampling of 274 wetlands stratified by land use. We used an information‐theoretic approach to evaluate the contributions of factors at multiple spatial extents in explaining the contemporary distribution of leopard frogs. Our results indicate leopard frogs have declined in Colorado, but this decline was regionally variable. The lowest proportion of occupied wetlands occurred in eastern Colorado (2–28%), coincident with urban development and colonization by non‐native bullfrogs (Lithobates catesbeianus). Variables at several spatial extents explained observed leopard frog distributional patterns. In low‐elevation wetlands introduced fishes, bullfrogs, and urbanization or suburbanization associated negatively with leopard frog occurrence, whereas wetland area was positively associated with occurrence. Leopard frogs were more abundant and widespread west of the Continental Divide, where urban development and bullfrog abundance were low. Although the pathogenic chytrid Batrachochytrium dendrobatidis (Bd) was not selected in our best‐supported models, the nearly complete extirpation of leopard frogs from montane wetlands could reflect the individual or interactive effects of Bd and climate patterns. Our results highlight the importance of considering multiple, competing hypotheses to explain species declines, particularly when implicated factors operate at different spatial extents.     

Testing the field of dreams hypothesis: functional responses to urbanization and restoration in stream ecosystems

As catchments become increasingly urban, the streams that drain them become increasingly degraded. Urban streams are typically characterized by high-magnitude storm flows, homogeneous habitats, disconnected riparian zones, and elevated nitrogen concentrations. To reverse the degradation of urban water quality, watershed managers and regulators are increasingly turning to stream restoration approaches. By reshaping the channel and reconnecting the surface waters with their riparian zone, practitioners intend to enhance the natural nutrient retention capacity of the restored stream ecosystem. Despite the exponential growth in stream restoration projects and expenditures, there has been no evaluation to date of the efficacy of urban stream restoration projects in enhancing nitrogen retention or in altering the underlying ecosystem metabolism that controls instream nitrogen consumption. In this study, we compared ecosystem metabolism and nitrate uptake kinetics in four stream restoration projects within urban watersheds to ecosystem functions measured in four unrestored urban stream segments and four streams draining minimally impacted forested watersheds in central North Carolina, U.S.A. All 12 sites were surveyed in June through August of 2006 and again in January through March of 2007. We anticipated that urban streams would have enhanced rates of ecosystem metabolism and nitrate uptake relative to forested streams due to the increases in nutrient loads and temperature associated with urbanization, and we predicted that restored streams would have further enhanced rates for these ecosystem functions by virtue of their increased habitat heterogeneity and water residence times. Contrary to our predictions we found that stream metabolism did not differ between stream types in either season and that nitrate uptake kinetics were not different between stream types in the winter. During the summer, restored stream reaches had substantially higher rates of nitrate uptake than unrestored or forested stream reaches; however, we found that variation in stream temperature and canopy cover explained 80% of the variation across streams in nitrate uptake. Because the riparian trees are removed during the first stage of natural channel design projects, the restored streams in this study had significantly less canopy cover and higher summer temperatures than the urban and forested streams with which they were compared.     

Incentive Structure of and Private Landowner Participation in an Endangered Species Conservation Program

Abstract: In the United States, voluntary incentive programs that aid conservation of plant and wildlife species on private lands provide a structural solution to the problem of protecting endangered species by reducing costs and enhancing benefits to landowners. We explored the potential for incentives to encourage landowners to manage land cover for the benefit of endangered songbirds in central Texas (U.S.A.) by asking landowners to indicate their preferences for financial incentives, technical assistance, and regulatory assurances. We identified owners of potential songbird habitat and collected data in face‐to‐face interviews and self‐administered questionnaires. We used a latent‐class stated‐choice model to identify 3 classes of landowners whose choices varied on the basis of their attitudes and perceived social norms: (1) strong positive attitude, perceived social pressure to participate, and willing to participate with relatively few incentives, (2) weak positive attitude, perceived no social pressure to participate, and required strongest incentives, and (3) negative attitude, perceived social pressure not to enroll, and unwilling to participate regardless of incentive structure. Given this heterogeneity in preferences, conservation incentives may increase management of land cover to benefit endangered species on private lands to some degree; however, exclusive reliance on incentives may be insufficient. Promoting conservation on private lands may be enhanced by integrating incentives into an approach that incorporates other strategies for conservation, including social networks and collaborative processes that reinforce social norms.