Application of the New Keystone-Species Concept to Prairie Dogs: How Well Does It Work?

Abstract: It has been suggested that the keystone-species concept should be dropped from ecology and conservation, primarily because the concept is poorly defined. This prompted Power et al. (1996) to refine the definition: keystone species have large effects on community structure or ecosystem function (i.e., high overall importance), and this effect should be large relative to abundance (i.e., high community importance). Using prairie dogs ( Cynomys spp.) as an example, I review operational and conceptual difficulties encountered in applying this definition. As applied to prairie dogs, the implicit assumption that overall importance is a linear function of abundance is invalid. In addition, community importance is sensitive to abundance levels, the definition of community, and sampling scale. These problems arise largely from the equation for community importance, as used in conjunction with removal experiments at single abundance levels. I suggest that we shift from the current emphasis on the dualism between keystone and nonkeystone species and instead examine how overall and community importance vary (1) with abundance, (2) across spatial and temporal scales, and (3) under diverse ecological conditions. In addition, I propose that a third criterion be incorporated into the definition: keystone species perform roles not performed by other species or processes. Examination of how these factors vary among populations of keystone species should help identify the factors contributing to, or limiting, keystone-level functions, thereby increasing the usefulness of the keystone-species concept in ecology and conservation. Although the quantitative framework of Power et al. falls short of being fully operational, my conceptual guidelines may improve the usefulness of the keystone-species concept. Careful attention to the factors that limit keystone function will help avoid misplaced emphasis on keystone species at the expense of other species.     

Coupling stable isotopes with bioenergetics to estimate interspecific interactions.

Interspecific interactions are often difficult to elucidate, particularly with large vertebrates at large spatial scales. Here, we describe a methodology for estimating interspecific interactions by combining stable isotopes with bioenergetics. We illustrate this approach by modeling the population dynamics and species interactions of a suite of vertebrates on Santa Cruz Island, California, USA: two endemic carnivores (the island fox and island spotted skunk), an exotic herbivore (the feral pig), and their shared predator, the Golden Eagle. Sensitivity analyses suggest that our parameter estimates are robust, and natural history observations suggest that our overall approach captures the species interactions in this vertebrate community. Nonetheless, several factors provide challenges to using isotopes to infer species interactions. Knowledge regarding species-specific isotopic fractionation and diet breadth is often lacking, necessitating detailed laboratory studies and natural history information. However, when coupled with other approaches, including bioenergetics, mechanistic models, and natural history, stable isotopes can be powerful tools in illuminating interspecific interactions and community dynamics.     

Competitive exclusion within the predator community influences the distribution of a threatened prey species

While much effort has been made to quantify how landscape composition influences the distribution of species, the possibility that geographical differences in species interactions might affect species distributions has received less attention. Investigating a predator-prey setting in a boreal forest ecosystem, we empirically show that large-scale differences in the predator community structure and small-scale competitive exclusion among predators affect the local distribution of a threatened forest specialist more than does landscape composition. Consequently, even though the landscape parameters affecting Siberian flying squirrel (Pteromys volans) distribution (prey) did not differ between nest sites of the predators Northern Goshawks (Accipiter gentilis) and Ural Owls (Strix uralensis), flying squirrels were heterospecifically attracted by goshawks in a region where both predator species were present. No such effect was found in another region where Ural Owls were absent. These results provide evidence that differences in species interactions over large spatial scales may be a major force influencing the distribution and abundance patterns of species. On the basis of these findings, we suspect that subtle species interactions might be a central reason why landscape models constructed to predict species distributions often fail when applied to wider geographical scales.     

Responses of microbial interactions to elevated salinity in activated sludge microbial community

● Salinity led to the elevation of NAR over 99.72%. ● Elevated salinity resulted in a small, complex, and more competitive network. ● Various AOB or denitrifiers responded differently to elevated salinity. ● Putative keystone taxa were dynamic and less abundant among various networks. Biological treatment processes are critical for sewage purification, wherein microbial interactions are tightly associated with treatment performance. Previous studies have focused on assessing how environmental factors (such as salinity) affect the diversity and composition of the microbial community but ignore the connections among microorganisms. Here, we described the microbial interactions in response to elevated salinity in an activated sludge system by performing an association network analysis. It was found that higher salinity resulted in low microbial diversity, and small, complex, more competitive overall networks, leading to poor performance of the treatment process. Subnetworks of major phyla (Proteobacteria, Bacteroidetes, and Chloroflexi) and functional bacteria (such as AOB, NOB and denitrifiers) differed substantially under elevated salinity process. Compared with subnetworks of Nitrosomonadaceae, Nitrosomonas (AOB) made a greater contribution to nitrification under higher salinity (especially 3%) in the activated sludge system. Denitrifiers established more proportion of cooperative relationships with other bacteria to resist 3% salinity stress. Furthermore, identified keystone species playing crucial roles in maintaining process stability were dynamics and less abundant under salinity disturbance. Knowledge gleaned from this study deepened our understanding of microbial interaction in response to elevated salinity in activated sludge systems.     

How a songbird with a continuous singing style modulates its song when territorially challenged

In songbirds of the temperate zone, often only males sing and their songs serve to attract females and to deter territorial rivals. In many species, males vary certain aspects of their singing behavior when engaged in territorial interactions. Such variation may be an honest signal of the traits of the signaler, such as fighting strength, condition, or aggressive motivation, and may be used by receivers in decisions on whether to retreat or to escalate a fight. This has been studied intensively in species that sing discontinuously, in which songs are alternating with silent pauses. We studied contextual variation in the song of skylarks (Alauda arvensis), a songbird with a large vocal repertoire and a continuous and versatile singing style. We exposed subjects to simulated territorial intrusions by broadcasting conspecific song and recorded their vocal responses. We found that males sing differently if they are singing spontaneously with no other conspecific around than if they are territorially challenged. In this last case, males produced lower-frequency syllables. Furthermore, they increased the sound density of their song: they increased the proportion of sound within song. They seem to do so by singing different elements of their repertoire when singing reactively. Furthermore, they increased the consistency of mean peak frequency: they repeated syllable types with less variability when singing reactively. Such contextual variation suggests that skylarks might use low frequencies, sound density, and song consistency to indicate their competitive potential, and thus, those song features might be important for mutual assessment of competitive abilities.     

Habitat complexity impacts persistence and species interactions in an intertidal whelk

Although experiments have shown that habitat structure may influence the distribution of species and species interactions, these effects are still not commonly integrated into studies of community dynamics. Since habitat structure often varies within and among communities, this may limit our understanding of how various factors influence communities. Here, we examined how mussel bed complexity (the presence and thickness of mussel layers) influenced the persistence of whelks (Nucella emarginata) and interactions with a top predator (ochre sea stars, Pisaster ochraceus) and prey (mussels, Mytilus californianus). Results from a mark?Crecapture experiment indicate that whelk recapture rates are higher in more complex habitats, and laboratory experiments demonstrate that habitat complexity affects whelk feeding, growth, and nonconsumptive interactions with a keystone predator. Habitat complexity therefore has direct effects on species and also may lead to trade-offs among feeding, refuge, and other factors, potentially influencing the distribution of whelks and the effects of both whelks and sea stars on intertidal communities. These results demonstrate that habitat structure may play an important role in intertidal communities and other habitats and should be further considered in the experimental design of future studies of community dynamics.     

Incorporating low-resolution historic species location data decreases performance of distribution models

Developing robust species distribution models is important as model outputs are increasingly being incorporated into conservation policy and management decisions. A largely overlooked component of model assessment and refinement is whether to include historic species occurrence data in distribution models to increase the data sample size. Data of different temporal provenance often differ in spatial accuracy and precision. We test the effect of inclusion of historic coarse-resolution occurrence data on distribution model outputs for 187 species of birds in Australian tropical savannas. Models using only recent (after 1990), fine-resolution data had significantly higher model performance scores measured with area under the receiver operating characteristic curve (AUC) than models incorporating both fine- and coarse-resolution data. The drop in AUC score is positively correlated with the total area predicted to be suitable for the species (R² = 0.163-0.187, depending on the environmental predictors in the model), as coarser data generally leads to greater predicted areas. The remaining unexplained variation is likely to be due to the covariate errors resulting from resolution mismatch between species records and environmental predictors. We conclude that decisions regarding data use in species distribution models must be conscious of the variation in predictions that mixed-scale datasets might cause.     

Potential of Marine Reserves to Cause Community-Wide Changes beyond Their Boundaries

Abstract:  Fishing and other human activities can alter the abundances, size structure, and behavior of species playing key roles in shaping marine communities (e.g., keystone predators), which may in turn cause ecosystem shifts. Despite extensive evidence that cascading trophic interactions can underlie community-wide recovery inside no-take marine reserves by protecting high-level predators, the spatial extent of these effects into adjacent fished areas is unknown. I examined the potential for community-wide changes (i.e., the transition from overgrazed coralline barrens to macroalgal beds) in temperate rocky reefs within and around a no-take marine reserve. For this purpose I assessed distribution patterns of predatory fishes, sea urchins, and barrens across the reserve boundaries. Predatory fishes were significantly more abundant within the reserve than in adjacent locations, with moderate spillover across the reserve edges. In contrast, community-wide changes of benthic assemblages were apparent well beyond the reserve boundaries, which is consistent with temporary movements of predatory fishes (e.g., foraging migration) from the reserve to surrounding areas. My results suggest that no-take marine reserves can promote community-wide changes beyond their boundaries.     

Modeling changes in the coastal ecosystem of the Pearl River Estuary from 1981 to 1998

The coastal ecosystem of the Pearl River Estuary (PRE) has been overfished and received a high level of combined pollution since the 1980s. Ecopath with Ecosim was used to construct two ecosystem models (for 1981 and 1998) to characterize the food web structure and functioning of the ecosystem. Pedigree work and simple sensitivity analysis were carried out to evaluate the quality of data and the uncertainty of the models. The two models seem reliable with regards to input data of good quality. Comparing the variations of outputs of these two models aimed to facilitate assessment of changes of the ecosystem during the past two decades.The trophic structure of the ecosystem has changed with an increase in the biomass proportion of lower trophic level (TL) organisms and a decrease in top predator biomass proportion. All the indices of ecosystem maturity examined show that the system was in a more mature condition in 1981 than in 1998, although the system has been in a condition of stress due to anthropogenic disturbances, such as environmental pollution and habitat destruction since 1981. The ecosystem was aggregated into six and seven integral TLs in 1981 and 1998, respectively, using the trophic aggregation routine of Ecopath. Most of the total system biomass and catch took place at TL II and III in both years. But the distribution of the total system biomass and catch at different TLs changed with decreasing proportions in higher TLs in 1998. The mean transfer efficiency was 9.1% and 10.2% in 1981 and 1998, respectively.Comparative network analysis allowed quantification of the importance of direct and indirect trophic interactions among functional groups. Moreover, a method derived from the mixed trophic impact (MTI) analysis allowed estimating importance of groups in terms of “keystoneness” and identifying the keystone species in the two models over the past two decades. The results indicate that there were no clear keystone species in 1998 but two keystone species at medium trophic levels were identified in 1981. Moreover, organisms located at low trophic levels such as phytoplankton, zooplankton and benthic invertebrates were identified to have relatively high keystoneness in the ecosystem.     

Role of the fish Astyanax aeneus (Characidae) as a keystone nutrient recycler in low-nutrient neotropical streams

Nutrient recycling by animals is a potentially important biogeochemical process in both terrestrial and aquatic ecosystems. Stoichiometric traits of individual species may result in some taxa playing disproportionately important roles in the recycling of nutrients relative to their biomass, acting as keystone nutrient recyclers. We examined factors controlling the relative contribution of 12 Neotropical fish species to nutrient recycling in four streams spanning a range of phosphorus (P) levels. In high-P conditions (135 microg/L soluble reactive phosphorus, SRP), most species fed on P-enriched diets and P excretion rates were high across species. In low-P conditions (3 microg/L SRP), aquatic food resources were depleted in P, and species with higher body P content showed low rates of P recycling. However, fishes that were subsidized by terrestrial inputs were decoupled from aquatic P availability and therefore excreted P at disproportionately high rates. One of these species, Astyanax aeneus (Characidae), represented 12% of the total population and 18% of the total biomass of the fish assemblage in our focal low-P study stream but had P excretion rates > 10-fold higher than other abundant fishes. As a result, we estimated that P excretion by A. aeneus accounted for 90% of the P recycled by this fish assemblage and also supplied approximately 90% of the stream P demand in this P-limited ecosystem. Nitrogen excretion rates showed little variation among species, and the contribution of a given species to ecosystem N recycling was largely dependent upon the total biomass of that species. Because of the high variability in P excretion rates among fish species, ecosystem-level P recycling could be particularly sensitive to changes in fish community structure in P-limited systems.     

Revisiting the classics: considering nonconsumptive effects in textbook examples of predator-prey interactions

Predator effects on prey dynamics are conventionally studied by measuring changes in prey abundance attributed to consumption by predators. We revisit four classic examples of predator-prey systems often cited in textbooks and incorporate subsequent studies of nonconsumptive effects of predators (NCE), defined as changes in prey traits (e.g., behavior, growth, development) measured on an ecological time scale. Our review revealed that NCE were integral to explaining lynx-hare population dynamics in boreal forests, cascading effects of top predators in Wisconsin lakes, and cascading effects of killer whales and sea otters on kelp forests in nearshore marine habitats. The relative roles of consumption and NCE of wolves on moose and consequent indirect effects on plant communities of Isle Royale depended on climate oscillations. Nonconsumptive effects have not been explicitly tested to explain the link between planktonic alewives and the size structure of the zooplankton, nor have they been invoked to attribute keystone predator status in intertidal communities or elsewhere. We argue that both consumption and intimidation contribute to the total effects of keystone predators, and that characteristics of keystone consumers may differ from those of predators having predominantly NCE. Nonconsumptive effects are often considered as an afterthought to explain observations inconsistent with consumption-based theory. Consequently, NCE with the same sign as consumptive effects may be overlooked, even though they can affect the magnitude, rate, or scale of a prey response to predation and can have important management or conservation implications. Nonconsumptive effects may underlie other classic paradigms in ecology, such as delayed density dependence and predator-mediated prey coexistence. Revisiting classic studies enriches our understanding of predator-prey dynamics and provides compelling rationale for ramping up efforts to consider how NCE affect traditional predator-prey models based on consumption, and to compare the relative magnitude of consumptive and NCE of predators.     

Asymmetric mate choice,hybridization, and hybrid fitness in two sympatric grasshopper species

Hybridization between animal species has long been regarded as unusual, but is meanwhile accepted as a widespread phenomenon. Typically, sexual interactions among species are studied in secondary contact zones of closely related species (hybrid zones) or between invasive and native species, whereas hybridization between sympatric congeners has received little attention. Here, we present a study on the hybridization potential of two grasshopper species, Chorthippus parallelus and Chorthippus montanus, which occur sympatric in large parts of Eurasia. We performed a mate choice experiment with specimens from an area of local parapatry in northwestern Germany. Most copulations were conspecific, but males of both species and females of C. parallelus were indiscriminate in their mate choice, while females of C. montanus clearly preferred conspecific males. Although these results suggest asymmetric introgression, a no-choice hybridization experiment revealed that hybridization is possible in both directions. The hatching success of the hybrids was intermediate between the parental species. Female hybrids showed no clear mate preferences, indicating that back crossing in both directions is possible in principle, but the fertility of the hybrids and the fitness of the F2 generation remain unknown. Our study suggests that hybridization between sympatric species might occur more often than assumed. It might affect the ecology and local distribution of animals in a similar way as competition does.     

Mangrove recruitment after forest disturbance is facilitated by herbaceous species in the Caribbean.

Plant communities along tropical coastlines are often affected by natural and human disturbances, but little is known about factors influencing recovery. We focused on mangrove forests, which are among the most threatened ecosystems globally, to examine how facilitation by herbaceous vegetation might improve forest restoration after disturbance. We specifically investigated whether recovery of mangrove forests in harsh environments is accelerated by nurse plants and whether the beneficial effects are species-specific. Quantification of standardized effects allowed comparisons across performance parameters and over time for: (1) net effect of each herbaceous species on mangrove survival and growth, (2) effects of pre- and post-establishment factors associated with each herbaceous species, and (3) need for artificial planting to enhance growth or survival of mangrove seedlings. Mangrove recruitment in a clear-cut forest in Belize was accelerated by the presence of Sesuvium portulacastrum (succulent forb) and Distichlis spicata (grass), two coastal species common throughout the Caribbean region. The net effect of herbaceous vegetation was positive, but the magnitude of effects on mangrove survival and growth differed by species. Because of differences in their vegetative structure and other features, species effects on mangroves also varied by mechanism: (1) trapping of dispersing propagules (both species), (2) structural support of the seedling (Distichlis), and/or (3) promotion of survival (Sesuviumn) or growth (Distichlis) through amelioration of soil conditions (temperature, aeration). Artificial planting had a stronger positive effect on mangrove survival than did edaphic conditions, but planting enhanced mangrove growth more in Sesuvium than in Distichlis patches. Our study indicates that beneficial species might be selected based on features that provide multiple positive effects and that species comparisons may be improved using standardized effects. Our findings are not only relevant to the coastal environments found in the Caribbean region, but our assessment methods may be useful for developing site-specific information to restore disturbed mangrove forests worldwide, especially given the large pool of mangrove associates (>45 genera) available for screening.     

The influence of size-specific indirect interactions in predator-prey systems

Nonlethal indirect interactions between predators often lead to nonadditive effects of predator number on prey survival and growth. Previous studies have focused on systems with at least two different predator species and one prey species. However, most predators undergo extreme ontological changes in phenotype such that interactions between different-sized cohorts of a predator and its prey could lead to nonadditive effects in systems with only two species. This may be important since different-sized individuals of the same species can differ more in their ecology than similar-sized individuals of different species. This study examined trait-mediated indirect effects in a two-species system including a cannibalistic predator with different-sized cohorts and its prey. I tested for these effects using larvae of two stream salamanders, Gyrinophilus porphyriticus (predator) and Eurycea cirrigera (prey), by altering the densities and combinations of predator size classes in experimental streams. Results showed that the presence of large individuals can significantly reduce the impact of density changes of smaller conspecifics on prey survival through nonlethal means. In the absence of large conspecifics, an increase in the relative frequency of small predators significantly increased predation rates, thereby reducing prey survival. However, with large conspecifics present, increasing the density of small predators did not decrease prey survival, resulting in a 14.3% lower prey mortality than predicted from the independent effects of both predator size classes. Small predators changed their microhabitat use in the presence of larger conspecifics. Prey individuals reduced activity in response to large predators but did not respond to small predators. Both predators reduced prey growth. These results demonstrate that the impact of a predator can be significantly altered by two different types of trait-mediated indirect effects in two-species systems: between different-sized cohorts and between different cohorts and prey. This study demonstrates that predictions based on simple numerical changes that assume independent effects of different size classes or ignore size structure can be strongly misleading. We need to account for the size structure within predator populations in order to predict how changes in predator abundance will affect predator-prey dynamics.     

Do males and females differ in the feeding of large and small siblings? An experiment with the bluethroat

Males and females have been reported to differ in their feeding of large and small siblings in several species of birds. According to recent hypotheses, this phenomenon may be related to a sexual conflict over avian hatching patterns. We designed an experiment to test for the existence of such a sex difference by manipulating nestling size hierarchies of the bluethroat (Luscinia s. svecica) in two directions; half the broods were “asynchronized” to yield large size-differences within broods and the other half were “synchronized” to yield small size-differences. In all broods, nestlings were categorized as being either large or small according to body mass. We recorded male and female food distribution by video early (day 4 after hatching) and late (day 8) in the nestling period. Males and females did not differ in their distribution of food among different-sized nestlings. With large size-differences, both males and females fed large nestlings nearly twice as often as small ones. In contrast, when the size-differences were small, food was more evenly distributed among nestlings. Early in the nestling period, males fed more nestlings during each feeding visit than did females. Our finding that male and female bluethroats do not differ in the feeding of large and small siblings is in contrast to most previous studies. Variation in costs and benefits to males and females from feeding different-sized nestlings, and restrictions to parental choice due to nestling interactions, may explain interspecific variation. Received: 27 June 1997 / Accepted after revision: 26 January 1998     

Can rare positive interactions become common when large carnivores consume livestock?

Livestock populations in protected areas are viewed negatively because of their interaction with native ungulates through direct competition for food resources. However, livestock and native prey can also interact indirectly through their shared predator. Indirect interactions between two prey species occur when one prey modifies either the functional or numerical responses of a shared predator. This interaction is often manifested as negative effects (apparent competition) on one or both prey species through increased predation risk. But indirect interactions can also yield positive effects on a focal prey if the shared predator modifies its functional response toward increased consumption of an abundant and higher-quality alternative prey. Such a phenomenon between two prey species is underappreciated and overlooked in nature. Positive indirect effects can be expected to occur in livestock-dominated wildlife reserves containing large carnivores. We searched for such positive effects in Acacia-Zizhypus forests of India's Gir sanctuary where livestock (Bubalus bubalis and Bos indicus) and a coexisting native prey (chital deer, Axis axis) are consumed by Asiatic lions (Panthera leo persica). Chital vigilance was higher in areas with low livestock density than in areas with high livestock density. This positive indirect effect occurred because lion predation rates on livestock were twice as great where livestock were abundant than where livestock density was low. Positive indirect interactions mediated by shared predators may be more common than generally thought with rather major consequences for ecological understanding and conservation. We encourage further studies to understand outcomes of indirect interactions on long-term predator-prey dynamics in livestock-dominated protected areas.     

Large Herbivores and the Design of Large-Scale Nature Reserves in Western Europe

Nature conservation in densely populated areas, as in western Europe, faces the continuous loss and fragmentation of suitable habitats for wild species. Current conservation measures have not been successful in countering this threat to biological diversity, so a new concept for nature conservation is required. I propose using the habitat requirements of large herbivores as a key to the design of large-scale nature reserves. Large herbivores can have a significant influence on vegetation composition and thus act as keystone species. Moreover, they require large tracts of land and can be considered an umbrella species group for the preservation of other plants and animals. I review the present status of nature reserves in the European Union concerning size distribution and the occurrence of large herbivores and their predators. I argue that habitat fragmentation and the increased management costs of maintaining seminatural areas necessitate an enlargement of existing reserves, a process-oriented management approach, and a minimization of human influence for the long-term preservation of biological diversity.     

Using Ecological Function to Develop Recovery Criteria for Depleted Species: Sea Otters and Kelp Forests in the Aleutian Archipelago

Abstract: Recovery criteria for depleted species or populations normally are based on demographic measures, the goal being to maintain enough individuals over a sufficiently large area to assure a socially tolerable risk of future extinction. Such demographically based recovery criteria may be insufficient to restore the functional roles of strongly interacting species. We explored the idea of developing a recovery criterion for sea otters (Enhydra lutris) in the Aleutian archipelago on the basis of their keystone role in kelp forest ecosystems. We surveyed sea otters and rocky reef habitats at 34 island‐time combinations. The system nearly always existed in either a kelp‐dominated or deforested phase state, which was predictable from sea otter density. We used a resampling analysis of these data to show that the phase state at any particular island can be determined at 95% probability of correct classification with information from as few as six sites. When sea otter population status (and thus the phase state of the kelp forest) was allowed to vary randomly among islands, just 15 islands had to be sampled to estimate the true proportion that were kelp dominated (within 10%) with 90% confidence. We conclude that kelp forest phase state is a more appropriate, sensitive, and cost‐effective measure of sea otter recovery than the more traditional demographically based metrics, and we suggest that similar approaches have broad potential utility in establishing recovery criteria for depleted populations of other functionally important species.     

Vulnerability and Resilience of Tropical Forest Species to Land‐Use Change

Abstract: We provide a cross‐taxon and historical analysis of what makes tropical forest species vulnerable to extinction. Several traits have been important for species survival in the recent and distant geological past, including seed dormancy and vegetative growth in plants, small body size in mammals, and vagility in insects. For major past catastrophes, such as the five mass extinction events, large range size and vagility or dispersal were key to species survival. Traits that make some species more vulnerable to extinction are consistent across time scales. Terrestrial organisms, particularly animals, are more extinction prone than marine organisms. Plants that persist through dramatic changes often reproduce vegetatively and possess mechanisms of die back. Synergistic interactions between current anthropogenic threats, such as logging, fire, hunting, pests and diseases, and climate change are frequent. Rising temperatures threaten all organisms, perhaps particularly tropical organisms adapted to small temperature ranges and isolated by distance from suitable future climates. Mutualist species and trophic specialists may also be more threatened because of such range‐shift gaps. Phylogenetically specialized groups may be collectively more prone to extinction than generalists. Characterization of tropical forest species’ vulnerability to anthropogenic change is constrained by complex interactions among threats and by both taxonomic and ecological impediments, including gross undersampling of biotas and poor understanding of the spatial patterns of taxa at all scales.     

Positive and Negative Effects of a Threatened Parrotfish on Reef Ecosystems

Species that are strong interactors play disproportionately important roles in the dynamics of natural ecosystems. It has been proposed that their presence is necessary for positively shaping the structure and functioning of ecosystems. We evaluated this hypothesis using the case of the world's largest parrotfish (Bolbometopon muricatum), a globally imperiled species. We used direct observation, animal tracking, and computer simulations to examine the diverse routes through which B. muricatum affects the diversity, dispersal, relative abundance, and survival of the corals that comprise the foundation of reef ecosystems. Our results suggest that this species can influence reef building corals in both positive and negative ways. Field observation and simulation outputs indicated that B. muricatum reduced the abundance of macroalgae that can outcompete corals, but they also feed directly on corals, decreasing coral abundance, diversity, and colony size. B. muricatum appeared to facilitate coral advancement by mechanically dispersing coral fragments and opening up bare space for coral settlement, but they also damaged adult corals and remobilized a large volume of potentially stressful carbonate sediment. The impacts this species has on reefs appears to be regulated in part by its abundance—the effects of B. muricatum were more intense in simulation scenarios populated with high densities of these fish. Observations conducted in regions with high and low predator (e.g., sharks) abundance generated results that are consistent with the hypothesis that these predators of B. muricatum may play a role in governing their abundance; thus, predation may modulate the intensity of the effects they have on reef dynamics. Overall our results illustrate that functionally unique and threatened species may not have universally positive impacts on ecosystems and that it may be necessary for environmental managers to consider the diverse effects of such species and the forces that mediate the strength of their influence. Efectos Positivos y Negativos de un Pez Loro Amenazado Sobre Ecosistemas Arrecifales