Potential trophic biomasses and trace-substance concentrations in unstructured marine food webs

It is proposed that unstructured food webs may more closely resemble real marine food webs than does the conventional, structured model. An unstructured food-web model leads to a set of very simple expressions for the potential partition of matter in the food web in steady state, including the potential fluxes of material and biomasses of trophic types and the concentration of trace substances in the members and materials of such a food web. The approach may explain some anomalies of relative predator-prey biomasses and of trace-element distribution, and may be of further use for analyzing and predicting (a) the tropho-dynamic parameters of marine systems, (b) the trophic positions, and the steadystate fluxes and biomasses of marine organisms, (c) the distribution of trace materials in marine biota; and for relating findings among these areas. Other matters, such as limitations of food conversion, indicated by concentration factors of trace substances, the possibilities of non-causal association of anomalously high levels of trace substances (including pollutants) with diseased or otherwise abnormal marine creatures, and an inverse relationship of early concentrations of newly introduced trace substances and their eventual concentrations in various organisms, are also developed in this approach.     

Trophic structure of coastal Antarctic food webs associated with changes in sea ice and food supply

Predicting the dynamics of ecosystems requires an understanding of how trophic interactions respond to environmental change. In Antarctic marine ecosystems, food web dynamics are inextricably linked to sea ice conditions that affect the nature and magnitude of primary food sources available to higher trophic levels. Recent attention on the changing sea ice conditions in polar seas highlights the need to better understand how marine food webs respond to changes in such broad-scale environmental drivers. This study investigated the importance of sea ice and advected primary food sources to the structure of benthic food webs in coastal Antarctica. We compared the isotopic composition of several seafloor taxa (including primary producers and invertebrates with a variety of feeding modes) that are widely distributed in the Antarctic. We assessed shifts in the trophic role of numerically dominant benthic omnivores at five coastal Ross Sea locations. These locations vary in primary productivity and food availability, due to their different levels of sea ice cover, and proximity to polynyas and advected primary production. The delta15N signatures and isotope mixing model results for the bivalves Laternula elliptica and Adamussium colbecki and the urchin Sterechinus neumeyeri indicate a shift from consumption of a higher proportion of detritus at locations with more permanent sea ice in the south to more freshly produced algal material associated with proximity to ice-free water in the north and east. The detrital pathways utilized by many benthic species may act to dampen the impacts of large seasonal fluctuations in the availability of primary production. The limiting relationship between sea ice distribution and in situ primary productivity emphasizes the role of connectivity and spatial subsidies of organic matter in fueling the food web. Our results begin to provide a basis for predicting how benthic ecosystems will respond to changes in sea ice persistence and extent along environmental gradients in the high Antarctic.     

Origin of compartmentalization in food webs

The response of an ecosystem to perturbations is mediated by both antagonistic and facilitative interactions between species. It is thought that a community's resilience depends crucially on the food web--the network of trophic interactions--and on the food web's degree of compartmentalization. Despite its ecological importance, compartmentalization and the mechanisms that give rise to it remain poorly understood. Here we investigate several definitions of compartments, propose ways to understand the ecological meaning of these definitions, and quantify the degree of compartmentalization of empirical food webs. We find that the compartmentalization observed in empirical food webs can be accounted for solely by the niche organization of species and their diets. By uncovering connections between compartmentalization and species' diet contiguity, our findings help us understand which perturbations can result in fragmentation of the food web and which can lead to catastrophic effects. Additionally, we show that the composition of compartments can be used to address the long-standing question of what determines the ecological niche of a species.     

Topological structure and interaction strengths in model food webs

We report the results of carrying out a large number of simulations on a coevolutionary model of multispecies communities. A wide range of parameter values were investigated which allowed a rather complete picture of the change in behaviour of the model as these parameters were varied to be built up. Our main interest was in the nature of the community food webs constructed via the simulations. We identify the range of parameter values which give rise to realistic food webs and give arguments which allow some of the structure which is found to be understood in an intuitive way. Since the webs are evolved according to the rules of the model, the strengths of the predator–prey links are not determined a priori, and emerge from the process of constructing the web. We measure the distribution of these link strengths, and find that there are a large number of weak links, in agreement with recent suggestions. We also review some of the data on food webs available in the literature, and make some tentative comparisons with our results. The difficulties of making such comparisons and the possible future developments of the model are also briefly discussed.     

Multiple anthropogenic stressors and the structural properties of food webs

Coastal environments are among the most productive on the planet, providing a wide range of ecosystem services. Development and exploitation mean that they are faced with stresses from a number of anthropogenic sources. Such stresses are typically studied in isolation, but multiple stressors can combine in unexpected ways to alter the structure of ecological systems. Here, we experimentally explore the impacts of inorganic nutrients and organic matter on a range of food web properties. We find that these two stressors combine additively to produce significant increases in connectance and mean food chain length. Such increases are typically associated with enhanced robustness to secondary extinctions and productivity, respectively. Despite these apparent beneficial effects, we find a simplification of web structure in terms of taxon richness and diversity, and altered proportions of basal and top species. These effects are driven by a reduction in community assembly and lower consistency in a range of system properties as a result of the multiple stressors. Consequently, impacted food webs are likely to be more vulnerable to human- or climate-induced perturbations in the long-term.     

Chemically mediated competition between microbes and animals: microbes as consumers in food webs

Microbes are known to affect ecosystems and communities as decomposers, pathogens, and mutualists. However, they also may function as classic consumers and competitors with animals if they chemically deter larger consumers from using rich food-falls such as carrion, fruits, and seeds that can represent critical windfalls to both microbes and animals. Microbes often use chemicals (i.e., antibiotics) to compete against other microbes. Thus using chemicals against larger competitors might be expected and could redirect significant energy subsidies from upper trophic levels to the detrital pathway. When we baited traps in a coastal marine ecosystem with fresh vs. microbe-laden fish carrion, fresh carrion attracted 2.6 times as many animals per trap as microbe-laden carrion. This resulted from fresh carrion being found more frequently and from attracting more animals when found. Microbe-laden carrion was four times more likely to be uncolonized by large consumers than was fresh carrion. In the lab, the most common animal found in our traps (the stone crab Menippe mercenaria) ate fresh carrion 2.4 times more frequently than microbe-laden carrion. Bacteria-removal experiments and feeding bioassays using organic extracts of microbe-laden carrion showed that bacteria produced noxious chemicals that deterred animal consumers. Thus bacteria compete with large animal scavengers by rendering carcasses chemically repugnant. Because food-fall resources such as carrion are major food subsidies in many ecosystems, chemically mediated competition between microbes and animals could be an important, common, but underappreciated interaction within many communities.     

Arsenic trophodynamics along the food chains/webs of different ecosystems: a review

In drinking water, arsenic (As) food chain accumulation may pose a risk to human health. An attempt was made to synthesise the published information concerning As trophic transfer along the food chain/web of various marine, freshwater, and terrestrial ecosystems. Some investigations included As speciation. Further objectives were to outline the factors potentially influencing As trophodynamics and to understand the consequence of As trophic transfer in the environment.     

Feeding by larvae of intertidal invertebrates: assessing their position in pelagic food webs

One of the leading determinants of the structure and dynamics of marine populations is the rate of arrival of new individuals to local sites. While physical transport processes play major roles in delivering larvae to the shore, these processes become most important after larvae have survived the perils of life in the plankton, where they usually suffer great mortality. The lack of information regarding larval feeding makes it difficult to assess the effects of food supply on larval survival, or the role larvae may play in nearshore food webs. Here, we examine the spectrum of food sizes and food types consumed by the larvae of two intertidal barnacle species and of the predatory gastropod Concholepas concholepas. We conducted replicated experiments in which larvae were exposed to the food size spectrum (phytoplankton, microprotozoan and autotrophic picoplankton) found in nearshore waters in central Chile. Results show that barnacle nauplii and gastropod veligers are omnivorous grazers, incorporating significant fractions of heterotrophs in their diets. In accordance with their feeding mechanisms and body size, barnacle nauplii were able to feed on autotrophic picoplankton (<5 microm) and did not consume the largest phytoplankton cells, which made the bulk of phytoplankton biomass in spring-summer blooms. Balanoid nauplii exhibited higher ingestion rates than the smaller-bodied chthamaloid larvae. Newly hatched C. concholepas larvae also consumed picoplankton cells, while competent larvae of this species ingested mostly the largest phytoplankton cells and heterotrophic protozoans. Results suggest that persistent changes in the structure of pelagic food webs can have important effects on the species-specific food availability for invertebrate larvae, which can result in large-scale differences in recruitment rates of a given species, and in the relative recruitment success of the different species that make up benthic communities.     

Sources of primary production supporting food webs in an arid coastal embayment

Penaeid shrimp can be useful ecological indicators of linkages between shallow tropical coastal habitats, acting as integrators of carbon and nitrogen sources due to their generalist feeding habits and their mobility between habitats and with tidal cycles. In the current study, the contribution of mangrove, seagrass and microbial mat to the nutrition of two penaeid shrimp species, Penaeus semisulcatus and Metapeneus ensis, in a shallow arid embayment in the Arabian Gulf was assessed through a combination of analysis of stomach contents and dual carbon and nitrogen stable isotope signatures. Shrimp tissue stable isotope signatures identified seagrass as a major source of carbon and nitrogen for both species, contributing 21–38?% (1–99?‰). Microbial mat was also detected as a significant nutritional source for early-stage Metapenaeus affinis postlarvae (1–27?%). However, mangroves were not identified as a significant source, with the range of results including the possibility of a zero contribution. Moreover, the greatest possible contribution of mangroves as source of carbon was less than for the other primary producers. This may be due the high salinity and wide temperature range limiting mangrove productivity as well as the low export of dissolved and particulate organic material out of the mangroves due to low rainfall.     

Native fish diversity alters the effects of an invasive species on food webs

Aquatic communities have been altered by invasive species, with impacts on native biodiversity and ecosystem function. At the same time, native biodiversity may mitigate the effects of an invader. Common carp (Cyprinus carpio) is a ubiquitous, invasive fish species that strongly influences community and ecosystem processes. We used common carp to test whether the potential effects of an invasive species are altered across a range of species diversity in native communities. In mesocosms, treatments of zero, one, three, and six native fish species were used to represent the nested subset patterns observed in fish communities of lakes in Illinois, USA. The effect of the invader was tested across fish richness treatments by adding common carp to the native community and substituting native biomass with common carp. Native species and intraspecific effects reduced invader growth. The invader reduced native fish growth; however, the negative effect was minimized with increasing native richness. The zooplankton grazer community was modified by a top-down effect from the invader that increased the amount of phytoplankton. Neither the invader nor richness treatments influenced total phosphorus or community metabolism. Overall, the invader reduced resources for native species; and the effect scaled with how the invader was incorporated into the community. Higher native diversity mitigated the impact of the invader, confirming the need to consider biodiversity when predicting the impacts of invasive species.     

Trophic ecology of large predatory reef fishes: energy pathways, trophic level, and implications for fisheries in a changing climate

Large predatory fishes are disproportionately targeted by reef fisheries, but little is known about their trophic ecology, which inhibits understanding of community dynamics and the potential effects of climate change. In this study, stable isotope analyses were used to infer trophic ecology of a guild of large predatory fishes that are targeted by fisheries on the Great Barrier Reef, Australia. Each of four focal predators (Plectropomus leopardus, Plectropomus maculatus, Lethrinus miniatus and Lutjanus carponotatus) was found to have a distinct isotopic signature in terms of δ¹³C and δ¹⁵N. A two-source mixing model (benthic reef-based versus pelagic) indicated that P. leopardus and L. miniatus derive the majority (72 and 62 %, respectively) of their production from planktonic sources, while P. maculatus and L. carponotatus derive the majority (89 and 74 %, respectively) of their production from benthic reef-based sources. This indicates that planktonic production is important for sustaining key species in reef fisheries and highlights the need for a whole-ecosystem approach to fisheries management. Unexpectedly, there was little isotopic niche overlap between three of four focal predators, suggesting that inter-specific competition for prey may be low or absent. δ¹⁵Nitrogen indicated that the closely related P. leopardus and P. maculatus are apex predators (trophic level > 4), while δ¹³C indicated that each species has a different diet and degree of trophic specialisation. In view of these divergent trophic ecologies, each of the four focal predators (and the associated fisheries) are anticipated to be differentially affected by climate-induced disturbances. Thus, the results presented herein provide a useful starting point for precautionary management of exploited predator populations in a changing climate.     

The probabilistic niche model reveals substantial variation in the niche structure of empirical food webs

The structure of food webs, complex networks of interspecies feeding interactions, plays a crucial role in ecosystem resilience and function, and understanding food web structure remains a central problem in ecology. Previous studies have shown that key features of empirical food webs can be reproduced by low-dimensional "niche" models. Here we examine the form and variability of food web niche structure by fitting a probabilistic niche model to 37 empirical food webs, a much larger number of food webs than used in previous studies. The model relaxes previous assumptions about parameter distributions and hierarchy and returns parameter estimates for each species in each web. The model significantly outperforms previous niche model variants and also performs well for several webs where a body-size-based niche model performs poorly, implying that traits other than body size are important in structuring these webs' niche space. Parameter estimates frequently violate previous models' assumptions: in 19 of 37 webs, parameter values are not significantly hierarchical, 32 of 37 webs have nonuniform niche value distributions, and 15 of 37 webs lack a correlation between niche width and niche position. Extending the model to a two-dimensional niche space yields networks with a mixture of one- and two-dimensional niches and provides a significantly better fit for webs with a large number of species and links. These results confirm that food webs are strongly niche-structured but reveal substantial variation in the form of the niche structuring, a result with fundamental implications for ecosystem resilience and function.     

The interaction of cannibalism and omnivory: consequences for community dynamics

Although cannibalism is ubiquitous in food webs and frequent in systems where a predator and its prey also share a common resource (intraguild predation, IGP), its impacts on species interactions and the dynamics and structure of communities are still poorly understood. In addition, the few existing studies on cannibalism have generally focused on cannibalism in the top-predator, ignoring that it is frequent at intermediate trophic levels. A set of structured models shows that cannibalism can completely alter the dynamics and structure of three-species IGP systems depending on the trophic position where cannibalism occurs. Contrary to the expectations of simple models, the IG predator can exploit the resources more efficiently when it is cannibalistic, enabling the predator to persist at lower resource densities than the IG prey. Cannibalism in the IG predator can also alter the effect of enrichment, preventing predator-mediated extinction of the IG prey at high productivities predicted by simple models. Cannibalism in the IG prey can reverse the effect of top-down cascades, leading to an increase in the resource with decreasing IG predator density. These predictions are consistent with current data. Overall, cannibalism promotes the coexistence of the IG predator and IG prey. These results indicate that including cannibalism in current models can overcome the discrepancy between theory and empirical data. Thus, we need to measure and account for cannibalistic interactions to reliably predict the structure and dynamics of communities.     

Meta-analysis: trophic level, habitat, and productivity shape the food web effects of resource subsidies

Studies of the effects of cross-habitat resource subsidies have been a feature of food web ecology over the past decade. To date, most studies have focused on demonstrating the magnitude of a subsidy or documenting its effect in the recipient habitat. Ecologists have yet to develop a satisfactory framework for predicting the magnitude of these effects. We used 115 data sets from 32 studies to compare consumer responses to resource subsidies across recipient habitat type, trophic level, and functional group. Changes in consumer density or biomass in response to subsidies were inconsistent across habitats, trophic, and functional groups. Responses in stream cobble bar and coastline habitats were larger than in other habitats. Contrary to expectation, the magnitude of consumer response was not affected by recipient habitat productivity or the ratio of productivity between donor and recipient habitats. However, consumer response was significantly related to the ratio of subsidy resources to equivalent resources in the recipient habitat. Broad contrasts in productivity are modified by subsidy type, vector, and the physical and biotic characteristics of both donor and recipient habitats. For this reason, the ratio of subsidy to equivalent resources is a more useful tool for predicting the possible effect of a subsidy than coarser contrasts of in situ productivity. The commonness of subsidy effects suggests that many ecosystems need to be studied as open systems.     

Trophic upgrading of food quality by protozoans enhancing copepod growth: role of essential lipids

 Protozoa are known for their intermediary trophic role in transferring organic matter from small size planktonic particles to mesozooplankton. This study concentrates on the possible addition of biochemical value during this transfer, by new production of compounds that are essential in copepod food. In laboratory experiments, copepods could not be raised on a diet of the chlorophycean Dunaliella sp., though they readily consumed this alga. Dunaliella sp. contained all essential amino acids, but was deficient in highly unsaturated fatty acids and in sterols. In contrast to copepods, the heterotrophic dinoflagellate Oxyrrhis marina grew well on Dunaliella sp., producing significant amounts of the long-chain fatty acids docosahexaenoic acid and eicosapentaenoic acid, in addition to cholesterol and brassicasterol. Using this O. marina grown on Dunaliella sp. to feed Temora longicornis and Pseudocalanus elongatus, both copepod species rapidly developed from young nauplius larvae to maturity on the dinoflagellate diet. Hence, in this experimental food-chain the inadequate chlorophycean food was biochemically upgraded by the protozoan to high-quality copepod food. The results indicate that highly unsaturated fatty acids and/or sterols are essential compounds, which can be produced by protozoans. Due to their intermediate size, the mechanism of trophic upgrading by protozoans may bridge the gap of essential nutrients between the microbial loop and higher trophic levels. Received: 11 January 1999 / Accepted: 3 June 1999     

Divergent composition but similar function of soil food webs of individual plants: plant species and community effects

Soils are extremely rich in biodiversity, and soil organisms play pivotal roles in supporting terrestrial life, but the role that individual plants and plant communities play in influencing the diversity and functioning of soil food webs remains highly debated. Plants, as primary producers and providers of resources to the soil food web, are of vital importance for the composition, structure, and functioning of soil communities. However, whether natural soil food webs that are completely open to immigration and emigration differ underneath individual plants remains unknown. In a biodiversity restoration experiment we first compared the soil nematode communities of 228 individual plants belonging to eight herbaceous species. We included grass, leguminous, and non-leguminous species. Each individual plant grew intermingled with other species, but all plant species had a different nematode community. Moreover, nematode communities were more similar when plant individuals were growing in the same as compared to different plant communities, and these effects were most apparent for the groups of bacterivorous, carnivorous, and omnivorous nematodes. Subsequently, we analyzed the composition, structure, and functioning of the complete soil food webs of 58 individual plants, belonging to two of the plant species, Lotus corniculatus (Fabaceae) and Plantago lanceolata (Plantaginaceae). We isolated and identified more than 150 taxa/groups of soil organisms. The soil community composition and structure of the entire food webs were influenced both by the species identity of the plant individual and the surrounding plant community. Unexpectedly, plant identity had the strongest effects on decomposing soil organisms, widely believed to be generalist feeders. In contrast, quantitative food web modeling showed that the composition of the plant community influenced nitrogen mineralization under individual plants, but that plant species identity did not affect nitrogen or carbon mineralization or food web stability. Hence, the composition and structure of entire soil food webs vary at the scale of individual plants and are strongly influenced by the species identity of the plant. However, the ecosystem functions these food webs provide are determined by the identity of the entire plant community.     

Carbon partitioning among the first trophic levels in the North Western Adriatic Basin

In the frame of PRISMA II Project samples for plankton analyses were carried out during four cruises (June, 1996; February and June, 1997; February, 1998) in order to assess the relative importance in term of biomasses of the three main size fractions (pico- <2 r µm, nano- 2-20 r µm and micro-plankton >20 r µm). Spatial and temporal distribution of the three plankton fractions were described as abundance and contribution to the total carbon content in an area between Po River mouth and Rimini. The relative contribution of picoplankton resulted higher in the offshore zone, while that of nanoplankton in the inshore waters. In February 1998 microphytoplankton, mainly constituted by diatoms, was very abundant in the inshore waters. Micro-zooplankton was always very scarce. Cluster analyses performed on these data grouped the stations on the basis of their community structure, and agreed with the hydrological features. Small size classes contributed more significantly to the total plankton carbon content in most of the situations. Microplankton fraction contribution was relevant only during spring diatom bloom of February 1998 and with a less extent in the confined coastal summer blooms.     

Mixed messages across multiple trophic levels: the ecology of bark beetle chemical communication systems

Summary. Chemical, physiological and behavioral components of pheromone communication have been described for a number of bark beetle species, yet our understanding of how these signals function under natural conditions remains relatively limited. Development of ecologically based models is complicated by the multiple functions and sources of variability inherent in bark beetle semiochemistry. This discussion addresses four ecological issues of chemical signaling in bark beetles: the effects of aggregation on individual fitness, the possibility of cheating, how plants can defend themselves against herbivores that employ aggregation pheromones, and the implications of variability in chemical communication systems to predator avoidance. An analysis of published data from thirteen scolytid – conifer systems indicates that the net benefit and optimal colonization density vary with host condition and beetle species. When beetles attack live trees, the benefit of cooperative host procurement exceeds losses due to competition for the limited substrate, at least up to moderate densities. When beetles colonize dead tissue, however, the effect of subsequently arriving beetles on initial colonizers is almost entirely negative. This suggests that aggregation originated as exploitation of senders, but evolved into manipulation of receivers. It is also proposed that the optimal colonization density which typifies each species or population may offer a more objective and less value–laden index of behavior than current labels such as “aggressiveness”. Beetles can maximize the relative benefits of group attack by incorporating instantaneous measures of host resistance into their colonization behavior, and by adjusting oviposition with colonization density. This system may provide opportunities for cheating. However a number of factors may select against a fixed strategy of cheating, including the linkage between tree allelochemistry and beetle semiochemistry, the reduced quality of substrate available to late arrivers, the short adult lifespans of most bark beetles, differential exposure to some predators, the difficulty of locating signalers during extensive endemic periods, and the low costs incurred during host assessment. However, the possibility that beetles employ flexible, density – dependent strategies deserves heightened attention. The ability of bark beetles to collectively exhaust host defenses poses a particular problem for plant defense. It is argued here that the ideal defense should include both direct resistance mechanisms against invading beetles, and indirect mechanisms that inhibit chemical communication. Evidence for the latter mechanism is explored. The ability of predators to efficiently exploit aggregation pheromones as kairomones in prey finding poses significant risk to bark beetles. It is proposed that minor alterations in pheromone components may provide colonizers with partial escape from such natural enemies while maintaining intraspecific functionality. Traditional interpretations emphasized the fidelity and consistency of pheromones, but under natural conditions chemical signals are modified by unpredictable features of the biotic and abiotic environment. Although we typically view variation in pheromonal signals as experimental noise or simple deviations from a population norm, such variation may reflect evolutionary dynamics. Complex ecological interactions may impose trade-offs between the clarity versus diversity of their signals. Received 3 July 2000; accepted 8 January 2001     

Predator diversity and identity drive interaction strength and trophic cascades in a food web

Declining predator diversity may drastically affect the biomass and productivity of herbivores and plants. Understanding how changes in predator diversity can propagate through food webs to alter ecosystem function is one of the most challenging ecological research topics today. We studied the effects of predator removal in a simple natural food web in the Sierra Nevada mountains of California (USA). By excluding the predators of the third trophic level of a food web in a full-factorial design, we monitored cascading effects of varying predator diversity and composition on the herbivorous beetle Chrysomela aeneicollis and the willow Salix orestera, which compose the first and second trophic levels of the food web. Decreasing predator diversity increased herbivore biomass and survivorship, and consequently increased the amount of plant biomass consumed via a trophic cascade. Despite this simple linear mean effect of diversity on the strength of the trophic cascade, we found additivity, compensation, and interference in the effects of multiple predators on herbivores and plants. Herbivore survivorship and predator-prey interaction strengths varied with predator diversity, predator identity, and the identity of coexisting predators. Additive effects of predators on herbivores and plants may have been driven by temporal niche separation, whereas compensatory effects and interference occurred among predators with a similar phenology. Together, these results suggest that while the general trends of diversity effects may appear linear and additive, other information about species identity was required to predict the effects of removing individual predators. In a community that is not temporally well-mixed, predator traits such as phenology may help predict impacts of species loss on other species. Information about predator natural history and food web structure may help explain variation in predator diversity effects on trophic cascades and ecosystem function.     

Matter accumulation and energy expenditure in planktonic ecosystems at different trophic levels

Functional peculiarities of pelagic communities from temperate and tropical zones of the ocean have been investigated in terms of food-web interrelations and balance sheets of matter and energy of the major populations and ecological groupings. Ranging from temperate or epiplankton ecosystems to tropical oceanic or deepwater ones, as well as from upwelling zones to stable oceanic oligotrophic regions, the following regular changes in the communities' main functional indices have been established: (1) enlargement of the food spectrum, omnivorousness and predatory activity; (2) reduction of rations and rates of organic matter accumulation in the lower heterotrophic levels with simultaneous increase of energy expenditure; (3) increased trophic complexity and stability of communities. Epiplanktonic systems of low stability proved to be richer and commercially more profitable.