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An ecological explanation for hyperallometric scaling of reproduction

In wild populations, large individuals have disproportionately higher reproductive output than smaller individuals. Some theoretical models explain this pattern - termed reproductive hyperallometry - by individuals allocating a greater fraction of available energy towards reproductive effort as they grow. Here, we propose an ecological explanation for this observation: differences between individuals in rates of resource assimilation, where greater assimilation causes both increased reproduction and body size, resulting in reproductive hyperallometry at the level of the population. We illustrate this effect by determining the relationship between size and reproduction in wild and lab-reared Trinidadian guppies. We show that (i) reproduction increased disproportionately with body size in the wild but not in the lab, where resource competition was eliminated and food availability restricted; (ii) in the wild, hyperallometry was greatest during the wet season, when resource competition is strongest; and (iii) detection of hyperallometric scaling of reproduction at the population level was inevitable if individual differences in assimilation were ignored. We propose that ecologically-driven variation in assimilation - caused by size-dependent resource competition, niche expansion, and chance - contributes substantially to hyperallometric scaling of reproduction in natural populations. We recommend that mechanistic models incorporate such ecologically-caused variation when seeking to explain reproductive hyperallometry.

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Roads as ecological traps for giant anteaters

Wildlife-vehicle collisions (WVC) represent a serious source of mortality for many species, threatening local populations persistence while also carrying a high economic and human safety cost. Animals may adapt their behaviour to road associated threats, but roadside resources can act as attractants, providing misleading signals about the quality of roadside habitats, ultimately acting as an ecological trap. Yet, the extent to which individuals modify their behaviour and space use to roads is largely unknown for most taxa. Using fine-scale movement data from 41 giant anteaters (Myrmecophaga tridactyla), tracked in the Brazilian Cerrado, we aimed to identify facets of movement behaviour that might exhibit plasticity to roads and traffic volume. Specifically, the analysis of daily and instantaneous movement speeds, home range characteristics, and crossing rates/times, allowed us to test for an effect of road proximity, traffic volume and natural linear features on movement behaviour. We found no effect of road proximity or traffic volume on space use or movement behaviour. While individuals tended to reduce their movement speed when approaching roads and crossed roads ~3 times less than would have been expected by random chance, none of the highways we monitored were impervious. The majority of tracked anteaters living near roads (<2km) crossed them, with higher crossing rates for males. Consequently, habitat near roads may function as an ecological trap where healthy individuals occupy the territories nearby or bisected by roads but eventually are road-killed given their regular crossings, leaving the territory vacant for subsequent occupation. Crucially, we found no evidence that anteaters actively searched for passage structures to cross the roads. This suggests that crossing structures alone are unlikely to mitigate WVC induced mortality. Our research reinforces the need to implement fencing, properly linked to existing passages, and minimising the amount of night-time driving to reduce the number of WVCs. Article Impact StatementAnalyses of giant anteater movement show how roads act as ecological traps, reinforcing the need for fencing and reduced night-time driving.

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Living Shorelines Achieve Functional Equivalence to Natural Fringe Marshes across Multiple Ecological Metrics

Nature-based features provide a welcome class of adaptations to promote ecological resilience in the face of climate change. Along coastlines, living shorelines are among the preferred adaptation strategies to both reduce erosion and provide ecological functions. As an alternative to shoreline armoring, living shorelines are viewed favorably among coastal managers, wetlands boards, and some private property owners, but they have yet to undergo a thorough examination of how their levels of ecosystem functions compare to their closest natural counterpart: fringing marshes. Here, we provide a synthesis of results from a multi-year, large-spatial-scale study in which we compared numerous ecological metrics measured in thirteen pairs of living shorelines and natural fringing marshes throughout coastal Virginia, USA. Overall, we found that living shorelines were functionally equivalent to natural marshes in nearly all measured aspects, except for a lag in sediment composition. These data support the prioritization of living shorelines as a coastal adaptation strategy.

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Metabarcoding and ecological interaction networks for selecting candidate biological control agents

O_LIClassical biological control can be used to decrease the density of invasive species to below an acceptable ecological and economic threshold. Natural enemies specific to the invasive species are selected from its native range and released into the invaded range. This approach has drawbacks, despite the performance of specificity tests to ensure its safety, because the fundamental host range defined under controlled conditions does not represent the actual host range in natura, and these tests omit indirect interactions within community. C_LIO_LIWe focus on Sonchus oleraceus (Asteraceae), a weed species originating from Western Palearctic that is invasive worldwide and notably in Australia. We explore how analyses of interaction network within its native range can be used to 1) inventory herbivores associated to the target plant, 2) characterize their ecological host ranges, and 3) guide the selection of candidate biocontrol agents considering interactions with species from higher trophic levels. Arthropods were collected from plant community sympatric to S. oleraceus, in three bioclimatic regions, and interactions were inferred by a combination of molecular and morphological approaches. C_LIO_LIThe networks reconstructed were structured in several trophic levels from basal species (diversified plant community), to intermediate and top species (herbivorous arthropods and their natural enemies). The subnetwork centered on S. oleraceus related interactions contained 116 taxa and 213 interactions. We identified 47 herbivores feeding on S. oleraceus, 15 of which were specific to the target species (i.e. Generality value equal to 1). Some discrepancies with respect to published findings or conventional specificity tests suggested possible insufficient sampling effort for the recording of interactions or the existence of cryptic species. Among potential candidate agents, 6 exhibited interactions with natural enemies. C_LIO_LISynthesis and applications: Adopting a network approach as prerequisite step of the CBC program can provide a rapid screening of potential agents to be tested in priority. Once ecological host range defined, we suggest that priority should be given to agent predated by a minimum species, and, when they exist, to an agent that possesses enemies from the most distant taxonomical group from those occurring in the range of introduction. C_LI

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Ecological network analysis of watershed meta-ecosystems: A new perspective on quantifying the integrated watershed ecosystem

A watershed is an integrated ecosystem. In different disciplines, a watershed has been described as a geomorphic unit, a hydrological unit, an ecological unit, or a socio-economic unit and has been quantitatively described using different indicator systems. Until now, no general indicator system has existed that could quantitatively describe the geomorphic features, hydrologic features, ecological features and socio-economic features of an integrated watershed ecosystem (IWE) simultaneously. Here, we proposed a quantitative descriptive framework for an IWE (QDFIWE). This QDFIWE involved three steps: (1) constructing a watershed meta-ecosystem (WME) based on the hierarchical system of the watershed; (2) constructing flow networks based on the WME; and (3) identifying the holistic properties (such as spatial throughput, spatial organization and spatial resilience) of the WME through analyzing its flow networks based on ecological network analysis (ENA). Then, we applied this method to study the geomorphic topological structure, geomorphic spatial structure, natural water resource system and integrated water resource system of the Yangtze River basin. The results suggested that based on the QDFIWE, (1) one could construct different WMEs and corresponding flow networks for different requirements; (2) one could obtain time series of the holistic properties of an IWE to analyze its evolution; (3) one could compare, classify and cluster any number of IWEs through identifying their holistic properties according to similar strategies; and (4) one could determine or create more indicators, which could provide additional information, based on the holistic properties of an IWE. This study demonstrates that the QDFIWE is a general method of quantifying the holistic properties of all subsystems of an IWE simultaneously. Thus, the method can improve the understanding of the IWE.

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Harnessing the power of metabarcoding in the ecological interpretation of plant-pollinator DNA data: strategies and consequences of reads filtering

DNA metabarcoding approaches to analyse complex mixtures of pollen has become the standard in pollination biology, especially in the light of recent threats affecting pollination. In spite of the increasing adoption of High Throughput Sequencing (HTS) approaches, these studies generate huge numbers of raw reads, some of which might be associated to false positives or infrequently recorded species with potentially little biological information. If these reads are not discarded (i.e. pruned), they can lead to changes in the ecological findings and lead to biased conclusions. In this study we reviewed 42 papers in the recent pollen DNA metabarcoding literature and focused on the type of pruning applied. We also tested whether the different types of those cut off threshold may leave a mark on the DNA metabarcoding data. To do so, we compared for the first time community composition, species richness and networks of species interactions (i.e. Connectace, Modularity, Connectivity and Shannon entropy) associated with the most relevant ways of treating HTS outputs: no cut (no reads filtering), or cutting levels obtained as proportional 1% of sample total reads, or as fixed amount of 100 reads, or from ROC (Receiver operator characteristic). Results clearly indicated that pruning type shapes species composition and that to apply or not a threshold dramatically impacts ecological indices, potentially increasing the risk of misinterpreting DNA metabarcoding data under an ecological point of view. Given the high methodological heterogeneity from the revised literature, we discuss in what conditions filtering types may be more appropriate, and also recommend to biologically justify the pruning threshold when analysing DNA metabarcoding raw reads, and to develop shared approaches to make future studies more comparable.

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easyFulcrum: An R package to process and analyze ecological sampling data generated using the Fulcrum mobile application

Large-scale ecological sampling can be difficult and costly, especially for organisms that are too small to be easily identified in a natural environment by eye. Typically, these microscopic floral and fauna are sampled by collecting substrates from nature and then separating organisms from substrates in the laboratory. In many cases, diverse organisms can be identified to the species-level using molecular barcodes. To facilitate large-scale ecological sampling of microscopic organisms, we used a geographic data-collection platform for mobile devices called Fulcrum that streamlines the organization of geospatial sampling data, substrate photographs, and environmental data at natural sampling sites. These sampling data are then linked to organism isolation data from the laboratory. Here, we describe the easyFulcrum R package, which can be used to clean, process, and visualize ecological field sampling and isolation data exported from the Fulcrum mobile application. We developed this package for wild nematode sampling, but it is extensible to other organisms. The advantages of using Fulcrum combined with easyFulcrum are (1) the elimination of transcription errors by replacing manual data entry and/or spreadsheets with a mobile application, (2) the ability to clean, process, and visualize sampling data using a standardized set of functions in the R software environment, and (3) the ability to join disparate data to each other, including environmental data from the field and the molecularly defined identities of individual specimens isolated from samples.

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Untangling the ecological signal in the dental morphology in the bat superfamily Noctilionoidea

Diet has been linked to the diversification of the bat superfamily Noctilionoidea, a group that underwent an impressive ecological adaptive radiation within Mammalia. For decades, studies have explored morphological adaptations and diversity of noctilionoid bats to reveal traits associated with their ecological diversity. Surprisingly, despite such interest and recent application of novel techniques, ecomorphological studies have failed to fully resolve the link between diet and a critical component of the feeding apparatus: dental morphology. Using multivariate dental topographic analysis and phylogenetic comparative methods, we examined the phylogenetic, biological and ecological signal in the dental morphology of noctilionoid bats. Analysing the lower first molars of 110 species, we explored relationships between diet and dental morphology, accounting for three different dimensions of diet (guild, composition and breadth). Phylogenetic and size-dependent structuring of the dental topography data shows it does not correlate only to diet, highlighting the need to account for multiple sources of variation. Frugivorous noctilionoids have sharper molars than other previously reported frugivorous mammals. Nectarivorous noctilionoids showed reduced lower molar crown height and steepness, whereas animalivorous species had larger molars. Dietary composition suggested that the intensity of exploitation of a resource is also linked to different dimensions of dental morphology. Increasing carnivory positively correlated with MA, explaining the highest proportion of its variation, and increasing frugivory explained the highest proportion of variation in all other variables. Dietary breadth showed generalist species have sharper, more topographically-complex molars, whereas specialist herbivores and specialist animalivores fell at opposite ends in the range of tooth steepness and crown height. Together, the results suggest that adaptations affecting different attributes of dental morphology likely facilitated the dietary diversity and specialisation found in Noctilionoidea.

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Ecological dynamics of the gut microbiome in response to dietary fiber

Dietary fibers are generally thought to benefit intestinal health. Their impacts on the composition and metabolic function of the gut microbiome, however, vary greatly across individuals. Previous research showed that each individuals response to fibers depends on their baseline gut microbiome, but the ecology driving microbiota remodeling during fiber intake remained unclear. Here, we studied the long-term dynamics of gut microbiome and short-chain fatty acids (SCFAs) in isogenic mice with distinct microbiota baselines fed with the fermentable fiber inulin compared to the non-fermentable fiber cellulose. We found that inulin produced generally rapid response followed by gradual stabilization to new equilibria, and those dynamics were baseline-dependent. We parameterized an ecology model from the timeseries data, which revealed a group of bacteria whose growth significantly increases in response to inulin. and whose baseline abundance and interspecies competition explains the baseline-dependence of microbiome density and community composition dynamics. Fecal levels of of SCFAs, such as propionate, is associated with the abundance of inulin responders, yet inter-individual variation of gut microbiome impedes the prediction of SCFAs by machine learning models. Finally, we showed that our methods and major findings are generalizable to dietary resistant starch. This study emphasizes the importance of ecological modeling to understand microbiome responses to dietary changes and the need for personalized interventions.

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Monitoring photogenic ecological phenomena: Social network site images reveal spatiotemporal phases of Japanese cherry blooms

Some ecological phenomena are visually engaging and widely celebrated. Consequently, these have the potential to generate large footprints in the online and social media image records which may be valuable for ecological research. Cherry tree blooms are one such event, especially in Japan where they are a cultural symbol (Sakura, ). For centuries, the Japanese have celebrated Hanami (flower viewing) and the historical data record of the festival allows for phenological studies over this period, one application of which is climate reconstruction. Here we analyse Flickr social network site data in an analogous way to reveal the cherry blossoms seasonal sweep from southern to northern Japan over a twelve-week period. Our method analyses data filtered using geographical constraints, multi-stage text-tag classification, and machine vision, to assess image content for relevance to our research question and use it to estimate historic cherry bloom times. We validated our estimated bloom times against official data, demonstrating the accuracy of the approach. We also investigated an out of season Autumn blooming that has gained worldwide media attention. Despite the complexity of human photographic and social media activity and the relatively small scale of this event, our method can reveal that this bloom has in fact been occurring over a decade. The approach we propose in our case study enables quick and effective monitoring of the photogenic spatiotemporal aspects of our rapidly changing world. It has the potential to be applied broadly to many ecological phenomena of widespread interest.

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The creation of ghost forests is driven by physical, ecological, and disturbance factors but remains a rare phenomenon

Marine transgression and the landward migration of coastlines in response to sea-level rise will determine the future of coastal ecosystems and species worldwide, yet no complete model of these processes exist. Ghost forests, areas where coastal forest has been killed by saltwater inundation, are an attention-grabbing early indicator of marine transgression. Research on marine transgression to date has largely been limited to place-based studies that, due to logistical constraints and foci of expertise, investigate subsets of the potential drivers of marine transgression. Here I take advantage of new open source datasets to provide the first systematic analysis of marine transgression across large scales, focusing on the creation of ghost forests across a region of ecological and scientific importance, the northeast U.S. My analysis provides the first synthesis of the physical, ecological, and disturbance factors that influence marine transgression. It also provides crucial regional context for more intensive studies that have a limited geographic scope. I found that ghost forests are a rare phenomenon in the landscape context: 95% of recent forest loss is concentrated in less than 3.86% of marshes, and between 2000 and 2018, only 0.88% of the entire forested area of the transgression zone experienced loss. As a result of this rarity, regional variation in marine transgression is largely driven by opportunities for rare events, which are more numerous when suitable conditions, such as shallow slopes, cover extensive areas. Quantifying recent trends in the rate of forest loss found little evidence of acceleration, with evidence instead suggesting that fewer ghost forests are being created. I also found that physical, ecological, and disturbance factors, including hurricane impacts, were all important for understanding recent trends in forest loss, suggesting that an interdisciplinary approach is warranted for future analyses and modeling of marine transgression. Such interdisciplinary research is urgently needed, as the current rate of marine transgression points to the likelihood of near-term losses of coastal wetlands, with dire implications for the species that depend on them.

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A multifaceted ecological assessment reveals the invasion of the freshwater red macroalga Montagnia macrospora (Batrachospermales, Rhodophyta) in Taiwan

Invasive freshwater macroalgae are rarely described. Recently, a freshwater red alga, Montagnia macrospora, was introduced from South America to East Asia via the global aquarium trade. The earliest occurrence record of this alga in Taiwan is dated 2005. To determine whether M. macrospora has become an invasive species in Taiwan and to understand its traits that facilitated its invasion, we took a total-evidence approach that combines examination of ecological background and population genetic analysis. Our island-wide survey showed that M. macrospora alga was widespread in the field across Taiwan, where the climate greatly differs from that of South America. Our population genetic analysis revealed that the cox2-3 sequences of all the specimens of M. macrospora from Taiwan were identical, consistent with the hypothesis that the alga expanded through asexual reproduction. Moreover, during our long-term ecological assessments and field surveys, we observed that M. macrospora is an ecological generalist that can self-sustain for a decade and bloom. Taken together, our data suggest that M. macrospora has successfully invaded the freshwater ecosystems in Taiwan due to its ability to disperse asexually and to grow under broad environmental conditions. We hope that our study brings attention to invasive freshwater algae, which have been overlooked in conservation planning and management.

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When are bacteria really gazelles? Comparing patchy ecologies with dimensionless numbers

From micro to planetary scales, spatial heterogeneity--patchiness--is ubiquitous in ecological systems, defining the environments in which organisms move and interact. While this fact has been recognized for decades, most large-scale ecosystem models still use spatially averaged "mean fields" to represent natural populations, while fine-scale, spatially explicit models are mostly restricted to particular organisms or systems. In a conceptual paper, Grunbaum (2012, Interface Focus 2: 150-155) introduced a heuristic framework, based on three dimensionless ratios quantifying movement, reproduction, and resource consumption, to characterize patchy ecological interactions and identify when mean-field assumptions are justifiable. In this paper, we calculated Grunbaums dimensionless numbers for 33 real interactions between consumers and their resource patches in terrestrial, aquatic, and aerial environments. Consumers ranged in size from bacteria to blue whales, and patches lasted from minutes to millennia, spanning spatial scales of mm to hundreds of km. We found that none of the interactions could be accurately represented by a purely mean-field model, though 26 of them (79%) could be partially simplified by averaging out movement, reproductive, or consumption dynamics. Clustering consumer-resource pairs by their non-dimensional ratios revealed several unexpected dynamic similarities between disparate interactions. For example, bacterial Pseudoalteromonas exploit nutrient plumes in a similar manner to Mongolian gazelles grazing on ephemeral patches of steppe vegetation. Our findings suggest that dimensional analysis is a valuable tool for characterizing ecological patchiness, and can link the dynamics of widely different systems into a single quantitative framework.

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Movement ecology of vulnerable lowland tapirs across a gradient of human disturbance

Animal movement is a key ecological process that is tightly coupled to local environmental conditions. While agriculture, urbanisation, and transportation infrastructure are critical to human socio-economic improvement, these have spurred substantial changes in animal movement across the globe with potential impacts on fitness and survival. Notably, however, human disturbance can have differential effects across species, and responses to human activities are thus largely taxa and context specific. As human disturbance is only expected to worsen over the next decade it is critical to better understand how species respond to human disturbance in order to develop effective, case-specific conservation strategies. Here, we use an extensive telemetry dataset collected over 22 years to fill a critical knowledge gap in the movement ecology of lowland tapirs (Tapirus terrestris) across a gradient of human disturbance within three biomes in southern Brazil: the Pantanal, Cerrado, and Atlantic Forest. From these data we found that the mean home range size across all monitored tapirs was 8.31 km2 (95% CI: 6.53 - 10.42), with no evidence that home range sizes differed between sexes nor age groups. Interestingly, although the Atlantic Forest, Cerrado, and Pantanal vary substantially in habitat composition, levels of human disturbance, and tapir population densities, we found that lowland tapir movement behaviour and space use were consistent across all three biomes. Human disturbance also had no detectable effect on lowland tapir movement. Lowland tapirs living in the most altered habitats we monitored exhibited movement behaviour that was comparable to that of tapirs living in a near pristine environment. Contrary to our expectations, we observed very little individual variability in lowland tapir space use and movement, and human impacts on the landscape also had no measurable effect on their movement. Lowland tapir movement behaviour thus appears to exhibit very little phenotypic plasticity. Crucially, the lack of any detectable response to anthropogenic disturbance suggests that human modified habitats risk being ecological traps for tapirs and this information should be factored into conservation actions and species management aimed towards protecting lowland tapir populations.

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Multi-marker metabarcoding resolves subtle variations in freshwater condition: Bioindicators, ecological traits, and trophic interactions

Freshwater systems are experiencing rapid biodiversity losses resulting from high rates of habitat degradation. Ecological condition is typically determined through identifying either macroinvertebrate or diatom bioindicator assemblages and comparing them to their known tolerance to stressors. These comparisons are typically conducted at family or genus levels depending on the availability of taxonomic keys and expertise for focal groups. The objective of this study was to test whether a more taxonomically comprehensive assessment of communities in benthic samples can provide a different perspective of ecological conditions. DNA metabarcoding was used to identify macroinvertebrates and diatoms from kick-net samples collected from sites with different habitat status. Sites with good condition were associated with higher beta diversity as well as slightly higher directed connectance and modularity indicating higher resilience compared with fair condition sites. Indicator value and correlation analyses used DNA metabarcoding data to detect 29 site condition indicator species consistent with known bioindicators and expected relative tolerances. DNA metabarcoding and trophic network analysis also recovered 11 keystone taxa. This study demonstrates the importance of taxonomic breadth across trophic levels for generating biotic data to study ecosystem status, with the potential to scale-up ecological assessments of freshwater condition, trophic stability, and resilience.

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Niche difference determines coexistence and similar underlying processes in four ecological groups

Understanding the drivers of species coexistence is an important objective in ecology. Yet, the multitude of methods to study coexistence hampers cross-community comparisons. Here, we standardized niche and fitness differences (i.e how species limit themselves compared to others and their competitive ability, respectively) across 1018 species pairs to investigate species coexistence across ecological groups and methodological settings (experimental setup, natural co-occurrence, population model used, and growth method). We find that, first, coexistence is driven by large niche differences, not by small fitness differences. Second, species group into clear clusters of coexisting and non-coexisting species along the niche axis. Finally, these clusters are not driven by ecological or methodological settings. This suggests differences between coexisting and non-coexisting communities transcending those measured in our empirical systems. Overall, our results show that species coexistence is mainly influenced by mechanisms acting on niche differences.

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Ecological interactions: Patterns of host utilization by tropical butterflies

Structural complexity of ecological networks facilitate the functional robustness of natural ecosystems. Threatened by the human actions such as habitat destruction and climate change, species may be more or less prone to ecological perturbations depending on the nature of their interactions. We examined the host network of tropical butterflies from the Indian region to see their level of interconnectedness. We manually curated larval host utilization data for 1053 butterflies of India. About 98.8% of species that occur pan-India and 90.6% of species exclusive to the Western Ghats had known hosts whereas it was only 25.9% for species exclusive to north-east India. There were 2589 unique butterfly-host interactions comprising 519 butterfly species and their 1091 known hosts. However, nearly 30% of the species had only single hosts. The Fabaceae and Poaceae were the key host families that accounted for 32.8% of the interactions. There were clear host preferences and monocots hosted disproportionately more butterfly species and interactions. Vanessa cardui had at least 39 known hosts while Ochlandra travancorica supported 19 butterfly species. There were 2693 species-pairs and 4226 interactions among 469 butterflies due to shared hosts. Many butterfly species that have relatively few/unique hosts might be vulnerable in the context of habitat destruction and climate change. This work has great relevance to the ecology and conservation of butterflies in India.

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Consumer-driven nutrient recycling of freshwater decapods: linking ecological theories and application in integrated multi-trophic aquaculture

AbstractThe Metabolic Theory of Ecology (MET) and the Ecological Stoichiometry Theory (EST) are central and complementary in the consumer-driven recycling conceptual basis. The comprehension of physiological processes of organisms at different levels of organizations is essential to explore and predict nutrient recycling behavior in different scenarios, and to design integrated productive systems that efficiently use the nutrient inputs through an adjusted mass balance. We fed with fish-feed three species of decapods from different families and with aquacultural potential to explore the animal-mediated nutrient dynamic and its applicability in productive systems. We tested whether physiological (body mass, body elemental content), ecological (diet), taxonomic and experimental (time of incubation) variables predicts N and P excretion rates and ratios across and within taxa. We also analysed body mass and body elemental content independently as predictors of N and P excretion of decapods across, among and within taxa. Finally, we verified if body content scales allometrically across and within taxa and if differed among taxa. Body mass and taxonomic identity predicted nutrient excretion rates both across and within taxa. When physiological variables were analysed independently, body size best predicted nutrient mineralization in both scales of analyses. Regarding body elemental content, only body P content scaled negatively with body mass across taxa. Results showed higher N-requirements and lower C:N of prawns than anomurans and crabs. The role of crustaceans as nutrient recyclers depends mainly on the species and body mass, and should be considered to select complementary species that efficiently use feed resources. Prawns need more protein in their feed and might be integrated with fish of higher N-requirements, while crabs and anomurans, with fish of lower N-requirements. Our study contributed to the background of MTE and EST through empirical data obtained from decapods and provided useful information to achieve more efficient aquaculture integration systems.

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