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Kelly, R. P.

Publications and source records attributed to Kelly, R. P..

At least 19 recordsLinked to original sources

Multi-omic Environmental Monitoring Reveals When Molecular Signals Reflect Living Biology

Environmental molecular monitoring has transformed biodiversity detection, yet most methods capture only species presence through amplified DNA fragments, missing the biological information encoded in native nucleic acids. Here we show that a single shotgun Oxford Nanopore sequencing library from water can be read through several independent biological lenses, and that each class of molecule (environmental RNA, CpG methylation, and bulk DNA) differs in its environmental persistence, which sets the window over which signal can be trusted. Sampling Pacific salmon spawning runs, we simultaneously recovered environmental RNA indicating fresh biological input, CpG methylation profiles reflecting age demographics, and pathogen signatures reflecting health. Environmental RNA tracked spawning phenology, qPCR corroborated the timing of fresh biological input over the weeks sampled, and methylation-based age inference held only while fresh DNA dominated, a condition we term the 'Freshness Gate.' Because environmental water is not a tissue sample, we detect functional transcripts as evidence of biological input but do not infer differential gene expression, and we read methylation state but do not infer biological age outside the Freshness Gate. Variance partitioning is consistent with biological freshness, rather than technical factors such as fragment length or sequencing depth, structuring environmental methylation, though these comparisons did not reach significance at nine time points. Comparable dynamics were observed in Chinook salmon, with timing offsets following their distinct spawning season. By defining when environmental molecules faithfully encode living biology and when degradation decouples signal from current state, our framework sets practical and theoretical boundaries for multi-omic environmental monitoring.

ecology↗

Long-range PCR amplification and nanopore sequencing of 8-10 kb mitochondrial fragments from environmental DNA

O_LILong-read sequencing data can provide increased taxonomic resolution and genomic linkage information that is otherwise impossible to obtain from short-read amplicon or shotgun sequencing. However, the use of long-read sequencing for environmental DNA (eDNA) analysis has thus far been limited by both the apparent rarity of long DNA molecules in eDNA samples and the lack of established bioinformatics workflows for long-read metabarcoding, particularly from mixed template samples. C_LIO_LIHere, we report nanopore sequencing of 8.0 - 9.5 kb mitochondrial fragments obtained from mesocosm and field eDNA samples, amplified with long-range PCR (LR-PCR) using primers designed to preferentially amplify teleost mitogenomes. C_LIO_LIWe recovered half-mitochondria from 13 fish species in field-collected samples (Puget Sound in Washington State, USA), as well as from approximately half of the fish species inhabiting the positive control mesocosm (the Seattle Aquarium). Among biological replicates, we observed consistent detection of read-abundant species, while there was greater stochasticity in the presence of rarer species. C_LIO_LIWe demonstrate that long fragments can be obtained from standard eDNA samples and successfully amplified and sequenced to obtain species identifications despite higher error rates characteristic of nanopore sequencing. We present both laboratory methods and an accessible bioinformatic pipeline for obtaining and analyzing LR-PCR amplified fragments from eDNA, providing a framework for future long-read metabarcoding studies. C_LI

bioinformatics↗

Modeling cetacean eDNA distribution along the Washington coast using metabarcoding from opportunistic samples and generalized additive models

Effective cetacean conservation depends on robust monitoring, yet traditional visual and passive-acoustic surveys have constraints. We evaluated environmental DNA (eDNA) metabarcoding coupled with species distribution modeling (SDM) as a tool to study habitat use of cetaceans along the Washington State coast, USA. Seawater was collected at the surface and at a 50 m depth from 43 sites (86 samples) during the 2019 U.S.-Canada Integrated Ecosystem & Acoustic-Trawl Survey. A partial section of the mitochondrial control region was amplified with cetacean - specific primers, sequenced on an Illumina MiSeq, and taxonomically assigned with a curated reference database. Nine species were detected; we modelled the three most frequent: Pacific white-sided dolphin (Lagenorhynchus obliquidens) (10 detections), humpback whale (Megaptera novaeangliae) (8 detections), and Rissos dolphin (Grampus griseus) (6 detections). Binomial generalized additive models related presence-absence to bathymetry, distance to shore, longitude, slope, and sea-surface temperature; model performance was assessed with stratified five-fold cross-validation. SDMs explained 17-51% of null deviance and presented high specificity ([≥] 0.80). The Pacific white-sided dolphin showed the highest eDNA presence probabilities offshore, beyond the shelf break. Humpback whale eDNA presence probabilities showed hotspots along the shelf break with secondary high-probability patches in near-shore waters. Rissos dolphin eDNA presence probabilities were elevated in offshore zones characterized by steep bathymetric gradients, particularly northwest of the sampled transect. These spatial patterns are consistent with historical visual-acoustic records, suggesting that eDNA-informed SDMs can capture cetacean habitat use. This proof of concept indicates that combining eDNA detections with flexible SDMs could provide a cost-effective, non-invasive complement to conventional surveys and may offer a scalable pathway for marine-mammal monitoring and spatial planning.

genomics↗

Quantitative, Multispecies Monitoring at a Continental Scale

Molecular data from environmental samples can reflect the abundance of species DNA, an index immediately relevant to natural-resources management at a broad geographic scale. These data commonly derive from assays designed and targeted for individual species (e.g., using qPCR or ddPCR), or from metabarcoding that use more general PCR primers to amplify many species simultaneously. Multispecies analyses make efficient use of field samples and laboratory time, and speak to inherently multispecies questions of management and ecological interest. However, unlike single-species techniques, metabarcoding alone generally reflects only the environmental DNA (eDNA) proportions of target-species DNA present, not their absolute quantities. Here, we combine qPCR and metabarcoding data derived from the same samples to estimate quantities of eDNA from many fish species along the US West Coast in three dimensions, demonstrating a technique of practical relevance for both management and ecology. We derive spatially explicit maps of eDNA abundances for 12 common species of ecological and management importance and point the way to quantitative surveys of wild species using molecules alone. We find that species distribution maps derived from eDNA largely mirror known species niches and known spatial distributions. Notably, our analyses indentify biodiversity hotspots that align with previously documented regions of ecological significance, such as the Columbia River plume and the Heceta Bank.

ecology↗

Estimating Organism Abundance Using Within-Sample Haplotype Frequencies of eDNA Metabarcoding Data

Environmental DNA (eDNA) metabarcoding provides powerful insights into species presence and community composition, but remains limited in its ability to quantify species abundance or structure. Here, we show that deviation between observed haplotype frequencies within a given sample and the population haplotype frequencies can be used to infer the number of individual contributors to an eDNA sample. We also lay out the theory for how population haplotype frequencies can be approximated from eDNA data alone, enabling broad applicability even in the absence of tissue-based references. We then present an estimator to derive the number of individual contributors to a given eDNA sample and validate its performance using simulations with variable allele frequencies and noise. Our framework demonstrates that differences between expected and observed frequencies carry meaningful biological information in eDNA data. Our results show that the number of contributors can be recovered under a range of conditions, particularly with hypervariable markers and sufficient sampling. This approach complements existing molecular methods and opens a new avenue for inferring abundance from eDNA metabarcoding datasets.

molecular biology↗

Vertebrate Biodiversity via eDNA at the air-water interface

Although aquatic, aerial, and terrestrial habitats are often treated as separate ecological systems, these environments exist along a continuum of connectivity: flows of biomass and energy routinely create linkages across water and air, blurring traditional boundaries. Here, we use eDNA sequencing to illustrate and quantify the movement of trace genetic information between water and air. We collected 27 paired air-and-water samples from two urban-wildland interface sites using passive air sampling and active water filtering. Metabarcoding with the MiFish-U 12S marker recovered 35 vertebrate taxa, 40% of which were detected in both water and air, ranging from the strictly aquatic salmon to wholly terrestrial cottontail rabbit. This reciprocal relationship suggests that eDNA pools form within each partition (water or air), with the probability of transfer governed by DNA concentration. In this view, detecting aquatic eDNA in the air is not contamination or stochastic noise but an expected, repeatable phenomenon. Logistic models confirm that higher abundance in one medium predicts spillover into the other. For instance, peaks in Coho and Chinook salmon eDNA align within 24 hours, demonstrating that passive air sampling reflects the temporal abundance trends of the most common aquatic species. In contrast, low-read-abundance taxa appear only sporadically, implying that rare detections are the first to drop out of cross-medium transfer and therefore demand intensified sampling in their primary habitat. Together, our findings bridge the conventional separation of aquatic and airborne eDNA and establish a unified, non-invasive framework for holistic vertebrate monitoring at the land-water interface. This approach offers transformative potential for conservation, invasive-species early warning, and One Health surveillance.

molecular biology↗

Integrating eDNA and acoustic-trawl data to provide small pelagic biomass estimates for fisheries assessment

Accurate abundance estimates of fisheries resources are essential for sustainable fisheries management. In response to the growing need for developing more accurate and cost-effective biomass estimation methods, the analysis of environmental DNA (eDNA) has recently emerged as an alternative for fish abundance quantification. However, practical approaches for integrating eDNA data into fisheries assessment remain limited. Here, we introduce a Bayesian joint model that combines acoustic-trawl and eDNA data to estimate fish biomass. Utilizing 209 water eDNA samples and 196 acoustic transects, the model was applied to estimate the distribution and abundance of the European anchovy (Engraulis encrasicolus) in the Bay of Biscay. The joint model produced estimates consistent with known spatial patterns of anchovy, with eDNA data suggesting a broader distribution and potentially higher abundance. This research demonstrates the value of incorporating eDNA data as a complement to acoustic-trawl for stock assessment and illustrates the versatility of joint Bayesian models and their potential application to various species and datasets. Ultimately, our work opens new avenues for more holistic fisheries assessment, underscoring the growing role of eDNA in that context.

genetics↗

Fish from the sky: Airborne eDNA tracks aquatic life

Water and air are generally treated as separate reservoirs of environmental DNA (eDNA) derived from the species resident in those respective environments. However, it is likely that eDNA routinely crosses the air-water boundary in both directions as a result of deposition, evaporation, or other processes. Here, we systematically tested methods of sampling eDNA at the air-water interface, showing for the first time that aquatic life can be reliably detected from passive air samples collected nearby. We deployed four simple air samplers -- three different kinds of filters and one open tray of deionized water -- alongside paired water samples and visual counts over a six-week peak run of Coho salmon (Oncorhynchus kisutch) at a local spawning stream. We then quantified eDNA concentrations in both air and water (air: copies/day/cm2; water: copies/L) using quantitative PCR, to estimate (1) the concentration of target eDNA in air vs. water, and (2) the capture performance of each filter type. Despite an approximate 25,000-times dilution versus water, passive air collectors captured quantitative airborne eDNA signals that closely paralleled salmon counts, although recovery varied with sampler design and orientation. We show the air-water interface is a quantifiable source of aquatic genetic information using simple, passive samplers that do not require electricity, making them appealing for biomonitoring in remote or resource-limited settings. This work points the way to using airborne eDNA as a robust pathway for biological information critical to conservation, resource management, and public-health protection.

molecular biology↗

Differential Decay of Multiple eNA Components from a Cetacean

Environmental nucleic acids (eNA), such as DNA and RNA, are powerful tools for monitoring biodiversity. Yet, interpreting eNA detections requires understanding of their environmental persistence. We conducted a decay experiment using seawater from an open enclosure to track degradation of six eNA components derived from Tursiops truncatus: mitochondrial eDNA of varying lengths, ribosomal eRNA, and messenger eRNA. Targets were quantified over seven days via digital droplet PCR (ddPCR). Decay followed a biphasic exponential model with rapid initial loss ([~]24 hours at 15 {degrees}C), followed by slower degradation. Cytb messenger eRNA was least stable, disappearing within four hours. Ribosomal eRNA persisted longer but degraded slightly faster than its eDNA counterpart ({lambda} = 0.236 vs. 0.165 hr-{superscript 1}). Longest eDNA fragments decayed more rapidly ({lambda} = 0.190 hr-1) than shorter ones ({lambda} = 0.114 hr-1). These findings support the use of eDNA fragment length as a proxy for degradation state and reinforce that combining multiple eNA components with distinct stabilities can potentially provide a molecular clock for inferring eNA age. This approach improves the spatiotemporal resolution of eNA-based monitoring, particularly for rare marine-mammal that act as point sources. We also emphasize the importance of explicitly distinguishing between RNA types (ribosomal vs. messenger) in environmental studies, given their divergent stability and interpretability.

molecular biology↗

Fast, Flexible, Feasible: A Transparent Framework for Evaluating eDNA Workflow Trade-offs in Resource-Limited Settings

Environmental DNA (eDNA) analysis enables biodiversity monitoring by detecting organisms from trace genetic material, but high reagent costs, cold-chain logistics, and computational demands limit its broader use, particularly in resource-limited settings. To address these challenges and improve accessibility, we systematically evaluated multiple workflow components, including four DNA extraction methods, two primer sets, three Nanopore basecalling models, and two demultiplexing pipelines. Across 144 workflow combinations tested in an aquarium with 15 fish species, we mapped trade-offs between cost, sensitivity, and processing speed using a hierarchical Bayesian model, to assess where time and resource savings are possible without compromising detection. Workflows using the Qiagen Blood and Tissue (BT) extraction kit provided the highest sensitivity, recovering all 15 species of the sampled fish community within 3-5 hours of Oxford Nanopore sequencing when paired with MiFish-U primer set and high-accuracy (HAC) basecalling. Chelex, an alternative lower-cost extraction method, required extended sequencing (>24 hours) to reach comparable species-detection rates. DirectPCR and QuickExtract offered field-friendly extraction alternatives that achieved comparable recovery in [~]10-12 hours, though their cost-effectiveness varied. While the MarVer1 primer was designed to broaden vertebrate detection, it recovered the same fish species as MiFish-U, though with fewer total reads. Real-time sequencing trials (0-61 hours) revealed that high-efficiency workflows (BT + HAC) reached detection plateaus rapidly, indicating sequencing time can be reduced without sacrificing accuracy. The OBITools4 bioinformatics pipeline enabled automated demultiplexing but discarded more reads than an alternative, ONTbarcoder2.3, which retained low-abundance taxa at the cost of manual curation. Rather than identifying a single "best" workflow, this study provides a transparent decision framework for prioritizing cost, speed, and sensitivity in eDNA applications, supporting scalable, cost-effective eDNA monitoring, in resource-limited settings.

molecular biology↗

Invasive European green crab (Carcinus maenas) predation revealed with quantitative DNA metabarcoding

Predation by invasive species can threaten local ecosystems and economies. The European green crab (Carcinus maenas), one of the most widespread marine invasive species, is an effective predator associated with clam and crab population declines outside of its native range. In the U.S. Pacific Northwest, green crab have recently experienced increases in abundance and expanding distributions, generating concern for estuarine ecosystems and associated aquaculture production. However, regionally-specific information on the trophic impacts of invasive green crab is highly limited. We compared the stomach contents of green crabs collected on aquaculture beds versus natural intertidal sloughs in Willapa Bay, Washington, to provide the first in-depth description of European green crab diet at a particularly crucial time for regional management. We first identified putative prey items using DNA metabarcoding. For key prey species, we also applied a quantitative model to calculate the true abundance of each species in stomach content DNA samples. From the stomach contents of 62 green crabs, we identified 56 unique taxa belonging to nine phyla. The stomach contents of crabs collected from aquaculture beds were significantly different from the stomach contents of crabs collected at natural intertidal sloughs. Across all sites, arthropods were the most frequently detected prey, with the native hairy shore crab (Hemigrapsus oregonensis) the single most common prey item. Of the eight species included in the quantitative model, two ecologically-important native species - the sand shrimp (Crangon franciscorum) and the Pacific staghorn sculpin (Leptocottus armatus) - were the most abundant in crab stomach contents, when present. In addition to providing timely information on green crab diet, our research demonstrates the novel application of a recently developed model for quantitative DNA metabarcoding. This represents another step in the ongoing evolution of DNA-based diet analysis towards producing the quantitative data necessary for modeling invasive species impacts.

ecology↗

Environmental DNA reveals patterns of biological invasion in an inland sea

Non-native species have the potential to cause ecological and economic harm to coastal and estuarine ecosystems. Understanding which habitat types are most vulnerable to biological invasions, where invasions originate, and the vectors by which they arrive can help direct limited resources to prevent or mitigate ecological and socio-economic harm. Information about the occurrence of non-native species can help guide interventions at all stages of invasion, from first introduction, to naturalization and invasion. However, monitoring at relevant scales requires considerable investment of time, resources, and taxonomic expertise. Environmental DNA (eDNA) metabarcoding methods sample coastal ecosystems at broad spatial and temporal scales to augment established monitoring methods. We use COI mtDNA eDNA sampling to survey a diverse assemblage of species across distinct habitats in the Salish Sea in Washington State, USA, and classify each as non-native, native, or indeterminate in origin. The non-native species detected include both well-documented invaders and species not previously reported within the Salish Sea. We find a non-native assemblage dominated by shellfish and algae with native ranges in the temperate western Pacific, and find more-retentive estuarine habitats to be invaded at far higher levels than better-flushed rocky shores. Furthermore, we find an increase in invasion level with higher water temperatures in spring and summer across habitat types. This analysis contributes to a growing understanding of the biotic and abiotic factors that influence invasion level, and underscores the utility of eDNA surveys to monitor biological invasions and to better understand the factors that drive these invasions.

ecology↗

Distinguishing Signal from Noise: Understanding Patterns of Non-Detections to Inform Accurate Quantitative Metabarcoding

Correcting for amplification biases in genetic metabarcoding data can yield quantitative estimates of template DNA concentrations. However, a major source of uncertainty in metabarcoding data is the presence of non-detections, where a technical PCR replicate fails to detect a species observed in other replicates. Such non-detections are an important special case of variability among technical replicates in metabarcoding data, particularly in environmental samples. While many sampling and amplification processes underlie observed variation in metabarcoding data, understanding the causes of non-detections is an important step in distinguishing signal from noise in metabarcoding studies. Here, we use both simulated and empirical data to 1) develop a qualitative understanding of how non-detections arise in metabarcoding data, 2) outline steps to recognize uninformative data in practice, and 3) identify the conditions under which amplicon sequence data can reliably detect underlying biological signals. We show in both simulations and empirical data that, for a given species, the rate of non-detections among technical replicates is a function of both the template DNA concentration and species-specific amplification efficiency. Consequently, we conclude metabarcoding datasets are strongly affected by (1) deterministic amplification biases during PCR and (2) stochastic sampling of amplicons during sequencing -- both of which we can model -- but also by (3) stochastic sampling of rare molecules prior to PCR, which remains a frontier for quantitative metabarcoding. Our results highlight the importance of estimating species-specific amplification efficiencies and critically evaluating patterns of non-detection in metabarcoding datasets to better distinguish environmental signal from the noise inherent in molecular detections of rare targets.

molecular biology↗

Message in a Bottle: Archived DNA Reveals Marine Heatwave-Associated Shifts in Fish Assemblages

Marine heatwaves can drive large-scale shifts in marine ecosystems but studying their impacts on whole species assemblages can be difficult. Here, we leverage the taxonomic breadth and resolution of DNA sequences derived from environmental DNA (eDNA) in the ethanol of a set of 23-year longitudinal ichthyoplankton samples, combining these with microscopy-derived ichthyoplankton identification to yield higher-resolution, species-specific quantitative abundance estimates of fish assemblages in the California Current Large Marine Ecosystem during and after the 2014-16 Pacific marine heatwave. This integrated dataset reveals patterns of tropicalization with increases in southern, mesopelagic species and associated declines in important temperate fisheries targets (e.g., North Pacific Hake (Merluccius productus) and Pacific Sardine (Sardinops sagax)). We observed novel assemblages of southern, mesopelagic fishes and temperate species (e.g., Northern Anchovy, Engraulis mordax) even after the return to average water temperatures. Our innovative preservative derived eDNA metabarcoding and quantitative modeling approaches open the door to reconstructing the historical dynamics of assemblages from modern and archived samples worldwide. SummaryNovel quantitative abundance estimates from archived DNA reveals marine heatwave-associated shifts in fish assemblages.

ecology↗

Toward Quantitative Metabarcoding

Amplicon-sequence data from environmental DNA (eDNA) and microbiome studies provides important information for ecology, conservation, management, and health. At present, amplicon-sequencing studies - known also as metabarcoding studies, in which the primary data consist of targeted, amplified fragments of DNA sequenced from many taxa in a mixture - struggle to link genetic observations to underlying biology in a quantitative way, but many applications require quantitative information about the taxa or systems under scrutiny. As metabarcoding studies proliferate in ecology following decades of microbial and microbiome work using similar techniques, it becomes more important to develop ways ot make them quantitative to ensure that their conclusions are adequately supported. Here we link previously disparate sets of techniques for making such data quantitative, showing that the underlying PCR mechanism explains observed patterns of amplicon data in a general way. By modeling the process through which amplicon-sequence data arises, rather than transforming the data post-hoc, we show how to estimate the starting DNA proportions from a mixture of many taxa. We illustrate how to calibrate the model using mock communities and apply the approach to simulated data and a series of empirical examples. Our approach opens the door to improve the use of metabarcoding data in a wide range of applications in ecology, public health, and related fields.

ecology↗

A visual tour of carbon export by sinking particles

To better quantify the oceans biological carbon pump, we resolved the diversity of sinking particles that transport carbon into the oceans interior, their contribution to carbon export, and their attenuation with depth. Sinking particles collected in sediment trap gel layers from 4 distinct ocean ecosystems were imaged, measured, and classified. The size and identity of particles was used to model their contribution to particulate organic carbon (POC) flux. Measured POC fluxes were reasonably predicted by particle images. Nine particle types were identified, and most of the compositional variability was driven by the relative contribution of aggregates, long cylindrical fecal pellets, and salp fecal pellets. While particle composition varied across locations and seasons, the entire range of compositions was measured at a single well-observed location in the subarctic North Pacific over 1 month, across 500 m of depth. The magnitude of POC flux was not consistently associated with a dominant particle class, but particle classes did influence flux attenuation. Long fecal pellets attenuated most rapidly with depth whereas certain other classes attenuated little or not at all with depth. Small particles (<100 m) consistently contributed [~]5% to total POC flux in samples with higher magnitude fluxes. The relative importance of these small particle classes (spherical mini pellets, short oval fecal pellets, and dense detritus) increased in low flux environments (up to 46% of total POC flux). Imaging approaches that resolve large variations in particle composition across ocean basins, depth, and time will help to better parameterize biological carbon pump models.

ecology↗

Environmental DNA Metabarcoding Reveals Winners and Losers of Global Change in Coastal Waters

2Studies of the ecological effects of global change often focus on one or few species at a time. Consequently, we know relatively little about the changes underway at real-world scales of biological communities, which typically have hundreds or thousands of interacting species. Here, we use monthly samples of environmental DNA to survey 222 planktonic taxa along a gradient of temperature, salinity, dissolved oxygen, and carbonate chemistry in nearshore marine habitat. The result is a high-resolution picture of changes in ecological communities using a technique replicable across a wide variety of ecosystems. We estimate community-level differences associated with time, space and environmental variables, and use these results to forecast near-term community changes due to warming and ocean acidification. We find distinct communities in warmer and more acidified conditions, with overall reduced richness in diatom assemblages and increased richness in dinoflagellates. Individual taxa finding greater suitable habitat in near-future waters are more taxonomically varied and include the ubiquitous coccolithophore Emiliania huxleyi and the harmful dinoflagellate Alexandrium sp. These results suggest foundational changes for nearshore food webs under near-future conditions.

ecology↗

Environmental DNA Metabarcoding for Simultaneous Monitoring and Ecological Assessment of Many Harmful Algae

Harmful algae can have profound economic, environmental, and social consequences. As the timing, frequency, and severity of harmful algal blooms (HABs) change alongside global climate, efficient tools to monitor and understand the current ecological context of these taxa are increasingly important. Here we employ environmental DNA metabarcoding to identify patterns in a wide variety of harmful algae and associated ecological communities in the Hood Canal of Puget Sound in Washington State, USA. We track trends of presence and abundance in a series of water samples across nearly two years. We find putative harmful algal sequences in a majority of samples, suggesting that these groups are routinely present in local waters. We report patterns in variants of the economically important genus Pseudo-nitzschia (family Bacillariaceae), as well as multiple harmful algal taxa previously unknown or poorly documented in the region, including a cold-water variant from the saxitoxin-producing genus Alexandrium (family Gonyaulacaceae), two variants from the karlotoxin-producing genus Karlodinium (family Kareniaceae), and one variant from the parasitic genus Hematodinium (family Syndiniaceae). We then use data on environmental variables and the biological community surrounding each algal taxon to illustrate the ecological context in which these species are commonly found. Environmental DNA metabarcoding thus simultaneously (1) alerts us to potential new or cryptic occurrences of harmful algae, (2) expands our knowledge of the co-occurring conditions and species associated with the growth of these organisms in changing marine environments, and (3) provides a tool for monitoring and management moving forward.

ecology↗