Percent cover and density measurements of sessile and mobile intertidal species from an Annual Colonization Experiment (ACE) in the rocky intertidal zone, Oregon Coast from 2011 to 2023

Website: https://www.bco-dmo.org/dataset/990939
Data Type: Other Field Results, experimental
Version: 1
Version Date: 2026-05-15

Project
» RAPID: Testing the rocky intertidal community consequences of the decimation of purple sea star populations along the Oregon coast by sea star wasting disease (Sea star wasting)
» Collaborative Research: Mechanisms of resistance and resilience to system-wide loss of a keystone predator in an iconic intertidal community (Keystone Species Loss)
» LTREB: Testing tipping points in a model rocky intertidal meta-ecosystem – Climate-change, increasing variances, and response mechanisms (LTREB Intertidal Tipping Points)

Programs
» Partnership for Interdisciplinary Studies of Coastal Oceans (PISCO)
» Partnership for Interdisciplinary Studies of Coastal Oceans (PISCO)
» Long Term Research in Environmental Biology (LTREB)
ContributorsAffiliationRole
Menge, Bruce A.Oregon State University (OSU)Principal Investigator
Gravem, SarahOregon State University (OSU)Co-Principal Investigator
York, Amber D.Woods Hole Oceanographic Institution (WHOI BCO-DMO)BCO-DMO Data Manager

Abstract
This dataset contains percent cover and density measurements of species identified from photographs during the experiment described below. Experiment/Study abstract: We tested the effects of predation, competition and facilitation on the annual colonization of rocky intertidal invertebrates from 2011-2023 at 6 sites along the Oregon coast. Mid zone factorial experiments testing the interactive effects of predation (primarily by whelks), competition, and facilitation on colonizers of annually cleared plots in the mid zone at core sites (ACE = Annual Colonization Experiment). The long-term goal was to determine if whelk-prey interactions varied with environmental change. Effects (interaction strengths) of each interaction were determined at the end of each experiment (fall of each year) by calculating the difference between cage controls and predator exclusion cages (predation effect, presumably by whelks in this mid-intertidal zone), exclusion cages with mussels removed after settlement (competition effect, primarily of mussels on barnacles), and exclusion cages with barnacles removed (facilitation effect of barnacles on mussels). ACE is now in its 14th year, and results through 2023 show that strengths of each of these interaction types have varied in space and time. The relative importance of interactions on prey abundance varied among types; facilitation by barnacles on mussels was the strongest effect, followed by predation effect of whelks on sessile invertebrates, with weak effects of competition between sessile invertebrates. Barnacle and mussel colonization rate was quantified using replicated marked plots that are cleared each spring prior to initiation of heavy settlement and then photographed in fall (October-December) after abatement of settlement and growth. 


Coverage

Location: Rocky Intertidal along the Oregon Coast. See accompanying site information in data
Spatial Extent: N:44.83863831 E:-124.05875 S:42.717772 W:-124.56512
Temporal Extent: 2011-06-01 - 2023-11-29

Dataset Description

This time series was funded by the Long Term Research in Environmental Biology [LTREB] program including the LTREB project listed on this page (Award DEB-2050017) and prior awards (Award DEB-1050694,DEB-1554702).

ACE = Annual Colonization Experiment


Methods & Sampling

Species interaction experiments called ACE (Annual Colonization Experiment) include six treatments to test the main and interactive effects of predation, competition and disturbance on annual colonization of rocky substrates in the mid-zone of the rocky intertidal along the Oregon Coast (see site list attached). The experiment began in 2011 and is repeated annually. All plots are 15 x 15cm and fixed,  and the experiment is repeated in the same location each year.  Each spring (usually April or May), all plots are cleared of all biomass using scrapers and oven cleaner to reset the community. The exception is the control plots  (CO) which were started in 2021 and are nearby plots that have never been intentionally disturbed.   There are 6 treatments that include 1) undisturbed controls of the climax community which is typically mussel-dominated, 2) marked plots that are cleared annually to capture annual prey colonization and serve as controls for the interaction experiments, 3) partial cages that serve as cage controls for the interaction experiments 4) predator exclusion cages where an open-topped metal cage is placed around the plot to deter entry by predatory whelks and sea stars and where mussel and barnacle colonization occurs undeterred (to test predation), 5)  predator exclusion + barnacle removal cages in which predator are excluded plus barnacle settlers removed (to test facilitation by barnacles of mussels), and 6)  predator exclusion + mussel removal cages  in which predator are excluded plus mussel settlers removed  (to test competition effects of mussels on barnacles). All plots are photographed approximately monthly, and treatments are maintained by removing predators that invaded the cages and by clearing the relevant competitors (mussels or barnacles). Photos are analyzed by overlaying a 5 x5 grid onto the photo in ImageJ (so that each square is 4% cover) and estimating the % cover of each major space holder to the nearest 1%.  Space holders include: free space, algal crusts, acorn barnacles, gooseneck barnacles, mussels, articulated coralline algae, foliose algae and other sessile invertebrates. We also count the number of sea stars, whelks, and other herbivores.  Each spring, all plots are cleared and biomass collected (except controls). Biomass of functional groups is measured as wet and dry mass of mussels, barnacles, gooseneck barnacles, mobile predators, herbivores, and other. 


BCO-DMO Processing Description

* Table within sheet 1 of the submitted file "BCODMOSubmission_ACEMasterWide_ByFunc_2025-07-16.xlsx" was imported into the BCO-DMO data system for this dataset. Values "NA" imported as missing data values. Table will appear as Data File: 990939_v1_ace-experiment.csv (along with other download format options).

Missing Data Identifiers:
* In the BCO-DMO data system missing data identifiers are displayed according to the format of data you access. For example, in csv files it will be blank (null) values. In Matlab .mat files it will be NaN values. When viewing data online at BCO-DMO, the missing value will be shown as blank (null) values.

* Site list exported from supplied SiteList_STARS.xlsx(version from 2026-05-11) Sheet1 after changing date format to ISO 8601. Attached to dataset as sitelist_stars.csv. Sheet2 contained column information which was added to the sitelist_stars.csv file description after modifications to match the data table (description for column "state" added, column name "LoggerCode_OSU" is "OldLoggerCode_OSU" in data table).


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Data Files

File
990939_v1_ace-experiment.csv
(Comma Separated Values (.csv), 928.78 KB)
MD5:7933c714b583ad27f918b16a63fdc745
Primary data file for dataset ID 990939, version 1. Annual Colonization Experiment Photo Date for % cover and density of focal species. All plots are 15 x 15cm.

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Supplemental Files

File
sitelist_stars.csv
(Comma Separated Values (.csv), 4.90 KB)
MD5:258008f1bac6f545aee9a1fdce6790b6
Site List.

Column information (in csv format):

Column Name,Description,Units
SiteCode_STARS,Code used in all STARS datasets to designate site,unitless
Site,Name of STARS Site,unitless
SiteNum,Number of site north to south,unitless
Latitude,Latitude of site,decimal degrees
Longitude,Longitude of site,decimal degrees
SiteCode_OSU,Matching Site Codes for some datasets used by OSU,unitless
SiteCode_PISCO,Matching Site Codes for some datasets used by PISCO,unitless
SiteCode_MARINe,Matching Site Codes for some datasets used by MARINe,unitless
SiteType,"Site type for the STARS Project. Core = full suite of experiments. Ancillary = Surveys and environmental data only, None = not focal but may be in some datasets",unitless
Cape,"Cape, Point or Headland closest to site",unitless
CapeNum,Number of cape north to south,unitless
Region,"Regional Designation of Site. Oregon, Norcal, CenCal or SoCal",unitless
State,State of Site.,unitless
GroupCode,"Group that led work at that site. OSU, UCSC, Concordia or ODFW",unitless
OldLoggerCode_OSU,Site code used in older temperature logger data,unitless
Offset_MSLtoMLLW,The difference between 0 meters above mean sea level (MSL) and mean lower low water (MLLW) at that site,meters (m)
YearOfSSWS,The year that the sea star wasting disease outbreak began at that site. (yyyy),unitless
DateofSSWS,The date that the sea star wasting disease outbreak began at that site. See Gravem et al. 2021 IUCN report on Pycnopodia helianthoides for more. ISO 8601 date format.,unitless
MarineReserve,The nearest Oregon Marine Reserve to the Site,unitless
Designation,Whether the site is in an Oregon Marine Reserve or can serve as a comparison area,unitless

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Parameters

ParameterDescriptionUnits
PhotoID

Identifier for each photo showing SiteCode, StartYear, Replicate, Treatment, and PhotoDate. E.g. CB_2011_5_MP_2012-04-10.

unitless
PlotID

Identifier for each plot showing SiteCode, Replicate, Treatment. E.g. CB_5_MP.

unitless
SurveyID

Identifier for each survey showing SiteCode, Project, Date E.g. CB_ACE_2012-04-10.

unitless
ProjectCode

ACE for Annual Colonization Experiment.

unitless
State

State experiment performed. All Oregon.

unitless
Region

Region experiment performed. All Oregon.

unitless
Cape

Cape experiment performed. [Foulweather, Perpetua, Blanco]

unitless
SiteCode_STARS

Abbreviated Site Code.

unitless
Site

Site Name.

unitless
Latitude

Latitude of Site.

decimal degrees
Longitude

Longitude of Site.

decimal degrees
Treat

Detailed cage treatment of the plot. [CO = Control, never cleared; MP = Marked Plot Control, plot cleared annually; FC = Partial Fence Control, plot cleared annually; CG = Predator exclusion cage, plot cleared annually; CG-C = Predator exclusion cage, barnacles removed ~monthly, plot cleared annually; CG-M = Predator exclusion cage, mussels removed ~monthly, plot cleared annually]

unitless
PredTreat

Predator treatment of the plot. Cages exclude predators, primarily whelks in this zone . [Predators present, Predators absent]

unitless
CompTreat

Competitor treatment of the plot. Competitors removed ~monthly. [No competitors, removed, Mussels removed, Barnacles removed]

unitless
CageTreat

Simple Cage Treatment of the plot. [Cage absent, Cage present]

unitless
ClearTreat

Annual clearing treatment of the plot. [Cleared, Not cleared]

unitless
StartYear

Year clearance started. Usually clearances are in spring and plots are monitored until the following spring.

unitless
StartDate

Date clearance started. Usually clearances are in spring and plots are monitored until the following spring.

unitless
DaysSinceClear

Days since clearance started for the site (even if plot wasn't cleared).

unitless
Season

Season photo taken.

unitless
Date

Date photo taken.

unitless
Year

Year photo taken.

unitless
Month

Month photo taken.

unitless
Day

Day photo taken.

unitless
Rep

Replicate. Each replicate has one of each treatment and is clustered in space. [1 to 5]

unitless
Tot_Free

Total % cover free space.

percent
Tot_Crust

Total % cover algal crusts.

percent
Tot_AcornBarns

Total % cover acorn barnacles.

percent
Tot_GooseBarns

Total % cover gooseneck barnacles.

percent
Tot_Muss

Total % cover mussels.

percent
Tot_OthSessInv

Total % cover other sessile invertebrates.

percent
Tot_ArtCoral

Total % cover articulated coralline algae.

percent
Tot_Foliose

Total % cover foliose algae.

percent
Tot_Stars

Total number of sea stars.

count
Tot_Whelks

Total number of whelks.

count
Tot_Herb

Total number of herbivores - snails and chitons.

count
Tot_OthMobInv

Total number of other mobile invertebrates.

count

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Project Information

RAPID: Testing the rocky intertidal community consequences of the decimation of purple sea star populations along the Oregon coast by sea star wasting disease (Sea star wasting)


Coverage: Oregon coast


This study will investigate the ecological consequences of the decimation of sea star populations by wasting disease along the Oregon coast. Hallmarks of wasting disease are the formation of sores on the sea star that progress to cause loss of arms, and ultimately death of the animal. Wasting disease was reported in sea star populations including those of the purple sea star, Pisaster ochraceus, in British Columbia, Washington, and California as early as April 2013. In Oregon, wasting was first observed in April 2014, and by June 2014 rates of infection ranged up to 80%, and sea star abundance had declined. At that rate, many populations may disappear by the end of summer 2014. Prior research has shown that in the absence of the purple sea star, mid-shore mussel populations increase, and ultimately overgrow the sea weeds and invertebrates that occur low on the shore, reducing biodiversity. However, because disease events of this magnitude have never occurred along the entire coastline, it is unclear if the small-scale expansion of mussels observed previously will be a general result of this event. One possibility is that predators unaffected by wasting, such as whelks and crabs, will increase their predation effects and blunt the expected invasion of mussels to the low shore. The research in this project will evaluate this possibility by testing the role of these alternative predators. Broader Impacts include the training of undergraduate and graduate students, the involvement of coastal residents and the production of microdocumentaries and video to document the changing context of this ecosystem.

The research project is designed to test three hypotheses. First, that in the absence of Pisaster ochraceus, predation by whelks will increase in strength through increases in whelk abundance and in whelk size, and at least partially compensate for the absence of Pisaster. Second, the small sea star Leptasterias spp. will also expand its role as a predator through increased size and abundance, and expansion of its habitat beyond mussel beds. Although individuals of this sea star have been observed to suffer from wasting as well, the frequency so far appears low, and it seems likely this species may persist. Third, the crab Cancer productus, normally mostly a subtidal species, will expand its range into the intertidal and help to compensate for the loss of Pisaster. Tests of these hypotheses will include manual removal experiments (whelk removal, Leptasterias removal, removal of both and of neither), cage exclusion experiments (whelk exclusions), cage inclusion-exclusion experiments (Leptasterias inclusion, Leptasterias exclusion). Experiments will be replicated with appropriate controls, and done at multiple sites on the central Oregon coast that vary naturally in population abundances, rates of prey and predator recruitment, and oceanographic conditions. Results obtained under this unprecedented set of circumstances will deepen and expand our empirical understanding of the dynamics of an iconic ecosystem, and will help parameterize community models.

Additional Project Information: Sea Star Wasting Map


Collaborative Research: Mechanisms of resistance and resilience to system-wide loss of a keystone predator in an iconic intertidal community (Keystone Species Loss)


Coverage: Temperate west coast of North America


This project is affiliated with the Partnership for Interdisciplinary Studies of Coastal Oceans (PISCO) and Multi-Agency Rocky Intertidal Network (MARINe).

NSF abstract:

Diseases that compromise the health of predators can lead to large, abrupt and sometimes unexpected changes in the structure of ecosystems. This project will combine field surveys, manipulative experiments and mathematical models to both understand and predict the ecosystem-level effects of the unprecedented sea star wasting disease (SSWD) outbreak that devastated populations of Pisaster ochraceus, a critical predator, across the West Coast of the United States. Specifically, the project will determine (1) the ecological and environmental factors that promote vs. compromise the resilience of intertidal ecosystems to sea star wasting disease, (2) document the pace and extent of recovery from this major disturbance across the West Coast of the United States, and (3) identify hotspots of resilience to sea star wasting disease that may serve as important conservation targets to preserve these iconic ecosystems. The research will address important societal needs by cross-training undergraduate and graduate students in disease ecology, marine biology, mathematical modeling and biostatistics. Students from underrepresented groups will be recruited broadly from West Coast states. Each summer, four undergraduate students will be trained in rocky intertidal field research techniques. SSWD-focused modules will be developed and used in ecology courses at each institution to emphasize the importance of quantitative and interdisciplinary training for addressing important questions in biology. Graduate students will work with the Oregon Migrant Leadership Institute (OMLI) for migrant workers and their children to create workshops for students about SSWD. The PIs will continue interacting with the media and public groups, and will expand outreach activities through The Nature Conservancy and CoastWatch-sponsored workshops for high school teachers interested in involving students in sea star monitoring to ensure that the results of this project are disseminated beyond traditional academic circles. Finally, a series of model-based interactive web modules will be created as part of this project to illustrate the ecosystem-level effects of sea star wasting disease to the broader public. The studies on this model system will lead to a better understanding of how other ecosystems may resist or be vulnerable to human activities (e.g., fishing, hunting and habitat destruction) that asymmetrically influence top predators.

Diseases that threaten the health of predators can reduce their top-down influence and thus lead to significant changes in ecosystem structure. In 2013-15, sea star wasting disease (SSWD) devastated populations of Pisaster ochraceus, the original keystone predator, along the west coast of North America in one of the most extensive marine disease events ever recorded. This project will leverage this unprecedented outbreak to test and extend keystone predation theory by documenting and explaining the temporal pace, geographical extent, and spatiotemporal co-occurrence of ecosystem recovery from SSWD. The disease event also provides an opportunity to test the resistance and resilience of a well-studied ecosystem at an unprecedented scale. At each of 14 sites, the investigators will quantify processes that underlie potential resistance of the system to loss of sea stars (prey recruitment and colonization, mussel growth, predation intensity, facilitative interactions among sessile organisms, and the effect of alternative predators). In the latter experiments, the PIs will conduct caging exclusion experiments to test the effects of both larger (e.g., birds) and smaller (e.g., whelks) alternative predators on prey recolonization of cleared plots. The investigators will also conduct a novel set of experiments to manipulate factors affecting facilitation of mussels by barnacles and turf-forming algae. All these empirical studies will be used to parameterize modeling efforts that will explore the longer-term and larger-scale implications of these processes, both for this system and for other ecosystems. Specifically, the PIs will fit a novel spatially-explicit metacommunity model to the empirical data in order to determine the relative importance of intraspecific and interspecific resistance vs. resilience mechanisms for the recovery of intertidal ecosystems following a historical, coastal-scale SSWD disturbance.


LTREB: Testing tipping points in a model rocky intertidal meta-ecosystem – Climate-change, increasing variances, and response mechanisms (LTREB Intertidal Tipping Points)

Coverage: West coast of North America


NSF abstract:

In recent decades, ocean ecosystems, long thought to be immune to change, have undergone disruptions to their structure, diversity, and geographic range, yet the actual underlying reasons for such changes in oceanic biota are often unclear. Coastal intertidal zones (i.e., the shore between high and low tides) have long served as important ecological model systems because of advantages in accessibility and ease of observation, occupancy by easily studied and manipulated organisms of relatively short lifespans, and exposure to often severe environmental conditions. This research will address the stability of a well-known rocky shore system along the Oregon and California coasts. Prior long-term research indicates that, although casual observation suggests these systems are stable, in fact, they may be on the cusp of shifting into another state, losing iconic organisms like mussels and sea stars, and becoming dominated by seaweeds. These changes might be comparable to losing trees and large predators from terrestrial systems. This study would result in the training of undergraduates and graduate students, including individuals from under-represented groups. Additionally, this study would include outreach to the general public.

The researchers will focus particularly on impacts of increasing and more variable warming on community recovery. For example, climate oscillations (e.g., El Niño), coastal upwelling, and particularly temperature have all changed in recent decades in ways leading to increased stress on intertidal biota. In apparent response, coastal ecosystems evidently have become less productive, organismal performance (growth, reproduction) has declined, and key dynamical processes (species interactions) have weakened. The new research will pursue these strong hints of an impending “tipping point” by (1) continuing the projects that led to the insights of increasing instability, (2) adding new projects that will pinpoint ecological changes, and (3) expanding the region of work to include locations in California. Research will assess whether or not sea stars recover from wasting disease, experimentally test if species interactions are indeed weakening, quantify the annual inputs of new prey and changes in abundance, diversity, stability, and resilience of intertidal communities, and document changes in the physical environment. Using field observations and experiments, the research will provide insight into impacts of environmental change, particularly warming, on the future of coastal ecosystems, and more generally, into possible future states of Earth’s ecosystems. Using these data, we will test the hypothesis that direct and indirect effects of climate change are driving, or may drive these systems into new, alternative states.

This award reflects NSF's statutory mission and has been deemed worthy of support through evaluation using the Foundation's intellectual merit and broader impacts review criteria.



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Program Information

Partnership for Interdisciplinary Studies of Coastal Oceans (PISCO)


Coverage: West coast of North America from Mexico to Alaska


The Partnership for Interdisciplinary Studies of Coastal Oceans is a long-term ecosystem research and monitoring program established with the goals of: 

  • understanding dynamics of the coastal ocean ecosystem along the U.S. west coast
  • sharing that knowledge so ocean managers and policy makers can make science based decisions regarding coastal and marine stewardship
  • producing a new generation of scientists trained in interdisciplinary collaborative approaches

Over the last 10 years, PISCO has successfully built a unique research program that combines complementary disciplines to answer critical environmental questions and inform management and policy. Activities are conducted at the latitudinal scale of the California Current Large Marine Ecosystem along the west coast of North America, but anchored around the dynamics of coastal, hardbottom habitats and the oceanography of the nearshore ocean – among the most productive and diverse components of this ecosystem. The program integrates studies of changes in the ocean environment through ecological monitoring and experiments. Scientists examine the causes and consequences of ecosystem changes over spatial scales that are the most relevant to marine species and management, but largely unstudied elsewhere.

Findings are linked to solutions through a growing portfolio of tools for policy and management decisions. The time from scientific discovery to policy change is greatly reduced by coordinated, efficient links between scientists and key decision makers.

Core elements of PISCO are:

  • Interdisciplinary ecosystem science
  • Data archiving and sharing
  • Outreach to public and decision-making user groups
  • Interdisciplinary training
  • Coordination of distributed research team

Established in 1999 with funding from The David and Lucile Packard Foundation, PISCO is led by scientists from core campuses Oregon State University (OSU); Stanford University’s Hopkins Marine Station; University of California, Santa Cruz (UCSC); and University of California, Santa Barbara (UCSB). Collaborators from other institutions also contribute to leadership and development of PISCO programs.  As of 2005, core PISCO activities are funded by collaborative grants from The David and Lucile Packard Foundation and the Gordon and Betty Moore Foundation. Core support, along with additional funding from diverse public and private sources, make this unique partnership possible.


Partnership for Interdisciplinary Studies of Coastal Oceans (PISCO)


Coverage: West coast of North America from Mexico to Alaska


The Partnership for Interdisciplinary Studies of Coastal Oceans is a long-term ecosystem research and monitoring program established with the goals of: 

  • understanding dynamics of the coastal ocean ecosystem along the U.S. west coast
  • sharing that knowledge so ocean managers and policy makers can make science based decisions regarding coastal and marine stewardship
  • producing a new generation of scientists trained in interdisciplinary collaborative approaches

Over the last 10 years, PISCO has successfully built a unique research program that combines complementary disciplines to answer critical environmental questions and inform management and policy. Activities are conducted at the latitudinal scale of the California Current Large Marine Ecosystem along the west coast of North America, but anchored around the dynamics of coastal, hardbottom habitats and the oceanography of the nearshore ocean – among the most productive and diverse components of this ecosystem. The program integrates studies of changes in the ocean environment through ecological monitoring and experiments. Scientists examine the causes and consequences of ecosystem changes over spatial scales that are the most relevant to marine species and management, but largely unstudied elsewhere.

Findings are linked to solutions through a growing portfolio of tools for policy and management decisions. The time from scientific discovery to policy change is greatly reduced by coordinated, efficient links between scientists and key decision makers.

Core elements of PISCO are:

  • Interdisciplinary ecosystem science
  • Data archiving and sharing
  • Outreach to public and decision-making user groups
  • Interdisciplinary training
  • Coordination of distributed research team

Established in 1999 with funding from The David and Lucile Packard Foundation, PISCO is led by scientists from core campuses Oregon State University (OSU); Stanford University’s Hopkins Marine Station; University of California, Santa Cruz (UCSC); and University of California, Santa Barbara (UCSB). Collaborators from other institutions also contribute to leadership and development of PISCO programs.  As of 2005, core PISCO activities are funded by collaborative grants from The David and Lucile Packard Foundation and the Gordon and Betty Moore Foundation. Core support, along with additional funding from diverse public and private sources, make this unique partnership possible.


Long Term Research in Environmental Biology (LTREB)



Long Term Research in Environmental Biology (LTREB)

Supports research for a period of 10 years or longer to generate an extended time series of data with a focus on evolutionary biology, ecology and ecosystem science.

Synopsis
The Long Term Research in Environmental Biology (LTREB) Program supports the generation of extended time series of data to address important questions in evolutionary biology, ecology, and ecosystem science. Research areas include, but are not limited to, the effects of natural selection or other evolutionary processes on populations, communities, or ecosystems; the effects of interspecific interactions that vary over time and space; population or community dynamics for organisms that have extended life spans and long turnover times; feedbacks between ecological and evolutionary processes; pools of materials such as nutrients in soils that turn over at intermediate to longer time scales; and external forcing functions such as climatic cycles that operate over long return intervals.



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Funding

Funding SourceAward
NSF Division of Ocean Sciences (NSF OCE)
NSF Division of Ocean Sciences (NSF OCE)
NSF Division of Environmental Biology (NSF DEB)
NSF Division of Environmental Biology (NSF DEB)
NSF Division of Environmental Biology (NSF DEB)

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