{
  "abstract": "The loose equilibrium (LE) hypothesis predicts that biotic assemblages will exhibit modest change from year-to-year but will not follow directed trajectories away from a stable baseline over longer periods of time. Using 8 years (2016–2023) of data from 21 National Ecological Observatory Network streams, we test the LE hypothesis for benthic insect assemblages. We also expand the traditional LE method, which focuses on a single sampling season, by using multi-season data to determine whether LE is influenced by seasonal change. Results indicate that most benthic insect assemblages may be in a state of LE. Seventeen of the 21 assemblages exhibited random, non-directional change among sampling events, consistent with the LE hypothesis. However, directional movement away from a baseline condition was detected for two insect assemblages, while directional movement from the baseline, followed by a return toward the baseline was detected for the remaining two assemblages. Supplementary taxonomic, feeding behavior, and environmental data suggest that the four instances of directional change may be responses to recent disturbances or altered physicochemical habitat. Distinct seasonal clusters (spring, summer, fall) of insects are also detected for 13 of the 21 assemblages. Among the 13 clustered assemblages, 11 have at least one significantly small cluster that reflects minimal interannual change within the respective season. We conclude by explaining how strong seasonality may support LE. Specifically, we hypothesize that seasonally distinct, yet predictable environments promote inter-seasonal consistency in assemblage structure, thereby constraining opportunities for insect assemblages to shift far from the baseline condition. This work adds to the broader field of equilibrium theory in ecology and may help anticipate future changes in benthic assemblage structure. Author summary: Assemblages of living organisms are never truly static. They all change through time, though change is often gradual and centered about a stable baseline. To characterize this dynamic, ecologists have developed a theory of ‘loose equilibrium’ (LE). LE predicts that biotic assemblages will exhibit modest change over short time intervals but will not follow continuous or directed trajectories away from a stable baseline over longer intervals. This study is a novel test of LE for freshwater insect assemblages. It begins with an 8-year record of seasonal insect samples from 21 streams in the National Ecological Observatory Network. Using this robust dataset in combination with multivariate analyses and Monte Carlo tests, we find that insect assemblages in 81% of the study streams are likely to be in a state of LE. We also show that seasonal clustering, defined by distinct spring, summer, and fall subsets of insects, is a common feature of the study streams. Finally, we hypothesize that seasonally variable, yet predictable environments may help maintain LE by creating a template that promotes intraseasonal consistency in assemblage structure while constraining change among seasons.",
  "abstract_shown": true,
  "affiliations": [
    "School of Life Sciences & Sustainability, Virginia Commonwealth University, Richmond, Virgina, United States of America",
    "National Ecological Observatory Network, Battelle, Boulder, Colorado, United States of America",
    "Department of Biological Sciences, Wayne State University, Detroit, Michigan, United States of America",
    "Odum School of Ecology, University of Georgia, Athens, Georgia, United States of America",
    "Institute of Environment, Florida International University, Miami, Florida, United States of America",
    "Padre Associates, Bakersfield, California, United States of America",
    "Department of Civil, Environmental & Architectural Engineering, University of Kansas, Lawrence, Kansas, United States of America",
    "Department of Biological Sciences, Florida International University, Miami, Florida, United States of America",
    "Missouri Department of Natural Resources, Jefferson City, Missouri, United States of America",
    "Land Resources and Environmental Sciences, Montana State University, Bozeman, Montana, United States of America",
    "Department of Biology, Georgia Southern University, Statesboro, Georgia, United States of America",
    "Department of Natural Resources & Environmental Science, University of Nevada at Reno, Reno, Nevada, United States of America",
    "Environmental Science, Policy & Management, University of California at Berkeley, Berkeley, California, United States of America",
    "Virginia Department of Wildlife Resources, Henrico, Virginia, United States of America",
    "City of Omaha Environmental Quality Control Division, Omaha, Nebraska, United States of America",
    "Eurofins, Albuquerque, New Mexico, United States of America",
    "Department of Geology and Environmental Science, University of Pittsburgh, Pittsburgh, Pennsylvania, United States of America"
  ],
  "article_type": "Research Article",
  "arxiv": null,
  "author_total": 19,
  "authors": [
    {
      "affs": [
        0
      ],
      "name": "Daniel J. McGarvey"
    },
    {
      "affs": [
        1
      ],
      "name": "Stephanie M. Parker"
    },
    {
      "affs": [
        2
      ],
      "name": "Héctor J. Esparra-Escalera"
    },
    {
      "affs": [
        3
      ],
      "name": "Natalia Vargas-López"
    },
    {
      "affs": [
        4
      ],
      "name": "Lauren K. Emer"
    },
    {
      "affs": [
        5
      ],
      "name": "Jonathan R. Juarez"
    },
    {
      "affs": [
        6
      ],
      "name": "Victoria Secondine"
    },
    {
      "affs": [
        7
      ],
      "name": "Liz D. Ortiz Muñoz"
    },
    {
      "affs": [
        3
      ],
      "name": "Viviana Bravo Ortiz"
    },
    {
      "affs": [
        8
      ],
      "name": "Alexis R. Reifsteck"
    },
    {
      "affs": [
        9
      ],
      "name": "Jordyn Solliday"
    },
    {
      "affs": [
        10
      ],
      "name": "Reginald K. Turner"
    },
    {
      "affs": [
        11
      ],
      "name": "Deandre Presswood"
    },
    {
      "affs": [
        12
      ],
      "name": "Patina K. Mendez"
    },
    {
      "affs": [
        13
      ],
      "name": "Makayla DeVivo"
    },
    {
      "affs": [
        14
      ],
      "name": "Rodrigo Meza Gonzalez"
    },
    {
      "affs": [
        15
      ],
      "name": "Hope K. Romero"
    },
    {
      "affs": [
        16
      ],
      "name": "Jason Aguirre"
    },
    {
      "affs": [
        3
      ],
      "name": "Breanna Ondich"
    }
  ],
  "bibtex": "https://research.draconitic.com/paper/akh8dcffrrdcb.bib",
  "cited_by": 0,
  "doi": "10.1371/journal.pesy.0000037",
  "fulltext_url": null,
  "id": "akh8dcffrrdcb",
  "issns": [
    "3070-8141"
  ],
  "landing_url": "https://doi.org/10.1371/journal.pesy.0000037",
  "language": null,
  "licence": "CC-BY-4.0",
  "licence_label": "CC BY 4.0",
  "licence_source": "journal",
  "licence_url": "https://creativecommons.org/licenses/by/4.0/",
  "metadata_licence": null,
  "native_subjects": [
    [
      "openalex",
      "2303|Freshwater macroinvertebrate diversity and ecology"
    ],
    [
      "plos",
      "/Biology and life sciences/Organisms/Eukaryota/Animals/Invertebrates"
    ],
    [
      "plos",
      "/Biology and life sciences/Organisms/Eukaryota/Animals/Invertebrates/Arthropoda/Insects"
    ],
    [
      "plos",
      "/Biology and life sciences/Organisms/Eukaryota/Animals/Vertebrates/Fish/Freshwater fish"
    ],
    [
      "plos",
      "/Biology and life sciences/Taxonomy"
    ],
    [
      "plos",
      "/Biology and life sciences/Zoology/Animals/Invertebrates"
    ],
    [
      "plos",
      "/Biology and life sciences/Zoology/Animals/Invertebrates/Arthropoda/Insects"
    ],
    [
      "plos",
      "/Biology and life sciences/Zoology/Animals/Vertebrates/Fish/Freshwater fish"
    ],
    [
      "plos",
      "/Biology and life sciences/Zoology/Entomology/Insects"
    ],
    [
      "plos",
      "/Computer and information sciences/Data management/Taxonomy"
    ],
    [
      "plos",
      "/Earth sciences/Marine and aquatic sciences/Aquatic environments/Freshwater environments/Fresh water"
    ],
    [
      "plos",
      "/Earth sciences/Seasons"
    ],
    [
      "plos",
      "/Earth sciences/Seasons/Spring"
    ],
    [
      "plos",
      "/Ecology and environmental sciences/Aquatic environments/Freshwater environments/Fresh water"
    ],
    [
      "plos",
      "/Physical sciences/Mathematics/Algebra/Polynomials"
    ]
  ],
  "openalex": "W7221550425",
  "pdf_url": "https://journals.plos.org/ecosystems/article/file?id=10.1371/journal.pesy.0000037&type=printable",
  "pmcid": null,
  "pmid": null,
  "published": "2026-10-12",
  "publisher": "Public Library of Science",
  "ris": "https://research.draconitic.com/paper/akh8dcffrrdcb.ris",
  "sources": [
    {
      "first_seen": "2026-10-30T12:17:30Z",
      "native_id": "10.1371/journal.pesy.0000037",
      "source": "plos",
      "url": "https://doi.org/10.1371/journal.pesy.0000037"
    }
  ],
  "subjects": [
    {
      "field": "biology",
      "origin": "plos:zoology",
      "subfield": "zoology-behaviour"
    }
  ],
  "title": "Tightening the loose equilibrium concept: New seasonal insights with benthic insect data from National Ecological Observatory Network streams",
  "title_text": "Tightening the loose equilibrium concept: New seasonal insights with benthic insect data from National Ecological Observatory Network streams",
  "type": "peer-reviewed",
  "updated_at": "2026-10-30T12:17:30Z",
  "url": "https://research.draconitic.com/paper/akh8dcffrrdcb",
  "venue": "PLOS Ecosystems",
  "versions": []
}