Traits and metabolic constraints affect marine animal forest structure

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Published: Jun 5, 2025
DOI: https://doi.org/10.12681/mms.39396
Keywords:
gorgonian sponge energy crowding morphology size
MARK JOHNSON
Ryan Institute and School of Natural Sciences, University of Galway, University Road, Galway, Ireland, H91 TK33
https://orcid.org/0000-0002-5078-1410
JESSICA FERREIRA
Ryan Institute and School of Natural Sciences, University of Galway, University Road, Galway, Ireland, H91 TK33
https://orcid.org/0009-0001-3419-1525
Abstract

The term marine animal forests (MAFs) describes a diverse range of 3-dimensional habitats dominated by benthic suspension feeders. The ecosystem functions of MAFs are broadly understood, but an overall framework remains to be developed. We present an equation that generalizes the relationships among species’ height, depth of occurrence and mean water temperature in terms of the metabolic theory of ecology (MTE). The applicability of the equation to field observations collated from the literature (n = 284) was tested using a multiple regression of colony density (log of numbers m-2) against transformed depth, height and temperature as predictors. Morphology was also examined both as a factor and when interacting with the effect of colony height. The model explained 39.7% of the variability in colony density with significant effects of morphology and height. Colony density decreased with taller individuals (slope -0.760, SE 0.1677). Typical mass-height scaling in MAF species suggests that the observed slope is less than the slope that would be predicted by the MTE. A detailed evaluation requires further estimates of metabolic rates in MAF species. Some morphologies, particularly planar and bushy sea rods, reach higher colony densities than expected for their size. These morphologies have been associated with environments where food supply may be higher due to currents or turbidity. Morphology-environment associations may therefore allow traits to be incorporated into an overall description of MAF ecosystems. With further improvements in observations of both colony density and metabolic scaling with size, a comprehensive description of energy flow in MAFs could be possible.

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References
Assis, J., Fernández Bejarano, S.J., Salazar, V.W., Schepers, L., Gouvêa, L. et al., 2024. Bio-ORACLE v3.0. Pushing marine data layers to the CMIP6 Earth system models of climate change research. Global Ecology and Biogeography, 33 (4), e13813.
Brown, J.H., Gillooly, J.F., Allen, A.P., Savage, V.M., West, G.B., 2004. Toward a metabolic theory of Ecology. Ecology, 85 (7), 1771-1789.
Burgess, S.C., Ryan, W.H., Blackstone, N.W., Edmunds, P.J., Hoogenboom M.O. et al., 2017. Metabolic scaling in modular animals. Invertebrate Biology, 136 (4), 456-472.
Cau, A., Bramanti, L., Cannas, R., Follesa, M.C., Angiolillo, M. et al., 2016. Habitat constraints and self-thinning shape Mediterranean red coral deep population structure: implications for conservation practice. Scientific Reports, 6 (1), 23322.
Cerrano, C., Danovaro, R., Gambi, C., Pusceddu, A., Riva, A. et al., 2010. Gold coral (Savalia savaglia) and gorgonian forests enhance benthic biodiversity and ecosystem functioning in the mesophotic zone. Biodiversity Conservation, 19, 153-167.
Chimienti, G., Angeletti, L., Rizzo, L., Tursi, A., Mastrototaro, F., 2018a. ROV vs trawling approaches in the study of ben thic communities: the case of Pennatula rubra (Cnidaria: Pennatulacea). Journal of the Marine Biological Association UK. 98 (8) ,1859-1869.
Chimienti, G., Tursi, A., Mastrototaro, F., 2018b. Biometric relationships in the red sea pen Pennatula rubra (Cnidaria: Pennatulacea). IEEE International Workshop on Metrology for the Sea; Learning to Measure Sea Health Parameters (MetroSea) Bari, Italy, 212-216.
Coma, R., Ribes, M., Zabala, M., Gili, J.-M.,1998. Growth in a Modular Colonial Marine Invertebrate. Estuarine, Coastal and Shelf Science, 47 (4), 459-470.
Coppari, M., Zanella, C., Rossi, S., 2019. The importance of coastal gorgonians in the blue carbon budget. Scientific Reports, 9 (1), 13550.
Currie, D.J., Fritz, J.T., 1993. Global patterns of animal abundance and species energy use. Oikos, 67, 56-68.
Damuth, J.,1987. Interspecific allometry of population density in mammals and other animals: the independence of body mass and population energy-use. Biological Journal of the Linnean Society, 31 (3), 193-246.
Derviche, P., Menegotto, A., Lana, P., 2022. Carbon budget trends in octocorals: a literature review with data reassessment and a conceptual framework to understand their resilience to environmental changes. Marine Biology, 169 (12), 159.
Ferreira, J.A., Johnson, M.P.,2025. Height and density data for marine animal forest forming species [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15276467.
Fox, J., Weisberg, S., 2018. Visualizing fit and lack of fit in complex regression models with predictor effect plots and partial residuals. Journal of Statistical Software, 87, 1-27.
Fox, J., Weisberg, S., 2019. An R Companion to Applied Regression, 3rd Edition. Thousand Oaks, CA
Francis, F.T., Burke, L., Marliave, J., Schultz, J., Borden, L. et al., 2024. Fishing damage to cloud sponges may lead to losses in associated fish communities in Pacific Canada. Marine Environment Research, 197, 106448.
Gillooly, J.F., Brown J.H., West, G.B., Savage, V.M. Charnov, E.L., 2001. Effects of Size and Temperature on Metabolic Rate. Science, 293(5538) 2248-2251.
Gjoni, V., Glazier, D.S., 2020. A Perspective on body size and abundance relationships across ecological communities. Biology, 9 (3), 42.
Gomes-Pereira, J.N., Carmo, V., Catarino, D., Jakobsen, J., Alvarez, H. et al., 2017. Cold-water corals and large hydrozoans provide essential fish habitat for Lappanella fasciata and Benthocometes robustus. Deep Sea Research II, 145, 33-48.
Gori, A., Rossi, S., Berganzo, E., Pretus, J.L., Dale, M.R.T. et al., 2012. Spatial distribution patterns of the gorgonians Eunicella singularis, Paramuricea clavata, and Leptogorgia sarmentosa (Cape of Creus, Northwestern Mediterranean Sea). Marine Biology, 158, 143-158.
Isaac, N.J.B., Storch, D., Carbone, C., 2017. Taxonomic variation in size–density relationships challenges the notion of energy equivalence. Biology Letters, 7 (4), 615-618.
Johnson, M.P., White, M., Wilson, A., Würzberg, L., Schwabe, E. et al., 2013. A vertical wall dominated by Acesta excavata and Neopycnodonte zibrowii, part of an undersampled group of deep-sea habitats. PLoS ONE, 8 (11), e79917. Kleiber, M., 1932. Body size and metabolism. Hilgardia, 6 (11), 315-351.
Linares, C., Coma, R., Garrabou, J., Díaz, D., Zabala, M., 2008. Size distribution, density and disturbance in two Mediterranean gorgonians: Paramuricea clavata and Eunicella singularis. Journal of Applied Ecology, 45 (2), 688-699.
Martin, J.H., Knauer, G.A., Karl. D.M., Broenkow, W.W., 1987. VERTEX: carbon cycling in the northeast Pacific. Deep Sea Research Part A. Oceanographic Research Papers, 34 (2), 267-285.
McClain, C.R., Allen, A.P., Tittensor, D.P., Rex, M.A., 2012. Energetics of life on the deep seafloor. Proceedings of the National Academy of Sciences, 109 (38), 15366-15371.
Murillo, F.J., MacDonald, B.W., Kenchington, E., Campana, S.E., Sainte-Marie, B. et al, 2018. Morphometry and growth of sea pen species from dense habitats in the Gulf of St. Lawrence, eastern Canada. Marine Biology Research, 14 (4), 366-382.
OBIS, 2025. Ocean Biodiversity Information System. Intergovernmental Oceanographic Commission of UNESCO. https://obis.org. (Accessed 18 April 2025).
OSPAR Commission, 2008. OSPAR List of Threatened and/or Declining Species and Habitats. Reference number 2008-6. https://www.ospar.org/documents?v=7099.
Ouyang, S., Xiang, W., Wang, X., Xiao, W., Chen, L. et al., 2019. Effects of stand age, richness and density on productivity in subtropical forests in China. Journal of Ecology, 107 (5), 2266-2277.
Parker, G.G., 2020. Tamm review: Leaf Area Index (LAI) is both a determinant and a consequence of important processes in vegetation canopies. Forest Ecology and Management, 477, 118496.
Pearman, T.R.R., Robert, K., Callaway, A., Hall, R.A., Mienis F. et al., 2023. Spatial and temporal environmental heterogeneity induced by internal tides influences faunal patterns on vertical walls within a submarine canyon. Frontiers in Marine Science, 10, 1091855.
Pedersen, R.O., Faurby, S., Svenning, J.C. 2017. Shallow size-density relations within mammal clades suggest greater intra-guild ecological impact of large-bodied species. Journal of Animal Ecology, 86 (5), 1205-1213.
Provoost, P., Bosch, S., 2022. _robis: Ocean Biodiversity Information System (OBIS) Client_. R package version 2.11.3, <https://CRAN.R-project.org/package=robis>.
R Core Team 2024. _R: A Language and Environment for Statistical Computing_. R Foundation for Statistical Computing, Vienna, Austria. <https://www.R-project.org/>.
Rossi, S., Bramanti, L., Broglio, E., Gili, J.M., 2012 Trophic impact of long-lived species indicated by population dynamics in the short-lived hydrozoan Eudendrium racemosum. Marine Ecology Progress Series, 467, 97-111.
Rossi, S., Bramanti, L., Gori, A., Orejas, C., 2017. Animal forests of the world: An overview. p. 1-28. In: Marine Animal Forests. Rossi, S., Bramanti, L., Gori, A., Orejas, C. (Eds). Springer, Switzerland.
Rossi, P., Ponti, M., Righi, S., Castagnetti, C., Simonini, R. et al., 2021. Needs and gaps in optical underwater technologies and methods for the investigation of marine animal forest 3D-structural complexity. Frontiers in Marine Science, 8, 591292.
Rossi, S., Bramanti, L., Horta, P,. Allcock, L., Carreiro-Silva, M. et al., 2022. Protecting global marine animal forests. Science, 376, 929-929.
Santavy, D.L., Courtney, L.A., Fisher, W.S., Quarles, R.L., Jordan, S.J., 2013. Estimating surface area of sponges and gorgonians as indicators of habitat availability on Caribbean coral reefs. Hydrobiologia, 707, 1-16.
Schönberg, C.H.L., 2021. No taxonomy needed: Sponge functional morphologies inform about environmental conditions. Ecological Indicators, 129, 107806.
Schuster, L. Cameron, H. White, C.R., Marshall D.J., 2021. Metabolism drives demography in an experimental field test. Proceedings of the National Academy of Science U.S.A., 118 (34) e2104942118.
Shokri, M., Cozzoli, F., Basset, A., 2025. Metabolic rate and foraging behaviour: a mechanistic link across body size and temperature gradients. Oikos, e10817.
Shokri, M., Lezzi, L., Basset, A., 2024. The seasonal response of metabolic rate to projected climate change scenarios in aquatic amphipods. Journal of Thermal Biology, 124, 103941.
Teixidó, N., Garrabou, J., Harmelin, J.-G., 2011. Low dynamics, high longevity and persistence of sessile structural species dwelling on Mediterranean coralligenous outcrops. PLoS One, 6 (8), e23744.
Tyberghein, L., Verbruggen, H., Pauly, K., Troupin, C., Mineur, F. et al., 2012. Bio-ORACLE: A global environmental dataset for marine species distribution modelling. Global Ecology and Biogeography, 21 (2), 272-281.
Tyler, E.H.M., Somerfield, P.J., Vanden Berghe, E., Bremner, J., Jackson, E. et al., 2012. Extensive gaps and biases in our knowledge of a well-known fauna: implications for integrating biological traits into macroecology. Global Ecology and Biogeography, 21 (9), 922-934.
WoRMS Editorial Board, 2024. World Register of Marine Species. Available from https://www.marinespecies.org at VLIZ. (Accessed 8 February 2024).