We searched multiple electronic bibliographic databases (web of Science, ASFA, Google Scholar), with the following terms: “macroalga(e)” and “fish(es)”, “seaweed(s)”, and “fish(es)”, “kelp” and fish(es), “kelp forests (and beds) and fish(es)”. The last search was conducted on 2 December 2022. We divided the records identified through database searches and records from other sources including references from the studies identified through the database searches. Personal reference collections and colleagues with active research interests in these areas were also included. We recorded the total number of studies for screening (>100 studies). We reviewed studies that manipulated habitat and subsequently sampled fish that were published to the end of 2020. In addition to manipulative experiments, we also reviewed observational studies that measured and compared the natural variability in fish(es) metrics (abundance, species richness, functional diversity) in kelp and non-kelp forests (i.e., urchin barrens, sponge gardens, turf algae, sand).
For articles that met selection criteria, we recorded type of study (experimental or observational), mean species richness, total species abundance, and/or abundances of single species in either treatments (kelp control versus removal) or habitat type (kelp forest versus non-kelp forest), sample size of groups, and variance in order to estimate the effect of kelp on fish. Where possible, we noted the ontogenetic state of fish communities or single species assessed (e.g., young of the year (YoY) or settlement phase, juveniles and/or adults). Individual values from most articles were found in graphs such as scatterplots and bar plots in which case a web plot digitizer was used for translation of plots into numerical values (Rohatgi 2020). Articles that did not provide relevant means, measures of variance, or other values in tables, means plots, boxplots, or scatterplots were not used in this meta-analysis.
Where possible, we recorded information about fish trophic groups and size to assess the variability of the effects of fish on kelp. The functional roles of reef fish species may be described using broad trophic groups or exploring more detailed information on traits. We extracted trait data for individual fish species from FishBase using rFishBase. We used broad trophic groups in order to compare species between studies. We categorized species as planktivores, herbivores, detritivores, omnivores, benthic microcarnivores (that feed mostly on small epifauna), benthic macroinvertivores, and general carnivores (including piscivores). These groupings were available for all taxa, regardless of taxonomic resolution. We also recorded maximum total length and a continuous measure of fish trophic level.
To assess the effects of kelp structural complexity on fish, we grouped kelp according to their numbers of stipes as single (i.e., Ecklonia) or multi-stipe (i.e., Macrocystis pyrifera). Studies spaned a total of 17 different kelp species: 13 Laminarian species (Macrocystis pyrifera, Ecklonia radiata, Saccharina latissima, L. digitata, L. farlowii, Eisenia arborea, Egregia menziesii, Nereocystis luetkeana, Pterygophora californica, Lessonia trabeculata, Undaria pinnatifida) and 4 Fucoid species (Carpophyllum maschalocarpum, C. flexuosum, Cystophora racemosa, C. retroflexa). Some studies (experimental and/or observational) have compared a complex set of kelp species against habitats devoid of kelp forests. When two kelp species with different morphologies were studied, we categorized the study using the species that was more structurally complex.
Papers Included
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