Functional diversity: a review on freshwater fish research

Gomes, Louise C.; Dias, Rosa M.; Ruaro, Renata; Benedito, Evanilde

doi:10.1590/1982-0224-2023-0022

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Original Article • Neotrop. ichthyol. 21 (02) • 2023 • https://doi.org/10.1590/1982-0224-2023-0022 copy

Functional diversity: a review on freshwater fish research

Authorship SCIMAGO INSTITUTIONS RANKINGS

Abstract

Functional diversity is an emergent approach in ecology that has been applied globally to better understand the relationships between organisms and the environment. However, assessing the functional diversity of freshwater fish is a challenge for scientists. Aiming to summarize the evolution of scientific knowledge on fish functional diversity, we performed a systematic review of the literature published from 1945 to 2021 using the Web of Science. Based on the 101 articles reviewed, we found that publications about functional diversity of fishes have increased over time, mainly in Neotropical, Indomalayan and Palearctic regions. Most studies were conducted in lotic ecosystems, especially to assess environmental impacts such as biological invasions and land use. Functional diversity has been assessed mainly by morphological traits that reflect feeding and locomotion dimensions. Functional richness was the most common index used in the studies. Our findings suggest that political neglect and lack of investments may hamper the research development in several places. The missing information about the functional traits of many species may limit the use of the functional approach. We also highlight the need for the incorporation of functional diversity in conservation programs once functional diversity is a key facet of biodiversity to maintain freshwater ecosystem functioning.

Keywords:
Aquatic ecosystems; Biogeographic realms; Functional ecology; Functional traits; Systematic review

Sociedade Brasileira de Ictiologia Neotropical Ichthyology, Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura, Universidade Estadual de Maringá., Av. Colombo, 5790, 87020-900, Phone number: +55 44-3011-4632 - Maringá - PR - Brazil
E-mail: neoichth@nupelia.uem.br

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[1] Edited-by

Lilian Casatti

Extracted data	Classification	Application
a) Year of publication	1945–2021	Used to determine the temporal trend of publications.
b) Biogeographic realm	Palearctic, Nearctic, Neotropical, Indomalayan, Australian, Afrotropical, and Global (when the study assessed more than one region).	Used to identify the distribution of research effort among biogeographic realms.
c) Freshwater environment	River, Stream, Lake, Reservoir, Floodplain, and several (when the study assessed more than one environment).	Used to verify the type of freshwater ecosystem most assessed.
d) General background	1) Biological invasion; 2) Climate change; 3) Conservation; 4) Environmental factors; 5) Environmental filtering; 6) Flood pulse; 7) Functional structure; 8) Habitat heterogeneity; 9) Impoundments; 10) Land use; 11) Lateral connectivity; 12) Methodological; 13) Multiple stressors; 14) Taxonomic and functional patterns	Classification based on the main objectives of the studies.
e) Functional trait	Ecological traits; morphological traits	Classification based on the type of trait measure. Used to identify the traits most applied to assess functional diversity.
f) Functional category	Feeding; Habitat use; Life History; Locomotion; and Physiology (Classified according to Villéger et al., 2017).	Identify which functional category with the greatest number of traits evaluated.
g) Functional diversity index	All indices found in the reviewed studies.	Used to identify the main index applied to quantify the functional diversity of fish.

Functional diversity index	Number of studies*	Description	References
Functional Richness (FRic)	64	Volume of the functional space occupied by the community	*Mason et al. (2005)Mason NWH, Mouillot D, Lee WG, Wilson JB. Functional richness, functional evenness and functional divergence: the primary components of functional diversity. Oikos. 2005; 111(1):112–18. https://doi.org/10.1111/j.0030-1299.2005.13886.x https://doi.org/10.1111/j.0030-1299.2005... , Villéger et al. (2008)*Villéger S, Mason NWH, Mouillot D. New multidimensional functional diversity indices for a multifaceted framework in functional ecology. Ecology. 2008; 89(8):2290–301. https://www.jstor.org/stable/27650754 https://www.jstor.org/stable/27650754...
Functional Evenness (FEve)	46	Sum of the minimum spanning tree branch length weighted by relative abundance of the two species	*Mason et al. (2005)Mason NWH, Mouillot D, Lee WG, Wilson JB. Functional richness, functional evenness and functional divergence: the primary components of functional diversity. Oikos. 2005; 111(1):112–18. https://doi.org/10.1111/j.0030-1299.2005.13886.x https://doi.org/10.1111/j.0030-1299.2005... , Villéger et al. (2008)*Villéger S, Mason NWH, Mouillot D. New multidimensional functional diversity indices for a multifaceted framework in functional ecology. Ecology. 2008; 89(8):2290–301. https://www.jstor.org/stable/27650754 https://www.jstor.org/stable/27650754...
Functional Divergence (FDiv)	34	Species deviance from the mean distance to the center of gravity weighted by relative abundance	*Mason et al. (2005)Mason NWH, Mouillot D, Lee WG, Wilson JB. Functional richness, functional evenness and functional divergence: the primary components of functional diversity. Oikos. 2005; 111(1):112–18. https://doi.org/10.1111/j.0030-1299.2005.13886.x https://doi.org/10.1111/j.0030-1299.2005... , Villéger et al. (2008)*Villéger S, Mason NWH, Mouillot D. New multidimensional functional diversity indices for a multifaceted framework in functional ecology. Ecology. 2008; 89(8):2290–301. https://www.jstor.org/stable/27650754 https://www.jstor.org/stable/27650754...
Functional Dispersion (FDis)	34	Mean distance in functional space of individual species to the centroid of all species	Laliberté, Legendre (2010)Laliberté E, Legendre P. A distance-based framework for measuring functional diversity from multiple traits. Ecology. 2010; 91(1):299–305. https://www.jstor.org/stable/25661046 https://www.jstor.org/stable/25661046...
Rao (Q)	19	Sum of species distance weighed by abundance	Botta-Dukát (2005)Botta-Dukát Z. Rao’s quadratic entropy as a measure of functional diversity based on multiple traits. J Veg Sci. 2005; 16(5):533–40. https://doi.org/10.1111/j.1654-1103.2005.tb02393.x https://doi.org/10.1111/j.1654-1103.2005... , Ricotta, Moretti (2011)Ricotta C, Moretti M. CWM and Rao’s quadratic diversity: a unified framework for functional ecology. Oecologia. 2011; 167:181–88. https://doi.org/10.1007/s00442-011-1965-5 https://doi.org/10.1007/s00442-011-1965-...
Functional Originality (FOri)	10	Mean distance between each species and its nearest neighbour in the functional space	*Mouillot et al. (2013)*Mouillot D, Graham NAJ, Villéger S, Mason NWH, Bellwood DR. A functional approach reveals community responses to disturbances. Trends Ecol Evol. 2013; 28(3):167–77. https://doi.org/10.1016/j.tree.2012.10.004 https://doi.org/10.1016/j.tree.2012.10.0...
Functional Specialization (FSpe)	8	Mean Euclidean distance between each species and the average position of all species in the functional space	*Mouillot et al. (2013)*Mouillot D, Graham NAJ, Villéger S, Mason NWH, Bellwood DR. A functional approach reveals community responses to disturbances. Trends Ecol Evol. 2013; 28(3):167–77. https://doi.org/10.1016/j.tree.2012.10.004 https://doi.org/10.1016/j.tree.2012.10.0...
Functional Redundancy (FRed)	8	Difference between species diversity (Gini-Simpson diversity index) and Rao; average number of species or as mean abundance or biomass per functional group	*de Bello et al. (2007)Bello F, Lepš J, Lavorel S, Moretti M. Importance of species abundance for assessment of trait composition: an example based on pollinator communities. Commun Ecol. 2007; 8(2):163–70. https://doi.org/10.1556/ComEc.8.2007.2.3 https://doi.org/10.1556/ComEc.8.2007.2.3... , Laliberté et al. (2010)*Laliberté E, Wells JA, DeClerck F, Metcalfe DJ, Catterall CP, Queiroz C et al. Land-use intensification reduces functional redundancy and response diversity in plant communities. Ecol Lett. 2010; 13(1):76–86. https://doi.org/10.1111/j.1461-0248.2009.01403.x https://doi.org/10.1111/j.1461-0248.2009...
Functional Diversity (FD)	3	Sum of the largest branch of the functional dendrogram	Petchey, Gaston (2002)Petchey OL, Gaston KJ. Functional diversity (FD), species richness and community composition. Ecol Lett. 2002; 5(3):402–11. https://doi.org/10.1046/j.1461-0248.2002.00339.x https://doi.org/10.1046/j.1461-0248.2002...
Functional Uniqueness (FUni)	3	Ratio between Rao index and the Simpson diversity index, relating functional diversity to the maximum dissimilarity value of the community	*Ricotta et al. 2016*Ricotta C, Bello F, Moretti M, Caccianiga M, Cerabolini BEL, Pavoine S. Measuring the functional redundancy of biological communities: A quantitative guide. Methods Ecol Evol. 2016; 7(11):1386–95. https://doi.org/10.1111/2041-210X.12604 https://doi.org/10.1111/2041-210X.12604...
Functional Identity	2	Expressed as the biomass-weighted mean trait value for a community	*Mouillot et al. (2011)*Mouillot D, Villéger S, Scherer-Lorenzen M, Mason NWH. Functional structure of biological communities predicts ecosystem multifunctionality. PLoS ONE. 2011; 6(3):e17476. https://doi.org/10.1371/journal.pone.0017476 https://doi.org/10.1371/journal.pone.001...
Functional Vulnerability	2	Sum of the total number of functional entities and the number of species in functional entity	*Mouillot et al. (2014)*Mouillot D, Villeger S, Parravicini V, Kulbicki M, Arias-Gonzáles JE, Bender M et al. Functional over-redundancy and high-functional vulnerability in global fish faunas on tropical reefs. Proc Natl Acad Sci USA. 2014; 111(38):13757–62. https://doi.org/10.1073/pnas.1317625111 https://doi.org/10.1073/pnas.1317625111...
Mean Pairwise Distance (MPD)	2	The average of the distances between pairs of species in the focal community	Webb (2000)Webb CO. Exploring the phylogenetic structure of ecological communities: an example for rain forest trees. Am Nat. 2000; 156(2):145–55. https://doi.org/10.1086/303378 https://doi.org/10.1086/303378...
Mean Nearest Taxon Distance (MNTD)	1	The average of distances between the species of focal community with the respective functionally most similar species (“neighbor closer”)	Webb (2000)Webb CO. Exploring the phylogenetic structure of ecological communities: an example for rain forest trees. Am Nat. 2000; 156(2):145–55. https://doi.org/10.1086/303378 https://doi.org/10.1086/303378...
Functional Regularity Index (FRO)	1	Species evenness in functional space weighted by species abundances	*Mouillot et al. (2005)*Mouillot D, Mason NWH, Dumay O, Wilson JB. Functional regularity: a neglected aspect of functional diversity. Oecologia. 2005; 142:353–59. https://doi.org/10.1007/s00442-004-1744-7 https://doi.org/10.1007/s00442-004-1744-...
Functional Distinctiveness	1	Functional dissimilarity of one species in relation to the other species of the community, representing functional redundancy	*Grenié et al. (2017)*Grenié M, Denelle P, Tucker CM, Munoz F, Violle C. funrar: an R package to characterize functional rarity. Divers Distrib. 2017; 23(12):1365–71. https://doi.org/10.1111/ddi.12629 https://doi.org/10.1111/ddi.12629...