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TITLE
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A compilation of quantitative functional traits for marine and freshwater crustacean zooplankton.
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AUTHORS
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Marie-Pier Hébert1‡*, Beatrix E. Beisner2*, and Roxane Maranger1*
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1
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Montréal, QC, H3C 3J7, Canada
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2
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ville, Montréal, QC, H3C 3P8, Canada
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*Groupe de Recherche Interuniversitaire en Limnologie et en Environnement Aquatique (GRIL)
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‡
Département des Sciences Biologiques, Université de Montréal, C.P. 6128, Succ. Centre-ville,
Department of Biological Sciences, University of Quebec at Montreal, C .P. 8888, Succ. Centre-
[email protected]
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ABSTRACT
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This data compilation synthesizes 8609 individual observations and ranges of 13 traits from 201
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freshwater and 191 marine crustacean taxa belonging to either Copepoda or Cladocera, two
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important zooplankton groups across all major aquatic habitats. Most data were gathered from
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the literature, with the balance being provided by zooplankton ecologists. With the aim of more
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fully assessing zooplankton effects on elemental processes such as nitrogen (N), phosphorus (P)
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and carbon (C) stocks and fluxes in aquatic ecosystems, this data set provides information on the
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following traits: body size (length and mass), trophic group, elemental and biochemical corporal
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composition (N, P, C, lipid and protein content), respiration rates, N- and P-excretion rates, as
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well as stoichiometric ratios. Although relationships for zooplankton metabolism as a function of
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body mass or requirements have been explored in the past three decades, data have not been
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systematically compiled nor examined from an integrative and large-scale perspective across
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crustacean taxa and habitat types. While this contribution likely represents the most
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comprehensive such assembly of traits for both marine and freshwater species, this data set is not
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exhaustive either. As a result, this compilation also identifies knowledge gaps; a fact that should
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encourage researchers to disclose information they may have to help complete such databases.
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This trait matrix is made available for the first time in this data paper; prior to its release, the data
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set has been analyzed in a meta-analysis published as a companion paper (Hébert et al. in press).
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These data should prove extremely valuable for aquatic ecologists for trait-based characterization
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of plankton community structure as well as biogeochemical modeling. These data are also well-
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suited for deriving shortcut relationships that predict more difficult to measure trait values, most
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of which can be directly related to ecosystem properties (i.e., effect traits), from simpler traits
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(e.g., body size), and for exploring patterns of trait variation within and amongst taxonomic units
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or ecosystem types. Overall, this data set is likely to provide new insights into the functional
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structure of zooplankton communities and increase our mechanistic understanding of the
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influence of these pivotal organisms on aquatic ecosystems.
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INTRODUCTION
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Over the past three decades, the importance of individuals in regulating ecosystem processes,
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such as consumer-driven nutrient cycling, has been increasingly recognized. In aquatic
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environments, zooplankton represent a central and key component of food webs, exerting a
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strong influence on other trophic groups and energy fluxes or material. Several species’
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characteristics have been pointed out as being good indicators, or predictors, of the effect of
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zooplankton on their environment, including individual body size, corporal stoichiometry, and
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specific physiological rates (Peters 1983, Ikeda 1985, Ikeda et al. 2001, Sterner and Elser 2002);
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most of such descriptors can also be termed “functional traits”. While the use of traits has
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recently gained popularity amongst aquatic community ecologists (e.g., for zooplankton, Barnett
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et al. 2007, Barnett and Beisner 2007, Merico et al. 2009, Kiørboe 2011, Litchman et al. 2013,
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Barton et al. 2013), few have concretely applied this approach to mechanistically link
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zooplankton community structure to ecosystem functioning (e.g., extrapolate relative effects of
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species traits on ecosystem processes). One obstacle toward developing a fully quantitative trait-
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based assessment of community structure is that many traits require much effort or investment to
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measure, especially for physiological traits such as respiration or excretion. An alternative
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approach to direct trait measurement consists of using published taxon-specific trait values from
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the literature (e.g., Barnett et al. 2007), but much of this trait information is dispersed in different
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journals and/or is published in literature that is less accessible (e.g., older journal volumes,
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undigitized books, institutional reports, Ph.D. theses, etc.).
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Here, we provide a compilation of (mostly) quantitative trait data for crustacean zooplankton
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species from both marine and freshwater habitats, gathering 8609 empirical observations on 392
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different taxa (including 341 copepod and cladoceran species) in a single database. We mostly
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focus on species-level trait values, contributing either directly or indirectly to elemental
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processes, i.e., N, P, and C cycling. Including trait information on species from both freshwater
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and marine ecosystems, this data set is of equal interest to limnologists and oceanographers. The
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wide scope of our trait matrix allows for the quantification of large-scale trait patterns among
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diverse taxa and between habitat types. For instance, in the companion meta-analysis (Hébert et
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al. in press), we describe significant differences in zooplankton allometry and respiration
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between major aquatic ecosystems, with freshwater species having lower length-specific mass
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and three times higher mass-specific respiration rates.
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While the main purpose of this database is to render trait information more readily available to
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aquatic ecologists, this data set is also useful to highlight knowledge gaps in the literature, such
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as an unbalanced research effort relative to trait type or aquatic field (Hébert et al. in press), and
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thus to direct future research efforts. For example, this data set could be used to identify
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underrepresented taxa within Copepoda and Cladocera in the trait literature.
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This metadata file outlines the structure of the data set and provides references for all reported
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observations. In spite of data limitations, this database allows for a more complete assessment of
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trait relationships and trade-offs, promotes novel functional grouping/classification of species,
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and may facilitate a trait-based characterization of community structure for further species-
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ecosystem linkages. The release of this trait compilation, along with its companion meta-analysis
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(Hébert et al. in press), aims to stimulate a search for general patterns amongst zooplankton taxa
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and aquatic habitats, and to provide a better integration of plankton community ecology and
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biogeochemistry through the use of effect traits.
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METADATA
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Class I. Data set descriptors
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A. Data set identity
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Title: A compilation of quantitative functional traits for marine and freshwater
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crustacean zooplankton.
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B. Data set and metadata identification codes
Suggested data set identity codes: zooplankton_traits.csv, references.csv.
C. Data set description
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Investigators: same names and addresses as above.
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Abstract: same as above.
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D. Key words
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Allometry, body size, carbon, corporal composition, effect traits, functional traits,
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metabolism, nitrogen, phosphorus, physiology, stoichiometry, zooplankton.
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Class II. Research origin descriptors
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A. Overall project description
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Identity: Classification and relationships of crustacean zooplankton functional traits:
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linking organisms to ecosystems.
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Originators: same names and addresses as above. Data were culled from the literature by
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Marie-Pier Hébert as part of her graduate studies; this project was part of a Fonds de
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Recherche Québécois sur la Nature et les Technologies (FQRNT)-team grant to RM and
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BB.
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Period of Study: 2012-2013.
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Objective: the overall objective of this study was to promote the integration of
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zooplankton community ecology and aquatic biogeochemistry through the use of traits.
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Using data compiled from a wide variety of literature on crustacean zooplankton traits
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contributing to elemental processes, we first quantified trait relationships amongst taxa
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and between aquatic habitats. The aim was then to provide equations to predict trait
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values to facilitate their use and propose a novel trait classification framework as a tool to
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more readily link zooplankton organisms to the ecosystem processes they are likely to
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influence.
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Abstract: same as above.
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Sources of funding: this project was funded by a Fonds de Recherche Québécois sur la
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Nature et les Technologies (FQRNT)-team grant to RM and BB, the Groupe de
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Recherche Interuniversitaire en Limnologie et en environnement aquatique (GRIL) as
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well as an excellence award from the Faculté des études supérieures et post-doctorales
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de l’Université de Montréal that supported MPH.
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B. Specific subproject description
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Site description: this data set primarily comprises data from peer-reviewed publications
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(i.e., 183 articles and books), spanning a wide array of taxa from both marine and
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freshwater habitats. It also contains information from gray literature, such as institutional
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reports or theses, and data provided by colleagues (acknowledged below). In few cases,
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inferences on the basis of knowledge on similar species were performed, e.g., for body
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size of the same genus. In particular, this was done for some species of Bosmina,
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Chydorus, Holopedium, and Skistodiaptomus (length and dry mass); and certain species
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of Alona, Daphnia, Diaphanosoma, and Tropocyclops (for dry mass only).
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Experimental/Sampling design: all data were obtained from the literature (articles,
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books, theses) and zooplankton ecologists. Research methods are outlined below;
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however, details on the statistical analysis of this data set as well as the emerging results
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form the core of a companion manuscript (Hébert et al. in press).
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Research methods: Web of Science (1945-2013) and Google Scholar databases were
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searched in 2012 to find data sources on zooplankton nitrogen and phosphorus content
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and excretion rates. Two separate searches were conducted including the following
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keywords: (i) zooplankton* AND (nitrogen* OR ammonia* OR phosphorus* OR
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phosphate*) AND excretion*, (ii) zooplankton* AND (elemental* OR nitrogen* OR
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phosphorus* OR lipid* OR protein*) AND body* AND composition*. Both databases
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were revisited in 2013 for information on zooplankton carbon content and respiration
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rates, using two combinations of search terms: (i) zooplankton* AND respiration*, (ii)
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zooplankton* AND (elemental* OR carbon*) AND body* AND composition*.
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The Abstract, Results, and when necessary, the full text of each publication were then
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manually searched to decide whether the study matched our selection criteria. References
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cited in these articles were also checked and considered in our literature search. We only
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selected studies that (i) included information for crustacean meso- and
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macrozooplankton, i.e., essentially copepods and cladocerans as these are the most
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studied groups (ostracods, mysids, amphipods, euphausiids were not included), (ii)
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focused on adult individuals of female sex (few observations on male individuals were
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retained but were marked accordingly), and (iii) provided species’ dry mass data from
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which we could estimate individual-level excretion and respiration rates (or, inversely,
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estimate mass-specific rates), and body composition as % elemental proportions of unit
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dry mass. When provided, trophic groups (i.e., preferred diet) of species were recorded.
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To minimize variance and potential bias, we focused on respiration rates expressed in
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volume of oxygen (O2) given the difficulty in unit conversion (i.e., mol versus litre) when
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environmental parameters are not provided (e.g., ambient pressure).
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Ambient or experimental temperatures in studies of zooplankton metabolism (i.e., for
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respiration and excretion rates) were recorded so as to further apply a standardized
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temperature correction. Data were sometimes extracted directly from figures in articles or
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reference books using the software Datathief III, (version 1.6, Bas Tummers ©).
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Furthermore, this trait compilation includes a sub-data set containing body size estimates
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and trophic groups for freshwater crustacean zooplankton species; the latter represents an
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extension to Barnett et al. (2007) initially elaborated upon during a workshop of the
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Canadian Institute of Ecology and Evolution (CIEE). The original estimates of body size
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in this sub-data set were enhanced by the inclusion of data from several North American
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datasets including the North Temperate Lakes Long Term Ecological Research (NTL-
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LTER) site in Wisconsin, U.S.A., the Dorset Environmental Research Centre (DESC),
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and data from the Experimental Lakes Area, Canadian Department of Fisheries and
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Oceans (DFO) until 2012 and now part of the International Institute for Sustainable
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Development (IISD-ELA). Dry mass estimates were based on taxon-specific length-mass
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allometric equations (McCauley 1984; Culver et al. 1985).
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Finally, details and description of the methods used to measure respiration rates provided
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by colleagues G. Darnis and L. Fortier is described in Darnis and Fortier (2012); methods
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used by T. Ikeda (unpublished data) to measure respiration and excretion rates are
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outlined in Omori and Ikeda (1984) and Ikeda et al. (2000).
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Class III. Data set status and accessibility
A. Status
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Latest updates: the formal literature search for trait information ended in December
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2013. In 2014-2015, the following information have been added to the data set: 58 values
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of total body lipid in marine species (data extracted from the references of Lee et al. 2006
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and Vijverberg and Frank 1976), total body lipid and protein content for 14 freshwater
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species (data extracted from the references of Vijverberg and Frank 1976, and Riccardi
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and Mangoni 1999), over 100 body size observations (from diverse data sources), and
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trophic group information for 59 marine species (most of which were borrowed from a
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trait compilation available through Pomerleau et al. 2015).
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Latest archive date: current.
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Metadata status: current.
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Data Verification: data were evaluated and double-checked for accuracy. Information
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outside normal operating ranges were checked and compared to the original data
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provenance. We outlined methods for treatment of data and outliers in the manuscript that
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reviews and evaluates this data compilation (Hébert et al. in press).
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B. Accessibility
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191
Storage location and medium: original data files are held by the authors. The data set
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published on Ecological Archives is the first release from this compilation of crustacean
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zooplankton quantitative functional traits.
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Contact person: queries about the entire data set or individual specific trait values can be
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directed to Marie-Pier Hébert, email: [email protected].
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Copyright and proprietary restrictions: none. When using the dataset, we kindly
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request that you cite this article, recognizing the work that went into gathering the data
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together and the original authors’ willingness to make it publicly available.
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Costs: none.
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Class IV. Data structural descriptors
A. (1) Data set file
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Identity: zooplankton_traits.csv
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Size: 1199 lines of data, excluding header row. Note that the number of individual
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observations does not equal to the number of species, where the number of observations
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for a single trait vary between 1 and 72 for a given species. As a result, when more than
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one trait estimate was available, individual data lines were numbered accordingly, with
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replicate numbers listed in a separate column (see section B). Overall, this data set
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contains 8609 individual observations and ranges (i.e., group or habitat type excluded) on
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201 freshwater and 191 marine crustacean zooplankton taxa.
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Format and storage mode: comma-delimited, no compression.
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Header information: headers are given as header name (e.g., type of trait). For each trait
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value found in the literature, the reference numerical codes are indicated in the following
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column (see section B for the description of trait abbreviation inserted in reference
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column names). Complete references and their corresponding reference numerical codes
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are provided in Class V, and the information on authors, year, title, journal, and
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publication type is also provided in a separate .csv file to facilitate the use of our
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reference list. For trait values expressed as ranges, the maximum and minimum values
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are indicated in the two (max., min.) columns following the column for mean trait values
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(see section B for trait abbreviation). For physiological traits, i.e., excretion and
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respiration rates, the recorded temperatures are also indicated in the following column.
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Alphanumeric attributes: mixed.
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Special characters/fields: none.
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(2) Reference file
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Identity: references.csv
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Size: 198 lines of reference data, excluding header row.
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Format and storage mode: semicolon-delimited, no compression.
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Header information: headers are given as header name, with separate columns for
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reference numerical code, authors, year, title of the publication (e.g. article, book
229
chapter), name of the journal or book, and publication type. The complete bibliographic
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information (e.g., volume, pages, edition) is however provided in Class V.
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Alphanumeric attributes: mixed.
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Special characters/fields: none.
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B. Variable definitions
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(1) Data set file
Column
name
Genus
Variable definition
Units
Scientific name (genus)
N/A
Storage
type
Character
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Species
Replicate.num
ber
Group
Habitat
Trophic.group
Body.length
Dry.mass
Cct.prop
Cct.total
Nct.prop
Nct.total
Pct.prop
Pct.total
N:Pct
Protein.ct
Lipid.ct
Respiration.rt.
ind
Respiration.rt.
mg
N-ammonia.
ex.rt.ug.ind
N-ammonia.
ex.rt.ug.dm
Scientific name (species);
“-(M)” denotes the few male
individuals.
Number of individual investigated
for a given trait, when N > 1
(ranging from 1 to 72).
Taxonomically classified as:
Copepoda;
Cladocera.
Freshwater habitat;
Marine habitat.
Preferred diet known as:
Carnivore;
Herbivore;
Omnivore;
Omnivore-Carnivore (OmniCarn);
Omnivore-Herbivore (OmniHerb).
Individual mean body length.
Individual mean body dry mass.
Mean proportion of C corporal
content.
Mean amount of C corporal
content.
Mean proportion of N corporal
content.
Mean amount of N corporal
content.
Mean proportion of P corporal
content.
Mean amount of P corporal
content.
Mean N:P molar ratio in corporal
content.
Mean proportion of protein in
corporal content.
Mean proportion of lipid in
corporal content.
Mean individual and mass-specific
respiration rate.
N/A
Character
N/A
Numeric
N/A
Character
N/A
Character
N/A
Character
mm
mg
%
Numeric
Numeric
Numeric
mg
Numeric
%
Numeric
mg
Numeric
%
Numeric
mg
Numeric
N/A
Numeric
%
Numeric
%
Numeric
µl O2
ind-1 h-1
µl O2
mgDM -1 h-1
Mean individual and mass-specific µg N-NH4+
ammonia (N-NH4+) excretion rate; ind-1 h-1
“ND” indicates values under
µg N-NH4+
detectable thresholds*.
mgDM-1 h-1
Numeric
Numeric
Numeric
Alphanumeric*
12
N-ammonia.
ex.rt.nmol.dm
N-ammonia.
ex.rt.nmol.ind
P-phosphate.
ex.rt.ug.ind
P-phosphate.
ex.rt.ug.dm
P-phosphate.
ex.rt.nmol.dm
P-phosphate.
ex.rt.nmol.ind
N:P.ex
Temp.trait
Ref.trait
Max.trait
nmol N-NH4+
mg DM-1 h-1
nmol N-NH4+
ind-1 h-1
Mean individual and mass-specific µg P-PO43phosphate (P-PO43-) excretion rate. ind-1 h-1
ug P-PO43- mg
DM-1 h-1
nmol P-PO43mg DM-1 h-1
nmol P-PO43ind-1 h-1
Numeric
Mean N:P molar ratio in
excretion;
“ND” indicates values under
detectable thresholds*.
Recorded temperature for
respiration (.resp.rt) and excretion
(.Nex.rt and .Pex.rt) data;
“RT” indicates measurements at
room temperature‡.
Corresponding reference
numerical code.
Trait abbreviations in column
names refer to the following traits:
.tg = trophic group;
.bl = body length;
.dm = dry mass;
.C = C corporal content;
.N = N corporal content;
.P = P corporal content;
.NP = N:P body ratio;
.prot = protein content;
.lip = lipid content;
.resp = respiration rate;
.N.ex = N excretion rate;
.P.ex = P excretion rate;
.NP.ex = N:P excretion ratio.
Maximum value for a given trait;
Trait abbreviations in column
names are listed above, with the
addition of:
.Cprop = proportion of C content;
.Nprop = proportion of N content;
N/A
Alphanumeric*
Celcius
Alphanumeric‡
N/A
Numeric
Subject to
variation
Numeric
Numeric
Numeric
Numeric
Numeric
Numeric
13
Min.trait
235
236
237
238
Subject to
variation
Numeric
Variable definition
Units
Corresponding reference
numerical code
Authors of the publication or data
source.
Year of publication, or
consultation for online data base
or unpublished data.
Title of the publication, book
chapter, or data set provided.
Name of the journal or book, if
published data.
Type of publication:
Article;
Thesis;
Book or book chapter;
Institutional report, bulletin or
memoirs;
Open access online repository;
Unpublished data.
N/A
Storage
type
Numeric
N/A
Character
N/A
Numeric
N/A
Character
N/A
Character
N/A
Character
Abbreviations:
DM: Dry mass; Ind: individual
(2) Reference file
Column
name
Ref.code
Authors
Year
Title
Journal.Book
Pub.type
239
240
.Pprop = proportion of P content;
.resp.mg = mass-specific
respiration rate;
.Nex.rt.ug.dm =
mass-specific ugN excretion rate;
.Nex.rt.nmol.ind =
individual nmolN excretion rate;
.Pex.rt.ug.dm =
mass-specific ugP excretion rate;
.Pex.rt.nmol.ind =
individual nmolP excretion rate.
Minimum value for a given trait;
Trait abbreviations in column
names: as above.
C. Data limitations
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241
Because of the large information gaps and data heterogeneity in the diverse forms of
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literature, this data set comprises several limitations that may restrict the scope of further
243
analyses. In addition to the limited number of empirical observations for some of the
244
targeted traits, or less-studied cladoceran and copepod species, there was a lack of
245
stoichiometric information on food supply. This precluded the inclusion of food supply
246
stoichiometry in the data set, although we recognize the relevance of this environmental
247
driver in the assessment of zooplankton N and P excretion. Furthermore, although we
248
included studies from a wide range of marine and freshwater environments, most studies
249
retained were from temperate regions, resulting in an unbalanced spatial coverage. There
250
was a lack of specific information regarding the geographic origins of collected
251
individuals in studies involving lab experiments. As a result, the only level of
252
geographical resolution, or habitat type, that could be reasonably assumed for all species
253
included in the data set was the classification between marine and freshwater habitats.
254
Another limitation important to acknowledge is the diversity of research methods used to
255
measure traits (such as body composition or metabolic rates) among studies, introducing
256
potential sources of variation (i.e., noisy data). The variance or standard error were often
257
not provided with the trait values reported, making it impossible to weight the estimates
258
by the number of observations for further statistical analyses. Although these sources of
259
variation may sometimes limit the possibilities in terms of data analyses, we still explored
260
and quantified trait relationships among traits by giving equal weight to all trait values in
261
the meta-analysis accompanying this data set (Hébert et al. in press). Finally, we are
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aware that more information on species body size, carbon content and trophic groups are
263
available in the literature; however, it should be noted that the focus of Hébert et al. (in
15
264
press) was initially on zooplankton excretion, with respect to body composition in terms
265
of nitrogen and phosphorus, and thus there was an unequal effort given to some non-focal
266
traits. This data set is expansive, but other traits and species could be added in the future;
267
for example, the inclusion of other crustacean zooplankton groups such as ostracods for
268
which there is likely less information in the trait literature. Nevertheless, we hope that
269
this start will stimulate the use of a trait-based approach to characterize community
270
structure influences on ecosystems, explorations of general patterns across ecosystem
271
types and trade-offs among species traits, prompting for more formal linkages between
272
effect traits and ecosystem function.
273
Class V. Data set references
Reference
numerical Complete reference
code
Canadian Institute of Ecology and Evolution (CIEE) working group on
zooplankton dynamics. 2012. Freshwater zooplankton database. Unpublished
1
data.
Lynch, M. 1980. The evolution of cladoceran life histories. Quarterly Review of
2
Biology 55:23–42.
Bottrell, H. H., Duncan, A., Gliwicz, Z. M., Grygierek, E., Herzig, A., HillbrichtIlkowska, A., Kurasawa, H., Larsson, P., and T. Weglenska. 1976. A review of
some problems in zooplankton production studies. Norwegian Journal of Zoology
3
24:419–456.
Tümpling, W. V., and G. Friedrich. 1999. Biologische Gewässeruntersuchung in
4
Methoden der Biologischen Wasseruntersuchung II, Gustav Fischer:352–360.
Adrian, R. 1988. Untersuchungen zur herbivore und carnivoren ernährungsweise
von Cyclops Kolensis und C. Vicinus (Crustacea, Copepoda) Ph.D. Thesis,
5
Fachbereich Biologie, Freie Universitat Berlin, Berlin (GER).
Dumont, H. J., de Velde, I. V., and S. Dumont. 1975. The dry weight estimate of
biomass in a selection of Cladocera, Copepoda and Rotifera from the plankton,
6
periphyton and benthos of continental waters. Oecologia 19:75–97.
Berne, T. V. 1991. Abundance, standing crop and production of microcrustacean
populations (Cladocera, Copepoda) in the littoral zone of Lake Biel, Switzerland.
7
Archiv für Hydrobiologie 123:165–185.
Frey, D. G. 1973. Comparative morphology and biology of three species of
8
Eurycercus (Chrydoridae, Cladocera) with a description of Eurycercus
16
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
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macrocanthus sp. Internationale Revue der gesamten Hydrobiologie 58:221–267.
Smirnov, N. N. 1964. Eurycercus lamellatus (chydoridae: Cladocera): Field
observations and nutrition. Hydrobiologia 20:280–294.
Green, J. 1956. Growth, size, and reproduction in Daphnia (Crustacea: Cladocera).
Journal of the Zoological society of London 126:173–204.
Burgis, M. J. 1973. Observations on the Cladocera of Lake George, Uganda.
Journal of the Zoological society of London 170:339–349.
Murugan, N., and K. G. Sivaramakrishnan. 1973. The biology of Simocephalus
acutirostratus King (Cladocera: Daphnidae) – Laboratory studies of life span,
instar duration, egg production, growth, and stages in embryonic development.
Freshwater Biology 3:77–83.
Branstrator, D. K. 2005. Contrasting life histories of the predatory cladocerans
Leptodora kindtii and Bythotrephes longimanus. Journal of Plankton Research
27:569–585.
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ACKNOWLEDGMENTS
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In addition to the aforementioned funding sources, we acknowledge discussion on zooplankton
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size and trophic groupings with the working group on zooplankton dynamics at the CIEE, and
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especially thank the Ontario Ministry of Environment - Dorset Environmental Research Centre
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(DESC), the North Temperate Lakes Long Term Ecological Research site (NTL-LTER), as well
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as the International Institute for Sustainable Development – Experimental Lakes Area (IISD-
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ELA) for contributing information integrated to calculate some freshwater zooplankton body
282
lengths. We thank M. Kainz for his willingness to provide data on zooplankton lipid composition
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as well as G. Darnis and L. Fortier for sharing their data on zooplankton respiration. We also
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wish to acknowledge Elise Lavieville, Gael Lainé-Panet, and Akash Sastri from the Beisner
285
laboratory for their help in assembling Pacific zooplankton traits. Lastly, we are grateful to all
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zooplankton ecologists making their data publicly available in open access, especially T. Ikeda
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for sharing his extensive work conducted over the past four decades. This data paper is part of a
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contribution to the Groupe de Recherche Interuniversitaire en Limnologie et en Environnement
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Aquatique (GRIL).
290
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