Freshwater Biology (2007) 52, 796–813 doi:10.1111/j.1365-2427.2007.01733.

Functional diversity of crustacean zooplankton

communities: towards a trait-based classification
ALLAIN J. BARNETT, KERRI FINLAY AND BEATRIX E. BEISNER
Department of Biological Sciences, University of Quebec at Montreal, Montreal, QC, Canada

SU M M A R Y
1. While studies of phytoplankton and terrestrial plant communities have increasingly
emphasised the use of functional traits in ecological research, few have yet to apply this
approach to zooplankton communities.
2. This study reviews laboratory and observational studies on zooplankton feeding and life
history and provides a series of functional trait tables for the North American freshwater
zooplankton. Qualitative and quantitative trait tables highlight areas where data were
more scarce and point to which types of studies could fill in gaps in our knowledge of
zooplankton niches.
3. Data were most complete for the Cladocera across most traits, while feeding information
for cyclopoids was most sparse. Qualitative data that distinguished congeneric species
were lacking for most groups.
4. A regional community dendrogram for common north-eastern North American
zooplankton species was generated and shows that taxonomic differences between species
do not capture fully functional differences based on the traits of body length, habitat,
trophic group and feeding type.
5. The data collected here, combined with readily measurable species attributes, can be
used to generate a multivariate measure of the functional niche of each species found in a
community. Armed with this information, functional relationships that are useful for
ecological studies of lake ecosystems can be more easily conducted.

Keywords: biodiversity, classification, functional diversity, niche, traits

simply the number of taxonomic species (Walker,

Introduction
The human proclivity towards the classification of
nature has led to the definition and cataloguing of
species in communities based traditionally on mor-
phological and more recently, on genetic descriptors.
While this taxonomic classification is essential, it may
not be the most appropriate scheme for many ecolog-
ical applications. For example, many studies on the
relationship between biodiversity and ecosystem
function have concluded that it is the ecological roles
of the species present that are important, and not
Correspondence: Beatrix E. Beisner, Department of Biological
Sciences, University of Quebec at Montreal, CP 8888, Succ.
Centre-Ville, Montreal, QC, Canada H3C 3P8.
E-mail: beisner.beatrix@uqam.ca
796  2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd
1991; Hooper & Vitousek, 1997; Tilman et al., 1997;
Walker, Kinzig & Langridge, 1999; Symstad, Siemann
& Haarstad, 2000). Thus, there have been recent calls
for the use of functional diversity measures – descrip-
tors of communities based on a new type of classifi-
cation scheme that is applicable to many ecological
questions (Petchey & Gaston, 2002; Norberg, 2004;
Mason et al., 2005; Mouillot et al., 2005) and inspired
by previous approaches (e.g. Grime, 1977; Reynolds,
1980) including the niche concept.
Functional diversity is a biodiversity measure based
on functional traits of the species present in a
community. Functional traits are those that define
species in terms of their ecological roles – how they
interact with the environment and with other species
(Diaz & Cabido, 2001). In phytoplankton, for example,

these traits usually include body size, tolerance and
sensitivity to environmental conditions, motility,
shape, N-fixation ability and Si requirements (Rey-
nolds et al., 2002; Weithoff, 2003). In terrestrial plant
communities, researchers have included more com-
plex traits like rates of growth, nutrient requirements
and water uptake (Walker & Langridge, 2002).
Similarly, in stream environments, a functional
approach to invertebrate communities have been
invoked for examining diversity patterns and for
biomonitoring purposes (e.g. Statzner et al., 2001;
Bady et al., 2005).
There are several methods for calculating functional
diversity of a community. Recent methods include
Functional Attribute Diversity, as used in a study of
Australian rangelands by Walker et al. (1999), and
Functional Diversity (FD) proposed more recently by
Petchey & Gaston (2002). Trait variance, measured as
the width of a trait distribution, has been proposed by
Norberg (2004). Beyond the simple measurement of
diversity, Mason et al. (2005) proposed also estimating
functional richness, functional evenness and func-
tional divergence, to enable descriptions of niche use
and competitive interactions in communities. In order
to take a functional approach and to use these new
measures, however, we must have descriptors of the
functional groups present in a community.
The effects of environmental changes on ecosystem
properties are thought to be largely determined by the
range of biological processes carried out by the suite
of species present in the community (Reynolds et al.,
2002). Studies that focus on whether biodiversity
affects the functioning of an ecosystem (e.g. produc-
tivity, total biomass) have yielded results that suggest
a link between the functional roles of species and
community responses to environmental changes. For
example, Tilman et al. (1997) found that measures of
functional diversity were more strongly related to
ecosystem response variables (e.g. productivity, soil
NO3, soil NH4) in a plant community than was
traditional species diversity. In another study, Walker
et al. (1999) observed that minor, functionally redund-
ant species in ungrazed grasslands increased in
abundance in heavily grazed communities, providing
support for the insurance hypothesis (McNaughton,
1977; Yachi & Loreau, 1999), whereby seemingly
redundant species contribute to ecosystem resilience
when environmental conditions change. Functional
traits of species describe how they respond to the
 2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd, Freshwater Biology, 52, 796–813
Functional diversity of zooplankton communities 797
environment, and if we know what functional traits
are present in a community, then we should be better
able to predict how communities will shift as envi-
ronmental conditions change and thereby understand
the role of biodiversity more fully (Norberg, 2004).
A functional classification of species should greatly
enhance our ability to predict how the pulsed and
gradual changes of environmental conditions will
affect communities. For terrestrial plant species, a
classification strategy has recently been proposed to
predict changes in vegetation in response to environ-
mental change (Lavorel, McIntyre & Landsberg, 1997).
A similar strategy has also been proposed for
phytoplankton to be applied to ecological questions
such as the intermediate disturbance hypothesis,
insurance hypothesis and alternative stable state shifts
in lakes (Reynolds et al., 2002; Weithoff, 2003). For
example, the intermediate disturbance hypothesis
predicts the highest diversity of species at intermedi-
ate magnitude or frequency of disturbance by per-
mitting species with different strategies and
competitive advantages to coexist (Connell, 1978).
Weithoff (2003) asserts that this hypothesis relies on
the assumption that species are functionally dissimilar
in response to disturbance. For a greater understand-
ing of the catastrophic shift with eutrophication from
a vegetation dominated clear state to an alga domin-
ated turbid one, a functional approach can be used to
determine the gradual changes in attributes of the
algal community in response to nutrient addition,
leading up to a shift to an alternative state (Weithoff,
2003). Finally, the use of functional diversity for
testing the ‘insurance hypothesis’ provides insight
into the potential ‘roles’ of seemingly minor or
redundant species in lake communities (Weithoff,
2003) as well as in terrestrial ones (Walker et al., 1999).
Therefore, the use of functional diversity enhances
our understanding of these important ecological
phenomena.
In aquatic environments, zooplankton play an
important role in structuring phytoplankton commu-
nities (McCauley & Briand, 1979; Sterner, 1989), and in
mediating energy flow to higher trophic levels in
pelagic habitats (Gliwicz & Pijanowska, 1989). Until
now, no thorough examination of the potential for
functional classification of zooplankton has been
attempted. We explore the possibility of determining
functional grouping strategies for zooplankton using
a review of the literature to determine for which
798 A.J. Barnett et al.
groups and traits knowledge exists and for which
information is still lacking. We develop here, a
functional classification tree for common North
American zooplankton and show that taxonomic
relationships are not always sufficient surrogates for
questions of function. The data reviewed and collec-
ted here will be useful both to expand our under-
standing of traits of zooplankton, and to determine
where future studies can focus their efforts in order to
achieve a more robust classification method of func-
tional diversity to help predict ecosystem function in
lakes.
Methods
We conducted an extensive search of the primary
literature for freshwater zooplankton functional traits.
Emphasis was on studies that focused on traits that
related to body size, feeding, growth and predator
avoidance. These traits were chosen because they
describe both an organism’s response to environmental
conditions (e.g. body size, life history, predator
avoidance, food availability), and their potential effects
on ecosystem processes (e.g. feeding rate).
The investigation focused on traits for zooplankton
species found in north-eastern North America, where
our research on plankton communities is based.
However traits for other common North American
species were included in the final analysis. Searches
were initiated using online databases: Biological
Sciences (1960–2006) and ISI’s Web of Science (1981–
2006) using keywords ‘freshwater’, ‘zooplankton’,
‘body size’, ‘feeding’, ‘growth’, ‘development’ or
simply the names of each species. The reference
sections of papers found in this manner were also
used to search for further data in older publications.
The average adult body length of each species
included in the trait table were obtained wherever
possible from published studies. When only ranges of
body sizes were found, we present the mid-point of
this range in the results tables. For values that we were
unable to obtain from published studies, we included
our own mean values from zooplankton communities
in 18 lakes in the Eastern Townships of Quebec,
Canada. Standard length measurements of zooplank-
ton (McCauley, 1984) were taken on a minimum of 10
individuals from each lake in July 2004 using an
Olympus dissection microscope and Evolution Qei
Monochrome camera using ImagePro software under
 2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd, Freshwater Biology, 52, 796–813
20· or 32· magnification. The overall mean across all
lakes for each species was calculated.
Only growth rates, feeding rates and feeding rate
regressions with body length that were measured in
published laboratory studies at temperatures between
15 and 25 C, were included in results tables as they
reflect mean summer lake temperatures in this region
of North America. For copepods, we included only
rates for adult females. Information found in the text
of studies that provided qualitative information, such
as habitat or food preferences, or presence/absence of
a defensive strategy was also included. All data were
compiled into trait tables and separated into quanti-
tative and qualitative data tables.
Community dendrogram
Qualitative and quantitative traits that were found for
a sufficient number of species (length, habitat, trophic
group and feeding type) were used to generate a
functional dendrogram for the regional species pool of
north-eastern North American lake zooplankton. In
addition to the full functional dendrogram based on
the four characteristics, a functional dendrogram
using only body length was generated for comparison
of what information can be gleaned using a very
limited estimate of species functional traits. Qualitative
measures were entered as ranked categories (i.e. from
herbivore to carnivore, and from more passive forms
of filtering type through to raptorial feeding). The
dendrogram of species for which sufficient traits
existed was generated using hierarchical clustering
analysis from standardised Euclidean distances as in
the method of FD calculation proposed by Petchey &
Gaston (2002). These dendrograms were generated
using average linkage clustering and unweighted pair
grouping (Krebs, 1999) using the statistical computer
program P R I M E R (Clarke & Gorley, 2001). FD can be
considered as a quantitative measure of the richness
of functional traits, or the degree of complementarity
within a community (Petchey & Gaston, 2002). FD for
a particular lake community of interest (containing a
subset of species presented) would be the sum of
branch lengths in the dendrogram of the regional
species pool calculated herein. Dendrograms were
compared with a simplified taxonomic tree for the
regional species pool that was constructed according
to major family and class designations for each species
according to Thorp & Covich (2001).

Results
The life history information presented was collected
from 126 sources spanning Frank (1952) to Frost et al.
(2006). The majority of references are from the primary
peer-reviewed literature, while some (particularly
qualitative) data were taken from the Great Lakes
Copepod key of the USGS Great Lakes Science Center
(http://www.glsc.usgs.gov/greatlakescopepods), and
other taxonomic keys (e.g. Pennak, 1989; Hebert, 1995;
Witty, 2004). Only traits for which data on more than
25% of all species were found were included. A
complete list of the primary literature used to generate
trait tables is given in Supplementary material.
Quantitative traits defined
Quantitative traits are listed in Tables 1 and 2. Two
measures of feeding were included in the trait table
for both copepods and cladocerans: food size range
(lm) and maximum clearance rate (lL ind.)1 h)1).
Mesh size (lm) and regressions of clearance rate with
body size were sufficiently abundant in the literature
on cladocerans to warrant inclusion.
Food size range represents the smallest and largest
particles consumed by each species calculated over all
cited studies. For non-spherical particles, sizes were
standardised as equivalent spherical diameter (lm
ESD) using the equation (length · width)0.5. The
studies used to calculate the range of food size
included only those that measured a feeding rate
explicitly, and excluded those that reported food size
in relation to some other measurement, such as
generation time. Whenever possible, values were
from studies that assessed preferred food sizes across
several species of zooplankton – the best way to
collect such comparative data. Further, for Cladocera,
the mesh size of the feeding appendages provided an
alternative measure of the size of the smallest particle
an organism is capable of capturing and this was also
recorded in Table 1.
Clearance rate is the volume of water swept clear
per individual zooplankter per unit time. This value
often remains high and constant at lower food
concentrations and then declines with increasing food
concentrations (Sterner, 1989; Mauchline, 1998). When
multiple clearance rate values were reported in a
single study, we included only the rate at the lowest
food concentration in order to estimate maximum
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Functional diversity of zooplankton communities 799
values better. Maximum values from each study are
reported in Table 1. Because cladocerans do not
undergo metamorphosis, it is often unclear for what
stage a clearance rate is being measured. Moreover, it
has been demonstrated that clearance rate in cladoc-
erans is affected by body size. For this reason, we also
included clearance rate versus body length regression
relationships in Table 1. For copepods, clearance rates
for adult females were used.
The measures of growth included in the trait table
are generation time for both cladocerans and cope-
pods, and rate of population increase (r) for cladoc-
erans. Generation time is defined as the number of
days from hatching until moult into the adult for
copepods, and from hatching until production of the
first clutch in cladocerans. Generation time can vary
with temperature and food concentration, so we
preferentially selected maximum generation times
determined at 20 C, exceptions are noted in the
Tables. The rate of population increase is typically
measured using either life tables or by population
growth studies, and results from both types were
included in the cladoceran trait table as separate
columns.
Stoichiometric ratios of C : P and C : N as meas-
ured for individual species of zooplankton were also
included in the trait table, as these values will affect
nutrient excretion rate and in general capture proces-
ses of material flow and cycling through the zooplank-
ton.
Quantitative data results
Data for most species were obtained for mean body
length, food size range and clearance rate, but were
lacking for calanoid generation time and for stoi-
chiometric ratios of many species. Food size range
tended to vary with major taxonomic grouping
since, for example, raptorially feeding cylopoids
generally fed on much larger food items. Cladocer-
ans, which generally filter food, fed on the smallest
food particles.
A large range in clearance rate was observed both
between and within species. The lowest observed
clearance rate was for the small herbivorous cladoceran
Bosmina longirostris, with a value of 13 lL ind.)1 day)1,
while the highest was observed for the carnivorous
cladoceran Polyphemus pediculus, at 20 000 lL ind.)1
day)1. Between studies on the same species, Cyclops
 Table 1 Quantitative functional traits of cladoceran zooplankton including sources in superscript numbers (see Supplementary material). Max r is the rate of population increase.
800

Parameters y and x in the regression equations represent clearance rate and body size (mm), respectively. Body lengths marked with an asterisk (*) are mid-points of the range
observed and those without a reference are personal observations of mean size.

Mean Food Max Regression Mesh size Fastest Max r

body size clearance CR versus (setulae generation Max r population
length range rate length (CR in distances) time life table growth expts
Species (mm) (lm) (lL ind.)1h)1) mL ind.)1h)1) (lm) (days) (day)1) (day)1) C : P[30] C : N[30]
[1]
Alona affinis 0.8 * 56[2] 17[32]‡
A.J. Barnett et al.

Leydig
Alona rectangula 10[33]§ 0.34[33] 0.28[34], 0.07[35]
Sars
[3]
Eubosmina corregoni 0.6 * 1.4–5[4] 0.68–1.6[4]
Baird
Bosmina longispina 0.44 196[6] 1.1–1.4[6] 0.32[36]a 156.0[37] 6.1[37]
Leydig
[7]
Bosmina longirostris 0.4 13[8]a, 40[9]b, y ¼ 0.487·1.83[15] 0.4–0.7[12] 6[1] 0.24[38], 0.29[39]– 0.23[41]†– 140.4[42))44] 10.0[42))44]
Müller 125[10], 146[11]b, 0.31[40]
150[12], 170[13],
190[14], 200[15]a,
230[16], 300[17]
Ceriodaphnia dubia 0.8[7] 18[45]b 0.27[45]b 0.27[34]
Richard
Ceriodaphnia 0.8[7] 0.2–0.6[12] 6[1] 0.44[46], 0.58[47]b–
reticulata
Jurine
Ceriodaphnia lacustris 0.85[7] 0.4–7[4] 106[10,11] 0.24–0.4[4]
O.F.M. 188[18], 321[9]
[1]
Chydorus sphaericus 0.46 * 0.4–2[4] 13[12], 131[2] 0.24–0.4[4] 5[1], 9[32]‡ 0.22[39]b–, 0.23[38], 0.28[36]a
Müller 0.27[48]b–
Chydorus brevilabris 0.29
Frey
Daphnia ambigua 0.88[7] y ¼ 0.41·3.63[19] 7[1], 8[49]‡ 0.33[39]–
Scourfield
[3]
Daphnia catawba 1.7 * y ¼ 0.085·1.74[20] 0.23–0.45[4]
Coker
[21]
Daphnia galeata 2 * 1.1–20[4]†† 258[22]a, 1375[23], y ¼ 0.26·1.58[20] 0.32–1.0[4]††, 7[1], 12[50] 0.14[38], 0.2[50], 97.5[31,55))57] 6.9[31,55))57]
mendotae Sars 1650[24]** 0.4–0.7[12], 0.29[51], 0.32[52],
0.42[24] 0.35[53,54]–,
0.37[39]b–
[1]
Daphnia 1.67 * 350[24], 688[6], 0.5–0.9[6], 7.2[1] 0.36[58], 0.38[59] 82.6[37,42] 6.1[37,42]
longispina 820[25] 0.38[24]
Müller

 2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd, Freshwater Biology, 52, 796–813
 Table 1 (Continued)

Mean Food Max Regression Mesh size Fastest Max r

body size clearance CR versus (setulae generation Max r population
length range rate length (CR in distances) time life table growth expts
Species (mm) (lm) (lL ind.)1h)1) mL ind.)1h)1) (lm) (days) (day)1) (day)1) C : P[30] C : N[30]
[21]
Daphnia magna 3.5 * 0.6–40[4] 2350[24], 3560[26] y ¼ 0.28·2.2[27]‡ 0.24–0.64[4], 6[60], 8[1], 0.25[38], 0.40[62], 0.29[60] 92.7[55,57,65,66] 5.9[55,57,65,66]
[63]– [64]
Straus 0.46[24] 16[61]‡ 0.44 , 0.58
Daphnia pulex 1.75[7] 326[28], 570[29], y ¼ 0.42·1.61[20], 0.40[24] 6.5[1], 7[67], 0.29[69]–, 0.35[38], 0.25[73] 106.0[74] 7.3[74]
Leydig 1920[24] y ¼ 0.41x3.37[19] 8[68] 0.4[70]b–, 0.50[71],
0.51[72]
Daphnia pulicaria 1.75 1.5–30[4] 2080[24], 2483[23] y ¼ 0.39·3.46[19] 0.45–1.4[4], 5[50] 0.33[50,75]– 154.0[55] 5.7[55]
Forbes 0.4–0.7[12], 0.47[24]
Daphnia rosea 1.4[7] 267[22]a, 867[9], y ¼ 0.23·2.74[15] 0.4–0.7[12] 6[76] 0.046[77], 123.5[31,78] 7.1[31,78]
Sars 1067[18], 1190[12], 0.22[41]†–
1200[15]
Diaphanosoma 0.85[7] 0.25–5.00[4] 213[18], 237[9], y ¼ 0.31·3.02[22], 0.16–0.24[4], 6[1], 9[79] 0.32[79] 0.20[34],
brachyurum 271[6], 389[10], y ¼ 0.26·1.27[20] 0.2–0.3[6] 0.25[41]†–
Liéven 390[12] 406[11]
[20]
Holopedium 1.04 * 4.0–25[4] 521[9], 2271[6], y ¼ 0.38·0.36[20] 1.2–2 0.155[77] 115.5[31,37] 6.4[31,37]
[18]
gibberum 4979 (gnathobasal setae)[4],
Zaddach 1.8–3.9[6]
Sida crystallina 2.55[7] 4.5–25[4] 2920[24], 3800[12], 0.9–2.3 11[1], 16[32]‡
[2] [4]
Müller 9929 (gnathobasal setae) ,
2.48[24]
Polyphemus 0.83* >20000[30]
pediculus Linnee

Superscript letters denote values at temperatures other than 20C. a ¼ 15–19 C, b ¼ 21–25 C.
†
Based on field data.
‡
As cited in Gillooly (2000).
§
As cited in Sarma et al. (2005)
–
As cited in Lynch (1980)
**Data are for Daphnia dentifera (part of the D. galeata mendotae complex)
††
Data are for Daphnia galeata (European species which hybridises with D. mendotae in North America).

 2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd, Freshwater Biology, 52, 796–813
Functional diversity of zooplankton communities
801
 802

Table 2 Quantitative functional traits of copepod species including sources in superscript numbers (see Supplementary material).
A.J. Barnett et al.

Mean body Food size Max CR Fastest generation

Species length (mm) range (lm) (lL ind.)1 h)1) time (days) C:P C:N

Cyclopoida
Acanthocyclops robustus Sars 1.14 114–562[80] 13.4[81], 13.9[82], 14.8[83]a,
16.5[84], 18[85],19.8[86]
Acanthocyclops vernalis Fischer 0.72 12[86]
Cyclops scutifer Sars 0.88 0.25–5.2[87]
[88]
Cyclops vicinus Ulianine 1.63 6–31.7[89))91] 28.5[92]a, 59[93], 1200[91]a 23[81], 25.1[82]b, 26.3[94],
27[86], 31[95]
Diacyclops bicuspifatus thomas Forbes 0.84 15–100[96] 213[97]a, 7500[96]a 92.4[31,98] 5.2[31,98]
Eucyclops speratus Lilljeborg 0.80
Mesocyclops edax Forbes 0.93 <900 lm rotifers[99] 2708[100] 17.2[101] 107.6[31] 5.7[31]
[102]
Tropocyclops prasinus Fischer 0.55 6.5–80[102] 380[102] 16.8[103]a, 41.4[104]
[88]
Thermocyclops crassus Fischer 0.89 9.7[101]b, 23.5[81], 27.7[86]
Calanoida
Diaptomus minutus Lilljeborg 0.89 [102] 6.5–80[10,11,102,105,106] 487[106], 571[102], 790[11]b, 805[10]
Diaptomus oregonensis Lilljeborg 1.14[102] 3–80[102,107] 1356[102], 1666[108]
Epischura lacustris Forbes 1.53 30–80[102,105,109] 667[102], 966[105] 116.6[31] 5.2[31]
Leptodiaptomus minutus Lilljeborg 0.84 131.3[31] 5.8[31]
[110]
Leptodiaptomus sicilis Forbes 1.07 5–50
Onychodiaptomus birgei Marsh 1.44 [102]
Skistodiaptomus oregonensis Lilljeborg 1.26 2.5–30[107]
Skistodiaptomus reighardi Marsh 1.06
a b
Superscript letters denote values at temperatures other than 20 C. ¼ 15–19 C, ¼ 21–25 C.

 2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd, Freshwater Biology, 52, 796–813

vicinus was observed to feed at 28–63 lL ind.)1 day)1
on algae (Adrian, 1991; Santer, 1993) and
1200 lL ind.)1 day)1on rotifers (Rabette, Thouvenot
& Lair, 1998).
Generation time tended to be shorter for cladocer-
ans than cyclopoids, but again observed values varied
considerably both between and within species. The
most extreme example was for Thermocylops crassus in
which generation time ranged from 9.7 days (Wyng-
aard et al., 2005) to 27.7 days (Maier, 1994).
Stoichiometric ratios (C : P, C : N) for freshwater
zooplankton have recently been summarised in two
major studies (Elser et al., 2000; Frost et al., 2006) and
these values were included here. Unfortunately, only
values for common cladoceran species and four
copepod species appear to have been determined so
far.
Qualitative traits defined
Qualitative traits found that were sufficiently com-
mon to include in Tables 3 and 4 were habitat, trophic
group, optimal productivity conditions, feeding type
and selectivity, for both copepods and cladocerans,
and preferred food size category, predatory escape
response and helmet forming capacity, for cladocer-
ans only. The habitat in which each organism was
most likely to be found was categorised as either
littoral or pelagic. Trophic group was divided into the
traditional herbivore, omnivore and carnivore categ-
ories with the addition of herbivore–omnivore and
carnivore–omnivore. This addition was used to dis-
tinguish between copepods such as Mesocyclops edax,
that is more carnivorous, and Tropocyclops prasinus
that is more herbivorous. In this study, feeding on
non-photosynthetic protists was considered herbivo-
ry, in order to distinguish between those species that
feed on protists (e.g. Daphnia spp.), and those that feed
on larger organisms like rotifers and crustaceans (e.g.
Polyphemus spp. and Mesocyclops spp.). Optimal pro-
ductivity conditions characterised the lake trophic
status at which optimal competitive ability is achieved
by each species. For example, chydorids are more
successful in very productive lakes while daphniids
are generally more successful in low productivity
lakes with high quality food (Tables 3 and 4). Food
size was used to describe broadly the relative food
size category preferred by each species. Predatory
escape response characterised how a species evades
 2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd, Freshwater Biology, 52, 796–813
Functional diversity of zooplankton communities 803
an attack from a predator. The ability to produce a
helmet in Cladocera was also included as an anti-
predation trait.
Feeding type is the manner in which species or
groups obtain their food. Cladocerans have been
divided into four classes based on DeMott & Kerfoot
(1982); (1) Daphnia-type (D-type), where filtering is
from a stationary position with filtering apparatus on
the third and fourth legs, (2) Sida-type (S-type), similar
to D-type with the exception that the filtering appar-
atus is located on the first five legs, (3) Bosmina-type
(B-type), characterised by a horizontal active swim-
ming and less developed filtering apparatus on
thoracic appendages and (4) Chydorus-type (C-Filtra-
tion) where feeding is predominantly by scraping
algal particles from periphyton. Amongst the cope-
pods, raptorial predators like cyclopoids actively
capture and kill prey, while stationary suspension
feeders are more passive with less frequent swim-
ming. Finally Epischura lacustris is differentiated from
both raptorial and stationary suspension feeding as it
swims continuously while creating feeding currents,
thus being characterised as a ‘current cruiser’.
The last qualitative trait in Tables 3 and 4 is
selectivity, a relative estimation of how selective an
organism is towards its food. Raptorial feeders are
generally considered to be more selective than filter
feeders, but there are also relative degrees of selec-
tivity exhibited within the filter feeding Cladocera
species. The degree of selectivity can be defined based
on how active a species is in seeking out food
of a particular quality, through selective filtering or
particle rejection.
Qualitative data results
The qualitative trait matrix contains a number of trait
variables available only for some taxonomic groups
(Tables 3 and 4). For example, the ability to form
helmets obviously does not apply to copepods. Based
on our biological knowledge of many of these species,
qualitative traits were often assumed to be the same
within a genus, or sometimes within a family or order.
The majority of trait information in Tables 3 and 4 is
considered to apply to the entire genus, although
exceptions mentioned specifically in the literature are
cited. The preferred food of most cladocerans is algae,
and thus there is no trophic group distinction given
between cladoceran species as all are thought to
 804

Table 3 Qualitative functional traits of cladoceran species including sources in superscript numbers (see Supplementary material). Traits without a reference are personal
A.J. Barnett et al.

observations. Trait attributes obtained by generalisation of a group is indicated with a ‘†’.

Trophic Optimal
group productivity
Species Predatory escape response Habitat conditions Food size Helmet Feeding type Selectivity

Alona affinis Leydig Littoral[1,2] Herbivore[2] No C-Filtration

Alona rectangula Sars
Eubosmina corregoni Baird Reduced swimming[111,112]† Pelagic[113]† Herbivore[15]† Large[5] No B-Filtration[41]† Moderate[15,16]†
Bosmina longispina Leydig Reduced swimming[111,112]† Pelagic[113]† Herbivore[15]† Large[5] No B-Filtration[41]† Moderate[15,16]†
Bosmina longirostris Müller Reduced swimming[111,112]† Pelagic[1,113]† Herbivore[15]† High[114] Large[5] No B-Filtration[41]† Moderate[15,16]†
Ceriodaphnia dubia Richard Rapid swimming[112]† Pelagic[115]† Herbivore[113]† No D-Filtration[41]†
Ceriodaphnia reticulataJurine Rapid swimming[112]† Pelagic[115]† Herbivore[113]† No D-Filtration[41]†
Ceriodaphnia lacustris O.F.M. Rapid swimming[112]† Pelagic[115]† Herbivore[113]† All[5] No D-Filtration[41]†
Chydorus sphaericus Müller Littoral[1] Herbivore[2] High[36]† Small[5] No C-Filtration
Chydorus brevilabris Frey Littoral[1,115] Herbivore[113]† High[36]† Large[5] No C-Filtration
Daphnia ambiguaScourfield Rapid swimming[112]† Pelagic[1,113] Herbivore[114] Low[29,114,116] Yes[21,117] D-Filtration[41]† Low[15,16]†
Daphnia catawba Coker Rapid swimming [111] Pelagic[3] Herbivore[112] No D-Filtration[41] Low[14,15]
Daphnia galeata Sars Rapid swimming[112]† Pelagic[1,113] Herbivore[113]† Low[29,116]† Medium[5] Yes[21,117] D-Filtration[41]† Low[15,16]†
Daphnia longispina Müller Rapid swimming[112]† Pelagic[1,113] Herbivore[113]† Low[29,116]† No[117] D-Filtration[41]† Low[15,16]†
Daphnia magna Straus Rapid swimming[112]† Ponds[21] Herbivore[113]† Low[29,116] Small[5] No[21] D-Filtration[41]† Low[15,16]†
Daphnia pulex Leydig Rapid swimming[112]† Pelagic[1,113] Herbivore[113]† Low[29,116] No[21] D-Filtration[41]† Low[15,16]†
Daphnia pulicaria Forbes Rapid swimming[112]† Pelagic[113]† Herbivore[113]† Low[29,100,116] Medium[5] No[21] D-Filtration[41]† Low[15,16]†
Daphnia rosea Sars Rapid swimming[112]† Pelagic[1,113] Herbivore[113]† Low[29,116]† No[21,117] D-Filtration[41]† Low[15,16]†
Diaphanosoma brachyurum Liéven Pausing and jumping[112]† Littoral[113]† Herbivore[113]† High[114] All[5] No S-Filtration[41]†
Holopedium gibberum Zaddach Pelagic[113]† Herbivore[113]† Large[5] No S-Filtration[41]†
Sida crystallina Müller Littoral[1,2] Herbivore[2] Large[5] No S-Filtration[41]†
Polyphemus pediculusLinnee Pelagic[30] Carnivore[30] No Raptorial[30]

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 Table 4 Qualitative functional traits of copepod species including sources in superscript numbers (see Supplementary material). Trait attributes obtained by generalisation of a
group is indicated with a ‘†’.

Species Habitat Trophic group Optimal productivity conditions Feeding type Selectivity

Cyclopoida
Acanthocyclops robustus Sars Pelagic[118] Omnivore[118] Raptorial[88]† High[102,119]†
Acanthocyclops vernalis Fischer Pelagic[88]† Omnivore[88]† Raptorial[88]† High[102,119]†
Cyclops scutifer Sars Pelagic[118] Omnivore[88]† Raptorial[88]† High[102,119]†
Cyclops vicinus Ulianine Pelagic[120] Omnivore[88,90] Raptorial[88]† High[102,119]†
Diacyclops bicuspifatus thomas Forbes Pelagic[96] Omnivore carnivore[96,121] Raptorial[96,121] High[96,121]
Eucyclops speratus Lilljeborg Littoral[118] Omnivore herbivore[122] Raptorial[88]†
Mesocyclops edax Forbes Pelagic[115] Omnivore carnivore[123] Raptorial[123] High [99,123]
Tropocyclops prasinus Fischer Pelagic [118] Omnivore herbivore[102] Raptorial[102] High[102,119]†
Thermocyclops crassus Fischer Pelagic[88]† Omnivore herbivore[88]† Raptorial[88]† High[102,119]†
Calanoida
Diaptomus minutus Lilljeborg Pelagic[115] Omnivore-carnivore[124] High[29]† Current-cruiser[102] High[102,119,125,126]†
Diaptomus oregonensis Lilljeborg Littoral[127] Herbivore[118] High[29]† Stationary suspension[102]† High[102,119,125,126]†
Epischura lacustris Forbes Herbivore[118] High[29]† Stationary suspension[102]† High[102,119,125,126]†
Leptodiaptomus minutus Lilljeborg Herbivore[118] High[29]† Stationary suspension[102]† High[102,119,125,126]†
Leptodiaptomus sicilis Forbes Pelagic[118] Omnivore[102] High[29]† Stationary suspension[102]† High[102,119,125,126,128]†
Onychodiaptomus birgei Marsh Pelagic[118] Herbivore[118] High[29]† Stationary suspension[102]† High[102,119,125,126]†
Skistodiaptomus oregonensis Lilljeborg Pelagic[115] Omnivore[129] High[29]† Stationary suspension[129]† High[102,119,125,126]†
Skistodiaptomus reighardi Marsh Pelagic[118] Omnivore[118] Stationary suspension[102]†

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Functional diversity of zooplankton communities
805
806 A.J. Barnett et al.
be predominantly herbivorous (Table 3), with the littoral herbivores (except for Leptodiaptomus sicilis)
exception of Polyphemus. Cyclopoids were all classi- while the bottom half of group 1 is mainly pelagic
fied as raptorial feeders in the five studies where herbivores (Eucyclops speratus excepted). It appears
qualitative data was obtained (Table 4). Other traits, that herbivorous habit and smaller body size distin-
such as preferred food size and productivity condi- guish group 1. Group 2 contains the remaining
tions for optimal growth, were found for some copepods, all of which display omnivorous–carnivor-
cladoceran species but not for cyclopoids. Data are ous tendencies. The cladoceran Polyphemus is also
sparse overall for some variables such as the produc- found in this group as it is carnivorous. Group 3
tivity conditions of optimal growth and qualitative consists of the larger pelagic cladocerans and contains
food size. all Daphnia species with the exception of D. pulex and
D. magna – both of which are pond/littoral species
and fall into group 4 along with the other littoral,
Functional community dendrogram
large Cladocera. Finally, Diaphanosoma brachyrum
The major groups in the taxonomy-based tree of 36 forms group 5, as it is an S-filtration species but its
species constructed through simple clustering of much smaller body size appears to warrant its
species by genus, family, class, etc. (Fig. 1) are similar separation from S-filtering Sida.
but not identical to the dendogram generated using While the dendrogram generated using body length,
length, habitat, trophic group and feeding type habitat and trophic group grouped some related taxa
(Fig. 2). Five major groups emerge from the FD together, the dendrogram generated using only body
dendogram (Fig. 2), with one species (Diaphanosoma length showed no tendency to group species of a
brachyurum) forming an entire group. Group 1 on particular genus or family together (Fig. 3).
Fig. 2 is a mixed group of smaller herbivorous
Cladocera and herbivorous (occasionally, herbivor-
Discussion
ous–omnivorous) calanoid and cyclopoid copepods.
The upper half of this group is composed of mainly In order to apply a functional perspective to
zooplankton communities, we must first establish
Sida crystallina
whether function can be adequately estimated for
Ctenopoda
Diaphanosoma brachyurum commonly encountered species. Given the large
Holopedium gibberum
Polyphemus pediculus
Onychopoda Chydorus brevilabris
number of studies on zooplankton life history, mor-
Chydorus sphaericus
Branchiopoda Alona affinis phology and physiology over the past few decades,
Bosmina longirostris
Bosmina longispina one would expect that a functional classification could
Eubosmina corregoni
Daphnia catawba be developed. In this review of the literature, we have
Amomopoda Daphnia ambigua
Daphnia rosea pieced together the various studies and data therein
Daphnia pulicaria
Daphnia pulex for common North American species of freshwater
Daphnia magna
Daphnia longispina
cladocerans, and calanoid and cyclopoid copepods. In
Daphnia galeata mendotae general, most data were available for cladocerans
Ceriodaphnia reticulata
Ceriodaphnia dubia while there were fewer studies on freshwater cope-
Ceriodaphnia lacustris
Skistodiaptomus reighardi pods. This bias reflects the ease of culturing and
Skistodiaptomus oregonensis
Onychodiaptomus birgei studying cladocerans, and especially Daphnia species,
Calanoida Leptodiaptomus sicilis
Leptodiaptomus minutus because of their relatively non-selective filter feeding
Epischura lacustris
Copepoda Mesocyclops edax behaviour and nutritional requirements (Pennak,
Eucyclops speratus
Thermocyclops crassus 1989). Their feeding differs from that of copepods,
Tropocyclops prasinus
Cyclopoida Cyclops vicinus which actively capture and ingest individual suspen-
Cyclops scutifer
Diacyclops bicuspifatus thomas ded food particles (Koehl & Strickler, 1981). Further,
Acanthocyclops vernalis
Acanthocyclops robustus longer generation times and sexual reproduction lead
to culturing difficulties which bias studies away from
Fig. 1 A taxonomic tree of the species used to create the
functional dendograms in Figs 2 & 3. Taxonomy is based on
freshwater copepod species.
Thorp & Covich (2001). Branch lengths are for representation Feeding preferences and rates are important char-
only and do not represent quantitative values. acteristics of species that ideally would be included in
 2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd, Freshwater Biology, 52, 796–813
 Functional diversity of zooplankton communities 807

Onchyodiaptomus birgei
Leptodiaptomus sicilis
Chydorus brevilabris
Chydorus sphaericus
1 Alona affinis
Bosmina longirostris
Bosmina longispina
Eubosmina corregoni
Eucyclops speratus
Thermocyclops crassus
Tropocyclops prasinus
Skistodiaptomus reighardi
Skistodiaptomus oregonensis
Leptodiaptomus minutus
Epischura lacustris
Diacyclops bicuspifatus thomas
2 Polyphemus pediculus
Mesocyclops edax
Cyclops scutifer
Acanthocyclops vernalis
Acanthocyclops robustus
Cyclops vicinus
Daphnia longispina
Daphnia catawba
Daphnia pulicaria
Daphnia galeata mendotae
Fig. 2 A functional dendrogram (FD) Daphnia rosea
Daphnia ambigua
generated by hierarchical clustering 3 Ceriodaphnia lacustris
Ceriodaphnia reticulata
analysis of the standardised Euclidean Ceriodaphnia dubia
distances of each species based on four Holopedium gibberum
Sida crystallina
4
functional traits (body length, habitat, Daphnia pulex
5 Daphnia magna
trophic group and feeding type). A Diaphanosoma brachyurum

dashed line indicates the split that defines 4 3 2 1 0

five groupings (numbered) of species as
discussed in the text. Distance

Chydorus sphaericus
Bosmina longispina
Bosmina longirostris
Tropocyclops prasinus
Eubosmina corregoni
Chydorus brevilabris
Leptodiaptomus minutus
Diacyclops bicuspifatus thomas
Diaphanosoma brachyurum
Ceriodaphnia lacustris
Polyphemus pediculus
Cyclops scutifer
Daphnia ambigua
Thermocyclops crassus
Eucyclops speratus
Ceriodaphnia reticulata
Ceriodaphnia dubia
Alona affinis
Mesocyclops edax
Acanthocyclops vernalis
Skistodiaptomus reighardi
Leptodiaptomus sicilis
Holopedium gibberum
Acanthocyclops robustus
Sida crystallina
Daphnia magna
Daphnia longispina
Daphnia catawba
Cyclops vicinus
Daphnia pulicaria
Daphnia pulex
Daphnia galeata mendotae
Fig. 3 A functional dendrogram (FD) Onychodiaptomus birgei
Daphnia rosea
generated by hierarchical clustering Epischura lacustris
Skistodiaptomus oregonensis
analysis of the standardised Euclidean
distances of each species based solely on 2.5 2.0 1.5 1.0 0.5 0
body length. Distance

calculations of functional diversity (e.g. FD). We chose tion (Peters & Downing, 1984; Sterner, 1989; Mauch-
to include clearance rate and food size range here, as line, 1998), body size (Peters & Downing, 1984) and
these were the two most commonly reported standard temperature (Burns, 1969; Peters & Downing, 1984).
measures in the literature. The wide range of We attempted to account for these differences by
clearance rates between studies, however, makes it selecting maximum clearance rate values at one
difficult to choose a single clearance rate to include in temperature (20 C) and low food concentrations,
FD calculations. Clearance rate is known to be affected and we accounted for variations with body size by
by a wide range of factors, including food concentra- reporting only values for adult female copepods and
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808 A.J. Barnett et al.
clearance rate versus body size regressions for cla-
docerans. Clearance rate, however, also varies with
food type (Bogdan & Gilbert, 1982; Sterner, 1989),
food quality (Sterner, 1989; Gulati & DeMott, 1997),
predation pressure (Williamson, 1986), hunger (Stem-
berger, 1986; DeMott, 1995; Mauchline, 1998), length
of experiment (Mauchline, 1998) and the time of day
the experiment was run (Schoeneck, Williamson &
Stoeckel, 1990). These various factors probably ac-
count for the wide range of clearance rates for several
species in the trait tables. Despite this considerable
variability, an examination of the range of observed
values may still provide valuable insights into the
relative feeding rates of individual species.
Ingestion rates (food quantity consumed per unit
time) may also prove to be a useful trait for FD
calculations. Many studies of copepods in particular,
however, report ingestion rates as the number of food
particles consumed per unit time, rather than a rate of
mass or carbon ingestion, limiting the comparibility of
studies. Ingestion rate also varies with experimental
conditions, as noted above for clearance rate, making
ingestion rate difficult to include in FD calculations at
present. Reported feeding rates in our review have not
been adjusted by the mean body size of each species.
This is because the goal of an FD calculation is to
provide a measure of the overall niche space occupied
by a community of organisms. The actual feeding
rates of zooplankton are thus necessary to determine
what the overall effect of a particular community
might be on its resource. In addition, FD calculations
are sensitive to highly correlated data (Mason et al.,
2005). Thus it would be inappropriate to include both
body length and biomass-adjusted feeding rates in a
single calculation since individual biomass is most
usually calculated through length–weight regressions.
As mean body length is the more reliable trait, we
have included that in our FD calculations.
In terms of feeding preferences, some issues arise
when data are collated as we have done. In comparing
across studies, it was clear that the characterisation of
cyclopoids as carnivores (Sterner, 1989) was not
supported. Studies on Tropocyclops prasinus and Meso-
cyclops edax reveal that, while both are omnivorous, the
former is more herbivorous (Peacock & Smyly, 1983)
while the latter is more carnivorous (Confer, 1971), and
algae are an important component of the diet of both
juvenile and adult cyclopoids (Brandl, 1998). The use of
newer food web determination techniques, such as
 2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd, Freshwater Biology, 52, 796–813
stable isotopes, may be particularly fruitful for the
characterisation of the food sources of zooplankton
within lakes of interest in the near future (Bearhop
et al., 2004), and such information could be included in
an expanded functional classification scheme.
The mesh size of filtering apparatus in cladocerans
(Table 1) can be useful as a predictor of food selec-
tivity, but this may not always be the case. While there
is a correlation between uptake efficiency for small
food sizes (bacteria) and filtering apparatus size
(Brendelberger, 1985), some exceptional species, like
Bosmina, do not feed efficiently on food sizes predic-
ted from mesh size (DeMott, 1985). Furthermore, ‘soft’
algae (naked, gelatinous flagellates) are more likely to
be ingested by filter feeders than ‘hard’ algae (diatoms
and dinoflagellates) (DeMott, 1995). An alternative to
using quantitative mesh size measurements is to
follow the more qualitative groupings suggested by
Geller & Müller (1981) and DeMott (1985); (1) high
efficiency bacteria feeders, (2) low efficiency bacteria
feeders and (3) presumptive macrofiltrators. How-
ever, we did not use this classification because mesh
size is a morphological feature that is more readily
measured on preserved zooplankton that would
typically be collected and used for FD calculations.
For FD calculations, a measure of population
growth rate is most informative, but this information
is only available at present for some species of
cladocerans. Generation time was the most common
standard measure of growth available in the litera-
ture. These values are also known to be dependent on
food quantity and temperature (Hart, 1990; Peterson,
2001; Sarma, Nandini & Gulati, 2005), predation
pressure and food quality (Sarma et al., 2005). By
selecting the shortest generation times at 20 C, we
attempted to account for these variations as much as
possible, but discrepancies between studies were still
apparent, sometimes making it difficult to select one
value of generation time per species for FD calcula-
tions. Clutch sizes was also available for many
zooplankton species although, per se, clutch size does
not reflect the rate of population increase and is of
limited value in FD calculations. Egg production rate
per female, in number of eggs produced per day or
carbon produced per day, would be more useful for
FD calculations, although these measures are not as
common as clutch size in the literature.
Stoichiometric ratios are increasingly being used to
characterise the role of species in communities, as

they give insight into nutrient cycling (e.g. Elser et al.,
2000; Frost et al., 2006). Given all the caveats and
issues with calculation of feeding and growth rates,
and given its strong connection with ecosystem effects
of species, stoichiometry seems particularly to offer a
good measure of functional traits in freshwater
zooplankton.
Productivity conditions for optimal growth was one
qualitative trait for which some information was
available. However, this trait should probably be
considered together with other habitat characteristics,
as there are also important top–down features that
will affect zooplankton. DeMott, Edington & Tessier
(2004) demonstrated, for example, that the presence of
predators, as well as refuges, played a more important
role in determining zooplankton species presence and
abundance in lakes than did bottom–up nutrient
effects. Thus, while these productivity conditions
remain useful for determining the food niches of
zooplankton species, studies determining the pred-
ator or refuge niches of zooplankton would be of
benefit.
Finally, an important difficulty with the collection
of traits from the literature stems from changes in
species classification after taxonomic re-evaluation.
For example, B. longirostris was considered a common
North American species until De Melo & Hebert
(1994) analysed allozymic traits and re-analysed
morphology with an electron microscope and showed
that this species was located only in California. In the
rest of the North American distribution, it has been
divided into two species; Sinobosmina liederi and
Sinobosmina freyi. Functional information on these
newly defined species is obviously lacking but prob-
ably overlaps significantly with that of B. longirostris.
In addition, due to a re-evaluation of the taxonomy of
the genus Daphnia, the taxonomic status of Daphnia
rosea is now under question (Hebert, 1995) and, thus,
data for this species may no longer be applicable in
future research. Furthermore, our study was limited
to the collection of traits of crustacean zooplankton
and did not include rotifers, the latter being an
important food web link to microbes, as well as
competitors and prey for crustacean zooplankton
(Nogrady, Wallace & Snell, 1993). Thus, the determi-
nation of similar parameters for rotifers may also be
useful for estimates of zooplankton functional diver-
sity and the study of the drivers of diversity, niche use
and ecosystem function.
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Functional diversity of zooplankton communities 809
The decision as to which traits to include in the FD
dendogram was therefore a difficult one. Quantitative
traits were plagued with the issues of discrepancies
between studies, and the lack of data for several
groups of species. Furthermore, as functional mea-
sures like Petchey & Gaston’s (2002) FD are affected
by the use of measures that are correlated, it would be
redundant to use correlated measures (Mason et al.,
2005) such as body size and clearance rate predicted
from body size regressions. It would be preferable to
use only one of these measures at a time. Because of
these concerns, we opted to include body length as the
only quantitative trait, and habitat, feeding type and
trophic group as qualitative traits.
The dendrograms in Figs 2 & 3 provide an example
of the how a suite of traits can be used to generate a
functional dendrogram for a regional species pool.
The dendrogram based on four functional traits
(Fig. 2) more closely resembled taxonomic groupings,
although with some significant exceptions, than did
the dendrogram constructed based on body size alone
(Fig. 3). This is not surprising as morphological
structure is often related to function in zooplankton
and, thus, although we argue that taxonomic identi-
fication (based largely on morphology) is not suffi-
cient, it should still have some relationship with
function. The important results lie in the exceptions
observed where closely related sister-genera and
other related taxa do not cluster together in the
functional dendrogram. Group 1 (Fig. 2) was a good
example of relatively unrelated taxonomic species that
were grouped together in the functional dendrogram.
Here a mixture of unrelated herbivores consisting
of smaller Cladocera (bosminids, chydorids) and
copepods (Eucyclops, Onchyodiaptomus, Leptodiapto-
mus) were classified together. Similarly, carnivorous
species made up another important group. The group
most closely related to taxonomic classification was
group 3, which contained most of the Daphnia species
but also other pelagic cladocerans. Thus, considering
a zooplankton community in terms of its ecological
functioning should lead to a different diversity calcu-
lation than one would obtain using traditional taxo-
nomically based diversity indices. Our dendrogram
based on four traits provides a starting point for such
functional calculations and usefully classifies species
based on their ecological roles.
The number of traits included can significantly
affect the FD measure (Petchey & Gaston, 2002). From
810 A.J. Barnett et al.
the dendrogram calculated using body size alone
(Fig. 3), the resulting classification does not resemble
at all the taxonomic one, and probably does not
provide an accurate functional descriptor of the
community either. It is therefore evident that the
functional dendrogram and estimate of FD can
depend on which characters are included in the
analysis. Petchey & Gaston (2002) demonstrated the
importance of choosing a suite of traits that function-
ally characterises the community without overempha-
sising functional complementarity; too few traits
underestimates the level of complementarity, while
too many traits results in a measure that is effectively
the same as species richness. We show here the two
extremes in what is currently possible to calculate for
zooplankton FD based on our review of the literature
to date. The ultimate test of which characters to use in
combination to calculate FD will be determined by the
combination that best represents a particular ecosys-
tem function of interest (e.g. total biomass of
zooplankton) or most responds to an environmental
gradient (such as a gradient of total phosphorus for
example, Barnett & Beisner, 2007). Blackburn et al.
(2005) conducted such an analysis for the purposes of
determining which functional characteristics of a
community of invasive predators were most respon-
sible for driving bird species extinct on islands. They
tested all 211 possible combinations of 11 functional
traits for predator communities to determine which
provided the highest explanatory power for extinc-
tions. Such an analysis can also be conducted for
zooplankton communities to determine which func-
tional traits contribute most to their responses or their
function in particular lake ecosystems (Barnett &
Beisner, 2007).
To obtain Petchey & Gaston’s (2002) FD of a lake
community containing only a subset of this regional
pool for north-eastern North America, one would
need only to sum the branch lengths of those species
found in the lake in our dendrogram. Not only do
these dendrograms provide an example of how these
traits can be used, they show how taxonomic related-
ness does not relate directly to function, even when
we have had to assume in some cases that species
within a genus have similar qualitative characteristics.
Large taxonomic differences may not necessarily
imply large difference in the ecological roles of species
in a community. Therefore, a more functional
approach to diversity of lake zooplankton communi-
 2007 The Authors, Journal compilation  2007 Blackwell Publishing Ltd, Freshwater Biology, 52, 796–813
ties may be very useful, especially when predicting
how community function, in addition to structure,
may change under future altered conditions.
A practical approach to constructing functional
classifications would combine traits that can be readily
measured. For example, multiple sampling through
time of a lake undergoing a seasonal succession
provides snapshots of the zooplankton community.
Measurements of length can be used to infer feeding
rates and food niches to a certain degree, while
generation time, population growth rate and age
structure can be useful measures of the responses to
predation and environmental changes. Stoichiometric
values can be applied to the community to determine
fluxes of major nutrients through the food web. With
the data compiled in this review such an approach can
now begin, with at least a minimum of characteristics.
Again, it will be important to test this classification
scheme experimentally, to see which combination of
traits most accurately captures changes in community
function under varied habitat conditions.
In summary, this study provides a synthesis of
available functional traits of zooplankton species
based on approximately five decades of laboratory
and observational studies. Data were most abundant
for cladocerans, particularly Daphnia species, while
most lacking for calanoid species. This information is
useful to those who wish to estimate functional
diversity for North American freshwater zooplankton
communities based on taxonomic information of
community composition. It also outlines the limits of
our current understanding of niche use by zooplank-
ton species, and points to a more integrative
functional approach to compiling new data on
zooplankton that would be beneficial to the ecological
study of lakes. The use of functional measurements of
diversity in capturing ecological function of aquatic
ecosystems has yet to be explored in detail, but work
from terrestrial systems shows that such an approach
should provide a greater understanding of the mech-
anisms which determine community composition and
response to major environmental shifts.
Supplementary material
The following supplementary material is available for
this article:
Appendix S1. List of references used to generate
functional trait matrices.

This material is available as part of the online article
from: http://www.blackwell-synergy.com/doi/abs/
10.1111/j.1365-2427.2007.01733.x (This link will take
you to the article abstract).
Please note: Blackwell Publishing are not respon-
sible for the content or functionality of any supple-
mentary materials supplied by the authors. Any
queries (other than missing material) should be
directed to the corresponding author for the article.
Acknowledgments
The authors thank NSERC and CFI (Canada), FQRNT
(Québec, Canada) for funding to BEB, the foundation
of the University of Quebec at Montreal for a post-
graduate scholarship to AJB, and a GRIL postdoctoral
fellowship to KF. Thanks to M.L. Longhi, M. Maezo
and G. Méthot for field and lab assistance and to
R. Vogt, W. DeMott, A. Hildrew and an anonymous
reviewer for helpful comments on the manuscript.
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(Manuscript accepted 5 January 2007)