Article
Biodiversity of aquatic insects: spatial gradients and environmental correlates of assemblage-level measures at large scales Jani Heino1,2 Finnish Environment Institute, Research Programme for Biodiversity, P.O. Box 413, FI-90014, University of Oulu, Finland
1
(correspondence). E-mail:
[email protected] Department of Biology, University of Oulu, P.O. Box 3000, FI-90014 University of Oulu, Finland.
2
Received 15 July 2008; accepted 29 October 2008; published 12 March 2009
Abstract Biodiversity embraces multiple facets of the variability of nature, although most research has dealt separately with population-, species- and assemblage-level measures. This review concentrates on aquatic insect biodiversity and the assemblage-level measures, such as species richness, assemblage compositional variation, taxonomic distinctness and functional diversity. Most studies on aquatic insects have considered biodiversity patterns based on surveys of local assemblages along geographical and environmental gradients, while there is a virtual lack of studies that have considered regional grain sizes (i.e. the size of the observational unit). Latitudinal gradients at both regional and local grain are ambiguous in aquatic insects, as different studies have found either higher or lower local diversity in the tropics than in the temperate zone. Other geographical patterns in aquatic insect diversity may also be relatively weak, as suggested by subtle among-ecoregion differences in both local species richness and assemblage composition. An ecological explanation for the absence of strong geographical gradients is that local environmental features may not necessarily vary with geography, and these factors may override the influences of historical and climatic influences on local diversity.
Evidence from
large-scale studies suggests, however, that not only various local habitat and ecosystem variables, but also those measured at the watershed, regional and geographical scales are needed to account for variation in the species richness and assemblage composition of aquatic insects. Within regional species pools, species richness and assemblage composition in both lotic and lentic ecosystems vary most strongly along gradients in habitat size and acidity.
Knowing how α-, β- and γ-
diversity of aquatic insects vary along geographical and environmental gradients has important implications for the conservation of biodiversity in freshwater ecosystems.
Such knowledge is
not yet well-developed, and two aspects should be considered in future research. First, further survey research on lotic and lentic ecosystems is necessary for improving our understanding of general biodiversity patterns.
DOI: 10.1608/FRJ-2.1.1
Second, given that freshwater ecosystems are facing a severe
Freshwater Reviews (2009) 2, pp. 1-29 © Freshwater Biological Association 2009
Heino, J.
biodiversity crisis, the implementation of representative networks of freshwater protected areas would certainly benefit from increased understanding of patterns in aquatic insect biodiversity. Keywords: Assemblage composition; congruence; environmental gradients; functional biodiversity; higher taxon surrogates; lakes; latitudinal gradient; ponds; rivers; species richness; streams.
Introduction
based on surveys of local ecosystems within small regions (Vinson & Hawkins, 1998; Jeppesen et al., 2000).
Biodiversity embraces multiple facets of the variability
A decade ago, Vinson & Hawkins (1998) reviewed
of nature, ranging from genes to species, and from
factors affecting stream insect biodiversity at local,
ecosystems to biomes (Gaston & Spicer, 1998).
Most
catchment and regional scales. While their review focused
research has traditionally been based on populations,
on patterns at different scales, especially those at the
species and assemblages (Angermeier & Schlosser, 1995).
within-stream scales, large scales received less attention.
This review will concentrate on assemblage-level measures
However, a large number of large-scale studies spanning
of biodiversity using aquatic insects as a model group.
catchments, ecoregions and countries have emerged since
Assemblage-level measures in this context include species
the publication of that paper. One aim of the present
richness, assemblage composition, taxonomic variability
paper is thus to update and complement the information
and functional diversity. The approach in this review is
provided by the previous reviews on stream insect
general, however, and most consideration will be grounded
biodiversity (Vinson & Hawkins, 1998; Meyer et al., 2007;
on general ecological knowledge.
Clarke et al., 2008). The present review concentrates on
General patterns of biodiversity include diversity–
large-scale patterns across natural environmental gradients,
latitude (e.g. Willig et al., 2003), diversity–productivity (e.g.
and the reader should consult previous reviews for further
Waide et al., 1999), diversity–disturbance (e.g. Petraitis et
insights into small-scale aspects of aquatic insect ecology
al., 1989) and diversity–heterogeneity relationships (e.g.
(Ward, 1992;Allan & Castillo, 2007; Vinson & Hawkins, 1998)
Vinson & Hawkins, 1998). These relationships have been
and anthropogenic influences on freshwater ecosystems
studied across a wide variety of spatial scales. Most of the
and biodiversity (Malmqvist & Rundle, 2002; Allan, 2004;
research on latitudinal gradients has dealt with macro-
Dudgeon et al., 2006). The emphasis in this review will
scales (i.e. wide spatial extents and large-sized grids), and
be on aquatic insects, because they typically account for
these studies have largely concentrated on terrestrial and
more than 80 % of taxa in freshwater macroinvertebrate
marine taxa. Many macro-scale studies on terrestrial and
assemblages (e.g. Sandin, 2003; Briers & Biggs, 2005).
marine taxa show strong latitudinal gradients in species richness (Rosenzweig, 1995; Gaston & Blackburn, 2000;
Importance of spatial scale
Willig et al., 2003; Hillebrand, 2004). By contrast, few studies have examined freshwater taxa in this respect,
Spatial scale embraces two components (Wiens, 1989).
and the studies that exist have considered latitudinal
First, grain refers to the size of the observational unit. In
variation in local richness (Vinson & Hawkins, 2003;
aquatic studies, grain may be related to units sampled by a
McCreadie et al., 2005). Similarly, although numerous
small sampling device (e.g. Surber sampler), parts of whole
studies in the terrestrial realm have concentrated on
ecosystems (e.g. a stream riffle) or whole ecosystems
macro-scales when examining diversity–productivity and
(e.g. a pond). In macroecology, grain sizes are typically
diversity–heterogeneity relationships (Rosenzweig, 1995),
much larger than these, and macroecological studies
freshwater studies have considered these relationships
are often based on large-scale grids, regions or countries
© Freshwater Biological Association 2009
DOI: 10.1608/FRJ-2.1.1
Aquatic insect biodiversity
(Gaston & Blackburn, 2000). Second, extent refers to the area
as α-diversity, and between-habitat diversity as β-
over which the sampling units are distributed. In aquatic
diversity. In this review, the first one refers to the number
studies, extent may be a drainage basin, an ecoregion or
of species in a local ecosystem, while the latter refers
even a continent. In the present review, emphasis will be
to variation in assemblage composition between local
given to grain sizes that comprise parts of ecosystems and
ecosystems. While α-diversity thus reflects the importance
whole ecosystems and will be directed at extents spanning
of local abiotic and biotic factors, β-diversity is related
from single drainage basins and ecoregions to countries
to the way in which species respond to environmental
and continents.
heterogeneity along ecological gradients.
Finally, γ-
The importance of considering grain and extent is
diversity refers to regional species richness, and is the
closely related to the fact that patterns and associated
sum of differences among habitats in species composition.
processes typically change with regard to the spatial
It is thought to be determined primarily by large-scale
scales considered (Levin, 1992; Whittaker et al., 2001).
evolutionary, historical and climatic factors. In this review,
For example, the determinants of species richness may
γ-diversity refers to the number of species in separate
differ considerably when streams versus large-scale grids
geographical regions, large-scale grids or drainage basins.
(e.g. 100 km × 100 km) are considered across the same
Another approach to understanding the effects of
spatial extent (Ricklefs, 1987). In the former case, not only
spatial scale relates to the importance of environmental
historical and climatic processes determine species richness,
variables measured at different scales. This approach is
but also local-scale factors, such as disturbance, stress and
closely associated with the idea of environmental filters
heterogeneity. By contrast, when large-scale grids are
(Fig. 1) (Tonn, 1990; Poff, 1997) and has been followed in
considered across the same spatial extent, it is unlikely that
a number of studies on freshwater macroinvertebrate
‘mean’ local habitat conditions override the influences of
assemblages (Johnson & Goedkoop, 2002; Sandin, 2003;
historical and climatic factors on species richness. These
Townsend et al., 2004; Heino et al., 2007a). These studies
scale-dependent differences mainly stem from the fact
have generally considered multiple nested scales, with the
that local-scale environmental factors are homogenised
assumption that factors at different scales are hierarchically
across large-scale grids.
Another reason is that local
structured, and that factors at larger scales determine, at
species richness is pooled over all local assemblages when
least in part, those at smaller scales (Frissell et al., 1986;
large-scale grids are considered, yielding a measure of
Hildrew & Giller, 1994). In this review, both of these
regional species richness. In this vein, it is also important
approaches will be considered, yet a clear distinction will
to note that latitudinal gradients are typically stronger
be made between (i) spatial scale in terms of grain and
when large-scale grids are examined, whereas weaker
extent and (ii) variables measured at different spatial scales.
species richness–latitude relationships may emerge when local ecosystems are considered (Hillebrand, 2004). This pattern may result from different species and species
Large-scale patterns in species diversity
groups responding differently to local-scale environmental factors that do not necessarily co-vary with latitude.
Geographical patterns in regional diversity
More generally, these notions suggest that variation in biodiversity may be best explained by different factors at
Few studies have considered latitudinal gradients in
different grain sizes and geographical extents (Rosenzweig,
aquatic insect diversity. Most of the existing studies have
1995; Whittaker et al., 2001; Blackburn & Gaston, 2002).
been based on surveys of local ecosystems, but a couple
Diversity measured at different scales has typically
of studies have examined patterns across large grain size
been expressed as α-, β- and γ-diversity (Whittaker, 1975).
and considered regional species richness. For example,
Within-habitat diversity has typically been considered
Boyero (2002) examined the regional species richness of
DOI: 10.1608/FRJ-2.1.1
Freshwater Reviews (2009) 2, pp. 1-29
Heino, J.
mayflies and dragonflies in the New World. A general finding was that of the four large regions compared, after region area was accounted for, the regional species richness
Continental species pool
of mayflies was lowest in tropical South America and highest in Central America. A corresponding analysis of dragonflies showed that Central America was again the most diverse region, whereas North America was home for History and Climate
the lowest number of species. When the area of each region was not taken into account a different pattern emerged.
Watershed
Ecosystem
Macrohabitat
Decreasing spatial scale of environmental filters
Regional species pool
Mayfly diversity was clearly highest in North America and lowest in Central America, whereas dragonfly diversity was by far highest in tropical South America and lowest in temperate South America. The degree to which the four regions comprised too large a grain size for a meaningful analysis is unknown, and more detailed analyses based on smaller grain size would allow more useful insights into the latitudinal patterns of aquatic insect diversity. A comparison of major aquatic insect groups across Europe shows that there are significantly negative species richness–latitude relationships in mayflies, dragonflies and stoneflies, while other groups do not exhibit significant latitudinal patterns (Table 1). However, most aquatic insect orders attain their highest levels of diversity in France, as opposed to the more southerly countries of Portugal and Greece. Lowest levels of diversity occur in Latvia,
Microhabitat
the Netherlands, Denmark, Norway and Finland. These among-country differences reflect two potential suites of factors behind geographical gradients in diversity. First,
ASSEMBLAGE
Fig. 1. A schematic model of environmental filters affecting local assemblages. The continental species pool is determined by speciation and extinction processes at very large spatial and temporal scales. Filters at the largest scale in this scheme are history (e.g. speciation, extinction, dispersal) and climate (e.g. temperature, precipitation, energy) which determine the structure of regional species pools. Within the limits of regional species pools, there are filters at four levels that eventually determine local assemblages. These are watershed (e.g. vegetation, hydrological regimes), ecosystem (e.g. temperature, water chemistry), macrohabitat (e.g. depositional vs. erosional habitats, macrophyte cover) and microhabitat (e.g. macrophyte structural complexity, substratum particle size) filters. These filters determine diversity, composition, as well as functional and taxonomic variability of assemblages through species traits. Only species with suitable traits are able to overcome the challenges presented by the filters at each scale. For original ideas, see Tonn (1990) and Poff (1997). © Freshwater Biological Association 2009
southern and central European countries with major mountain ranges, such as France, Germany, Spain, Italy and Austria, support high diversity, whereas lowland countries, such as the Netherlands, Latvia and Denmark, support low diversity in most aquatic insect groups. This suggests that large altitudinal variation increases the heterogeneity of freshwater habitats and variability of thermal conditions, which has positive effects on diversity.
Also, if geographically separated mountain
ranges are present, each mountain range may support unique species, contributing to high diversity per country. Second, diversity decreases to the north of Europe, with diversity being clearly higher in the southerly countries, such as Spain and Italy, than in the northerly countries, DOI: 10.1608/FRJ-2.1.1
DOI: 10.1608/FRJ-2.1.1
158
-
148
-
114
58
67
142
358
France
Bulgaria
Italy
Greece
Portugal
Spain
63
84
87
-
97
-
75
558
138
57
91
146
82
147
70
-
110
125
101
126
101
117
50
33
12
25
37
10
35
36
-0.692**
Stoneflies
Plecoptera
124
-
54
60
89
-
79
-
-
67
61
37
73
62
66
-
56
65
60
64
23
52
53
-0.437
Bugs
Heteroptera
1113
-
231
269
342
-
579
-
-
-
358
349
480
287
419
-
410
201
294
354
202
269
-
-0.193
Beetles
Coleoptera
1173
355
216
150
382
-
465
-
-
305
307
176
322
251
266
-
197
188
166
219
188
190
214
-0.448
Caddisflies
Trichoptera
1309
-
-
-
-
-
635
-
-
342
556
327
703
239
411
-
520
197
528
506
-
503
539
-0.127
Midges
Chironomidae
Note: The number of Odonata species in Finland is actually 54. For comparability purposes, the figure given is that from http://www.freshwaterecology.info.
* P < 0.05; ** P < 0.01
Europe
-
-
Romania
77
77
71
84
69
73
Ukraine
114
Poland
-
52
85
-
Belgium
Switzerland
51
UK
53
116
36
Netherlands
52
59
122
43
Denmark
Austria
60
Sweden
54
Slovakia
49
Latvia
45
46
97
46
Norway
143
53
Finland
-0.805*
Dragonflies
Germany
-0.629**
Correlation
Odonata
Czech
Mayflies
Country
Ephemeroptera
Table 1. Numbers of species in major aquatic insect orders in European countries. Data were compiled from http://www.freshwaterecology.info (web site accessed on December 28, 2007); note that data for some countries and orders may have been updated on the website since. Information for many orders was not available for a number of countries (-). Note that information for the family of non-biting midges (Chironomidae: Diptera) may still be inadequate even in countries for which the numbers of species are given. Spearman rank correlations between species richness and latitude (south latitude of each country) are also shown. Countries are arranged on an approximate north–south gradient. Aquatic insect biodiversity
Freshwater Reviews (2009) 2, pp. 1-29
Heino, J.
such as Finland and Norway. This suggests that large-
preferences, with mid-latitude regions possibly providing
scale historical and climatic factors have some effects on
a wide range of habitats suitable to various kinds of taxa.
aquatic insect diversity along long geographical gradients.
However, the fact that other studies have found either
For example, Pleistocene glaciations may have had some
higher (Stout & Vandermeer, 1975; Lake et al., 1994;
effects on the present-day patterns of diversity, with some
Jacobsen et al., 1997) or lower (Stanford & Ward, 1983;
species having not yet reached all suitable areas. One caveat
Arthington, 1990; McCreadie et al., 2005) taxa richness
should be considered in this comparison. Taxonomic
in tropical than in temperate streams suggests that more
and faunistic traditions in each country may affect the
comprehensive studies, especially in tropical streams and
degree to which observed diversity reflects clear patterns.
spanning several tens of degrees of latitude, are needed
For some insect groups, such as non-biting midges,
to resolve whether stream insects do or do not show clear
historical faunistic work may differ considerably among
latitudinal gradients.
countries. This is critical for comprehensive comparisons
At a scale smaller than continents, some studies have
of aquatic insect diversity between countries and regions.
examined latitudinal patterns in the local species richness
Latitudinal species richness patterns may vary with
and assemblage composition of freshwater macroinverte-
the habitat types of the considered taxa. Ribera et al. (2003)
brates. In North Europe, where ecoregional delineations
found that the diversity of lotic beetles was negatively
largely follow latitude, there are differences between
related to latitude across countries in Europe, whereas
ecoregions in both species richness (Sandin & Johnson,
lentic beetles did not show a significant latitudinal pattern.
2000; Heino et al., 2002) and assemblage composition
The authors interpreted this finding as an indication
(Sandin & Johnson, 2000; Heino et al., 2007a). However,
that the higher historical persistence of lotic than lentic
these patterns are typically rather weak, and species
habitats contributed differently to diversity patterns
richness does not show a linear response to latitude
in the two groups of species. Proving the generality
from southern to northern ecoregions. Studies that have
of this hypothesis clearly awaits further studies on
directly examined species richness–latitude relationships
other aquatic insect groups and geographical regions.
in stream macroinvertebrates in Sweden and Finland have corroborated these findings (Sandin, 2003; Heino,
Geographical patterns in local diversity
unpublished manuscript). By contrast, it has been found that the species richness of stream macroinvertebrates
Patterns detected at regional scales may change when local
declines with latitude across Alaska (Wrona et al., 2006).
ecosystems are considered. Based on global literature on
This finding is in accordance with a previous study
local genus richness, Vinson & Hawkins (2003) found no
that found that stream insect genus richness declines
clear linear latitudinal richness trends for stream mayflies,
linearly from 40°N to 70°N in North America (Vinson &
stoneflies and caddisflies. By contrast, the highest levels
Hawkins, 2003). Similarly to lotic macroinvertebrates,
of genus richness were generally attained at mid-latitudes
studies of lentic macroinvertebrates have generally found
in North America and South America, although there was
no linear declines in species richness with latitude and
much scatter around the relationships. Secondary peaks
ecoregional delineations (Johnson, 2000; Vamosi et al.,
occurred in the tropics. While these patterns may result
2007). By contrast, part of the variability in the assemblage
partly from inadequate data and taxonomic knowledge in
composition of lake macroinvertebrates is related to
tropical regions, they also suggest that stream insects may
latitude (Johnson & Goedkoop, 2002; Johnson et al., 2004).
deviate from patterns detected for many major terrestrial
The two most probable reasons for the weak
taxa (Willig et al., 2003). A reason for this discrepancy may
relationships between latitude and species richness or
be that mayflies, stoneflies and caddisflies contain genera
assemblage composition are that (i) the local environmental
and species showing highly differential temperature
features of freshwater ecosystems do not necessarily
© Freshwater Biological Association 2009
DOI: 10.1608/FRJ-2.1.1
Aquatic insect biodiversity
show clear latitudinal variation (e.g. substratum particle
been directed to the relationship between local species
size, moss cover, current velocity, gradient), and (ii) local
richness (LSR) and regional species richness (RSR). Two
environmental factors override the influences of large-
general patterns have been suggested for LSR–RSR plots
scale historical and climatic factors on local assemblages.
(Cornell & Lawton, 1992). First, LSR may increase linearly
This reasoning is undeveloped, however, because single
with RSR, suggesting that local factors do not limit LSR,
surveys of local assemblages do not typically span large
assemblages being mainly under regional control. Second,
latitudinal gradients.
Thus, new large-scale datasets
if LSR attains an asymptote with increasing RSR, then it
may change our views about the latitudinal diversity
is suggested that local assemblages are saturated, being
patterns shown by freshwater macroinvertebrates.
mainly under local control. Although the approach in
In addition to patterns along latitudinal gradients, other
general has been criticised, particularly because highly
geographical patterns are also of interest. For example, east-
interactive assemblages may show linear LSR–RSR
to-west gradients in species richness have been found for
relationships (Hillebrand, 2005) and because such plots
stream insects across North Africa (Beauchard et al., 2003).
cannot be considered as evidence of competition (He et al.,
This pattern was explained by a combination of climatic,
2005), the examination of LSR–RSR relationships may at least
hydrological and biogeographic factors, with the latter
provide hints as to whether regional factors are important at
relating to historical dispersal processes. Similar historical
all (Witman et al., 2004; Harrison & Cornell, 2008).
influences of dispersal after the last glacial period may
LSR–RSR relationships for freshwater macroinverte-
also be related to east–west and south–north gradients of
brates have typically been derived using a riffle site as the
stream macroinvertebrate species richness in other regions,
local arena and a drainage basin as the region. Research
although the combination of geographical gradients with
on Finnish streams revealed that mean LSR was linearly
local environmental features may be more influential.
related to RSR, suggesting that regional species pools
More complex geographical gradients in species richness
limited LSR at least to some degree (Heino et al., 2003a).
and assemblage composition have also been suggested
A similar linear relationship was found for southeastern
by findings from surveys of streams across ecoregions
Australian streams (Marchant et al., 2006). The Australian
in New Zealand (Harding et al., 1997), North America
dataset contained a much larger number of species than the
(Hawkins & Vinson, 2000) and North Europe (Heino
Finnish dataset, suggesting that even at very high levels of
et al., 2002). While stream and lake macroinvertebrate
LSR and RSR, a linear relationship may prevail. In contrast,
assemblages show some differences between ecoregions,
based on a much more limited number of species, a strongly
differences are typically rather weak (Hardin et al.,
asymptotic relationship between LSR and RSR was found
1997; Hawkins et al., 2000; Johnson, 2000; Hawkins
for stream blackflies in the New World (McCreadie et al.,
& Vinson, 2000).
However, even weak relationships
2005). A study on Swedish streams and lakes yielded a
suggest that local assemblages are, at least in part,
weakly asymptotic mean LSR to RSR relationship in both
constrained by large-scale climatic and historical factors.
habitat types (Stendera & Johnson, 2005). The patterns detected in this study were probably confounded by the
Local–regional species richness
fact that RSR estimates for each ecoregion were based on
relationships
different numbers of streams and lakes, and a correction for the number of sampled sites might have been necessary
Theoretical considerations and empirical evidence suggest
for valid comparisons of LSR–RSR relationships (Marchant
that species richness in local assemblages is constrained
et al., 2006). Finally, a study on Palaearctic predaceous
not only by local factors, but also by regional species pools
diving beetles showed that LSR was positively related to
(Ricklefs, 1987; Cornell & Lawton, 1992; Griffiths, 1997;
RSR, even when compensations were made for differences
Angermeier & Winston, 1998). Much recent interest has
in local pond characteristics (Kholin & Nilsson, 1998). This
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Freshwater Reviews (2009) 2, pp. 1-29
Heino, J.
finding suggested a regional enrichment of LSR, and that
in local diversity. However, proving the predominance
a suite of historical and environmental explanations might
of weak latitudinal diversity patterns would necessitate
be needed to account for variation in the species richness of
comprehensive surveys of tropical freshwater ecosystems
predaceous water beetles in ponds. The findings from the
and those that span several tens of degrees of latitude
above studies clearly indicate that there is no single type
across major biomes. A fruitful approach in this context
of LSR–RSR relationship in freshwater macroinvertebrates.
would be to determine the relative influences of latitude, and of regional, watershed, ecosystem and habitat factors
Relative influences of variables at different scales
on local aquatic insect diversity, as has been done in studies conducted within smaller geographical extents.
Such
studies should not only be directed towards lotic systems, Conceptual models suggest that local assemblages are
for which some data already exist, but also towards lentic
structured by environmental factors at multiple scales
systems, for which our knowledge of geographical diversity
(Fig. 1) (Tonn, 1990; Poff, 1997). These suggestions have
patterns is more limited. Understanding geographical
been corroborated by empirical evidence. For example, the
diversity patterns would be a valuable asset in determining
assemblage composition of both stream and lake macroin-
and implementing geographically representative networks
vertebrates has been found to be determined by multiple
of protected areas for the conservation of freshwater
variables, ranging from local habitat and ecosystem features
biodiversity.
through watershed characteristics to regional climate and geographical factors (Townsend et al., 2003; Johnson et al., 2004; Sandin & Johnson, 2004; Heino et al., 2007a).
Species diversity–environment relationships within regions
In general, variables at small scales have been suggested to be the most important, both in terms of explained
The drainage basin is a natural regional unit in lotic ecology
variability in assemblage composition and in the ecological
(e.g. Frissell et al., 1986), and lake districts have similarly
sense, as small-scale variables are directly associated with
been given considerable recent attention in lentic ecology
the organisms. However, in a study covering a large
(e.g. Kratz & Frost, 2000). It is not surprising, therefore, that
geographical extent in Europe, geographical position
there have been many within-region surveys of lotic and
and ecoregion overrode the influences of local factors on
lentic assemblages. In the following, I will review findings
stream assemblages (Johnson et al., 2007). Also, species
from surveys of aquatic insect assemblages at geographical
richness in streams has been found to be best explained by
extents smaller (i.e. single ecoregions, drainage basins and
models including variables acting at different spatial scales
lake districts) than those considered above (i.e. several
(Sandin, 2003).
ecoregions and countries). I will first review research on (i) diversity–disturbance, (ii) diversity–productivity,
Synthesis and implications
(ii) diversity–acidity, (iv) diversity–heterogeneity and (v) diversity–habitat size relationships separately in both
Aquatic insect biodiversity varies geographically, although
lotic and lentic ecosystems. I will also consider similarity
present evidence suggests that these patterns are complex.
in the environmental relationships of species richness
Both species richness and assemblage composition
and assemblage composition. I will end this section by
typically show some variability among ecoregions and
addressing variation in species richness and assemblage
drainage basins, yet among-site heterogeneity within
composition with regard to the nested subset pattern.
regions largely masks the influences of climate and history
Biotic interactions, including predation and competition,
on local assemblages. Such a weak influence of large-scale
will be considered below only if there are shifts in species
factors is likely to contribute to weak latitudinal gradients
interactions associated with these major environmental
© Freshwater Biological Association 2009
DOI: 10.1608/FRJ-2.1.1
Aquatic insect biodiversity
gradients. Readers should consult previous reviews for
than the latter type of streams (Lake et al., 1986; Williams,
the importance of species interactions at small within-
1996). This is not surprising, because drying out of streams
ecosystem scales (Giller & Malmqvist, 1998; Allan &
presents severe challenges to the life history adaptations
Castillo, 2007).
and physiological performance of aquatic insects.
A
comprehensive recent review of the effects of drought Species diversity and major environmental
on stream biota can be found elsewhere (Lake, 2003).
gradients in lotic systems Diversity–productivity relationships Diversity–disturbance relationships
In addition to diversity–disturbance relationships, the
Floods are major natural causes of disturbance in running
effect of productivity on diversity has been studied
waters (Power et al., 1988; Resh et al., 1988; Poff, 1992)
extensively.
and constitute important determinants of diversity (Resh
diversity–productivity relationship takes many forms,
et al., 1988; Lake, 2000). Diversity is typically negatively
ranging from positive, unimodal and negative to non-
related to flow disturbance intensity and frequency
significant (Waide et al., 1999; Mittelbach et al., 2001). It is
(Robinson & Minshall, 1986; Death, 2002) and positively
possible that productivity in terms of stream water nutrient
related to benthic habitat stability (Death & Winterbourn,
concentration or periphyton biomass may also show
1995; Death, 2002).
Recent reviews have suggested that the
By contrast, none of the studies
positive, unimodal or negative relationships to the species
reviewed by Vinson & Hawkins (1998) suggested that the
richness of stream macroinvertebrates. When productivity
intermediate disturbance hypothesis would fit the species
ranges from very low to moderate, a positive relationship
diversity patterns of stream macroinvertebrates, although
may occur (Perry & Sheldon, 1986). Such a relationship
intermediate levels of disturbance may be associated with
may also be common in arctic and subarctic streams, which
higher diversity in other systems (Connell, 1978; Petraitis
have generally very low to moderately low nutrient levels.
et al., 1989). However, more recent studies have found
When nutrient levels range from very low to very high, a
highest diversity at sites perturbed by intermediate levels of
unimodal relationship may be found. Such a relationship
disturbances, leading to a unimodal diversity–disturbance
may be common in temperate regions with a wide range in
relationship (Townsend et al., 1997). This is due to the fact
stream water nutrient levels. Finally, when nutrients range
that high-level disturbances prevent abundant species from
from moderate to very high levels, a negative relationship
attaining a dominant position and competitive advantage
may be found. Such a relationship may also be common
(Huston, 1994), yet disturbances of intermediate level do
in temperate regions, where anthropogenic effects have
not lead to the exclusion of most disturbance-sensitive
increased nutrient inputs to streams. However, a negative
species. However, very few studies on lotic macroinver-
relationship may also occur at relatively low levels of
tebrates have examined these relationships across multiple
productivity (Malmqvist & Eriksson, 1995). Malmqvist
streams (Townsend et al., 1997; Death, 2002), whereas
& Eriksson (1995) suggested that the negative relationship
a number of within-stream studies have also examined
resulted, at least in part, from productive lake outlets being
the effects of disturbance on macroinvertebrate diversity
dominated by few abundant and competitively dominant
(Matthaei et al. 1999, 2000; McCabe & Gotelli, 2000). These
species, such as highly aggressive net-spinning caddis
small-scale studies, however, are out of the scope of this
larvae (see also Giller & Malmqvist, 1998). It should be
review.
emphasised, however, that nutrient levels may not be
Another form of disturbance affecting stream macroin-
the sole source of productivity affecting stream insects,
vertebrates is drought (Lake, 2003). In this regard, some
although nutrients are most closely associated with
studies have compared intermittent and permanent
autochthonous production in these systems. It should also
streams, and found that the former support fewer species
be noted that the above predictions are largely theoretical
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Heino, J.
rather than proven correct by empirical evidence.
Diversity–acidity relationships
Empirical studies are thus needed to ascertain if the form of
Among the most important stress factors in stream
the diversity–productivity relationship varies with regard
ecosystems is acidity. Species richness–acidity relationships
to regional range in potential productivity. These studies
have been studied in regional studies in widely-separated
should not only consider in-stream productivity, but also
geographical areas, ranging from Sweden (Malmqvist &
the productivity of the riparian zone.
Hoffsten, 2000) and Britain (Townsend et al., 1983) to the USA (Rosemond et al., 1992) and New Zealand (Collier et
Interactive effect of disturbance and productivity
al., 1990). Acidity typically leads to a low diversity of stream
on diversity
macroinvertebrates and such patterns have been observed
Disturbance and productivity may act in concert in affecting
in a number of studies spanning gradients from pH values
diversity, and this relationship is typically expressed as
4 to 7. In general, considerable reduction in the occurrence
the dynamic equilibrium model (Huston, 1994).
This
and richness of macroinvertebrates is typically seen at
model predicts that the unimodal relationship between
pH values below 5.5 (Otto & Svensson, 1983; Townsend
disturbance and diversity in the intermediate disturbance
et al., 1983; Hämäläinen & Huttunen, 1990; Larsen et al.,
hypothesis moves in relation to the productivity of a
1996). However, the findings of general reductions of
system, producing a contour of decreasing diversity with
species richness with acidity have also been challenged
high disturbance and high productivity. Such patterns
(Winterbourn & Collier, 1987; Collier et al., 1990; Dangles
of diversity result from two opposing forces: population
et al., 2004; Petrin et al., 2007). These discrepancies may be
growth rate and disturbance. First, local processes, such
related to the fact that acidity has both direct and indirect
as competition, dominate and reduce diversity under high
effects through food availability on species richness (Otto
population growth rates and low disturbances. Second,
& Svensson, 1983), or that species from different regions
other local processes, such as extinction, are important
may possess differing sensitivity to acid stress (Petrin et
at very low levels of growth rate and high disturbance
al., 2007). Furthermore, Petrin et al. (2007) suggested that
frequencies that thus reduce diversity. Very few studies
macroinvertebrate species richness decreases with acidity
have tested this model, because it necessitates surveys at
if it is of anthropogenic origin, while species richness may
large among-stream scales, making experiments unfeasible.
remain virtually unaltered if acidity is of natural origin.
However, Death & Zimmerman (2005) tried to overcome
Based on surveys in southern (acidity of anthropogenic
this problem, and examined simultaneously the effects of
origin) and northern Sweden (acidity of natural origin),
both disturbance and productivity on macroinvertebrate
they showed that the species richness of mayflies decreased
species richness in a set of streams. In general they found
with acidity in both regions, whereas the species richness
weak support for the two above-mentioned diversity
of caddisflies showed a negative response to acidity in
models and suggested instead that species richness was
the former region. The species richness of stoneflies did
a function of time since the last disturbance, this being
not vary along acidity gradients. Thus, the overall species
associated with the recovery of the autotrophic production
richness–acidity relationships for lotic macroinvertebrates
base for macroinvertebrates (Death & Zimmerman, 2005;
as a whole may be driven by a few taxonomic groups, while
see also Death & Winterbourn, 1995; Bond & Downes 2000;
others are more or less indifferent with regard to acidity.
Death, 2002).
For example, stoneflies in general may be rather indifferent with regard to acidity, and their species richness typically shows no clear response to acidity (Malmqvist & Hoffsten, 2000). However, despite no effects on species richness, the species composition of stonefly assemblages may, as
© Freshwater Biological Association 2009
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Aquatic insect biodiversity
11
in many other insect groups, change along gradients of
(Minshall et al., 1985a; Hildrew & Townsend, 1987; Allan
acidity (Petrin et al., 2007).
& Castillo, 2007; Cummins, 1996), and suggest that species richness increases from the headwaters (orders 1 to 3) to
Diversity–heterogeneity relationships
mid-order rivers (orders 4 to 6) and then declines with
Habitat heterogeneity has a positive effect on species
increasing river size (orders 7 to 12). These mid-order peaks
richness in general (Huston, 1994), and the same is true for
in species richness have been explained by higher levels
stream macroinvertebrates at various spatial scales (Vinson
of habitat heterogeneity and higher numbers of thermal
& Hawkins, 1998). Studies conducted at small scales, for
niches in mid-sized rivers than in smaller or larger rivers
example, within streams based on samples of stones, have
(Minshall et al., 1985b). In general, empirical evidence
associated variation in species richness to various measures
corroborates the prediction of the RCC that the species
of habitat heterogeneity, diversity and complexity. For
richness of stream macroinvertebrates increases with
example, species richness is typically positively related to
stream size up to mid-order rivers. Such patterns have been
habitat complexity in terms of stone surface features and
found in widely-separated geographical regions, including
plant cover at small scales (Hart, 1978; Hart & Horwitz,
Europe (Malmqvist & Hoffsten, 2000; Céreghino et al., 2003;
1991; Douglas & Lake, 1994; Dudley et al., 1986; Downes
Arscott et al., 2005), North America (Minshall et al., 1985b;
et al., 2000; Heino & Korsu, 2008). Positive relationships
Grubaugh et al., 1996) and South America (Tomanova et al.,
between stream substratum heterogeneity and species
2007). However, in studies where only small to mid-sized
richness have also been found among sites within streams
streams are sampled, species richness may be positively
(Cowie, 1985). By contrast, although habitat heterogeneity
related to stream size (Brönmark et al., 1984; Ward, 1986;
could be envisaged to account for among-stream variation
Malmqvist & Mäki, 1994; Malmqvist & Eriksson, 1995;
in species richness, evidence for such relationships is scant
Wiberg-Larsen et al., 2000; Paller et al., 2006), although
(Vinson & Hawkins, 1998). Reasons for the virtual absence
exceptions to these patterns also exist (Winterbourn et al.,
of a species richness–habitat heterogeneity relationship at
1981; Malmqvist & Hoffsten, 2000; Voelz & McArthur,
the among-stream scale are unknown, but possibly result
2000; Heino et al., 2008). When species richness attains
from stronger impacts of other physical and chemical
an asymptote or decreases in mid-sized rivers, a possible
factors that override the influences of habitat heterogeneity
reason for such patterns is that sampling effort has not
on species richness. Alternatively, stream ecologists may
been sufficient in mid-sized reaches, as opposed to more
not have measured heterogeneity at the among-stream
adequate effort in the headwater reaches (Malmqvist
scale as perceived by, or is important for, aquatic insects.
& Hoffsten, 2000; Heino et al., 2008). An alternative
It is also unknown if habitat heterogeneity is positively
explanation is that localised changes in environmental
related to stream size. It is possible, however, that habitat
conditions (e.g. substratum particle size, macrophyte
heterogeneity increases with stream size in small to mid-
cover) disrupt expected species richness–habitat size
sized rivers, but larger rivers do not necessarily possess
relationships (Wright & Li, 2002; Heino & Paasivirta, 2008).
more heterogeneous habitats (see below).
It should also be emphasised that data from the short and steep catchments in New Zealand and Australian drainage
Diversity–habitat size relationships
basins do not support the RCC predictions (Winterbourn
The river continuum concept (RCC) provides a theoretical
et al., 1981). Note that habitat size for lotic ecosystems in
basis to lotic ecology, for example, by relating river size
the above-mentioned studies has been based on a number
and longitudinal position to variation in functional feeding
of measures, including stream order, catchment area,
group composition and macroinvertebrate species richness
distance to source and stream width. Another important
(Vannote et al., 1980). The various predictions of the RCC
aspect to note is that not all studies have examined a
have been summarised several times since its formulation
single river continuum, but have been based on surveys of
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Heino, J.
different-sized streams scattered across a drainage basin
Diversity–productivity relationships
or an ecoregion. This may increase the importance of
Productivity in lentic ecosystems has typically been
localised environmental influences, causing patterns to
described by proxy variables, such as nutrient levels and
differ from the predictions of the RCC.
chlorophyll-a concentration (Dodson et al., 2000; Jeppesen et al., 2000; Declerk et al., 2005; Barnett & Beisner, 2007).
Species diversity and major environmental
However, very few studies have directly tested diversity–
gradients in lentic systems
productivity relationships in lentic macroinvertebrates. Brodersen et al. (1998) found that the species richness and
Diversity–disturbance relationships
Shannon’s diversity index of littoral macroinvertebrates
Disturbance in terms of hydroperiod has both direct and
were negatively related to these measures of productivity.
indirect influences on diversity in lentic littoral and pond
Given that the lakes they studied ranged from mesotrophic
ecosystems. Permanent ponds typically support higher
to highly eutrophic, negative relationships can be easily
numbers of species than temporary ponds, and diversity
envisaged. Chase & Leibold (2002) observed two different
increases with increasing degree of permanency (Spencer
types of species richness–productivity relationships in
et al., 1999; Urban, 2004; but see Larson, 1985). This is
pond macroinvertebrates with varying spatial scale. When
likely to be due to physical limitation on the life history
analysed with ponds as replicates, a unimodal relationship
characteristics of various taxa (Schneider & Frost, 1996;
was found in α-diversity. By contrast, when analysed over
Spencer et al., 1999), although the determination of species
ponds in a watershed, a positive relationship emerged for
richness and assemblage composition along the lentic
γ-diversity. They explained these differing relationships as
permanency gradient may be more complex (Wellborn
an indication that, in watersheds with highly productive
et al., 1996). Wellborn et al. (1996) reviewed literature on
ponds, β-diversity was higher due to multiple stable
invertebrate assemblages along this gradient and found that
states, while β-diversity was lower in low-productivity
various species occurred in either only temporary habitats,
watersheds.
only permanent but fishless habitats, or only permanent
contributed to differences in γ-diversity. Despite these
fish containing habitats. These patterns were suggested
rigorous studies, more studies in lentic ecosystems, with
to be driven by the relative roles of: (i) drying that selects
macroinvertebrates as the model group, are needed before
for traits allowing rapid development and physiological
any general conclusions about the diversity–productivity
capacity to cope with disturbance; (ii) winter oxygen stress
relationship can be made.
These differences in species turnover
that eliminates fish; and (iii) types of top predators that vary in different types of lakes, being dragonfly larvae in
Diversity–acidity relationships
permanent fishless lakes and fish in least disturbed lakes.
Diversity–acidity relationships have been relatively
The interaction between disturbance and predation should
well studied in lentic ecosystems. A typical finding has
also be manifest in overall assemblage composition and
been that the overall species richness of lentic macroin-
species richness, and different groups of aquatic insects
vertebrates decreases strongly with natural (Bradford
are likely to show differing diversity patterns along the
et al., 1998) and anthropogenic (Schindler et al., 1989)
permanency–predation gradient. This freshwater habitat
acidity. However, to my knowledge, no study has tried to
gradient offers interesting possibilities for further studies
determine the actual roles of these two sources of acidity
trying to tease out the effects of disturbance and predation
in affecting lentic macroinvertebrates. Decreasing trends
on the species richness and assemblage composition of
between species richness and acidity have been found for
aquatic insects.
both ponds (Friday, 1987) and lakes (McNicol et al., 1995; Hynynen & Meriläinen, 2005; Arnott et al., 2006). These trends are typically driven by the negative responses of
© Freshwater Biological Association 2009
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Aquatic insect biodiversity
species in a few taxonomic groups, with mayflies among
only organic bottoms and sparse cover of macrophytes at
aquatic insect taxa being generally highly sensitive to
best supported much lower species richness than forest
acidity in lakes (Meriläinen & Hynynen, 1990; Carbone et
lakes with a heterogeneous bottom of both organic and
al., 1998). However, some taxa may show no significant
inorganic material, as well as more developed macrophyte
relationships to acidity (Pollard & Berrill, 1992), and
beds.
diversity patterns opposite to those of whole assemblages
relationship between the species richness of diving beetles
may even be detected due to species benefiting indirectly
and macrophyte cover and complexity in lakes (Nilsson et
from acidic conditions (Bendell & McNicol, 1987). These
al., 1994; Nilsson & Söderberg, 1996).
This finding is in agreement with a positive
species may not necessarily prefer acidic conditions, but are excluded from neutral lakes due to biotic interactions.
Diversity–habitat size relationships
For example, fish are often excluded from acidic lakes,
Species–area relationships have been studied extensively
this providing an opportunity for macroinvertebrates
in various systems, and positive relationships have
sensitive to fish predators to thrive in acidic lakes (Eriksson
been found in a large number of studies (Rosenzweig,
et al., 1980; Bendell & McNicol, 1987; McNicol et al., 1995).
1995; Gaston & Blackburn, 2000). Habitat size has also
Among the most well-known examples in this regard are
been found to be important for the species richness and
larval dragonflies, waterstriders and waterboatmen, some
assemblage composition of aquatic insects in temporary
species of which may be more frequent and abundant in
ponds (Nilsson & Svensson, 1995; Kholin & Nilsson, 1998),
fishless habitats (Macan, 1965; Bendell & McNicol, 1995;
permanent ponds (Oertli et al., 2002) and lakes (Allen et
McNicol et al., 1995). Thus, similarly to disturbance, acidity
al., 1999; Heino, 2000). However, non-significant species
has both direct and indirect effects on species richness and
richness–habitat size relationships have also been found
assemblage composition in lentic ecosystems.
for some groups of aquatic insects in ponds (Larson, 1985; Oertli et al., 2002). For lakes, habitat size and habitat
Diversity–heterogeneity relationships
heterogeneity are typically correlated, and a positive
Habitat heterogeneity, diversity and complexity are
relationship between species richness and lake size may
potentially important determinants of diversity in lentic
thus result from heterogeneity effects. Given that habitat
littoral and pond ecosystems (Eadie & Keast, 1984; Rahel,
heterogeneity may account for more variation in species
1984). Similarly to lotic ecosystems, a number of studies
richness than lake size, heterogeneity is likely to be behind
have examined diversity–heterogeneity relationships
strong species–area relationships in surveys of lakes.
at small within-ecosystem scales.
Not surprisingly,
Similarly to species richness, the assemblage composition
these studies have typically found positive relationships
of lentic macroinvertebrates may vary strongly with lake
between macroinvertebrate diversity, abundance and
size and habitat structure (Heino, 2000).
habitat complexity, including that of macrophyte beds (Carpenter & Lodge, 1986; Brown et al., 1988), although
Multiple environmental variables determine
exceptions also exist (Cyr & Downing, 1988; McAbendroth
diversity
et al., 2005a).
Evidence about diversity–heterogeneity
relationships based on among-ecosystem surveys is more
Vinson & Hawkins (1998) called for multi-factor analyses in
limited. However, Heino (2000) found that the species
studies on aquatic insect biodiversity. Since the publication
richness of littoral macroinvertebrates was strongly related
of their review, a number of studies have examined the
to a composite habitat heterogeneity variable, comprising
relative importance of multiple environmental variables
information about substratum diversity, heterogeneity
on aquatic insect biodiversity. Such studies have been
of organic material distribution, and heterogeneity and
conducted in both lotic (Brosse et al., 2003; Sandin,
diversity of macrophytes.
2003) and lentic ecosystems (Nilsson & Svensson, 1995;
DOI: 10.1608/FRJ-2.1.1
In general, bog lakes with
Freshwater Reviews (2009) 2, pp. 1-29
14
Heino, J.
Heino, 2000). In general, most studies have noted that a
pH and water colour were the most important variables
comprehensive understanding of variation in the diversity
determining variability in both species richness and
of aquatic insects requires the measurement of multiple
assemblage composition of streams in northern Sweden.
environmental variables, including ecosystem-scale (e.g.
Wiberg-Larsen et al. (2000) also found that both the species
pH, nutrients, temperature) and habitat-scale variables (e.g.
richness and assemblage composition of caddisflies were
substratum characteristics, current velocity, macrophytes).
strongly related to stream size in Denmark. In all of these
This reasoning may be true in most cases, but exceptions
cases, the environmental gradients were apparently strong
may occur when one dominant environmental gradient
enough to modify diversity, and variability in species
has been surveyed and others are not influential. In such
richness and assemblage composition followed the same
cases, variation in diversity may be explained adequately
gradients. By contrast, in situations where there are several
by a single environmental gradient (Townsend et al.,
equally strong and uncorrelated environmental gradients,
1983; Wiberg-Larsen et al., 2000). Studies that have not
species richness and assemblage composition may vary
found significant relationships between species richness
differently along environmental gradients. This is true in
and environmental features may be complicated by a
situations where α- and β-diversity vary independently, i.e.
simultaneous survey of multiple ecosystem types (e.g.
showing an anti-nested pattern (see below).
springbrooks, headwater streams, large rivers, lake outlets; Malmqvist et al., 1999).
Surveys of different
Nestedness
ecosystem types, with each harbouring a certain suite of characteristic species, may thus lead to non-significant
Ecological assemblages within the limits of a regional
diversity–environment relationships.
Alternatively, if
species pool show a number of generalised patterns (Leibold
significant diversity–environment relationships have not
& Mikkelson, 2002; Heino, 2005a). One such pattern is
been detected, it is possible that influential environmental
nestedness. A perfectly nested pattern occurs if species in
variables were not measured at all. Biodiversity surveys
depauperate assemblages are proper subsets of those in
should therefore include a large number of explanatory
progressively more diverse assemblages, with common
variables and use multivariate methods to help to identify
species occurring in all assemblages and rare species
various factors of potential importance.
tending to occur only in diverse assemblages (Patterson & Atmar, 1986). Thus, if a set of assemblages is perfectly
Do species richness and assemblage composition vary along the same gradients?
nested, the most species-rich site includes all species detected at lower-richness sites.
Several studies have
found a significant nested subset pattern in various taxa This question is of relevance to conservation, as not only
and environments, implying that nestedness is a common
species richness (α-diversity), but also spatial changes
pattern in nature (D.H. Wright et al., 1998). However,
in assemblage composition (β-diversity) should be
the virtual ubiquity of nestedness has recently been
considered in conservation planning.
Reports of the
challenged, and only 10 % to 40 % of the matrices studied
environmental relationships of both species richness and
by Ulrich & Gotelli (2007) were actually significantly
assemblage composition have been rather unambiguous.
nested.
Not surprisingly, both these components of diversity
analyses stem from the fact that, in the former study, null
appear to respond similarly to environmental gradients.
models for significance testing were liberal (D.H. Wright et
For example, Townsend et al. (1983) found that pH was
al., 1998), whereas the latter study comprised multiple and
the most important variable for both species richness and
typically more conservative null models (Ulrich & Gotelli,
assemblage composition in southern English streams.
2007). Earlier studies on the nestedness of aquatic insects
Malmqvist & Hoffsten (2000) found that stream size,
have typically been based on the liberal null models, such
© Freshwater Biological Association 2009
These discrepancies between the two meta-
DOI: 10.1608/FRJ-2.1.1
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Aquatic insect biodiversity
as that of the nestedness calculator (Atmar & Patterson,
these environmental and organismal relationships of
1993), yet the results have been ambiguous. The published
nestedness could be easily envisaged based on information
studies on aquatic insects have found either that these
from terrestrial systems (D.H. Wright et al., 1998) and other
assemblages are significantly nested (Nilsson & Svensson,
aquatic taxa (Soininen, 2008), further studies are required
1995; Malmqvist 1999; Malmqvist & Hoffsten, 2000; Baber
to prove the generality of these findings in aquatic insects.
et al., 2004; Heino, 2005a; McAbendtroth et al., 2005b; Monaghan et al., 2005; Heino et al., 2009) or non-nested
Synthesis and implications
(Malmqvist & Eriksson, 1995; Nilsson & Svensson, 1995; Malmqvist et al., 1999; Urban, 2004). However, even when
Aquatic insect assemblages within the limits of a
significant nestedness has been found in studies of aquatic
regional species pool show many interesting patterns.
insects, it has often been rather weak. Boecklen (1997)
The relationships of species richness and assemblage
noted earlier the same pattern for aquatic invertebrates in
composition to major environmental gradients may
general, and suggested that, because these assemblages
sometimes be rather predictable, although the explanatory
contain various kinds of organisms many of which possess
power in statistical models is typically rather low. In
high colonisation rates, there is only weak nestedness at
general, habitat size and acid stress appear to structure
best. Weak nestedness means that there are somewhat
aquatic insect assemblages most strongly in both lotic
anti-nested patterns, which might mean checkerboard
and lentic ecosystems, although there are some notable
distributions of species pairs and high β-diversity (Gotelli &
differences between streams, rivers, ponds and lakes.
McCabe, 2002; Heino et al., 2009).
Understanding the species richness–environment and
Although a considerable number of studies have
assemblage
composition–environment
relationships
assessed the degree of nestedness, very few studies have
has a number of implications for the conservation and
considered the determinants of this pattern in aquatic
management of α-, β-, and γ-diversity. First, conservation
insects. McAbendroth et al. (2005b) found that nestedness
of α-diversity benefits from the understanding of species
in pond macroinvertebrates was primarily driven by
richness–environment relationships, although the degree
habitat size, and secondarily by habitat features and
of nestedness should also be considered (Boecklen, 1997).
isolation.
Interestingly, species contributing to perfect
If assemblages are perfectly nested, then the conservation
nestedness (i.e. nested species) varied in a number of
of the most species rich site protects all species. By contrast,
organismal characteristics from species not contributing
if nestedness is weak, as has been found in many studies
to nestedness (i.e. idiosyncratic species). Nested species
on aquatic insects, then rare species are not restricted
showed stronger spatial structure, weaker dispersal ability
to species rich sites only. In such cases, an alternative
and narrower ecological tolerances than idiosyncratic
conservation strategy should be followed, with interest in
species. These suggestions corroborate well the findings
both idiosyncratic species and species-rich sites. Second,
from boreal streams (Heino et al., 2009). In this study, the
given the rather predictable assemblage composition–
most important correlate of nestedness in stream insect
habitat size relationships, different-sized streams and lakes
assemblages was stream size. Furthermore, idiosyncratic
are needed to conserve β-diversity. Thirdly, the protection
species occurred, on average, at more sites than nested
of sites supporting characteristic assemblage types and
species, mirroring the restricted distributions of several
sites at different parts of environmental continuums is also
nested species that were inclined towards species-rich
important for the conservation of γ-diversity.
sites. Idiosyncratic and nested species also differed in niche position and niche breadth, with idiosyncratic species having less marginal niche positions and wider niches than nested species (Heino et al., 2009). Although DOI: 10.1608/FRJ-2.1.1
Freshwater Reviews (2009) 2, pp. 1-29
16
Heino, J.
Among-taxon congruence in diversity
invoked
similar
environmental
responses
as
the
mechanism contributing to co-varying patterns of assemblage composition in pond insects. By contrast,
Understanding the degree to which different taxonomic
Heino et al. (2003b) stated that, due to differing responses
groups show congruent biodiversity patterns is an
to geographical location and major environmental
important field of study in conservation biology (Gaston,
gradients, stream insect taxa showed weak congruence in
1996b) and environmental assessment (McGeogh, 1998).
biodiversity. Thus, the responses of different taxonomic
However, despite its importance, relatively few studies
groups to the same environmental gradients are most likely
have examined congruence in the diversity patterns of
to be responsible for the degree of congruence. Responses
aquatic organisms in general (Allen et al., 1999; Paavola et
to correlated environmental factors may also contribute to
al., 2003; Heino et al., 2005) and aquatic insects in particular
congruence in some situations, whereas biotic interactions
(Briers & Biggs, 2003; Heino et al., 2003b; Bilton et al.,
and random draw are less likely explanations at the grain
2006). The studies that have appeared thus far have led to
of local ecosystems and the extent of single drainage basins.
differing conclusions. Heino et al. (2003b) found that the
However, random draw from the regional species pool
species richness and assemblage composition patterns of
may contribute to strong congruence if local ecosystems are
stream mayflies, stoneflies, caddisflies and midges were
surveyed across long geographical gradients (Heino et al.,
weakly, though sometimes significantly, correlated at the
2003b). These suggestions are speculative, and given that
across-ecoregions scale in Finland. Within ecoregions, a
very few studies on aquatic insects have considered among-
few cross-taxon correlations were slightly higher, but even
taxon congruence, further studies in various regional and
these correlations lacked the predictive power required
environmental settings are needed to resolve this question.
for them to be useful in practice. These findings generally
Knowing the degree to which different taxonomic
comply with those for the same four insect groups in
groups show congruent diversity patterns is potentially
a single boreal drainage basin (Heino & Mykrä, 2009).
valuable in the conservation context.
Bilton et al. (2006) also studied among-taxon congruence
showing strongly congruent patterns with other taxonomic
in species richness and assemblage similarity in several
groups, a good biodiversity indicator taxon should possess
pond macroinvertebrate taxa. They found that, although
a number of additional qualities. These include occurrence
species richness was weakly correlated among taxonomic
in various environmental conditions, cost-efficiency,
groups, assemblage composition patterns showed stronger
applicability at different scales, and it should be easily
among-group correlations. Sánchez-Fernández et al. (2006)
sampled, easily identified and ecologically well-known
found that aquatic beetles were relatively good indicators
(Noss, 1990; McGeogh, 1998). The concept of biodiversity
of species richness in various stream macroinvertebrate
indicator taxa has only rarely been considered in aquatic
taxa, with correlations ranging from non-significant for
insect studies. Briers & Biggs (2003) found that coenagriid
molluscs and mayflies to significant and rather strong for
damselflies and limnephilid caddisflies were the most
aquatic bugs, stoneflies, caddisflies and total richness of the
promising groups as indicators of pond biodiversity.
studied taxonomic groups.
Bilton et al. (2006) considered aquatic beetles as the most
Why should some taxonomic groups show strong correlations in biodiversity?
In addition to
useful indicator group for pond biodiversity assessment,
There are four main
because they conform best of the studied taxa to the
mechanisms behind strong among-taxon congruence:
qualities listed above. Similarly, Sánchez-Fernández et al.
(i) similar environmental responses; (ii) responses to
(2006) also regarded aquatic beetles as a good indicator
different, but correlated environmental features; (iii) biotic
taxon. However, the degree to which aquatic beetles
interactions; and (iv) random draw from the regional
perform as surrogates in the biodiversity assessment of
species pool (Gaston & Williams, 1996). Bilton et al. (2006)
streams and lakes is uncertain, and recent research on
© Freshwater Biological Association 2009
DOI: 10.1608/FRJ-2.1.1
17
Aquatic insect biodiversity
boreal bog and forest lakes suggests that aquatic beetles
A few studies have examined functional biodiversity
are so sporadically distributed that they cannot be used
along
multiple
non-anthropogenic
environmental
effectively as an indicator taxon (Heino et al., unpublished).
gradients.
It is thus likely that there is no single, universal indicator
describing variation in the functional structure of stream
taxon among aquatic insect taxa, and that indicator
insect assemblages of mountain streams. They found that
groups need to be adjusted according to regional and
functional structure varied primarily with periphyton
environmental settings. This may also mean that indicator
biomass and particle size, although the longitudinal
groups are unlikely to be useful in most situations.
position was also important. Heino (2005b) concentrated
Finn & Poff (2005) used nine traits for
on the macroinvertebrates of boreal lowland streams,
Patterns of functional diversity
and used a combination of functional feeding groups (Cummins, 1973) and habit trait groups (Merritt &
Although the functional approach has a long history in
Cummins, 1996). Functional feeding groups refer to the
ecology (Statzner et al., 2001a), there has been increasing
feeding modes and approximate food type of macroinver-
interest in the topic recently due to greater recognition of
tebrates, whereas information on where food is obtained
the importance of functional biodiversity to ecosystem
(e.g. on stones versus within sediments) is used to derive
functioning (Kinzig et al., 2002). Early examinations of
habit trait groups.
macroinvertebrate functional feeding groups (Cummins,
spatially-structured and environmentally-related variation,
1973; Cummins & Klug, 1979) in the context of the river
with pH, stream size and moss cover being among the
continuum concept provided first glimpses of the utility
most important variables. Three measures of functional
of the functional approach when studying the longitudinal
diversity (functional richness, diversity and evenness)
gradient in lotic ecosystems (Vannote et al., 1980; Minshall
decreased with water acidity, canopy cover and water
et al., 1985a; Minshall, 1988). By contrast, few studies have
colour, and increased with moss cover. In a similar study
examined patterns in the functional structure and diversity
on the functional biodiversity of lake littoral macroinverte-
of freshwater macroinvertebrate assemblages along
brates, a general finding was that functional structure and
multiple non-anthropogenic environmental gradients,
diversity varied strongly along gradients in macrophyte
although the need for such studies has been clearly
cover and lake area (Heino, 2008a). The above findings
identified (Poff, 1997; Usseglio-Polatera et al., 2000). Using
were somewhat surprising, because some authors have
functional structure, or more generally species traits, as
suggested that the trait structure of assemblages varies only
response variables instead of taxonomic composition may
little along natural environmental gradients, and virtually
increase our understanding of the relationship between
no variation is related to geographical location (Charvet et
freshwater biodiversity and ecosystem functioning.
al., 2000; Statzner et al., 2001b). These discrepancies between
However, although most recent studies have certainly
studies are apparently due to the types of functional traits
addressed multiple important traits (Gayraud et al.,
used and analytical methods. However, despite these
2003; Lamouroux et al., 2004; Poff et al., 2006), there has
methodological differences, theory suggests that functional
typically been rather little division between the traits that
traits should be strongly related to environmental gradients,
are truly functional in terms of ecosystem functioning
be they of natural or anthropogenic origin (Poff, 1997).
Functional structure showed both
and those that portray various life history characteristics
A question pertaining to the utility of functional
of stream macroinvertebrates. Yet, directly emphasising
traits in the description of freshwater macroinvertebrate
the relationships between particular traits and ecosystem
biodiversity has been tackled recently: do functional traits
functioning might further increase our understanding of
provide more and complementary information to that
freshwater ecosystems.
provided by species? This question is important, because if there are no clear differences in assemblage patterns
DOI: 10.1608/FRJ-2.1.1
Freshwater Reviews (2009) 2, pp. 1-29
18
Heino, J.
between the functional trait and taxonomic approaches,
found that the taxonomic distinctness of aquatic beetle
then it may not be necessary to allocate taxa to various
assemblages responded more weakly than species richness
functional categories.
Recent studies have suggested
to anthropogenic alteration of inland waters in Spain. This
that both functional groups and taxa show relatively
finding is, in part, in accordance with those from Finland,
similar assemblage patterns along major environmental
where neither taxonomic distinctness nor species diversity
gradients (Finn & Poff, 2005; Heino et al., 2007a).
indices varied between near-pristine and anthropogenically
Furthermore, diversity measures based on functional
altered streams (Heino et al., 2007b). However, because
groups (e.g. number of functional groups) have been
taxonomic distinctness and species diversity indices were
found to be strongly related to similarly derived taxonomic
at best weakly correlated, both groups of indices should
measures (e.g. number of species) in both lentic and lotic
be used in biodiversity assessment. By contrast, Marchant
macroinvertebrates (Heino, 2008a; Heino et al., 2008).
(2007) found that taxonomic distinctness values of stream insect assemblages from anthropogenically altered sites
Patterns in the taxonomic variability of assemblages
were often below expectations of random draws of species from the species pools (see also Clarke & Warwick, 1998), and taxonomic distinctness was also negatively correlated
Species richness and diversity indices have traditionally
with a water quality gradient that described variation in
been used to assess variability in biodiversity across
turbidity and total phosphorus. Thus, it was suggested that,
near-pristine and anthropogenically altered ecosystems
due to a high number of species and taxonomic variability
(Magurran, 2004). However, it has been suggested that
in the dataset, a taxonomic distinctness index was able to
biodiversity measures should not only portray variability in
separate near-pristine and anthropogenically altered sites
species richness and relative abundances, but also take into
(Marchant, 2007). Accordingly, taxonomic distinctness
account the evolutionary relatedness of species (Harper &
indices in the other two studies on aquatic insect assemblages
Hawksworth, 1994). For this purpose, Clarke & Warwick
may not have performed well due to a more limited
(1998) devised a set of taxonomic distinctness indices that
number of species or higher taxa. Given that measures
take into account the taxonomic relatedness of species.
of taxonomic variability of assemblages are important
Thus, an assemblage that harbours distantly-related
in biodiversity assessment, not only environmental
species from, for example, different phyla is more diverse
researchers but also biodiversity researchers should
than an assemblage with the same number of species from
include taxonomic distinctness indices in their toolbox.
just one family. Taxonomic distinctness indices have been widely used in marine studies (Warwick & Clarke, 1998; Rogers et al., 1999; Ellingsen et al., 2005), but research on
Higher taxon surrogates for species-level patterns
the taxonomic variability of assemblages has just begun with aquatic insects.
The higher taxon approach in biodiversity research has
Three recent studies on aquatic insects have addressed
usually been directed at examining genus and family
taxonomic variability based on the taxonomic distinctness
richness as surrogates for species richness. These studies
indices. These studies have not only considered variability
have generally concluded that the numbers of higher taxa
in near-pristine ecosystems, but have also examined
are typically strongly correlated with species richness
biodiversity along anthropogenic gradients. Campbell &
(Williams & Gaston, 1994; Balmford et al., 2000; Gaston,
Novelo-Gutierrez (2007) studied the response of taxonomic
2000). By contrast, very few studies have directly examined
distinctness of odonate assemblages to dam impoundment
cross-taxonomic level patterns in assemblage composition,
and detected reduced values after the construction
so it is uncertain if patterns in assemblage composition are
of hydroelectric impoundment.
similarly strongly congruent between taxonomic levels as
Abellán et al. (2006)
© Freshwater Biological Association 2009
DOI: 10.1608/FRJ-2.1.1
19
Aquatic insect biodiversity
with taxon richness patterns. It has been suggested that
Conclusions
assemblage patterns between different taxonomic levels are strongly congruent in regions where there are low
Biodiversity
species-to-genus and species-to-family ratios, whereas
assemblages are complex. This complexity is seen in
patterns
exhibited
by
aquatic
insect
congruence diminishes in regions with high species
the environmental relationships of both species richness
diversity, resulting from high degrees of adaptive radiation
and assemblage composition.
(Hawkins & Norris, 2000). These suggestions remain to be
environmental variables are needed to understand
studied rigorously with aquatic insects.
variation in the diversity of aquatic insects. Although
In general, multiple
Freshwater research has traditionally concentrated
a few major environmental gradients, such as habitat
on examining the importance of taxonomic resolution
size and acidity, typically account for a modest amount
in bioassessment, whereas few biodiversity studies
of variability in aquatic insect assemblages, other minor
have addressed higher taxa as surrogates of species-
factors are also often required to understand variation in
level diversity. A number of lessons can be drawn from
diversity. This is especially true in cases where there is no
bioassessment studies. First, species richness has often
single dominant environmental gradient which has an
been found to be correlated with genus and family richness
overriding effect on biodiversity. Knowing assemblage–
(Guerold, 2000; Hill et al., 2001; Marshall et al., 2006). Also,
environment relationships is valuable for conservation
patterns of assemblage composition of different taxonomic
planning, and generally sites along different parts of major
levels are typically strongly correlated (Bowman &
environmental gradients should be protected to conserve
Bailey, 1997; Marshall et al., 2006; Heino, 2008b). Second,
α-, β- and γ-diversity of aquatic insect assemblages (Clarke
higher taxa often respond similarly to species to major
et al., 2008).
environmental gradients (Furse et al., 1984; Marchant
However, our knowledge of various aspects of aquatic
et al., 1995; Feio et al., 2006). Third, although genus- and
insect biodiversity is woefully incomplete. For example,
family-level identifications of stream macroinvertebrates
understanding one of the oldest patterns in ecology, the
are less expensive and time-consuming than species-
latitudinal diversity gradient (Hawkins, 2001), is severely
level identifications (Marshall et al., 2006), the latter are
hindered by the lack of both distributional data at broad
necessary when the interest is in community ecology and
scales and comprehensive surveys of aquatic insect
species distribution for conservation purposes (Furse et
assemblages across long south–north gradients. Achallenge
al., 1984; Bailey et al., 2001; Lenat & Resh, 2001). Studies
to future studies is thus to survey local assemblages along
addressing the higher taxon approach for biodiversity
latitudinal gradients. More than that, researchers should
and conservation purposes have generally corroborated
aim at comparing the relative importance of geographical
the findings from bioassessment research, with patterns
position, regional factors, watershed characteristics, and
in taxonomic richness and assemblage composition being
ecosystem and habitat variables for aquatic insect diversity
highly similar at species, genus and family levels (J.F. Wright
across large geographical extents. There are emerging
et al., 1998; Heino & Soininen, 2007). Thus, because higher
possibilities for such studies in Europe, and recent research
taxa show patterns similar to those of species, they can be
on streams has already touched this topic (Johnson et al.,
used in rapid biodiversity assessments, the goal of which
2007). However, there are severe problems in studies of
is to assess and rank a large number of sites according to
local ecosystems across large geographical extents. One
their relative biodiversity value. Such approaches may
such problem relates to the species-level identification of
be the only option for studies in tropical regions lacking
the aquatic larval stages of insects. Given that taxonomic
taxonomic information at the species level, and higher taxon
knowledge is unevenly distributed among countries,
surrogates have thus been used as estimates of species-level
large-scale surveys in poorly known areas could utilise
biodiversity patterns (Jacobsen et al., 1997; Jacobsen, 2004).
higher taxa as surrogates for local species-level diversity
DOI: 10.1608/FRJ-2.1.1
Freshwater Reviews (2009) 2, pp. 1-29
20
Heino, J.
patterns at least in the early phases of inquiry into large-
headwater lakes. Streams and ponds have been used as
scale patterns (Jacobsen, 2004).
Such surrogates have
arenas for testing ecological theory for a long time, and
proved promising in previous environmental assessment
given the easier sampling of small than large freshwater
and biodiversity studies (J.F. Wright et al., 1998).
ecosystems, this emphasis on streams and ponds is likely
In addition to species richness and assemblage
to continue. However, because freshwater biodiversity
composition, other measures of biodiversity should
is in severe crisis (Dudgeon et al., 2006), we should
be given more attention in research on aquatic insect
have no excuse for neglecting surveys of large rivers
assemblages. For example, it is still a largely unsettled
and headwater lakes across extensive regions.
question as to whether measures of the taxonomic variability
by knowing biodiversity patterns in various types of
of assemblages are useful in assessing environmental
freshwater ecosystems, will we be able to predict, conserve
degradation of biodiversity in freshwater ecosystems.
and manage aquatic insect biodiversity effectively enough.
Only
Both negative and positive results have emerged from studies that have used this approach (Heino et al., 2007b;
Acknowledgements
Marchant, 2007). However, even though measures of taxonomic variability of assemblages would not perform
The author wishes to thank Kai Korsu, Timo Muotka,
well in determining anthropogenic degradation of
Janne Soininen, Colin S. Reynolds, and two anonymous
freshwater ecosystems, the fact that they are at best weakly
reviewers for constructive comments on earlier versions of
correlated with various other measures of biodiversity
this paper. The preparation of this paper was supported by
suggests that they should be utilised more widely in
a grant from Maj and Tor Nessling Foundation.
studies on aquatic insect assemblages (Heino et al., 2008). Functional biodiversity should also be given more
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Author Profile Jani Heino is a researcher working at the Finnish Environment Institute, Research Programme for Biodiversity. He received his PhD from the University of Jyväskylä in 2002. His current research lies at the nexus of large-scale ecology, community ecology, landscape ecology and conservation biology, with the main aim to understand spatial and temporal variation of biodiversity across and within aquatic ecosystems. He also uses the information on ecological patterns in guiding the conservation and management of aquatic ecosystems. His study subjects comprise a wide range of aquatic ecosystems (e.g. streams, rivers, littorals) and groups of organisms (e.g. invertebrates, bryophytes, algae). Freshwater Reviews (2009) 2, pp. 1-29