Hydrobiologia 462: 115–129, 2001.

© 2001 Kluwer Academic Publishers. Printed in the Netherlands.

115

Distribution of macroinvertebrates in relation to physical and biological

variables in the littoral zone of nine New Zealand lakes

Mark A. Weatherhead & Mark R. James1

National Institute of Water and Atmospheric Research Ltd, P.O. Box 8602, Christchurch, New Zealand
Tel: +64-3-348-8987. Fax: +64-3-348 5548. E-mail: m.weatherhead@niwa.cri.nz
1 Present Address: NIWA, P.O. Box 11-115, Hamilton, New Zealand

Received 9 October 2000; in revised form 24 July 2001; accepted 5 August 2001

Key words: macroinvertebrates, lake littoral zone, macrophytes, substrate, regression models, New Zealand

Abstract
Macroinvertebrates play a key role in the littoral zone of lakes. Macroinvertebrate community composition is
closely linked to habitat conditions. To date, there have been few attempts to relate macroinvertebrates to habitat
factors in lakes. In this study, nine mainly oligotrophic lakes from throughout New Zealand were surveyed for
macroinvertebrates. The lakes were selected to represent a range of suspended sediment loading and lake level
regimes. Within each lake, several sites were selected to provide a range of exposure to wave action. A multiple
regression approach was taken to relate macroinvertebrate community composition and habitat characteristics. The
results of the analysis suggest that the littoral zone of the lakes we studied could be divided into four general
habitats. The first is the wave wash zone characterised by coarse substrates and macroinvertebrate taxa usually
associated with lotic environments, such as Ephemeroptera and Plecoptera. The second habitat is associated with
macrophytes and is limited at the top by wave action and at depth by light attenuation. In this zone, the snail
Potamopyrgus antipodarum is dominant, along with Trichoptera and Odonata. At the base of the macrophytes is
the detrital habitat characterised by fine, organic rich sediments and dominated by chironomids, oligochaetes and
Trichoptera. At depths below the macrophyte zone, fine sediments are found, and bivalves such as the freshwater
mussel Hyridella menziesi are common. While macroinvertebrate abundance can be highly variable, some general
predictions of community structure can be made based on a few key environmental factors. Abundance of snails
Odonata and Trichoptera was positively related to macrophyte biomass. Some macroinvertebrate groups such as
oligochaetes, chironomids, snails and bivalves were more common in fine substrates, while Ephemeroptera were
characteristic of coarse substrates. Detrital biomass was important for most of the macroinvertebrate groups studied
showing a positive relationship for oligochaetes and Trichoptera and a negative relationship for Ephemeroptera and
Plecoptera.

Introduction ditions such that detritivores (oligochaetes and certain

Macroinvertebrates play a key role in the littoral zone
of lakes. They potentially control the biomass of
periphytic algae (Hawes & Schwarz, 1996; James et
al., 2000a), recycle detrital material (Kornijów et al.,
1995) and provide a critical link from primary produc-
tion through to fish, thus underpinning lake fisheries
(Graynoth et al., 1986). The macroinvertebrate com-
munity composition is closely linked to habitat con-
chironomid taxa) are characteristic of organically rich
sediments under macrophytes, while Ephemeroptera
and hydroptilids are more characteristic of shallow
wave-washed cobbled habitats (James et al., 1988).
These characteristic communities, however, can be
modified by other physical and biological factors.
Physical factors such as exposure to waves and the
range of lake level fluctuations can have direct effects
on the macroinvertebrate community through physical
116
Figure 1. Hypothetical physical and biological interactions in the littoral zone of lakes.
disturbance and indirect effects which are mediated
through effects on the distribution of macrophytes and
associated epiphytic algae.
There have been a number of attempts to relate
macroinvertebrate distribution and community com-
position to physical and biological variables, but these
have generally been limited to only a few variables.
For example, Biggs & Malthus (1982) and Van den
Berg et al. (1997) showed a relationship between
macroinvertebrate community composition and bio-
mass and the type of macrophyte (vascular vs. char-
acean) present. In an earlier paper on the macroin-
vertebrate community in Lake Coleridge, South Island
(James et al., 1998), it was determined that substrate,
macrophyte biomass and shoreline profiles were im-
portant determinants of macroinvertebrate distribution
but this could be modified by slope and exposure
to wave action. Brodersen (1995) found periphyton
abundance and slope were more important than ex-
posure to wave activity in determining macroinver-
tebrate abundance and distribution in the surf zone of
mesotrophic Lake Esrom, Denmark.
There have been very few attempts to define the
major variables influencing macroinvertebrate distri-
bution. This has limited our ability to predict the im-
pacts of various anthropogenic activities on macroin-
vertebrate communities and ultimately on lake fisher-
ies that depend on this food source. Habitat suitability
models have been developed for some of the major
macroinvertebrate taxa found in lotic systems (e.g.
Bovee, 1986; Morin et al., 1986; Jowett & Richardson,
1990). The best models for these systems generally
showed that substrate, water velocity and depth were
the most useful predictors for taxa such as Delea-
tidium. Despite our increased knowledge of the in-
teraction of physical variables and macroinvertebrate
communities in rivers, there have been no attempts
that we are aware of to develop similar habitat suit-
ability models for lakes where factors such as wave
exposure are likely to play a major role and other
factors such as velocity are not so important.
The way in which some of the physical processes
are mediated through physical and biological interac-
tions to influence macroinvertebrates is demonstrated
schematically in Figure 1. From this we have hypo-
thesised that:
1. The key direct determinant of macroinvertebrate
community composition is substrate type, which
in turn is determined by variables such as exposure
to wave activity.
2. Macrophyte biomass is the second most import-
ant determinant with distribution of macrophytes
largely determined by substrate type and macro-
phyte biomass and bottom limits by light attenu-
ation from suspended sediments.
3. Lake level history is a secondary factor.
To test these hypotheses, we have surveyed a num-
ber of oligotrophic and mesotrophic lakes throughout
New Zealand covering a matrix of suspended sediment
levels and lake level fluctuations. Within each lake,

Figure 2. The location of the lakes used in this study. Further detail
on the physical parameters of the lakes is shown in Table 1.
several sites were selected to cover a range of exposure
to wave activity, proximity to river inputs and slope.
The specific aims of this paper were to:
1. Describe the physical and biological variables
important in structuring macroinvertebrate com-
munities in the littoral zone of oligotrophic lakes
in New Zealand.
2. Develop simple models that could be used to
predict community structure and abundance of
different taxonomic groups.
Site descriptions
Physical characteristics and bottom limits for macro-
phytes in the lakes surveyed are given in Table 1 and
their location is shown in Figure 2. Nine lakes were
sampled; three in the North Island (Taupo, Waikare-
mona, and Rotoaira) and six in the South Island
(Coleridge, Tekapo, Hawea, Wanaka, Te Anau and
117
Ruataniwha). The lake sizes ranged from 3.5 km2 (Ru-
ataniwha) up to 622 km2 (Lake Taupo, New Zealand’s
largest lake). All of the South Island lakes studied had
some degree of glacial input including glacial flour,
while the North Island lakes had riverine or stream
inputs. In the case of Lakes Tekapo and Ruataniwha
(which receives water from Lake Tekapo), the glacial
flour load reduced secchi disk depth to no more than
2 m during the study period. All of the lakes studied
were oligotrophic except for Lake Rotoaira which was
mesotrophic.
Methods
General
Up to 11 sites were surveyed in each of the nine lakes
(a total of 51 sites) that were chosen to cover a range
of lake level regimes and suspended sediment concen-
trations (see Table 1). The sites surveyed in each lake
were chosen to include different degrees of exposure
to prevailing winds and strong wave action, differing
proximity to diversions for hydroelectric storage and
varied shoreline topography. Sampling depths varied
between the sites and lakes depending on the type of
habitats encountered in the littoral zone down to a
maximum depth of 18 m below mean water level at
any site. Details of sampling are given in Table 2.
Macroinvertebrates
Three to five replicate macroinvertebrate samples were
taken at each site and depth. Three types of sampling
devices were used to collect macroinvertebrates: in
shallow regions where the substrate was mostly
cobbles Surber samplers (0.10 m2 , mesh 0.4 mm) were
used; for finer sediments and mud/macrophyte hab-
itats, Wisconsin grab samplers (0.10 m2 , mesh 0.4
mm) were used; at sites with predominantly cliffs or
large boulders, samples were scraped from an area of
33 cm2 into jars (‘Pottle’ method in Table 2). Surber
samplers are generally used in flowing waters, but
in the present study we carefully collected cobbles
from a known area into the mesh bag. All debris and
rocks collected in the area enclosed by the Surber
sampler were brushed clean and dislodged material
was collected on a 0.5 mm mesh sieve.
All samples were preserved in formalin (4% final
concentration) in the field. Samples containing large
volumes of sand were sorted in the field by repeated
washing of sediments and passing the water through
118
Table 1. Physical characteristics and bottom limits for macrophytes in the lakes surveyed

Lake Area Altitude Max. Water Secchi Macrophte Water Trophic Status
(km2 ) (masl) depth level depth bottom sourceb
(m) range (m) (m) limit (m)a

Coleridge 32.9 507 200 1.13 13.0 33 Glacial Oligotrophic

Hawea 137.6 347 384 2.73 11.7 34.4 Glacial Oligotrophic
Rotoaira 15.3 564 15 0.40 4.5 15 Riverine Mesotrophic
Ruataniwha 3.4 460 28 0.30 0.6 3.7 Glacial Oligotrophic
Taupo 611 357 163 1.24 12.5 16 Riverine Oligotrophic
Te Anau 347.5 203 417 2.74 10.0 13.7 Glacial Oligotrophic
Tekapo 86.8 708 120 2.94 2.0 0 Glacial Oligotrophic
Waikaremoana 55.7 583 248 2.55 11.5 17.3 Riverine Oligotrophic
Wanaka 180.1 277 176 2.11 9.9 23.6 Glacial Oligotrophic
a After Schwarz et al. (2000).
b From Livingston et al. (1986a, b).

Table 2. Sampling regime and number of sites for each lake surveyed

Lake Date No. sites Depth range Replicates Sampling methoda

Coleridge Feb 95 8 0.5–18 3 S,W

Hawea Mar 99 5 0.5–14 3 S,W
Rotoaira Oct 98 4 0.5–10 4 W,P
Ruataniwha Feb 97 5 0.5–18.3 3 S
Taupo Sep 98 11 0.5–4 5 S,W,P
Te Anau Feb 99 5 0.5–5 3 S,W
Tekapo Mar 99 4 0.5–8 3 S,W
Waikaremoana Mar 98 4 0.5–20.5 3 S,W
Wanaka Feb 99 5 0.5–9 3 S,W

a S = Surber, W = Wisconsin, P = Pottle.

a 0.5 mm sieve to collect organisms. On return to the
laboratory, benthic macroinvertebrates were separated
from macrophytes and detritus using an inclined, run-
ning water trough that drained through a 0.5 mm mesh
sieve (Allan, 1951). Macrophytes and detritus were
removed from the preserved samples and the macro-
phytes separated into vascular plants and charophytes.
The dry weights (DW) of each type of macrophyte and
detritus were determined after drying at 80 ◦ C for 24
h. Large samples were subdivided (12.5–50%) using
a barrel sample splitter with a median vertical parti-
tion. Insect larvae were identified from keys in Cowley
(1978) and Winterbourn & Gregson (1989). Number
of taxa per site, total abundance, and abundances of
individual taxa per m2 were calculated.
Substrate
Substrate composition was recorded for particles col-
lected in the Wisconsin grabs and from diver obser-
vations. Substrates were classified after Jowett et al.
(1991) according to the following criteria: <0.5 mm
mud/silt, 0.5–2 mm sand, 2–64 mm gravel, 65–256
mm cobbles and >256 mm boulders. For statistical
analyses, substrate composition was converted to a
substrate index (SI) (Suren, 1996) where:
SI = 0.07×% boulder + 0.06×% cobble +
0.05×% gravel + 0.04×% sand + 0.03×% mud/silt
This gives a value ranging from 3 for mud/silt to 7
for boulders.
Exposure
Exposure to wind-generated wave action was calcu-
lated using a wind-wave hindcast model designed for
restricted fetch, enclosed waterways (e.g. lakes and
harbours). Wave generation is based on the NARFET
(NARrow FETch) model (Smith, 1991). Shorelines
were extracted from GIS layers at 50–200 m resol-

ution. Wind information was collected from national
wind station data records. Where possible, data from
the previous 12 months was used from the site closest
to the lake. In some cases, recent data was not avail-
able, so historic data (typically 3–5 years of data)
was used to estimate typical conditions. We used his-
torical data from the lake where possible rather than
current data from a more distant point. In the case of
Lake Waikaremoana, there was no present or historical
wind station within 50 km of the lake so wind data
from Lake Taupo was used. For each point around
the lakeshore, a number of wave parameters were
derived from the model. For this analysis, we used
average wave-base depth as a measure of the depth of
disturbance caused by waves.
Secchi depth and water level fluctuations
Secchi disk depths were taken from the literature and
unpublished sources and are shown in Table 1. Lake
level ranges were obtained from NIWA’s lake level
database. Where possible secchi depths and lake level
ranges were obtained for the year prior to the surveys
with lake level range for previous 3, 6 and 12 months.
The complex inter-relation between depth, sub-
strate, plant biomass, detritus, and the macroinverteb-
rates shown schematically in Figure 1 is reflected in
Table 3 showing the results of a correlation analysis of
the physical factors.
Statistical analyses
The macroinvertebrates were collected over several
years as part of a number of different studies. Most of
the sampling was carried out in February and March
(Table 2). In order to minimise the variation caused
by using data from different studies, the inverteb-
rates were grouped to order. The exceptions were the
gastropod Potamopyrgus antipodarum which is very
common and easily identified, and the Hydroptilidae
family of caddisflies which are often found at the top
of the shore in the wave zone and again are very easy
to identify.
The statistical analysis was approached in two
ways. First, a multiple regression analysis of total
macroinvertebrate abundance vs. the physical factors
was carried out following log (x + 1) transforma-
tion using Statistica 5.5 (Statsoft, Tulsa Okl.). The
aim of this analysis was to indicate those factors
which regulated the macroinvertebrate community.
Further regression analysis was undertaken for the
most common groups: oligochaetes, Trichoptera, P.
119
antipodarum (the most common gastropod), chiro-
nomids, bivalves and Ephemeroptera. This analysis
was directed at determining the relationships between
the most common macroinvertebrate groups and the
physical variables.
The second analysis examined macroinvertebrate
community relationships using DECORANA (PC-
ORD). This analysis showed which macroinvertebrate
groups were most similar in their habitat preferences.
The resulting ordination axis values were also com-
pared to the physical variables using Pearson cor-
relations to determine which physical factors were
correlated to the observed clusters based on macroin-
vertebrate communities.
Results
Physical variables
Average wave base depth (AWBD) was highly vari-
able over the lakes surveyed ranging from 0.2 m to 1.9
m (see Table 4). As would be expected, the smallest
lake (Ruataniwha) had the shallowest AWBD reflect-
ing the short fetch at this lake, interestingly the other
small lake (Rotoaira) had a similar AWBD range to the
other much larger lakes. The greatest AWBD (1.9 m)
was found at the southern most end of Lake Coleridge
which is exposed to the very strong north-west winds
characteristic of that part of New Zealand.
Substrates were generally coarser at depths less
than 4 m on most shorelines. When we compared
substrate index within the wave zone (i.e. at depths
less than the AWBD) to those below this zone, we
found that substrate index was significantly higher (i.e.
coarser) within the wave zone (t = −11.8, df = 382,
P <0.0001) averaging 5.6 ± 0.8 compared to 3.9 ± 1.1
below the wave zone. Figure 3 shows a box-whisker
plot of substrate index vs. depth for all of the lakes and
sites surveyed – note that depth has been binned into 2
m bands. The substrate at depths below 4 m was either
fine silt/mud or silt overlying coarser substrates. This
is a feature that was common to most sites although
a few sites in Lakes Coleridge and Taupo had coarse
substrate below the macrophyte zone.
Maximum macrophyte biomass ranged from 7 g
m−2 at Lake Tekapo up to over 2100 g m−2 at Lake
Waikaremoana (Table 4). The depth of maximum
macrophyte biomass in general ranged from 2 to 3.8
m although at Lakes Coleridge and Hawea maximum
biomass was at 10.6 and 8.8 m, respectively. Max-
imum detrital biomass ranged from 66 g m−2 in Lake
120
Table 3. Pearson correlations between the physical variables studied. n = 384, ∗ = p < 0.05

Depth SI Lake Secchi Average Charo- Vascular

range wave phytes
base

SI −0.474∗

Lake range 0.045 −0.057

Secchi 0.114∗ −0.071 0.171∗

Average wave base −0.013 0.171∗ 0.016 0.343∗

Characeans 0.037 −0.269∗ 0.171∗ 0.113∗ −0.033

Vascular −0.115∗ −0.132∗ −0.080 −0.094 −0.001 −0.017

Detritus −0.049 −0.111∗ −0.111∗ 0.126∗ 0.103∗ 0.051 0.166∗

Table 4. Summary of physical characteristics found in the surveys of each lake

Lake Substrate Average Maximum Average Maximum Average

index wave base macrophyte depth of detrital depth of
range range (m) biomass macrophyte biomass detrital
(g m−2 ) maxima (m) (g m−2 ) Maxima (m)

Coleridge 3.0–6.8 0.3–1.9 534 10.6 1289 5.8

Hawea 3.0–6.7 0.6–1.5 115 8.8 259 7.2
Rotoaira 3.0–6.5 0.7–1.0 1788 3.8 618 3.1
Ruataniwha 3.0–5.0 0.2–0.5 118 2.7 - -
Taupo 3.0–7.0 0.7–1.8 1299 2.3 929 1.7
Te Anau 3.5–6.3 0.6–0.7 661 3.8 2519 2.2
Tekapo 3.0–6.6 0.5–1.0 7 2.0 66 5.6
Waikaremoana 3.0–5.9 0.3–1.4 2123 3.0 460 2.6
Wanaka 3.0–6.8 0.7–1.2 541 2.4 289 1.8

NB: No detrital biomass assessments were made at Lake Ruataniwha.

Tekapo to 2500 g m−2 in Lake Te Anau where large
amounts of allochthanous beech leaves can be found
at some sites. The depth of maximum detrital biomass
ranged from 1.7 m to 7.2 m.
Invertebrate communities
The invertebrate communities encountered in the sur-
veys were typical of those in New Zealand lakes. The
snail Potamopyrgus antipodarum was the dominant
macroinvertebrate in most lakes, both in terms of num-
bers and biomass. P. antipodarum was particularly
common in the macrophyte beds and was found in
very high numbers in Lakes Rotoaira (up to 180 000
per m2 ) and Waikaremoana (up to 73 000 per m2 ).
Mayflies were rare and only present in Lakes Col-
eridge (up to 93 per m2 ), Tekapo (up to 141 per m2 ),
Taupo (up to 52 per m2 ) and Waikaremoana (up to
14 per m2 ). The number of taxa found ranged from
2 to 13 taxa per site, and was especially low in Lake
Tekapo (2–5 taxa on average). Plant biomass was also
very low in Lake Tekapo with a very patchy distribu-
tion. This is probably due to the high turbidity caused
by input of glacial flour to the lake. At the other end
of the scale, Lakes Waikaremoana and Rotoaira had
very high macrophyte biomass with some plant beds
up to 3 m deep (G. Kelly, NIWA, pers. comm.). These
 121

Figure 3. The relationship between substrate index and depth. Note that depths have been pooled into 2 m bands.

Table 5. Summary of the Multiple Regression analyses e.g. Chironomids = 7.781 – 0.190×Substrate – 0.466×Average Wave Base +
0.037×Secchi + 0.216×Detritus. All macroinvertebrate taxa, macrophyte biomass and detrital biomass have undergone log (x+1) transformation.
∗ = P <0.05, ∗∗ = P < 0.01

Taxa Intercept B coefficient values (only significant shown) r2

Substrate Depth Average Secchi Macrophyte Detritus
index wave biomass
base

Ephemeroptera 2.646∗∗ 0.057∗∗ −0.013∗∗ 0.013∗ 0.215

Plecoptera −0.010∗∗ 0.015∗∗ 0.104
Chironomids 7.781∗∗ −0.190∗∗ −0.466∗∗ 0.037∗∗ 0.216∗∗ 0.248
Oligochaetes 8.004∗∗ −0.392∗∗ −0.601∗∗ −0.060∗∗ 0.497∗∗ 0.336
Bivalves 12.596∗∗ −0.220∗∗ 0.154∗∗ 0.125∗ 0.292
Acarina −13.081∗∗ −0.025∗∗ 0.135∗ 0.177
Snails 2.193∗∗ −0.352∗∗ 0.836∗∗ 0.301∗∗ 0.691
Odonata −5.151∗∗ 0.225∗∗ 0.083∗ 0.344
Trichoptera 1.035∗∗ 0.236∗∗ 0.288∗∗ 0.189
Hydroptilidae 0.862∗∗ −0.024∗ −0.042∗∗ 0.128∗ 0.244∗∗ 0.100
Ostracoda −6.925∗∗ 0.185∗∗ 0.132
Axis 1 694.863∗∗ 4.392∗ −10.518∗∗ −7.986∗∗ 0.302
Macrophytes −7.616∗∗ −0.527∗∗ −0.049∗∗ −0.033∗ 0.402

beds supported high densities of snails, especially P. Substrate

antipodarum.
Multiple regression analysis
The results of the multiple regression analysis are
shown in Tables 5 and 6. Table 5 shows the multiple
regression equation coefficients and Table 6 shows the
Beta coefficients which indicate the relative contribu-
tion of each independent variable to the prediction of
the dependent variable.
The three most abundant macroinvertebrate groups
that showed a significant relationship with substrate
size in the multiple regression analysis were Ephemer-
optera, chironomids and oligochaetes. Figure 4 shows
the relationships between the abundance of these
groups and substrate index. Ephemeroptera were most
commonly associated with coarse substrates (higher
substrate index) while the other two groups were
generally found in finer mud/silt and sand.
122
Table 6. Beta coefficients for the multiple regression analysis. NB Beta coefficients are equivalent to B coefficients if all of the data was
standardised to mean of 0 and standard deviation of 1. The magnitude of the values indicates their importance in the regression equation

Taxa Beta coefficient values

Substrate Depth Average Secchi Macrophyte Detritus
index wave biomass
base

Ephemeroptera 0.174 −0.169 0.117 −0.265

Plecoptera −0.186 0.216 −0.139
Chironomids −0.242 0.207 0.144 0.191
Oligochaetes −0.407 −0.218 −0.190 0.358
Bivalves −0.282 0.166 0.111
Acarina −0.137 0.126
Snails −0.291 0.576 0.172
Odonata 0.383 0.118
Trichoptera 0.244 0.247
Hydroptilidae −0.123 −0.159 0.134 0.213
Ostracoda 0.215
Axis 1 0.125 −0.252 −0.159
Macrophytes −0.631 −0.239 0.119

Table 7. Results of the Pearson Correlation between DECOR-

Macrophytes ANA Axis scores and the physical and biological variables. n =
Although several macroinvertebrate groups were asso- 384, ∗ = p < 0.05
ciated with macrophyte biomass, only snails (almost
exclusively the small P. antipodarum) and Trichoptera Variable Axis 1 Axis 2 Axis 3
were found in abundance in these habitats. Figure Depth −0.103∗ −0.048 −0.204∗
5 shows the relationship between macrophyte bio- Substrate index 0.293∗ 0.078 0.579∗
mass and abundance of gastropods and Trichoptera. Lake range −0.001 −0.169∗ −0.046
A strong positive relationship was found for gastro- Secchi disc −0.124∗ −0.207∗ 0.081
pods but the relationship with Trichoptera was much Average wave depth −0.226∗ −0.100∗ 0.143∗
weaker. Characeans −0.360∗ 0.020 −0.208∗
Vascular −0.289∗ 0.018 −0.143∗
Detritus −0.055 0.014 −0.121∗
Detritus
Four major macroinvertebrate groups were associated
with detritus; Trichoptera, chironomids, Oligochaeta
and hydroptilid caddisflies. The regression analysis for
these four groups is shown in Figure 6. There were likely that substrate size was the controlling factor for
statistically significant regression functions found for Ephemeroptera.
these groups, however there was a large amount of
variation around these lines indicating that factors Wave action and lake level range
other than detrital biomass are influencing these faunal Chironomids and oligochaetes showed negative rela-
groups. tionships to average wave base depth indicating that
they were less common at wave-exposed sites. Neither
macroinvertebrate taxa abundances nor macrophyte
Depth
biomass showed a relationship with lake level range.
Both Ephemeroptera and Plecoptera had a negative
relationship with depth in the multiple regression Ordination analysis
analysis. However, Ephemeroptera showed a much
stronger relationship with coarser substrates, which The DECORANA ordination and subsequent correl-
are also negatively correlated with depth. It seems ation analysis showed similar results to the multiple

Figure 4. The relationship between substrate index and the three
major macroinvertebrate groups associated with substrate in the
analysis.
regression analysis (see Fig. 7 and Table 7). Axis 1
was positively correlated with substrate index and neg-
atively correlated with depth and macrophyte biomass.
Axis 2 was negatively correlated with Secchi disc, lake
level range and average wave base depth. Axis 3 (not
shown on Fig. 7) showed similar patterns to Axis 1.
123
Figure 5. The relationship between macrophyte abundance and the
two major macroinvertebrate groups associated with macrophytes in
the analysis.
Discussion
Based on the analysis of macroinvertebrate communit-
ies in a range of lakes, this study has demonstrated that
substrate, macrophyte biomass and detritus were the
most important factors directly controlling macroin-
vertebrate distribution and abundance in the lakes
that we studied. Of lesser importance to macroinver-
tebrates were water depth, exposure to wave action,
secchi depth and recent lake level history. However,
this second group of variables could have a consider-
able indirect effect on macroinvertebrate distribution
by controlling macrophyte distribution and abundance,
and substrate characteristics. Schwarz et al. (2000)
compiled macrophyte distribution data from 63 New
Zealand lakes and found a very strong relationship
between light attenuation and the bottom limit of
macrophytes. The most important control on upper
depth limit of macrophytes is exposure to wave ac-
124
Figure 6. The relationship between detritus and the four major
macroinvertebrate groups associated with substrate in the analysis.
tion (Howard-Williams et al., 1995; Hudon et al.,
2000) although this can be highly variable depending
on the wind conditions, fetch and orientation of the
lakeshore. In addition to controlling macrophyte dis-
tributions, wave action increases substrate coarseness
by washing away fine sediments. As would be expec-
ted coarse substrates were generally found in the wave
wash zone at shallow depths (see Fig. 3).
The results of our analysis showed that Ephemer-
optera showed a preference for the coarse gravel and
cobble substrates in the shallow wave zone. Ephem-
eroptera were found in four of the nine lakes that
we studied (Coleridge, Tekapo, Taupo and Waikare-
moana). While the numbers of individuals was low,
especially in Lake Waikaremoana, these results con-
trast with the findings of Timms (1982) who studied
20 lakes in the South Island and found no Ephemerop-
tera. It must be pointed out that while Timms studied
lakes of a range of sizes, the five largest lakes studied
were only sampled at depths greater than 3 m, so the
higher energy zones more analogous to rivers were not
represented. Our findings were more consistent with
those of Jowett & Richardson (1990) and Jowett et al.
(1991) in lotic systems. They found that Deleatidium
spp. abundance was greatest in habitats with gravel
and cobble size substrate classes, and also in areas
of high water velocity. Quinn & Hickey (1990), in
a study of 88 New Zealand streams, also found that
Ephemeroptera and Plecoptera were associated with
streams with steep gradients (and therefore high water
velocities) and cobble beds.
For Potamopyrgus antipodarum, which is usually
the most common macroinvertebrate in New Zealand
lakes, abundance was closely related to macrophyte
biomass. P. antipodarum are known to graze on epi-
phytic periphyton growing on macrophytes rather than
the macrophytes themselves (James et al., 2000a).
Large stands of macrophytes provide a large surface
area for periphyton growth and, therefore, are cap-
able of supporting large numbers of grazers. Quinn &
Hickey (1990) found that P. antipodarum was found
in high numbers in streams with high periphyton bio-
mass. P. antipodarum is also known to prefer lower
water velocities (Jowett et al., 1991; Holomuzki &
Biggs, 1999) and the macrophyte beds provide a
refuge from both turbulence and possibly predation.
Unlike P. antipodarum, which is common on mac-
rophytes, oligochaetes, chironomids and bivalves were
associated with the fine, detritus rich sediments found
under the macrophyte beds and below the macrophyte
zone. This is in agreement with a study by Quinn &

Figure 7. DECORANA plot of the major macroinvertebrate groups. Correlations of physical factors with the axis scores is shown in Table 7.
Hickey (1990) who found that oligochaetes and chiro-
nomids were dominant in rivers with organic enrich-
ment reflecting their detritivorous feeding behaviour.
These taxa are known to be tolerant of low oxygen
levels (McLachlan, 1970, Van den Berg, 1997) and
fine suspended sediments (Biggs & Malthus, 1982).
Our first hypothesis was that the key direct de-
terminant of macroinvertebrate community composi-
tion is substrate type, which in turn is determined by
variables such as exposure to wave activity. Our res-
ults indicate that substrate was certainly an important
factor for determining community composition and
for the distribution for taxa such as Ephemeroptera,
chironomids and oligochaetes. This most likely re-
flects a greater exposure to wave action, resulting in
a coarsening of the substrate. In many ways, shallow,
wave-swept lake shores are analogous to streams and
as such provide another habitat for aquatic insect taxa
more commonly associated with lotic environments to
colonise. Barton & Hynes (1978) in a study of the
Great Lakes found a shift from lentic to lotic taxa in
exposed sites. These disturbance tolerant taxa though
are often not as abundant in the wave zone of lakes as
they are in streams and this is probably due to the less
than optimal, intermittent disturbance regime (Broder-
125
sen, 1995). The snail P. antipodarum, however, which
is the dominant macroinvertebrate in the littoral zone
of most of the lakes surveyed, was most common in
deeper waters, reflecting its inability to cope with the
higher energy, wave swept lake shores.
The second hypothesis was that macrophyte bio-
mass is the second most important determinant of
macroinvertebrate community composition with dis-
tribution of macrophytes largely determined by sub-
strate type and macrophyte biomass determined by
light attenuation from suspended sediments.
Macrophyte biomass was the most important con-
trolling factor for some of the macroinvertebrate taxa,
most notably P. antipodarum, but also some of the
less abundant taxa such as Odonata and Trichoptera.
Light attenuation certainly controls the bottom limit
of macrophytes in the majority of New Zealand lakes
(Schwarz et al., 2000) although in a few cases this
can be raised by the presence of herbivorous fresh-
water crayfish (Parenephrops planifrons). The role
of substrate size in determining macrophyte biomass
is not clear from this study. While it is true that
macrophytes were associated with fine sediments, the
causative relationship was not firmly established as the
macrophyte stands themselves are ideal environments
126
Figure 8. Stylised lake profile showing the four habitats indicated by the combined analyses.
for the settling of fine suspended material. In addi-
tion, the large populations of snails usually present on
the macrophytes can ‘groom’ the macrophytes (James
et al., 2000a) and produce large quantities of faecal
material which enrich the organic content of the sed-
iments. This material also settles out amongst the
macrophytes, although this is unlikely to significantly
contribute to sediment build up. With the exception
of freshwater crayfish, very few of the macroinverteb-
rates present in New Zealand lakes consume macro-
phytes, rather they graze the epiphytic algae (James
et al., 2000b) so in effect the macrophytes provide
another form of substrate for macroinvertebrates to
live on which is independent of the substrate the
macrophytes are growing in.
We did not originally hypothesise about the role
of the detritus in the control of macroinvertebrate
abundance. Although the role of detritus in structur-
ing stream invertebrate communities is a fundamental
component of stream ecological theory (e.g. Vannote,
1980), in New Zealand lakes primary production, es-
pecially epiphytic algae, provides the major source of
food for herbivorous macroinvertebrates in the littoral
zone (Hawes & Schwarz, 1996; James et al., 2000b). It
was surprising then that several major groups includ-
ing oligochaetes, chironomids and Trichoptera were
strongly correlated to detrital biomass. Much of the
detrital material comes from macrophytes although
a significant proportion can come from periphyton
sloughed off coarse substrates in the wave zone. Oc-
casionally, large amounts of allochthanous material
were also present in the detritus (for example several
of the sites on Lake Te Anau had large quantities of
Beech leaves). Much of this detrital material accumu-
lates within the macrophyte beds and also below the
macrophyte zone in the deeper parts of the lake where
it forms the basis of the benthic food web. The accu-
mulation of detritus provides a substrate for the matrix
of bacteria, algae, and fungi supplying a rich source of
organic material for detritivorous macroinvertebrates.
Our final hypothesis was that lake level history
is only a secondary factor controlling macroinverteb-
rate community composition. We found that the range
of lake level fluctuations were not important in con-
trolling macroinvertebrate abundance. However, the
range of lake levels fluctuations was not particularly
large in the lakes we surveyed (<3 m). The difficulty
with examining the effect of lake level fluctuation
is that to a certain extent these fluctuations are also
surrogates for the different lakes, which makes sep-
arating the differences between lake level fluctuation
and other lake characteristics difficult. It also seems
likely that the abundance of certain macroinvertebrate
groups could be influenced by biogeographic factors.
The analysis of lake ranges is also biased to a certain
extent because the lakes with the smallest ranges are
also the smallest lakes and therefore have the shortest
wave fetch and exposure. Despite the lack of a re-
lationship between macroinvertebrate abundance and
lake level fluctuation in this study, it is still possible
that this factor could be important. For example, Lake
Ruataniwha has a very high turbidity (secchi disk =

0.6 m, Table 1) due to the input of glacial flour. This
low water clarity results in a very narrow band around
the edge of the lake close to the surface where there is
sufficient light to support macrophytes (E. Graynoth,
NIWA, pers. comm.). The effect of even a moderate
shift in lake level in this lake then could potentially
have important consequences. Lake Tekapo also has a
very low secchi disk (2 m) but a much larger lake level
range (2.97 m). This, combined with the larger wave
action associated with a larger fetch, may have com-
bined to result in very low biomass of macrophytes,
the epiphytes which grow on them, and the associated
paucity of macroinvertebrates.
Figures 3–6 showed some of the key relation-
ships found between the macroinvertebrate groups and
the environmental variables identified by our analysis.
These figures could be used to make general pre-
dictions of the type of macroinvertebrate community
structure that could be expected under these condi-
tions. The relationships are consistent with what is
known about these organisms, however there is a large
amount of variability associated with the data and
the results should be treated with caution. For ex-
ample we get a strong log–log relationship between
P.antipodarum and macrophytes biomass, but with a
range of up to two orders of magnitude. Despite the
large number of sites and samples, the data set could
only be analysed for a few of the more common groups
as many taxa were rarely found. The presence of some
groups in the lakes surveyed (such as Plecoptera and
amphipods) was patchy, possibly reflecting the influ-
ence of other factors such as proximity to river mouths,
catchment use and substrate stability.
The high variability in the data reflects the broad
range of sites spread over the nine lakes, however by
working at the Order level we have attempted to look
at the larger scale differences. While there will be
some differences between lakes, especially for some
variables such as the degree of wave impact, factors
which act over smaller scales, such as substrate type
and macrophyte biomass, are more important in these
lake systems. This contrasts with rivers where large-
scale factors such as flood frequency, landuse and
source of flow exert a strong control on the biota
(Quinn & Hickey, 1990; Biggs, 2000). The exception
to this is the wave zone where the controlling factors
are much more analogous to lotic systems, especially
in larger lakes.
The major problem with understanding the pro-
cesses in the lake littoral zone is the inter-relation
between depth, substrate, plant biomass, detritus and
127
the macroinvertebrates. These factors form a complex
web shown schematically in Figure 1. Macrophytes
are limited by wave action at the top of their range and
by light limitation at depth. In addition, macrophytes
require suitable substrates for their root systems. Mac-
rophytes provide habitat for both macroinvertebrates
and epiphytic algae, which in turn forms the basis of
the littoral food web (James et al., 2000b). Detrital ma-
terial from decomposing macrophytes, sloughed off
periphyton, faecal pellets and other allochthanous ma-
terial recycles carbon through the detrital food web.
At various points, feedback loops occur, for example
snails groom the leaves of the macrophytes, reducing
sediment and algal films and improving photosyn-
thetic efficiency (James et al., 2000a).
The findings from this study have shown that the
littoral zone of the lake can be divided into four gen-
eral habitats shown schematically in Figure 8. They
are: (1) The shallow zone where wave action is the
dominant physical force that scours fine sediments and
restricts macrophytes to short turf species. The depth
and intensity of the wave zone depends on wave fetch
(and therefore the size and geometry of the lake), wind
direction and shore profile. In the wave zone, which
has a mainly cobble sized substrate, macroinvertebrate
abundance is low and characterised by Ephemerop-
tera, Plecoptera, Coleoptera and Hydroptilidae. At
more sheltered sites, detritivores such as oligochaetes
and some types of chironomids can be common among
the cobbles if there is an accumulation of detrital ma-
terial. (2) Below the wave wash zone, macrophytes
become more common, tend to be dominated by vas-
cular species and typically reach their highest biomass
from 5 to 9 m depth. The deep macrophyte beds
create stable water conditions where fine sediment
readily settles out. The macrophytes also provide a
substrate for periphyton growth and this can result in
very high grazer biomass – typically dominated by
populations of P. antipodarum, which can occasion-
ally reach densities of more than 150 000 per m2 when
the macrophytes beds are very deep. An example is
the 3 m tall macrophyte beds in Lake Rotoaira. Other
macroinvertebrate groups present include Trichoptera
and Odonata. (3) Underneath the macrophytes fine
suspended sediment and plant material dislodged by
grazers settles along with large quantities of faeces
to produce a thick layer of fine mud often rich in or-
ganic material which can support a large detritivorous
community of oligochaetes, nematodes and chironom-
ids. (4) At greater depths, plant growth is limited by
light, and thus the depth range where many macroin-
128
vertebrates can be found in high numbers. Sediments
in this zone are characterised by fine mud or sand. In
such conditions, bivalves such as the large freshwater
mussel Hyridella menziesi are common, along with
oligochaetes. In some lakes, crayfish (Parenephrops
planifrons) are present and they can reduce the bottom
limit of macrophytes through grazing (Schwarz et al.,
2000).
While this study only covered a limited number
of mostly oligotrophic New Zealand lakes, it assists
lake management by providing a better understand-
ing of the potential impacts of anthropogenic effects
such as increased loadings of fine sediments and large
shifts in lake levels. This study shows that while
macroinvertebrate abundance can be highly variable,
some general predictions of community structure can
be made based on a few key physical and biological
factors.
Acknowledgements
This work was funded by The Foundation for Research
Science and Technology (contract CO1816). Genesis
Power and Mighty River Power provided funding for
the studies on Lakes Rotoaira, Waikaremoana and
Taupo. We thank Carmen Stanger, Greg Kelly, Mark
Gall, Julian Sykes, Neil Blair, Don Forsyth and Kathy
Walter for technical assistance. Dr Murray Hicks pro-
grammed the NARFET model. Dr Alastair Suren and
two anonymous referees reviewed the manuscript and
provided valuable advice.
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