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International Journal of Sediment Research 25 (2010) 203-220

Influence of bed heterogeneity and habitat type on


macroinvertebrate uptake in peri-urban streams
REID, H. E.1, BRIERLEY, G. J.2, and BOOTHROYD, I. K. G.3

Abstract
The role of geomorphic structure, referred to as physical heterogeneity, and its influence upon the
colonization of habitat by macroinvertebrates was analysed in the peri-urban, Twin Streams Catchment,
in West Auckland, New Zealand. Using a cross-scalar approach, 4 riffle-run assemblages were analysed
in each of 2 River Styles (a confined, low sinuosity, gravel bed river and a partly confined, low sinuosity,
bedrock, cobble, and gravel bed river). Each of these 8 locations comprised 2 distinct sampling areas; the
upstream zone had a more heterogeneous river bed with a high diversity of physical features and flow,
whilst the downstream area had a more homogeneous structure. Microhabitat features sampled at each
site included streambed material, bank margins, fine grained organic debris, wood, and boulders. Habitat
mosaics and their associated macroinvertebrate relationships followed a semi-predictable but interrupted
pattern, supporting the view that river systems are a patchy discontinuum. Homogeneous zones were
more frequently characterised by higher proportions of Trichoptera than heterogeneous zones, whilst
heterogeneous zones were frequently characterised by Plecoptera and Ephemeroptera. Diversity was
maximised when the species pools from heterogeneous and homogeneous sites were combined for any
given site. Functional habitats influenced macroinvertebrate assemblages in non-linear and complex
ways. Wood and organic debris habitats were associated with high diversity, abundance, and sensitive
species whereas streambed habitat was usually associated with low diversity. A diverse range of physical
zones that approximates the ‘natural range of behaviour’ for the given type of stream was considered to
provide a more effective platform for rehabilitation planning than emphasising heterogeneity of physical
structure in its own right.

Key Words: Heterogeneity, Functional habitat, Urban stream, Rehabilitation, Geomorphology,


Macroinvertebrates

1 Introduction
Biophysical templates can be used to contextualise and explain patterns in ecological data (Clarke et al.,
2003; Montgomery, 2001). As noted by Harper and Everard (1998): “…the dynamic physical processes
occurring within a river create the physical (habitat) structure of the river: the diversity and dynamics of
this habitat structure is the basis for river biodiversity…”. Although retention of physical integrity does
not guarantee that ecological integrity will be maintained (Hilderbrand et al., 2005), a system with poor
physical structure is almost certain to have a highly disturbed ecosystem (Cullum et al., 2008). Restoring
ecological integrity to systems that retain their physical integrity is far easier and less expensive than
trying to restore ecosystems once abiotic thresholds have been crossed (Fryirs et al., 2008; Hobbs and
Harris, 2001).
Various frameworks have been developed to characterize the spatial organization of river systems
(Cullum et al., 2008). The Process Zone Concept separates rivers into different geomorphic process zones
1
Doctoral student, School of Environment, University of Auckland, 10 Symonds St, Auckland, New Zealand,
E-mail:h.reid@auckland.ac.nz
2
Prof., School of Environment, University of Auckland, 10 Symonds St, Auckland, New Zealand
3
Dr., Research Scientist, Golder Associates LtD, PO Box 33-849, Takapuna 0622, Auckland New Zealand
Note: The original manuscript of this paper was received in March, 2010. The revised version was received in May
2010. Discussion open until Sept. 2011.
International Journal of Sediment Research, Vol. 25, No. 3, 2010, pp. 203–220 - 203 -
with differing disturbance regimes which influence ecosystem structure and dynamics (Montgomery,
1999). Poole (2002) used this idea to develop the “hierarchical patch dynamics”-model, in which
processes operating over multiple scales create, destroy, or alter patch characteristics and their ecosystems.
Patches are distributed as a discontinuum across the catchment, forming a mosaic of river character, with
one patch not necessarily more related to its neighboring patch than other patches in the system. Similarly,
the Riverine Ecosystem Synthesis (Thorp et al., 2006, 2008) combined eco-geomorphology with the
hierarchical patch dynamics model, viewing river systems as complex arrays of hydrogeomorphic patches
largely influenced by climate and vegetation zonation. These patches have unique ‘functional process
zones’ reflecting differences in ecological and geomorphological character, rather than progressive
downstream changes (cf., Vannote et al., 1980).
Heterogeneity, defined as “variability in a process or pattern over space and time” (Palmer and Poff,
1997), provides a basis to interpret rivers as dynamic and complex systems. Physical heterogeneity of a
river is a product of geomorphic processes such as sediment sorting, erosion, deposition, and hydraulic
variability along with vegetation interactions (Allan, 2004). Townsend (1989) developed a conceptual
framework that links the heterogeneity of functional habitat patches to community structure and dynamics.
Ward et al. (2002) postulated that a wide range of habitats provides capacity to support a variety of
different species. Poff and Ward (1990) identified heterogeneity as being a key component of the physical
template that acts to increase the recovery of biota to anthropogenic and natural disturbance. Hence,
finding ways to analyse and reinstigate heterogeneity could aid river rehabilitation (e.g., Tullos et al.,
2009). Numerous studies have indicated that the functional habitat scale is the most appropriate scale at
which to apply such thinking.
Functional habitats have been defined as “…structural components of the substrate and the vegetation,
objectively identified as distinct by the assemblage of invertebrates which they hold” (Kemp et al., 1999).
This micro-scale concept, also referred to as biotypes or habitat patches, combines geomorphic,
hydrologic, and ecological processes, thereby linking biodiversity to geomorphic processes (e.g., Wohl et
al., 1995, Harper and Everard, 1998; Kemp et al., 1999; Newson and Newson, 2000; Buffagni et al.,
2002; Thomson et al., 2004; Storey and Lynas, 2007). Local scale macroinvertebrate communities have
been found to be influenced by many variables including velocity, Froude number, and Reynolds number
(Brooks et al., 2005; Clifford et al., 2006); habitat types such as organic debris and leaf litter (Mackay and
Kalff, 1969; Dobson et al., 1992); riparian vegetation type (Death, 2000); wood (Scealy et al., 2007;
Collier et al., 2000); depth, percent sand, and percent silt (Collier et al., 2000); substrate size (Quinn and
Hickey, 1990; Jowett and Richardson, 1990); periphyton abundance (Jowett and Richardson, 1990); bank
morphology (Armitage et al., 2001); organic composition (Sudduth and Meyer, 2006); and the degree of
bank modification (Erba et al., 2006). In addition, Lamouroux et al. (2004) found that macroinvertebrate
traits including maximum size, body form, mode of attachment to substrate, feeding habitats,
reproduction, lifespan, and strategies of dissemination were significantly correlated to microhabitat
variables that described either flow, substrate, or trophic condition. Hence, while biotic factors such as
competition and predation are key determinates of macroinvertebrate assemblages, the physical integrity
of the river exerts a primary role in its own right.
Various studies have analysed macroinvertebrate assemblages in relation to the physical heterogeneity of
a river. Townsend et al. (1997) noted that generalist species tended to be correlated with unstable sites,
while specialist species favoured more stable sites. This highlights how species traits have adapted to
physical heterogeneity in an effort to survive disturbance events. Beisel et al. (2000) established that
heterogeneous river beds have a higher number of species than homogeneous beds, which are dominated
by only one or two species. Species richness is highest in patchy and heterogeneous environments that
contain a diversity in substrate sizes and hence a greater number of niches. Brooks et al. (2002)
manipulated stream bed variability to represent high and low levels of heterogeneity and measured the
impact upon macroinvertebrate community characteristics. They concluded that conditions at each site
affected the viability of each bed variability treatment group. However, further replication with sites of a
different character is required to derive statistically significant results. Sullivan et al. (2004) found that the
percentage of macroinvertebrate community comprising the sensitive Ephemeroptera, Plecoptera, and
Trichoptera taxa (%EPT) was significantly correlated with more stable habitats that exhibit better
geomorphic condition and better quality habitats. However, some studies have found little correlation
between physical heterogeneity and biotic assemblages. Lepori et al. (2005) found that restored reaches
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with greater heterogeneity had minimally different fish and macroinvertebrate communities compared to
the unrestored disturbed homogeneous reaches at the reach and patch scale. They concluded that “habitat
changes caused by channelization and restoration have no substantial consequence for the structure of fish
and invertebrate assemblages in the study area” (Lepori et al., 2005) and suggested that restoration at a
scale relevant to humans may not be relevant to biological assemblages. Brown (2007) found that areas
with higher habitat heterogeneity had lower temporal community variability, from which it was inferred
that stability and refugia are increased in more heterogeneous environments. Although large storms
diminished this trend, it reappeared soon after disturbance.
These findings indicate that increasing physical integrity in the hope that improved ecological
functioning will follow, also referred to as the “field of dreams” hypothesis, does not always work. As
noted by Hilderbrand et al. (2005): “effective restoration of the physical variables will create the template
for biotic recovery, but physical structure does not always beget biotic structure, and biotic structure does
not necessarily result in similar ecosystem functions across sites”. Tullos et al. (2009) found that taxa
sensitive to disturbance were characteristic of upstream control reaches (‘natural’ streams), whereas
insensitive taxa were characteristic of restored streams (reconfigured, previously channelized reaches). As
restoring physical structure has become a major technique to improve ecosystem health, analysis of
physical and biotic interactions in different settings is required to appraise the likely successes of different
restoration techniques.
This study investigates the response of macroinvertebrate communities to different physical habitat
attributes along river courses. Macroinvertebrate samples are compared from heterogeneous and
homogeneous zones as well as different functional habitats: wood, organic debris, streambed material
(excluding boulders), boulders, and bank margins were compared. The study is performed in a peri-urban
setting, incorporating sites in native forest, pasture, and urban land uses, which enabled interpretation of
the importance of the natural range of geomorphic diversity and functional habitat types upon
macroinvertebrate communities along a continuum of increasingly degradational land use. This differed
from other studies in that it used natural diversity, not created diversity, and looked at the how habitat
diversity interacted with streambed diversity to influence communities. In addition, the influence of land
use intensity was implicitly considered, and sites were characterised using % land use, water quality,
periphyton abundance, geomorphic habitat assessment and the abundance of functional habitats. This
allowed the intensity and effects of land use at a site to be understood, so that the relationship between
macroinvertebrate assemblage and habitat could be seen, even in degraded sites. Rehabilitation is
unnecessary in pristine locations, therefore understanding how physical habitat can influence ecology in
areas with differing intensity of land use, is important for understanding the success rehabilitation of
habitat may have at different points in the catchment. This also investigates the role different types of
habitat may have in rehabilitation in areas of differing intensity of land use.

2 Study area
The Twin Streams Catchment consists of the Oratia, Opanuku, and Waikumete trunk streams, which is
located in Waitakere City, Auckland, New Zealand (Fig. 1). Most catchments in the Auckland Region are
small (less than 100 ha) resulting in relatively short (first or second order) and narrow (< 2 m wide)
streams. The catchment drains approximately 61 km², divided among three subcatchments, which drain
into Henderson Creek and out into Waitemata Harbour. The Opanuku drains a catchment area of 27 km²
and has a channel length of 15.5 km compared to the Oratia’s smaller area of 17 km² and length of 9.7 km.
The Waikumete was not included in this study.
The headwater reaches of the Oratia and Opanuku Streams are confined, low sinuosity, gravel bed rivers
comprised of riffles, runs, pools, steps, and bedrock outcrops. Bed material consists of a mix of gravels,
boulders, cobbles, bedrock, and sand. As gradient decreases downstream, floodplain pockets are evident
along partly confined, low sinuosity, bedrock, gravel and cobble bed rivers. Single channels locally split
around bars. Geomorphic units include pools, runs, riffles, and lateral and mid-channel bars. Bedrock
outcrops are common on both the bed and the banks, locally forcing pool and riffle sequences.
Upstream areas of the Opanuku and Oratia catchments are comprised of fine grained volcanistic bedded
sandstone or siltstone with localised areas of lava and stratified andesitic boulder and cobble-pebble
conglomerate. Middle sections contain bedded, graded sandstones and laminated mudstones with patches
of alluvium. The lower reaches of the streams flow through pumiceous and gravely sandstone (Edbrook,
International Journal of Sediment Research, Vol. 25, No. 3, 2010, pp. 203–220 - 205 -
2001).

Fig. 1 Location of the Twin Streams catchment and surveyed sites in this study

The headwaters areas in the Waitakere Ranges retain native vegetation with low density housing. The
middle region of the catchment is characterised by rolling foothills which are now mainly pasture and
orchards. The lowland alluvial floodplain is urban. The catchment has experienced rapid and sustained
land use change since the time of European settlement in the 1840s (Gregory et al., 2008). Today, there is
a continuum in the intensity of land use from low in the forested headwaters, medium in the light pastoral
area of the rolling foothills, and high in the urbanised area of the lower catchment. Despite the various
land use impacts, streams retain a relatively good geomorphic structure and function across most of the
catchment (Reid et al., 2008).
The climate in Auckland is temperate. Mean temperatures fluctuate between 11oC and 20oC, with
extremes of -2.5oC and 30oC (NIWA, 2006). Annual average rainfall is 1,240 mm, increasing to 2,000
mm in headwater areas. Other than in a few ephemeral sections in the headwaters, streams flow all year.
Mean monthly flow for Oratia and Opanuku streams in the winter is up to 1.3 m3/s and summer monthly
flows are between 0.3 and 0.4 m3/s (ARC, 2006).

2.1 Site characterization


Eight sites were selected across the catchment that reflected the downstream continuum in land use from
native bush to pasture to peri-urban development. Upstream land use, water quality, geomorphic habitat
assessment, and functional habitat score are summarised in Table 1 (Note: heterogeneous sites are
referred to as US sites as they were always upstream and homogeneous as DS sites as they were always
downstream). Native forest was the primary land use for all sites, but was notably lower for Cochrane
Stream and Border Road (where the percentage of pasture was higher) and Millbrook Road (where the
percentage of urban was higher). Scores for geomorphic habitat condition declined down-catchment (see
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Reid et al., 2008). For Pipeline and Candia Road, both the heterogeneous (US in Table 1) and
homogeneous (DS in Table 1) zones were in good condition, at Cochrane Stream both were moderate and
at Border Road both were poor. The heterogeneous site was in better condition than the homogeneous site
at Grassmere Stream, Otimai, Driving Stream and Millbrook Road. All heterogeneous sites had higher
functional habitat total scores (the sum of the ranks for each habitat), with the exception of Otimai (19 at
the heterogeneous and 24 at the homogeneous) and Candia Road, which had the same scores at both sites.

Table 1 Site characterisation. US (short for upstream) refers to the Heterogeneous sites
and DS (short for downstream) to the Homogeneous sites
Grass- Driving Candia Border Millbrook
Site Locations Pipeline Otimai Cochrane
mere stream Rd Rd Rd
1- Land Use (%)
Native forest 98.6 95.5 91 56.5 75 78 65 53
Exotic forest 0 0 1 3 0.5 0.25 2 2
Pasture/Agriculture/
Horticulture 1.3 4.5 4 38.5 23 20 28 19
Urban 0.1 0 4 2 1.5 1.75 5 26
2- Water Quality
Temperature (°C) 16.1- 21.5 15.5 17.0 16.5 16.4 16.4 16.5 17.4 17.7
pH 7.2-7.8 7.4 7.6 7.9 7.4 7.5 7.4 7.3 7.4
Conductivity (ȝs)
129.6 162.2 212.0 181.8 158.5 140.6 145.9 195.2
> 175*
Turbidity (NTU) 5.6 2.3 3.7 1.8 2.0 4.2 3.4 6.5 14.3
Dissolved oxygen
96.8 89.7 115.2 92.8 88.3 90.5 88.6 88.8
(%) 98-105
NOx (ug/L) 444 1366.7 866.7 1066.7 700.0 600.0 733.3 900.0 400.0
FRP (ug/L) 33 189.2 134.2 203.5 331.1 907.5 888.8 112.2 262.9
Dissolved Copper
0.87 0.79 2.38 1.13 1.07 1 1.39 1.51
(ppb) 1.4
Dissolved Zinc
3.17 3.46 8.05 6.4 5.36 4 6.06 10.99
(ppb) 8
Periphyton
3.7 8.8 3.3 25.8 9.9 1.5 102.8 13.3
(AFDW- g/m2) 35*
US DS US DS US DS US DS US DS US DS US DS US DS
3- Geomorphic
Habitat Assessment
Channel geometry 9 9 9 9 O 8 9 8 9 O 9 9 8 8 9 8
Geo-diversity O 8 9 8 9 8 O 8 9 8 9 O O 8 O 8
Riparian vegetation 9 9 8 O 9 O 8 O 9 8 9 9 8 8 8 8
Urban modification 9 9 9 9 9 9 9 9 9 O 9 9 O O O 8
Total 9 9 9 O 9 O O O 9 8 9 9 8 8 O 8
4- Functional
habitat
Boulders 4 3 5 3 2 2 4 2 2 3 1 1
Cobbles 5 4 4 4 2 2 3 2 5 3 4 3 2 1 3 3
Gravel 3 5 3 4 5 5 5 4 4 5 4 4 4 3 4 3
Sand 3 4 4 1 3 3 3 3 4 4 3 4
Silt 2 2 2 3
Marginal plants 2 4 2 1
Floating plants 2
Mosses 2 3 1 2 2 2 2 3 2
Filamentous algae 2 3 2 3 3 3 2
Leaf litter 3 3 3 2 2 2 2 1 3 5 2 2 4 1 2 2
Woody debris 2 1 3 3 1 1 1 2 2 5 4 2 4
Tree roots 2 2 2 2 3 4 3 2 1 5 1 2 2 3 1
Undercut banks 3 3 1 2 1 3 3 2 1 3 3 2
Total 24 22 23 21 19 24 26 20 24 23 22 22 28 24 28 25
1- Percent land use values sourced from Terralink International Limited (2001).
2- Bold refers to ANZECC (2000) guidelines for ecosystem function other then periphyton (Biggs, 2000).
Bold italics refer to the sites that exceed the ANZECC/Biggs guideline.
3- 9indicates good, O moderate and 8 poor condition.
4- 1- rare, 2-few, 3- common, 4- very common, and 5- abundant. Blank spaces indicate that habitat was absent.

International Journal of Sediment Research, Vol. 25, No. 3, 2010, pp. 203–220 - 207 -
Water quality was analysed at all sites over two days at ‘normal’ flows using the ANZECC guidelines
(2000; see Table 1). Some water quality variables (Dissolved Oxygen (DO) and phosphate) were outside
ANZECC (2000) guidelines for all sites. Pipeline and Grassmere Stream were upstream sites with
minimal water quality degradation. Water quality at Otimai was relatively poor, with high conductivity,
pH, DO, and dissolved zinc. Cochrane Stream had slightly high conductivity and low DO. Driving Stream
and Candia Road had below guideline levels for DO and much higher filtered reactive phosphate than
other sites. Water quality was most degraded at the two urban sites, Border Road and Millbrook Road,
where DO was low and turbidity was high. Millbrook Road had high dissolved copper and zinc and
conductivity, while Border Road had very high periphyton growth.

3 Methods

3.1 Site selection


Using a cross-scalar approach, 8 sites were analysed consisting of 4 riffle-run assemblages within each
of 2 river styles (1. Confined, low sinuosity, gravel bed river and 2. Partly confined, low sinuosity,
bedrock, cobble and gravel bed river). Each of these 8 locations comprised 2 distinct sampling areas: the
upstream zone had a more heterogeneous river bed and a high diversity of physical features and flow,
whilst the downstream area had a simpler, more homogeneous structure. Sites were between 8 and 15 m
in length and separated from each other by a distance of between 40 and 100 m. This distance was based
on the locations of riffle-run units that had attributes that were desirable for this study. These riffle-run
zones had similar average flows.
Physical habitats at each site were assessed using two techniques: Geomorphic Habitat Assessment and
Functional Habitat Assessment (Reid et al., 2008; see Table 2). The Geomorphic Habitat Assessment
analysed the condition and functioning of the system, whilst the Functional Habitat Assessment provided
a count of the amount and types (organic versus inorganic) of habitats present at a site. Together, these
procedures provide a picture of geomorphic and ecological conditions in each zone.

Table 2 Brief overview of the variables analysed in each of the habitat assessment techniques. Geomorphic habitat
assessment is derived from Brierley and Fryirs (2005) and Reid et al. (2008). In this assessment, points are
awarded based on qualitative assessment where descriptions are provided for different scores for each
variable. The list of Functional Habitats was sourced from Newson et al. (1998). Each habitat type was
awarded a score out of 5 with the following interpretations: 1- rare, 2-few, 3- common, 4- very common,
and 5- abundant.
Geomorphic habitat assessment Functional habitats
Bed level stability Boulders
Largest clast size Cobbles
Bank erosion Gravel
Channel geometry Sand
Diversity of geomorphic units Silt
Diversity of flow Marginal plants
Wood Emergent plants
Floodplain diversity Floating leaved plants; submerged broadleaved plants
Natives vs. exotic riparian vegetation Submerged, fine leaved plants
Continuous strip of riparian vegetation Mosses
In-stream vegetation Filamentous algae
Landuse- impervious surfaces Leaf litter
Direct urban effects Wood
Rehabilitation efforts Tree roots, trailing vege
Undercut banks

3.2 Functional habitats


Samples of benthic macroinvertebrates were taken from five common functional habitats: streambed
material (excluding boulders), bank margins, boulders, trapped accumulations of organic debris, and
wood. Macroinvertebrates samples from the streambed are the most commonly used for monitoring
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programmes (Carter and Resh, 2001; Stark et al., 2001). The streambed sampling habitat comprised of
cobbles, gravels, and bedrock, and were sampled separately from areas consisting only of sand or fine
grained sediment, as these are known to support different species (Stark et al., 2001; Stark and Maxted,
2007). Boulders were sampled as a separate habitat, as they create a local diversity of flow and sediment
distributions as well as providing refuge areas during high flows (Townsend, 1989). Boulders have been
found to be associated with higher macroinvertebrate diversity and abundance (Quinn and Hickey, 1990).
Bank margins are diverse entities where complex changes in flow, vegetation, and bank morphology
create a mosaic of habitat patches (Armitage et al., 2001). Bank margins retain a higher taxa richness in
chemically and physically disturbed streams, where adjacent riffles showed a decrease in richness and
density (Roy et al., 2003). As organic debris and wood provide habitat and food to the stream, they have
been found to exhibit high densities of macroinvertebrates (Mackay and Klaff, 1969; Scealy et al., 2007).

3.3 Macroinvertebrate survey


Benthic macroinvertebrate samples were taken in duplicates of 3 from each habitat at each zone or until
that habitat type was exhausted (i.e., a maximum of 15 samples from each zone). Each sampling site was
marked on a site map and flow was recorded. The homogeneous (downstream) zone was always sampled
first to ensure that sampling did not impact the heterogeneous (upstream) sampling zone. All samples
were taken by the same person to ensure consistency of procedure. A 30 cm D-net with a mesh size of 0.5
mm was used to collect the samples (except the organic debris). Sample material was disturbed both by
hand (rubbing the surface to dislodge organisms) and through use of a brush to remove organisms from
crevices.
Differing sampling quadrants were used for each habitat. Streambed samples were taken for a sampling
area of 0.125 m2. The same sampling area was used for bank margins but was distributed differently to
enable the sampling of shallow sections of the channel. As organic debris provides habitat on two sides,
the sample area was halved (i.e. 0.0625 m2 quadrant). The organic debris was coarsely washed over the
collecting net, leaving the residual macroinvertebrates. The entire surface of wood and boulders were
sampled and the area of the feature recorded so that it could be standardised for comparability.
Taxa were sorted in the field and identified in the lab. Samples were stored in 70% ethanol to ensure
long-term storage (Stark et al., 2001). A microscope was used to identify invertebrates to the species level,
if possible. For invertebrates where only one species was present the genus level was deemed sufficient.
Macroinvertebrate indicators were calculated as follows. The macroinvertebrate community index (MCI)
attributes a value between 1 and 10 to each genus depending on its sensitivity to water quality and habitat
disturbance. This results in a score between 20 and 200 (Stark et al., 2001). Quality classes are as follows;
scores over 119 indicates that the stream has excellent quality with clean water, 100-119 means water
quality is good with possible mild pollution, 80-99 is fair quality with probable moderate pollution, and
less than 80 indicates poor quality with probable severe pollution (Stark and Maxted, 2007). Taxa
richness was assessed based on the number of species in a sample. Abundance was analysed by dividing
the number of organisms by the area sampled to give an output of the number of organisms per m2.

3.4 Treatment of data


Prisim 6 was utilised to perform non-parametric Mann-Whitney U tests to compare means of the
heterogeneous and homogeneous sites for the summary statistical variables including %EPT, number EPT,
MCI, number of species, and number of organisms.
Multivariate analysis was carried out using Primer-E (Plymouth Routines in Multivariate Ecological
Research) V.6 statistical software. This method makes few assumptions about normality (a vital attribute
for biological count matrices) and is apt at dealing with the high numbers of zeros found in this data type
(Clarke and Gorley, 2001). Data were transformed using the square root and standardised to allow for
samples with different areas to be compared on the same scale. Similarity matrices were created using the
Bray-Curtis coefficient. This output was presented using non-metric Multidimensional Scaling (MDS)
diagrams that display (three-dimensional) data in a non-dimensional (two-dimensional) space. This is a
robust and widely used technique for interpreting ecological population matrices, which allows the
relationships between samples based on their community composition to be viewed (Clarke, 1993). SPSS
was used to correlate macroinvertebrate community descriptors to the axis so that the characteristics of
samples based on their distribution could be understood.
International Journal of Sediment Research, Vol. 25, No. 3, 2010, pp. 203–220 - 209 -
4 Results
MDS diagrams show that samples from heterogeneous and homogenous sites formed clusters at most of
the sites, with minimal overlap (Fig. 2). This indicates that samples taken in each zone were more similar
to each other than those taken in the neighbouring zone. However, these patterns differed from site to site,
indicating non-linear and complex relationships.
Samples from the heterogeneous sites at Pipeline, Grassmere Stream and Driving Stream were
significantly linked to more sensitive species. Samples from the heterogeneous site at Pipeline had a
significantly higher number of EPT taxa per unit area (p = 0.0034) and higher abundance of organisms (p
= 0.0034) than samples from the homogeneous site. Whilst samples from Grassmere Stream’s
heterogeneous site had a significantly higher average % EPT from all sample types (p = < 0.0001) and
higher % EPT (p = <0.0095) and MCI (p = <0.0095) scores for streambed samples than the samples from
the homogeneous site. The heterogeneous site at Driving Stream had higher %EPT per average sample
than the homogeneous site (p = 0.011). All of these sites displayed relatively tight clustering of samples
from the heterogeneous and homogeneous sites on the MDS diagrams. Candia Road’s heterogeneous site
streambed samples had higher species diversity than the homogeneous site streambed samples (p =
0.0435). These sites were all located in the Opanuku subcatchment in areas with relatively low intensity
land use.
Samples taken from the heterogeneous and homogeneous sites at Otimai and Cochrane Stream showed
minimal overlap on the MDS diagram (Fig. 2). Despite this, the number of sensitive species (seen in the
% EPT and MCI scores) was similar for samples from the heterogeneous and homogeneous sites,
displaying a different relationship from that seen in the headwater sites in the Opanuku subcatchment.
Sample clusters at the urbanised Border Road and Millbrook Road sites had little or no overlap. Both
heterogeneous sites were characterised by high abundance of organisms relative to the homogeneous sites
(p=0.0256 at Border Road and p= 0.138 at Millbrook Road). The number of sensitive species was similar
from samples taken at both sites, though the number of species was higher in the samples from the
heterogeneous site (4 more at Border and 8 more at Millbrook).
River Style was found to exhibit a minimal influence upon the relationship between macroinvertebrate
communities and heterogeneity in comparison with land use intensity and sub-catchment differences. The
confined, low sinuosity, gravel bed rivers in the Opanuku sub-catchment (Pipeline and Grassmere Stream)
displayed a larger difference in macroinvertebrate communities between samples from the heterogeneous
and homogeneous sites, especially with regard to more sensitive species compared to sites of the same
river style in the Oratia sub-catchment (Otimai and Cochrane Stream). The partly confined, low sinuosity,
gravel bed rivers were influenced by urbanisation, with samples from the heterogeneous sites located
within the urban area (Border Road and Millbrook Road) being characterised by higher abundances of
organisms than the homogeneous sites, whilst samples from the heterogeneous sites of the same River
Style within pasture were linked to higher sensitive species (Driving Stream) and species diversity
(Candia Road) than samples from the homogeneous sites.
The number of species found at a site was typically higher at the heterogeneous site than the
homogeneous site. Otimai and Candia Road were exceptions with higher diversity at the homogeneous
site than the heterogeneous site. However, when the samples from only the streambed were compared,
Candia Road had a statistically significant increase in species collected the heterogeneous site relative to
the homogeneous site. This indicates that habitat features other than streambed contained most of the
species pool at the homogeneous site. Species numbers were very similar from the heterogeneous and
homogenous sites at Cochrane Stream and Driving Stream. In all instances, diversity was maximised
when the species pools from the heterogeneous and the homogeneous sites were combined (Fig. 3).
Combining species pools added between 2 (at the most urbanised site) and 10 (at a light pastoral site)
additional species.
Relationships between heterogeneous and homogeneous sites and macroinvertebrate community
descriptors are summarised in Table 3. Samples from heterogeneous sites had a positive relationship with
MCI, number of EPTs in a sample, abundance of organisms, and percentage of Plecoptera in all instances.
Weaker positive relationships were evident for percent Ephemeroptera, Mollusca, and Diptera. Diversity
and %EPT were positively related to heterogeneous and homogeneous sites 50% of the time.
Heterogeneous sites were only related to Trichoptera at 33% of the sites, Oligochaete at 20% of the sites
and Crustacea at 25% of the sites. Samples from homogeneous sites were not positively related to any
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Fig. 2 Ordination diagrams of macroinvertebrate samples taken from heterogeneous and homogeneous sites by
non-metric Multidimensional Scaling (MDS). Axis arrows contain the macroinvertebrate community
descriptors that were correlated to the axis and the direction of this correlation. Sites are A- Pipeline, B-
Grassmere Stream, C- Otimai, D- Cochrane Stream, E- Driving Stream, F- Candia Road, G- Border Road and
H- Millbrook Road.

International Journal of Sediment Research, Vol. 25, No. 3, 2010, pp. 203–220 - 211 -
Fig. 3 The number of species for the heterogeneous and homogeneous sites and then combined. The dotted
line indicates the highest species pool that was present when only one site was looked at

Table 3 Summary of relationships between macroinvertebrate samples and the descriptors with which
they were correlated on the MDS diagrams. The hetero and homo columns refer to the
percentage of heterogeneous and homogeneous sites at which the variable was found to be
positively correlated to the heterogeneous and homogeneous samples. Black blocks indicate
positive correlation at heterogeneous sampling sites, grey positive correlation at the
homogeneous sampling site and white indicates no positive correlation.
Hetero Homo Pipe Grass Otim Coch Driv Cand Bord Mill
Diversity 50 50
MCI 100 0
% EPT 50 50
# EPT 100 0
# organisms 100 0
% Ephemeroptera 60 40        
% Plecoptera 100 0        
% Trichoptera 33 66        
% Mollusca 66 33        
% Diptera 66 33        
% Oligochaete 20 80        
% Crustacea 25 75        

variable 100% of the time, showing higher variation in the macroinvertebrate community characteristics
at a site. Weaker positive relationships were seen with more robust species Oligochaete (80%) and
Crustacea (75%) and more sensitive Trichoptera (66%). Weaker positive relationships were seen for %
Ephemeroptera (40%) and Mollusca and Diptera (33%). In summary, samples from the heterogeneous
sites were characterised by higher numbers of EPT, MCI, abundance, and Plecoptera, while
homogeneous sites had more robust species (Oligochaete and Crustacea) but importantly more of the
sensitive Trichoptera.

4.1 Habitat types


The MDS diagrams in Fig. 4 show that despite some overlap, samples from a specific habitat tend to
cluster together. The Pipeline, Otimai, and Driving Stream sites had the least overlap and showed the
clearest clustering of samples from the same habitat. Overlap was high for Millbrook Road, Candia Road
and Grassmere stream, though Grassmere Stream showed quite tight clustering. Streambed and boulder
samples at Cochrane Stream were well spread, with bank margin and wood samples forming tight clusters
within this. Border Road had streambed samples spread across much of the diagram, with wood only on
the left and bank margins on the right. Overall wood, organic debris, and bank margins had tighter
clustering, followed by streambed and sometimes boulder samples.

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Fig. 4 Ordination diagrams of macroinvertebrate samples taken from different habitat types by non-metric
Multidimensional scaling (MDS). Axis arrows contain the macroinvertebrate community descriptors that were
correlated to the axis and the direction of this correlation. Dotted lines indicate separation between
heterogeneous (US) and homogeneous (DS) samples. Sites are A- Pipeline, B- Grassmere Stream, C- Otimai,
D- Cochrane Stream, E- Driving Stream, F- Candia Road, G- Border Road and H- Millbrook Road
International Journal of Sediment Research, Vol. 25, No. 3, 2010, pp. 203–220 - 213 -
Relationships between habitat type and macroinvertebrate communities are summarised in Table 4.
Samples at sites with wood and organic debris were characterised by high diversity, organism abundance,
and number of sensitive species at between 80% and 100% of sites. Macroinvertebrate communities on
wood commonly consisted of Oligochaete, Mollusca, and to a lesser extent Ephemeroptera, and Diptera
and were not characterised by Plecoptera. Communities on organic debris commonly had Diptera,
Crustacea, Mollusca, and again to a lesser extent Ephemeroptera. Plecoptera were more common on
organic debris than wood, with a positive correlation 66% of the time.

Table 4 Summary of functional habitat associations from the correlations shown in the MDS diagrams
Functional Macroinvertebrate associations
habitat
Correlated with high abundance and MCI scores at all sites, high diversity at 80% of sites, and
Wood %EPT at 67% of sites. Taxa commonly found on wood include Oligochaete, Mollusca (100% of
sites), Ephemeroptera (83% of sites), and Diptera (80% of sites). LWD was positively correlated
with Trichoptera at only 50% of sites, Crustacea at 40% of sites, and Plecoptera at no sites.
Positively correlated with diversity, abundance, and MCI scores at all sites. At no sites was it
Organic correlated with %EPT. Macroinvertebrate communities were found to consist of Diptera,
debris Crustacea, and Mollusca at all sites and Ephemeroptera (80% of sites) and Plecoptera (66% of
sites). Both Trichoptera and Oligochaete were positively correlated with organic debris only 25%
of the time.
Positively correlated at 50% or more of sites for just 4 variables: %EPT (100%), Crustacea (100%),
Bank Trichoptera (75%), and Oligochaete (66%). However, negative correlations at 100% of the sites
margins were common, including diversity, abundance, and the percent of Plecoptera. Positive correlations
at only some sites were seen for Mollusca (33%), Ephemeroptera (33%), Diptera (20%), and 50%
of the sites were correlated with high MCI scores.
Streambed Positively correlated with abundance (88% of sites), percent of Oligochaete (70%), Trichoptera
(excluding (60%) and EPT taxa (57%), diversity (30%), MCI scores (30%), and proportions of Ephemeroptera
boulders) (42% sites), Plecoptera (33%), Diptera (25%), Crustacea (13%) and Mollusca (33%).
Positively correlated to the percent of Crustacea (at 100% of sites), percent Trichoptera (77%); and
diversity, abundance, and Mollusca (all 66%). 50% of the sites were positively correlated with the
Boulders percent of Oligochaete. Boulder samples were only positively correlated with EPT taxa at 30% of
sites, Ephemeroptera at 42% of sites, Diptera at 36% of sites, MCI scores at 25% of sites, and
Plecoptera at no sites.

Bank margins had low diversity, abundance, and Plecoptera, but high proportions of Trichoptera,
Oligochaete, and Crustacea. Samples taken from the streambed had a positive relationship with
abundance and a weaker relationship with Oligochaete, Trichoptera, and EPT taxa. Macroinvertebrate
communities taken from the streambed were strongly influenced by the relative diversity of geomorphic
structure and flow in the heterogeneous and homogenous zones. Streambed samples had a high number of
organisms, but a low diversity of species and a limited proportion of sensitive species. Communities in
this habitat type appeared to be more easily altered by differences in velocity and habitat heterogeneity in
a way that was not apparent for more organic habitats. Boulder samples were positively associated with
Crustacea, Trichoptera, diversity, abundance, and Molluscs. Boulders supported communities that were
productive and had a high diversity of species. However, few of the sensitive families were associated
with this habitat, other than Trichoptera.
Extra interest was paid to the types of communities associated with streambed samples, as these are
most commonly used for monitoring (Carter and Resh, 2001). In this study streambed samples were
found to have a high abundance of organisms but usually a low diversity. Figure 5 shows the portion of
the species pool that was found on streambed samples, related to the total number of species found across
all habitat types. As noted on MDS diagrams, streambed samples only covered a portion of the area
covered by samples from other habitats. Wood and organic debris were the habitat type that is most likely
to be outside of the area covered by streambed samples.

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Fig. 5 The number of species at a site found in the streambed samples compared with the total found
from sampling all habitat types. US refers to the heterogeneous site and DS to the homogeneous site

5 Discussion

5.1 Heterogeneity
Although heterogeneity influences macroinvertebrate communities, relationships vary in a non-linear
and complex way at different locations in Twin Streams catchment. Samples from the homogeneous
zones were more frequently characterised by higher proportions of Trichoptera than the heterogeneous
zones, whilst heterogeneous zones were frequently characterised by Plecoptera and Ephemeroptera. This
suggests that homogeneous zones support different types of species that have less complex habitat
requirements. Ecologically this makes sense. Heterogeneous environments may be colonised by habitat
specialist species whilst homogenous environments may be preferentially selected by general species that
can out-compete the specialist species (Townsend et al., 1997). Heterogeneous and homogeneous sites
may also have different macroinvertebrate disturbance histories, as heterogeneity may increase prospects
for macroinvertebrate recovery (Poff and Ward, 1990).
Findings from this study convey a more complex relationship between physical heterogeneity and
macroinvertebrate communities to that noted by Beisel et al. (2000), Sullivan et al. (2004), and Brown
(2007). In this instance, homogenous zones were found to contribute significantly to the overall diversity
of macroinvertebrate assemblages at any given site. As noted by Muhar et al. (1995), in their comments
on restoration programs that increased riffle habitat, little is to be gained in creating uniformly
heterogeneous reaches. A diverse range of physical zones that approximates the ‘natural range of
behaviour’ for the given type of stream is the key platform for rehabilitation planning. Just as ‘natural’
rivers have a mix of heterogeneous and homogeneous zones, so too should rehabilitation programmes that
target biotic diversity.

5.2 Habitat types


Findings from this study also indicate how habitat type influences macroinvertebrate communities.
Although overlap in species ranges is evident, samples from a specific habitat type tended to cluster
together on MDS plots. The strength of the clustering and the characteristics of the community vary from
site to site, indicating complex, non-linear relationships.
Wood and organic debris were associated with high diversity, abundance, and sensitive species. These
habitats provide habitat and food and are a major energy source for freshwater ecosystems (Webster and
Benfield, 1986; Collier et al., 2000; Parkyn and Winterbourn, 1997; Mackay and Kalff, 1969; Hax and
International Journal of Sediment Research, Vol. 25, No. 3, 2010, pp. 203–220 - 215 -
Golladay, 1993). The character of the wood also is important, as more complex wood supports more taxa
and individuals than simpler pieces (Scealy et al., 2007). Mutual interactions among geomorphic
attributes and wood/organic matter are an integral component of a well-functioning river system
(Corenblit et al., 2007; Gorecki et al., 2006). The use of wood as a rehabilitation technique has been
shown to increase geomorphic heterogeneity and increase litter retention, whilst causing minimal damage
to the system (Brooks et al., 2004; Muotka et al., 2002). As such, wood and organic debris must be
viewed on top of geomorphic structure in determining measures of the physical heterogeneity of a river
that provide a basis to develop river rehabilitation initiatives.
The streambed is the most common habitat type sampled for monitoring and assessment of the success
of restoration projects (Stark et al., 2001; Carter and Resh, 2001). Results from this study showed that
sampling the streambed in isolation would yield only a portion of the species pool, ignoring some of the
more sensitive species that reside on wood and leaf matter. However, the community composition of
streambed samples more closely reflected the characteristics of heterogeneous and homogeneous zones
than other habitats and is, therefore more likely to provide an opportunity to detect biotic changes.
The diversity and pattern of functional habitats are an essential component of heterogeneity. Key
habitats may be missing in some areas, while other areas may have a pattern of habitats that is unable to
support elements of ecosystem functionality. For example, the homogeneous zone at the Otimai site had a
higher diversity, number of sensitive species, and productivity. This zone also had a higher number and
diversity of functional habitats than the heterogeneous zone, because of marginal plants, tree roots, and
undercut banks. This indicates that the diversity of functional habitats is a key component in assessments
of the ecological integrity of a site. In some instances, zones with simpler geomorphic structure may have
greater biotic diversity due to the greater range of habitat types. The opposite may also be true. Candia
Road had very low macroinvertebrate abundance in both the heterogeneous and homogeneous zones
despite only minimally degraded water quality. However, periphyton was found to be scarce at the site,
limiting the number of macroinvertebrates that could be supported (as seen in Jowett and Richardson,
1990). In such situations, the diversity of geomorphic structure may not be a major determinant of
community structure, indicating that a high level of physical integrity does not guarantee ecological
integrity (Hilderbrand et al., 2005). Muotka et al. (2002) found that the recovery of moss post-restoration
was an important determinant on the recovery of macroinvertebrate communities, as it provided essential
primary production. In this study, heterogeneous zones with no wood may be less biodiverse than
homogenous zones with high wood loadings. These complex sets of associations indicate that biological
attributes such as riparian vegetation, primary production, and functional habitats must build upon a
geomorphic template to interpret patterns of macroinvertebrate assemblages. Although physical structure
provides a critical starting point for rehabilitation planning, many other considerations must be addressed
to facilitate river repair.

5.3 Patterns of physical structure and ecological communities


Functional habitats do not provide habitat for biota in isolation. Patchiness results from multiple forcing
variables acting over multiple scales. Findings from this study confirm the operation of river
environments as patchy systems (Montgomery, 1999; Poole, 2002; Thorp et al., 2006). The assumption
that neighbouring sections are alike and exhibit similar relationships is not always valid.
In this study, patchy relationships were noted within the catchment, within sub-catchments, for reaches
of the same geomorphic type of river, from heterogeneous to homogenous zones at each site, and at the
functional habitat scale. For example, Driving Stream and Candia Road are both reaches of the same river
style, have similar land use and are located close to each other in the middle of the same catchment.
However, differences among macroinvertebrate communities between heterogeneous and homogenous
zones were much more pronounced at Driving Stream. Notable differences are also evident for the same
river style elsewhere. Confined reaches in Opanuku subcatchment (Pipeline and Grassmere Stream) had
much greater species diversity in the heterogeneous zone relative to the homogenous zone, but trends for
the same river style in the Oratia subcatchment (Otimai and Cochrane Stream) were much weaker (indeed,
reverse for some variables). Although reaches may have a similar physical structure, their biotic
functionality may be quite different.
Heterogeneity is conceptualised differently at different scales. Catchment-wide heterogeneity reflects the
distribution of different types of rivers, whereas micro-scale heterogeneity looks at individual niches,
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local velocity, and types of habitats. Whilst these two scales should be considered, heterogeneity at the
reach scale is likely to be particularly relevant in rehabilitation planning (Wohl et al., 2005). This scale
integrates insight into channel and floodplain landforms, fashioning the range of micro-habitats (e.g.,
flow depth, velocity, and functional habitat). Framing restoration in terms of geomorphic building blocks
at the reach scale provides a platform to integrate natural variability in heterogeneity for a given type of
river (Petersen et al., 1992; Brierley and Fryirs, 2005; Fryirs and Brierley, 2009).
The complex array of factors that influence the viability of aquatic habitat makes comparing treatments
a difficult task. Field sites are far removed from laboratory situations with controlled conditions, making
scientific ‘proof of concept’ extremely challenging when viewed simplistically in terms of statistical
relationships. Indeed, the quest for such notional rigour can impede our capacity to implement
rehabilitation plans (e.g., Lemons and Victor, 2008).
Rehabilitation of physical processes is an integral component in biodiversity management (Jansson et al.,
2005). Different river systems are likely to have different relationships between habitats and geomorphic
diversity. Hence, it is critical to establish these fundamental relationships in a region, viewing them in
light of system complexity. Enhanced understanding of the interactions between environmental factors
and ecosystems is essential if the success of restoration projects is to improve (Bernhardt et al., 2005;
Darby and Sear, 2008). Understanding the physical template is a necessary first step in efforts to
rehabilitate ecosystem health. Physical heterogeneity across different scales, tied to appraisals of
functional habitat types at different points in a catchment and variable land uses, are key considerations in
this process.

6 Conclusion
Findings from this study indicate that the heterogeneity of the physical template of a river system
influences macroinvertebrate communities in non-linear, unpredictable ways. As differing functional
habitats, especially organic units, are able to support a greater range of species than single habitat systems,
the more heterogeneous the units, the greater the prospective range of macroinvertebrates. However,
heterogeneous and homogeneous reaches support different types of macroinvertebrates, making them
both essential components of riverine systems. In addition, the ecological functioning and habitat quality
of a site must be assessed in relation to the surrounding land use. The influence of physical structure on
macroinvertebrate distributions has been shown to be very complex and patchy. Multiple processes
operating at multiple scales result in a mosaic of patches that may display different relationships to a
seemingly similar neighbouring patch. Heterogeneity can provide a useful concept in rehabilitation
planning so long as procedures value uniqueness and difference within a river system, rather than
generalising and simplifying river character and behaviour (Simon et al., 2008).

Acknowledgements
We would like to thank Waitakere City Council, in particular Anil Karan and Graham Leonard for their
support and funding of the project. Funding was also provided by the School of Environment at the
University of Auckland. Technical staff including David Jenkinson, Peter Crossley, Rebecca Bibby,
Sandra Anderson, and Nevil Hudson are thanked for field support. Thanks are given to Liza Inglis, Nick
Carter, Robin Gardner-Gee, and Kohmei Kadowaki for helping with statistics, macroinvertebrate
identification, and providing the software necessary for this work. Thanks are given to Robyn Reid for
proof reading. This work would not have been possible without the many people who helped in the field
including Nadine Trahan, Ethel-Mae Seaman, Claire Gregory, Richard Mairs, Hiroki Ogawa, James
Waugh, Petra Vanlimburg, Robyn Reid, Suzy Reid, Michaela Cowie, Rebecca Bibby, Zac Brierley, and
Charis Wong, and last but by no means least special thanks are given to Iain McGillvary.

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