867 Stark & PhillipsSeasonal variability in biotic indices New Zealand Journal of Marine and Freshwater Research, 2009,

Vol. 43: 867882

00288330/09/43040867 The Royal Society of New Zealand 2009
M08058; Online publication date 15 July 2009
Received 16 October 2008; accepted 15 April 2009
Seasonal variability in the Macroinvertebrate Community Index:
are seasonal correction factors required?
JOHN D. STARK
*
Cawthron Institute
Private Bag 2
Nelson, New Zealand
NGAIRE PHILLIPS
National Institute of Water and Atmospheric
Research Limited
P.O. Box 11-115
Hamilton, New Zealand
*
Present address: Stark Environmental Limited,
P.O. Box 1831, Nelson 7040, New Zealand. email:
jdstark@paradise.net.nz
Abstract Freshwater macroinvertebrate data
collected from streams in Taranaki, New Zealand
(19812006) were used to examine seasonality of
biological indices including taxon richness, %EPT
(Ephemeroptera, Plecoptera, and Trichoptera)
richness, and the hard- and soft-bottomed (-sb)
stream versions of the Macroinvertebrate Community
Index (MCI). All indices tested showed modest but
statistically signifcant seasonal variation in hard-
bottomed streams. Seasonal means for the MCI
were within 3.0% of the annual mean. The Semi-
Quantitative MCI (SQMCI) (4.3%), %EPT richness
(7.4%) and taxon richness (7.7%) showed greater
seasonal variability than the MCI. MCI, SQMCI, and
%EPT richness were signifcantly higher in spring and
winter compared with autumn and summer. Taxon
richness was lower in winter than in other seasons.
In soft-bottomed streams, SQMCI-sb showed least
seasonality (within 3.6% of the annual mean, non-
signifcant), followed by MCI-sb and taxon richness
(4.7%), and %EPT richness (11.2%). Spring and
winter MCI-sb and SQMCI-sb values tended to be
signifcantly higher than those calculated from summer
or autumn samples, although most values were within
5% of the annual site means. Examination of seasonal
variation in species traits and their relationships to
observed seasonal patterns of biotic indices did not
support the suggestion that seasonal variations in
life histories of aquatic macroinvertebrates affect
index values. Rather, seasons with higher biotic index
values were associated with a greater frequency
of flow disturbance, which is consistent with the
view that the character of New Zealands stream
macroinvertebrate communities with their poorly-
synchronised life histories, has been shaped by the
unpredictable physical environment. We recommend
the MCI and MCI-sb for State of Environment
reporting in New Zealand for cost-effectiveness and
because data requirements and seasonal variation
were less than for quantitative alternative indices.
We suggest that seasonal variability is unlikely to
confound interpretation, and does not need to be
considered.
Keywords macroinvertebrates; monitoring;
seasonality; biotic indices; New Zealand
INTRODUCTION
Stream health assessments using macroinvertebrates
are based on the assumed relationships between
stream health and the pollution tolerances or
habitat preferences of invertebrates (Ministry for
the Environment 1999; Stark & Maxted 2007a,b),
but the rapid adoption of some biotic indices has
occurred without appreciation of factors that may
confound interpretation. For example, New Zealands
Macroinvertebrate Community Index (MCI) (Stark
1985), was used for several years before Stark (1993)
recommended that sampling should be restricted to
riffe or run habitat within defned ranges of depth
(0.10.4 m), velocity (0.21.2 m/s), and substrate
(median rock dimension 60140 mm) to improve
assessment reliability. Furthermore, MCI values
(or the semi-quantitative (SQMCI) or quantitative
(QMCI) variants) obtained from single samples
often were compared without any consideration of
868 New Zealand Journal of Marine and Freshwater Research, 2009, Vol. 43
estimation error until Stark (1993, 1998) determined
how much biotic index values from single samples
need to differ, for the difference to be statistically
significant (e.g., MCI values at least 11 units
apart were likely to be significantly different).
The detectable differences determined by Stark
(1993) provided a basis for determining whether
differences in MCI were likely to be significant
based on cost-effective monitoring programmes
involving collection of single hand-net samples
per site.
Whether based on biotic indices or predictive
models (Wright 1995; Davies 1997; Reynoldson
et al. 1997; Coysh & Norris 1999; Joy & Death
2000, 2002, 2003), reliable assessments require that
potential confounding influences are minimised,
eliminated, or are sufficiently small not to be
considered. Often spatial variability is of interest (e.g.,
decreasing MCI values at sites progressively further
downstream, or below point-source discharges)
(Maxted et al. 2005; Niyogi et al. 2007), but also may
be a confounding infuence (e.g., variations in MCI
between riffes, runs, or pools within a single stream
reach, or differences between soft-bottomed or hard-
bottomed habitats). Similarly, changes in stream
health over time may be of interest (e.g., using the
MCI to monitor the improvement in stream health
resulting from elimination of a point source nutrient
discharge), but temporal variability potentially could
confound interpretation of long-term monitoring
programmes such as State of the Environment (SoE)
monitoring if sampling has been undertaken over
several weeks or in different seasons.
It should be possible to undertake biological
monitoring at any time of year and obtain a
stream health assessment that is comparable to
other sampling times (Murphy 1978). However,
reporting on the state of New Zealands freshwater
environment using samples collected from different
regions of New Zealand at different times of year
could be compromised if there is strong seasonality
in macroinvertebrate community composition and
the biotic indices used for reporting the results. We
would need to determine, for example, whether MCI
values from one region collected in spring were
higher than those from another region collected in
summer owing to a real difference in stream health
between regions or the effect of season.
Several studies have found that season may
influence biomonitoring assessments based on
predictive models (Reynoldson et al. 1997; Reece
et al. 2001; Feio et al. 2006), and have concluded
that models should be run using test data collected
in the same season/s as the available reference
data (and that more data collected from different
seasons should be added to the reference database
to encompass suffcient natural variation to allow
the use of the models year round).
River fow, water temperature, dissolved oxygen
levels, and invertebrate life histories, have all
been implicated as causes of seasonal variability
in biotic indices, with biotic indices indicative of
poorer stream health recorded in warmer seasons
(Murphy 1978; Armitage et al. 1983; Hilsenhoff
1988; Bickerton et al. 1993; Lenat 1993; Zamora-
Munoz et al. 1995; Linke et al. 1999; Pires et
al. 2000; Pastuchova 2006; Sporka et al. 2006).
Seasonal variability in biotic indices is undesirable,
unless it refects water quality changes, as it makes
comparisons between different years or different
seasons in the same year diffcult (Murphy 1978).
Nevertheless, correction for the confounding effects
of season has been investigated for relatively few
biotic indices. For example, Hilsenhoff (1988) and
Lenat (1993) derived additive seasonal correction
factors for adjusting Family Biotic Index (FBI)
and the closely related North Carolina Biotic Index
(NCBI), respectively.
Examination of species traits is one means to
determine whether invertebrate life histories infuence
biotic indices. Key life history, physiological and
behavioural characteristics of stream invertebrates
(or species traits) can be used to describe the
functional structure of a stream community, based
on taxa abundances and an array of biological traits
describing size, reproductive characteristics, mobility,
body form, and feeding habits (Charvet et al. 1998,
2000; Gayraud et al. 2003). Analysis of community
responses to disturbance based on species traits
offers an alternative or complementary approach
to community structure for assessing disturbance
responses in stream communities (Corkum &
Ciborowski 1988; Poff & Allan 1995; Poff 1997;
Lamouroux et al. 2002; Doldec et al. 2006). This
approach provides insight into the mechanisms
causing observed changes in functional and structural
characteristics of communities, as biological traits
refect the functional relationships between biota and
various environmental characteristics (Poff 1997).
In this study, we examined seasonal variability
in several biotic indices that are commonly used
in New Zealand to determine the implications for
biological assessments, and to determine whether
seasonal correction factors are required to facilitate
valid comparisons of monitoring data collected
in different seasons. We examined also the roles
869 Stark & PhillipsSeasonal variability in biotic indices
of species traits and flow variability as possible
explanations for biotic index seasonal variation.
METHODS
Taranaki macroinvertebrate data
Taranaki Regional Council (TRC) maintains an
extensive database of macroinvertebrate samples
collected from rivers and streams in the Taranaki
region, North Island, New Zealand (Taranaki Regional
Council 2004, 2006, 2007). Sites are assigned to one
of five different stream types based primarily on
the source of fow (viz., RPOP = Taranaki ringplain
streams with source of fow on upper ringplain but
outside the Egmont National Park boundary; RPNP =
Taranaki ringplain streams with source of fow on Mt
Taranaki above the Egmont National Park boundary;
LOWL = lowland non-ringplain streams with source
of fow below 350 m above mean sea level; LOWC
= coastal ringplain streams with source of fow below
350 m above mean sea level; HILL = non-ringplain
streams with source of fow in hill-country east of Mt
Taranaki) (Table 1). Data analysed were from samples
collected between 23 June 1981 and 15 September
2006 by kick-sampling using a 0.5 mm mesh D-
net, in a manner consistent with Protocol C1 (Stark
et al. 2001), since this protocol was based on the
sampling method used in Taranaki. Before February
1986, only presence-absence data were recorded, and
from then until October 1998, a three-level coded
abundance system (rare, common, abundant) was
used. Thereafter, sample processing was consistent
with standard Protocol P1 (Stark et al. 2001), with
fve levels of coded abundances (viz., rare (R) = 1 to
4 animals/sample; common (C) = 5 to 19; abundant
(A) = 20 to 99; very abundant (VA) = 100 to 499; and
very, very abundant (VVA) = 500+).
Samples were assigned to seasons based on
sampling dates (viz., spring SeptemberNovember,
summer DecemberFebruary, autumn MarchMay,
winter JuneAugust) (Table 1).
Data used were based on 2617 samples collected
at 390 different locations in hard-bottomed streams
in Taranaki. Sampling effort was greatest in spring
(32%) and summer (31%), and least in winter (13%)
(Table 1). In addition, data were analysed from 400
samples collected from 65 different locations in soft-
bottomed streams. Sampling effort was greatest in
summer (45%), and least in winter (19%) (Table 1).
MCI, SQMCI, MCI-sb, and SQMCI-sb values were
calculated by Microsoft Excel user-defned functions
using the taxon tolerance values and equations
described by Stark (1985, 1993, 1998), and Stark
& Maxted (2007a,b). Other indices calculated were
taxon richness (number of macroinvertebrate taxa
per sample) and %EPT richness (the percentage of
taxon richness comprising Ephemeroptera (mayfies),
Plecoptera (stonefies), and Trichoptera (caddisfies)).
All indices were standardised as percentages
of overall site means to enable data from sites of
differing stream health to be analysed together and
seasonal variation in index values to be expressed
as percentages. Data were unsuitable for repeated
Table 1 Numbers of sampling sites and samples included in Taranaki Regional Council (New Zealand)
macroinvertebrate data (19812006). Seasons: spring (SeptemberNovember), summer (DecemberFebruary), autumn
(MarchMay), winter (JuneAugust). Source of fow: RPOP = ringplain outside Egmont National Park; RPNP =
ringplain within Egmont National Park; LOWL = lowland non-ringplain; LOWC = coastal ringplain; HILL = non-
ringplain hill country.
Source No. of No. of samples
Stream type of fow sites Spring Summer Autumn Winter Total
Hard-bottomed RPOP 88 134 151 125 45 455
RPNP 198 553 537 363 185 1638
LOWL 54 61 54 56 48 219
LOWC 10 22 21 17 12 72
HILL 40 80 56 44 53 233
Sub-total 390 850 819 605 343 2617

Soft-bottomed RPOP 12 12 19 14 17 62
LOWL 40 41 81 45 29 196
LOWC 13 45 47 20 30 142
Sub-total 65 98 147 79 76 400
Total 455 948 998 684 419 3017
870 New Zealand Journal of Marine and Freshwater Research, 2009, Vol. 43
measures ANOVA (because the number of replicates
per sampling site was highly variable), so we used
one-way ANOVA followed by post hoc Tukey HSD
tests to determine whether seasonal variation was
statistically significant (P < 0.05) (Winer et al.
1991; Lenat 1993; StatSoft 2007). We assumed that
samples were independent, which we considered a
reasonable assumption given that the most frequent
sampling at any sites was seasonal, with 6-monthly,
or annual sampling at other sites.
Flow data
Flow data (Taranaki Regional Council unpubl. data)
from 14 sites that had fow recorded continuously
for 20 to 36 years were used to determine median
fows for each site. For each river, we calculated
the number of days (overall) and for each season
the number of days on which mean daily flows
exceeded three times median fow as a measure of
the disturbance regime. Apart from the magnitude of
the fow, high fow frequency variables were found
by Clausen & Biggs (1997, 2000) to be the single
most direct constraint on biota in temperate perennial
rivers. Consequently, we used the frequency of
foods exceeding three times median fow (FRE
3
) as
a measure of the effect of disturbance on periphyton
and macroinvertebrate communities.
Species traits analyses
We examined species trait composition in relation
to season as a possible explanation for the observed
seasonal variation in MCI and SQMCI values. For
each macroinvertebrate species, we documented 55
categories (modalities) of 15 biological traits in a
similar manner to Usseglio-Polatera et al. (2000) and
Gayraud et al. (2003). The selected traits refected
the life history of organisms, the resilience or
resistance potential beyond that of life history traits,
and general biological and physiological features
(see Doldec et al. 2006).
As we used multiple sources of information, we
used a fuzzy coding approach to quantify the affnity
of each taxon for each category within a trait. We
gave each taxon an affnity score ranging between 0
and 3 that refected the relative strength of association
between a taxon and a given trait category. For
example, a species that is predominantly a predator,
but feeds as early instars by scraping, was given an
affnity of 3 for the feeding category, predator, and
a 1 for the category, scraper. Using this approach,
we were able to incorporate the variability of traits
that often occurs at different life stages. We scored
traits as 0 for any category for which information
was not available. For most traits, the percentage of
missing information was 213 %.
Affnity scores were standardised so that their sum
for a given species and a given trait was 1. We then
calculated the relative abundance of traits at each
site by multiplying the proportion of each category
per trait by the relative abundances of species at the
site. The resulting trait-by-site array contained the
relative abundance of each trait category at each
site (for further details see Chevenet et al. 1994).
These data were compared with SQMCI values.
In addition, we calculated trait frequencies based
on presence/absence data and compared them with
MCI values. We examined seasonal patterns in trait
modality frequencies using Batch (ByGroup) Main
Effects ANOVA (STATISTICA v.8.0, StatSoft Inc.
2007) followed by the Benjamini & Hochberg (1995)
False Discovery Rate (FDR) procedure to reject
signifcant correlations that could have occurred by
chance when undertaking multiple comparisons.
Signifcant results (P < 0.05) remaining after FDR
analysis were compared with observed patterns in
biotic indices.
Trait frequencies were computed with the ade4
library implemented in the R freeware package
(Ihaka & Gentleman 1996).
RESULTS
Hard-bottomed streams
Taxon richness, %EPT richness, MCI, and SQMCI
values covered the entire expected ranges for these
indices (Fig. 1), and all showed statistically signifcant
seasonal variation in Taranaki hard-bottomed streams
(Table 2). In winter, taxon richness was c. 7.7% lower
than the annual average and signifcantly lower than
in the remaining seasons (which were within 13%
of site annual average values). %EPT richness was
signifcantly higher in spring and winter compared
with summer and autumn, with the former 3.87.4%
higher than average and the latter 4.37.1% lower
than average. MCI values were signifcantly higher
in spring (3% above average), than winter (0.6%
above average), with values in these seasons higher
than those in autumn (1.7% below average) and
summer (2.1% below average). SQMCI values were
signifcantly higher in winter and spring (3.34.3%
above average) than in autumn or summer (1.83.7%
below average). Overall, for hard-bottomed streams,
MCI showed least seasonal variation (within 3.0%),
followed by SQMCI (within 4.3%), %EPT richness
(within 7.4%), and taxon richness (within 7.7%).
871 Stark & PhillipsSeasonal variability in biotic indices
Fig. 1 Summary box plots
of t axon ri chness, %EPT
(Ephemeroptera, Plecoptera,
Trichoptera) richness, Macroin-
vertebrate Community Index
MCI/MCI-sb, and Semi-Quanti-
vative SQMCI/SQMCI-sb values
from 390 hard-bottomed stream
sites (2617 samples), and 65 soft-
bottomed (-sb) stream sites (400
samples) in the Taranaki region,
North Island, New Zealand (23
June 1981 15 September 2006).
Source of fow: RPOP = ringplain
outside Egmont National Park;
RPNP = ringplain within Egmont
National park; LOWL = lowland
non-ringplain; LOWC = coastal
ringplain; HILL = non-ringplain
hill country.
Similar results were obtained for the MCI when
data from four stream quality classes (i.e., poor (MCI
<80), fair (8099), good (100119), and excellent
(120+)) were analysed separately. Spring and/or
winter were the seasons with highest MCI values
with autumn and summer the lowest. The ranges
between seasonal means were 5.1% overall, 4.7%
for poor, 3.9% for fair, 5.3% for good, and 4.6% for
the excellent quality class.
Soft-bottomed streams
The ranges of taxon richness, %EPT richness, MCI-
sb, and SQMCI-sb values were more restricted
compared with hard-bottomed streams (Fig. 1).
Taxon richness, % EPT richness, and MCI-sb, all
showed statistically signifcant seasonal variation in
Taranaki soft-bottomed streams but SQMCI-sb did
not (Table 3). In summer, taxon richness was c. 4.7%
higher than the annual average and signifcantly
higher (by 6.28.2%) than in the remaining seasons.
%EPT richness showed greatest seasonal variation
with highest values in winter (111.2%) and spring
(107.8%) compared with summer (93.2%) or autumn
(92.1%). MCI-sb values were highest in winter
(104.7%) and spring (100.8%) followed by summer
(99.5%) and autumn (95.5%), with all seasonal
differences statistically signifcant. SQMCI-sb values
ranged from 2.5% higher than average (spring) to
3.6% lower than average (autumn) although seasonal
variation was not statistically signifcant. Overall, for
soft-bottomed streams, SQMCI-sb showed the least
seasonal variation (within 3.6% of average, not
Hard-bottomed streams
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873 Stark & PhillipsSeasonal variability in biotic indices
Table 4 Signifcance (P) values derived from bygroup analysis of seasonal differences in trait modality frequency
for macroinvertebrate communities from hard-bottomed (HB) and soft-bottomed (SB) Taranaki (New Zealand)
rivers and streams based on species presence/absence and coded-abundance data. Signifcant results (P<0.05), after
application of the Benjamini & Hochberg (1995) False Discovery Rate, in bold. (a, same seasonal rank order as biotic
index; b, reverse order).
Trait Presence-absence Coded-abundance
No. Biological trait Trait modality HB SB HB SB
1 Maximum potential size 5 mm <0.001 0.019 <0.001 0.072
2 >510 mm <0.001 0.824 <0.001 0.008
3 >1020 mm <0.001 <0.001 <0.001 0.002
4 >2040 mm <0.001 0.001 <0.001 0.009
5 >40 mm 0.004 0.030 <0.001 0.366
6 Maximum potential number of semivoltine 0.252 0.363 <0.001 0.290
7 reproductive cycles per year univoltine <0.001 <0.001 <0.001 <0.001
8 plurivoltine <0.001 <0.001 <0.001
b
<0.001
9 Number of reproductive cycles per 1 <0.001 0.109 <0.001 0.625
a
10 individual 2 <0.001 0.073
b
<0.001 0.549
11 Life duration of adults 1 day <0.001 0.001 <0.001 0.146
12 >110 days <0.001 0.005 <0.001 0.400
13 >1030 days <0.001 0.554 <0.001 0.024
14 >30365 days <0.001 0.029 <0.001 0.166
b
15 >365 days <0.001 0.689 <0.001 0.477
16 Reproductive technique asexual 0.002 0.055 <0.001 0.517
b
17 hermaphroditic <0.001 0.025 <0.001 <0.001
18 sexual <0.001 0.113
a
<0.001 <0.001
19 Oviposition site submerged <0.001 0.085 <0.001 <0.001
20 water surface <0.001 0.008 <0.001 0.083
21 terrestrial <0.001 0.002 <0.001 <0.001
22 eggs endophytic <0.001 0.697 0.393 0.126
23 Egg/egg mass free <0.001 0.002 <0.001 0.105
24 cemented <0.001 0.334 0.007 0.095
25 female bears eggs in/on body <0.001 0.002 <0.001 0.004
26 Dissemination potential low (10 m) <0.001 0.039 <0.001
a
0.188
27 medium (1 km) <0.001 0.103
a
<0.001 0.114
a
28 high (>1 km) <0.001 0.020 <0.001 0.057
29 Attachment of aquatic stages swimmers (water column) 0.003 0.307 <0.001
a
0.487
30 to substrate crawlers (epibenthic) <0.001 0.948 <0.001 0.021
31 burrowers (infauna) <0.001 0.037 0.327 0.099
32 attached <0.001 0.135 <0.001 0.002
33 Body fexibility none (<10) <0.001 <0.001 0.005 0.720
34 low (1045
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35 high (>45
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a
<0.001 0.034
36 Body form streamlined 0.118 <0.001 0.168 <0.001
37 fattened (dorso-ventral or lateral) <0.001 0.217 0.092
a
0.182
38 cylindrical <0.001 0.081 0.038 0.002
39 spherical <0.001 0.693 <0.001 0.063
40 Feeding habits shredder 0.936 0.030 <0.001
a
0.095
41 scraper <0.001 0.034 <0.001 <0.001
42 deposit-feeder 0.028
a
0.023 0.681 <0.001
43 flter-feeder <0.001 0.058 <0.001 0.064
44 predator <0.001
b
0.014 <0.001 <0.001
45 algal piercer <0.001 0.939 <0.001 0.989
46 Dietary preferences strong (specialist) <0.001 0.066 <0.001 <0.001
47 moderate <0.001 0.058 0.429 <0.001
48 weak (generalist) <0.001 0.069 0.360
a
<0.001
49 Respiration of aquatic stages tegument 0.024 0.072 <0.001 0.008
50 gills <0.001
a
0.602b <0.001 0.038
51 plastron <0.001 0.135 <0.001 0.046
52 aerial <0.001 0.809 0.240 0.558
53 Aquatic stages adult, larva <0.001 0.024 <0.001 0.574
54 adult or larva <0.001 <0.001 <0.001 0.035
b
55 larva, pupa <0.001 0.113 <0.001 0.079
874 New Zealand Journal of Marine and Freshwater Research, 2009, Vol. 43
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875 Stark & PhillipsSeasonal variability in biotic indices

signifcant) followed by taxon richness and MCI-sb
(both within 4.7%), and %EPT richness (11.2%
of average).
Similar results were obtained for the MCI-sb
when data from four stream quality classes were
analysed separately. As for the MCI, spring and/or
winter were the seasons with highest MCI-sb values
with autumn and summer the lowest. The ranges
between seasonal means were 9.3% overall, 12.5%
for poor, 8.6% for fair, 5.1% for good, and 12.0%
for the excellent quality class, although seasonal
variability in MCI-sb was not statistically signifcant
for the good and excellent quality classes.
Species traits
Hard-bottomed streams
Most trait modality frequencies showed statistically
significant differences across seasons based
on presence/absence and coded-abundance data
collected from hard-bottomed streams (Table 4). Of
55 traits tested, 52 showed statistically signifcant
seasonality based on analysis of presence-absence
data, but only three of them showed the same (deposit
feeders, gill respirers) or inverse (predators) seasonal
pattern as the MCI. Forty-six of 55 traits showed
signifcant seasonality based on analysis of coded-
abundance data, but only four showed the same
(low dispersal, swimmers, shredders) or inverse
(plurivoltine) pattern as the SQMCI (Table 4).
Soft-bottomed streams
In contrast to hard-bottomed streams, markedly
fewer trait modality frequencies showed statistically
significant differences across seasons based on
presence/absence (13/55) and coded-abundance
(21/55) data collected from soft-bottomed streams
(Table 4). Although fve trait modalities showed
the same (sexual reproduction, medium dispersal,
fexible bodies) or inverse (multiple reproduction,
gill respirers) seasonal pattern as the MCI-sb based
upon presence-absence data, in all of these examples
the trait modality seasonality was not statistically
signifcant. Similarly, fve trait modalities showed
the same (reproduces once, medium dispersal) or
inverse (lifespan of up to a year, asexual, aquatic
adult or larva) seasonal pattern as the SQMCI-sb,
but none was statistically signifcant (Table 4).
Infuence of source of fow on metric seasonality
Although taxon richness, % EPT richness, MCI,
MCI-sb, and SQMCI (but not SQMCI-sb) showed
signifcant seasonal variation when data from all
hard-bottomed (Table 2) and soft-bottomed (Table 3)
Taranaki rivers and streams were analysed together,
statistically signifcant seasonal patterns were not
always consistent when rivers were classifed by
source of fow (Table 5).
Taxon richness showed significant seasonal
variation only for hard bottomed streams on the
Taranaki ringplain (RPOP, RPNP) (Table 5). For both
river classes, taxon richness was highest in summer,
followed by spring, autumn, and winter (Table 6).
Taxon richness in winter was signifcantly lower
than in the other three seasons at approximately
8.211.3% less than the annual average.
%EPT richness showed statistically signifcant
seasonal variation only in hard-bottomed ringplain
streams with a national park source of fow (RPNP),
and soft-bottomed non-ringplain lowland streams
(LOWL) (Table 5). In both examples, %EPT
richness was highest in spring, followed by winter,
summer, and autumn (Tables 6 and 7). Spring %EPT
richness values were 8.3% (hard-bottomed RPNP)
to 12.7% (soft-bottomed LOWL) higher than the
annual average (Tables 57).
MCI showed significant seasonal variation in
ringplain streams with a national park source of
fow (RPNP) and non-ringplain hill country (HILL)
streams (Table 5). In these source of fow classes,
spring and winter MCI values were signifcantly
greater than those recorded in summer or autumn
(Table 6). In RPNP streams, seasonal variation was
within 3.5% of the annual mean (cf. within 5.8%
for HILL streams) (Tables 5 and 6).
MCI-sb showed signifcant seasonal variation only in
lowland non-ringplain (LOWL) soft-bottomed streams,
with seasonal mean values within 3.7% of the annual
mean (Table 5). Winter, spring, and summer MCI-sb
values were all higher than average and signifcantly
greater than autumn values (Table 7).
SQMCI showed signifcant seasonal variation only
in ringplain streams with a national park source of fow
(RPNP) with seasonal mean values within 6.2% of
the annual mean (Table 5). Winter and spring SQMCI
values were higher than average and signifcantly
greater than autumn and summer values (Table 6).
SQMCI-sb showed signifcant seasonal variation
only in lowland non-ringplain (LOWL) soft-bottomed
streams, with seasonal mean values within 8.1% of
the annual mean (Table 5). Spring, summer, and
winter MCI-sb values were all signifcantly greater
than autumn values (Table 7).
Flow variability
Analysis of the disturbance regimes of 14 hard-
bottomed streams in Taranaki revealed signifcant
876 New Zealand Journal of Marine and Freshwater Research, 2009, Vol. 43
seasonal differences in the number of days between
floods defined as three times median flow or
greater (F(3,1072) = 35.757, P < 0.0001). Post hoc
testing (Tukey HSD test) indicated that the time
between foods was signifcantly less in winter
(11.7 days) or spring (12.8 days), than it was in
summer (22.4 days) or autumn (20.5 days). The
biotic indices that showed signifcant seasonality
(viz., MCI, SQMCI, MCI-sbsee Tables 2 and
3) tended to exhibit higher than annual average
values in seasons with more frequent signifcant
freshes. Examination of MCI versus time since
the last signifcant fresh indicated that sampling
approximately 1520 days after a fresh that
Table 6 Statistically signifcant seasonal variation in taxon richness, %EPT (Ephemeroptera, Plecoptera, Trichoptera)
richness, Macroinvertebrate Community Index (MCI), and Semi-Quantitative (SQ)MCI from hard-bottomed streams in
Taranaki (New Zealand) assessed by ANOVA with signifcant differences identifed using post hoc Tukey HSD multiple
comparisons tests. n = no. of samples. Lower case letters (ac) span seasons that did not differ signifcantly.
Seasonal mean (as % of overall site means) ANOVA
Spring (Sp) Summer (Su) Autumn (Au) Winter (Wi ) Tukey HSD
Hill country streams (HILL)
MCI 105.83 95.07 94.57 101.01 Sp Wi Su Au
SE (n) 1.31 (79) 1.47 (56) 1.84 (44) 1.32 (53) a a b b
Ringplain streams with source of fow within Egmont National Park (RPNP)
Taxon richness 100.35 103.74 98.39 91.78 Su Sp Au Wi
SE (n) 0.92 (553) 0.92 (537) 1.10 (362) 1.68 (185) a b b c
%EPT richness 108.31 95.03 92.10 105.01 Sp Wi Su Au
SE (n) 0.91 (553) 0.83 (537) 1.17 (361) 1.81 (185) a a b b
MCI 103.45 97.49 98.03 100.82 Sp Wi Au Su
SE (n) 0.39 (553) 0.36 (537) 0.53 (362) 0.75 (185) a b c c
SQMCI 104.92 97.87 97.36 106.22 Wi Sp Au Su
SE (n) 0.94 (539) 0.77 (824) 1.10 (357) 1.60 (157) a a b b
Ringplain streams with source of fow outside Egmont National Park (RPOP)
Taxon richness 101.29 102.34 99.85 88.73 Su Sp Au Wi
SE (n) 1.78 (134) 1.14 (151) 1.72 (125) 3.12 (45) a a a b
Table 7 Statistically signifcant seasonal variation in %EPT (Ephemeroptera, Plecoptera, Trichoptera) richness,
Macroinvertebrate Community Index (MCI)-sb, and Semi-Quantitative (SQ)MCI-sb from soft-bottomed streams
in the Taranaki region (New Zealand) assessed by ANOVA with signifcant differences identifed using post hoc
Tukey HSD multiple comparisons tests. n = no. of samples. Lower case letters (ac) span seasons that did not differ
signifcantly.
Seasonal mean (as % of overall site means) ANOVA
Spring (Sp) Summer (Su) Autumn (Au) Winter (Wi ) Tukey HSD
Lowland non-ringplain streams (LOWL)
%EPT richness 112.73 95.29 92.02 107.55 Sp Wi Su Au
SE (n) 5.57 (41) 3.86 (81) 4.18 (45) 7.76 (29) a ab b b
MCI-sb 103.22 100.56 93.72 103.64 Wi Sp Su Au
SE (n) 1.22 (41) 1.17 (81) 1.19 (45) 2.02 (29) a a a b
SQMCI-sb 105.49 102.37 91.87 98.76 Sp Su Wi Au
SE (n) 2.34 (35) 1.74 (81) 2.14 (43) 2.15 (28) a a a b
exceeded three times median fow was likely to
yield biotic index values that were within 2% of
the annual average values.
DISCUSSION
Biotic indices used for measuring stream health
should increase or decrease in response to changes
in the quality or degree of pollution of a site, and
be relatively unaffected by other environmental
factors. A useful index should be consistent over
time at a site if no change in quality occurs during
the sampling period (Karr et al. 1987).
877 Stark & PhillipsSeasonal variability in biotic indices
Seasonality in some overseas biotic indices has
been attributed to macroinvertebrate life histories,
as many indicator taxa for high water quality (e.g.,
various stonefy and mayfy taxa) favour cold water
and those that indicate moderate or poor water
quality (e.g., some Diptera) are less prominent at
colder times of year in a given stream (Linke et al.
1999). Hilsenhoff (1988) found that Biotic Index (BI)
values in winter were lower in warmwater streams
and higher in coldwater streams. He attributed the
differences in community composition (and hence BI
values) to the infuences of temperature seasonality
on invertebrate life histories, but noted that stream
temperature does not appear to affect the BI, except
for its infuence on seasonal differences (Hilsenhoff
1988). Soulsby et al. (2001) found that invertebrate
communities change seasonally largely as a result
of variation in their species life cycles. However, we
expected that biotic index seasonality in New Zealand
would not be strongly linked to the presence, absence,
or relative abundances of invertebrates owing to their
life histories. As Winterbourn et al. (1981) noted,
many New Zealand stream insects have broad habitat
requirements, lack feeding niche differentiation, and
possess fexible, poorly synchronised life histories.
They contended that abiotic factors have been of
paramount importance in the shaping of New Zealand
stream communities, via selection for opportunism
owing to the prevalence of steep, unstable streams
and the unpredictability of the physical environment
(Winterbourn et al. 1981). Death (1995) considered
that habitat stability acted as a bottleneck and was the
principal structuring force in New Zealand stream
macroinvertebrate communities, and that unstable
streams had similar faunas that can survive and/or
recolonise after food events. However, when the
bottleneck was absent (e.g., in more stable lake- or
spring-fed systems) communities tuned to other
abiotic or biotic factors could become established
(Death 1995).
Our fndings are consistent with those of Bche et
al. (2006), who examined seasonal variation in trait
composition of intermittent and perennial streams
in northern California, United States, and concluded
that biological traits varied less and exhibited greater
seasonal stability in unpredictable and harsh stream
environments than those in more moderate stream
environments.
Flow variability
Floods and low flows are known to influence
riverine macroinvertebrate community composition
densities, taxonomic composition, and taxon
In Aucklands soft-bottomed streams, Stark &
Maxted (2007a) recorded MCI-sb values from 28
to 157 (cf. 32123 in Taranaki), and SQMCI-sb
values from 0.838.61 (cf. 0.928.39 in Taranaki).
The comparatively low maximum values of %EPT
richness (58.33%) and MCI-sb (123), in particular,
reflect the fact that the soft-bottomed streams
sampled in Taranaki tended to be within farmland
or at low altitudes (<350 m above mean sea level)
rather than on the upper ringplain near Egmont
National Park boundary or in pristine native forest
catchments (as were the healthiest soft-bottomed
streams in Auckland).
New Zealands MCI is relatively unaffected by
water depth, current velocity, and substrate, provided
that samples are collected from stony riffes (Stark
1993). However, our research showed that biotic
indices exhibited statistically signifcant variation at
sites unaffected by anthropogenic disturbance. For
example, in Taranaki rivers and streams, the MCI,
MCI-sb, SQMCI, and SQMCI-sb showed seasonal
variation within 3.04.7% of the annual site mean
values. Seasonal variability of other indices, such as
taxon richness (4.77.7%) and %EPT richness (
7.411.2%) was greater.
Spring and winter MCI and SQMCI values tended
to be signifcantly higher than those calculated from
samples collected in summer or autumn. However,
most values were within 5% of the annual site
mean.
Possible explanations
for seasonal variation in biotic indices
Species traits
Although seasonal variation in the life histories of
aquatic macroinvertebrates has been shown to infuence
biotic index values (e.g., Hilsenhoff 1988; Linke et
al. 1999), our examination of seasonal variation in
species trait modalities and their relationships to the
observed seasonal patterns of biotic indices did not
support this explanation. In soft-bottomed streams,
no statistically signifcant links between species traits
and biotic indices (MCI-sb and SQMCI-sb) were
detected. In hard-bottomed streams, plurivoltine life-
histories, low dissemination potential, swimming
aquatic stages, dorso-ventrally fattened body form,
feeding habitats (deposit feeding, shredder, predator),
and weak dietary preferences all showed the same (or
inverse) seasonal pattern as the MCI or the SQMCI
(but not both). However, there was no commonality
between the analyses undertaken using presence-
absence data (for the MCI) and coded-abundance
data (for the SQMCI).
878 New Zealand Journal of Marine and Freshwater Research, 2009, Vol. 43
richness (McElravy et al. 1989; Clausen & Biggs
1997, 2000; Suren & Jowett 2006). It follows that
biotic indices, which are a measure of community
structure, may vary in response to changes in river
fow, especially in streams with highly variable fow
regimes. When rivers are in recession (especially
during warmer months when periphyton growth
is more rapid) invertebrate communities can
become dominated by aquatic worms, snails, and
chironomids, while densities of mayfies, stonefies,
and caddisflies decline or are eliminated (Biggs
2000; Jowett 2000). Signifcant freshes may reset
aquatic communities by fushing away periphyton
accumulations and associated macroinvertebrates,
and restoring communities dominated by high scoring
taxa such as mayfies, stonefies and caddisfies,
which are washed downstream from better-quality
tributaries or habitat upstream. Since periphyton-
associated macroinvertebrates tend to be lower
scoring than thin-flm diatom community-associated
mayfies, stonefies and most caddisfies, these fow-
mediated changes in macroinvertebrate community
composition are refected in biotic indices such as
the MCI, SQMCI, and %EPT.
The fnding that samples collected from rivers
during seasons in which signifcant foods were more
frequent tended to have higher biotic index values
(and vice versa), supports the hypothesis that the
fow disturbancestability gradient infuences biotic
indices.
Macroinvertebrate biomonitoring programmes
involve collection of samples from natural populations,
processing these samples, and calculating indices for
reporting results. Error or imprecision is inherent in
every stage of this process. Communities show high
spatial variability and samples (which seldom collect
the entire population) provide estimates (of differing
precision) of population characteristics such as taxon
richness, density, and various indices. Random errors
arise when taxa are missed, and systematic errors
may occur when sampling methods are selective
(Ostermiller & Hawkins 2004). Errors may arise
when processing samples, although standard
methods and quality control procedures (Stark et
al. 2001) aimed at minimising processing errors are
in widespread use throughout New Zealand (J.D.
Stark pers. obs.).
Stark (1998) used an ANOVA procedure
(developed by Narf et al. 1984) to determine
detectable differences for both hand-net and Surber
samples based on MCI, SQMCI, and QMCI values
from replicate samples. These analyses suggested
that MCI values calculated from single hand-net
samples collected according to Protocol C1 (Stark et
al. 2001) would need to differ by 10.83 MCI units (or
approximately 11% for an average MCI of 100) to
be considered signifcantly different. The detectable
differences for the SQMCI from single hand-net
samples (c. 17%), and the MCI (c. 22%) and QMCI
(c. 27%) from single Surber samples were somewhat
higher. MCI estimates based upon data from single
samples are more precise than for the other indices
(named above) in New Zealand (Stark 1993), more
precise than for some widely used indices elsewhere
(e.g., Hilsenhoffs (1988) FBI (c. 25%) and EPT
richness (c. 50%) in the San Pedro River in south-
eastern Arizona, United States (Gebler 2004)), but
comparable to others (e.g., Montana Multimetric
Index (1015%) (Stribling et al. 2008)).
Stark & Maxted (2007b) recommended that
the MCI and MCI-sb (but not the SQMCI, QMCI,
SQMCI-sb, or QMCI-sb) should be used for SoE
reporting in New Zealand. The MCI and MCI-sb show
less temporal variability than the semi-quantitative
and quantitative variants (and normally are highly
correlated with themStark 1993, 1998; Stark &
Maxted 2007a) and are also more cost-effective
(Stark 1993, 1998) because sample collection and
processing takes less time. The MCI and MCI-sb are
preferred over taxon richness and %EPT richness
because they showed less seasonal variability.
Furthermore, taxon richness is not strictly an index
of stream health since highest values tend to be in
slightly enriched habitats, and some pristine habitats
are species-poor (J.D. Stark pers. obs.). Furthermore,
estimates of taxon richness are highly dependent
upon sampling effort or sample size, a disadvantage
that is not shared with biotic indices such as the
MCI, SQMCI, or QMCI (because they are average
tolerance values per taxon) (Stark 1993).
Collection of single hand-net samples from
hard-bottomed (Protocol C1), or soft-bottomed
(Protocol C2) stream habitat (Stark et al. 2001) is
widespread for SoE monitoring undertaken in New
Zealand (Stark & Maxted 2007a; Taranaki Regional
Council 2007). Most regional councils sample once
per year (often during summer), but some sample
twice (during spring and autumn, or spring and
summer).
Seasonal variability is unlikely to confound
interpretation of SoE monitoring results for an
individual regional council that may undertake SoE
monitoring in one season, but for councils that sample
in two seasons (e.g., TRC: spring and summer), it
could be important to know whether or not the results
from different seasons are comparable. Information
879 Stark & PhillipsSeasonal variability in biotic indices
from the present study (Table 2) suggests that spring
MCI values were approximately 3.0% higher and
summer values were 2.1% lower than the annual
mean, and that averaging spring and summer MCI
values was likely to produce a mean that was within
0.5% of the annual mean. In this instance, we consider
that the infuence of season on index values does
not need to be considered. Seasonal effects in soft-
bottomed streams are likely also to be insignifcant
since spring and summer MCI-sb values were within
1% of the annual mean (Table 3).
If seasonal variability in biotic indices was
signifcant it could introduce spurious differences
if data from different regions, collected at different
times of year, were used for reporting on the state
of New Zealands rivers and streams. The purpose
of SoE monitoring is to collect accurate and reliable
information on the environment so that informed
decisions can be made about natural resource
management, and to provide a sound basis for setting
environmental policy (Ministry for the Environment
2007). Since it is impossible to measure every aspect
of the environment, environmental reporting relies
on indicators to measure and report on the health
of the environment in a cost-effective, practical,
and meaningful way (Ministry for the Environment
2007). SoE or consent monitoring can be used to
track environmental trends over time, detect activities
that have an impact on the environment, and to
evaluate the effectiveness of environmental policies
and management actions (Stark & Fowles 2006;
Taranaki Regional Council 2006; Taranaki Regional
Council 2009). The need for SoE monitoring to
be cost-effective and practical means that some
compromise is necessary between the intensity of
investigation that can be undertaken at each site and
the number of sites that can be included. To achieve
representative coverage of the region, the intensity of
effort expended at each site must be limited. A biotic
index, like the MCI, is an appropriate indicator of
river health for use in SoE monitoring because it can
be applied cost-effectively, is a proven indicator of
nutrient enrichment and anthropogenic disturbance
(Quinn & Hickey 1990; Maxted et al. 2003; Doldec
et al. 2006), and provides an easily understandable
summary of stream health for water managers and
the general public (Ministry for the Environment
1999; Taranaki Regional Council 2009). However,
the error associated with calculating the MCI from
single samples (11%) could result in some sites
being misclassifed. For example, the true value of
a measured MCI of 100 could lie between 89 and
111 and the site could be classifed into either the
good (100119) or fair (8099) quality class. To
avoid apparent misclassifcation, Wright-Stow &
Winterbourn (2003) suggested a return to fuzzy
boundaries between stream health classes similar to
those suggested originally by Stark (1985). However,
an apparent misclassifcation (i.e., one or two MCI
values higher or lower than most in the time series)
is unlikely to have an appreciable impact on trends
analyses, or on the overall assessment of stream
health within the region. Similarly, in our view, the
infuence of biotic index seasonality is minor and we
believe that the MCI can be used for reporting on the
health of New Zealands freshwater environments
using data collected in different seasons, without
the need for seasonal correction factors.
AckNOWLEdgmENts
Funding for this research was obtained from the Foundation
for Research, Science and Technology via a subcontract
under NIWAs Water Allocation research programme
(C01X0308). We thank Taranaki Regional Council
(TRC) for generously providing macroinvertebrate and
hydrological data from their extensive databases. Without
these data and the assistance of Chris Fowles, Peter Nolly,
and Fiona Jansma from TRC, this work would not have
been possible. Advice on statistical analyses was provided
by Darrel Amarasekera (StatSoft, Australia). John Hayes
(Cawthron Institute), Chris Fowles (TRC), and three
anonymous referees provided helpful comments on the
draft manuscript.
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