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Species traits in relation to temporal and spatial heterogeneity in streams: A

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Article  in  Freshwater Biology · April 1997

DOI: 10.1046/j.1365-2427.1997.00166.x

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Freshwater Biology (1997) 37, 367–387

Species traits in relation to temporal and spatial

heterogeneity in streams: a test of habitat templet
theory

C O L I N R . T O W N S E N D , * † S Y L VA I N D O L É D E C ‡ A N D M I K E R . S C A R S B R O O K * §
*Zoology Department, University of Otago, PO Box 56, Dunedin, New Zealand
‡URA CNRS 1974, Ecologie des Eaux Douces et des Grands Fleuves, Université Claude Bernard, Lyon 1, 69622 Villeurbanne
Cedex, France

†Author to whom correspondence should be sent

§Present address: National Institute of Water and Atmospheric Research, Hamilton, New Zealand

S U M M A RY
1. The habitat templet approach depends on defining templet axes appropriate to the
organism(s) of interest, predicting the traits of species associated with different parts of
the templet, and testing these predictions in a range of habitats whose positions in the
templet have been determined.
2. In this study of thirty-five benthic insect taxa at fifty-four tributary sites of the Taieri
River on the South Island of New Zealand, we chose as the temporal axis the intensity/
frequency of disturbance, defined in terms of bed movement during high discharge
events. As the spatial axis, we postulated that three features would provide refugia and
therefore ameliorate disturbance—percentage of the bed with low shear stress,
percentage of the bed made up of large substratum particles and availability of
interstitial space in the bed—from which we derived a combined multivariate refugium
axis.
3. More disturbed communities contained a significantly higher percentage of
individuals possessing the following traits: small size, high adult mobility, habitat
generalist (each predicted to confer resilience in response to disturbance), clinger,
streamlined/flattened and with two or more life stages outside the stream (each
predicted to confer resistance in the face of disturbance). When analyses were performed
on the percentage of taxa having particular traits, the predicted positive relationships
with average bed movement were found for high adult mobility and habitat generalist
traits.
4. The percentage of variance in trait scores explained by intensity of disturbance was
generally higher in sites with less refugia available and lower in sites further from the
headwaters. The percentage of variance explained was higher in sites recently subject to
a major high discharge disturbance, suggesting that disturbances tend to strengthen the
pattern of preponderance of resilience/resistance traits.
5. We mapped insect taxa onto the two-dimensional templet, following Grime et al.’s
triangular terrestrial plant classification. The full variety of resistance and resilience
traits were represented in insect species throughout the templet, but taxa associated with
more disturbed conditions generally displayed a larger number of resilience and
resistance traits, combined, than taxa associated with more stable stream beds.

© 1997 Blackwell Science Ltd 367

368 C.R. Townsend, S. Dolédec and M.R. Scarsbrook
Introduction
Ecology has to deal with a bewildering variety of resistance traits might still persist in moderately dis-
organisms and environments, and a basic aim is to turbed habitats.
understand the relationship between them (Begon, These predictions received their first multisite test
Harper & Townsend, 1996). In this paper we follow using physicochemical and taxonomic data accumu-
Southwood (1977, 1988) and focus on the premise that lated, for various purposes, over a period of 17 years
the habitat provides the templet on which evolution from a wide variety of habitat types in the River
forges characteristic life-history strategies. On an Rhône in France (Statzner, Resh & Dolédec, 1994).
evolutionary timescale, the levels of spatial and tem- Species traits for a range of taxa (including macro-
poral variation inherent in an environment act as phytes, oligochaetes, ostracods, crustaceans, insects,
selective forces for species traits conferring the ability fish, amphibians and floodplain birds) generally failed
to survive and reproduce there. In ecological time, the to conform to the predictions, indicating either that
habitat templet acts also to filter out unsuccessful the theory is wrong or that the test was inadequate
strategists from the potential pool of colonists, thereby because of methodological limitations. Resh et al.
controlling community composition (Scarsbrook & (1994) concluded that future studies should deal with
Townsend, 1993). smaller taxonomic groups, and concentrate on measur-
The success of the habitat templet approach depends ing temporal and spatial heterogeneity appropriate
on defining templet axes that relate to the scales of to the group in question in a narrower range of
space and time relevant to the organism(s) of interest, habitat types.
predicting the traits of species associated with different The second multisite test of Townsend & Hildrew’s
parts of the templet, and testing these predictions in (1994) templet theory, described here, concerns aquatic
a range of habitats whose positions in the templet insects living on the bed of fifty-four tributary sites of
have been determined. Townsend & Hildrew (1994) the Taieri River on the South Island of New Zealand.
proposed for river habitats a templet with axes of Our temporal and spatial axes deal specifically with
temporal heterogeneity and spatial heterogeneity. the habitat factors assumed to be of importance by
They assumed that temporal variation would bear a Townsend & Hildrew (1994). Thus, we chose as the
relationship with the disturbance regime to which temporal axis a measure of disturbance defined in
organisms are subjected, while increased spatial het- terms of bed movement caused by high discharge
erogeneity would ameliorate or modify the influence events. As a spatial axis, we postulated that three
of disturbances by provision of refugia within which features would provide refugia and therefore amelior-
survival was more likely. Disturbance was defined as ate disturbance: percentage of the bed with low shear
a relatively discrete event in time that tends to remove stress (Lancaster & Hildrew, 1993a,b), percentage of
organisms and opens up space or other resources that the bed made up of large substratum particles (which
can be utilized by individuals of the same or other provide firm attachment and microsites with low shear
species (Townsend, 1989). In the face of a disturbance, stress) and availability of interstitial space in the bed
populations may exhibit resilience (the capacity for a (extent of the hyporheos, which may act as a refugium
rapid return towards the density prevailing before the during high discharge events). Using data for these
disturbance) or resistance (the capacity to withstand three categories we derived a combined multivariate
the disturbance without appreciable loss of indi- refugium axis. Our limited knowledge of the biology
viduals). Along the temporal heterogeneity axis, of the species allowed us to test predictions about
species with traits conferring resilience (e.g. early the proportion of individuals and of taxa in each
age at first reproduction, short reproductive cycles, community showing six resilience or resistance traits.
potential for regeneration, ability to recolonize from We tested the proposition that potential refugium
refugia) or resistance (e.g. firm attachment, stream- availability ameliorates the influence of disturbance
lined body form, invulnerable life stages) were pre- by dividing the habitats into two subsets, those with
dicted to increase in importance. The predicted effect least and those with greatest refugium availability,
of increasing spatial heterogeneity was to reduce the and by testing the predicted relationship with disturb-
effect of disturbances so that species lacking resilience/ ance in each case. If refugia ameliorate the influence
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
 A habitat templet for streams 369
Table 1 Species traits of thirty-five insect taxa (blank 5 trait absent, 1 5 trait present). Names of expert entomologists who
supplied relevant information are given (see footnote) as are key references

of disturbance, we expected a stronger relationship of the majority of fully grown larvae 10 mm or less;
between the representation of species traits and dis- (b) high adult mobility—capacity for intercatchment
turbance in the subset of habitats with least refugia. flight; (c) habitat generalist (weak habitat preferences).
2 Resistance traits: (d) clinger—possesses behavioural
or morphological means of clinging to the substratum;
Materials and methods
(e) streamlined/flattened—possesses a morphological
Selection of species traits means of reducing drag and therefore dislodgement
in the flow (often accompanied by appropriate behavi-
A total of fifty-eight aquatic insect taxa were identified our); (f) at least two developmental stages spent
in the study area. Sufficient data from published outside the stream environment (usually adult plus
papers, unpublished theses and personal communica- egg or adult plus pupa, but in the case of mayfly
tions from expert entomologists were available to nymphs (Ephemeroptera) subimago plus adult; note
characterize life history traits for thirty-five species that the vast majority of stream insects have aquatic
(Table 1). larvae but terrestrial adults).
Each species was coded as possessing or lacking Traits (a)–(c) can be thought of as having the poten-
each of the following traits (Table 1). tial to provide resilience. In this connection, body size
1 Resilience traits: (a) small body length—body length is assumed to be negatively correlated with generation
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
370 C.R. Townsend, S. Dolédec and M.R. Scarsbrook
time (a characteristic for which we lack reliable data); at high discharge, providing potential refugia
thus, small species are expected to be more resilient (Lancaster & Hildrew, 1993a,b).
because the period from hatching to oviposition is The second variable is the percentage of large par-
shorter. Traits (d)–(f) can be thought of as having the ticles (exceeding 128 mm in width). The positive rela-
potential to provide resistance. tionship between critical shear stress required for
The set of traits available for this analysis was incipient motion and particle size (Carling, 1983;
restricted because of the limited knowledge of the Komar, 1988) suggests that large particles will be
New Zealand benthic fauna. It should be noted that more likely than small ones to retain or accumulate
our choice of traits, and the categorization of species individual organisms during high discharge events.
as possessing or lacking them, were made a priori, Evidence on this point is meagre, but the results of
before carrying out any analyses to test our predictions. McAuliffe (1984) indicated that large stones were
In every case, the relative importance of a trait (meas- more likely to retain certain sedentary species, and
ured as the proportion of individuals or taxa in the Scarsbrook (1995) showed in a tributary included in
community possessing it) was predicted to increase the present study (no. 24) that invertebrate density
with disturbance frequency/intensity but to decrease associated with large particles actually increased after
with refugium availability. a high discharge event in 1991. Substratum size distri-
bution was estimated using the method of Wolman
(1954), in which 100 randomly chosen particles are
Selection and characterization of stream sites
measured at each site and arranged in size classes
Two sites (30 m reaches) were chosen in each of corresponding to the Wentworth–Lane scale (Mosely,
twenty-seven similarly sized tributaries of the Taieri 1982).
River (Fig. 1). Distance between the upper and lower The third variable is an index of hyporheic volume
sites ranged from µ 200 m to 15 km (usually greater (interstitial space in the bed inhabitable by insects).
than 1 km). The physical habitat at each site, including An estimate of the potential depth of the hyporheic
the spatial axis of our templet, was measured during zone was obtained by averaging the depth to which
the period from September 1993 to March 1994. The a metal sounding rod (16 mm diameter, 1.5 m length)
temporal dimension was assessed from measurements could be hammered into the substratum at ten ran-
made on five occasions during a period from Sep- domly chosen points. Studies in eleven of the fifty-
tember 1993 to June 1994. four tributaries have shown that a diverse hyporheic
Three variables were used to characterize the spatial fauna exists in gravel sediments (whose extent is
axis of our templet. Our intention was to obtain measured by the sounding rod) to the depth limit of
measures that are likely to be related to refugium the sampling methods used (45 cm; Scarsbrook, 1995;
availability, but it was not possible to quantify refugia R. Montgomery, personal communication).
directly or to be certain that insects actually use them To estimate bed movement during high discharge
(this point is considered further in the Discussion). events, painted tracer particles corresponding to the
The first spatial variable is the proportion of the 50th, 75th and 90th percentiles of the substratum
stream bed within the site that is subject to low shear particle size distribution (excluding bedrock) were
stress, measured as the percentage of 100 random arranged on the bed of the stream in regular arrays
point shear stress measurements, measured at base- (five rows of three particles in each size class randomly
flow, where the value did not exceed 0.771 dynes assigned to transects at 1 m intervals). The movement
cm–2. This value corresponds to the critical shear stress of these particles was monitored on five occasions
required to move a hemisphere (3.85 cm radius) with from September 1993 to June 1994, during which
density 1.015 g cm–3, the lightest in a set of twenty- period some unusually high discharge events
four hemispheres differing only in density which are occurred. On each occasion, particles that had moved
used to estimate near-bed shear stress (Statzner & since the previous monitoring occasion were noted
Müller, 1989; Statzner, Kohmann & Hildrew, 1991). It and replaced. Our index of bed movement was calcu-
was not possible to estimate shear stress under high lated as the average of the percentage of the painted
discharge conditions; however, some fraction of the tracer particles of all size classes that had moved
low shear stress locations can be expected to remain between consecutive sampling occasions.
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
 A habitat templet for streams 371

Fig. 1 Location of stream sites in

tributaries of the River Taieri on the
South Island of New Zealand. Two
sites were selected in each of twenty-
seven tributaries. U, upper site; L,
lower site.

We had two choices for the temporal/disturbance proved to be strongly and positively correlated with
dimension of the templet: the frequency of disturb- each other (correlation coefficients of 0.767, 0.889, 0.874
ances of a given magnitude at each site or the intensity and 0.765 for frequencies of 20, 40, 60 and 80%,
(average magnitude) of disturbance. The first respectively). The two variables measure essentially
approach, measuring the frequency of occurrence of a the same thing. Very similar relationships were
disturbance of arbitrary magnitude, has two limita- obtained between species traits and both mean per-
tions: first, there is no obvious rationale for choosing centage bed movement and frequency of bed move-
the percentage movement to consider (20, 40, 60, 80% ment (arbitrarily taken to be 40%), except that
of particles?) and, second, since only five periods correlations were generally stronger in the former
had been monitored for bed movement the resulting case. Throughout the results, we use the continuous
frequency would be a discrete rather than continuous mean bed movement variable as our disturbance axis.
variable. On the other hand, the mean percentage To provide a context for our study, though generally
movement during the five periods is a continuous not to produce data to be used in the analyses, the
variable. In fact, intensity and frequency of disturbance following were also measured for each site at baseflow
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
372 C.R. Townsend, S. Dolédec and M.R. Scarsbrook
conditions: (i) stream width—measured at six cross- (NIWA Dunedin, personal communication). In con-
sections; (ii) stream depth—measured at three points trast, the 1990 sampling followed a period of low flows
across each cross-section; (iii) channel slope—meas- without major flood disturbances (mean discharge for
ured over the 30 m reach with an Abney level; the period 30 November 1989 to 31 March 1990 5
(iv) distance downstream from furthest headwater— 6.2 m3 s–1, maximum discharge 5 110.3 m3 s–1). Thus,
estimated from topographical maps (scale 1 : 50 000); an opportunity was presented for assessing the influ-
(v) median substratum particle size; (vi) average dis- ence on invertebrate communities of the timing of the
charge; (vii) coefficient of variation of discharge; most recent major disturbance.
(viii) flood frequency—taken to be frequency per year
of flows at least five times as great as the median flow.
Discharge was continuously monitored on the main Analysis
stem of each catchment over the course of the study Each independent a priori hypothesis (percentage of
by the National Institute of Water and Atmospheric individuals showing one given trait vs. one templet
Research (NIWA). In addition, each tributary had axis) was studied using a simple linear regression
maximum flow indicators and was regularly gauged model. The significance of the regression coefficient
by NIWA staff to allow the correlation of flows in the (slope) against zero was tested using analysis of
tributaries with those of the main stem. variance (null hypothesis rejected if Fobs ù F1 – α with
an alpha of 0.05). We further explored the relationships
Invertebrate sampling of percentage of individuals showing traits vs. templet
axes by reference to the proportion of variance
While the physical measurements at each site were all explained by the linear regression and the sign of the
taken during 1993–94, the biological measurements correlation coefficient. The use of a linear regression
derive from two periods. On tributaries 1–21, four model requires data to be normally distributed. Tests
Surber samples (area 0.06 m2, 300 µm mesh, two each of normality (Lilliefors, 1967) showed that a log trans-
in a pool and a riffle) were taken from random formation was required for average bed movement,
locations in the 30 m study reaches in the austral index of hyporheic volume and percentage of large
summer between 2 January and 7 February 1990 as particles.
part of a large-scale, long-term community-level study A multivariate ordination of three variables (index
of sites within the Taieri River system. Frequent visits of dead space, percentage of large particles and index
confirmed that size class composition of the bed of hyporheic volume) was used to define our spatial
material at these sites has not changed between 1990 axis. Each of our three variables was scaled between
and 1994 (authors’ observations). Therefore, we 0 and 1 as follows:
assumed our index of disturbance, derived in 1993–
94, is an acceptable representation of conditions in xij 5 (zij – min.)/(max. – min.)
1990. In order to increase the number of sites in
our study, we sampled macroinvertebrates at sites in where zij is the value of the jth variable for the ith
tributaries 22–27 using the same techniques between site, min. is the minimum value of the jth variable,
17 January and 1 March 1994. We intended that for and max. is the maximum value of the jth variable. A
analytical purposes the two sets of samples would be non-centred principal components analysis (Lebart,
equivalent (invertebrate community species composi- Morineau & Fénelon, 1982) then ordered the sites
tion within a site shows little variation in different according to their refugium availability. The first com-
summers; authors’ observations). However, a few ponent of such an analysis is always unipolar (Noy-
weeks before the 1994 sampling some very high dis- Meir, 1973); that is, all values for sites and/or variables
charge events occurred in the Taieri catchment, the are positive or negative. Consequently, this analysis
largest of which (1467 m3 s–1 measured below the takes into account the additive effect of the refugium
confluence of tributaries 5–27) occurred on 22–23 variables (Whittle, 1952).
December 1993 and caused extensive damage to low- We used principal components analyses (PCA) to
lying areas of the Taieri River drainage. The return investigate the covariations among the six selected
period for this event was in the order of 30 years traits. Prior to PCA, data were centred as follows:
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
 A habitat templet for streams 373
xij 5 (zij – z̄j)/sj The physical differences (spatial and temporal), both
within and between tributaries of the Taieri River,
where zij is the value (% individuals or % taxa) of the form the physical habitat templet which we have
jth trait for the ith site, z̄j is the mean percentage and summarized in terms of the intensity of bed movement
sj is the standard deviation of the jth trait. and the availability of potential refugia.
The locations of thirty-five taxa in the habitat templet
were computed as the weighted average distribution
along the intensity of disturbance axis (or the spatial Definition of the habitat templet
axis) as follows (ter Braak, 1987): The locations of the fifty-four stream sites in the
k templet are shown in Fig. 2. Note that the positions
uk 5 Σ y x /y
ik i .k of sites from within the same tributary are not necessar-
i51 ily in close proximity in the templet. Thus, pairs of
where uk is the weighted average of the kth species sites vary considerably in disturbance intensity (both
(k ø 35), y.k is the total abundance of species k, and xi because of different discharge regimes, if other tribu-
is the disturbance or refugium availability value for tary streams enter between the two sites, and because
the ith site (i ø 54). The explanatory power of a different substratum size distributions influence the
score (intensity of disturbance or spatial axis) was extent to which a given discharge event moves the
investigated using a dispersion index (δ of ter Braak, stream bed) and/or refugium availability (refer to
1987) as follows: Table 2).
m
δ5 Σ y u /y
k51
2
.k k .. (m ø 35)
Species traits
where y.. is the overall total abundance. The statistical Compositions of the fifty-four communities, in terms
significance of the dispersion of species was checked of the thirty-five taxa included in the analyses, are
using a random permutation test. In this test, 1000 delta described in Table 3. Note that where specific names
indices were computed by randomly permutating the could not reliably be assigned, taxa have been grouped
values of intensity of disturbance (or the spatial axis). at the genus level. Likewise, where species have not
We used ADE version 4.0 (Thioulouse et al., 1995) been formally described, family or subfamily names
to compute multivariate ordinations and dispersion have been given. From the taxon-by-site table (Table 3)
indices, and to produce the graphics. and the table of species traits (Table 1) we calculated
the percentage of total individuals per site and the
percentage of total taxa per site demonstrating each
Results trait.
Physical nature of the stream sites A typology of traits using the percentage of
individuals in sites possessing each trait was pro-
The sites chosen for study reflect the range of stream cessed by a PCA centred by trait (Fig. 3a). The first
habitats to be found in headwaters of the Taieri River two axes successfully described the covariance among
catchment (Table 2). Mean discharge at the sites ranged traits (explaining 73% and 13% of variance, respect-
from 9 to 1760 l s–1, with flood frequency ranging from ively). The positive scores for all traits on axis 1
three to twenty-six times during 1993–94. Channel indicates that they tend to co-occur in sites. The
morphology also varied widely between sites (slope strongest correlations were found among streamlined
varied from 0.11% to 9.92%; stream width : depth ratio, or flattened bodies, two or more stages outside the
a measure of channel cross-sectional shape, ranged stream, high adult mobility and habitat generalist
from 4.52 (U-shaped) to 49.03). Median particle size traits; thus, resistant and resilient traits tend to occur
ranged from medium gravel (–4.0φ 5 16 mm) to large together. An equivalent analysis of traits using the
cobble (–8.0φ 5 256 mm). Sites ranged from being percentage of taxa possessing each trait (Fig. 3b;
situated in the extreme headwaters (0.45 km of stream respectively, 36, 26 and 16% of variance explained
above the site) to being at the downstream end of by the first three axes) indicates that the traits high
long catchments (161.29 km of stream above the site). adult mobility and habitat generalist occur together
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
374 C.R. Townsend, S. Dolédec and M.R. Scarsbrook
Table 2 Variables measured at fifty-four stream sites (refer to Fig. 1) describing (i) the spatial axis of the templet [1, % low shear
stress (,0.771 dyn cm–2); 2, % large particles (diam. .128 mm); 3, mean depth of hyporheic zone (m); 4, combined multivariate
refugia axis); (ii) the temporal axis of the templet (5, average % bed movement; 6, frequency of bed movement events .40%); (iii)
general site characteristics (7, mean discharge (1 s–1); 8, coefficient of variation of discharge; 9, flood frequency (flood 5 discharge
greater than 53 median); 10, channel slope (%); 11, mean stream width (m); 12, mean stream width/depth ratio; 13, median
particle size (phi); 14, mean shear stress (dyn cm–1); 15, length of stream above study site (km)].

in the same species (high positive scores on axis 1), axis 2 and negative scores on axis 3). Small size is
as do two or more stages outside the stream and quite independent of the other traits (negative scores
streamlined or flattened bodies (positive scores on on axes 2 and 3).
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
 A habitat templet for streams 375
1 the predicted positive relationships (though not all
with P , 0.05) were evident for every trait in the
complete data set and for all traits except clingers when
the set was divided into 1990 and 1994 components;
2 the percentages of variance in trait scores explained
by intensity of disturbance (R2) were generally higher
in the 1994 than the 1990 data (specifically for high
adult mobility, habitat generalist, clingers, streamlined
or flattened and two or more stages outside the
stream), but lower for small size;
3 the positive relationships evident for all traits in the
complete data set were also seen when the set was
divided into upper and lower sites (though not all
with P , 0.05);
4 the percentages of variance explained were generally
similar in upper and lower sets, except that a stronger
relationship between trait score and intensity of dis-
turbance was evident in lower sites for high adult
Fig. 2 The positions of fifty-four stream sites in a two- mobility and in upper sites for streamlined or flattened
dimensional templet with axes ‘intensity of disturbance’
(average % of the bed that moved in five successive periods in
bodies and two or more stages outside the stream.
a year) and ‘refugium index’ (a composite of % low shear The hypothesis that refugium availability amelior-
stress, % large particles and hyporheic volume). The numbers ates the influence of disturbance was tested by arbitrar-
refer to tributaries shown in Fig. 1; in each case the upper site ily dividing the habitats into two half-sets (n 5 27 in
in the tributary is depicted by a solid square and the lower
site by an open circle.
each case), those with smaller (refugium index , 0.69)
and those with greater refugium availability (refugium
index . 0.69), and testing the predicted relationship
with disturbance in each case. The percentages of
Relationship between templet axes and species traits
variance in trait scores explained by intensity of dis-
The proportional representation of traits among turbance (R2) were higher, for every trait except cling-
individuals in the fifty-four communities is shown ers, for the sites with smaller refugium availability
for each templet axis in Fig. 4. All six traits showed (indicating that refugia ameliorate the effect of disturb-
positive relationships with disturbance, having slopes ance; Table 4). These differences were most obvious for
significantly different from zero at P , 0.05. Two small size, habitat generalist, streamlined or flattened
traits also showed the predicted negative relationship bodies and two or more stages outside the stream.
with refugium availability: high adult mobility (R2 5 A further analysis was performed after sites had
0.229, P , 0.001) and habitat generalist (R2 5 0.248, been arbitrarily divided into half-sets (n 5 27 in each
P , 0.001). The predicted relationships for intensity case), irrespective of the tributary in which they
of disturbance are positive in each case and for the occurred, according to their distance downstream (dis-
refugium index are negative. A combined trait tance to furthest headwater , 6.2 km or . 6.2 km). All
measure, defined as the sum of percentages of all of the predicted positive relationships were evident
traits for each site in Fig. 4, was significantly in both sets (though not all with P , 0.05), but the
positively related to intensity of disturbance and percentages of variance in trait scores explained by
negatively to the refugium index (Fig. 5a,b). intensity of disturbance were higher for the upstream
Data derived from 1990 and 1994 and from upper set in all cases except small size.
and lower sites in each tributary are distinguished by One taxon, the leptophlebiid mayfly genus Deleatid-
different symbols in Figs 4(a) and (b). An exploration ium, frequently dominates New Zealand stream com-
of the relationships between trait scores and intensity munities (Winterbourn, Rounick & Cowie, 1981).
of disturbance in these different cases (Table 4) When analyses were repeated having excluded Deleati-
shows that: dium individuals, two of the predicted positive rela-
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
Table 3 Densities (m2) of thirty-five taxa (key in Table 1) in fifty-four sites.

376 C.R. Townsend, S. Dolédec and M.R. Scarsbrook

 A habitat templet for streams 377

Fig. 3 Typologies of species traits

produced using a centred principal
components analysis: (a) according to
the percentage of individuals per site
that possess the traits; (b) according to
the percentage of taxa per site that
possess the traits (arrangement of traits
according to axis 2 v 3 is inset).

tionships with disturbance were still evident (two or measure (sum of percentage of all traits at a site) was
more stages outside the stream: R2 5 0.074, P 5 0.046; recalculated after excluding Deleatidium, the predicted
habitat generalist: R2 5 0.096, P 5 0.032), as were two positive relationship with intensity of disturbance and
negative relationships with refugium availability (high negative relationship with the refugium index were
adult mobility: R2 5 0.165, P 5 0.002; habitat general- still evident but with lower R2 values (Fig. 5c,d).
ist: R2 5 0.185, P 5 0.001). When the combined trait Analyses of individual traits were also run separ-
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
378 C.R. Townsend, S. Dolédec and M.R. Scarsbrook

Fig. 4 Percentage of individuals per

site that possess each of six traits (a–f)
in relation to (A) intensity of
disturbance and (B) the refugium
index. A regression line is shown in
cases where the slope is significantly
different from zero at P , 0.05. Upper
and lower sites in each tributary are
shown as closed and open symbols,
respectively, while sites sampled in
1990 and 1994 are shown as circles and
triangles, respectively.

ately for three taxonomic groups. In the case of 0.133, P 5 0.007) and habitat generalist traits (R2 5
Ephemeroptera (including Deleatidium), positive rela- 0.128, P 5 0.008). No trait showed a relationship with
tionships were recorded with disturbance for small refugium availability. Plecoptera (stoneflies) failed to
size (R2 5 0.128, P 5 0.008), high adult mobility (R2 5 reveal any relationships. In the case of Trichoptera
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
 A habitat templet for streams 379
Fig. 5 Relationship between a
combined score of species traits (sum
of percentage of all traits at a site) and
(a) intensity of disturbance (best fit
regression: Y 5 3.227 1 1.774X; R2 5
0.205, P 5 0.0006) and (b) refugium
index (Y 5 5.045 – 1.233X; R2 5 0.145,
P 5 0.005) for the complete data set.
After excluding Deleatidium the
relationships remain significant but
with lower R2 values: (c) intensity of
disturbance (Y 5 2.615 1 0.959X; R2 5
0.105, P 5 0.017) and (d) refugium
index (Y 5 3.628 – 0.713X; R2 5 0.084,
P 5 0.033). Upper and lower sites in
each tributary are shown as closed and
open symbols, respectively, while sites
sampled in 1990 and 1994 are shown as
circles and triangles, respectively.

(caddisflies), a positive relationship with disturbance to move the median particle diameter at each site).
was evident for the habitat generalist trait (R2 5 0.091, They noted that more frequently disturbed sites tended
P 5 0.027), and negative relationships with refugium to be dominated by low-stature, potentially rapidly
availability occurred for high adult mobility (R2 5 colonizing bryophyte species, while large perennial
0.099, P 5 0.020) and habitat generalist traits (R2 5 species were characteristic of more stable sites. It
0.165, P 5 0.002). seems that to understand the distribution of benthic
When analyses were performed on percentage of species, disturbance needs to be measured in terms of
taxa (rather than individuals) in a community having bed movement. In contrast, discharge variation may
particular traits, the predicted positive relationships matter directly to fish that occupy the water column.
with average bed movement were found for high Thus, Poff & Allan (1995) found that hydrologically
adult mobility and habitat generalist traits; negative variable river habitats (high coefficient of variation of
relationships with refugium availability were found daily flows, moderate frequency of spates) in Wiscon-
for the same traits (Table 5). sin and Minnesota, U.S.A., tended to support fish
that are feeding and habitat generalists while stable
habitats were characterized by a higher proportion of
Discussion
specialists.
Many of our predictions about the prevalence of traits Our results, like those of Death & Winterbourn
in different parts of the habitat templet have been (1994), Biggs (1995), Muotka & Virtanen (1995) and Poff
supported. Thus, it seems that our a priori choice of & Allan (1995), among others, indicate that temporal
intensity of disturbance as the temporal templet axis variables are equally or more important in the deter-
was appropriate; the intensity of bed movement really mination of community composition than the more
seems to matter to the distribution of organisms in traditionally measured spatial ones (e.g. Townsend,
tributaries such as those studied in the Taieri River. Hildrew & Francis, 1983; Wright et al., 1989). We have
Since we were working with benthic insects associated expressed temporal variation in terms of the intensity
with the stream bed, it seemed likely that bed move- of disturbance to the bed (average percentage of
ment would be a critical factor. In their study of the representative particles moved during successive time
community composition of bryophytes living on the periods), having noted that in our study the frequency
beds of river sites in north-eastern Finland, Muotka of disturbance and the intensity of disturbance are
& Virtanen (1995) also found that species composition closely correlated. Whether frequency or intensity is
was related to the frequency of bed disturbance (not the more influential in determining insect distributions
measured directly, but estimated from discharge is a moot point. A greater frequency of disturbance
records and the calculated critical tractive force needed may affect the representation of species by excluding
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
 Table 4 Relationship between the percentage of individuals per site that possess each of six traits and intensity of disturbance for the total data set and for various subsets of
the data: sites sampled in 1990 (n 5 42) or 1994 (n 5 12); upper (n 5 27) and lower (n 5 27) sites in each tributary; sites with lower refugium availability (n 5 27) and higher
refugium availability (n 5 27); sites closest to (n 5 27) and furthest from the headwaters (n 5 27). R2, percentage of variation explained; P, probability that slope of the
relationship is significantly different from zero; s, sign of the best fit regression line

Distance down- Distance down-

380 C.R. Townsend, S. Dolédec and M.R. Scarsbrook

Total 1990 1994 Lower sites Upper sites Low refuge High refuge stream ,6.2 km stream .6.2 km

R2 P s R2 P s R2 P s R2 P s R2 P s R2 P s R2 P s R2 P s R2 P s

Small size 0.103 0.018 1 0.137 0.016 1 0.105 0.303 1 0.088 0.132 1 0.101 0.106 1 0.241 0.009 1 0.009 0.644 1 0.084 0.142 1 0.140 0.054 1
High adult mobility 0.108 0.015 1 0.038 0.216 1 0.746 0.000 1 0.157 0.041 1 0.054 0.242 1 0.051 0.255 1 0.049 0.265 1 0.257 0.007 1 0.003 0.391 1
Habitat generalist 0.174 0.002 1 0.101 0.041 1 0.733 0.000 1 0.187 0.024 1 0.135 0.059 1 0.189 0.024 1 0.035 0.353 1 0.251 0.008 1 0.135 0.059 1
Clingers 0.093 0.025 1 0.063 0.110 1 0.564 0.005 1 0.114 0.085 1 0.070 0.182 1 0.066 0.197 1 0.120 0.077 1 0.162 0.037 1 0.055 0.241 1
Streamlined or flattened 0.132 0.007 1 0.017 0.408 1 0.552 0.006 1 0.058 0.225 1 0.269 0.006 1 0.190 0.023 1 0.022 0.465 1 0.258 0.007 1 0.058 0.226 1
Two1stages outside stream 0.148 0.004 1 0.062 0.111 1 0.662 0.001 1 0.099 0.111 1 0.202 0.019 1 0.181 0.027 1 0.032 0.374 1 0.171 0.032 1 0.127 0.068 1

© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387

 A habitat templet for streams 381
Table 5 Relationships between the percentage of taxa per site that possess each of six traits and intensity of disturbance. R2,
percentage of variation explained; P, probability that slope of the relationship is different from zero; s, sign of the best fit
regression line

Intensity of disturbance Refugia index

Species traits R2 P s R2 P s

Small size 0.044 0.126 1 0.008 0.510 –

High adult mobility 0.078 0.041 1 0.129 0.008 –
Habitat generalist 0.137 0.006 1 0.198 0.001 –
Clingers 0.021 0.300 1 0.006 0.567 1
Streamlined or flattened 0.000 0.954 – 0.011 0.448 1
Two1 stages outside stream 0.000 0.998 1 0.006 0.574 1

those that cannot reliably recolonize in the intervals between our two templet axes (r 5 –0.35, P , 0.009),
between disturbances (not resilient at the site scale), something that is evident in Fig. 2. Thus, sites with a
whereas a greater intensity of disturbance may exclude high disturbance intensity score tend to have low
species that cannot take advantage of the small refugium availability. On the other hand, sites with
remaining area (refugium) of undisturbed bed (not intermediate or low disturbance show a wide range
resistant at the site scale). However, it seems that of refugium scores. The correlation is only partial
the two cannot meaningfully be disentangled where because the intensity of disturbance depends not only
disturbance intensity is measured as percentage of on the substratum but also on the discharge regime.
space unaffected and disturbance frequency as number A case could be made to collapse the two axes into one
of occasions when more than an arbitrarily defined disturbance intensity axis, or to derive two compound
percentage of area is disturbed. More frequently dis- axes that are orthogonal, but we have resisted these
turbed sites are also more intensely disturbed. options. First, our approach was intended to follow
Our attempts to quantify physical refugia must be the classical formulation of a priori hypotheses, rather
considered speculative; first because we do not have than the a posteriori identification of patterns. Second,
direct measures of the three refugium categories and the axes are not perfectly correlated and we believe
second because we do not know for certain whether that they contain the essence of two truly distinct
insects use the postulated refugia. Thus, we could components. Most important, the hypothesis that refu-
not directly measure amount of dead space at high gia ameliorate the effect of disturbance was supported
discharge (our index derives from low discharge condi- by showing that the percentage of variance in trait
tions) and we have no observations of whether insects scores explained by intensity of disturbance (R2) was
are less likely to be dislodged if they are in a dead generally higher, as predicted, for the sites with less
space. Our measures relating to large particles and refugium availability.
hyporheic space are similarly suspect. Nevertheless, The support gained for our predictions also indicates
the a priori choice of a multivariate refugium score for that the choice of species traits was appropriate (des-
the spatial templet axis, incorporating these imperfect pite the inevitably imprecise nature of the information
measures, seems to have some foundation because a upon which our classification is based). In their theor-
majority of species traits behaved predictably with etical formulation, Townsend & Hildrew (1994)
respect to this axis. The multivariate index was most expected the predicted traits to occur as alternative
strongly correlated with the amount of dead space at suites of characters in real species. Furthermore, they
a site (r 5 0.82, P , 0.0001) and with the percentage noted that no a priori predictions could be made about
of large particles on the bed (r 5 0.53, P , 0.0001), how many of the potential traits might be evident in
while hyporheos volume showed a positive but insig- a particular taxon, pointing out that, for example, an
nificant relationship (r 5 0.23, P . 0.05). Since the pro- especially effective holdfast might ensure the success
portion of the bed that is not disturbed (unity minus of an organism in the face of disturbances without the
the intensity of disturbance) provides some of the ‘need’ for further traits to be represented. In our
refugia that we identify, there is inevitably a correlation analysis, the traits high adult mobility and habitat
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
382 C.R. Townsend, S. Dolédec and M.R. Scarsbrook
generalist tended to occur together in the same species, stream, we would expect tighter relationships between
as did two or more stages outside the stream and the prevalence of species traits and intensity of disturb-
streamlined or flattened bodies. An equivalent analysis ance in the upper set of sites. In fact, the predicted
based on percentage of individuals having each trait positive relationships and the percentages of variance
showed that streamlined or flattened bodies, high explained were generally similar in the upper and
adult mobility, habitat generalist and two or more lower sets of sites (Table 4a); it seems that sites within
stages outside the stream tended to co-occur in sites. tributaries are largely independent of each other, at
Thus, individuals from species with different suites of least when separated by the distances established in
traits, each conferring an advantage in highly dis- this study. Thus, even at the local (intratributary) scale,
turbed locations, occurred together in predictable asso- the habitat templet serves to filter out unsuccessful
ciations; in other words, it seems that different traits strategists from the available colonists. However, an
can incur the same adaptive advantage. Townsend intriguing pattern emerges at a slightly larger scale,
& Hildrew (1994) also made the point that while when sites closer to the headwaters were compared
resilience/resistance traits might be expected to pre- with those further downstream (irrespective of
dominate in highly disturbed sites, a mixture of resili- whether they were in the same tributary). Upstream
ence/resistance and non-resilience/non-resistance sites showed the predicted positive relationships with
traits would be expected in more stable locations, five of the six traits (all except small size), while
since species with resilience/resistance traits would downstream sites showed relationships with none (at
not necessarily be excluded from the latter. This is P , 0.05). It may be that drift into downstream sites
what was observed. of invertebrates with a range of species traits from a
The mayfly genus Deleatidium often dominates, variety of types of upstream locations tends to blur
numerically, New Zealand stream communities, par- the predicted relationships between species traits and
ticularly those that are often disturbed (Winterbourn disturbance. Perhaps downstream sites are character-
et al., 1981; Scrimgeour & Winterbourn, 1989; ized not only by refugee types reflecting local, micro-
Scarsbrook & Townsend, 1993). Since this taxon pos- scale refugia but also, and predominantly, by those
sesses all the traits predicted to be associated with that use what could be called the ‘drift refugium’
disturbed situations, it might be imagined that it is provided by upstream tributaries. The results of
responsible for the set of significant correlations we Lancaster, Hildrew & Gjerlov (1996) indicate that the
found with disturbance. However, after excluding return rate to the stream bed of individuals drifting
Deleatidium two of the predicted positive relationships from upstream locations varies with both species and
with disturbance remained evident, and the combined hydraulic transport characteristics of the stream.
trait measure (sum of percentage of all traits at a site) The occurrence of a major Taieri-wide flood a few
was still significantly related to intensity of disturb- weeks before the 1994 but not the 1990 invertebrate
ance, though with a smaller percentage of variation sampling provided an opportunity to assess the influ-
explained (lower R2). Moreover, analyses performed ence on representation of species traits of the timing
on percentage of taxa, rather than percentage of indi- of the most recent major disturbance. (Note, however,
viduals, provided support for two of the six predicted that the same sites were not included in 1990 and 1994
relationships. Thus, while Deleatidium is particularly samplings, a circumstance that would have allowed
influential in providing the predicted outcomes, it is greater confidence in the results.) The predicted rela-
by no means solely responsible. tionships were generally tighter in the post-flood set
In our study, we included two sites from each of sites (R2 values ranging from 0.55 to 0.75) than in
tributary because we expected that disturbance regime the pre-flood set (largest R2 0.14). It seems that given
and availability of refugia would vary as a result time without disturbances the predicted patterns
of different substratum compositions at sites within become blurred by the arrival (either in the drift or
tributaries. Perusal of Fig. 2 confirms that there are via upstream movements or via ovipositing adults) of
often substantial differences within tributaries in dis- colonists with a wider array of traits. Stated another
turbance intensity and/or refugium availability. If way, disturbances reset the pattern of preponderance
lower sites were not independent of upper sites of resilience/resistance traits in more intensely dis-
because of the tendency for insects to drift down- turbed sites.
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
 A habitat templet for streams 383
The general hypothesis to be tested here was that An alternative approach, followed here, involves an
present-day habitat conditions are matched by present- attempt to map traits or trait combinations directly
day traits in the organisms. In this hypothesis, there- on to habitats (Southwood, 1977, 1988; Alerstam &
fore, historical events and limitations caused by Högstedt, 1982; Grime, Hodgson & Hunt, 1988; Taylor,
phylogenetic relationships were assumed not to con- Aarssen & Loehle, 1990; Silvertown, Franco &
strain the match between organism and environment. McConway, 1992; Silvertown et al., 1993). Such efforts
Nevertheless, a hint of phylogenetic limitations on have been criticized due to the difficulty they have in
what is possible is provided by the finding that the deriving generalizations; given that the definition of
Plecoptera displayed none of the six predicted trait the habitat changes with the organism under study, it
relationships, the Trichoptera displayed only one, that is simpler and more general to deal with explanations
of generalist habitat use, while the Ephemeroptera couched in terms of mortality schedules rather than
displayed a broad array. It is also important to remem- habitats (Stearns, 1992). Stearns (1992) argues that to
ber that in our analysis the dynamics of populations relate habitat to life history satisfactorily it is necessary
were assumed to be governed essentially by autecolog- to understand the underlying mechanism—not simply
ical processes, in the sense that a direct match might by relating habitat to life history but by understanding
be expected between traits and the physicochemical the links between habitat, mortality schedule and life
environment, rather than one mediated indirectly history. The disturbed habitats in the present study
through interactions with other species. The support can be viewed as imposing large, frequent, unpredict-
for a variety of hypotheses indicates that this simplify- able and density-independent mortalities on the
ing assumption was not without merit. However, we
stream inhabitants. This establishes, in turn, an advant-
already know that communities inhabiting many of
age to species possessing traits that make it less likely
the Taieri tributaries have been influenced by the
they will be killed or removed during a disturbance
arrival of an exotic fish species, the brown trout (Salmo
(resistant traits) and on species with the capacity to
trutta L). In general, only those tributaries which are
utilize available refugia (within or outside the stream)
inaccessible to trout, because of barrier waterfalls,
from where they can recolonize and rapidly reproduce
contain native fish in the genus Galaxias (Townsend &
(resilient traits).
Crowl, 1991), and there is evidence that the abund-
Finally, the positions of insect species in our two-
ances of some invertebrate species have been affected
dimensional templet are shown in Fig. 6, providing a
by this change in predation regime (Flecker &
map of species distributions in stream habitat space
Townsend, 1994; McIntosh & Townsend, 1994). Abiotic
akin to Grime et al.’s (1988) well-known triangular
factors, in particular those related to the disturbance
terrestrial plant classification. The full variety of resist-
regime, are clearly important in determining the com-
ant and resilient traits are represented in insect species
position of the invertebrate community, but biotic
interactions must certainly provide a modifying influ- throughout the templet. However, the half-set of
ence. However, if certain species are excluded by species associated with more disturbed conditions (to
their interactions with predators or competitors, what the right in Fig. 6) generally display a larger number
remains is likely to be a subset of those with traits of resilient and resistant traits, combined, than those
that match the physical environment. associated with more stable stream beds (3.00 6 0.38
Trying to understand the similarities and differences and 1.89 6 0.28 traits, respectively, t33 5 2.36, P 5
in life histories is one of the fundamental challenges 0.02).
of modern ecology, and there is a rich theory of life- Community composition in streams has tradition-
history evolution (Begon et al., 1996). The mainstream ally been defined either according to species composi-
of research has concerned optimality procedures tion (e.g. Townsend et al., 1983; Wright et al., 1989) or
which, first, deal with trade-offs limiting the set of functional feeding role (shredders of large particles,
possible trait combinations (e.g. between current collector–gatherers or collector–filterers of small par-
reproduction and survival or between number and ticles, etc.; Cummins, 1973; Vannote et al., 1980). Classi-
quality of offspring) and, second, search for the com- fication according to the representation of resistant/
bination of traits within that set which maximizes resilient traits provides a further perspective which
fitness (Lessells, 1991; Smith, 1991; Stearns, 1992). can usefully be combined with the others to provide
© 1997 Blackwell Science Ltd, Freshwater Biology, 37, 367–387
384 C.R. Townsend, S. Dolédec and M.R. Scarsbrook
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Death R.G. & Winterbourn M.J. (1994) Environmental
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The associated dispersion indices (δ, of ter Braak, 1987) are perspective. Journal of the North Americam Benthological
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We thank Chris Arbuckle for his management of the In: An Introduction to the Aquatic Insects of North America
Taieri database and for help in the field, Murray (eds R.W. Merritt and K.W. Cummins). Kendall/Hunt,
Mckenzie for making a set of Statzner’s hemispheres, Dubuque, IA.
Luc Corstens, Femke Jansma and Richard Mont- Flecker A.S. & Townsend C.R. (1994) Community-wide
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who provided access to stream sites. Special thanks streams. Ecological Applications, 4, 798–807.
to Alex Huryn, Brian Patrick, Bob Pilgrim, Richard Glasgow J.P. (1936) The bionomics of Hydropsyche colonica
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Daniel Chessel for statistical advice. The work was
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