Freshwater Biology (2011) doi:10.1111/j.1365-2427.2011.02648.

Spatial variability in macroinvertebrate assemblages

along and among neighbouring equatorial glacier-fed
streams
J E S P E R K U H N * , P A T R I C I O A N D I N O †, R O G E R C A L V E Z ‡, R O D R I G O E S P I N O S A †,
L A D I S L A V H A M E R L I K * , § , S I G N E V I E * , O L I V I E R D A N G L E S †, – A N D D E A N J A C O B S E N *
*
Department of Biology, Freshwater Biological Section, University of Copenhagen, Helsingørsgade, Hillerød, Denmark
†
Laboratorio de Entomologı́a, Escuela de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador, Quito, Ecuador
‡
IRD, Institut de Recherche pour le Développement, UMR G-EAU, Cemagref, 361 Rue Jean-Francois Breton, Montpellier Cedex 5,
France
§
Institute of Zoology, Slovak Academy of Sciences, Dubravska cesta, Bratislava, Slovakia
–
Institut de Recherche pour le Développement (IRD), UR 072, LEGS, UPR 9034, CNRS 91198 Gif-sur Yvette Cedex, France and
Université Paris-Sud, Orsay Cedex, France

SUMMARY
1. During the past two decades, understanding of the structure and function of glacier-fed
stream ecosystems at temperate latitudes has increased substantially. In contrast,
information on their tropical counterparts is very limited. We studied three neighbouring
glacier-fed streams in the tropical Andes of Ecuador. Our main goals were (i) to determine
overall longitudinal patterns in density, taxon richness and the composition of macroin-
vertebrate assemblages and driving factors in equatorial glacial streams and (ii) to examine
variability among replicate streams in faunal metrics and assemblages, and stream-specific
effects of supposed environmental key factors.
2. We measured four geographical and 17 environmental factors and collected five Surber
samples (500 cm2) of macroinvertebrates at each of nine sites, three sites along three
streams. The streams were located 1–5 km apart. In each stream, the three sites were
placed at comparable distances from the glacier and were grouped as ‘upper’ (50–200 m),
‘middle’ (1.5 km) and ‘lower’ sites (3.5–5.6 km).
3. In total, 2200 individuals (64% chironomids) were collected and 47 taxa (30 dipterans, 18
of these Chironomidae) identified. Density ranged from 176 to 372 ind. m)2, and the
number of taxa ranged from 2 to 6 at the upper sites and 868–3044 ind. m)2 and 21–27 taxa
at the lower sites. Density, number of taxa, rarefied richness and axis-1 coordinates from a
MDS ordination increased logarithmically with distance from the glacier. These faunal
metrics were equally related to altitude and glacier per cent of catchment and correlated
with maximum conductivity, mean temperature, mean daily maximum temperature and a
channel stability index. As expected, the mean difference in distance decay in similarity
was higher at the upper (47% km)1) than at the lower reaches (20% km)1) of the streams.
4. The number of taxa varied among sites within the upper and middle groups, but not
among the lower sites. In contrast, but in accordance with our expectation, assemblage
composition did not differ among upper sites but did so at middle and lower sites,
following a supposed decrease in environmental harshness along the streams. Relation-
ships between faunal metrics and the four environmental variables mean temperature, the

Correspondence: Dean Jacobsen, Freshwater Biological Section, Department of Biology, University of Copenhagen, Helsingørsgade 51,
DK-3400 Hillerød, Denmark. E-mail: Djacobsen@bio.ku.dk

 2011 Blackwell Publishing Ltd 1

2 J. Kuhn et al.

stability index, chlorophyll a and coarse particulate organic matter also varied among the
three streams. Generalised linear model analyses revealed that temperature interacted
with stream on macroinvertebrate density, while chlorophyll a had a significant effect on
the number of taxa in interaction with stream and stability.
5. The basic predictions of the Milner et al. (2001a), model regarding longitudinal faunal
patterns and temperature and stability as main driving factors were met by our three
replicate equatorial glacial streams. Qualitative departures from the model were mainly
because of zoogeographical differences. We demonstrated that variability in assemblages
between comparable sites in closely situated streams was considerable, and the effect of
key environmental factors varied among streams and interacted with other factors.
Quantifying spatial variation in benthic assemblages may help us foresee possible
consequences for biodiversity as a result of glacial retreat.

Keywords: Ecuador, glacier-fed streams, macroinvertebrates, spatial variability, stream-specific

effects, tropical Andes

largely confirmed by a longitudinal study on a glacial

Introduction
Glacier-fed streams are probably one of the harshest
environments in nature and therefore usually show a
characteristic spatial pattern in the longitudinal dis-
tribution of diversity and species assemblages down-
stream and away from the constraining influence of
the glacier (Sæther, 1968; Steffan, 1971). Diversity of
benthic macroinvertebrates is typically low close to
the glacier, but increases, often logarithmically, with
increasing distance from the glacier and decreasing
glacial contribution to stream flow [Milner et al.
(2001a)]. The increase in channel stability and maxi-
mum stream temperature (Tmax) with distance from
the glacier are thought to be the main factors affecting
the biota along these streams (Milner et al., 2001a).
The general model of longitudinal zonation in the
benthic fauna proposed by Milner et al. (2001a) was
mostly based on European and Alaskan studies
(Milner & Petts, 1994) but has been supported by
studies in other temperate regions of the world such
as New Zealand (Milner, Taylor & Winterbourn,
2001b; Cadbury, Milner & Hannah, 2010). However,
the composition of macroinvertebrate assemblages in
temperate glacier-fed streams changes seasonally over
the year, with density and diversity reaching maxima
during periods of low flow (e.g. Füreder et al., 2001;
Lods-Crozet et al., 2001; Robinson, Uehlinger & Hie-
ber, 2001). In spite of differences in seasonal dynamics
between temperate and equatorial glacier-fed streams,
the Milner et al. (2001a) model has also recently been
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
stream in equatorial Ecuador (Jacobsen et al., 2010), in
which stream temperature and channel stability also
explained most of the faunal distribution.
Even though glacier-fed streams are considered
harsh and constrained habitats with low spatial
heterogeneity (Ilg & Castella, 2006), there is consider-
able variation among streams in assemblage compo-
sition and longitudinal diversity patterns (e.g. Brittain
et al., 2001; Milner et al., 2001b). These differences do
not seem to be explicable in terms of simple regulation
by specific factors such as temperature and stability
because relationships to these factors also vary among
streams. Local assemblages are not regulated by each
environmental factor independently, but by a com-
plex interacting effect of many driving variables. For
example, the influence of temperature and stability is
clearly interdependent in determining the outcome of
competitive interactions and thus community compo-
sition (Flory & Milner, 1999). Likewise, the impor-
tance of available food sources such as periphyton
depends on the habitat stability (Death, 2002). Further,
in systems where environmental variables often are
interrelated, interpretation of mechanisms may be
difficult. For instance, in the Swiss glacier-fed Mutt
stream, maximum water temperature and periphyton
biomass had parallel gradients and an almost similar
effect on the macroinvertebrate assemblage composi-
tion (Lods-Crozet et al., 2001), while in the New
Zealand glacier-fed Rob Roy stream, these two
variables were uncorrelated and maximum water

temperature explained much more of variance in
macroinvertebrate assemblages than did algae
(Cadbury et al., 2010). These differences have been
observed comparing streams separated by large dis-
tances and may therefore be influenced by different
biological (regional species pool) and ⁄ or environ-
mental (e.g. water chemistry, stream discharge, topo-
graphy and seasonality) settings.
Assemblages, longitudinal patterns and environ-
ment–biota relationships are all expected to be largely
similar in closely situated streams. Within catchments,
we assume that stream macroinvertebrates are not
strongly dispersal limited (Thompson & Townsend,
2006; Heino & Mykrä, 2008). Consequently, in the
harshest environments close to the glaciers, we expect
strong niche selection to filter out many species from
the regional pool, so that community composition is
more predictable and the similarity between commu-
nities is higher than in more benign habitats further
away from the glacier (Leibold et al., 2004; Chase,
2007). In addition, assemblage similarity and distance
between sites typically follow a logarithmically
decreasing model (Soininen, McDonald & Hillebrand,
2007), and one of the reasons for the distance decay in
similarity between communities is a result of a
competitive sorting of species with different physio-
logical tolerances (i.e. a deterministic niche model;
Nekola & White, 1999). Along glacier-fed streams, this
means that the high rate of environmental change per
unit distance close to the glaciers is expected to
produce even larger differences in similarities per unit
distance between sites close to the glaciers than
further downstream. Comparing sites along closely
situated replicate streams gives us a good opportunity
to examine these predictions.
We sampled the macroinvertebrate fauna and
measured environmental factors at three sites along
each of three replicate, glacier-fed, neighbouring
streams on the Antisana volcano in Ecuador. Our
main goals were (i) to determine overall longitudinal
patterns in density, taxon richness and the compo-
sition of macroinvertebrate assemblages and driving
factors in equatorial glacial streams and (ii) to
examine variability among replicate streams in fau-
nal metrics and assemblages, and stream-specific
effects of supposed environmental key factors such
as water temperature, physical stream stability and
available food resources. Our a priori hypotheses
were (i) that longitudinal patterns would confirm an
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
Spatial variability in equatorial glacier-fed streams 3
earlier study in Ecuador (Jacobsen et al., 2010) as
well as the general model for glacial streams
proposed by Milner et al. (2001a) and that distance
decay in similarity would be highest closest to the
glaciers and (ii) that differences in fauna between
neighbouring streams would be minor and follow
similar relationships with environmental variables
and that similarity among replicate sites would
decrease with decreasing harshness further away
from the glaciers.
Materials and methods
Study area
The study streams were tributaries to the Rı́o Anti-
sana, a headwater of the River Napo, which in turn is
a main tributary of the upper Amazon River. It
originates at the glaciers on Mt. Antisana in the
eastern cordillera of the Andes of Ecuador (028¢S,
7809¢W, 5760 m a.s.l.), c. 50 km south of the equator.
The ablation zone extends from c. 5150 m to the
glacier snouts at 4730–4850 m a.s.l. For more infor-
mation on the study area, see the study by Jacobsen
et al. (2010).
Three streams originating side by side from three
glaciers on Antisana were selected (Fig. 1): stream
G12 (originating from glacier 12, also called Crespo,
extension 2.13 km2), stream G14 (originating from
glacier 14, extension 0.65 km2) and stream G15 (orig-
inating from glacier 15, extension 0.46 km2). On each
of these streams, three sites were selected at varying
distance from the glacier snout and situated between
4196 and 4835 m a.s.l. Stream sites were grouped
according to their location along the streams; site
group one (‘upper sites’) included those closest to the
glaciers (sites G12.1, G14.1 and G15.1), site group two,
the ‘middle sites’ (sites G12.2, G14.2 and G15.2) and
site group three, the ‘lower sites’ farthest away from
the glaciers (sites G12.3, G14.3 and G15.3). Only
stream G14 had some small intermittent, non-glacial
tributaries along the study reach; the three streams
were thus ideal for studying longitudinal glacial
influence. All stream sites were confined within an
area (c. 10 km2) small enough to be expected to
contain the same species pool and subject to similar
meteorological conditions and temporal variations.
Differences in the presence and composition of spe-
cies at a specific site should thus be due to differences
4 J. Kuhn et al.
2
1
2
1
3 2 1
3
in environmental conditions. The field work was
carried out during April–May 2009.
Environmental variables
In addition to four geographical variables (altitude,
catchment extension, per cent of catchment glacier-
covered and distance from glacier), we measured 17
environmental site characteristics. However, referring
to the conceptual model for the zonation of macroin-
vertebrates in glacier-fed streams (Milner et al.,
2001a), we paid special attention to the effect of
temperature and physical channel stability as driving
forces and to two measures of available food sources,
benthic periphyton biomass and coarse particulate
organic matter, as additional explicative factors.
At each site, electronic temperature and water-level
loggers (HOBO U20) were placed inside a plastic tube
(part of a drainpipe, length: 60 cm and diameter:
8 cm) and fastened to the bank with string and metal
tubes. The exact position in reference to the bottom of
the streambed and water surface was noted and used
in calculating the exact water level and water-level
variations. The loggers were left in the field to gather
data and retrieved 1–2 months later. From these
recordings, several variables were obtained: mean
temperature, mean daily maximum temperature,
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
Fig. 1 Map of the Antisana stream sys-
tem. Numbers 1–3 indicate the three study
sites along each of the three Antisana
streams marked by the numbers G12, G14
and G15.
mean daily minimum temperature, coefficient of
variation in temperature (=CV temp.) and CV water
depth.
Instead of the commonly used Pfankuch index
(Pfankuch, 1975), we developed and applied a new
stability index based on four measurable parameters:
CV temp., CV water depth, skewness of a conductiv-
ity versus time curve from a dilution gauging and
tractive force divided by mean substrate particle size
(from transects). The values of each of these four
parameters were ranked from 1 to 9, one being given
to the highest values of CV temp., CV depth and
tractive force ⁄ mean substrate particle size and to the
lowest value of skewness, and then the four ranks
were summed for each site (the stability index was
therefore ranked from 4 to 36). We believe this index
captures the instability caused by glacial melt water
contribution to the study region. Glacial melt water is
close to 0 C, and the daily melting and discharge
regime determines the variability in temperature and
depth. The components of the stability index and
some additional variables are explained later.
Stream slope at each site was measured using a
transparent plastic tube carrying flowing water from
the upstream to the downstream end of the reach. The
slope was calculated as the difference between the
water level inside the tube when raised until flow

stopped and that of the surface of the stream water at
the downstream end, divided by the distance between
the upstream and downstream end of the tube
(c. 25 m). At each of the nine study sites, 10 transects
with three-metre intervals were laid across the stream
and used to record width, depth and mineral sedi-
ment type. Depth and bottom material were recorded
at five points along each transect, giving a total of 50
measurements per stream. The sediment was
additionally grouped according to grain size: silt
(0.004–0.062 mm), sand (0.063–2 mm), small gravel
(2–30 mm), large gravel (30–64 mm), small cobble
(64–128 mm), large cobble (128–256 mm) and boulder
(256 mm<) following the Wentworth Scale (Giller &
Malmqvist, 1998). Based on these 50 points, the
substrate type diversity was calculated (Shannon–
Wiener index).
Tractive force was calculated as (TG = qgRS), where
TG is the tractive force (N m)2), q is the density of
water (1000 kg m)3), g is the gravitational acceleration
(9.81 m s)1), R is the hydraulic radius (m) (estimated
as width + depth) and S is the slope of the energy line.
The tractive force is a measure of the power per-
formed by the water on the streambed; with higher
values, larger particles will be moved by the water
and the requirement for the macroinvertebrates to
hold on increases. In periods with high discharge, the
level of tractive force increases.
Current velocity and discharge were measured
twice at each site at the end of our field work
during morning ‘base flow’ by means of dilution
gauging (White, 1978). A bucket of a known amount
of dissolved salt (volume and conductivity) was
added at the upstream end of the 15- to 25-m
stream reach, and we measured the conductivity
every 5 or 10 s at the downstream end of the reach.
Large skewness of the conductivity versus time
curve from the dilution gauging is considered a
measure of hydraulically dead space and thus of the
extent of low-stress refugia for macroinvertebrates.
Mean current velocity was calculated as the time
elapsed for half of the salt to pass the stream reach
divided by the length of the reach. Discharge (Q)
was calculated from these same measurements by
the formula: Q = V · C ⁄ A, where V is the volume of
salt water in the bucket, C is the conductivity of the
salt water in the bucket and A is the integrated area
beneath the curve from the conductivity versus time
plot.
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
Spatial variability in equatorial glacier-fed streams 5
Stones were collected for the quantification of
chlorophyll a as a measure of benthic algal biomass.
Five samples, each containing three stones (size: large
gravel), were collected at random, covered with 96%
ethanol and placed in the dark until processing in the
laboratory. There, the containers were given a 10-min
ultrasonic bath to increase the extraction efficiency.
After settlement for a few hours, a sample was
transferred to a spectrophotometer and absorption
measured at 665 and 750 nm. Stone surface area was
estimated using the formula: A = ((LW) + (LH) +
(WH)) · 1.15 proposed by Graham, McCaughan &
McKee (1988), where A is the area, L is length, W is
width, H is height of the stone and 1.15 is a fixed
factor to corrugate for the irregular shape of the
stones. Chlorophyll a was calculated using this
formula: Chlorophyll a (mg m)2) = Abs(665–750) ·
V ml · 10 000 cm2 m)2 ⁄ 83.4 ml mg)1 · A cm2, where
V is the volume of ethanol used, 83.4 is the absorption
coefficient for chlorophyll a in 96% ethanol and A is
the summarised area of the stones.
Macrobenthos sampling
At each site, five quantitative Surber samples
(500 cm2; mesh size 200 lm) were collected randomly
from pebble–cobble substratum in riffle ⁄ run habitats.
All samples were collected during daytime and
preserved in the field in 70% ethanol. In the labora-
tory, samples were rinsed through a 200-lm sieve and
sorted without the use of magnification. No subsam-
pling was applied. Benthic coarse particulate organic
matter (CPOM) was obtained by filtering once more
the remaining material from Surber samples through
a 200-lm sieve. CPOM was oven-dried (80 C, 24 h),
weighed, burned in a mufle furnace (for 4 h at 550 C)
and reweighed to calculate ash-free dry mass
(AFDM).
Identification of the Ecuadorian stream fauna to
species level is not possible. Invertebrates other than
non-insects were identified mostly to genus according
to Roldán (1996), Merritt & Cummins (1996), Fernán-
dez & Domı́nguez (2001) and separated into morpho-
species. Larvae and pupae of Chironomidae were
sorted under a stereoscopic microscope at 10 · mag-
nification, dehydrated in 96 and 99% ethanol and
mounted in Euparal. Larvae were identified to genus
and morphospecies under a compound microscope at
maximum 400 · magnification using current taxonomic
6 J. Kuhn et al.
literature (Wiederholm, 1983; Epler, 2001). Larvae of
the subfamily Diamesinae were identified according
to Ruiz-Moreno et al. (2000).
Data treatment
We applied four different univariate metrics to cha-
racterise the fauna: density of individuals, number of
taxa encountered in the five samples (also denoted
taxon density), rarefied richness and Pielou’s index of
evenness. To correct for the effect of varying number
of individuals in samples on the number of taxa
recorded, we included a finite, individual-based
rarefied richness (performed without replacement on
44 individuals; the lowest number in the five pooled
Surber samples of any site). These were calculated
using the software ‘Species diversity and richness’
version 3.02, Pisces Conservation Ltd, Hants, UK.
One-way A N O V A s were performed to analyse for
differences in faunal density and taxon richness
between the three sites within groups, using individ-
ual Surber samples. A N O V A s and Pearson’s product
moment correlation analyses between log (x + 1)-
transformed environmental variables and faunal met-
rics were performed in Excel.
Non-metric multidimensional scaling (NMDS) based
on Bray–Curtis similarities calculated on square-root-
transformed assemblage data was used to examine
spatial patterns in macroinvertebrate assemblage com-
position among sites. This ordination technique has no
assumptions with repect to distributions and repre-
sents samples as points in low-dimensional space, such
that the relative distances among points reflect the
relative similarities of the samples with respect to both
abundance and composition (Gucht, Vandekerckhove
& Vloemans, 2005). The NMDS goodness of fit was
estimated with a stress function (which ranges from 0 to
1), with values close to zero indicating a good fit. The
Bray–Curtis index was also used to illustrate the
similarities between faunal assemblages at different
sites: 0% indicating no species in common and 100%
indicating complete species match occurring in the
same abundances. These procedures were performed
using Primer 5 Ltd. version 5.2.4, 2001, Plymouth, UK.
Differences in the composition of the macroinver-
tebrate assemblages among site groups were further
tested using an analysis of similarities (ANOSIM). The
ANOSIM tested the null hypothesis that within-group
similarity was equal to between-group similarity.
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
ANOSIM generates a statistical parameter R that is
indicative of the degree of separation between groups:
a score of 1 indicates complete separation and a score
of 0 indicates no separation (Gucht et al., 2005).
Monte-Carlo randomisation of the group labels was
used to generate null distributions to test the hypoth-
esis that within-group similarities are higher than
would be expected by chance. These analyses were
performed using PAST (Paleontological statistics,
version 1.79, Oslo).
Generalised linear model analysis (GLM) was used
to build Poisson log-linear models to test the effect of
mean temperature, stability, benthic chlorophyll a and
CPOM on the faunal metrics density, number of taxa,
rarefied richness and Pielou’s evenness. We focussed
on the effect of temperature and stability as explan-
atory variables because they are the two basic factors
in the conceptual model by Milner et al. (2001a,b) and
because an earlier study along stream G12 confirmed
these to be the factors primarily related to the
distribution of the fauna (Jacobsen et al., 2010). We
also included a ‘stream’ variable to allow for inter-
stream comparisons of the effect of these variables.
Stream was treated as a fixed effect, but we obtained
similar results by means of the generalised linear
mixed model function (glmmPQL; MASS library for
R), with stream as a random effect. Change in density,
richness and evenness owing to each factor was
modelled considering each factor independently and
in combination with other factors, including biologi-
cally reasonable two-way interactions, and squared
variables. Indeed, the relationships between density
and taxon richness, and environmental variables are
often curvilinear (Austin, 1980). The more parsimoni-
ous model was identified using the corrected Akaike’s
Information Criterion (AICc), which is recommended
for small sample sizes (AIC, see Venables & Ripley,
2002) in likelihood ratio tests to find the difference
between the initial model and the reduced model,
dropping an ‘effect’ term. All analyses were per-
formed on log-transformed data using the MASS
library for R (R Development Core Team, 2008).
Results
Environmental variables
The nine stream sites could all be characterised as
small (width 0.39–2.04 m; depth 0.03–0.19 m) with
 Spatial variability in equatorial glacier-fed streams 7
Table 1 Environmental characteristics measured at the nine sites along the Antisana streams G15, G14 and G12

Site Variable G15.1 G15.2 G15.3 G14.1 G14.2 G14.3 G12.1 G12.2 G12.3

Altitude (m.a.s.l.) 4835 4521 4335 4789 4535 4196 4728 4496 4225
Catchment (km2) 0.46 1.02 1.47 0.65 1.12 14.27 2.13 3.21 4.33
Glacier cover in catchm. (%) 94 45 31 92 67 10 100 66 49
Distance from glacier (m) 100 1500 3500 200 1500 5600 50 1500 4300
Min. turbidity (NTU) 812 211 65 181 111 58 285 442 411
Max. pH 7.98 7.07 6.44 6.31 7.04 7.96 7.47 6.36 6.39
Max. Conduc. (lS cm)1) 3 8 24 11 10 47 5 10 14
Mean temp. (ºC) 1.56 4.20 5.05 2.48 4.31 7.42 1.59 2.86 5.38
CV temp. (%) 111 102 63 126 80 34 180 46 51
Mean daily max. temp. (ºC) 4.38 12.35 10.65 9.47 10.70 11.26 5.44 5.21 10.94
Mean daily min. temp. (ºC) 0.47 0.30 1.56 0.00 0.60 4.32 0.00 1.36 2.62
Mean depth (cm) 5 6 10 6 10 18 3 13 19
CV depth (%) 52 84 69 18 20 30 110 35 27
Substrate types (nr) 6 8 6 5 6 8 5 6 10
Substrate type div. (H¢) 1.63 1.88 1.52 1.60 1.70 1.95 1.05 1.60 2.11
Mean current (m s)1) 0.24 0.11 0.13 0.25 0.24 0.19 0.22 0.38 0.21
Mean width (cm) 67 59 63 72 133 133 39 204 158
Slope (cm m)1) 11.7 9.7 7.3 24.0 10.8 2.2 1.4 8.7 9.0
Stability index 9 18 25 15 21 31 16 24 21
Chlorophyll (mg m)2) 0.14 3.92 2.63 0.55 6.17 1.85 0.27 3.15 0.99
CPOM (g AFDM m)2) 7.98 13.26 145.50 8.74 21.18 22.04 8.68 43.55 33.26

AFDM, ash-free dry mass; CPOM, coarse particulate organic matter; CV, coefficient of variation.

medium flow velocity (0.11–0.38 m s)1), diverse sub-
stratum composed mainly of gravel, pebble and
cobbles, high turbidity (58–812 NTU), low conductiv-
ity (3–47 lS s)1) and circum-neutral pH (6.31–7.96)
(Table 1). Other variables differed more among
sites, e.g. slope (1.4–24.0%), benthic chlorophyll a
(0.14–6.17 mg m)2) and benthic CPOM (8–145 g
AFDM m)2).
The geographical variables such as altitude, glacier
% of catchment and distance from the glacier were all
highly correlated (Appendix 1). All environmental
variables, except pH, CV depth, mean current, mean
width and slope, were significantly correlated with
one or more of these geographical variables. In
general, many of the environmental variables were
intercorrelated (Appendix 1).
All temperature parameters increased downstream
from the glacier (temperature variation decreased),
but mean daily maximum temperature showed an
irregular longitudinal pattern. Overall, the proposed
stability index increased with the distance from the
glaciers, even though G12.3, the lowermost site,
scored the lowest stability index. Benthic algal bio-
mass (chlorophyll a) was quite similar in the three
streams and peaked at middle sites in all three
streams. There was an overall downstream increasing
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
tendency in CPOM pools, but with large local devi-
ations, and even at the upper sites close to the glaciers,
biomasses of c. 10 g AFDM m)2 were found.
General longitudinal patterns and driving factors
In the 45 Surber samples from the nine sites, we
collected a total of 2200 individuals, 64% of them
chironomids. A total of 47 taxa, of which 30 were
dipterans, 18 of these Chironomidae, were identified
(Appendix 2). Density varied between 176 and 3044
ind. m)2. The number of taxa ranged from 2 to 6 at the
upper sites and from 21 to 27 at the lower sites.
Chironomidae was the most abundant group at all
sites, except at site G15.2 that had a high abundance of
Simulium. In particular, Podonominae was completely
dominant at the upper sites (>98% of ind.). Single
individuals of a few other groups were found at the
upper sites (Muscidae and Staphylinidae). In all three
streams, Podonominae became less dominant and
Orthocladiinae more abundant with increasing
distance from the glaciers. The two genera Atopsyche
and Cailloma (Hydrobiosidae and Trichoptera) were
found in all three streams at middle and lower sites,
while the ephemeropteran Andesiops (Baetidae) did
not appear until the lowest sites. The plecopteran
8 J. Kuhn et al.
Claudioperla (Gripopterygidae) was rare and appeared G15 G14 G12
in low numbers only at site G14.3. Elmidae (Coleop-
(a) 3500
tera) were common at two of the three lower sites.
3000
Overall, with increasing distance from the glacier,

Density (ind. m–2)

there was a logarithmic increase in the density of 2500
individuals (y = 86.697Ln(x) ) 341.88; r2 = 0.46; P = 2000
0.0085), number of taxa (y = 4.6863Ln(x) ) 16.939; 1500
r2 = 0.85; P = 0.0010) and rarefied richness (y =
1000
2.0877Ln(x) ) 5.9076; r2 = 0.87; P = 0.0005) (Fig. 2),
500
and these three faunal metrics were equally related
to other geographical factors such as altitude and 0
0 1000 2000 3000 4000 5000 6000
glacier per cent of catchment (Table 2). They were also
significantly correlated with maximum conductivity, (b) 30
mean temperature, mean maximum temperature and
25
the stability index (P < 0.05), but not with chlorophyll

Number of taxa
a, and only rarefied richness with CPOM. In addition, 20
the number of taxa and rarefied richness were 15
significantly correlated with CV temp., mean depth
10
and the number of substrate types. Pielou’s evenness
did not correlate with any of the environmental 5
variables (Table 2). 0
The NMDS ordination had a low stress (0.02), 0 1000 2000 3000 4000 5000 6000
indicating a good degree of representation (Fig. 3).
(c) 14
The diagram revealed a clear longitudinal pattern in
the composition of the macroinvertebrate assem- 12
Rarefied richness

blages; correlation coefficients between NMDS axis-1 10

coordinates and log (x + 1)-transformed distance from 8
the glacier (0.94), altitude ()0.91) and per cent of 6
glacier cover in catchment ()0.69) were all significant
4
(P < 0.05). All other environmental variables that
were significantly correlated with distance from the 2

glacier (Appendix 2) were also correlated with NMDS 0

0 1000 2000 3000 4000 5000 6000
axis-1 coordinates, except substrate type diversity. In
general, assemblage composition was largely related (d) 1
to the same environmental variables as the univariate
fauna metrics. 0.8
Pielou evenness

Mean similarity was lower comparing upper and

0.6
middle sites (1 versus 2) than comparing middle and
lower sites (2 versus 3) (Fig. 4). Divided by distance 0.4
between sites, the difference in distance decay in
similarity is even more pronounced, 47% km)1 from 0.2
upper to middle sites and 20% km)1 from middle to
0
lower sites. 0 1000 2000 3000 4000 5000 6000
Distance from glacier (m)
Among-stream faunal variability and stream-specific
Fig. 2 Density of individuals (a), number of taxa (b), rarefied
effects richness (c) and Pielou’s evenness (d) as a function of distance
from the glacier, showing the three study sites along each of the
Somewhat differing longitudinal patterns in faunal three Antisana streams.
density, number of taxa and rarefied richness were
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
 Spatial variability in equatorial glacier-fed streams 9

0.17 )0.05 0.03 )0.23 )0.32 )0.33 )0.01 )0.17 )0.05 )0.07 1.00
Pielou’s evenness
revealed in the three streams (Fig. 2). Following
Rarefied richness
logarithmic relationships between faunal metrics and

0.73* 0.89** 0.97** 1.00

distance from glacier, the variability among streams
Number of taxa (residuals) can be converted into distance. For the

0.94** 1.00
three middle sites (all situated at 1500 m from the
Density ðind: m2 Þ glaciers), this variability among streams equalled

1.00
2300 m (200–2500 m) for density, 1700 m (400–
CPOM ðg AFDM m2 Þ 2100 m) for taxon richness, 800 m (700–1500 m) for

0.38
0.63
rarefied richness and 1800 m (800–2600 m) for assem-
Chlorophyll a ðmg m2 Þ blage composition expressed as axis-1 coordinates
)0.24 0.62 0.58 )0.43 0.39 )0.30 0.81** 0.63
)0.12 0.68* 0.59 )0.50 0.38 )0.26 0.84** 0.62

0.81** 0.80** )0.15 0.67* 0.55 )0.37 0.47 )0.28 0.89** 0.57
from the NMDS ordination (Fig. 5). A N O V A s con-
Stability index
firmed that density varied significantly among the
Table 2 Pearson’s correlation coefficients between fauna metrics and environmental variables. Bonferroni correction of P not applied

sites within the middle and lower groups (P < 0.003),

Slope ðcm m1 Þ
but not among the group of upper sites (P = 0.240).
Mean width (cm) The number of taxa differed significantly among sites
within the upper and middle groups (P < 0.018), but
Mean current ðm s1 Þ
not among the lower sites (P = 0.059).
Substrate type div. (H’)
Overall, the three streams did not differ in assem-
blage composition (R = )0.366, P = 0.996, ANOSIM),
but the three site groups delimited on Fig. 3 (grouped
0.33

Substrate types
according to their location as upper, middle or lower
0.27

CV depth (%)
sites) were significantly different (R = 0.951,
P = 0.003, ANOSIM). Further, ANOSIM analyses
0.72*

Mean depth (cm)

0.65

0.08

performed separately on the 15 individual Surber

AFDM, ash-free dry mass; CPOM, coarse particulate organic matter; CV, coefficient of variation.

samples from each of the three site groups showed

)0.86** 0.92** )0.71* )0.09 0.84** 0.98** )0.72* 0.80** 0.75*

Mean daily min. temp. (C)

)0.83** 0.83** )0.74* 0.04 0.81** 0.94** )0.64 0.83** 0.66

)0.41 0.36

that the three upper sites did not differ significantly

(R = 0.187, P = 0.081), while the middle sites
)0.22 0.86** 0.95** )0.79* 0.73*

Mean daily max. temp. (C)

(R = 0.663, P < 0.0001) and lower sites (R = 0.639,
P < 0.0001) did. This was confirmed by the mean
)0.20 )0.08 )0.19

CV temp. (%)
similarity between replicate sites which decreased
Mean temp. (C) slightly with increasing distance from the glaciers
(Fig. 4).
Max. conductivity ðlS cm1 Þ Relationships between the four univariate faunal
metrics, the NMDS axis-1 coordinates and the focal
environmental variables mean temperature, the sta-
0.60

Max. pH
bility index, chlorophyll a. and CPOM also varied
)0.98** 0.68* )0.82** 0.94** )0.64

Min. turbidity (NTU) among the three streams (Fig. 6). Recognising that
)0.03 0.40

these fauna-environment patterns varied among

Distance from glacier (m) streams we performed GLM analyses to test for
stream-specific effects and interaction effects between
streams and environmental variables (Table 3). Apart
)0.13

Glacier % of catchment (%)

from the relationships obtained in the simple corre-
lation analyses (Table 2), the GLM analyses revealed a
)0.83** 0.65
)0.93** 0.62

0.13

Catchment ðkm2 Þ

few new patterns. Temperature interacted with

*P < 0.05; **P < 0.01.

stream on density, while stability interacted with

)0.08

Altitude (m.a.s.l.)

stream on evenness. This indicates that the effects of

evenness

temperature and stability were quantitatively differ-

richness
Number

Rarefied
of taxa

Pielou’s
Density

ent in the three streams. Chlorophyll a had a signif-

icant effect on the number of taxa and evenness in
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
10 J. Kuhn et al.
Stress: 0.02
1.0 14.3

0.8

0.6
12.3
0.4
15.1 14.1
0.2
Fig. 3 Bray–Curtis similarity-based Non-
0 metric multidimensional scaling ordina-
15.3 tion showing square-root-transformed
–0.2 12.1 faunal data from the three Antisana
streams. Results are based upon sum-
–0.4
marised data from five Surber samples
15.2
from each site. Site groups 1, 2 and 3 are
–0.6 14.2
12.2 marked. Stress value is shown. Environ-
–0.8 mental variables that show significant
Pearson’s correlations with the x-coordi-
–1.6 –1.4 –1.2 –1.0 –0.8 –0.6 –0.4 –0.2 0 0.2 0.4 0.6 0.8 1.0 1.2 nates in the ordinations diagram are
Distance to glacier Mean temperature Stability index included.

70

60
Mean similarity (%)

50

40

30

20

10

0 1800 m
1 versus 1 2 versus 2 3 versus 3 1 versus 2 2 versus 3 1 versus 3
Sites

Fig. 4 Mean Bray–Curtis similarity (square-root-transformed

faunal data) between upper (1), middle (2) and lower (3) sites
along and between the three Antisana glacial streams. All
columns n = 3 and error bars denote max-min values.
interaction with stream, and with respect to the effect
on evenness, chlorophyll a interacted with stability.
Hence, the effect of chlorophyll a differed among
Fig. 5 A logarithmic fit to the relationship between axis-1
coordinates from a MDS ordination of the nine Antisana stream
sites as a function of their longitudinal distance from the glacier
(y = 0.4865Ln(x) ) 3.293; r2 = 0.89; P = 0.0002). The diagram
illustrates that following the general relationship, the variation
among the three replicate sites located at 1500 m from the
glaciers corresponds to a longitudinal distance of 1800 m
(800–2600 m).

streams and depended on stability levels.

is Diamesa, while it is Maoridiamesa in New Zealand,

Discussion where Diamesa does not occur (Milner et al., 2001b).
We did not find Diamesa and, according to Sæther &
The fauna of Neotropical glacier-fed streams
Spies (2004), the genus does not occur in the Neo-
The very low numbers of Diamesinae, and their tropical region either. The upper Antisana sites were
complete absence at the three stream sites closest to instead dominated by Podonominae. The Podonom-
the glacier, differed from the longitudinal distribution inae are a feature of Gondwana land masses and are
of faunal groups typically observed in north-temper- common in high Andean streams (Brundin, 1966;
ate glacier-fed streams (Milner et al., 2001a). The Jacobsen, 2008). The few sporadic specimens of other
typical meta-kryal Diamesinae in the holarctic zone taxa (Muscidae and Staphylinidae) at the highest sites
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
 Spatial variability in equatorial glacier-fed streams 11
G15 G14 G12

3500 3500 3500

3500
Density (ind. m–2)

3000 3000 3000 3000

2500 2500 2500 2500

2000 2000 2000 2000

1500 1500 1500 1500

1000 1000 1000 1000

500
(a) 500
(b) 500
(c) 500
(d)
0 0 0 0
0 1 2 3 4 5 6 7 8 0 5 10 15 20 25 30 35 0 1 2 3 4 5 6 7 0 20 40 60 80 100 120 140

30 30
30 30
25 25
Number of taxa

25 25

20 20 20 20

15 15 15 15

10 10 10 10

5 5 5 5
(e) (f) (g) (h)
0 0 0 0
0 1 2 3 4 5 6 7 8 0 5 10 15 20 25 30 35 0 1 2 3 4 5 6 7 0 20 40 60 80 100 120 140

14 14 14 14
Rarefied richness

12 12 12 12

10 10 10 10

8 8 8 8

6 6 6 6

4 4 4 4
2 (i) 2 (j) 2 (k) 2 (l)
0 0 0 0
0 1 2 3 4 5 6 7 8 0 5 10 15 20 25 30 35 0 1 2 3 4 5 6 7 0 20 40 60 80 100 120 140

1 1 1 1
0.9 0.9
Pielou evenness

0.8 0.8 0.8 0.8

0.7 0.7
0.6 0.6 0.6 0.6
0.5 0.5
0.4 0.4 0.4 0.4
0.3 0.3
0.2 0.2 0.2 0.2
(m) (n) 0.1 (o) 0.1 (p)
0 0 0 0
0 1 2 3 4 5 6 7 8 0 5 10 15 20 25 30 35 0 1 2 3 4 5 6 7 0 20 40 60 80 100 120 140

1.5 1.5 1.5 1.5

MDS axis-1 coord.

1 1 1 1

0.5 0.5 0.5 0.5

0 0 0 0

–0.5 –0.5 –0.5 –0.5

–1
–1
(q) (r) –1
(s) –1
(t)
–1.5 –1.5 –1.5 –1.5
0 1 2 3 4 5 6 7 8 0 5 10 15 20 25 30 35 0 1 2 3 4 5 6 7 0 20 40 60 80 100 120 140

Mean temp. (ºC) Stability index Chlorophyll a (mg m–2) CPOM (g AFDM m–2)

Fig. 6 Density of individuals (a,b,c,d), number of taxa (e,f,g,h), rarefied richness (i,j,k,l), Pielou’s evenness (m,n,o,p) and Non-metric
multidimensional scaling x-coordinates (q,r,s,t) of macroinvertebrate assemblages as a function of mean temperature, stability
index, chlorophyll a and coarse particulate organic matter in each of the three Antisana glacial streams.

were probably due to sporadic immigrant individuals Patagonian streams (Miserendino & Pizzolon, 2000).
from lower sites, rather than representing perma- Even though the family is relatively common in the
nently reproducing populations. high Andes, it has not been found above 3900 m a.s.l.
Other Diamesinae genera (probably Paraheptagyia) in Ecuador (Jacobsen, 2004). The model of Milner et al.
appeared lower down the streams, but they never (2001a) further predicts that chloroperlid and
became numerically important. In New Zealand, the nemourid stoneflies should appear as soon as Tmax
Diamesinae were largely replaced by other taxa such exceeds 4 C. These two families do not occur in the
as the ephemeropteran Deleatidium (Leptophlebiidae), Ecuadorian highlands, where Gripopterygidae
at the sites nearest to the glacier, even though these (Claudioperla) is the only family of stoneflies and was
sites were the coldest (Milner et al., 2001b; Winterbo- found at the lowest sites. This Austral family is also
urn et al., 2008; Cadbury et al., 2010). Leptophlebiidae common in the streams of glacial origin in Bolivia
have been collected in a glacial stream at 4400 m a.s.l. (Molina, 2004), Patagonia (Weis & Bonetto, 1988) and
in Bolivia (Molina, 2004) and are also common in cold New Zealand (Milner et al., 2001b). Hydrobiosidae
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
12 J. Kuhn et al.
Table 3 Results of the generalized linear model deviance anal-
ysis on on the fauna metrics density, number of taxa, rarefied
richness and Pielou’s evenness
Effect AICc D AICc LRT P-value
Density
Stream · Temperature 147.2 6.1 4.321 <0.001
Number of taxa
Stream · Chlorophyll a 56.9 2.2 3.631 0.023
Rarefied richness
None
Pielou evenness
Stream · Stability 74.3 2.3 3.865 0.013
Stream · Stability 79.7 7.7 4.633 0.002
· Chlorophyll a
Only significant effects are shown. AICc is the corrected
Akaike’s Information Criterion for the initial model after
removal of the ‘effect’ term. DAICc corresponds to the difference
between the AICc of the initial model and that of the reduced
model. Likelihood ratio test (LRT) and associated P-value test
the hypothesis that the suppression of the ‘effect’ term provides
no better fit than the initial model.
(Trichoptera) were found in all three streams, and the
family is abundant and widespread in high Andean
(Jacobsen, 2004, 2008) as well as in New Zealand
alpine streams (Milner et al., 2001b; Cadbury et al.,
2010). This family of predatory free-living larvae
seems to occupy the equivalent ecological niche to
Rhyacophilidae in European glacial streams (Castella
et al., 2001). Coleoptera are usually not present in
alpine streams at temperate latitudes, and even
Elmidae are rarely abundant (Ward, 1994). Elmidae
(Coleoptera) were found in two of the three lower
sites on Antisana, and this family is very common in
high Andean streams (Jacobsen, 2008). Thus, qualita-
tive departures from the Milner et al. (2001a) model
were mainly attributable to zoogeographical differ-
ences between the Neotropical and the north-temper-
ate (and New Zealand) stream fauna.
General longitudinal patterns and driving factors
Overall, our data support the general conceptual
model of logarithmic increase in density and richness
of fauna with increasing distance from a glacier
(Milner et al., 2001a) and confirm the longitudinal
patterns reported earlier by Jacobsen et al. (2010) from
one of the streams. The greater distance decay in
similarities between sites close to the glaciers than
further downstream was in accordance with our
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
expectations and further supported this logarithmic
development of macroinvertebrate communities
along glacier-fed streams. These relationships were
basically the same with other geographical variables
such as altitude and per cent of glacier cover in the
catchment.
Temperature and stability were the two main
variables structuring faunal patterns (Table 2, Fig. 3).
Temperature parameters and the stability index were
the only environmental variables that correlated
significantly with the density of individuals, the
number of taxa and geographical variables. Like
Jacobsen et al. (2010), we found that mean daily
maximum temperature showed an irregular longitu-
dinal pattern, and no truly kryal zone (Tmax < 4 C
sensu Steffan, 1971; Ward, 1994; Milner et al., 2001a)
was identified, as even mean daily maximum tem-
perature exceeded 4 C at all localities. This could be
explained by the high insolation at the equator,
heating up the small amount of melt water in the
morning faster than the discharge of cold melt water
increases (Jacobsen et al., 2010). High water tempera-
ture in proglacial reaches has also been reported from
streams with low discharge from small glaciers in the
European Alps (Maiolini & Lencioni, 2001). In con-
trast, mean temperature increased systematically with
increased distance from the glaciers, and this param-
eter was also more closely related to the faunal
metrics than was maximum temperature. This sug-
gests that mean temperature might be an ecologically
more meaningful parameter in setting physiological
limits for the distribution of species.
The proposed stability index correlated significantly
with density, the number of taxa and rarefied
richness. Following the intermediate disturbance
hypothesis, both taxon richness (Townsend, Scars-
brook & Dolédec, 1997) and assemblage evenness
(Death & Winterbourn, 1995) are expected to peak at
intermediate levels of disturbance (or stability)
because a certain level of disturbance should impede
dominance from competitively superior species while
allowing the existence of species able to cope with
such environmental conditions in a space with low
interspecific competition (McAuliffe, 1984). Positive
rather than peaked relationships between faunal
metrics and stability are generally found in glacier-
fed streams, probably because glacial streams only
cover the lower end of a stability gradient. Surpris-
ingly, we found no relationship between stability and

evenness. Further studies are needed to evaluate the
wider usefulness of our proposed stability index, but
here it seemed to work at least as well as the
commonly used, but nevertheless subjective, Pfan-
kuch index.
Benthic algal biomass (chlorophyll a) varies greatly
within and among glacial streams; in some cases,
biomass increases steadily with increasing distance
from the glacier (Lods-Crozet et al., 2001), but this is
not always so (Gı́slason et al., 2001; Cadbury et al.,
2010). In our three streams, algal biomasses were quite
similar and peaked at middle sites in all three streams,
a result confirming the findings of Jacobsen et al.
(2010). This was probably due to high insolation at
these sites compared with the generally more incised
and therefore more shaded lower sites. No overall
relationship between chlorophyll a and faunal metrics
was found.
Our levels of benthic CPOM were very high
compared with other studies (Lods-Crozet et al.,
2001; Cadbury et al., 2010). Even at the upper sites
close to the glaciers, we found biomasses of c. 10 g
AFDM m)2, confirming earlier results reported by
Jacobsen et al. (2010). CPOM seemed to be extremely
patchy, and at sites with high values, the CPOM was
primarily composed of moss and filamentous algae
detached from stones during the Surber sampling.
Thus, general patterns and correlations between the
fauna and CPOM levels are difficult to demonstrate.
Among-stream faunal variability and stream-specific
effects
In our study, spatial location and environmental
features were obviously very strongly related, so we
could not test for the importance of niche processes
versus dispersal limitation as drivers of local diversity
and community patterns (Thompson & Townsend,
2006; Heino & Mykrä, 2008), neither was this our goal.
Nevertheless, as mean similarity among replicate sites
decreased from upper to lower sites, following a
supposed decrease in environmental harshness along
the streams, this seems to support an expected
stronger niche selection close to the glaciers than
further downstream (Leibold et al., 2004; Chase, 2007).
Even though our three streams appeared very similar,
we found considerable variation in faunal metrics
among replicate sites (Fig. 5), and the quantitative
response of fauna metrics to environmental variables
 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x
Spatial variability in equatorial glacier-fed streams 13
such as temperature and stability also varied among
streams (Table 3, Fig. 6). If we discard the possibility
that at some sites variables were not correctly
measured, we may conclude that their regulating
effect is stream specific and depends on other,
perhaps unknown, factors. The three streams differed
somewhat with respect to exposure, insulation and
morphometry, and similar hydrological events could
therefore cause different effects in the three streams,
perhaps implying unsynchronised temporal variabil-
ity in macroinvertebrate assemblages. We suspect that
these circumstances are responsible for the varying
relationships between fauna and environmental fac-
tors such as temperature and stability among the
streams.
Overall, chlorophyll a did not relate significantly to
any of the faunal metrics, but it did show a stream-
specific effect on the number of taxa and evenness in
the GLM analyses. Chlorophyll a as such is not a very
precise measure of either quality or quantity of
benthic algae as food for macroinvertebrates, as the
taxonomic composition of the periphyton community
may vary greatly among streams (Biggs, 1996; Rott
et al., 2006). For instance, both filamentous blue-green
algae and diatoms possess chlorophyll a, but their
value as a food source for macroinvertebrates is very
different. Nor was it surprising that chlorophyll a
interacted with stability on evenness, because the
biomass and type of algae in streams is closely
controlled by disturbance regime (Biggs, Goring &
Nikora, 1998; Hieber et al., 2001). In addition, benthic
algal biomass is temporally variable (Cadbury et al.,
2010).
Local assemblages were thus structured by the
complex interaction of the many variables constituting
the entire environmental setting. Our results under-
line that assemblages distributed along spatial gradi-
ents such as glacial streams with multiple
environmental factors varying simultaneously are
not regulated independently by each environmental
factor in a similar and straightforward way. This
could explain why different studies on glacier-fed
streams have reached different conclusions on the
importance of the same regulating factors, such as
chlorophyll a having a greater effect than CPOM on
macroinvertebrate assemblages in the Swiss Mutt
stream (Lods-Crozet et al., 2001), while the opposite
was true in the New Zealand Rob Roy stream
(Cadbury et al., 2010).
14 J. Kuhn et al.
The current rapid glacial retreat (Francou & Cou-
drain, 2005; Bradley et al., 2006; Ceballos et al., 2006)
is, in the near future, expected to affect natural
communities and biodiversity in glacial streams
(Brown, Hannah & Milner, 2007; Milner, Brown &
Hannah, 2009). Quantifying spatial (and temporal)
variation in glacier-fed stream communities and
relationships with environmental factors will help us
interpret future changes in macroinvertebrate assem-
blages as sentinels of glacial retreat.
Acknowledgments
The funding by a WWF-Novozymes grant 2008 to DJ
and an Ecofondo grant nr. 034-ECO8-inv1 to OD is
greatly appreciated. We thank Narcis Prat for help
with the Chironomid identifications and José Delgado
from the Hacienda Antisana for access to the study
area.
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 Appendix 1
Pearson’s correlation coefficients between log (x + 1) transformed environmental variables.

Max. pH
CV temp. (%)
CV depth (%)
Stability index

Slope ðcm m1 Þ

Substrate types

Mean temp. (C)

Mean depth (cm)
Mean width (cm)

Catchment ðkm2 Þ

Altitude (m.a.s.l.)
Mean current ðm s1 Þ

Min. turbidity (NTU)

CPOM ðg AFDM m2 Þ

Chlorophyll a ðmg m2 Þ

Substrate type div: ðH0 Þ

Distance from glacier (m)

Mean daily min. temp. (C)

Max: conductivity ðScm1 Þ

Mean daily max. temp. (C)

Glacier % of catchment (%)

Altitude (m.a.s.l.) 1.00
Catchment (km2) )0.77 1.00
Glacier % of 0.83 )0.77 1.00
catchment (%)
Distance from )0.92 0.57 )0.77 1.00
glacier (m)
Min. turbidity 0.51 )0.36 0.70 )0.52 1.00
(NTU)
Max. pH 0.16 0.19 )0.22 )0.29 0.01 1.00
Max. conductivity )0.83 0.73 )0.89 0.80 )0.80 )0.15 1.00
(lS cm)1)
Mean temp. (ºC) )0.92 0.63 )0.86 0.95 )0.69 )0.13 0.88 1.00
CV temp. (%) 0.85 )0.74 0.76 )0.88 0.33 0.13 )0.76 )0.78 1.00
Mean daily )0.62 0.24 )0.58 0.70 )0.71 )0.26 0.67 0.83 )0.34 1.00
max. temp. (ºC)

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Mean daily )0.90 0.83 )0.82 0.82 )0.34 0.04 0.77 0.79 )0.94 0.34 1.00
min. temp. (ºC)
Mean depth (cm) )0.85 0.67 )0.65 0.89 )0.31 )0.29 0.74 0.81 )0.95 0.45 0.90 1.00
CV depth (%) 0.16 )0.16 0.07 )0.31 0.12 0.31 )0.32 )0.31 0.41 )0.28 )0.27 )0.58 1.00
Substrate types )0.75 0.53 )0.56 0.71 )0.02 0.01 0.40 0.70 )0.61 0.53 0.69 0.65 )0.12 1.00
Substrate type )0.60 0.34 )0.52 0.73 )0.10 )0.10 0.44 0.70 )0.66 0.59 0.61 0.73 )0.52 0.83 1.00
div. (H¢)
Mean current (m s)1) 0.28 0.05 0.42 )0.25 0.48 )0.19 )0.26 )0.42 )0.12 )0.64 )0.05 0.13 )0.45 )0.35 )0.18 1.00
Mean width (cm) )0.55 0.51 )0.31 0.65 0.01 )0.29 0.41 0.50 )0.82 0.14 0.66 0.87 )0.70 0.46 0.63 0.54 1.00
Slope (cm m)1) 0.32 )0.63 0.41 )0.01 0.25 )0.53 )0.24 )0.13 0.11 0.12 )0.31 0.03 )0.52 )0.08 0.36 0.16 0.18 1.00
Stability index )0.85 0.74 )0.75 0.82 )0.70 )0.27 0.87 0.82 )0.76 0.57 0.70 0.73 )0.19 0.37 0.30 )0.09 0.52 )0.37 1.00
Chlorophyll a (mg m)2) )0.44 0.09 )0.35 0.67 )0.49 )0.26 0.37 0.59 )0.45 0.53 0.24 0.43 )0.17 0.23 0.35 )0.12 0.44 0.11 0.62 1.00
CPOM (g AFDM m)2) )0.69 0.27 )0.46 0.72 )0.43 )0.50 0.59 0.58 )0.65 0.32 0.59 0.62 0.00 0.23 0.19 )0.12 0.37 )0.04 0.68 0.50 1.00

Correlations with P < 0.05 in bold.

Spatial variability in equatorial glacier-fed streams 17
 18
J. Kuhn et al.

Appendix 2
Complete taxon list for the five Surber samples from each of three sites in three glacial streams on Mt Antisana in Ecuador.

Stream 15 14 12

Site 1 2 3 1 2 3 1 2 3

Surber nr. 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5

Family Taxa

Nematoda – sp1 1 1
– sp2 2 3 8 2 1 3 1 2 9 15 3
– sp3 2 2
Oligochaeta Naididae sp1 9 2 20 3 5 4 1 3 3 8 3 3 7 10 9 7 20 5 2 3 5 3 1
Coleoptera Elmidae Neoelmis sp1 3 2 10 3 1
Elmidae Neoelmis sp2 1 5 11 2 10 2 1 2 2
Scirtidae Prionocyphon sp1 1 1 2 2
Dytiscidae sp1 1
Staphylinidae sp1 1 1
Staphylinidae sp2 1
Ephemeroptera Baetidae Andesiops sp 1 3 10 1 26 16 2 18 10 3 12 15 4 14
Plecoptera Gripopterygidae Claudioperla sp 1 1
Trichoptera Hydroptilidae Ochrotrichia sp 1 10 3 3 2 3 1
Hydrobiosidae Atopsyche sp1 1 1 4 1 3 2 2
Hydrobiosidae Cailloma sp1 1 1 1 1 1 2 1 1 1 5 4 1 1 2

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 Appendix 2
(Continued)

Stream 15 14 12

Site 1 2 3 1 2 3 1 2 3

Surber nr. 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5

Family Taxa

Diptera Empididae Hemerodromia 1 1 1 1

sp
Empididae Chelifera sp 1 1 1 1 1 1 1 1 1 5 3
Empididae sp3 3
Muscidae Limnophora sp1 1 1 1 3 5 7 1 2
Muscidae sp4 1 1
Simuliidae Simulium sp1 23 20 45 9 27 3 4 2 3 3 2 7 3 7 2 1 1 2 1 8 5 2 6
Tipulidae Tipula sp 1
Limoniidae Geranomyia sp 1
Limoniidae indet. 1 1
Limoniidae Molophilus sp1 1
Ceratopogonidae Alluaudomyia sp1 1 3 3 10 12 16 16 2 1 6 2 2
Blephariceridae Blepharicera sp1 1
Chironomidae Parochlus sp. A 5 5 1 5 9 2 6 2 5 12 2 4 2 2 5 2 2 2 23 22 75 14 43 7 8 12 9 1 29 8 1 8 3 9 9 10 14 4 1 1 1
Chironomidae Parochlus sp. B 27 1 1 1 2 1 16 22 18 15 1 1 4 2 9 3 11 3 1 1 4 1 1 1 1
Chironomidae Podonomus sp. 1 2 3 1 1 1 2 11 3
Chironomidae Tanypodinae indet. 1
Chironomidae Paraheptagyia sp. A 1 1 1 1
Chironomidae Paraheptagyia sp. B 1
Chironomidae Corynoneura lobata gr. 4 5
Chironomidae Cricotopus sp. A 6 13 6 24 6 12 11 20 35 20 2 2 2 7 4 12 5 7 14 1 10 2 8 9 18 2 6 1 8 1
Chironomidae Cricotopus sp. B 1 1 4 3 1 5 1 4 4 31 39 40 71 49 1 4 53 16 1 14

 2011 Blackwell Publishing Ltd, Freshwater Biology, doi:10.1111/j.1365-2427.2011.02648.x

Chironomidae Cricitopus sp. C 4 2 1 11 3 5 14 7 2 5 1 1 2 1 2 3
Chironomidae Cricotopus sp. D 1 28 17 43 19 19 1 1
Chironomidae Limnophyies sp. 1 1 1 1
Chironomidae Metriocnemus fuscipes gr. 4 3 9
Chironomidae Parakiefferiella sp. 1 2
Chironomidae Pseudosmittia sp. 1
Chironomidae Polypedilum sp. 1 2
Chironomidae Tanytarsus sp. 1
Chironomidae Chironomini indet. 1
Hirudinea – sp 1 8 3 10 1
– sp 3 1 1 2

(Manuscript accepted 3 June 2011)

Spatial variability in equatorial glacier-fed streams 19