Freshwater Biology (2010) 55, 20622076

doi:10.1111/j.1365-2427.2010.02462.x

Low river flow alters the biomass and population

structure of a riparian predatory invertebrate
MICHELLE J. GREENWOOD*, AND ANGUS R. MCINTOSH*
*School of Biological Sciences, University of Canterbury, Christchurch, New Zealand

National Institute for Water and Atmospheric Research, Ltd, Christchurch, New Zealand

SU M M A R Y
1. Low flows in rivers are predicted to increase in extent and severity in many areas in the
future, yet the consequent impacts of river drying on terrestrial communities via (i)
changes to riparian microclimatic conditions and (ii) the identity and abundance of
emerging aquatic insects available to riparian predators have not been quantified.
2. We investigated the influence of low river flow on a riparian fishing spider, Dolomedes
aquaticus, in five New Zealand rivers containing permanently flowing and drying reaches
and, in one river, along a longitudinal drying gradient.
3. The biomass of aquatic insects, potential prey for D. aquaticus, declined with low river
flows while the abundance of potential terrestrial prey remained similar at all sites. In the
replicate rivers, and along the longitudinal drying gradient, spider biomass was lower, and
size classes were skewed towards more small and fewer large spiders, in drying sites. A
desiccation experiment in the laboratory indicated high sensitivity of the spiders, with
prey presence increasing spider survival.
4. Differences in the spatial distribution, biomass and population size structure of spiders
were observed along the longitudinal drying gradient and disappeared within 16 days of
the water returning to all sites.
5. In total, low river flow affected the biomass of D. aquaticus, as well as their size class
structure and spatial distribution. This indicates that low river flows have the potential to
affect adjacent terrestrial ecosystems.
Keywords: distribution, Dolomedes aquaticus, drought, size structure, spider

Introduction
Flow regime has an overriding influence on the
physical environment of river channels and the
adjacent riparian zone (Bendix, 1997; Amoros &
Bornette, 2002; Stromberg et al., 2007). Flow-related
abiotic factors, such as substratum movement and
hydrological connectivity, also often have strong
influences on the composition and stability of aquatic
communities (Winterbourn, Rounick & Cowie, 1981;
Resh et al., 1988; Williams, 1996; Hart & Finelli, 1999;
Correspondence: Michelle J. Greenwood, National Institute of
Water and Atmospheric Research, P.O. Box 8602, Christchurch
8011, New Zealand. E-mail: michelle.j.greenwood@gmail.com

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Amoros & Bornette, 2002). Furthermore, flow regimes

can impact terrestrial communities via changes in the
abundance and identity of potential prey and the
suitability of the riparian zone as a habitat for
terrestrial predators (Bell, Petts & Sadler, 1999;
Greenwood & McIntosh, 2008; Sperry & Weatherhead, 2008). Many riparian predators, such as insectivorous birds, bats, lizards, beetles and spiders, live
and or forage within the riparian zone, often depending on emerging winged aquatic insects for a large
proportion of their diet (Nakano & Murakami, 2000;
Collier, Bury & Gibbs, 2002; Sanzone et al., 2003;
Baxter, Fausch & Saunders, 2005). A subsidy of
aquatic prey can increase the abundance of terrestrial
predators and sometimes result in strong top-down
 2010 Blackwell Publishing Ltd

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M. J. Greenwood and A. R. McIntosh

between-river differences in the rate and direction of

river drying and the degree of change in wetted width
will have on the response of D. aquaticus to low river
flows. Further research is needed to clarify how
terrestrial predators will respond to the pattern and
speed of river drying, and to investigate whether
aquatic and terrestrial communities, by becoming
adapted, can maintain higher abundances in rivers
that dry regularly or predictably. The fact that we saw
significant and consistent effects of river drying on
D. aquaticus biomass in five different drying rivers
suggests that low flows are a strong determinant of the
spider biomass that can be supported. Moreover, our
study design highlights the value of multiriver comparisons, as opposed to the traditional approach of
studying one river intensively.

Dolomedes aquaticus population size structure and

spatial aggregation
We predicted that the general decline in aquatic
invertebrate biomass (and, by implication, food availability) would affect the size structure of D. aquaticus
populations. Larger individuals (or those in better
condition) should have been able to survive longer
under food stress. However, drying sites were dominated by smaller, juvenile spiders, with very few or
no large spiders occurring. A strategy of depending
on damp soil near the river to resist desiccation is
used commonly in riparian spiders (Carico, 1973;
DeVito & Formanowicz, 2003), and smaller spiders
can presumably use smaller or deeper interstitial
refuges and remain in contact with damp rocks for
longer. In addition, desiccation mortality is likely to
vary with body size, as body surface area to volume
ratios alter (Willmer, Stone & Johnston, 2000), with
smaller spiders often surviving longer than adults
(DeVito & Formanowicz, 2003). However, our desiccation mortality experiment showed no evidence of
size-selective desiccation mortality where death was
very rapid for all size classes at a moderately high
temperature (24 C) and low humidity (30%). In
addition, during the field survey, water was still
present within the river channel at all sites, and
D. aquaticus was often under rocks very close to the
waters edge, which presumably allows them to avoid
desiccation. Thus, it is unlikely that any effects of sizedependent refuge use on D. aquaticus population size
structure were occurring at the time of sampling.

Other size-specific sources of mortality that may

occur at drying river reaches include increased sizespecific predation as D. aquaticus and other terrestrial
predators congregate around the decreasing aquatic
habitat. Dolomedes aquaticus became aggregated as
sites on the Selwyn River dried. This may also have
occurred at the drying river reaches on the replicate
rivers because the river drying had not progressed far
enough to encourage spider aggregations. Spider
aggregations at the drying sites on the Selwyn River
disappeared only 1 day after the water returned,
possibly indicating that high small-scale densities
incur a cost, such as an increased likelihood of
intraspecific encounters and competition. Size-selective predation on large D. aquaticus by other predators
congregating around the drying pools of the river
may also lead to the low proportion and number of
large spiders found at drying reaches.
In conclusion, the changes in biomass and size class
structure of D. aquaticus found at river reaches that
flowed permanently, or had declining flow, were
probably related to a reduction in aquatic prey
abundance as the river dried. Desiccation thermal
stress may also play a role in rivers that dry
completely, as D. aquaticus was shown to be prone
to desiccation mortality. Furthermore, the combination of aggregated spider distributions and low
aquatic food availability provided ideal conditions
for increased predatory or competitive interactions
between conspecifics. Predicted alterations to the
global climate, including an increased occurrence of
droughts in many areas (Arnell et al., 1996), mean that
understanding the impact of river low flows on
adjacent ecosystems is of considerable importance
(e.g. Harper & Peckarsky, 2006). Our results indicate
that the drying of rivers has serious consequences, not
only for the aquatic organisms but also for the
abundance and biomass of terrestrial consumers that
are supported (at least in part) by the river reach. The
nature of the drying regime, the relative dependence
of the consumer on aquatic prey and water, and its
scale of dispersal relative to the scale of drying will
largely determine specific consumer responses and
merits further investigation.

Acknowledgments
Jon Harding, Russell Death, Pete McHugh, Mary
Power, Mike Winterbourn and two anonymous
 2010 Blackwell Publishing Ltd, Freshwater Biology, 55, 20622076