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AQUATIC CONSERVATION: MARINE AND FRESHWATER ECOSYSTEMS

Aquatic Conserv: Mar. Freshw. Ecosyst. 19: 113125 (2009)


Published online 9 October 2008 in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/aqc.895

VIEWPOINT

Integrating ecology with hydromorphology: a priority for river


science and management
I.P. VAUGHANa,*, M. DIAMONDb, A.M. GURNELLc, K.A. HALLb, A. JENKINSd, N.J. MILNERe,
L.A. NAYLORf, D.A. SEARg, G. WOODWARDh and S.J. ORMERODa
a

Catchment Research Group, Cardi School of Biosciences, Cardi University, Cardi CF10 3US, UK
Environment Agency, North West Region, Richard Fairclough House, Knutsford Road, Warrington WA4 1HG, UK
c
Department of Geography, Kings College London, Strand, London WC2R 2LS, UK
d
Centre for Ecology and Hydrology, Wallingford, Oxfordshire, OX10 8BB, UK
e
National Fisheries Technical Team, Environment Agency, University of Bangor, School of Biological Sciences, Deiniol Road,
Bangor, Gwynedd LL57 2UW, UK
f
Department of Geography, University of Exeter, Cornwall Campus, Treliever Road, Penryn, Cornwall, TR10 9EZ, UK
g
School of Geography, University of Southampton, Higheld, SO17 1BJ, UK
h
School of Biological and Chemical Sciences, Queen Mary University of London, London, E1 4NS, UK
b

ABSTRACT
1. The assessment of links between ecology and physical habitat has become a major issue in river research and
management. Key drivers include concerns about the conservation implications of human modications (e.g.
abstraction, climate change) and the explicit need to understand the ecological importance of hydromorphology
as prescribed by the EUs Water Framework Directive. Eorts are focusing on the need to develop ecohydromorphology at the interface between ecology, hydrology and uvial geomorphology. Here, the scope of
this emerging eld is dened, some research and development issues are suggested, and a path for development is
sketched out.
2. In the short term, major research priorities are to use existing literature or data better to identify patterns
among organisms, ecological functions and river hydromorphological character. Another early priority is to
identify model systems or organisms to act as research foci. In the medium term, the investigation of pattern
processes linkages, spatial structuring, scaling relationships and system dynamics will advance mechanistic
understanding. The eects of climate change, abstraction and river regulation, eco-hydromorphic resistance/
resilience, and responses to environmental disturbances are likely to be management priorities. Large-scale
catchment projects, in both rural and urban locations, should be promoted to concentrate collaborative eorts,
to attract nancial support and to raise the prole of eco-hydromorphology.
3. Eco-hydromorphological expertise is currently fragmented across the main contributory disciplines (ecology,
hydrology, geomorphology, ood risk management, civil engineering), potentially restricting research and
development. This is paradoxical given the shared vision across these elds for eective river management based
on good science with social impact. A range of approaches is advocated to build sucient, integrated capacity
that will deliver science of real management value over the coming decades.
Copyright # 2008 John Wiley & Sons, Ltd.
Received 23 February 2007; Revised 11 June 2007; Accepted 1 July 2007
KEY WORDS:

ecology; geomorphology; hydrology; hydromorphology; rivers; Water Framework Directive

*Correspondence to: Dr Ian Vaughan, Catchment Research Group, Cardi School of Biosciences, Cardi University, Cardi CF10 3US,
UK. E-mail: vaughanip@cardi.ac.uk

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I.P. VAUGHAN ET AL.

INTRODUCTION
For over 40 years, issues of water quality have dominated river
research, management and conservation } driven by seminal
publications (e.g. Hynes, 1960), by widespread problems from
point or diuse pollution sources, and by major legislation (e.g.
the UK Water Acts of 1973, 1983; the US Clean Water Act
1977). Although interest in relationships between river organisms
and their physical habitat is also long-standing (e.g. Riley, 1921;
Percival and Whitehead, 1929), emphasis on this has generally
been less. This balance is currently being re-dressed for several
reasons. Globally, there is a need to understand the ecological
eects of a wide range of changes in physical habitat, as rivers are
increasingly exploited, regulated or otherwise modied through
ood-defence engineering, impoundment, river restoration,
climate change and the spread of alien species. Across Europe,
the Water Framework Directive (WFD; 2000/60/EC) has been
the major legislative driver by specifying that hydromorphology
should underpin good ecological status (European Commission,
2000). The improvements in water quality in Europe and North
America over recent decades mean that hydromorphic limits on
ecological quality are becoming increasingly apparent. Finally,
the recognition that these problems require multi-disciplinary
solutions has stimulated dialogue between physical scientists and
biologists whose shared vision is of more eective river science
and management.
River conservation has much to gain in this renewed push
for an improved understanding of ecologyphysical habitat
relationships. Hydromorphological integrity is central to
conservation since it provides the template upon which all
other ecological structures and functions are built. Furthermore,
in seeking good ecological status by sensitive management at
whole basin scales, rather than in the channel or riparian zone
alone, the WFD has become a highly signicant element in
wider river conservation. With hydromorphology an explicit
component of the Directive, the need to understand links to
ecology and conservation are clear.
Three important observations can be made regarding the links
between river ecology and hydromorphology. First, current
scientic understanding is generally poor, especially at the
quantitative levels required for eective prediction and
management. This is despite scientic literature stretching back
more than 80 years (Riley, 1921; Percival and Whitehead, 1929),
and comprising many thousands of peer-reviewed publications.
Numerous } mainly observational } studies have described
links between biological pattern, ecological processes, and river
form and physical processes, yet the underlying mechanisms are
often known only in outline. Relationships in riparian and
oodplain environments are less widely studied than those in the
wetted channel, highlighting the need to consider whole
catchments and river landscapes in the development of ecohydromorphic research (Eyre et al., 2002).
Secondly, improved understanding of ecologyhydromorphology is a pressing need if the timetable and aims of key
river legislation are to be met (e.g. for statutory regulation or
programmes of measures under the WFD). Major challenges
arise in distinguishing the inuences of hydromorphic
modications on organisms or processes from other potentially
confounding eects such as pollution (Allan, 2004). Biological
indicators of physical modication are still preliminary, rarely
described or poorly founded, while few biological models
diagnose how physical eects contribute to biological

Copyright # 2008 John Wiley & Sons, Ltd.

departures from expected conditions (Davies et al., 2000). The


denition of expected or reference conditions as required by the
WFD is challenging given the inherent variability in both physical
habitat and biology (Nijboer et al., 2004).
Finally, the need to understand ecologyhydromorphology
linkages is accentuated by the prospects of climate change,
altered ow regimes and increased water consumption
(Jackson et al., 2001). It is vital that such changes are both
understood and clearly communicated to practitioners if river
environments are to be managed eectively in future. River
environments appear to be highly sensitive to climatic eects,
but features that mitigate impacts or increase ecological
resilience are poorly understood (Durance and Ormerod,
2007).
In this viewpoint paper, we consider the interface between
river ecology and hydromorphology: its scope, major research
and development issues, and sketch out a path for the
development of eco-hydromorphic research. Our aims are
to draw attention to the need for better science that links the
physical and ecological dimensions of rivers; to encourage
better collaboration; to prompt sponsoring agencies to
recognize the gaps; and to prompt bodies responsible for
river management to recognize the need. Some areas are
identied in which collaboration could be most eective.
Naturally, the issues raised reect the perspectives of the
authors, and are indicative rather than denitive. It is hoped,
nevertheless, that this stimulates a wider debate.

ECO-HYDROMORPHOLOGY: THE INTERDISCIPLINARY INTERFACE


Several terms have been applied to the relationships between
organisms and physical habitats in rivers. Examples include
biogeomorphology (Viles, 1988), ecogeomorphology (Parsons
et al., 2003), ecohydrology or hydroecology (Wassen
and Grootjans, 1996), eco-hydromorphology (Clarke et al.,
2003) and geobiology (Noke, 2005). Here, we adopt ecohydromorphology, since it captures the main contributory
disciplines (ecology, hydrology and geomorphology). More
pertinently, this term is consistent with the WFD which uses
hydromorphology to describe the hydrologic and geomorphic
elements of river habitats (European Commission, 2000).
Eco-hydromorphology can be dened as the interactions of
the biological entities and ecological processes of a river with
the hydrological and geomorphological form and dynamics. In
this context:
*

eco- encompasses riverine biota at all levels of


organization
(from
genes,
through
individuals,
populations and communities, to whole ecosystems), all
taxonomic levels and across all functional groupings (e.g.
primary producers, detritivores). It includes ecological
processes manifested in individuals through to entire
ecosystems (e.g. dispersal, reproduction, decomposition),
and which act over a wide range of timescales from the
immediate to the evolutionary (Townsend and Hildrew,
1994) and all spatial scales from local to lifetime
movements (Durance et al., 2006).
hydromorphology describes the geomorphology and
hydrology of a river system, their interactions, and their
arrangement and variability in space and time. Key
elements include the ow (sensu Po et al., 1997) and

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DOI: 10.1002/aqc

INTEGRATING ECOLOGY WITH HYDROMORPHOLOGY

sediment regimes; channel and oodplain dimensions,


topography and substratum; continuity and connectivity
(longitudinal, lateral, vertical and temporal); hydrological
and geomorphological processes (e.g. sediment transport);
and the spatio-temporal arrangement of the hydromorphological components (European Commission, 2000;
Gilvear et al., 2004). Articial features (e.g. bank
protection works, weirs) and human modications to
processes are also included.
interactions } the mechanisms by which hydromorphology
and ecology aect one another. Hydromorphology may
inuence ecology, such as the eects of ow velocity on
macrophyte photosynthesis (Madsen et al., 1993) or the role of
hydromorphology in selecting assemblages of organisms with
appropriate traits (Townsend and Hildrew, 1994). Conversely,
organisms may aect hydromorphology, such as the impact of
biolms on ow characteristics (Battin et al., 2003) or the
impacts of invertebrates on sediment stability and release
(Edwards, 1968).

Eco-hydromorphology extends beyond ecology, geomorphology and hydrology into other contributing elds (e.g.
civil engineering, economics, social sciences) and the majority
of research is not labelled as being eco-hydromorphic (or any
of the other phrases coined) per se. Similarly, it spans both
pure and applied science, academia and regulatory agencies.
This diversity has increasingly been recognized, along with
the consequent needs to engage a disparate research and
management community, and to foster greater interdisciplinary collaboration (Gurnell et al., 2000; Hannah
et al., 2004). Unfortunately, there is little evidence that these
developments are reected in the composition of research
programmes. Hannah et al. (2004) assessed the authorship of
research papers involving the term ecohydrology (and
derivatives thereof) and found that collaborating university
researchers very rarely came from more than one academic
department. It seems that while the eco-hydromorphic
interface can be clearly dened, it is poorly developed, and
unless this situation can be addressed it could seriously
handicap the development of the science.

KEY RESEARCH AND DEVELOPMENT ISSUES


IN ECO-HYDROMORPHOLOGY
Process and causality
Most eco-hydromorphic research has relied upon correlating
static ecological and hydromorphological patterns, using
space-for-time substitutions. Documenting such patterns is
typically the rst stage in the development of a research area,
from which more detailed mechanistic understanding can
develop (Gaston and Blackburn, 1999). In isolation, such
research creates a relatively weak science base, and so
eco-hydromorphology needs to move towards the use of
stronger inference (e.g. experimentation) and studying
underlying mechanisms wherever possible. The next step in
this direction involves the study of dynamics and process rates,
rather than static patterns. Ultimately, there is a need to
understand how eco-hydromorphic processes generate
observed patterns, and in turn how patterns inuence
processes. An example is the way in which vegetation and

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115

sediments interact to shape river channels and ow


characteristics, and, in turn, vegetation recruitment and
sediment transport (Gran and Paola, 2001; Gurnell and
Petts, 2002). The ultimate aim is to achieve a causal
understanding of how the components of the ecology
hydrologygeomorphology interface interact.
River management would benet greatly from
understanding eco-hydromorphic mechanisms, as this would
provide a rm foundation for predicting the eects of
both management interventions and more gradual, longerterm environmental changes. Currently, the challenge for
prediction stems from the novel combinations of
hydromorphic conditions that may be created. In the
UK, for example, climate change is generally predicted to
increase the seasonality of rainfall and runo (Kay et al.,
2006). This may produce novel environmental conditions,
which could have major ramications for ecohydromorphology. Similarly, the ecological responses to river
restoration may be dicult to predict, because specic
structural changes to geomorphology may be made without
altering the formative processes (Sear, 1994) } in eect this
partially or temporarily decouples recognized hydromorphic
processes and forms. To predict the eects of such changes,
models are asked to extrapolate outside currently observed
conditions, and these are the circumstances in which
correlation-based science and models often fail (Sutherland,
2006). In contrast, the underlying mechanisms that link
ecology, hydrology and geomorphology should remain
fundamentally unchanged, and so provide a foundation for
more reliable predictions.
Studying mechanisms, as opposed to correlations, presents a
major challenge. Temporal changes in patterns (e.g. channel
form or invertebrate assemblages) can provide detailed
information about process rates. Fluvial geomorphology
provides good examples, such as quantifying sediment
transport rates using temporal series of channel form records
(Fuller et al., 2002; Lane et al., 2003). However, the nancial and
logistical demands of research spanning many years frequently
restricts opportunities, despite widespread recognition of the
value of research carried out over large spatial and/or temporal
extents, such as the Long Term Ecological Research Network
(Symstad et al., 2003). Equally, management often requires rapid
appraisals (ecological or environmental), which may preclude
temporal monitoring. Progress requires concerted research
eort to address the two-way patternprocess interaction,
helping to identify, for example, how valuable patterns
recorded at a single point in time are for inferring information
about eco-hydromorphic processes.

The role of spatial structure


The spatial conguration of river hydromorphology, and how
it changes over time, has diverse eects upon river ecology
(Wiens, 2002). As a consequence, eco-hydromorphology is
more than an inventory of parts and processes. Potentially
important spatial structures can be identied across a wide
range of scales, from the properties of individual sediment
particles, through the arrangement of biotopes in a reach, to
the large-scale spatial context, including the conguration of
drainage networks and neighbouring catchments. Salmonids
are among the best-studied organisms in the context of spatial
structure, because of the long-standing recognition of both

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I.P. VAUGHAN ET AL.

their migratory movements and the use of dierent habitat


types for spawning and for dierent developmental stages
(Bardonnet and Baglinere, 2000). The role of barriers }
especially articial features (e.g. weirs) } is an important topic
in salmonid research (Morita and Yokota, 2002; Woord
et al., 2005). Spatial structure is a key characteristic, as the
signicance of such barriers depends not only on their
particular characteristics } their permeability to sh under
dierent ow conditions } but also their location within the
river network (Fagan, 2002).
To elucidate the role of spatial structure in eco-hydromorphology, there is a need to identify: (i) the key spatial
characteristics for organisms and how these dier between
life-stages; (ii) how the activities of organisms spatially
structure the hydromorphology; (iii) how physical structure
inuences ecological, physical and eco-hydromorphic
processes; and (iv) ecologically meaningful ways of
quantifying structure. This applies both within the wetted
channel, and to the linkages between the channel and the wider
riverine landscape (Wiens, 2002; Brierley et al., 2006). An
example of these issues is provided by patch boundaries
and ecotones, which are thought to play important roles in
modulating eco-hydromorphic processes and the movements
of organisms (Ward and Wiens, 2001). The most obvious
boundary is the river shoreline, with terrestrialaquatic
interchanges making important contributions to riverine
ecosystems (Nakano and Murakami, 2001). Recent research
has focused upon shoreline length and complexity (Van der
Nat et al., 2002) and how it relates to a range of organisms
and ecological processes, through such mechanisms as increased
water retention and availability of lentic habitats (Reckendorfer
et al., 1999; Schiemer et al., 2001). This research suggests that
shoreline length could be a relatively simple, yet ecologicallyrelevant measure for a range of organisms. In the opposite
direction, ecological processes aect shoreline structure (e.g. via
bank stabilization by growing vegetation and hydraulic eects of
large wood in the channel; Gurnell et al., 2002), providing an
example of ecological processes interacting with hydromorphic
patterns.
The capacity to study spatial structure has increased
dramatically. Technological developments make it possible to
capture patterns over larger areas and at higher resolutions
than ever before, and to analyse the large, complex data sets
that result (Ehlers et al., 2006). The accompanying theory
has also developed, and disciplines such as landscape
ecology provide valuable ideas (ONeill et al., 1988;
Gustafson, 1998).

Scale and variability


The changing uxes of organisms, materials and energy in
space and time, diering across scales, presents a research
challenge. This complexity is not discussed here, but some
generic points are made about scale and variability in ecohydromorphic research.

Scale
The results of research are conditional upon the scale(s) of
observation (Wiens, 1989). For example, the relationships
between organisms and their environment may appear to
change with the scale of observation in both space and time
(Wiley et al., 1997; Malmqvist, 2002). Hence, to understand a

Copyright # 2008 John Wiley & Sons, Ltd.

process, the correct scale(s) must be chosen when studying it.


Organism body size and life cycle have major inuences, such
that smaller organisms with rapid generations (e.g. bacteria,
fungi and algae) interact with river structure over very dierent
spatio-temporal extents from mammals, birds and sh. At one
extreme, large populations of some organisms are contained
within just one river biotope. At the other, many individual
biotopes can be contained within the home range of a single
organism (Woodward and Hildrew, 2002).
Scaling eects also have ramications for monitoring,
assessment and management, all of which are likely to be
enhanced by selecting appropriate scales for the interactions
of interest. To support these aims, research designs should
include scales of management relevance (Vaughan and
Ormerod, 2003; Durance et al., 2006). In the UK, this could
include reach-scale (ca 500 m) units, water bodies, catchments
and river basin districts. Equally, monitoring and
bioassessment need to be aware of the timescales over which
ecological, hydrological and geomorphological patterns and
processes respond to environmental change. An obvious
example is the potential lag in ecological responses to
hydromorphic changes, such as those in the ow regime
following impoundment (Martinez et al., 1994; Kruk and
Penczak, 2003). Management needs to be exible, adopting
scales that research reveals to be critical for eco-hydromorphic
processes.
In lieu of an a priori knowledge of the important scales,
multi-scale studies (in time and/or space) provide a way of
discovering them or insuring against missing critical scales.
Multi-scale studies are relatively well established in river
systems as a consequence of the explicitly hierarchical
organization of rivers (Frissell et al., 1986). Studies focusing
on individual sites and reaches could benet from the addition
of larger-scale factors, as they provide context for individual
sites and may allow the conclusions from small-scale studies
to be more readily generalized (Wiens, 2002). Equally, short
duration studies may benet from longer-term monitoring
data to provide context (Bradley and Ormerod, 2002).
The ultimate aim is to achieve cross-scale understanding in
eco-hydromorphology, revealing the potential ramications of
processes or management interventions on organisms and
processes across spatial and temporal scales. Damming a river,
for example, has eects both locally (e.g. habitat
fragmentation and altered physical structure) and across the
entire river system (e.g. eects on river regime or migratory
organisms) (Allan, 1995). New methods for data analysis may
need to be devised or introduced to eco-hydromorphology to:
(i) reveal cross-scale pattern and process relationships; and (ii)
facilitate scaling up of results from the small spatio-temporal
scales often amenable to research, to larger scales relevant to
management (Phillips, 2005).

Variability
The variability and dynamics of river environments present a
serious research challenge, yet need to be understood for
successful river management (Thoms, 2006). In the rst
instance, empirical study is required to characterize the
variation that occurs (including rates and magnitudes of
change) and this needs to be followed by an understanding of
the role of variability in eco-hydromorphic processes. From a
conservation viewpoint, the observed variability is important

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INTEGRATING ECOLOGY WITH HYDROMORPHOLOGY

to the functioning of river ecosystems (Thoms, 2006), and so


needs to be preserved as part of conservation management.
Eco-hydromorphic variation has been studied over a wide
range of spatial and temporal extents } the latter often in the
course of long-term monitoring, such as ow gauging (Sheer
et al., 2003), retrospective aerial photograph/map analyses of
geomorphology (Tiegs and Pohl, 2005) and annual river bird
surveys (Carter, 1989). Long-term monitoring can provide
invaluable information about the behaviour of river systems,
as can palaeohydrological investigations that extend the study
of river variability over hundreds or thousands of years, or
even into longer geological and evolutionary time-scales (Sear
and Arnell, 2006). Methodological improvements, both in
terms of data collection (e.g. increased accuracy and precision
of remote sensing) and analysis (e.g. spatio-temporal statistics)
have greatly increased the potential to study variability, yet
major gaps in understanding remain (Thoms, 2006).
Alongside the characterization of variation, frameworks/
paradigms
are
needed
to
incorporate
it
into
eco-hydromorphology. Concepts such as meta-stability,
where a system is considered to vary within certain
boundaries } unless subjected to major perturbations } are
valuable (ONeill et al., 1989). Much emerging theory in ecohydromorphology places variability at its core, such as the
concepts of ood pulses (Junk et al., 1989), ow pulses
(Tockner et al., 2000) and the natural river regime (Po et al.,
1997). This trend needs to continue, acknowledging variability
in river ecology, hydrology and geomorphology, and their
interactions.
The ramications of eco-hydromorphic variability are less
clearly understood. In part, this stems from the diculty of
capturing temporal variation in the short time-scales employed
in most research. When spatio-temporal variability is
examined explicitly, more detail can be revealed about the
underlying processes. For example, Langhans and Tockner
(2006) revealed marked dierences in leaf litter decomposition
rates on the river Tagliamento oodplain according to the
durations and frequencies of inundation. In the Colorado
River basin, Cooper et al. (2003) found variations in the
patterns of Populus and Tamarix recruitment that
corresponded to the frequency and magnitude of high and
low ow events revealed by long-term gauging data. Making
the links between variability and processes is therefore a clear
research priority.
The spatio-temporal variability observed in rivers has been
greatly altered by several thousand years of anthropogenic
changes in catchment vegetation cover and land use (Birks
et al., 1988), and } more recently } by direct modications to
hydromorphology (e.g. ood defences). For most research and
management processes, the dynamics within these existing,
modied systems are of great interest. However, understanding
natural systems is also important, for example in studying
hydromorphic processes and acting as references against which
modied dynamics can be compared (Tockner et al., 2003), but
to do so requires the eects of widespread modication to be
circumvented. Coupled palaeohydrological and palaeoecological studies examining the behaviour of rivers prior to
major human modications provide one possibility (Brown,
2002). Another approach is through the direct observation of
relatively undisturbed rivers. In Europe, such systems are rare
} and therefore valuable (e.g. Fiume Tagliamento; Tockner
et al., 2003). Both approaches have weaknesses, stemming from

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117

incomplete information in reconstructing the past or questions


over the generality of model rivers, but nevertheless can
provide unique insights into the behaviour of natural rivers.

Eco-hydromorphic responses to environmental change


An understanding of how river systems respond to environmental (external) changes is of fundamental research interest
and vital for successful management. Changes may consist
either of short-term pulse disturbances of varying magnitudes
(e.g. weed cutting, ood events), or longer-term press
changes, such as shifts in land use or climatic changes
(Brunsden and Thornes, 1979). Depending upon the
particular
river
system
and
disturbance,
ecohydromorphology could show no change, temporary
displacement followed by a return to pre-disturbance
conditions, progressive adjustment to the new levels of the
drivers or a rapid shift to a dierent set of conditions/stability
domain (Brunsden and Thornes, 1979). Understanding the
ecological responses to such disturbances should enable
practitioners to appraise alternative policies based on their
likely impacts on the river system and to set upper limits to
potentially damaging activities (Groman et al., 2006). An
example of the latter are the attempts to nd an acceptable
level of urban development within a catchment before changes
in ecology are observed (Allan, 2004).
The component disciplines of eco-hydromorphology have
made valuable progress in studying river responses to
environmental change. Eco-hydromorphic responses to high
ow events and ooding provide a case in point. Numerous
studies document how channel forms have changed following
ooding, and then shift to a dierent state or recover their
original form (e.g. Schumm and Lichty, 1963; Myers and
Swanson, 1996; Sloan et al., 2001). Similar documentation of
ood eects has been carried out on river ecology (e.g. Power
and Stewart, 1987; Boulton et al., 1992), as well as
experimental disturbances intended to mimic ooding (e.g.
Melo et al., 2003). Ecological or hydromorphological
responses to ooding (rapid recovery or shift to a dierent
form) have been conceptualized over recent decades, invoking
ideas including stability, sensitivity, resistance, resilience,
stability domains separated by thresholds, nonlinear
dynamics and complexity theory (Leopold and Wolman,
1957; Holling, 1973; Schumm, 1979; Phillips, 1992; Downs
and Gregory, 1995; Gunderson, 2000; Stallins, 2006). By
combining empirical observations and conceptual models, the
mechanisms underlying these behaviours are being elucidated,
such as the roles of feedback mechanisms and thresholds in
maintaining or switching between alternative system states
(Dent et al., 2002).
Developments in ecology, hydrology and geomorphology
are tempered by the limited development at their interface.
While much of the work on environmental change focuses on
hydromorphic and ecological interactions, it often suers from
the common problem of expertise concentrated in a single
discipline (Hannah et al., 2004). The conceptual frameworks
for studying the responses to change have developed in
isolation, such that analogous physical and ecological
concepts are given dierent terms and classied in dierent
ways. For example, while thresholds and process (or stability)
domains are dened and used in similar ways, the concept of
landscape sensitivity encompasses the ecological notions of

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I.P. VAUGHAN ET AL.

resistance and resilience in a more holistic concept (Brunsden


and Thornes, 1979; Gunderson, 2000). One conclusion is that
there is a clear need to unite dierent disciplines in the course
of studying environmental change, to share a common
language and concepts (Benda et al., 2002). In terms of
specic research and management aims, there is a need to
identify: (i) the responses of dierent elements of ecohydromorphology to dierent forms of environmental
change (e.g. brief perturbations or prolonged changes); (ii)
the nature of any thresholds in the external drivers (e.g.
climate) of river systems beyond which major changes occur;
(iii) the proximate drivers and more gradual changes that
reduce resistance/resilience (or increase the sensitivity) to
environmental changes (e.g. reductions in water quality
reducing the ecological resilience to hydromorphic
disturbance); (iv) the trajectories of eco-hydromorphic
changes, both in response to perturbations and in recovering
to the previous state; and (v) indicators of when ecohydromorphology is in the vicinity of a threshold, to trigger
management interventions (Groman et al., 2006).

Covariance between eco-hydromorphic elements


Many elements of the riverine environment covary. Urban land
adjoining a channel, for example, may be associated with
modied water quality, altered ow regime, structural changes
to the channel (e.g. channelization, bank reinforcement) and
disruption of processes such as sediment supply (Paul and
Meyer, 2001). Concomitant ecological changes in such
situations (e.g. reduced taxonomic diversity or increased
decomposition rates; Paul and Meyer, 2001) could be a
response to any or all of the changes associated with the land
use. Their eects could be additive, subtractive or synergistic.
Covariance between elements of river systems is both a
challenge and a benet. The challenge occurs in establishing
causal relationships among correlated variables. In the case of
urban development, this may include separating the roles of
hydromorphic modications from reduced water quality.
Where feasible, experimentation could break some of the
correlations and identify key factors, such as the distinct
inuences of substratum and ow velocity on macrophytes }
two variables that are normally correlated (Chambers et al.,
1991). However, experimentation creates articial conditions,
potentially limiting generalizations unless all of the important
factors are addressed.
The potential benet of covariance stems from the opportunities to describe general eco-hydromorphic relationships.
To pursue the urbanization example, if consistent relationships
between water quality and hydromorphic variables could be
characterized, it might be feasible to combine them in general
ecologyurbanization relationships (Paul and Meyer, 2001).
In this way, the observed covariance may simplify ecohydromorphic study. A sound principle underlies this idea,
because, assuming that correlations have been correctly
identied, the covariance between eco-hydromorphic
components is indicative of some common causal process(es)
} albeit unresolved and potentially far removed from the
observed variables (Shipley, 2000). Studying the covariances
between eco-hydromorphic components could assist in the
generation of causal hypotheses.
Two implications of multicollinearity for data analysis are
noteworthy. The rst is that many studies will benet from

Copyright # 2008 John Wiley & Sons, Ltd.

being placed in a broader context, identifying a wider set of


covarying eco-hydromorphic elements to avoid drawing
premature conclusions about the relevant variables. Second,
careful choice of methods is crucial. Factor analysis or
ordination methods can help to describe collinearity and
identify underlying structures/variables that have common
causes (Vaughan and Ormerod, 2005). Path analysis and
structural equation modelling may provide a more relevant
framework for dealing with inter-correlated variables (Pugesek
et al., 2003). Statistical variable selection methods (e.g. stepwise
regression) should be avoided, as they are unreliable in the
presence of collinearity and present the temptation to make
inferences about the main variables based upon statistical
signicance (Graham, 2003).

Development of tools and methods


The need to develop novel or more rened tools and methods
reects (i) the complexity and uncertainty surrounding river
research and management, (ii) the diverse audience } in terms
of opinions, interests and technical understanding } that
needs to be engaged, and (iii) attempts to link ecology and
hydromorphology more directly, rather than simply
correlating patterns. Numerous issues could be suggested:
here, this is restricted to three examples.

Linking ecological and hydromorphic processes in


research
Identifying and redressing technical limitations is a clear
priority to facilitate stronger coupling of physical and
ecological processes. At present there is often a mismatch in
the capacity to capture the ecological features and the physical
environment at the empirical level, and this may limit progress
in some aspects of research. For example, laser scanning can
rapidly quantify channel morphology over several biotopes
(hundreds of square metres) at high resolutions (potentially
sub-centimetre), and using repeat imaging, quantify sediment
transport (McCarey et al., 2005). Matching biological
sampling to such data presents problems, both in terms of
obtaining thorough coverage of such large areas and capturing
how organisms such as benthic invertebrates interact with
physical habitat at such high resolution (Peckarsky, 1991).
Developing the temporal dimension of eco-hydromorphic
measurement is vital if the links between processes are to be
made more directly than using simple space-for-time
correlations between patterns. Signicant technological
advances have been made, such as the potential applications
of passive integrated transponder (PIT) tagging to monitor
organism or sediment particle movements at higher temporal
resolutions than previously (Lucas and Baras, 2000; Lamarre
et al., 2005).
Nevertheless, fundamental limits remain on the ability to
measure eco-hydromorphic patterns and processes in both
space and time. For example, in a rapidly changing ow
regime with concurrent change in bed morphology, it is at
present impossible to quantify spatial patterns at meso-habitat
scales (ca 0.5 m resolution), let alone microhabitat (Cliord
et al., 2005). In lieu of such abilities, modelling presents a way
forward. Physically based numerical modelling frameworks are
capable of accurately representing ow structures in 13
dimensions and over time (Parsons et al., 2004). Sear et al. (in

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INTEGRATING ECOLOGY WITH HYDROMORPHOLOGY

press) advocate the use of coupled ecologicalgeomorphic


models within an experimental design to explore the
relationships between hydromorphic processes and the
resulting habitats. Agent-based models provide a versatile basis
on which to relate the behaviour of organisms or ecological
processes to physical habitat structure or processes (Booker
et al., 2004; Rashleigh and Grossman, 2005). Many modelling
frameworks are adept at handling nonlinear systems. Using
observations of ecological patterns and physical forms to
calibrate process rates and system behaviour in models,
followed by further observation to test the models, provides a
powerful way of reconciling the desire to study processes with
the relative ease of recording patterns.

Addressing uncertainty
The importance of uncertainty in research and management
has long been recognized, yet rarely addressed adequately.
Uncertainty derives from a range of sources, including
measurement errors, the weak science-base for much of ecohydromorphology, conicting evidence about a phenomenon,
and issues } especially in the future } that can never be
known (Van Asselt and Rotmans, 2002). It is vital that
uncertainty be addressed explicitly in many situations, to avoid
over-estimating condence in conclusions or predictions, or
setting unrealistic goals for management (Clark, 2002). River
restoration projects provide a good example, being inherently
complex and involving a high degree of uncertainty from a
range of sources } especially when projects are viewed over
geomorphologically relevant timescales (Sear et al., in press).
Explicitly acknowledging uncertainties provides a way of
managing unrealistic stakeholder and societal expectations
(Clark, 2002; Rogers, 2006).
Frameworks are required that consider uncertainty, along
with tools with which to describe or quantify it (Clark, 2002).
On one level, analyses that handle known uncertainties such as
measurement error can be adopted readily. Recent
developments using Bayesian statistics, information-gap
theory and other methods illustrate how this may be possible
in eco-hydromorphology (Regan et al., 2005; Halpern et al.,
2006). At a higher level, frameworks for placing management
within the context of all uncertainties } known and unknown
} are being developed (Johnson and Brown, 2001).

Decision making and communication tools


The need for decision making and communication tools is a
consequence of: (i) the diversity of stakeholders involved in
river management; (ii) acknowledging the complexity and
inherent uncertainty in managing river systems; and (iii) the
concomitant capacity for conicts of interest (Clark and
Richards, 2002). Potential conicts occur frequently, such as
the desire to increase water abstraction for drinking and
agriculture, compared with the desire to minimize human
impacts. This has driven the development of a range of
decision support and communication tools. For example, costbenet analyses are increasingly being performed, often in
terms of improvements in water quality (Eisen-Hecht and
Kramer, 2002; Mourato et al., 2005), but also relating to ecohydromorphology. Kuby et al. (2005) developed a modelling
method for dam removals in the Willamette River basin in
Oregon, which compared the potential benets for salmonid

Copyright # 2008 John Wiley & Sons, Ltd.

119

migration against socio-economic losses (water storage and


hydropower) under dierent scenarios, and which suggested
signicant habitat connectivity benets for relatively low levels
of dam removal. Willis and Garrod (1999) considered the
balance between water abstraction and the potential losses of
recreational income associated with low ows, and showed
that recreational angling in particular could provide nancial
justication for many types of low ow alleviation.
Developments in decision-making need to be coupled to
initiatives aimed at making the tools, and the outputs from
them, readily accessible to the relevant parties and policy
makers. Arnold et al. (2000) described education programmes
for making GIS-based decision support systems for catchment
land-use more accessible to local planners. Ultimately,
decision-making partnerships are required that involve all of
the stakeholders } scientists, managers, land owners and
wider society (Rogers, 2006) } in the same way as research
requires an inter-disciplinary approach.

THE DEVELOPMENT OF ECOHYDROMORPHOLOGY


Eco-hydromorphology is in its infancy, despite decades of
relevant research, and it is valuable to consider the framework
within which it could develop (Figure 1). Except for specic
eco-hydromorphic interactions that have been well studied,
much of the research eld still appears to be concerned with
identifying gaps in understanding and framing key research
questions, with expert opinion and some initial literature
reviewing helping to clarify the gaps (Figure 1). In the
preceding discussion some general research gaps have been
highlighted, and in the conclusions some key research aims and
questions are suggested.
Three approaches could be employed to address the research
questions: formal literature reviewing, use of existing data
resources, and dedicated data collection and experimentation
(Figure 1). With research spread across at least three separate
disciplines and many decades, reviews of the riverine ecohydromorphic literature are extremely important to synthesize
current understanding and to introduce ideas from the
individual disciplines across eco-hydromorphology. The
opportunities for data mining and other post hoc analyses are
considerable, given the extensive data resources }
ecological, hydrological and geomorphological } that have
been collected over many years. The overlaps between such data
sets in space and time provide a basis for valuable ecohydromorphic investigations, along the lines of the rebecca
project (relationships between ecological and chemical status of
surface waters), which uses existing data from across Europe,
mainly to examine the linkages between water quality and
benthic macroinvertebrates. Dedicated research programmes
are likely to be the only way of addressing many of the research
questions, but should be supported by literature reviewing and
opportunistic data analyses wherever possible.
Evidence appraisal is vital (Figure 1). The three basic
research methods can be arranged around a simple hierarchy
(Figure 2), representing a ladder of evidence in terms of the
depth of understanding and strength of the evidence.
Literature reviews and analyses of existing data can be
assigned to broad sections of the ladder, the precise positions

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120

I.P. VAUGHAN ET AL.

Informal literature
search

Literature
reviewing

Knowledge gap

Legislation

Key questions

Expert
opinion

New data
collection

Analysis of
existing data

Evidence appraisal filter

Science base

Basic science

Tools

Detailed
policy

Practice &
management

Figure 1. A framework for the development of eco-hydromorphic


research and management.

depending upon the rigour of both the methods used and those
of the contributing studies or data resources (Figure 2).
Literature reviews are potentially far more valuable than any
individual study owing to the accumulated mass of evidence,
especially if comparable studies are performed under nonidentical conditions, helping to reveal how general the ndings
are over space and time (Lindsay and Ehrenberg, 1993).
These should result in a critically appraised body of ecohydromorphic research, which could, in turn, feed into both
pure and applied outputs (Figure 1). Eco-hydromorphology
could contribute to basic science in a wide range of areas, both
in terms of general theory (e.g. diversity and ecosystem
functioning, ecologyphysical habitat interactions) and in the
ways that rivers dier from terrestrial and marine systems
(FBA, 2005). In applied areas, eco-hydromorphic research
should help to generate an evidence base for river management
(Sutherland et al., 2004) and underpin management tools (e.g.
decision support frameworks or bioassessments of
hydromorphological pressures). From all of these outputs,
and indeed from preceding stages in the framework, feedback is
anticipated to the early stages, representing further renement
of the knowledge gaps and key questions (Figure 1).

CONCLUSIONS: PRIORITIES FOR THE


DEVELOPMENT OF ECOHYDROMORPHOLOGY

discussed. Some of the most valuable short-term work could


focus on making the best use of existing resources } literature
or data. Multi-disciplinary reviews would not only help to
clarify the extent of eco-hydromorphic science, but could also
stimulate development of the area through the combination of
initiatives from the dierent disciplines. Opportunistic data
analyses and data mining will generally be limited to static,
correlative research, yet this should help to provide initial
answers to questions and provide quantitative hypotheses to
guide more detailed research. Another early priority is to
identify model systems or organisms that could act as research
foci, as many research questions will require time series data,
creating a minimum start-up period before they can be
addressed.
In the longer-term, numerous research lines requiring new
data collection can be identied. In this paper, patternprocesses linkages, spatial structuring, scaling relationships,
system dynamics and responses to environmental change have
been considered, and a few of the many research possibilities
within them suggested } many others could be proposed.
Over-arching priorities are to: (i) aim for mechanistic
understanding wherever possible (cf. simple, correlative
investigations); and (ii) try to approach research areas from
multiple disciplines, to draw expertise from dierent areas and
avoid unnecessary duplication of research eort resulting from
a lack of communication between elds. Given current
emphasis on climate change, abstraction and river
regulation, research into eco-hydromorphic resistance/
resilience and responses to environmental change are likely
to be particular priorities.

Monitoring and methods


Numerous issues can be identied relating to the methods and
tools required for research and management (Table 1). In the
short term, most of the aims identied focus on the appraisal
of current methods and ways in which they could be improved
(Table 1). In the longer term, the focus moves to the
development of specic eco-hydromorphic methods, rather
than the adaptation of methods developed for more specialized
purposes.
Carefully designed river monitoring programmes are
extremely important and have much wider value than their
monitoring role. The opportunistic analysis of existing data
sets can address a range of questions, yet is heavily dependent
upon the data that are available (Figure 2). Routine
monitoring programmes are potentially the most important
resource in this respect, with national coverage and temporal
time series; indeed, they may be the only such resource,
unequalled in their sample sizes and coverage. Unfortunately,
many monitoring programmes may have developed in a
piecemeal fashion and involve biases in eort or sampling
scheme (e.g. biases toward road networks). Appraising and
rening existing monitoring programmes, with a view to
optimizing their scientic rigour, would permit more valid
conclusions to be drawn for river monitoring and management,
as well as bringing widespread benets to eco-hydromorphology.

Research

Management and practice

Numerous research gaps can be identied, and several that are


considered priority areas for eco-hydromorphology have been

Suggested management priorities are a combination of


organizational and opportunistic measures, coupled with

Copyright # 2008 John Wiley & Sons, Ltd.

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INTEGRATING ECOLOGY WITH HYDROMORPHOLOGY

Literature
reviewing

New data
collection

Systematic
review

Strong, general evidence-base

121

Analysis of
existing data

Dedicated, replicated
experimental manipulations

Before-after comparisons

Experiments e.g.
data collected
before and after
flood defence works

Stronger correlative methods e.g.


independent testing,validation in
other rivers or with other
organisms
Qualitative
literature
review
Exploratory data analysis, expert
opinion.
HYPOTHESIS GENERATION

Data mining

No evidence-base

Figure 2. A ladder of evidence describing the relative strengths of literature reviewing, analyses of existing data and new data collection. New
investigations form the main body of the ladder, with stronger research methods and scientic inference encountered on higher rungs. Results from
most investigations can contribute to the overall evidence base (dotted lines). Literature and data-based research are plotted on the same scale, such

Table 1. Some monitoring and methodological priorities in riverine eco-hydromorphology


Short term
* Review current ecological and hydromorphological monitoring programmes:
what potential is there to add (i) hydromorphological measurements to water quality or ecological monitoring schemes, and (ii) ecological
variables to hydromorphological surveys (e.g. uvial audit)?
coverage, sample sizes and other concerns.
* Appraisal of hydromorphological survey methods (e.g. River Habitat Survey, physical habitat components of the US EPAs rapid
bioassessment protocol):
how relevant are they to ecology? Do they capture underlying processes and spatial structure?
do they operate at ecologically relevant scales?
do dierent methods complement one another? e.g. eld surveys, remote sensing and ow gauging
what elements are missing and could be added to increase the eco-hydromorphic relevance?
* Improved description of eco-hydromorphic variability, to help distinguish background variation from overall system changes.
* Evaluate appropriate research methods in eco-hydromorphology: opportunities for experimentation, eld survey designs, data analysis tools
for stronger inference (e.g. structural equation modelling)
* Strengths and weaknesses of dierent surveys and analysis methods, and guidance regarding when each should be used.
Longer term
* Targeted environmental sampling to document ecologyphysical habitat relationships. e.g. along gradients of hydromorphological pressures
* Comparison of eco-hydromorphological assessment tools and models in demonstration catchments
* Long-term monitoring at both near-pristine and modied sites } quantify variability and compare between natural and modied systems
* Developing specic eco-hydromorphological methods cf. modifying existing ecological or hydromorphological methods:
measures that address causal mechanisms and processes
multi-scale analysis tools
frameworks for up-scaling from small-scale experiments/monitoring to scales of management relevance
* Adopt more mechanistic/dynamic approaches to modelling and data analysis e.g. agent-based models to simulate ecological interactions with
physical habitat

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I.P. VAUGHAN ET AL.

Table 2. Some priorities for management and practice in eco-hydromorphology


Short term
* Dening the role for ecohydromorphology in dierent areas of catchment management
* Collate best available evidence for ecologyhydromorphology relationships } mainly literature on correlative ecologyhydromorphology
relations
* Establish a widely accessible repository for best practice and guidance for assessing habitat status, managing hydromorphological pressures
(e.g. ood defence works), and restoration and conservation schemes
* Devise forums for involving the full range of stakeholders and wider society in decision making
* Develop standard monitoring protocols for interventions (e.g. weir installation, river restoration) to ensure that as much information as
possible can be derived about the eects on ecology and hydromorphology
* Develop tools for handling uncertainty (e.g. decision support tools) and ways of setting management priorities (e.g. identifying important reaches
for the functioning of hydromorphic processes or those readily improved according to socio-economic criteria)
Longer term
* Causal understanding of ecologyhydromorphology relationships (cf. simple speciesenvironment correlations):
natural eco-hydromorphic relationships
responses to point hydromorphological modications (e.g. weirs, local engineering works)
responses to diuse or distal hydromorphological modications (e.g. land use, sediment loading, climate change)
* Appraise conservation interventions and river restoration using long-term monitoring
* Research the concepts of resilience and thresholds as they apply to ecology, hydromorphology and their interactions, e.g. extent of urban
development or impaired connectivity (longitudinal, channeloodplain or channelhyporheic zone)
* Integrate socio-economic concerns, e.g. cost-benet analyses
* Understanding the eects of combined stressors (e.g. morphological, chemical and climatic) on river ecology

some specic research and development aims (Table 2). Much


of current management, developing environmental standards
for hydromorphology and trying to identify actions that might
threaten or enhance river ecological quality, relies upon expert
opinion, with relatively little underpinning science. In the more
medium term, there is a need for research and development
that will enhance the capability for detecting, diagnosing and
predicting hydromorphological eects on ecological status.
Across Europe, this understanding will be vital into the next
WFD cycle (i.e. from 2015 onwards). The hope is that with
the development of eco-hydromorphology, management will
evolve into an increasingly evidence-based, science-led process.

Final point: capacity development


Eco-hydromorphological expertise is widely scattered across
the three main contributory disciplines (ecology, hydrology
and geomorphology) and beyond (e.g. civil engineering,
chemistry, social science). In some areas there has been a
dramatic loss of research capacity, such as among freshwater
ecologists in the UK (FBA, 2005; Raven, 2006). In other areas,
policy or regulatory requirements have tended to focus on
site-specic management (e.g. engineering or hydraulic works),
rather than considering the wider catchment. A change
in emphasis, through such initiatives as Making Space for
Water (in the UK) and the WFD, requires the skills of
geomorphologists, sedimentologists, hydrologists and allied
elds working at catchment scales to deliver eective riverine
planning and management (European Commission, 2000;
Defra, 2004). The dispersed nature of this wider ecohydromorphic community } both research and management
} has the potential to restrict both its research and
development capacity, and its ability to inuence policy.
A major priority in developing eco-hydromorphology, and
river research and management more generally, is to reverse
the declines and overcome the fragmented distribution of
expertise. A thriving community needs to be built and
sustained, linking the expertise that is available across
disciplines and institutions (including universities, research
institutes, regulatory bodies and conservation organizations).
Regular conferences and workshops, with broad invitations,

Copyright # 2008 John Wiley & Sons, Ltd.

are one way to achieve this. Web-based initiatives could also


prove to be an eective way of building and maintaining links
across subject and geographical boundaries (Foote, 1999).
There is a strong case for promoting large-scale projects to act
as foci for collaborative eorts, to attract nancial support and
to raise the prole of eco-hydromorphology. The studies based
in the Hubbard Brook Forest, USA (Bormann and Likens,
1979) and at Llyn Brianne in upland Wales (Edwards et al.,
1990; Durance and Ormerod, 2007) provide illustrations of
how eective and enduring demonstration catchments can be.
The addition of urban catchments for this purpose would not
only improve understanding of urbanization eects, but also
make it easier for the value of the research to be appreciated by
the wider public. Focusing resources on such catchments is a
long-term commitment, but this brings the benets of
understanding processes and change over a range of timescales.
Finally, it is vital to recognize the international nature of the
eco-hydromorphological community. In Europe, the WFD
provides an obvious focus for international collaborations, as
member states need to address similar challenges. Other
European developments, such as the EU Water Initiative,
could also play a role. Globally, similar questions are being
asked and challenges faced. This is evident from a series of
United Nations initiatives, including the International
Hydrological Programme, the World Water Assessment
Programme and Millennium Development Goals. All of
these involve consideration of sustainable use of water
resources: eco-hydromorphology is a key aspect in achieving
this. The extent of global interest reveals both the
opportunities for international collaboration, and the degree
to which such collaboration could benet research at the
ecologyhydrologygeomorphology interface.

ACKNOWLEDGEMENTS
This paper resulted from a workshop (Linking Physical
Habitat Structure to Riverine Biodiversity) held as part of the
UK Population Biology Network (UK PopNet), funded by the
Natural Environment Research Council (Agreement R8-H1201) and English Nature. Further funding was provided by the

Aquatic Conserv: Mar. Freshw. Ecosyst. 19: 113125 (2009)


DOI: 10.1002/aqc

INTEGRATING ECOLOGY WITH HYDROMORPHOLOGY

Environment Agency. We wish to thank all of the workshop


participants for valuable discussion and ideas, as well as
Professor Philip Boon and two anonymous referees, all of
whose comments enabled us to make valuable improvements
to the manuscript.

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