Вы находитесь на странице: 1из 15


Hydrol. Process. 21, 1749 1763 (2007)

Published online 14 February 2007 in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/hyp.6313

The concept of hydrological connectivity and its contribution

to understanding runoff-dominated geomorphic systems
Louise J. Bracken1 *, and Jacky Croke2

1 Department of Geography, University of Durham, Durham DH1 3LE, UK

School of Physical, Environmental and Mathematical Sciences, UNSW@ADFA, Northcott Drive, Canberra, ACT 2601, Australia

The term connectivity is increasingly being applied in hydrological and geomorphological studies. Relevant research
encompasses aspects of landscape connectivity, hydrological connectivity and sedimentological connectivity. Unlike other
disciplines, notably ecology, published studies show no consensus on a standard definition. This paper provides an overview
of how existing research relates to the concept of connectivity in both ecology and hydrology by proposing and evaluating
a conceptual model of hydrological connectivity that includes five major components: climate; hillslope runoff potential;
landscape position; delivery pathway and lateral connectivity. We also evaluate a proposed measure of connectivity called the
volume to breakthrough to quantify changing connectivity between different environments and catchments. Copyright 2007
John Wiley & Sons, Ltd.

connectivity; runoff generation; hillslope; catchment; volume to breakthrough; roads

Received 17 March 2005; Accepted 24 March 2006

The concept of connectivity is increasingly being
applied within a range of disciplines in the Earth and
Environmental Sciences as researchers recognize the need
to move beyond the traditional view that runoff is generated by either Hortonian infiltration excess or by the
variable source area (VSA) model. Recently, the VSA
model itself has been the focus of critique, with calls
for a new theory of runoff generation (e.g. McDonnell, 2003; Ambroise, 2004). Hydrological connectivity
is one possible concept of runoff generation and flood
production that could provide a way forward, but currently there is much confusion in the literature about
how the term is used and how it relates to existing
In hydrology and geomorphology we can identify
three major types of connectivity, broadly classified
as: (1) landscape connectivity, which relates to the physical coupling of landforms (e.g. hillslope to channel)
within a drainage basin; (2) hydrological connectivity,
which refers to the passage of water from one part of
the landscape to another and is expected to generate some
catchment runoff response; and (3) sedimentological connectivity, which relates to the physical transfer of sediments and attached pollutants through the drainage basin
and may vary considerably with, amongst others, particle
size. Work on coupling of hillslopechannel systems (e.g.
Harvey, 1996; Michaelides and Wainwright, 2002) has
* Correspondence to: Louise J. Bracken, Department of Geography, University of Durham, Durham DH1 3LE, UK.
E-mail: l.j.bracken@durham.ac.uk
e Bull.
Copyright 2007 John Wiley & Sons, Ltd.

alerted us to important differences in catchment response

when hillslopes are directly connected to a drainage line
or are disconnected, e.g. through the presence of an adjacent floodplain (Figure 1). There has also been considerable research on aspects of hydrological connectivity,
although not always referred to as such, including runoff
generation at the patch, hillslope and catchment scales
that makes it difficult to draw work together into a single
theoretical model of runoff connectivity. For example,
detailed studies on runoff connectivity between areas of
varying soil conductivity and vegetation at the patch or
band scale (Morgan, 1995; Fitzjohn et al., 1998; Cammeraat and Imeson, 1999; Ludwig et al., 1999a,b, 2000)
highlight the complex spatial and temporal patterns of
runoff connectivity even at this micro-scale. Other work
has focused on identifying natural or artificial factors
that affect hydrological connectivity, highlighting features such as dams that may reduce flow connectivity
or compacted surfaces of roads that may enhance runoff
connectivity through the physical integration of the road
and stream network (Wemple et al., 1986; Croke and
Mockler, 2001).
In terms of sediment, recent work has adopted the concept of connectivity to describe the the transfer of sediment from one zone or location to another and the potential for a specific particle to move through the system
(Hooke, 2003). This work has focused on the physical
connection and transfer of sediment through a reach and
basin (Fryirs and Brierley, 2000; Harvey, 2001; Hooke,
2003). Sediment connectivity as a concept has always
been implicit in such empirical approaches as the sediment delivery ratio, which assumes the physical transfer
of material from hillslopes to the channel and through



Coupled - hillslope leads directly into the channel

De-coupled - hillslope and channel separated by the floodplain





Deposited fine
Mean river level


(belo roughflow
w wa
ter ta

Figure 1. Illustrations of connectivity: (a) hypothetical models to illustrate simple channel coupling (modified from Michaelides and Wainwright
(2003)); (b) a more complex floodplain environment that may limit connectivity

the drainage basin. Typically, a lack of connectivity is

implied. Such an approach, however, has traditionally
been black box, with little real understanding of the
spatial and temporal patterns of sediment movement from
source areas to sinks, especially as they relate to varying
particle sizes. Although such approaches are clearly not
suitable to measure connectivity, it is of interest to note
that many early concepts implicitly include some representation of connectivity. The application of sediment
tracers and particle tagging techniques have recently identified some of the complexities of sediment transport and
storage and suggest that sediment connectivity is driven
by dynamic physical processes that reflect the interaction
of a large number of factors.
A consistent definition of the term, however, remains
difficult to discern from published studies in hydrology
and geomorphology. This may now be limiting the application of the concept to catchment management and modelling studies. The lack of a clear definition in hydrology
is in contrast to other disciplines, where there is a more
consistent application of the term connectivity, which
is often used to explore the dynamics of movement of
species (e.g. Pringle, 2001, 2003). The question, therefore, is whether we can standardize a definition that will
allow us to quantify and model hydrological connectivity. This quest seems to be in accordance with the view
of many ecologists who also call for a more predictive understanding of hydrologic connectivity (Pringle,
2003). The question comes at a time when hydrology
and geomorphology appear to be in need of refined concepts and new theoretical models to move beyond the
VSA concept. McDonnell (2003) argues for a post-VSA
Copyright 2007 John Wiley & Sons, Ltd.

model based on non-linearities and proposes flexible, box

models as a concept to support this. Ambroise (2004) suggests more accurate definitions and argues for differentiation between variable active and variable contributing
areas, but also importantly, variable active and variable
contributing periods.
The purpose of this paper is threefold. First, we discuss how the concept of connectivity has been used in
ecology to explore the experiences and lessons of another
discipline. Second, we develop a framework for hydrological connectivity which we use to synthesize the large
body of existing research on hydrological connectivity.
The resultant review focuses primarily on runoff generation and delivery as the key mechanisms in promoting
catchment connectivity. As such, infiltration and runoff
production are intimately linked, but our discussion goes
beyond the spatial variability of infiltration, which some
may consider the key issue influencing connectivity. By
focusing on runoff, a spatial pattern of connectivity will
emerge that corresponds closely with the movement of
fine particles. However, we do not discuss runoff thresholds (e.g. Wolman and Gerson, 1978; Ritter et al., 1999);
rather, we assume, first, that runoff thresholds must be
exceeded for runoff to be produced and, second, that all
factors that influence runoff thresholds are important for
hydrological connectivity. Third, we establish one empirical measure called the volume to breakthrough, which
can be used as a starting point in quantifying hydrological
connectivity between different catchments within a range
of different environments. Finally, we provide an example of this approach using a road network as a landscape
feature that enhances hydrological connectivity.
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp



Ecologists have been using the concept of connectivity
for a number of years as a critical property in the persistence of spatially structured populations (Metzger and
Decamps, 1997). In this context, hydrological connectivity is defined by Pringle (2001) as the water-mediated
transfer of matter, energy, and/or organisms within or
between elements of the hydrologic cycle. This definition, although encompassing other material fluxes as
well as water, bears some similarity to the application
of the term hydrological connectivity. Using this more
rigorous definition, numerous ecological studies have
contributed to a clearer understanding of why hydrological connectivity is essential to the ecological integrity
of the landscape, where alterations that either reduce or
enhance connectivity are seen to have potentially significant environmental impacts (Pringle, 2003). Likewise,
biologists define landscape connectivity as the degree to
which a landscape facilitates or impedes the movement
of individuals (Taylor et al., 1993). Hence, the concept
is both a landscape- and species-specific characteristic
(Wiens et al., 1997; Brooks, 2003). Important landscape
characteristics include shape, size and location of feature in the landscape, whilst key species qualities are
behaviour and patterns of gene flow (Brooks, 2003). No
index allows cross-system comparisons (Davidson, 1998)
and no empirical studies consider the various scales that
are implicit in exploring connectivity (Brooks, 2003).
As with geomorphology, ecology faces conceptual and
empirical issues with heterogeneity between landscapes
and species (Wiens et al., 1997; Brooks, 2003). Interestingly, an ecologists definition of connectivity combines
a measure of spatial integration of the landscape with the
movement of species (Table I).
Overall, there appears widespread recognition that
the term connectivity in any of these geo-ecological
senses is useful in promoting the interconnection between
the morphological components of the landscape and
the material fluxes that move across, and through, the
drainage basin. The term has also allowed us to think
of dis-connectivity (or isolation as used in ecological
studies) between both landscape parts and sources of

water and sediment. As a result, catchment managers

have found the term useful in identifying the non-linear
relationship between on-site disturbances and off-site
response. It appears that the word connectivity is used
to convey a similar meaning in ecology and biology,
including something about the spatial variation in flows
across landscape surfaces that result in the development
of habitats and communities.

At its most general, connectivity describes all the former

and subsequent positions, and times, associated with the
movement of water or sediment passing through a point in
the landscape. This definition makes no assumption about
process. Hydrology is then about describing the regularities in the observed distributions. The starting point
for developing a useful conceptual framework for hydrological connectivity, therefore, lies in resolving the issue
of spatial scale and observed distributions of connectivity. Figure 2 presents one framework for understanding
how characteristics at a point relate to both hillslope and
catchment runoff response, with patterns established at
smaller scales nesting into large-scale response. Previous work on runoff production has also provided some
useful insights into the heterogeneity of landscapes and
the variability in runoff connectivity even at the patch
scale. Cammeraat (2002), for example, identified a set of
key processes at the plot, hillslope and catchment for a
semi-arid and temperate environment (Figure 3). One of
his most important findings is establishing the relationship between rainfall intensity and the rainfall amount
required to generate runoff at different scales. The variation in response between rainfall and runoff was found
to be strongly related to the recurrence period of rainfall,
with the fine scale (patches/plots) producing many more
runoff events than the hillslope and catchment scales
(Cammeraat, 2002).
Likewise, Ambroise (2004) recently used the example
of the tiger-bush pattern of vegetation banding (cf.
Thiery et al., 1995) to illustrate significant differences
between what has previously been termed contributing
Table I. Methods used to determine ecological connectivity
area and what he proposes should more accurately be
called active areas. Surface runoff occurred over greater
than 50% of the area (active area >50%) but none of this
runoff was connected to the downslope outlet (contributStructural connectivity
Patch cohesion
Schumaker (1996)
ing area, 0%). Similar problems have been identified at
Proximity index
Whitcomb et al. (1981)
the hillslope scale, and this has resulted in the introducAverage nearest neighbour
ONeill et al. (1988)
tion of terms such as effective hillslope length (Aryal
et al., 2003) or dynamic contributing areas (Barling,
Fractal dimensions
ONeill et al. (1988)
1992; Beven, 1997). Approaches to resolving such hetCorridors
Andreassan et al. (1996a,b),
Habbad (1999a,b), Habbad
erogeneity in hillslope runoff production are considered
et al. (2000)
in more detail below. Fundamentally, it is such variability
Empirical studies
in governing factors that has been one limiting factor in
Stamps et al. (1987)
the accurate implementation of hydrological connectivSpecies richness
Golden and Crist (2000)
Demaynadier and Hunter (1988) ity in many physically-based models (Beven and Freer,
2001, Ambroise, 2004).
Copyright 2007 John Wiley & Sons, Ltd.

Hydrol. Process. 21, 1749 1763 (2007)

DOI: 10.1002/hyp




Soil / Lithology

Profile form
Plan form

Connected area

Storm profile over time

Areal distribution
Intensity scaled to runoff
Travel distances

Catchment response
Storm volume
Dist'n of HRUs
Effective intensity

Figure 2. The nesting of factors influencing hydrological connectivity at

different spatial scales

In a broad sense, we start with a conceptual framework

for hydrological connectivity that contains five major
components: (1) climate, (2) hillslope runoff potential,
(3) landscape position (4) delivery pathway and (5) lateral buffering (Figure 4). Within each of these components, there are a number of factors that may influence
the extent to which a catchment may be regarded as connected. The following sections review work on each of
these components and factors and their relative influence
on runoff production and catchment connectivity. This
overview is by no means exhaustive; as such, this framework is best viewed as a structure for exploring potential
gaps in the process understanding and, importantly, data
necessary to quantify connectivity.

0 metres 4

Response order source areas:

First response

Gutter system
Bare area (runoff source)

Second response
Drainage divide


Climatic environment
Climate. A key control on the pattern and distribution of runoff within a catchment is climate, specifically



Figure 4. The components of catchment connectivity
Copyright 2007 John Wiley & Sons, Ltd.

Third response

Flow line

Figure 3. An example showing connectivity modified from Cammeraat

(2002). Measured runoff comes from the first response areas, but with
increasing rainfall intensity and duration the second and third response
areas start to contribute

the runoff regime as determined largely by the nature

and distribution of rainfall. Runoff is mainly composed
of saturation overland flow (SOF) in humid-temperate
regions, whereas Hortonian overland flow (HOF) is more
common in semi-arid and arid regions. The VSA concept was developed for humid regions in the 1960s
(Betson, 1964; Hewlett and Hibbert, 1967; Dunne and
Black, 1970). Connectivity in humid temperate environments thus depends on the connection between saturated
areas, namely hillslope hollows and stream channels.
Antecedent conditions are important and usually there is
some base flow operating in the catchment and stores of
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp


subsurface flow waiting to be reconnected during rainfall. Hence, wet area expansion occurs frequently and
connectivity in runoff can be relatively easily achieved.
The direction of connectivity within a catchment may be
working headwards from the channels as saturated areas
expand, but also downslope from saturated hollows to
stream channels (Figure 5). A review of issues related to
connectivity in temperate areas can be found in Burt and
Pinay (2005).
The VSA concept does not hold in semi-arid and arid
environments because most runoff is produced by HOF
(Bryan and Yair, 1982). There are no areas of saturation
to initiate quick runoff because storms are infrequent and
of short duration. Run-on infiltration and transmission
losses are also high, which limits the development of
continuous hydrological pathways. Key runoff-producing
areas tend to be found on steep slopes, either abandoned
after agriculture or with sparse vegetation, and are
composed of runoff-promoting soils such as marls (e.g.
Bull et al., 2000). These areas do not necessarily relate
to the channel network, and the mosaic pattern they form
is the key to producing floods in ephemeral channels.
Hence, in semi-arid and arid areas, the conceptual
model of connectivity is very different from humidtemperate areas. As a general rule, it is more difficult for
connectivity to be achieved in drylands and it occurs less
frequently than in humid-temperate areas. Antecedent
conditions are relatively unimportant in drylands and
there is no base flow or stores of through flow waiting to
be reactivated. Connectivity occurs by the interaction and
response of a mosaic of patches to rainfall, and depends
on the rainfall duration and intensity being great enough
to allow transmission of water over hillslopes and into
channels, and then to propagate down channels overcoming transmission losses to connect whole catchments
(Figure 5). For example, Lavee et al. (1998) found that
in semi-arid areas there was a mosaic of arid watercontributing areas and moist water-accepting patches.
The two scenarios outlined above represent the opposite ends of a continuous spectrum of runoff response for
a range of different environments. In most areas, both
SOF and HOF will be produced in different proportions
depending on rainfall intensity and surface characteristics. Hydrological connectivity depends on the interaction


Runoff producing patch

First expansion of runoff area

Final runoff producing area

Channel network

Figure 5. Saturated areas expanding in humid-temperate environments

and mosaic patches of runoff that connect to produce flooding in dryland
Copyright 2007 John Wiley & Sons, Ltd.


of saturated areas that follow the river channels and hillslope hollows, and the mosaic of areas that are sensitive
to intense rainfall. In such areas, therefore, the response
to rainfall and hydrological connectivity is dynamic and
will change depending on the nature of rainfall input,
antecedent conditions and catchment characteristics.
Storm intensity and duration. It has been proposed that
rainfall intensities are of prime importance for flood generation in all catchment sizes (Costa, 1987; Schick, 1988;
Pitlick, 1994), and it has been appreciated that high discharges are produced by an optimal combination of basin
morphometry and storm intensity (Costa, 1987). Storm
intensity and duration are thus crucial to producing connectivity, and are the inputs that combine with the spatial
pattern of integration to produce connected flow. Other
storm attributes that influence connectivity are the temporal variation in rainfall throughout the storm and the
variation of rainfall spatially within a catchment. For
example, the timing of a high-intensity pulse of rain during a storm event can be crucial for connectivity. Light
rainfall can have the effect of wetting up a catchment, so
that when a high-intensity pulse occurs the runoff generation is rapid and transmission losses low, thus encouraging connected flow. However, a high-intensity pulse on
a dry catchment may have little impact, since infiltration
rates are high and thresholds for runoff generation and
connected flow are not exceeded. Schick (1988) demonstrated that two-thirds of runoff events in the Negev and
Sinai are generated in small to medium catchments by
high-intensity, short-duration storm events, which have
extremely flashy hydrographs and have dramatic effects
on geomorphology. The remaining one-third are more
long-term floods, produced by low rainfall intensities
that may last a few days. In contrast, research undertaken on the Colorado Plateau in Utah by Dick et al.
(1997) showed that rapid runoff was produced during
short-duration storms, which resulted in complex hydrographs that reflected channel network geometry. The
largest flow event recorded in the area was produced by
a low-intensity storm of long duration, which occurred
when antecedent moisture was high (Dick et al., 1997).
Romkens et al. (2001) suggest that storms with initially
high intensities have the potential for the early onset of
concentrated flow, which has obvious implications for the
probability of connectivity.
The temporal pattern of high-intensity rainfall is also
important for determining flow distances of runoff which
connect source areas and result in connectivity at the
hillslope and catchment scale. Relationships are complex and difficult to investigate, so there is not a large
a body of literature to draw upon. Yair and Lavee (1985)
conducted a series of experiments on hillslope runoff
response using varying rainfall intensity. Uniform runoff
generation across slopes was only found for storms with
rainfall intensities greater than 9 mm h1 and for durations greater than 45 min. They found that the properties of storm events explained discontinuities in runoff
from semi-arid and arid areas. Wainwright and Parsons
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp



(2002) investigated the relationship between varying rainfall and spatial variation in infiltration capacity, slope
length and runoff coefficients. It was found that the temporal structure of the storm influenced the runoff coefficient because runoff produced during the start of the
storm could infiltrate as the storm continued. Temporal
variation in rainfall intensity was significant in controlling the scale dependency of the runoff coefficient. It was
suggested that using constant rainfall intensities in modelling underpredicts runoff generation. These two studies
are supported by Van de Giesen et al. (2000), who found
that the runoff coefficient varied between storms with
different spatial and temporal dynamics.
Figure 6 presents an example of the variability in
storm characteristics that produce floods in the semiarid environment in southeast Spain. The dominant
relationship is between the total rainfall in a spell and
the size of the flood peak. In this analysis, spells were
defined as periods of rainfall without breaks greater
than 12 h. Thus, a new spell begins once it rains and
has been dry for over 12 h. The seven largest spells
of rainfall all exceeded 50 mm in total, and produced
flood peaks in the main channel; however, smaller events
can also produce a hydrograph peak. The higher flood
peaks fit more closely with the trend identified compared
with the smaller flood peaks, which have more scatter
(Figure 6). There is also a slight relationship between
the duration of a spell of rain and the flood peak,
although this relationship is likely to be caused by the
correlation between spell duration and spell total rainfall.
More results such as these are necessary to understand
fully the impact of storm intensity and duration on
Catchment-scale hydrological connectivity and flooding, therefore, overall requires prolonged, high-intensity
storms, whereas hillslope hydrological connectivity can
be initiated by shorter duration, or lower intensity events.

Each catchment has a base spatial pattern in terms of connectivity, depending on key runoff-generating areas, and
a response curve as the catchments wets up.
Hillslope runoff potential
Related factors. It is now commonly accepted that
catchments have a non-uniform and non-linear hydrological response to rainfall (e.g. Cerd`a, 1995; Bergkamp
et al., 1996). This is caused by spatial variability in
hydrological properties due to complex geological, pedalogical and management histories (Fitzjohn et al., 1998)
combined with temporally and spatially variable rainfall
(Thornes, 2001). The hillslope is the major landscape
unit and is the scale at which most research on runoff
generation has taken place. The processes of runoff and
run-on lead to dynamic interactions between different
zones of spatially variable hillslopes, and these interactions affect the large-scale response of catchments to
produce floods (Fiedler et al., 2002). As both Lal (1990)
and Bryan (2000) highlight, there is no single, measurable soil property that can fully represent the integrated
response that constitutes soil surface. There are many
factors that influence hillslope runoff, including crusting
and surface roughness (Auzet et al., 1993; Helming et al.,
1998; Singer and Le Bissonnais, 1998), heterogeneity
within the soil (Fitzjohn et al., 1998), the impact of variable density and type of vegetation (Imeson et al., 1992;
Bergkamp et al., 1996), changing catchment morphometry, transmission losses in tributaries and main channels
(Reid and Frostick, 1997), and the impact of land use
(Bull et al., 2000; Lasanta et al., 2000). We do not discuss
each variable in detail; rather, we specifically highlight
the role of each factor for connectivity.
Infiltration is a key process in runoff generation and,
hence, connectivity. The infiltration rate is probably
the most important hillslope characteristic for runoff
generation at the plot scale. In contrast, the spatial
Mean storm intensity

Total storm rainfall











Peak stage (m)


Storm duration (days)

Maximum storm intensity


Peak stage (m)







Peak stage (m)




Peak stage (m)


Figure 6. The relationship between floods and storm characteristics for the Rambla de Torrealvilla, southeast Spain
Copyright 2007 John Wiley & Sons, Ltd.

Hydrol. Process. 21, 1749 1763 (2007)

DOI: 10.1002/hyp


variation in runoff, related to the spatial variation in

infiltration, is more important for hillslope and catchment
connectivity. Once runoff is produced, a key factor
for connectivity is whether runoff can continue over
a hillslope surface or proceed down channel without
infiltrating and, hence, produce run-on and connected
flow. Hence, run-on, infiltration and transmission losses
have an important role in determining runoff pathways
and flood production. Both infiltration (on the hillslope)
and transmission losses (in channel) are influenced by
gradient, although increased gradient can either increase
(Kirkby et al., 2002) or decrease runoff (Poesen, 1984;
Govers, 1991). In terms of connectivity, there must be a
sufficient volume of water to run on and reach the base
of a hillslope and, hence, provide connected flow over
an area. If the volume of water is not sufficient to do
this, then runoff may decrease downslope whatever the
gradient. Raindrop impact can either dislodge particles or
seal the surface to reduce infiltration and increase runoff
(De Ploey, 1984; Govers, 1991; Bradford et al., 1987a,b).
Yair et al. (1980) found that, in the Zin Valley, Israel,
owing to the high stability and strong flocculation of
clay-rich aggregates, rainsplash is ineffective in surface
sealing. Infiltration capacities, therefore, remain high
despite prolonged periods of rain. Crusting can result in
a marked decrease in infiltration and possible protection
against erosion by rainsplash and overland flow. Crusts
influence connectivity at the plot scale; but, as with
infiltration, it is the spatial variation at the hillslope and
catchment scales that is more important for connectivity.
Antecedent conditions are important with respect to
the dynamic variation of connectivity at all scales,
and result in an expansion of saturated areas before
storm rainfall. The spatial pattern of antecedent soil
moisture is hard to predict and is highly spatially and
temporally variable, although pockets of soil moisture
remain after rainfall (Puidefabregas et al., 1998). As soil
moisture increases, the likelihood of connection being
made between sources of runoff increases (Fitzjohn et al.,
1998). Antecedent conditions have a large impact on the
spatial patterns of infiltration and runoff production and,
hence, connectivity. However, the impact is complex and
there is little research to assist in defining conceptual or
mathematical models of connectivity.
Surface roughness is complex, because it operates
at a number of spatial scales, can be highly dynamic
and influences runoff pathways. For instance plough furrows are relatively large and can channel flow in certain directions (form roughness) (Govers et al., 2000;
Kirkby et al., 2002), whereas individual stones and soil
aggregates can be larger than the flow depth (hydraulic
roughness), encourage infiltration and reduce the velocity
of run-on. At the small scale, disintegration of aggregates, differential swelling, surface sealing and crusting
are dynamic over the duration of a storm, while providing hydraulic roughness (Slattery and Bryan, 1992).
Micro-relief patterns due to micro-rills, crustose lichens,
pedestals and pinnacles can also result in the development
Copyright 2007 John Wiley & Sons, Ltd.


of efficient pathways (Imeson and Verstraten, 1988; SoleBenet et al., 1997). Hydrological connectivity produced
by surface roughness is influenced by slope. Smooth surfaces are not greatly affected by gradient, but for rough
surfaces the surface area changes dramatically and can
affect depression storage, runoff and the development of
rills as effective coupling between areas of a hillslope.
Runoff is usually concentrated along channels determined
by topography or agricultural practices (Ludwig et al.,
1995), and some research has shown that, as roughness
increases, flow concentration occurs, thus increasing erosion (Abrahams and Parsons, 1990; Helming et al., 1998;
Romkens et al., 2001) and, in turn, increasing connectivity.
Vegetation has a complex relationship with all the
factors mentioned above and is a major influence on
hydrological connectivity at all scales. Indeed, Imeson
and Prinsen (2004) suggest that patterns of vegetationbare soil can be used as indicators of the extent, distribution and connectivity of runoff and sediment sources
and sinks. Vegetation has a positive effect on infiltration by increasing organic matter, lowering bulk density
(Thornes, 1976; Boix-Fayos et al., 1998; Wilcox et al.,
1998) and increasing hydraulic conductivity (Nicolau
et al., 1996). At the plot scale, research has shown that
infiltration rates beneath bushes can increase infiltration
up to three times that of the inter-shrub area (Lyford and
Qashu, 1969). Vegetation also reduces rainsplash and,
hence, crust formation, but increases surface roughness
(hence hydraulic resistance) and ponding. Many investigations have noted patterning of vegetation at the hillslope scale (Sanchez and Puidefabregas, 1994; Dunkerley,
1999; Valentin et al., 1999; Puigedefabregas, 2005; Boer
and Puigedefabregas, 2005), which can greatly increase
infiltration compared with the inter-vegetated areas. During storms, therefore, runoff is generated on the bare
areas, but runoff infiltrates when it reaches a vegetated
area, limiting connectivity.
However, the spatial configuration of bare and vegetated areas controls runoff, not just the actual areas of
each (Morgan, 1995; Fitzjohn et al., 1998; Cammeraat
and Imeson, 1999; Ludwig et al., 2005). Within ecology
there has been a lot of research based on spatial variations in vegetation and how this is related to hydrological
processes. Banding in vegetation and resulting patchiness has been explored and evaluated in terms of theory
(Wiens, 1995), causes (Belsky, 1995), genesis (Thiery
et al., 1995), dynamics (Mauchamp et al., 1994) and
function (Ludwig et al., 1999a,b, 2005). There have been
many different terms for vegetation patchiness, varying
from two-phase (Archer, 1990; Belsky, 1995), two-phase
mosaics (Montana, 1992), stripes (Cornet et al., 1991;
Thiery et al., 1995) and banding (Slatyner, 1961; Boyland, 1973; Mabbutt and Fanning, 1987). These different
patterns are related to soil depth and soil fertility (Tongway and Ludwig, 1990; Ludwig and Tongway, 1995).
Loss of patchiness has been shown to be the greatest
influence on the ability of hillslopes to capture rainfall
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp



(which is related to plant productivity), with the difference between different types of patchiness (stripes,
strands or patches) being small compared with the differences between vegetation patch types (Ludwig et al.,
1999a). Ludwig et al. (1999b) also found that the redistribution of runoff into run-on spaces between vegetation
patches is greater in drier environments relative to wetter ones. The ratio of bare : patch length decreased as
the rainfall increased in intermediate textured soils (Ludwig et al., 1999b). Ludwig et al. (2000) also propose
that scaling equations determined for the relationships
between bare areas and vegetation patches can be used
in computer simulation models or in hypothetical situations to collapse the complexity of scaling issues into
practical management frameworks.
This type of research in ecology benefits the discussion of hydrological connectivity in two ways. First, it
develops our understanding of factors influencing hillslope infiltration and runoff. Second, and possibly more
important, it also examines the spatial variability in hillslope characteristics. One obvious benefit may be the
development of spatial scaling equations for use in upscaling simulation models.
Land management affects runoff response and the spatial integration of source areas at all scales of the landscape and is a key factor in connectivity. Many studies
have shown that changing from natural vegetation to
cultivation can increase soil erosion rates by an order
of magnitude or more (Abernethy, 1990; Morgan, 1995;
Walling, 1999). Increasing soil erosion by rill development has a direct relationship with increased connectivity.
Land management includes crop planting, road building,
land-use change and reservoir construction. Ploughing is
one of the major controls of surface properties, removing
vegetation, destroying crusting, and moving rock fragments to the surface (Oostwoud Wijdenes et al., 1997),
but even abandonment of previously farmed land has a
complex relationship with runoff (Lasanta et al., 2000).
Ludwig et al. (1995) found that interactions between agricultural and topographic features resulted in a complex
temporary hydrological hillslope structure, which consisted of a runoff collector network composed of topographic or agricultural depressions and the distribution of
runoff contribution areas, that varied over time.
Temporal variability in the factors affecting connectivity is important, because over the duration of a storm,
as well as for longer time periods, many factors affecting
connectivity are subject to change. For instance, during an
individual storm the infiltration capacity decreases as the
soil becomes saturated and fines block pores. The microscale surface roughness also changes as soil aggregates
are broken down and crusts develop, also decreasing the
infiltration capacity. However, erosional processes, such
as rill development, operate during a storm and, once
formed, rills result in increased form roughness and act
as efficient pathways that channel runoff. Clay minerals
also swell during wet periods and shrink during drying, also affecting the infiltration capacity, and hence
runoff. Intense cracking can also result in marcopores. At
Copyright 2007 John Wiley & Sons, Ltd.

the larger temporal and spatial scale, vegetation changes

also occur, resulting in seasonal differences in vegetation cover, surface roughness and runoff production, and
hence connectivity.
Landscape position
Landscape position in the context of hydrological connectivity is included to reflect the relationship between
runoff source and distance to the outlethillslope or
catchment. Intuitively, the probability of hydrological
connectivity will be enhanced if the transport distance for
water is short relative to the effective contributing area.
In its simplest sense, this can be expressed as distance to
stream or outlet.
Slope length influences connectivity at the hillslope
and catchment scales, and is relatively unimportant
at the plot scale. On longer slopes it is more likely
that the slope will cease to generate runoff before
runoff reaches the slope base, or channel. There is
a complicated relationship between slope length and
rainfall duration and intensity that produces connected
flow at the outlet. At the hillslope scale, a range of
investigations have proposed a decrease in runoff per
unit area with increasing slope length due to increased
opportunity for infiltration (Yair and Lavee, 1985; Lal,
1997; Van de Giesen et al., 2000; Wainwright and
Parsons, 2002). This is also affected by crust formation,
vegetation and land management practices. However,
Lal (1997) suggested that, on longer slopes with low
infiltration capacities, the runoff coefficient increases with
distance. The issue of slope length is complicated by the
formation of rills. Rills may increase infiltration capacity
if crusts are broken, but they also operate as channels and
increase the velocity of runoff and effectively reduce the
hillslope length (Truman et al., 2001).
Delivery pathway
Each runoff source has its own very specific delivery pattern that is dependent upon its landscape position
within the catchment and, in many instances, the management practices employed. The delivery of this water
downslope involves flow pathways of variable width,
depth and velocity (Figure 7). Broadly speaking, runoff
flowpaths can be described as incisional (such as rills and
gullies, which are represented in the landscape as a definable channel that acts to concentrate overland flow) and
dispersive (which can involve some concentrated flow
but are typically wide and shallow). Runoff delivered in
channellized pathways is characterized by high-energy
flow with little, or no, potential for runoff infiltration
and sediment deposition. Where these incisional features
are fully connected to a drainage line, the probability of
hydrological connectivity is high. Diffuse overland flow
paths have received less attention than the more obvious morphological pathways associated with rill or gully
development. This is partly related to the difficulty of
quantifying attributes of the overland pathways during
storm events and the lack of appropriate techniques to
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp




# # ##

# ## #


Connected Dispersive
Forest Road
Natural Drainage

#S # #
# #
# ##


Scale -













Figure 7. Pattern of hydrological connectivity in a forested catchment

in southeastern Australia (adapted from Croke et al. (2005)). Connected
dispersive dots illustrated the locations where the volume of runoff
discharging from the drain exceeds the available hillslope length and
connectivity is predicted to occur. Gullied points represent locations
where the road and stream network are connected by fully developed

quantify pathway width, depth, and velocity relationships

of surface runoff.
Dominant controls on the type of runoff pathway
include such factors as topography, especially the effects
of steepest slope, convergent hillslopes and hillslope hollows, but increasingly the effects of anthropogenic structures are emphasized. For example, a number of studies have demonstrated the significance of concentrated
flow paths at road outlets with respect to channel initiation and gully development (Montgomery, 1994; Wemple
et al., 1996; Croke and Mockler, 2001; La Marche and
Lettenmaier, 2001). Likewise, the presence and spatial
organization of certain linear features, such as tracks, furrows and ditches (Auzet et al., 1993; Ludwig et al., 1995,
1996; Desmet and Govers, 1997; Souch`ere et al., 1998;
Takken et al., 2001; Moussa et al., 2002) that collect and
guide runoff, have been noted to affect runoff flow paths
and hydrological connectivity.
Lateral buffering
The term lateral connectivity is well defined in ecological studies to describe the nature of flood inundation
between a channel and the adjacent floodplain (Pringle,
2001; Brooks, 2003). It has been recognized as a fundamental control on nutrient and organic matter transfer
between the main channel and adjacent areas of the floodplain. In terms of the conceptual framework proposed
here, we use the term later buffering, because hydrological connectivity will be significantly influenced by the
degree to which (a) hillslopes are physically connected
to channels and (b) the degree to which lateral buffering
acts to limit runoff and sediment delivery to the channel.
An example of the first instance occurs where hillslopes
Copyright 2007 John Wiley & Sons, Ltd.


feed directly into streams, especially low stream order.

For example, Michaelides and Wainwright (2002) proposed that decoupling reduced the relative importance of
the sensitivity of hillslope parameters to runoff, and that
sensitivity decreased with increasing magnitude of runoff
event. In the case of the latter, we refer to the presence
of wet areas or extensive riparian vegetation that may act
to limit runoff connectivity with the main channel. For
example, a recent review of the impacts of riparian vegetation on hydrological processes (Tabacchi et al., 2000)
describes some of the influences of both living and dead
plants on the physical transfer of water through the riparian zone. Following the redistribution of runoff on the
hillslope, overland flow enters a riparian buffer strip to
encounter an environment with the capability of greatly
improving water quality. Riparian zones are normally
characterized by a very rough soil surface, often with an
intact litter layer. The soil is porous with many macropores and the rooting zone is frequently deep. Deposition
by infiltration relies on there being an unsaturated surface
in the riparian zone so that the volume of overland flow
is reduced. The very porous nature of the undisturbed
riparian zone soil assists in this process, whereas the presence of a wet zone or capillary fringe from a water table
inhibits it. The zone may also be characterized by the
soil being wet due to subsurface redistribution of excess
rainfall from the hillslope (Dunne and Black, 1970). In
some circumstances, the riparian zone can remain moist
throughout the year due to the influence of the water table
and associated capillary fringe from an adjacent stream
(Abdul and Gillham, 1989). There is also the alternative
view that riparian zones can increase connectivity. For
example, in lowland Britain, agricultural drainage systems can result in the rapid transfer of storm water and
increase the efficiency of connectivity (Burt and Haycock, 1996; Burt 1997; Pinay et al., 1998; Burt and Pinay


Existing research on hydrological connectivity investigates two slightly different, but equally important, aspects
of runoff generation within natural landscapes: static and
dynamic representations of hydrological connectivity. By
static we mean spatial patterns, such as hydrological
runoff units (HRUs), that can be categorized, classified
and estimated. Figure 8 shows areas identified as key
regions of potential runoff within the Rambla Nogalte
in southeast Spain (Bull et al., 2003). These were predicted using 50 m resolution data on topography, land
use and geology. Comparison of this map with data from
a 1 in 7-year flood in September 1997 demonstrated that,
although some of the areas did produce runoff, there
was considerable variability between sites. For example,
the area labelled A is a hill, which did not produce
enough runoff to overcome run-on infiltration at the base
of the slope to produce connected flow into the main river
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp



channel. Likewise, the area labelled B is an area of tributaries disconnected by human-built drainage ditches to
channel flood water into wheat fields. However, with different input rainfall, such as a 1 in 10-year event, the areas
labelled A and B may produce runoff that connects with
the main channels to produce flood flow. Hence, there are
different responses with varying amounts of hydrological
runoff and resulting connectivity for different events.
In an attempt to characterize the complex distribution of spatial and temporal factors better, research
has focused on approaches to group or classify similar
hydrological surfaces. For example, for a storm to initiate catchment-scale runoff it must overcome the spatial
arrangement and threshold values of hydrologically similar surfaces (HYSS) at all scales (Kirkby et al., 2002).
Similarly, HRUs are defined as distributed, heterogeneously structured entities having a common climate,
land use and underlying pedologicaltopographical
geologic association controlling their hydrological
dynamics (Flugel, 1995). For hillslopes and channels to
be connected and produce floods, HRUs need to be interconnected and connect to main channels. Therefore, the
overland flow contribution from responsive areas depends
on their location within the catchment and their spatial
arrangement with respect to each other (Sharma et al.,
1987; Yair, 1992; Fitzjohn et al., 1998). Where there is no
connection between runoff-producing areas, only those
areas located adjacent to the channel contribute to catchment outflow, and runoff from spatially isolated areas
will be reinfiltrated (Yair, 1992; Fitzjohn et al., 1998).
Hence, hillslopes and catchments can be spatially isolated or spatially interactive (Sharma et al., 1987; Cerd`a,
1995; Fitzjohn et al., 1998); however, connectivity also
depends on the input of rainfall (duration and intensity)
to the system to produce connected flow.
By dynamic hydrological connectivity, we mean both
longer term landscape development, such as changes
following abandonment of agriculture, and short-term
variation in antecedent conditions and rainfall inputs
to systems that result in non-linearities in hillslope
and catchment response to rainfall. Dynamic aspects of
connectivity are more difficult to quantify, but they are
possibly more important than static elements because
it is the combination of static and dynamic elements
High potential runoff


Figure 8. Areas of high runoff potential for the Rambla Nogalte in

southeast Spain. The catchment area is 171 km2 and the main channel is
33 km in length
Copyright 2007 John Wiley & Sons, Ltd.

of connected hillslopechannel systems that provides

a realistic representation of hydrological connectivity
in natural landscapes. For instance, Wainwright and
Parsons (2002) found that the combination of timevarying rainfall intensity during a storm and run-on
infiltration provided an alternative explanation for scale
dependency in runoff coefficients, rather than the more
traditional explanation of spatial variation in infiltration
capacity (e.g. Yair et al., 1980; Lal, 1997; Wilcox et al.,
1997; Van de Giesen et al., 2000; Yair and Kossovsky,
One proposed measure of hydrological connectivity can
be expressed using the concept of volume to breakthrough, which means the accumulated runoff volume
per unit width to be applied at a point before flow appears
at a downslope point. In this sense, hydrological connectivity is seen as a function of the actual runoff produced
by an area, transmission losses due to such factors as
infiltration, depression storage, etc., slope length, and the
existence of features (natural or constructed) that either
prevent runoff reaching channels (such as dams and terraces) or those that encourage runoff connecting with
channels (such as roads and tracks). Hence, the strength
and development of connectivity are controlled by characteristics of both the storm and the landscape. Furthermore, the term can be applied at a range of spatial scales
from patch, hillslope length and within a catchment.
However, because of the general lack of hydrologic data
documenting conditions promoting connectivity, there are
some limitations inherent in such a definition. For example, we have limited data on the range of runoff volumes
required to initiate breakthrough and specifically on variability in connection with key aspects such as antecedent
conditions along the potential pathway. Likewise, such
a definition describes only the establishment of connection, not its continuity over time or with the end-of-storm
behaviour. In spite of these limitations, the volume to
breakthrough could form a unifying concept in quantifying connectivity at a range of spatial scales. Using the
specific example of hydrological connectivity from road
outlets, we demonstrate below the collection and interpretation of data that can be used to quantify connectivity.
Road-to-stream connectivity is a specific form of catchment connectivity that identifies the potential impact of
runoff from a specific source, such as a road or track, on
in-stream water quality and catchment response. Road
surfaces have been identified as highly compacted surfaces that generate rapid overland flow for the majority of
low to moderate rainfall events (Reid and Dunne, 1984;
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp



Luce and Cundy, 1994; Ziegler et al., 2000). As such,

we define their runoff potential as high, both in terms of
HOF and saturated subsurface flow, as road cuts intercept the hillslope (Megahan, 1983; Wemple and Jones,
2003). The connectivity of the road drainage to the stream
network determines the efficiency by which runoff intercepted by road cutslopes and road surfaces is routed to the
stream via drainage outlets (La Marche and Lettenmaier,
2001). Road runoff may (a) infiltrate into the soil directly,
(b) enter the stream directly at a stream crossing culvert,
(c) infiltrate below a gully that does not extend to the
stream channel, or (d) enter a stream channel indirectly
through the formation of a gully below a relief culvert. In
case (a) or (c), the road drainage is not connected to the
stream network. In cases (b) and (d), the road network is
connected to the drainage network directly and indirectly
The probability of road-to-stream connectivity is
strongly influenced by the nature of the delivery pathway.
On road and track surfaces, the spatial pattern of runoff
delivery is determined by the drain spacing employed. A
number of studies have found gully development at road
outlets where contributing road area is high (drain spacing
is inadequate) and the discharge hillslope is steep (Montgomery, 1994; Croke and Mockler, 2001). Both the incisional and dispersive delivery pathways described previously may be connected to the stream network to varying
degrees. These are best described using the framework
and terminology of Wemple et al. (1996), as later used
by Croke and Mockler (2001) and La Marche and Lettenmaier (2001) to describe the nature of road to stream
connectivity. Where overland flow pathways are diffuse,
the degree of connectivity depends on the relative runoff
source strength and the available length of hillslope over
which water can infiltrate (Figure 9). Where the available pathway is long below the runoff source there is
more opportunity to reduce the volume of overland flow
and, in some cases, to disconnect the runoff source from
the stream.
Using several rainfall simulator experiments in forested
catchments in southeastern Australia, the volume of

overland flow required to make a surface connection

at set distances downlsope (5 and 10 m) was measured
(Hairsine et al., 2002; Lane et al., 2004) (Figure 10).
Volume to breakthrough in these studies is defined as
the volume of runoff that may enter an area before
a discharge is observed at the downslope boundary of
that area (Hairsine et al., 2002). The distribution of
measured values was used to predict the length and
volume of overland flow in runoff plumes from road
and track outlets. This approach assumes that segments
of the pathway have a finite capacity to infiltrate water
that is filled before the runoff plume moves on to
the next segment. In this way, the movement of the
plume to the stream edge is controlled by the ratio of
the volume of water leaving the runoff generation area
and the length of the diffuse pathway to the stream.
The proposed model uses a statistical description of
the storage capacity of each pathway segment, so that
predictions of the length and likelihood of the plume
reaching the stream are probabilistic. Predictions of
overland flow plumes reaching a stream segment from
road drainage structures are illustrated in Figure 9 for a
60 km2 catchment in southeastern Australia. Stochastic
predictions of plume length and volume of runoff can be
used to determine runoff and sediment delivery hotspots
within the catchment.
Although the example used here is relatively simple in
terms of modelling catchment hydrological connectivity,
as it focuses on a specific runoff source within catchments
and makes several simplifying but conservative assumptions regarding overland flow processes across a segment
of hillslope, the methodology allows us to focus on quantifying volume to breakthrough. We suggest that, by using
similar experiments in areas of contrasting climate, terrain and soil type, we could build a more accurate pattern of hillslope hydrological connectivity, the thresholds
involved and any observed discontinuities in the downslope transfer of water. For example, it is possible to

Vbt (l)



U. Tyers

Hairsine et al.

Study area

Figure 9. Observed pattern of overland flow plumes following a highintensity storm on a severely burnt hillslope in southeastern Australia
Copyright 2007 John Wiley & Sons, Ltd.

Figure 10. Actual volumes of runoff in litres measured as those required

to observe runoff at the downslope portion of a 5 m length of hillslope.
The box plot of Hairsine et al. (2002) records values determined from
12 sites in New South Wales, Australia, which were later compared with
values for sites in Victoria in Lane et al. (2006). Of note is the fact that
the distributions of Vbt are not significantly different for the two locations
in spite of some obvious differences in soil type, depth and rainfall
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp



calculate the volume to breakthrough for an agricultural

environment where the runoff volume contained within
the concentrated flow paths of furrows and rills is critical in determining any breakthrough of runoff and any
alteration of the flow path direction. Where such furrows
overtop and redirect runoff, the probability of hydrological connectivity may be enhanced. It is probable that
such measurements of runoff volume have been derived
in hydrological experiments, but not specifically with a
view to quantifying hillslope connectivity. By standardizing the definition and application of the term connectivity
we may develop a more accurate understanding of the
factors governing the probability of connection.


The discussion and classification of previous research in
this paper confirms the need to move beyond a dichotomous representation of runoff generation. Research has
demonstrated that there are non-linearities in hillslope
response to rainfall in all environments, and that our
conceptual and computer simulation models now need
to be updated to recognize this. To do this effectively,
hydrologists and geomorphologists need to focus debate
on new models of hillslope response that can be applied
at both the hillslope and catchment scales. The conceptual
framework for connectivity proposed in this paper is one
such approach, and one which supports recent reviews
of hydrological processes (McDonnell, 2003; Ambroise,
The acceptance of connectivity as a model fits well
within the trajectory of current research. Much research
has been undertaken in a range of environments to help
characterize and understand the static element of connectivity, which is the pattern of hillslope response to
rainfall. These investigations have been used to classify elements of the drainage basin as similar surfaces
(HYSS, HRUs, grouped response units), some of which
have been coupled with the dynamic elements of hillslope response to rainfall and used to model catchmentwide runoff and flood production (e.g. Still and Shih,
1985; Karnoven et al., 1999). Ongoing research is also
focusing on developing models of pathways for runoff
and sediment within catchments. Lane et al. (2004) have
developed a methodology for predicting the risk of connectivity based upon the estimation of the probability that
a location in the landscape will be hydrologically connected to the river network. This is based upon the analysis of high-resolution digital topographic data in relation
to distributed rainfall data to optimize and prioritize land
management decisions. In a similar vein, McHugh et al.
(2002) have modelled sediment connectivity (the efficiency of sediment delivery between the land surface and
the river system) for management purposes. Calculation
of the connectivity ratio was based on runoff potential,
slope steepness, slope shape, drainage pattern, and land
use and sediment characteristics. Calculations were raster
based within a geographical information system, utilized
Copyright 2007 John Wiley & Sons, Ltd.

a 50 m DEM and were used to produce annual average

values for a 1 km2 map. Whilst the connectivity ratio provides a quantitative estimate of the fraction of sediment
mobilized by water erosion from the land surface within
a grid cell that reaches a stream, it can only be used as a
relative estimate compared with the values obtained from
the national-scale data set within this particular study.
Other limitations of this approach include issues of data
quality, the use of simplifying assumptions, that not all
sediment sources are included in the estimates, and that
the results include spatial averaging. However, this study
does demonstrate a viable technique that is useful for
catchment management.
Hence, it could be suggested that our understanding
of factors influencing the static aspect of connectivity
is quite well advanced. However, for connectivity to
be agreed as the main concept of runoff response a
more detailed understanding of the dynamic aspects of
connectivity is needed. One particular knowledge gap
appears to be the relationship between storm inputs
and connectivity. Reaney (2003) developed a computer
simulation model of runoff production for semi-arid areas
to analyse the impacts of surface roughness, slope and
vegetation and found that the temporal fragmentation
of high-intensity rainfall is important for determining
the travel distances of overland flow, and hence the
amount of runoff that leaves the slope as discharge.
The results showed that, if the high-intensity rainfall is
fragmented, runoff will be infiltrated a short distance
downslope, whereas longer periods of high-intensity
rainfall allowed runoff to travel further, and hence
become discharge. This work was undertaken for semiarid areas using a distributed, dynamic hydrology model
which considers the hydrological processes relevant for
a semi-arid environment at the temporal scale of a
single storm event. Therefore, storms may have the
same amount of high-intensity rainfall but produce vastly
different amounts of discharge, and this has important
implications for connectivity.
For the proposed framework of connectivity to be
as useful as possible, it needs to be supported by a
quantitative variable that can be measured and estimated
in the field. The method proposed in this paper is the
volume to breakthrough. Although the term has implicit
assumptions, it is an easily understood measurement and
not difficult to test, although there may logistical issues
of water supply in dryland environments. However, the
measure volume to breakthrough inverts the concept that
a large number represents a good connectivity and could
be re-inverted by the following. If V is the volume to
breakthrough per contour width, then the average depth
of storage of water at two points a distance x apart is
h D V/x. This can then be scaled to the soil hydrological
depth z. It follows that z/z C h varies from zero for poor
connectivity with h being infinite, to one for excellent
connectivity where h D 0. This example only considers
a single pair of points, which may be more dynamically
described rather than being prescribed a single number.
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp


This paper has introduced a framework for connectivity
that identifies some of the key factors that affect the
physical linkage of water and, indirectly, sediment in
catchments. We have also proposed and illustrated a
working definition of hydrological connectivity that we
suggest can be measured and thus incorporated into
future studies of runoff generation and sediment delivery.
Connectivity is one way forward in the discussion of postVSA models of hillslope response to rainfall and provides
a basis for integrating processes of runoff generation with
landscape characteristics.
One of the key aspects of connectivity is the combination of both static and dynamic process responses.
After several decades of hydrological research, current
understanding is well developed in terms of the static elements, but we know less about the dynamic components
of hydrological connectivity. This is one significant reason for developing a definition that can be measured and
estimated in the field, but also incorporated into computer
simulation models. Validation needs to be more than just
a comparison with stream hydrographs, because whereas
hydrographs are representative of process interactions,
they do not enable us to discern the complex interplay of
factors that result in flow generation. For connectivity to
be accepted as a viable post-VSA and Hortonian framework, we should acknowledge the benefits experienced
by ecologists in using a standard terminology and use this
to modify how we conceptualize runoff relationships.

Abdul AS, Gillham RW. 1989. Field studies of the effects of the
capillary-fringe on streamflow generation. Journal of Hydrology 112:
(1-2) 1-18.
Abernethy C. 1990. The use of river and reservoir sedimentation
data for the study of regional erosion rates and trends. In
International Symposium on Water Erosion Sedimentation and Resource
Conservation, India.
Abrahams AD, Parsons AJ. 1990. Determining the mean depth of
overland flow in field studies of flow hydraulics. Water Resources
Research 26: 501 503.
Ambroise B. 2004. Variable active versus contributing areas
or periods: a necessary distinction. Hydrological Processes 18:
1149 1155.
Andreassen HP, Halle S, Ims RA. 1996a. Optimal design of movement
corridors in root volesnot too wide and not too narrow. Journal of
Applied Ecology 33: 6370.
Andreassen HP, Ims RA, Steinset OK. 1996b. Discontinuous habitat
corridors: the effect on male root vole movements. Journal of Applied
Ecology 33: 555560.
Archer S. 1990. Development and stability of grasswoody mosaics
in a subtropical parkland Texas, USA. Journal of Biogeography 17:
453 462.
Aryal SK, Mein RG, OLoughlin EM. 2003. The concept of effective
length in hillslopes: assessing the influence of climate and topography
on the contributing area of catchments. Hydrological Processes 17:
131 151.
Auzet AV, Boiffin J, Papy F, Ludwig B, Maucorp J. 1993. Rill erosion
as a function of the characteristics of cultivated catchments in the north
of France. Catena 20: 41 62.
Barling RD. 1992. Saturation zones and ephemeral gullies on arable land
in south-eastern Australia. PhD thesis, The University of Melbourne,
Victoria, Australia.
Belsky AJ. 1995. Spatial and temporal landscape patterns in arid and
semi-arid Africa savannas. In Mosaic Landscapes and Ecological
Copyright 2007 John Wiley & Sons, Ltd.


Processes, Hansson L, Fahrig L, Meeriam G (eds). Chapman and Hall:

London; 3156.
Bergkamp G, Cammeraat LH, Mertinez-Fernandez J. 1996. Water
movement and vegetation patterns on shrubland and an abandoned
field in two desertification threatened areas in Spain. Earth Surface
Processes and Landforms 21: 1073 1090.
Betson RP. 1964. What is watershed runoff? Journal of Geophysical
Research 69: 1541 1552.
Beven K. 1997. Topmodel: a critique. Hydrological Processes 11:
1069 1085.
Beven K, Freer J. 2001. A dynamic TOPMODEL. Hydrological
Processes 15: 1993 2011.
Boer M, Puigdefabregas J. 2005. Effects of spatially structured vegetation
patterns on hillslope erosion in a semiarid Mediterranean environment:
a simulation study. Earth Surface Processes and Landforms 10:
149 167.
Boix-Fayos C,
Calvo-Cases A,
Imeson AC,
Soriano-Soto MD,
Tiemessen IR. 1998. Spatial and short-term temporal variations in
runoff, soil aggregation and other soil properties along a Mediterranean
climatological gradient. Catena 33: 123138.
Boyland EE. 1973. Vegetation of the mulga lands with special reference
to south-western Queensland. Tropical Grasslands 7: 3542.
Bradford JM, Remley PA, Ferris JE. 1987a. Interrill soil erosion
processes: I. Effect of surface sealing on infiltration, runoff and soil
splash detachment. American Journal of Soil Science 51: 1566 1571.
Bradford JM, Ferris JE, Remley PA. 1987b. Interrill soil erosion
processes: II. Relationship of splash detachment to soil properties.
American Journal of Soil Science 51: 1571 1575.
Brooks CP. 2003. A scalar analysis of landscape connectivity. Oikos 102:
433 439.
Bryan RB. 2000. Soil erodibility and processes of water erosion on
hillslope. Geomorphology 32: 385415.
Bryan RB, Yair A. 1982. Perspectives on studies of badland
geomorphology. In Badland Geomorphology and Piping, Bryan RB,
Yair A (eds). GeoBooks: Norwich; 112.
Bull LJ, Kirkby MJ, Shannon J, Dunsford HD. 2003. Predicting
hydrologically similar surfaces (HYSS) in semi-arid environments.
Advances in Environmental Monitoring and Modelling 2: 113.
Bull LJ, Kirkby MJ, Shannon J, Hooke JM. 2000. The variation in
estimated discharge in relation to the location of storm cells in SE
Spain. Catena 38: 191209.
Burt TP. 1997. The hydrological role of buffer zones within the drainage
basin system. In Buffer Zones: Their Processes and Potential in Water
Protection, Haycock NE, Burt TP, Goulding K, Pinay G (eds). Quest
Environmental: UK; 2132.
Burt TP, Haycock NE. 1996. Linking floodplains to rivers. In
Floodplain Processes, Anderson MG, Walling DE, Bates P (eds).
Wiley: Chichester; 461 492.
Burt TP, Pinay G. 2005. Linking hydrology and biogeochemistry in
complex landscapes. Progress in Physical Geography 29: 297316.
Cammeraat LH. 2002. A review of two strongly contrasting
geomorphological systems within the context of scale. Earth Surface
Processes and Landforms 27: 1201 1222.
Cammeraat LH, Imeson AC. 1999. The significance of soilvegetation
patterns following land abandonment and fire in Spain. Catena 37:
107 127.
Cerd`a A. 1995. Spatial distribution of infiltration on the matorral slopes in
a Mediterranean environment, Genoves, Spain. In Desertification in a
European Context: Physical and Socio-Economic Impacts, Fantechi R,
Peter D, Blabanis P, Rubio JL (eds). European Commission: Brussels;
427 436.
Cornet AF, Montana C, Delhoume JP, Lopez-Portillo J. 1991. Water
flows and dynamics of desert vegetation stripes. In Landscape
BoundariesConsequences for Biotic Diversity and Ecological Flows,
Hansen AJ, di Castri F (eds). Springer-Verlag: New York; 327 345.
Costa JE. 1987. A comparison of the largest rainfallrunoff foods in
the United States and the Peoples Republic of China and the world.
Journal of Hydrology 96: 101 115.
Croke J, Mockler S. 2001. Gully initiation and road-to-stream linkage in
a forested catchment, south eastern Australia. Earth Surface Processes
and Landforms 26: 205 217.
Croke J, Mockler S, Fogarty P, Takken I. 2005. Sediment concentration
changes in runoff pathways from a forest road network and the
resultant spatial pattern of catchment connectivity. Geomorphology 68:
257 268.
Davidson C. 1998. Issues in measuring landscape fragmentation. Bulletin
of the Wildlife Society 26: 3237.
De Ploey J. 1984. Hydraulics of runoff and loess loam deposition. Earth
Surface Processes and Landforms 9: 533 539.
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp



Demaynadier PG, Hunter ML. 1998. Effects of silvicultural edges on

the distribution and abundance of amphibians in Maine. Conservation
Biology 12: 340352.
Desmet PJJ, Govers G. 1997. Two-dimensional modelling of the withinfield variation in rill and gully geometry and location related to
topography. Catena 29: 283 306.
Dick G, Anderson RS, Sampson DE. 1997. Controls on flash flood
magnitude and hydrograph shape, Upper Blue Hills badlands, Utah.
Geology 25: 4548.
Dunkerley DL. 1999. Banded chenopod shrublands of arid Australia:
modelling responses to interannual rainfall variability with cellular
automata. Ecological Modelling 121: 127 138.
Dunne T, Black RD. 1970. Partial area contributions to storm runoff
in a small New England watershed. Water Resources Research 6:
1296 1311.
Fiedler FR, Frasier GW, Ramirez JA, Ahuja LR. 2002. Hydrologic
response of grasslands; effects of grazing, interactive infiltration and
scale. Journal of Hydrologic Engineering 7: 293301.
Fitzjohn C, Ternan JL, Williams AG. 1998. Soil moisture variability in a
semi-arid gully catchment: implications for runoff and erosion control.
Catena 32: 5570.
Flugel WA. 1995. Delineating hydrological response units by geographical information system analyses for regional hydrological modelling
using PRMS/MMS in the drainage basin of the River Brol, Germany.
Hydrological Processes 9: 423 436.
Fryirs K, Brierley G. 2000. A geomorphic approach to the identification
of river recovery potential. Physical Geography 21: 244 277.
Golden DM, Crist TO. 2000. Experimental effects of habitat fragmentation on rove beetles and ants: patch area or edge. Oikos 90: 525538.
Govers G. 1991. Rill erosion on arable land in central Belgiumrates,
controls and predictability. Catena 18: 133 155.
Govers G, Takken I, Helming K. 2000. Soil roughness and overland flow.
Agronomie 20: 131146.
Habbad NM. 1999a. Corridor and distance effects in inter-patch
movement: a landscape experiment with butterflies. Journal of Applied
Ecology 9: 612 622.
Habbad NM. 1999b. Corridor use predicted from behaviours at habitat
boundaries. American Naturalist 153: 215 227.
Habbad NM, Rosenberg DK, Noon BR. 2000. On experimentation and
the study of corridors: response to Beir and Noss. Conservation Biology
14: 1543 1545.
Hairsine PB, Croke JC, Matthews H, Fogarty P, Mockler SP. 2002.
Modelling plumes of overland flow from roads and logging tracks.
Hydrological Processes 16: 2311 2327.
Harvey AM. 2001. Coupling between hillslope and channels in upland
fluvial systems; implications for landscape sensitivity, illustrated from
the Howgill Fells, NW England. Catena 42: 225250.
Harvey AM. 1996. Holocence hillslope gully systems in the Howgill
Fells, Cumbria. In Advances in Hillslope Processes, vol. 2,
Anderson MG, Brooks SM (eds). Wiley: Chichester; 247270.
Helming K, Romkens MJM, Prasad SN. 1998. Surface roughness related
processes of runoff and soil loss: a flume study. Journal of the Soil
Science Society of America 62: 243250.
Hewlett JD, Hibbert AR. 1967. Factors affecting the response of small
watersheds to precipitation in humid areas. In Forest Hydrology,
Sopper WE, Lull HW (eds). Pergamon: New York; 345360.
Hooke J. 2003. Coarse sediment connectivity in river channel systems: a
conceptual framework and methodology. Geomorphology 56: 7994.
Imeson AC, Prinsen HAM. 2004. Vegetation patterns as biological
indicators for identifying runoff and sediment source and sink areas for
semi-arid landscapes in Spain. Agriculture Ecosystems & Environment
104: 333 342.
Imeson AC, Verstraten JM. 1988. Rills on badlands slopes; a physicochemically controlled phenomenon. In Geomorphic Processes in
Environments with Strong Seasonal Contrasts, Vol. I: Hillslope
Processes, Imeson AC, Sala M (eds). Catena Supplement 12. Catena
Verlag: Reiskirchen; 139 150.
Imeson AC, Verstraten JM, van Mulligan EJ, Sevink J. 1992. The
effects of fire and water repellency on infiltration and runoff under
Mediterranean type forest. Catena 19: 345361.
Karnoven T, Koivusalo H, Jauhianinen M, Palko J, Weppling K. 1999.
A hydrological model for predicting runoff from different land use
areas. Journal of Hydrology 217: 253265.
Kirkby MJ, Bracken LJ, Reaney S. 2002. The influence of landuse, soils
and topography on the delivery of hillslope runoff to channels in SE
Spain. Earth Surface Landforms and Processes 27: 1459 1473.
La Marche JL, Lettenmaier DP. 2001. Effects of forest roads on flood
flows in the Deschutes River, Washington. Earth Surface Processes
and Landforms 26: 115 134.
Copyright 2007 John Wiley & Sons, Ltd.

Lal R. 1990. Soil Erosion in the Tropics: Principles and Management.

McGraw-Hill: New York.
Lal R. 1997. Soil degradative effects of slope length and tillage methods
on alfisols in western Nigeria. 1. Runoff, erosion and crop response.
Land Degradation and Development 8: 201219.
Lane SN, Brookes CJ, Kirkby AJ. 2004. A network-index based version
of TOPMODEL for use with high-resolution digital topographic data.
Hydrological Processes 18: 191201.
Lane PJ, Hairsine PB, Croke JC, Takken I. 2006. Quantifying diffuse
pathways for overland flow between roads and streams of the mountain
ash forests of central Victoria, Australia. Hydrological Processes 20:
1875 1884.
Lasanta T, Garcia-Ruiz JM, Perez-Rontome C, Sancho-Marcen C. 2000.
Runoff and sediment yield in a semi-arid environment: the effect of
land management after farmland abandonment. Catena 38: 265 278.
Lavee H, Imeson AC, Sarah P. 1998. The impact of climate change on
geomorphology and desertification along a Mediterranean-arid transect.
Land Degradation & Development 9: 407422.
Luce CH, Cundy TW. 1994. Parameter identification for a runoff model
for forest roads. Water Resources Research 30: 1057 1069.
Ludwig B, Auzet AV, Boiffin J, Papy F, King D, Chadoeuf J. 1996. Etats
de surface, structure hydrographique et e rosion en rigole de bassins
versants cultives du Nord de la France. Etude et Gestion des Sols 3:
Ludwig B, Boiffin J, Chadoeuf J, Auzet AV. 1995. Hydrological
structure and erosion damage caused by concentrated flow in cultivated
catchments. Catena 25: 227252.
Ludwig JA, Tongway DJ. 1995. Spatial organisation of landscapes and
its function in semi-arid woodlands, Australia. Landscape Ecology 10:
Ludwig JA, Tongway DJ, Eager RW, Williams RJ, Cook GD. 1999b.
Fine-scale vegetation patches decline in size and cover with increasing
rainfall in Australian savannas. Landscape Ecology 14: 557 566.
Ludwig JA, Tongway DJ, Marsden SG. 1999a. Stripes, strands or
stipples: modelling the influence of three landscape banding patterns on
resource capture and productivity in semi-arid woodlands, Australia.
Catena 37: 257273.
Ludwig JA, Wiens J, Tongway DJ. 2000. A scaling rule for landscape
patches and how it applies to conserving soil resources in savannas.
Ecosystems 3: 8297.
Ludwig JA, Wilcox BP, Breshears DD, Tongway DJ, Imeson AC. 2005.
Vegetation patches and runoff-erosion as interacting ecohydrological
processes in semiarid landscapes. Ecology 86: 288 297.
Lyford FP, Qashu HK. 1969. Infiltration rates as affected by desert
vegetation. Water Resources Research 5: 1373 1376.
Mabbutt JA, Fanning PC. 1987. Vegetation banding in arid Western
Australia. Journal of Arid Environments 12: 4159.
Mauchamp A, Rambal S, Lepart J. 1994. Simulating the dynamics
of a vegetation mosaic: a spatialized functional model. Ecological
Modelling 71: 107130.
McDonnell JJ. 2003. Where does water go when it rains? Moving
beyond the variable source area concept of rainfall-runoff response.
Hydrological Processes 17: 1869 1875.
McHugh M, Wood G, Walling DE, Morgan R, Zhang Z, Anthony S,
Hutchins M. 2002. Prediction of sediment delivery to watercourses
from land, Phase II . R&D Technical Report no. P2-209, National Soil
Resource Centre, Cranfield University.
Megahan WF. 1983. Hydrological effects of clear cutting and wildfire on
steep granitic slopes in Idaho. Water Resources Research 19: 811819.
Metzger JP, Decamps H. 1997. The structural connectivity threshold:
a hypothesis in conservation biology at the landscape scale. Acta
Oecologica 18: 112.
Michaelides K, Wainwright J. 2002. Modelling the effects of hillslope
channel coupling on catchment hydrological response. Earth Surface
Processes and Landforms 27: 1441 1457.
Montana C. 1992. The colonization of bare areas in two-phase mosaics
of an arid ecosystem. Journal of Ecology 80: 315327.
Montgomery DR. 1994. Road surface drainage, channel initiation, and
slope instability. Water Resources Research 30: 1925 1932.
Morgan RPC. 1995. Soil Erosion and Conservation. Longman.
Moussa R, Voltz M, Andrieux P. 2002. Effects of the spatial organization
of agricultural management on the hydrological behaviour of a farmed
catchment during flood events. Hydrological Processes 16: 393 412.
Nicolau JM, Sole A, Puidefabregas J, Gutierrez L. 1996. Effects of
soil and vegetation on runoff along a catena in semi-arid Spain.
Geomorphology 15: 297309.
ONeill RV, Milne BT, Turner MG, Gardner RH. 1988. Resource
utilization scales and landscape patterns. Landscape Ecology 2: 6369.
Hydrol. Process. 21, 1749 1763 (2007)
DOI: 10.1002/hyp


Oostwould Wijdenes D, Poesen J, Vandekerckhove L, De Luna E. 1997.
Chiseling effects on the vertical distribution of rock fragments in the
tilled layer of a Mediterranean soil. Soil and Tillage Research 44:
55 66.
Pinay G, Ruffinoni C, Wondzell S, Gazelle F. 1998. Change in
groundwater nitrate concentration in a large river floodplain:
denitrification, uptake or mixing? Journal of the North American
Benthological Society 17: 179189.
Pitlick J. 1994. Relation between peak flow, precipitation, and
physiography for five mountainous regions in western USA. Journal
of Hydrology 53: 219 240.
Poesen J. 1984. The influence of slope gradient on infiltration rate
and Hortonian overland flow volume. Zeitschrift fur Geomorphologie
Supplement Band 49: 117 131.
Pringle CM. 2001. Hydrological connectivity and the management of
biological reserves: a global perspective. Ecological Applications 11:
981 998.
Pringle CM. 2003. What is hydrological connectivity and why is it
ecologically important? Hydrological Processes 17: 2685 2689.
Puidefabregas J. 2005. The role of vegetation patterns in structuring
runoff and sediment fluxes in drylands. Earth Surface Processes and
Landforms 30: 133 147.
Puidefabregas J, Barrio G, Boer M, Gutierrez L, Sole A. 1998. Differential responses of hillslope and channel elements to rainfall events in
semi-arid areas. Geomorphology 23: 337 351.
Reaney S. 2003. Modelling runoff generation and connectivity for semiarid hillslopes and small catchments. PhD thesis, Leeds University.
Reid I, Frostick LE. 1997. Channel form, flows and sediments in deserts.
In Arid Zone Geomorphology: Process, Form and Change in Drylands,
Thomas DSG (ed.). Wiley: Chichester; 205 229.
Reid LM, Dunne T. 1984. Sediment production from forest road surfaces.
Water Resources Research 20: 1753 1761.
Ritter DF, Kochel RC, Miller JR. 1999. The disruption of Grassy
Creek: implications concerning catastrophic events and thresholds.
Geomorphology 29: 323338.
Romkens MJM, Helming K, Prasard SN. 2001. Soil erosion under
different rainfall intensities, surface roughness and soil water regimes.
Catena 46: 103123.
Sanchez G, Puidefabregas J. 1994. Interactions of plant growth
and sediment movement on slopes in a semi-arid environment.
Geomorphology 9: 243 260.
Schick AP. 1988. Hydrologic aspects of floods in extreme arid
environments. In Flood Geomorphology, Baker VR, Kochel RC,
Patton PC (eds). Wiley: New York; 189 203.
Schumaker NH. 1996. Using landscape idiocies to predict habitat
connectivity. Ecology 77: 1210 1225.
Sharma ML, Luxmore RJ, DeAngelis R, Ward RC, Yeh GT. 1987.
Subsurface water flow simulated for hillslopes with spatially
dependent soil hydraulic characteristics. Water Resources Research 23:
1523 1530.
Singer MJ, Le Bissonnais YL. 1998. Importance of surface sealing in the
erosion of some soils from a Mediterranean climate. Geomorphology
24: 79 85.
Slattery MC, Bryan RB. 1992. Hydraulic conditions for rill incision under
simulated rainfall: a laboratory experiment. Earth Surface Processes
and Landforms 17: 127146.
Slatyer RO. 1961. Methodology of a water balance study conducted on a
desert woodland (Acacia aneura) community. Vegetation 91: 105120.
Souch`ere V, King D, Darroussin J, Papy F, Capillon A. 1998. Effects of
tillage on runoff direction: consequences on runoff contributing area
within agricultural catchments. Journal of Hydrology 206: 256267.
Sole-Benet A, Calvo A, Cerd`a A, Lazar R, Piny R, Barber J. 1997.
Influence of micro-relief patterns and plant cover on runoff related
processes in badlands from Taverns (SE Spain). Catena 31: 2338.
Stamps JA, Buchner M, Krishnan VV. 1987. The effects of edge
permeability and habitat geometry on emigration from patches of
habitat. American Naturalist 129: 533552.
Still DA, Shih SF. 1985. Using Landsat data to classify land use for
assessing the basin wide runoff index. Water Resources Bulletin 21:
931 940.
Tabacchi E, Lambs L, Guilloy H, Planty-Tabacchi AM, Muller E,
Decamps H. 2000. Impacts of riparian vegetation on hydrological
processes. Hydrological Processes 14: (16 17) 2959 2976.

Copyright 2007 John Wiley & Sons, Ltd.


Takken I, Govers G, Steegen A. 2001. The prediction of runoff flow

directions on tilled fields. Journal of Hydrology 248: 113.
Taylor PD, Fahrig L, Henein K, Merriam G. 1993. Connectivity is a vital
element of landscape structure. Oikos 68: 571573.
Thiery JM, dHerb`es JM, Valentin C. 1995. A model simulating the
genesis of banded vegetation patterns in Niger. Journal of Ecology
83: 497 507.
Thornes JB. 1976. Semi-arid erosional systems: case studies from Spain.
Geographical Papers No. 7, London School of Economics.
Thornes JB. 2001. Environmental crises in the Mediterranean. In
Geography, Environment and Development in the Mediterranean,
King R, de Mas P, Mansvelt Beck J (eds). Academic Press: Brighton;
261 280.
Tongway DJ, Ludwig JA. 1990. Vegetation and soil patterning in semiarid mulga lands of eastern Australia. Journal of Ecology 15: 2334.
Truman CC, Wauchope RD, Sumner HR, Davis JG, Gasho GH,
Hook JE, Chandler LD, Johnson AW. 2001. Slope length effects on
runoff and sediment delivery. Journal of Soil and Water Conservation
56: 249 256.
Valentin C, dHerb`es JM, Poesen J. 1999. Soil and water components of
banded vegetation patterns. Catena 37: 124.
Van de Giesen NC, Stomph TJ, de Ridder N. 2000. Scale effects of
Hortonian overland flow and rainfall-runoff dynamics in a West African
catena landscape. Hydrological Processes 14: 165175.
Wainwright J, Parsons AJ. 2002. The effect of temporal variations in
runoff on scale dependency in runoff coefficients. Water Resources
Research 38: 1267 1271.
Walling DE. 1999. Linking land user, erosion and sediment yields in river
basins. Hydrobiologia 410: 223 240.
Wemple BC, Jones JA. 2003. Runoff production on forest roads in a
steep, mountain catchment. Water Resources Research 39: 117.
Wemple BC, Jones JA, Grant GE. 1996. Channel network extension
by logging roads in two basins, Western Cascades, Oregon. Water
Resources Bulletin 32: 1195 1207.
Whitcomb RF, Lynch JJ, Klimkeiwwicz MK, Robbins CS, Whitcomb BL, Bystrak AD. 1981. Effects of forest fragmentation on avifauna of the eastern deciduous forest. In Forest Island Dynamics in
Man-Dominated Landscapes, Burgess RL, Sharpe DM (eds). SpringerVerlag: New York; 125 205.
Wiens JA. 1995. Landscape mosaics and ecological theory. In
Mosaic Landscapes and Ecological Processes, Hansson L, Fahrig L,
Meeriam G (eds). Chapman and Hall: London; 126.
Wiens JA, Schooley RL, Weeks RD. 1997. Patchy landscapes and animal
movements: do beetles percolate? Oikos 78: 257264.
Wilcox B, Newman BD, Brandes D, Davenport DW, Reid K. 1997.
Runoff from a semi-arid ponderosa pine hillslope in New Mexico.
Water Resources Research 33: 2301 2314.
Wilcox B, Wood M, Tromble J. 1998. Factors influencing infiltrability
of semi-arid mountain slopes. Journal of Rangeland Management 41:
197 206.
Wolman MG, Gerson R. 1978. Relative scales of time and effectiveness
of climate in watershed geomorphology. Earth Surface Processes and
Landforms 3: 189208.
Yair A. 1992. The control of headwater area on channel runoff in a small
arid watershed. In Overland Flow: Hydraulics and Erosion Mechanics,
Parsons AJ, Abrahams AD (eds). UCL Press: London; 5368.
Yair A, Kossovsky A. 2002. Climate and surface properties: hydrological
response of small semi-arid watersheds. Geomorphology 42: 4357.
Yair A, Lavee H. 1985. Runoff generation in arid and semiarid zones.
In Hydrological Forecasting, Anderson GM, Burt TP (eds). Wiley:
Chichester; 183220.
Yair A, Sharon D, Lavee H. 1980. Trends in runoff and erosion processes
over an arid limestone hillside, northern Negev, Israel. Journal of
Hydrology 25: 243255.
Ziegler AD, Sutherland RA, Giambelluca TW. 2000. Partitioning total
erosion on unpaved roads into splash and hydraulic components: The
roles of interstorm surface preparation and dynamic erodibility. Water
Resources Research 36: 2787 2791.

Hydrol. Process. 21, 1749 1763 (2007)

DOI: 10.1002/hyp