See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/227042307

Perspectives on biogeomorphology, ecosystem engineering and self-

organisation in island-braided fluvial ecosystems

Article  in  Aquatic Sciences · October 2009

DOI: 10.1007/s00027-009-9182-6

CITATIONS READS

90 186

3 authors:

Robert A Francis Dov Corenblit

King's College London Université Clermont Auvergne
94 PUBLICATIONS   2,298 CITATIONS    93 PUBLICATIONS   2,738 CITATIONS   

SEE PROFILE SEE PROFILE

Peter J. Edwards
ETH Zurich
321 PUBLICATIONS   16,308 CITATIONS   

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Increased competitive abilities in introduced species View project

Consilpop View project

All content following this page was uploaded by Robert A Francis on 17 November 2015.

The user has requested enhancement of the downloaded file.

Aquat. Sci.
DOI 10.1007/s00027-009-9182-6 Aquatic Sciences
 Birkhuser Verlag, Basel, 2009

Research Article

Perspectives on biogeomorphology, ecosystem engineering and

self-organisation in island-braided fluvial ecosystems
Robert A. Francis1,*, Dov Corenblit1,2 and Peter J. Edwards3

1
Department of Geography, Kings College London, Strand, London, WC2R 2LS, United Kingdom
2
Clermont Universit, GEOLAB – Laboratoire de gographie physique et environnementale, UMR 6042
CNRS/UBP, Maison des Sciences de lHomme, 4 rue Ledru, 63057 Clermont-Ferrand Cedex 1, France
3
Institute of Integrative Biology, CHN H 12.1, Universittstrasse 16, 8092 Zrich, Switzerland

Received: 25 October 2008; revised manuscript accepted: 9 February 2009

Abstract. Using the River Tagliamento, Italy, as an
example, we examine the role of self-organisation in
the formation and dynamics of vegetated islands in
fluvial ecosystems. We consider how various biogeo-
morphic processes, such as feedbacks between tree
growth and sedimentation, influence island self-as-
sembly, as well as the potential influences of island
landforms on resource distribution and shifts in
ecosystem state. Despite the abundance of island
landforms of different sizes and ages in island-braided
reaches along the River Tagliamento, island formation
is only found within a specific hydrological and
sedimentary envelope, and depends upon a delicate
balance of biotic-abiotic feedbacks. As a result, island
landforms tend to be lost when river functioning is
altered by human interventions. We argue that the
specific biogeomorphic processes and self-organisa-
tion associated with river island dynamics offer an
example of biogeomorphic inheritance, in which
reciprocal feedbacks between species and geomorphic
processes favour engineer species and promote the
future development of the landforms. Thus, islands
represent extended phenotypes – or external expres-
sions of genetic traits – of key riparian ecosystem
engineers. This capacity to modify the physical
environment has important implications for landform
evolution and riparian biodiversity. In conclusion, we
propose several topics that merit investigation to
improve our understanding of the biogeomorphology
and self-organisation of river island systems.

Key words. Self-organisation; river island; ecological engineer; biogeomorphology; River Tagliamento.

Introduction between factors controlling the cohesiveness of

The differing morphologies of fluvial systems are
physical expressions of particular functional states
resulting from interactions between the hydrogeo-
morphic and biological ecosystem components. In
particular, the fluvial style depends on the balance
* Corresponding author phone: +44 (0)20 7848 1233;
fax: ++44 (0)20 7848 2287; e-mail: robert.francis@kcl.ac.uk
Published Online First: 2009
landforms (sediment texture and structure, vegeta-
tion roots and roughness) and those inducing remo-
bilisation (i.e. water flow). Braided gravel-bed sys-
tems, for example, are characterised by high stream
power, variable discharge, erodible banks and wide
shallow channels; they also have wide active zones
that may or may not support patches of vegetation,
wood jams or river islands (e.g. Schumm, 2005;
Charlton, 2008). Wandering or transitional gravel-
bed systems are also associated with high loads of
2 R. A. Francis, D. Corenblit and P. J. Edwards
predominantly coarse sediments, but stream powers
are lower, leading to greater sediment aggradation,
fewer, more stable channels, and bars that support
wood jams and are frequently vegetated (e.g. Charl-
ton, 2008). Thus, these contrasting systems are
different expressions of the same basic ecosystem
components and processes. However, a change in
ecosystem state may occur when there is a total or
partial interruption to component structures and
dynamics; for example, during the last century
considerable morphological and ecological changes
have been observed following flood regulation,
gravel mining, and installation of dams, weirs and
bank protections (Johnson, 1994, 1997; Bravard et
al., 1997; Petts and Gurnell, 2005).
In Europe, island braided systems are now rare due
to extensive river regulation and engineering over the
last 700 years (Petts, 1989; Gurnell and Petts, 2002;
Francis et al., 2008a). Extant river islands within
braided systems have been noted as indicators of
ecosystem integrity (Ward et al., 2000). Certainly
evidence exists that many European and North
American rivers previously supported river islands,
but that the ecosystem components (e.g. intact ripar-
ian woodlands, large populations of species capable of
vegetative reproduction) and processes (e.g. dynamic
hydrology, flow variability, sediment transport, tree
entrainment, deposition and establishment) that allow
such systems to develop have been compromised,
leading to loss of these landforms from river systems
(Gurnell and Petts, 2002; Ward et al., 2002; Tockner et
al., 2003; Francis et al., 2008a). Recently, a suite of
studies in aquatic ecology and fluvial geomorphology
has focused on fluvial systems that maintain river
islands (e.g. Osterkamp, 1998; Edwards et al., 1999;
Gurnell and Petts, 2002; Tockner, 2003; Karaus et al.,
2005; Francis et al., 2008b). Such investigations are
important to increase our understanding of ecosystem
functioning, dynamics and integrity, and to support
appropriate river conservation and restoration meth-
odologies (see Tockner et al., 2003).
Recent research has highlighted the important role
of feedbacks between organisms and physical proc-
esses in determining the spatial structure and dynam-
ics of ecosystems, both terrestrial and aquatic (Hast-
ings et al., 1993; Phillips, 1999). Such feedbacks occur
when an organism regulates some aspect of its growth
or reproduction in response to one or more physical
processes, or when physical processes are to some
extent regulated by the activities of organisms. In
practice, such feedbacks are often mutual: for exam-
ple, some riparian plants are not only able to grow
through deposited sediments but grow faster where
there is sedimentation. The increased biomass may
then act as a roughness element that further increases
River islands and self-organisation
sediment deposition, in a reciprocal (mutual) feed-
back (e.g. Sand-Jensen, 1998; Kent et al., 2001;
Francis, 2006).
The term biogeomorphology has been used to
describe situations in which such feedbacks lead to the
development, maintenance or modification of geo-
morphic landforms (see Viles, 1988); and the term
functional ecomorphology may also be used to
encompass more general functional processes, such
as soil development and biogeochemical cycling at
broader spatial scales (see Fisher et al., 2007).
Examples of biogeomorphic interactions and feed-
backs have been demonstrated in many ecosystems,
from interactions between plant growth and repro-
duction and sediment deposition and stabilisation in
coastal dunes (e.g. Kent et al., 2001) to biofilms
stabilising sediments with polymers to create ridge
formations in intertidal mudflats (Rietkerk and van de
Koppel, 2008).
There is increasing evidence that ecosystem state
transitions due to biogeomorphic interactions are
commoner than was previously supposed (Pahl-Wostl,
1995; Dent et al., 2002; Francis, 2009). In the case of
natural fluvial ecosystems, for example, it has been
shown that state transitions are influenced not only by
changes or disruption of key hydrogeomorphological
drivers (e.g. regulation of flows of water and sedi-
ment), but also by feedbacks between flow regimes,
sediment, and vegetation dynamics. These feedbacks
result in characteristic biogeomorphic patterns and
strongly influence ecosystem functioning and biodi-
versity (e.g. Edwards et al., 1999; Dent et al., 2002;
Ward et al., 2002; Gurnell and Petts, 2002; Francis,
2006; Corenblit et al., 2007a). Often the expression of
particular ecosystem patterns is due to feedbacks by
one or more engineering species upon physical
processes, and can be considered as self-organisation,
(Rietkerk and van de Koppel, 2008). The phenomena
of self-organisation and self-organised criticality –
implying that a system develops to a point at which
small events lead to drastic change (Bak et al., 1988;
Bak, 1996; Hooke, 2007) – have been studied inten-
sively in both physical and social systems (e.g. Baas,
2007; Turcotte, 2007; Coco and Murray, 2007; Keller,
2008). In this paper we are concerned with the
implications of biogeomorphic engineering on self-
organisation within fluvial island-braided systems. We
propose that within natural alpine fluvial ecosystems
keystone ecosystem engineers (sensu Jones et al.,
1994) may have a significant effect on river dynamics
that deserves further investigation (e.g. Edwards et
al., 1999; Gurnell et al., 2005; Gurnell and Petts,
2006).
Although the mechanisms of island formation on
the River Tagliamento and other systems have been
Aquat. Sci.
explained in detail elsewhere, this paper aims to place
these mechanisms within the context of self-organisa-
tion at the landform and ecosystem scales, as well as
noting the significance of these processes for the
future existence and evolution of such landforms and
their engineers. We aim to encourage a better under-
standing of the biogeomorphic linkages involved, the
variability and dynamics of such linkages over varying
spatio-temporal scales, and the implications of both
self-organisation and the interruption of self-organ-
isation on functional and morphological states of
fluvial braided ecosystems. Finally, we note that
further research is needed as a basis for developing
strategies to preserve or restore the natural dynamics
of these systems. We propose several topics that merit
investigation.
The nature of ecosystem self-organisation
Self-organisation in general refers to the develop-
ment of organised structures that are far from
thermodynamic equilibrium as a result of interac-
tions occurring at a much smaller scale within the
system. Although self-organisation has been studied
in a range of physical and abstract systems – by
physicists, mathematicians, computer scientists and
geomorphologists to name just a few (see e.g.
Murray, 1989; Kauffman, 1993; Murray and Fonstad,
2007) – the concept is potentially relevant to bio-
logical systems, from the molecular to the ecosystem
level. It was applied to ecosystems as it was realised
that feedbacks between different components of an
ecosystem (e.g. between species or between species
and habitat) often induce a hierarchy of self-rein-
forcing states (Perry et al., 1989; Perry, 1995; Phillips,
1995). Self-organisation within ecosystems may be
considered as the emergence of large-scale structures
or patterns from local-scale interactions between
ecosystem components, usually associated with lo-
calised feedback loops (e.g. Camazine et al., 2001;
Rietkerk and van de Koppel, 2008). Such patterns
have been described at spatio-temporal scales rang-
ing from individual plants to extensive landscapes
(e.g. Aguiar and Sala, 1999; Wetzel et al., 2005;
Rietkerk and van de Koppel, 2008). In this paper, we
are mainly concerned with the causes and expression
of self-organisation in fluvial ecosystems at scales
ranging from the landform to landscape.
Self-organisation of plant or animal communities,
with or without associated landforms, has been found
to produce patterns at the landscape scale in a range of
ecosystems (see Table 1). In each of these ecosystems,
an initial emergent pattern based on the distributions
of key environmental variables is reinforced and
Research Article 3
maintained by positive feedbacks in resource acquis-
ition at the local scale, and negative feedbacks at
larger scales. In arid ecosystems, for example, the
tiger bush stripe and leopard bush spot patterns
described by Aguiar and Sala (1999) and Rietkerk et
al. (2002) are caused by vegetation establishing in
areas where soil moisture availability is locally higher.
Once vegetation establishes, a positive feedback
occurs, with higher plant biomass leading to more
humid conditions, increased infiltration of water into
the soil, and a redistribution of overland flow through
and alongside the vegetation (Rietkerk et al., 2002). In
turn, this leads to higher growth rates and biomass
production of the vegetation, reinforcing the feed-
back.
This local feedback occurs at the expense of
resource distribution throughout the rest of the land-
scape, as stands of vegetation effectively act as sinks,
drawing resources into the vegetation patches and
limiting plant establishment elsewhere (see Fig. 1;
Rietkerk and van de Koppel, 2008). The combination
of a short-distance positive feedback (abundant
resources within and around the vegetation patch)
and long-distance negative feedback (reduced resour-
ces away from the vegetation patch) is a prerequisite
for self-organisation within ecosystems (Rietkerk and
van de Koppel, 2008). It has been suggested that
ecosystems exhibiting self-organised patterns may be
less vulnerable to allogenic disturbances (e.g. fire
spread, pathogen invasion) due to their high biotic and
abiotic heterogeneity. However, the stability and
resilience characteristics of such ecosystems have
scarcely been studied (Forman, 1995; Drever et al.,
2006; Rietkerk and van de Koppel, 2008).
Figure 1. Scale-dependent feedback in ecosystem processes. Plant
communities increase resource availability and/or decrease stress
at the local scale leading to facilitation of plant survival and growth,
potentially in association with one or more mechanisms of
ecosystem engineering, while decreasing resources and/or increas-
ing stress at longer distances. Modified from Rietkerk and van de
Koppel, 2008.
4 R. A. Francis, D. Corenblit and P. J. Edwards River islands and self-organisation

Table 1. Examples of landscape patterns formed by self-organisation in real ecosystems, noting the scale-dependent feedback mechanisms
responsible for pattern formation. Summarised from Gumbricht et al., 2004; Wetzel et al., 2005; Van Hulzen et al., 2007; Larsen et al., 2007;
Rietkirk and van de Koppel, 2008.
Ecosystem Type of Reason for landscape patterning Scale-dependant feedback mechanism
landscape
pattern
Arid Stripes, Higher soil water availability in soil patches Higher vegetation density allows higher local water
labyrinths, infiltration into the soil, more humid local
spots microclimate, and redistribution of overland flow
Wetlands String Vegetation establishing on less waterlogged soils in Transpiration of the denser vegetation helps to draw
strings perpendicular to slopes then leads to ponding nutrients from groundwater, while interception of
behind the strings due to reduced hydraulic surface flows help to accumulate nutrients on the
conductivity in densely vegetated areas surface, at the expense of less vegetated areas
Savanna Spots Isolated trees and shrubs related to patches of higher Plants accumulate nutrients for themselves and highly
soil nutrient content local vegetation while competing with each other
through extensive shallow root systems that extend
away from the plants
Intertidal Stripes, Diatom biofilms establish on elevated ridges or Biofilms produce extracellular polymeric substances
mudflats patches patches within mudflats that increase sediment cohesion and stability and lead
to sedimentation and localised increases in elevation.
This diverts flowing water into surrounding areas
(runnels), preventing diatom establishment at lower
elevations.
Marsh Spots Regularly spaced clumps of Carex stricta grow on C. stricta produces abundant dead material that
tussocks higher elevations within freshwater wetlands accumulates in gaps between plants, preventing
seedling establishment in these areas via shading.
Growth of C. stricta on higher elevations keeps it above
the layer of dead material and reduces shading.
Everglades Patches Trees establish on higher elevations within freshwater Phosphorus is redistributed to tree islands by
tree islands wetlands subsurface flows generated by higher
evapotranspiration on islands, and deposition from
both the atmosphere and guano, as islands are useful
bird habitat.
Okavango Patches Termites form mounds above the level of flooding in Termite mounds accumulate nutrients that support
Delta suitable locations plant establishment and growth. Plants attract
island browsers which contribute further nutrients and
promote island growth.

Biogeomorphology and self-organisation of river

islands: perspectives from the River
Tagliamento, NE Italy
In some ecosystems, self-organisation of vegetation
communities is associated with biogeomorphic engi-
neering of physical landform features by one or a few
key plant species, though detailed studies are limited
(see Hughes, 1997; Gurnell et al., 2005; Pettit and
Naiman, 2005; Wetzel et al., 2005; Stallins, 2006;
Heffernan, 2008). In this context, biogeomorphic
engineering refers to a specific kind of landform-
building due to the presence of ecosystem engineers,
defined as organisms that directly or indirectly make
resources available to other species and thereby
modify, maintain or create habitat (Jones et al.,
1994). This interpretation places the engineering
species as central to the process, and argues that the
landform creation is formed via bottom-up processes
driven by the presence of particular species. However,
island landforms may also be placed within the
concept of functional ecomorphology (Fisher et al.,
2007), implying that the emergence of island forms is a
result of a range of processes including not only the
influence of organisms but also various geomorphic
processes. Since we are particularly concerned with
the traits of engineering species, we prefer to use the
term biogeomorphic engineering, thereby emphasis-
ing the link with the ecological literature on ecosys-
tem engineering. As is the case in many complex
adaptive systems (including ecosystems), we regard
engineering and self-organisational processes as un-
planned and leading to self-organisation without
intent (see Walker et al., 2004).
One fluvial ecosystem in which biogeomorphic
ecosystem engineering appears to be an essential
part of the self-organisation process is the island-
braided gravel bed River Tagliamento in Northeast
Italy (see Gurnell et al., 2001; Tockner et al., 2003;
Fig. 2). In this system, large pieces of wood deposited
on gravel bars, in particular whole trees that have the
capacity to survive vegetatively (see Edwards et al.,
1999; Gurnell et al., 2005; Opperman et al., 2008),
form the nucleus for the accumulation of a fine-scale
ecological community. Following initial deposition
during the falling limb of a flood event, both dead and
Aquat. Sci.
living riparian trees create an area of increased
sediment stabilisation and hydraulic roughness on
the gravel bar, leading to further changes during
subsequent inundation events. Thus, scour areas –
typically in the form of a crescent-shaped scour pool
– are formed in the surrounding bar as flows are
redirected around the root bole and forced down-
wards into the bar (see Abbe and Montgomery, 1996
for a more detailed description of this process), while
organic detritus, fine sediments and organisms (e.g.
plant propagules, fish, invertebrates) are trapped in
and around the deposited tree (e.g. Karaus et al.,
2005).
Figure 2. Aerial photo of a pre-alpine island-braided reach of the
River Tagliamento near Flagogna, illustrating patterning of
pioneer and building (self-assembled) islands within the active
zone, surrounded by a gravel matrix at low flow. Image taken from
Google Earth.
In this way, a stranded tree creates a distinct
arrangement of microhabitats on the bar, with scour
features generally positioned upstream and fine sedi-
ment extending in a plume downstream, and with
organic material deposited throughout (see Abbe and
Montgomery, 1996; Edwards et al., 1999; Fig. 3). These
engineered microsites form sites suitable for further
plant establishment from propagules deposited either
at the same time or during subsequent inundation
events (which typically occur 3 – 4 times per year but are
highly variable; Tockner et al., 2003). This deposition of
propagules has been directly linked to changes in plant
diversity during the first three years following tree
deposition (Francis et al., 2008).
Although these processes are common to all river
systems with large wood, most research has been
performed in single channel montane systems in which
out-of-channel engineering scarcely occurs (Keller
and Swanson, 1979; Fetherston et al., 1995; Abbe and
Montgomery, 1996; Opperman et al., 2008). Except
Research Article 5
on the Tagliamento, much less attention has been paid
to braided systems in which out-of channel deposition
of large wood is an important process.
Figure 3. Simple illustration of spatial patterning of habitats
formed in association with fluvially-deposited trees and pioneer
islands along the River Tagliamento. Taken from Francis et al.
(2008b).
This biogeomorphic ecosystem engineering is ac-
celerated if the tree survives following deposition, or if
the tree is large and therefore represents a more
substantial obstacle to flow; indeed, these factors are
not independent of each other, as larger trees have been
shown to have significantly greater survival than small-
er trees (Francis, 2007). Surviving trees produce
adventitious roots and shoots that help to anchor the
tree onto the bar and allow it to form a more substantial
and stable hydraulic roughness element to flow, further
facilitating the trapping of sediments and propagules
and the creation of scour features (Edwards et al., 1999;
Francis et al., 2006; Opperman and Merenlender, 2007;
Opperman et al., 2008). Providing subsequent disturb-
ance does not remove the deposited tree and its
associated vegetation, the patch grows to a relatively
dense community of riparian species that is then
considered to be a pioneer island (sensu Edwards et
al., 1999). As these islands grow in elevation and area
via sediment and biomass accumulation (Fig. 4), they
amalgamate with other islands to form large building
islands that may remain stable for decades (Gurnell
and Petts, 2002; Zanoni et al., 2008). This joining of
islands is a process predicted from modeling studies of
self-organisation in other ecosystems (e.g. Gumbricht
et al., 2004; Rietkerk et al., 2004).
In fact, large wood is not a requirement for plant
establishment on river bars, and simple vegetated bars
are common on many rivers, their dynamics regulated
by hydrogeomorphic processes (e.g. Gilvear and Will-
by, 2006; Gilvear et al., 2007; Dixon et al., 2002;
6 R. A. Francis, D. Corenblit and P. J. Edwards
Figure 4. Photograph showing a cross-sectional view of a river
island revealing the sedimentary stratigraphy coupled with exten-
sive vertical and lateral roots of the plants growing on the island.
The cumulative layers of fine sediment deposited by flood events
are held together in part by the root systems which add strength to
the deposits and provide protection from erosion during subse-
quent flood events. The growth of the plants adds organic matter to
the duff layer, which also collects further flood deposits and wind
blown sediment. Collectively these processes and material depo-
sition create what is here considered a self-organising biogeomor-
phologically engineered landform due to the nature of the feed-
back mechanisms between physical and biological processes.
Dixon, 2003). However, the building of large islands
on braid bars via the accumulation of plant biomass
and sediment (rather than via floodplain avulsion) has
so far only been demonstrated in association with
(predominantly living) large wood (Gurnell and Petts,
2002). This process of island formation may also be
considered a form of expression of river functioning
within complex fluvial ecosystems (see Fisher et al.,
2007).
River islands act as physical attractors or sinks for a
variety of resources and biological materials. These
include: 1) water, with islands supporting relatively
well developed riparian soils that store groundwater;
2) organic material, particularly large wood (Gurnell
et al. (2001) calculated an average storage of 293 to
1664 tonnes ha–1 of large wood on pioneer islands
compared to 1–6 tonnes ha–1 on gravel bars); 3) fine
sediment, with large islands having elevations up to
3 m above base flow level due to stratified layers of
fine sediments deposited in repeated inundation
events (Fig. 4); and 4) propagules, which tend to be
deposited on and around islands along with fine
sediments and wood. This is a clear example of local
positive feedback occurring at the expense of more
distant parts of the river system, both within the same
reach and further downstream; to the extent that as
these resources are limiting in a river system, their
removal necessarily restricts community development
in other areas.
River islands and self-organisation
Islands develop as a result of biogeomorphic
interactions in which the dominant plants play a
crucial role; thus, plants trap the sediment, anchor the
sediment, redirect flows, store water, and enable
landforms to build in area and elevation. Plants act
as dynamic elements during this self-organisation
process as they continue to grow and to adapt their
needs according to the variation of the balance
between disturbance, stress and competition that
accompanies island accretion. Distinct types of inter-
actions predominate at different scales, from microsite
creation around individual deposited trees to redis-
tribution of resources and island landform creation at
the landscape scale. The sink effect is probably closely
related to island size and area, though this relationship
has only been studied quantitatively for large wood
(Gurnell et al., 2001), and overall effects on landscape
functioning are not known. The area of islands within
the active zone of some sections of the River
Tagliamento can be substantial, with up to 400 islands
per km2 of active zone, and islands have been shown to
occupy 17 % of active zone area in one study (Tockner
et al., 2003). In braided reaches with islands, riparian
foreshore length is 50 % higher than in similar reaches
without islands, and these highly heterogeneous,
shifting island mosaics greatly increase available
riparian habitat for a range of species (Tockner et
al., 2003; Gurnell et al., 2005). The presence of
abundant and heterogeneous island landforms repre-
sents a difference in ecosystem functioning that
almost certainly translates into higher species diver-
sity in reaches containing islands, though this effect
has never been quantified.
Self-organisation and catastrophic shifts in ecosystem
state
The tendency for self-organised systems to undergo
catastrophic shifts (an abrupt dramatic transition) to
alternative states has been demonstrated for systems
as diverse as insect populations and information
networks, and at scales ranging from crystal formation
to climate regimes (see e.g. Graf, 1988; Bonabeau et
al., 1997; Scheffer et al., 2001; Rietkerk et al., 2004;
Walker et al., 2004; Fisher et al., 2007).
Within ecosystems, a key characteristic of self-
organised patch configurations is the tendency of such
configurations to undergo catastrophic shifts between
patchy and homogeneous stable states (metastability)
(Rietkerk et al., 2004). Fluvial ecosystems are meta-
stable critical systems in the sense that abrupt
changes and hysteresis cycles can occur (Bak et
al.,1988; Fig. 5). When thresholds are crossed, the
effects of higher hierarchical levels on lower levels
changes dramatically and can induce a rapid cascading
effect through the entire system, producing a new
Aquat. Sci.
stable domain expressed both ecologically and geo-
morphologically (Fisher et al., 2007). Shifts towards a
patchy state can occur within fluvial corridors be-
cause, although the ecosystems are open, resources
are not replenished or redistributed at a rate sufficient
to prevent the long-distance negative feedback from
occurring, and so the gradual emergence of contigu-
ous vegetated resource-rich patches is inevitable given
the presence of ecosystem engineers (Rietkerk et al.,
2002). The shift from a patchy state to a homogeneous
state occurs when the main limiting resource decreas-
es or increases beyond a threshold where it becomes
either too limiting or unlimited. An example of this
may be seen in arid landscapes in which striped or
spotted patchy states can shift to a bare homogeneous
state if rainfall (and therefore water resources)
decreases beyond a threshold, as the local positive
feedback mechanism fails (Rietkerk et al., 2004).
Conversely, a substantial increase in rainfall may
overcome the long-distance negative feedback that
maintains the self-organised patchy state and allow
vegetation to establish throughout the ecosystem; in
reality, however, this shift to a homogeneous vege-
tated state is unlikely to occur. Such changes in
ecosystem state have occasionally been inferred from
field observations (see Rietkerk et al., 2004), but they
have rarely been observed or quantified due to the
spatial and temporal scale of the systems and the
transitions to be measured (see van Wesenbeeck et al.,
2008). For this reason, most evidence for them has
come from modelling studies, e.g. cellular automaton
models (Rietkerk et al., 2004).
Figure 5. Cusp catastrophe model (Thom, 1975) for fluvial land-
form stability. This topology indicates that when the system is
within the metastable zone two potential states are possible,
corresponding to a position on one or the other side of the two
wings. When approaching the fold curve, a small change in the
controlling variables (i.e. hydrogeomorphic level of disturbance or
vegetation dynamics) may produce a sudden and drastic change in
state, demonstrated here by a sudden switch to the other wing.
Research Article 7
Island-braided fluvial ecosystems may represent
an important example of self-organised patchiness
dependent on biogeomorphic processes in a relatively
heterogeneous ecosystem with highly directional
flows and maintained primarily by hydrological dis-
turbance. Like other patterned ecosystems, fluvial
ecosystems are open dissipative systems within which
key resources are spatially and temporally limited;
what makes them unusual is that the flows of biotic
(species, propagules, organic detritus) and abiotic
(water, sediment, energy) resources are highly direc-
tional and driven by disturbance, usually in the form of
floods of varying severity and frequency (see Junk et
al., 1989; Tockner et al., 2000; Wiens, 2002; Karaus et
al., 2005; Fisher et al., 2007). In fluvial systems such as
the island-braided reaches of the River Tagliamento,
the hydrological regime serves to maintain the self-
organised patchiness of the ecosystem at a point where
a catastrophic shift does not occur in several ways.
These include: 1) disturbing the vegetation and
reconfiguring the spatial arrangement of vegetated
patches (islands); 2) removing or limiting the short-
distance positive feedbacks of some patches while
enabling it in others, as variation in flood effects lead
to both disturbance and (in part) determination of
where resources are deposited and in what quantity;
3) supplying resources to the reach, both longitudi-
nally from reach to reach and laterally between
channels, islands and floodplain; and 4) redistributing
existing resources over a range of spatial and temporal
scales. Consequently, the balance of autogenic bio-
geomorphic processes and allogenic disturbance
maintains the system along with its particular mor-
phological and functional characteristics and associ-
ated level of complexity and biodiversity. This is
observed within the active zones of other fluvial
systems (see e.g. Hughes, 1997; Baker and Walford,
1995; Dixon, 2003; Stanford et al., 2005; Gilvear and
Willby, 2006), but in the absence of biogeomorphic
engineers such as vegetatively regenerating trees,
stable islands and overtly patchy states do not emerge.
Most examples of ecosystem self-organisation
involve the redistribution of single key limiting
resources (e.g. moisture, nutrients). In the case of
river islands, however, interactions among several
ecosystem components are involved, including fine
sediment, water, trees and large wood, and plant
propagules. Without disturbance, the availability of
these various components may either increase or
decrease, leading to a shift in state (Fig. 6). Such
effects were demonstrated for the Tagliamento by
Arscott et al. (2002) and Zanoni et al. (2008) who used
a time series of aerial photographs to investigate the
dynamics of various riverine habitats within an island-
braided section of the river. Both studies demonstrat-
8 R. A. Francis, D. Corenblit and P. J. Edwards
ed that the island/gravel area ratio was relatively
stable over short timescales, while there was a high
turnover of island habitats due to hydrological dis-
turbances. However, Zanoni et al. (2008) also dem-
onstrated that the development of building islands was
limited when key ecosystem flows were interrupted.
For example, during the 1930s and 1940s much of the
riparian woodland was removed, limiting the supply of
large wood to the system; consequently island self-
assembly became much rarer, with island patches
disappearing from images of the river during this
period. Subsequent recovery of the riparian zone has
restored both the self-organisation of islands and the
emergent patchy state (Zanoni et al., 2008). This trend
has also been found for large scour pools (concave
islands) often associated with river islands, with this
habitat type being absent from reaches where flows of
large wood and water are interrupted and islands do
not develop (Karaus et al., 2005).
Figure 6. Self-organisation potential for island landforms in fluvial
systems. In scenarios where disturbance is high and ecosystem
resources (fine sediments, engineering species, organic matter) are
low, self-organisation does not occur and a relatively homogeneous
unvegetated system state is dominant. In scenarios with less severe
and frequent disturbance and medium levels of resource avail-
ability, ecosystem engineers (riparian trees) activate local-scale
feedbacks of resource acquisition self-organised pattern formation.
With infrequent disturbance and abundant resources, the effects of
the local positive and negative long-distance feedbacks are reduced
and vegetation can establish throughout the ecosystem, leading to a
loss of self-organised patchiness and a more homogeneous
vegetated state. Self-organisation of island landforms therefore
relies on a balance of resource availability and disturbance that
may be interrupted by anthropogenic activity.
Ecosystem variability and maintenance of river island
landscapes
Despite the abundance of island landforms of differ-
ent sizes and ages in island-braided reaches along the
River Tagliamento (Tockner et al., 2003), this expres-
sion of biogeomorphic self-organisation is only found
within a specific envelope of environmental condi-
tions, and relies on a series of integral biotic and
River islands and self-organisation
abiotic processes that may vary over time and are
spatially discontinuous through the river network, but
which nevertheless represent a delicate balance of
reciprocal interactions (Poole, 2002; Gurnell et al.,
2005; Corenblit et al., 2007a; Fisher et al., 2007).
There is now increasing recognition that the varia-
bility of natural processes is essential for maintaining
ecosystem functional integrity, in particular by main-
taining non-equilibrium conditions (Poff et al., 1997;
Stromberg, 2001; Naiman et al., 2008). It is also
recognised that the failure to maintain the diversity of
natural processes has been a major cause of ecosystem
degradation (Landres et al., 1999; Francis, 2009). For
instance, attempts to control the natural variability of
fluvial ecosystems inevitably lead to ecosystem deg-
radation, and often pose a substantial risk to humans
(for example the many problems associated with the
management and prediction of flood risks; Apel et al.,
2004). Recent research has shown natural river
systems to be far more dynamic than was originally
supposed (e.g. Fisher et al., 2007), and the River
Tagliamento is considered a reference system because
it is one of the few rivers in Europe that maintains a
natural hydrological dynamism along most of its
length (Tockner et al., 2003). The fact that river
islands are one of the first landforms to disappear
when rivers are managed (Tockner et al., 2003) is
probably because the processes maintaining them are
delicately balanced, with any disruption reducing the
capacity for self-organisation.
At the landscape scale, self-organising river islands
along the Tagliamento are only found where unit
stream power in high flows is not too high to preclude
tree establishment and growth, where the alluvial
aquifer during low flows is sufficient to support tree
growth, and where fine sediments are deposited; and
all of these conditions are determined, at least in part,
by the width of the active zone and gradient of the
river bed (Gurnell and Petts, 2006). That these islands
have developed within the active zone rather than
being excised from the floodplain via avulsion has
been proved by analysing historical photographs
(Zanoni et al., 2008) and by observing changes in
the elevation and extent of islands (Gurnell et al.,
2001). Furthermore, it is known that the active zone of
the intensively studied pre-alpine reaches has not
shifted sufficiently during the past 60 years to form the
islands that are present there. In reaches outside this
broad environmental envelope, only islands formed
by avulsion are found, and the processes of self-
assembly and self-organised patchiness described
above are absent.
Within this environmental envelope, perhaps the
most important process in island formation is the
initial inception of vegetation patches, primarily
Aquat. Sci.
around deposited wood, that leads to the development
of pioneer islands (Edwards et al., 1999; Gurnell et al.,
2001). Research has demonstrated that it is primarily
livewood (surviving deposited trees) that grow fast
enough to establish before subsequent disturbance,
because shoots from whole trees grow up to an order
of magnitude faster than seeds and smaller asexual
propagules such as branches (Francis et al., 2006). The
probability of a deposited tree surviving depends on
several factors:
1) Tree species. Only a limited number of tree species
evolutionarily adapted to fluvial ecosystems have
the capacity for vegetative regeneration following
uprooting, mainly species within the genera Salix,
Populus and Alnus; though regrowth from branch-
es and fragments is most often reported in other
systems (e.g. Barsoum, 2002; Rood et al., 2003a).
Along the River Tagliamento, Populus nigra L.,
Salix alba L. and Salix elaeagnos (Scop.) are the
most common asexually-regenerating species, al-
though regeneration from Salix daphnoides Vill.,
Salix purpurea L. and Alnus incana L. has also been
observed (Tockner et al., 2003; Francis, 2007).
These are early to mid-sere riparian trees, popu-
lations of which are only naturally maintained in
dynamic river systems where disturbance creates
mosaics of different seral stages within the riparian
zone (e.g. Fenner et al., 1985; Hupp and Oster-
kamp, 1996; Johnson, 2000; Karrenberg et al.,
2002). Due to the extent of the riparian zone along
the River Tagliamento, stochastic variation in
populations of these species are unlikely to have
significant long-term repercussions, but any mod-
ification to the river that removes natural dyna-
mism or reduces the extent of the riparian wood-
land may reduce the number of individuals capable
of regeneration that are conducted through the
system, and may therefore prevent island incep-
tion.
2) Tree condition. Floods are naturally destructive
and any tree transported through a (particularly
gravel-bed) system is likely to sustain damage from
collisions with landforms as well as transported
materials such as cobbles, boulders and other trees
(Pigay and Gurnell, 1997). No research has been
conducted into the influence of tree damage on
survival probabilities, but it is likely that fragment-
ed trees, trees which are stripped of branches and
bark, and trees that have lost their root boles are
less likely to survive than intact trees.
3) Internal resources. Production of adventitious
roots and shoots from tree nodes relies on the
tree containing sufficient internal resources such as
carbohydrates to initiate regrowth (Burgess et al.,
Research Article 9
1990), which will vary depending on the size and
age of the tree, the time of year at which entrain-
ment and deposition occurs, and tree health. Size
appears to be particularly important in determin-
ing which trees survive, with longer trees being
significantly more likely to survive than shorter
trees (Francis, 2007), a trend also demonstrated
with tree fragments (see Burgess et al., 1990; Rood
et al., 2003a). The presence of mature riparian
woodland is probably important for ensuring a
supply of trees capable of surviving on the gravel
bars.
4) Depositionary location. Trees may be deposited
across the active zone, from channel edges to bar
tops (Fetherston et al., 1995; Abbe and Montgom-
ery, 1996; Pigay et al., 1999), and the location of
deposition is likely to notably affect whether trees
survive or not. Elevation of depositionary site has
been shown to influence tree survival, with trees
generally surviving better at lower elevations, as
the adventitious roots of such trees can reach the
water table more easily (Francis, 2007). There is
some disadvantage to being deposited at low
elevations however, as trees that are deposited
too close to the channel edge are likely to be re-
entrained more easily than those deposited higher
up the bars; greatest survival is generally at mid-
elevations (around 0.5 – 1.5 m below the level of
riparian woodland on the River Tagliamento;
Francis, 2007). Sedimentary conditions at the
location of deposition are also likely to be impor-
tant for survival, with finer sediments providing
more moisture and nutrient-rich substrates to
promote root growth and penetration, and to
allow a deposited tree to grow beyond the level
that internal resources allow initially. These gen-
eral patterns are similar to those shown for
regeneration from seedlings in many other systems
(e.g. Dixon, 2003; Gilvear and Willby, 2006); but,
as noted before, in this system island inception is
largely dependent on the regeneration of relatively
intact trees (see Francis et al., 2006).
Once it has resprouted, the survival of a tree depends
on it not being significantly disturbed. In fluvial
environments, the key disturbances are erosion of
substrate and associated root structures, that could
lead to removal of the tree, and sediment deposition,
leading to burial (e.g. Kent et al., 2001; Pollen et al.,
2004). Both processes are highly variable spatially and
temporally, though little research has been conducted
to quantify such variability and potential impacts,
particularly for sedimentation and vegetation burial
(see Keddy, 2000; Kent et al., 2001). Fine-scale
thresholds are likely to exist for erosion and burial
10 R. A. Francis, D. Corenblit and P. J. Edwards
that determine whether the level of substrate scour or
deposition have a net positive or negative effect on
survival. Sediment scour is usually negative in its
effects, exposing root systems and reducing access to
water; however, if a scour feature develops into a pool
(e.g. Karaus et al., 2005; Francis et al., 2008b), it may
improve water availability. Burial can have both
positive and negative effects for plants: a layer of
fine sediment around a deposited tree provides a
moisture-retentive medium that can encourage vege-
tative sprouting, while too much sediment may
prevent shoots from reaching the surface (Kent et
al., 2001). These biogeomorphic thresholds are key
areas for future investigations into ecosystem engi-
neering.
Consequently, the probability of an individual tree
being the right species, being entrained and trans-
ported downstream, being deposited relatively intact
in a suitable location, having access to enough internal
resources to initiate regrowth, and then lasting long
enough to engineer the environment and contribute to
island development is probably quite low, and the
processes that lead to island development are only
maintained by the sheer number of trees moving
through the system (Edwards et al., 1999; Gurnell et
al., 2001). Once trees and islands are established
within the system, their ability to act as resource sinks
(including the trapping and storage of trees and wood
on and around the islands) creates a positive feedback
that leads to more island growth, and increases the
probability of deposited tree survival and island
inception, particularly in sheltered locations down-
stream of large islands. At the landscape scale, thresh-
olds probably exist whereby a critical mass of islands
is needed to facilitate this process and maintain the
island landscapes. Likewise, the loss of trees being
transported through the system, e.g. from the removal
of riparian woodland (Zanoni et al., 2008), may
diminish the island area to the point where this
positive feedback mechanism has little influence on
landscape patterns.
Island self-organisation and associated biogeo-
morphic processes as discussed above can only really
be maintained by natural dynamism, the most
important being hydrological dynamism, sediment
and wood transport, and riparian woodland succes-
sion (Rood et al., 1995; Tabacchi et al., 1998;
Stromberg, 2002; Rood et al., 2003b; Lytle and
Merritt, 2004). Interruption of any of these will
significantly reduce wood entrainment and trans-
portation through the system as well as the resources
required for island formation and development (e.g.
Jacobson et al., 1999; Karaus et al., 2005). This is
different from other ecosystems displaying self-
organised patchiness, where the ecosystem engineers
River islands and self-organisation
are assumed to be not limited; in this case, the highly
directional flows associated with fluvial ecosystems
change the nature of self-organisation and ecosystem
function.
Biogeomorphic inheritance and self-organisation of
river islands
The self-organizing dynamics of river islands take
place in fluvial corridors at the scale of ecological
succession. Corenblit et al. (2007a) considered self-
organisation at this scale as a biogeomorphic succes-
sion, defined as the synergy of ecological succession
and landform and habitat changes. The concept of
biogeomorphic succession encapsulates feedback
mechanisms between biological and physical compo-
nents of the riparian ecosystem (see also Fisher et al.,
2007). In this model, reciprocal adjustments between
riparian plant community and fluvial landform struc-
tures are initiated and driven by pioneer plant species
which, as detailed above, contribute to stabilising the
substrate and trap large amounts of fine sediment
within exposed zones of the active tract (see also e.g.
Hughes, 1997; Naiman and Decamps, 1997; Tooth and
Nanson, 2000). This biogeomorphic aggregative proc-
ess occurs according to a certain range of flood
magnitude and frequency, mainly during low to
intermediate magnitude and high to intermediate
frequency floods. Thus, the domain, or the envelope,
where abiotic-biotic feedbacks can reach sufficient
levels and sufficient continuity in time to produce self-
organisation may be restrictive within fluvial systems.
However, this restricted envelope may play an im-
portant role at the evolutionary scale for the develop-
ment of specific adaptations of plants to hydraulic
constraints (see also Reich et al., 2003; Lytle and Poff,
2004).
It is interesting to consider whether the self-
organisation processes that are observed at the
ecosystem level (Rietkerk and van de Koppel, 2008)
have an evolutionary basis and may essentially be
extended expressions of genes of engineering organ-
isms (Corenblit et al., 2007b, 2008); and whether the
characteristic landform patterns associated with en-
gineers may therefore maintain and replicate them-
selves by providing advantages to the engineering
organisms and/or other species which are facilitated
by the process of island and floodplain construction.
River islands on the Tagliamento are an expression of
the reproductive and vegetative properties of partic-
ular species, notably species of Salicaceae (Karren-
berg et al., 2002), that are in turn determined by
specific genes. The selections of such specific engi-
neering genes may have taken place in the long term
according to limits imposed by natural flow regimes
(see Lytle and Poff, 2004; Fig. 7).
Aquat. Sci.
Figure 7. Adaptive zone for ecosystem engineering traits within
fluvial systems. Related fluvial styles are indicated in the black
boxes.
In fluvial systems where the level of hydrogeo-
morphic disturbance is too weak, specific morpho-
logical, biomechanical and life-history adaptations to
water flow variability such as identified by Lytle and
Poff (2004) may not take place as competitive traits
or strategies may be more suitable for accessing
resources. In situations where notable hydrogeomor-
phic disturbances are too frequent, plants (in partic-
ular woody species) may not have time to complete
their life cycles and produce a significant number of
viable descendants. Specific morphological, biome-
chanical and life-history adaptations to hydrogeo-
morphic disturbances (e.g. Kozlowski, 1984; Islam
and MacDonald, 2004), in particular those related to
the mechanical constraints imposed by flows (Kar-
renberg et al., 2003; Puijalon et al., 2005), may only
have developed under conditions of intermediate
disturbance; under these conditions, disturbances
are sufficiently frequent and intense to act as an
evolutionary agent while preserving the continuity of
abiotic-biotic feedbacks. The selection of engineer-
ing biological traits producing self-organisation at
the ecosystem scale may only take place in systems
with such a regime (well illustrated by the Taglia-
mento), while the island braided fluvial style could
not exist without vegetation. In contrast to stabilised
rivers, dynamic rivers such as the Tagliamento may
be considered, at least in Europe, as relics of the
adaptive zone for riparian trees; thus, they are
essentially refuges for evolutionary processes asso-
ciated with flow variability.
The traits of engineering species (e.g. Populus
nigra, Salix spp., Alnus spp.) that enable them to
Research Article 11
reproduce vegetatively and engineer habitats and
landforms according to specific morphological and
biomechanical adaptations are likely to influence not
just river functioning and morphology at varying
scales but also the populations, distributions and
genetic diversity of the engineering species over time
(Braatne et al., 1996; Karrenberg et al., 2002; Rood
et al., 2003a, 2003b). This is an example of niche
engineering that may lie somewhere between pos-
itive niche construction (i.e. extended phenotype
sensu Dawkins, 1982) and negative niche construc-
tion (i.e. niche changing sensu Dawkins, 2004), as the
environmental modifications help to maintain pop-
ulations of the engineering species while at the same
time facilitating establishment of other plant and
animal organisms (Day et al. 2003). Taken together,
the specific biogeomorphic processes and self-organ-
isation associated with river island dynamics offer an
example of biogeomorphic inheritance (Corenblit et
al., 2007a), whereby the reciprocal processes found
on island landforms support these engineer species
and promote the future development of the land-
forms. This concept supports the observed absence of
self-assembled river islands and equivalent engineers
in some parts of the world, as the evolutionary
background is not there to drive the creation of such
landforms. The historical prevalence of river islands
in many river systems (Gurnell and Petts, 2002) may
have supported these species and helped to maintain
specific adaptations (e.g. flexible stems, vegetative
reproduction, rapid growth rates) that are advanta-
geous in highly disturbed environments; conversely,
the loss of islands may reduce the distribution of
these species in the future. No studies have looked at
the comparative success of sexual and asexual
reproduction in natural systems such as the Taglia-
mento. This is an urgent priority to ensure that the
role of river islands within fluvial biogeomorphology
at a range of spatial and temporal scales is fully
understood.
Future directions for research into
self-organisation of fluvial ecosystems
Based on this critical discussion of river islands as
examples of self-organisation, and using perspectives
drawn from studies on the River Tagliamento in Italy,
we suggest the following research areas as priorities
for observations, experimentation and modelling to
illuminate ecological and biogeomorphic dynamics in
rivers that still maintain island landforms and asso-
ciated biocomplexity and biodiversity. These research
areas may aid ecosystem management and restora-
tion:
12 R. A. Francis, D. Corenblit and P. J. Edwards
• Comparison of braided, single channel and island
braided river reaches in terms of biodiversity,
community composition, landscape metrics, spe-
cies functional typologies and resource distribu-
tions.
• Investigation of characteristics of self-organised
islands and island landscapes, including the relative
balance of sexual and asexual reproductive strat-
egies, genotypic variability, and quantification of
feedbacks at different scales.
• Identification and quantification of thresholds as-
sociated with self-organisation at scales ranging
from fine-scale biogeomorphic processes (e.g. how
sedimentation and erosion affect plant growth) to
landscape-scale thresholds (e.g. the number and
area of island landforms required to create a
positive feedback of resource retention and island
development).
• Assessment of the role of environmental hetero-
geneity in biogeomorphic self-organisation as a
basis for evaluating the range of conditions under
which self-organisation can occur, and determining
how such landforms can be restored.
• Evaluation of links between self-organised patchi-
ness and ecosystem resilience to natural and
anthropogenic disturbances.
• Investigation of the implications of self-organisa-
tion (and its loss) for evolution and genetic diversity
of riparian species, in particular those associated
with ecosystem engineering.
Acknowledgments
This paper presents perspectives that resulted from
discussions held by the authors during the European
Science Foundation workshop Linkages and feed-
backs in highly dynamic, alpine, fluvial systems held
in Cornino, Italy, from 1st-3rd July 2008. Two anon-
ymous reviewers are thanked for their constructive
comments that greatly helped to improve the manu-
script.
References
Abbe, T. B. and D. R. Montgomery, 1996. Large woody debris jams,
channel hydraulics and habitat formation in large rivers.
Regulated Rivers 12: 201 – 221.
Aguiar, M. R. and O. E. Sala, 1999. Patch structure, dynamics and
implications for the functioning of arid ecosystems. Trends in
Ecology and Evolution 14: 273 – 277.
Apel, H., A. H. Thieken, B. Merz and G. Bloschl, 2004. Flood risk
assessment and associated uncertainty. Natural Hazards and
Earth System Sciences 4: 295 – 308.
Arscott, D. B., K. Tockner and J. V. Ward, 2002. Geomorphic
River islands and self-organisation
dynamics along a braided-river corridor in the Alps (Fiume
Tagliamento, NE Italy). Ecosystems 5: 802 – 814.
Baas, A. C. W., 2007. Complex systems in aeolian geomorphology.
Geomorphology 91: 311 – 331.
Bak, P., 1996. How Nature Works: The Science of Self-organized
Criticality. Springer-Verlag, New York, 212 pp.
Bak, P., C. Tang and K. Wiesenfeld, 1988. Self-organised criticality.
Physical Review A 38: 364 – 374.
Baker, W. L. and G. M. Walford, 1995. Multiple stable states and
models of riparian vegetation succession on the Animas River,
Colorado. Annals of the American Association of Geogra-
phers 85: 320 – 338.
Barsoum, N., 2002. Relative contributions of sexual and asexual
regeneration strategies in Populus nigra and Salix alba during
the first years of establishment on a braided gravel bed river.
Evolutionary Ecology 15: 255 – 279.
Bonabeau, E., G. Theraulaz, J-L. Deneubourg, S. Aron and S.
Camazine, 1997. Self-organization in social insects. Trends in
Ecology and Evolution 12: 188 – 193.
Braatne, J. H., S. B. Rood and P. E. Heilman, 1996. Life history,
ecology, and conservation of riparian cottonwoods in North
America. In: R. F. Stettler, H. D. Bradshaw, P. E. Heilman and
T. M. Hinckley (eds.) Biology of Populus and its implications
for management and conservation, National Research Council
of Canada, Ottawa, pp. 57 – 85.
Bravard, J. P., C. Amoros, G. Pautou, G. Bornette, M. Bournaud,
M. Creuz des Chtelliers, J. Gibert, J. L. Peiry, J. F. Perrin, and
H. Tachet, 1997. River incision in south-east France, morpho-
logical phenomena and ecological effects. Regulated Rivers-
Research and Management 13: 75 – 90.
Burgess, D., O. Q. Hendrickson and L. Roy, 1990. The importance
of initial cutting size for improving the growth performance of
Salix alba L. Scandinavian Journal of Forest Research 5: 215 –
224.
Camazine, S., J.-L. Deneubourg, N. R. Franks, J. Sneyd, G.
Theraulaz and E. Bonabeau, 2001. Self-organisation in Bio-
logical Systems, Princeton University Press, New Jersey, 560
pp.
Charlton, R., 2008. Fundamentals of Fluvial Geomorphology,
Routledge, London, 234 pp.
Coco, G. and A. B. Murray, 2007. Patterns in the sand: From forcing
templates to self-organization. Geomorphology 91: 271 – 290.
Corenblit, D., E. Tabacchi, J. Steiger and A. M. Gurnell, 2007a.
Reciprocal interactions and adjustments between fluvial land-
forms and vegetation dynamics in river corridors: A review of
complementary approaches. Earth-Science Reviews 84: 56 –
86.
Corenblit, D., J. Steiger, A. M. Gurnell and E. Tabacchi, 2007b.
Darwinian origin of landforms. Earth Surface Processes and
Landforms 32: 2070 – 2073.
Corenblit, D., A. M. Gurnell, J. Steiger and E. Tabacchi, 2008.
Reciprocal adjustments between landforms and living organ-
isms: extended geomorphic evolutionary insights. Catena 73:
261 – 273.
Dawkins, R., 1982. The Extended Phenotype, Freeman, Oxford,
336 pp.
Dawkins R., 2004. Extended phenotype – but not too extended. A
reply to Laland, Turner and Jablonka. Biology and Philosophy
19: 377 – 396.
Day, R. L., K. N. Laland and J. Odling-Smee, 2003. Rethinking
adaptation – the niche-construction perspective. Perspectives
in Biology and Medicine 46: 80 – 95.
Dent, C. L., G. S. Cumming and S. R. Carpenter, 2002. Multiple
states in river and lake ecosystems. Philosophical Transactions
of the Royal Society of London Series B – Biological Sciences
357: 635 – 645.
Dixon, M. D., M. G. Turner and C. Jin, 2002. Riparian tree seedling
distribution on Wisconsin River sandbars: controls at different
spatial scales. Ecological Monographs 72: 465 – 485.
Dixon, M. D., 2003. Effects of flow pattern on riparian seedling
recruitment on sandbars in the Wisconsin River, Wisconsin,
USA. Wetlands 23: 125 – 139.
Aquat. Sci.
Drever, C. R., G. Peterson, C. Messier, Y. Bergeron and M.
Flannigan, 2006. Can forest management based on natural
disturbances maintain ecological resilience? Canadian Journal
of Forest Research 36: 2285 – 2299.
Edwards, P. J., J. Kollmann, A. M. Gurnell, G. E. Petts, K. Tockner
and J. V. Ward, 1999. A conceptual model of vegetation
dynamics on gravel bars of a large Alpine river. Wetlands
Ecology and Management 7: 141 – 153.
Fenner, P., W. W. Brady and D. R. Patton, 1985. Effects of
regulated water flows on regeneration of Fremont cottonwood.
Journal of Range Management 38: 135 – 138.
Fetherston, K. L., R. J. Naiman and R. E. Bilby, 1995. Large woody
debris, physical process, and riparian forest development in
montane river networks of the Pacific Northwest. Geomor-
phology 13: 133 – 144.
Fisher, S. G., J. B. Heffernan, R. A. Sponseller and J. R. Welter,
2007. Functional ecomorphology: Feedbacks between form
and function in fluvial landscape ecosystems. Geomorphology
89: 84 – 96.
Forman, R. T. T., 1995. Land Mosaics: The Ecology of Landscapes
and Regions, Cambridge University Press, Cambridge, 632 pp.
Francis, R. A., 2006. Allogenic and autogenic influences upon
riparian vegetation dynamics. Area 38: 453 – 464.
Francis, R. A., 2007. Size and position matter: riparian plant
establishment from fluvially-deposited trees. Earth Surface
Processes and Landforms 32: 1239 – 1243.
Francis, R. A., 2009. Ecosystem prediction and management. In: N.
Castree, D. Demeritt, D. Liverman and B. Rhoads (eds.), A
Companion to Environmental Geography, Blackwell, London,
pp. 421 – 441.
Francis, R. A., A. M. Gurnell, G. E. Petts and P. J. Edwards, 2006.
Riparian tree establishment on gravel bars: interactions
between plant growth strategy and the physical environment.
In: G. Sambrook-Smith, J. Best, C. Bristow, and G. E. Petts
(eds.), Braided Rivers: Process, Deposits, Ecology and Man-
agement, International Association of Sedimentologists Spe-
cial Publication 36, Blackwell, London, pp. 361 – 380.
Francis, R. A., G. E. Petts and A. M. Gurnell, 2008a. Wood as a
driver of past landscape change along river corridors. Earth
Surface Processes and Landforms 33: 1622 – 1626.
Francis, R. A., P. Tibaldeschi and L. McDougall, 2008b. Fluvially-
deposited large wood and riparian plant diversity. Wetlands
Ecology and Management 16: 371 – 382.
Gilvear, D., R. Francis, N. Willby and A. Gurnell, 2007. Gravel
bars: A key habitat of gravel-bed rivers for vegetation. In: H.
Habersack, H. Piegay and M. Rinaldi (eds.), Developments in
Earth Surface Processes 11: Gravel-Bed Rivers VI: From
Process Understanding to River Restoration, Elsevier,
pp. 677 – 702.
Gilvear, D. and N. Willby, 2006. Channel dynamics and geomorphic
variability as controls on gravel bar vegetation; River Tummel,
Scotland. River Research and Applications 22: 457 – 74.
Graf, W. L., 1988. Fluvial Processes in Dryland Rivers. Springer-
Verlag, New York, 346 pp.
Gumbricht, T., J. McCarty, T. S. McCarty, 2004. Channels, wetlands
and islands in the Okavango Delta, Botswana, and their
relation to hydrological and sedimentological processes. Earth
Surface Processes and Landforms 29: 15 – 29.
Gurnell, A. M. and G. E. Petts, 2002. Island-dominated landscapes
of large floodplain rivers, a European perspective. Freshwater
Biology 47: 581 – 600.
Gurnell, A. M. and G. E. Petts, 2006. Trees as riparian engineers:
the Tagliamento River, Italy. Earth Surface Processes and
Landforms 31: 1558 – 1574.
Gurnell, A. M., G. E. Petts, D. M. Hannah, B. P. G. Smith, P. J.
Edwards, J. Kollmann, J. V. Ward and K. Tockner, 2001.
Riparian vegetation and island formation along the gravel-bed
Fiume Tagliamento, Italy. Earth Surface Processes and Land-
forms 26: 31 – 62.
Gurnell, A. M., K. Tockner, G. E. Petts and P. J. Edwards, 2005.
Effects of deposited wood on biocomplexity of river corridors.
Frontiers in Ecology and Environment 3: 377 – 382.
Research Article 13
Hastings, A., C. L. Hom, S. Ellner, P. Turchin and H. C. J. Godfray,
1993. Chaos in ecology: is mother nature a strange attractor?
Annual Review of Ecology and Systematics, 24: 1 – 33.
Heffernan, J. B., 2008. Wetlands as an alternative stable state in
desert streams. Ecology 89: 1261 – 1271.
Hooke, J. M., 2007. Complexity, self-organisation and variation in
behaviour in meandering rivers. Geomorphology 91: 236 – 258.
Hughes, F. M.R., 1997. Floodplain biogeomorphology. Progress in
Physical Geography 21: 501 – 529.
Hupp, C. R. and W. R. Osterkamp, 1996. Riparian vegetation and
fluvial geomorphic processes. Geomorphology 14: 277 – 295.
Islam, M. A. and S. E. Macdonald, 2004. Ecophysiological adapta-
tions of black spruce (Picea mariana) and tamarack (Larix
laricina) seedlings to flooding. Trees – Structure and Function
18: 35 – 42.
Jacobson, P. J., K. M. Jacobson, P. L. Angermeier and D. S. Cherry,
1999. Transport, retention, and ecological significance of
woody debris within a large ephemeral river. Journal of the
North American Benthological Society 18: 429 – 444.
Johnson, W. C., 1994. Woodland expansion in the Platte River,
Nebraska, patterns and causes. Ecological Monographs 64:
45 – 84.
Johnson, W. C., 1997. Equilibrium response of riparian vegetation
to flow regulation in the Platte River, Nebraska. Regulated
Rivers-Research and Management 13: 403 – 415.
Johnson, W. C., 2000. Tree recruitment and survival in rivers:
influence of hydrological processes. Hydrological Processes
14: 3051 – 3074.
Jones, C. G., J. H. Lawton and M. Shachak, 1994. Organisms as
ecosystem engineers. Oikos 69: 373 – 386.
Junk, W. J., P. B. Bayley and R. E. Sparks, 1989. The flood pulse
concept in river-floodplain systems. In: D. P. Dodge (ed.),
Proceedings of the International Large River Symposium,
Canadian Special Publication of Fisheries and Aquatic Scien-
ces, 106: 110 – 127.
Karaus, U., L. Alder and K. Tockner, 2005. Concave islands:
diversity and dynamics of parafluvial ponds in a gravel-bed
river. Wetlands 25: 26 – 37.
Karrenberg, S., P. J. Edwards and J. Kollmann, 2002. The life
history of Salicaceae living in the active zone of flood plains.
Freshwater Biology 47: 733 – 748.
Karrenberg, S., S. Blaser, J. Kollmann, T. Speck and P. J. Edwards,
2003. Root anchorage of saplings and cuttings of woody
pioneer species in a riparian environment. Functional Ecology
17: 170 – 177.
Kauffman, S. A., 1993. The Origins of Order, Oxford University
Press, Oxford, 734 pp.
Keddy, P., 2000. Wetland Ecology: Principles and Conservation,
Cambridge University Press, Cambridge, 614 pp.
Keller, E. A. and F. J. Swanson, 1979. Effects of large organic debris
on channel form and fluvial process. Earth Surface Processes 4:
361 – 380.
Keller, E. F., 2008. Organisms, machines, and thunderstorms: A
history of self-organization, part one. Historical Studies in the
Natural Sciences 38: 45 – 75.
Kent, M., N. W. Owen, P. Dale, R. M. Newnham and T. M. Giles,
2001. Studies of vegetation burial: a focus for biogeography
and biogeomorphology? Progress in Physical Geography 25:
455 – 482.
Kozlowski, T. T., 1984. Responses of woody plants to flooding. In:
T. T. Kozlowski (ed.) Flooding and plant growth, Academic
Press, Orlando, pp. 129 – 163.
Landres, P. B., P. Morgan and F. J. Swanson, 1999. Overview of the
use of natural variability concepts in managing ecological
systems. Ecological Applications 9: 1179 – 1188.
Larsen, L. G., W. Judson, J. P. Harvey and A. Crimaldi, 2007. A
delicate balance: ecohydrological feedbacks governing land-
scape morphology in a lotic peatland. Ecological Monographs
77: 591 – 614.
Lytle, D. A. and D. M. Merritt, 2004. Hydrologic regimes and
riparian forests: A structured population model for cotton-
wood. Ecology 85: 2493 – 2503.
14 R. A. Francis, D. Corenblit and P. J. Edwards
Lytle, D. A. and N. L. Poff, 2004. Adaptation to natural flow
regimes. Trends in Ecology and Evolution 19: 94 – 100.
Murray, J. D., 1989. Mathematical Biology, Springer-Verlag, New
York, 736 pp.
Murray, B. and M. A. Fonstad, 2007. Preface: Complexity (and
simplicity) in landscapes. Geomorphology 91: 173 – 177.
Naiman, R. J. and H. Dcamps, 1997. The ecology of interfaces:
riparian zones. Annual Review of Ecology and Systematics 28:
621 – 658.
Naiman, R. J., J. J. Latterell, N. E. Pettit and J. D. Olden, 2008.
Flow variability and the biophysical vitality of river systems.
Comptes Rendus Geosciences 340: 629 – 643.
Opperman, J. J. and A. M. Merenlender, 2007. Living trees provide
stable large wood in streams. Earth Surface Processes and
Landforms 32: 1229 – 1238.
Opperman, J. J., M. Meleason, R. A. Francis and R. Davies-Colley,
2008. Livewood: geomorphic and ecological functions of
living trees in river channels. BioScience 58: 1069 – 1078.
Osterkamp, W. R., 1998. Processes of fluvial-island formation, with
examples from Plum Creek, Colorado, and Snake River, Idaho.
Wetlands 18: 530 – 545.
Pahl-Wostl, C., 1995. The Dynamic Nature of Ecosystems: Chaos
and Order Entwined, John Wiley and Sons Ltd., Chichester,
267 pp.
Perry, P. D., 1995. Self-organizing systems across scales. Trends in
Ecology and Evolution 10: 241 – 244.
Perry, A. A., M. P. Amaranthus, J. G. Borchers, S. L. Borchers and
R. E. Brainerd, 1989. Bootstrapping in ecosystems. BioScience
39: 230 – 237.
Pettit, N. E. and R. J. Naiman, 2005. Flood-deposited wood debris
and its contribution to heterogeneity and regeneration in a
semi-arid riparian landscape. Oecologia 145: 434 – 444.
Petts, G. E., 1989. Historical analysis of fluvial hydrosystems. In:
G. E. Petts (ed), Historical Change of Large Alluvial Rivers:
Western Europe, John Wiley and Sons Ltd., Chichester, pp. 1 –
18.
Petts, G. E. and A. M. Gurnell, 2005. Dams and geomorphology:
research progress and new directions. Geomorphology 71: 27 –
47.
Phillips, J. D., 1995. Self-organisation and landscape evolution.
Progress in Physical Geography 19: 309 – 321.
Phillips, J. D., 1999. Earth Surface Systems: Complexity, Order and
Scale, Blackwell, Oxford, 192 pp.
Pigay, H. and A. M. Gurnell, 1997. Large woody debris and river
geomorphological pattern: Examples from SE France and S
England. Geomorphology 19: 99 – 116.
Pigay, H., A. Thvnet and A. Citterio, 1999. Input, storage and
distribution of large woody debris along a mountain river
continuum, the Dr me River, France. Catena 35: 19 – 39.
Poff, N. L., J. D. Allan, M. B. Bain, J. R. Karr, K. L. Prestergaard, B.
D. Richter, R. E. Sparks and J. C. Stromberg, 1997. The natural
flow regime. BioScience 47: 769 – 781.
Pollen, N., A. Simon and A. Collison, 2004. Advances in assessing
the mechanical and hydrologic effects of riparian vegetation on
streambank stability. Riparian Vegetation and Fluvial Geo-
morphology 8: 125 – 139.
Poole, G. C., 2002. Fluvial landscape ecology: addressing unique-
ness within the river discontinuum. Freshwater Biology 47:
641 – 660.
Puijalon, S., G. Bornette and P. Sagnes, 2005. Adaptations to
increasing hydraulic stress: morphology, hydrodynamics and
fitness of two higher aquatic plant species. Journal of Exper-
imental Botany 56: 777 – 786.
Reich, P. B., I. J. Wright, J. Cavender-Bares, J. M. Craine, J.
Oleksyn, M. Westoby and M. B. Walters, 2003. The evolution
of plant functional variation: Traits, spectra, and strategies.
International Journal of Plant Sciences 164: S143 – S164.
Rietkerk, M., M. C. Boerlijst, F. van Langevelde, R. Hillerislam-
bers, J. van de Koppel, L. Kumar, H. H. T. Prins, and A. M. de
Roos, 2002. Self-organisation of vegetation in arid ecosystems.
American Naturalist 160: 524 – 530.
Rietkerk, M., S. C. Dekker, P. C. de Ruiter and J. van de Koppel,
River islands and self-organisation
2004. Self-organised patchiness and catastrophic shifts in
ecosystems. Science 305: 1926 – 1929.
Rietkerk, M. and J. van de Koppel, 2008. Regular pattern formation
in real ecosystems. Trends in Ecology and Evolution 23: 169 –
175.
Rood, S. B., J. M. Mahoney, D. E. Reid and L. Zilm, 1995. Instream
flows and the decline of riparian cottonwoods along the St.
Mary River, Alberta. Canadian Journal of Botany 73: 1250 –
1260.
Rood, S. B., A. R. Kalischuk, M. L. Polzin and J. H. Braatne, 2003a.
Branch propagation, not cladoptosis, permits dispersive, clonal
reproduction of riparian cottonwoods. Forest Ecology and
Management 186: 227 – 242.
Rood, S. B., J. H. Braatne and F. M. R. Hughes, 2003b. Ecophysi-
ology of riparian cottonwoods: stream flow dependency, water
relations and restoration. Tree Physiology 23: 1113 – 1124.
Sand-Jensen, K., 1998. Influence of submerged macrophytes on
sediment composition and near-bed flow in lowland streams.
Freshwater Biology 39: 663 – 679.
Scheffer, M., S. Carpenter, J. A. Foley, C. Folke and B. Walker,
2001. Catastrophic shifts in ecosystems. Nature 413: 591 – 596.
Schumm, S. A., 2005. River Variability and Complexity, Cam-
bridge University Press, Cambridge, 220 pp.
Stallins, J. A., 2006. Geomorphology and ecology: Unifying themes
for complex systems in biogeomorphology. Geomorphology
77: 207 – 216.
Stanford, J. A., M. S. Lorang and F. R. Hauer, 2005. The shifting
habitat mosaic of river ecosystems. International Association
of Theoretical and Applied Limnology 29: 123 – 136.
Stromberg, J. C., 2001. Restoration of riparian vegetation in the
south-western United States: importance of flow regimes and
fluvial dynamism. Journal of Arid Environments 49: 17 – 34.
Stromberg, J. C., 2002. Flood flows and population dynamics of
Arizona sycamore (Platanus wrightii). Western North Amer-
ican Naturalist 62: 170 – 187.
Tabacchi, E., D. L. Correll, R. Hauer, G. Pinay, A.-M. Planty-
Tabacchi and R. C. Wissmar, 1998. Development, maintenance
and role of riparian vegetation in the river landscape. Fresh-
water Biology 40: 497 – 516.
Thom, R., 1975. Structural Stability and Morphogenesis, Benjamin
Addison Wesley, New York, 400 pp.
Tockner, K., F. Malard and J. V. Ward, 2000. An extension of the
flood pulse concept. Hydrological Processes 14: 2861 – 2883.
Tockner, K., J. V. Ward, D. B. Arscott, P. J. Edwards, J. Kollmann,
A. M. Gurnell, G. E. Petts and B. Maiolini, 2003. The
Tagliamento River: A model ecosystem of European impor-
tance. Aquatic Sciences 65: 239 – 253.
Tooth, S. and G. C. Nanson, 2000. The role of vegetation in the
formation of anabranching channels in an ephemeral river,
Northern plains, arid central Australia. Hydrological Processes
14: 3099 – 3117.
Turcotte, D. L., 2007. Self-organized complexity in geomorphol-
ogy: Observations and models. Geomorphology 91: 302 – 310.
Van Hulzen, J. B., J. Van Soelen and T. J. Bouma, 2007. Morpho-
logical variation and habitat modification are strongly corre-
lated for the autogenic ecosystem engineer Spartina anglica
(Common cordgrass). Estuaries and Coasts 30: 3 – 11.
Van Wesenbeeck, B. K., J. van de Koppel, P. M.J. Herman, M. D.
Bertness, D. van der Wal, J. P. Bakker and T. J. Bouma, 2008.
Potential for sudden shifts in transient systems: Distinguishing
between local and landscape-scale processes. Ecosystems 11:
1133 – 1141.
Viles, H. A., 1988. Biogeomorphology, Basil Blackwell Ltd.,
Oxford, 1142 pp.
Walker, B., C. S. Holling, S. R. Carpenter and A. Kinzig, 2004.
Resilience, adaptability and transformability in social – eco-
logical systems. Ecology and Society 9: 5.
Ward, J. V., K. Tockner, P. J. Edwards, J. Kollmann, G. Bretschko,
A. M. Gurnell, G. E. Petts and B. Rossaro, 2000. Potential role
of island dynamics in river ecosystems. Verhandlungen inter-
nationale Vereinigung fuer theoretische und angewandte
Limnologie 27: 2582 – 2585.
 Aquat. Sci. Research Article 15

Ward, J. V., K. Tockner, D. B. Arscott and C. Claret, 2002. Riverine
landscape diversity. Freshwater Biology 47: 517 – 539.
Wende, R. and G. C. Nanson, 1998. Anabranching rivers: ridge-
form alluvial channels in tropical northern Australia. Geo-
morphology, 22: 205 – 224.
Wetzel, P. R., A. G. van der Valk, S. Newman, D. E. Gawlik, T. G.
Troxler Gann, C. A. Coronado Molina, D. L. Childers and
F. H. Sklar, 2005. Maintaining tree islands in the Florida
Everglades: nutrient redistribution is the key. Frontiers in
Ecology and the Environment 3: 370 – 376.
Wiens, J. A., 2002. Riverine landscapes: taking landscape ecology
into the water. Freshwater Biology 47: 501 – 515.
Zanoni, L., A. Gurnell, N. Drake and N. Surian, 2008. Island
dynamics in a braided river from analysis of historical maps and
air photographs. River Research and Applications 24: 1141 –
1159.

To access this journal online:

http://www.birkhauser.ch/AS

View publication stats