Cent. Eur. J. Geosci. • 4(3) • 2012 • 376-382
DOI: 10.2478/s13533-011-0075-6
Central European Journal of Geosciences
The emergence of an ‘evolutionary geomorphology’?
Commentary
Johannes Steiger1,2∗ , Dov Corenblit1,2
1 Clermont Université, Maison des Sciences de l’Homme, 4 rue Ledru, 63057 Clermont-Ferrand Cedex 1, France
2 CNRS, UMR 6042, GEOLAB – Laboratoire de géographie physique et environnementale, 63057 Clermont-Ferrand, France
Received 23 March 2012; accepted 1 July 2012
Abstract: Earth surface processes and landforms are modified through the actions of many microorganisms, plants and
animals. As organism-driven landform modifications are sometimes to the advantage of the organism, some
of these landform features have become adaptive functional components of ecosystems, concurrently affecting
and responding to ecological and evolutionary processes. These recent eco-evolutionary insights, focused on
feedback among geomorphologic, ecological and evolutionary processes, are currently leading to the emergence
of what has been called an ‘evolutionary geomorphology’, with explicit consideration of feedbacks among the
evolution of organisms, ecosystem structure and function and landform organization at the Earth surface. Here
we provide an overview in the form of a commentary of this emerging sub-discipline in geosciences and ask
whether the use of the term ‘evolutionary geomorphology’ is appropriate or rather misleading.
Keywords: evolutionary geomorphology • biogeomorphology • ecosystem engineer • niche construction • eco-evolutionary
dynamics
© Versita sp. z o.o.
1. The concept of evolutionary geomorphology
Organisms respond and contribute to the modification of
their physico-chemical environment from the micro- to the
global scale. As living organisms evolved over geological timescales, these biotic evolutionary changes affected
Earth surface processes and a variety of landforms adjusted to the new evolving life forms [1, 2]. In turn, certain
geomorphic modifications fed back to community structure
and function as well as organism evolution.
Thus, long-term feedback systems comprise an evolution∗
ary dimension, i.e. the effects on and responses of vegetation, microorganisms and multicellular animals [2]. The
responses of living organisms to long-term geomorphic and
ecological feedback have to be considered as the result of
passive biotic evolutionary processes acting on individuals. The development of new adaptations to modify environmental constraints should not be considered as an ‘active choice’ of building an improved physical environment.
Biotic evolution, in Darwin’s sense, strictly occurs as a
consequence of natural selection at the organism level,
where genotypic mutations, biotic and abiotic environmental selection pressures (e.g. physical disturbances and
stress, presence of predators, inter- and/or intra-species
competition) and species adaptation (i.e. the development
of biological traits beneficial for the survival and reproduction of the organism or population) are involved [3].
Above the level of organisms, i.e. at the community and
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Figure 1.
Feedback model of reciprocal interactions and adjustments between organisms and Earth surface processes and landforms. The model
integrates in an eco-evolutionary framework geomorphic, ecological and evolutionary processes. Black colour represents physicochemical elements and controls. Grey colour represents living organisms and biological controls. Earth surface processes, landforms
and living organisms co-adjust according to feedback mechanisms.
ecosystem levels, self-adjustment processes are involved
in the organization of structures (e.g. community assemblages, landforms) and in the regulation of matter and
energy fluxes.
The concept of ‘evolutionary geomorphology’ with explicit
consideration of feedback among the evolution of organisms, ecosystem structure and function, as well as landform organization, was proposed earlier by the authors
[4]. At first, the focus was on the role played by vegetation dynamics as a biotic force in addition to the main
abiotic forces (i.e. gravity, solar energy, tectonics, chemical weathering) in shaping the Earth’s surface, but it is
now being extended to other organisms. Overall, this can
be considered as an eco-evolutionary one, i.e. a framework that examines the interplay between ecology and
evolution at multiple scales (i.e. eco-evolutionary dynamics, sensu [5]), which also comprises geomorphological coadjustments. It aims (i) to describe niche constructing
activities (sensu [6]) with evolutionary consequences for
engineer species that modify Earth surface landforms to
their advantage or to the advantage of other species in the
ecosystem, and (ii) to explain the long-term (i.e. organism adaptation) and short-term biological responses (i.e.
community structure and function) to changes which engineer species can induce in the geomorphic dimensions and
dynamics of their niche. This biogeomorphological framework is therefore closely linked to the ecological concept
of ‘engineer species’ or ‘ecosystem engineers’ developed
by ecologists [7, 8] and not to be confounded with ‘civil engineering’, even though the authors of the concept consider
Homo sapiens to be an ecosystem engineer. According to
[8], physical ecosystem engineers are organisms that directly or indirectly control the availability of resources to
other organisms by causing physical state changes in biotic or abiotic materials. Physical ecosystem engineering
by organisms is the physical modification, maintenance or
creation of habitats. The concept of ecosystem engineers
was more recently integrated into the emerging concept
of ‘evolutionary geomorphology’ [4].
We tentatively define ‘evolutionary geomorphology’ as the
creation, modulation and adjustment of landforms involving physical ecosystem engineers (sensu [8]), which implicate natural selection pressures on organisms through
niche construction (sensu [6]) with positive or negative
evolutionary consequences. The conceptual basis of
‘evolutionary geomorphology’ is presented in the feedback model of reciprocal interactions and adjustments
between organisms, Earth surface processes and landforms (Fig. 1). The feedback model differentiates between
physico-chemical elements and controls on the one hand
and living organisms and associated biotic controls on the
other. According to eco-evolutionary dynamics, biological
traits and community structure and function are respectively selected and modulated: (i) through organism evolution at the species level (e.g. genotypic mutations, coevolution between predator and prey); (ii) through Earth
surface processes, e.g. erosion or sedimentation, which
modify the habitat and constitute environmental selection
pressures on organisms; and (iii) through landforms which
constitute characteristic habitats for organisms in conjunction with specific geomorphic characteristics (e.g. texture, relative altitude, slope, exposure to regular flooding).
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The emergence of an ‘evolutionary geomorphology’?
From a geomorphological perspective, change induced by
organisms, notably ecosystem engineers, is (i) an active
construction or modification of landforms (e.g. through
burrowing, digging, construction of structures, weathering of rocks and substrate); and (ii) a passive modulation
of Earth surface processes through the modulation of morphogenetic factors, such as gravity, water flow, wind, frost,
chemical alteration (e.g. stabilization of soil by roots, retention of fine sediment by vegetation on hillslopes) [9].
The term ‘modulation’ implies a modifying or controlling
influence, which can be biotic or abiotic.
2. ‘Evolutionary geomorphology’ –
an appropriate term?
The perspective presented here has recently contributed
to the consideration of many Earth surface landforms not
only as abiotic structures adjusting to intrinsic and extrinsic physico-chemical constraints but also potentially
functional components of ecosystems – ‘functional’ in its
ecological meaning, i.e. referring to what an element
does in the context of its ecosystem - integrated in a network of reciprocal interactions and adjustments between
physico-chemical, ecological and evolutionary processes
[1, 10]. However, it could still be argued that describing landscapes as results of evolutionary processes seems
inappropriate and that the term ‘evolutionary geomorphology’ might be misleading or even constitute an oxymoron.
Without the presence of biota, landforms clearly are physical features at both local and regional scales, and morphodynamics are expressions of a trade-off between physical and chemical forces of resistance [4], exhibiting change
over different timescales. The central question here is
whether this change over time – when it is not solely controlled by physico-chemical forces but also by evolving
engineer species (i.e. within bioclimatic contexts favorable
to living organisms) – can be referred to as ‘evolutionary
geomorphology’.
Following [2], we point out that, in the presence of
biota, the consideration of the interrelationships between
landform geomorphology and organisms on evolutionary timescales provides an argument for employing the
term ‘evolutionary geomorphology’. This is in the restricted context proposed here, with a strict boundary between natural selection acting at the organism level and
other processes, such as self-adjustment, acting at higher
ecosystem levels. Thus, the juxtaposition of the two words
‘evolutionary’ and ‘geomorphology’, which in themselves
have strong connotations and seemingly contradict each
other initially, becomes appropriate when considering the
evolution of organisms in the Darwinian sense (natural selection restricted to the organism level) on the one hand
and landscape adjustments on the other hand. The twoway coupling between natural selection and landform adjustment over evolutionary timescales finally leads to the
notion and concept of ‘evolutionary geomorphology’.
However, we definitely refute the term ‘co-evolution’ when
considering feedback between organisms and landforms,
since evolution occurs in biota and not in landforms, which
adjust to abiotic and biotic controls. Co-evolution sensu
stricto, i.e. in its biological meaning referring to species
adaptations, corresponds to co-evolving traits between
different living organisms connected to each other within
the ecosystem (e.g. predator-prey, host-symbiont or hostparasite relationships). Landforms do not reproduce and
evolve in the Darwinian sense and, as pointed out by [6],
abiotic components lack genes or any equivalent heritable information. Furthermore, natural selection does not
apply to the selection of efficient structures and functions
above the organism level, therefore excluding communities,
ecosystems and landforms. Organisms evolve through natural selection and landforms adjust to physico-chemical
and biotic constraints. However, landforms affect natural
selection and in return are affected by natural selection
through the evolution of engineering traits of organisms.
This feedback mechanism leads to the emergence of dynamic and adjusting biogeomorphic structures at various
spatial scales.
3. Co-adjustment of river styles and
vegetation dynamics at geological
timescales
The consideration of the relationship between vegetation and geomorphic dynamics at aquatic-terrestrial interfaces has been particularly fruitful for linking geomorphology and biota over ecological and evolutionary timescales
[4, 11, 12]. Indeed, plants progressively evolved biomechanical and physiological traits and strategies that favored their establishment and propagation within coastal
and fluvial environments [13]. The latter are subject to persistent sediment, water and energy fluxes, which require
plants to withstand hydraulic selection pressures. Biological traits developed to better survive under these constraints are characterized by: (i) the resistance to breakage and uprooting, and development of root physiology
plasticity (e.g. large stem cross sections, strengthening
tissues, deep rooting systems and respiration facilities
during floods); (ii) the passive prevention of physical damage (e.g. stem and leave flexibility, size and shape, canopy
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reconfiguration abilities to reduce the hydrodynamic force
during floods, brittle twig bases that enable living stems
to break free); and (iii) the resilience to mechanical destruction or sediment burial (e.g. resprouting from both
roots and damaged shoots, clonal growth, buoyant seeds
and fragments) [14]. By favoring vegetation establishment within river margins, these resistance and resilience
strategies contribute to increase river bank cohesiveness
and fine sediment and nutrient retention within stable islands and floodplains [15]. An extensive and decisive modification of fluvial dynamics and river styles at geological
timescales associated with the evolution of plants occurred
during the Early Palaeozoic period [12, 16]. Evolving
plants and large woody debris converted shallow braided
river styles into single-thread meandering channels and
into island braided and anastomosing channels. Concomitantly, engineering populations of plants evolved new life
history traits (e.g. new timing of their life cycle and morphology) as a result of the selection pressures they induced via their hydrogeomorphic modifications within fluvial corridors (e.g. decrease of exposure to hydraulic constraints).
4. Biotic signatures in the Earth’s
topography
As suggested by Dietrich and Perron [17], the success in
identifying unique signatures of life on topography, i.e.
a landform that could only exist in the presence of life,
certainly depends on the occurrence scale and frequency
of certain landform properties and the technical possibility to detect them through high-resolution topographic
data. Unique signatures, such as animal and insect nests,
mounds, burrows and galleries, very frequently occur at
the sub-metre scale. A significant proportion of the Earth
surface is subject to such activities with long-term cumulative impacts. Furthermore, life itself can constitute unique
signatures on the topography at large scales, through autogenic bioconstruction activities. For example, Rudist
and coral reefs respectively represent fossil and existing topographic signatures of life on the Earth surface
at a large scale and have varied in the extent of their
development at different periods in Earth history. Dietrich and Perron [17] excluded such constructional features from their approach, defining these structures as
‘life itself’ rather than the influence of life on topography
through the mediation of sediment erosion, transport and
deposition processes. In line with that restriction, they
argued that unique signatures of life on the topography
of the Earth’s surface may not exist above the sub-metre
scale. However, even though certain landforms, such as
dunes, alluvial bars, single meandering channels, rounded
hillslopes and hydrological networks, would exist without life [17], many structural and dynamic properties of
these landforms are modulated by the effects of engineer
species.
5. Constructed
landforms
as
adaptive functional components of
ecosystems
Even if we accept the conservative delineation of Dietrich
and Perron [17], we suggest that the role of ecosystem
engineers played in landform evolution can be significant.
In biotic contexts, within river corridors [18], dune systems
[19], and salt marshes [20], the texture, size, shape and
temporal changes of alluvial or intertidal bars and dunes
are modified by the engineering morphological, biomechanical and life history attributes of pioneer plants. The
above-mentioned fact is also illustrated by animals influencing river systems and saltmarshes, for example dam
constructing beavers [21], crayfish constructing pits and
mounds on gravel substrates [22], aquatic mollusks adding
physical structure to the environment via shell production
and resulting reefs [23] or burrowing crabs [24]. These
modulations divert landform texture, morphology and dynamics from the state they would reveal in strictly abiotic
conditions. We point out that in certain cases these landform modulations are favorable for the engineer species
themselves and at the same time for many other species
which have adapted to the constructed ecosystem [2, 7].
We suppose that ecosystem engineer dynamics may have
positive, negative or no specific effects on the engineering organism and/or other organisms. The examples discussed here are positive ecosystem engineer effects, where
ecosystem engineering provides a benefit to the engineer
or other species [8]. Nevertheless, positive niche construction as defined by [6] – i.e. on average increasing
the fitness of the niche-constructing organism itself – can
have more or less negative impacts on other species in
the ecosystem. These impacts may enhance evolutionary
responses of the affected species, or lead to their exclusion from the ecosystem if they lack the capacity to adapt
to the modified environmental conditions. The construction of a beaver dam, for example, shows such negative
impacts when terrestrial organisms are submerged by the
newly formed pond or lake. However, the question of to
which extent ecosystem engineering has negative effects
on the ecosystem engineer itself seems more difficult to
answer. [6] defines negative niche construction as niche-
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The emergence of an ‘evolutionary geomorphology’?
constructing activities that change environments in such
a way that engineer specie fitness is reduced, e.g. by
inducing a discordance between the organism and its environment.
We shall now attempt to discuss this point by using the example of pioneer riparian trees as ecosystem engineers in
fluvial corridors [25]. In order to colonize and reproduce
within hydrogeomorphologically disturbed river systems,
pioneer riparian trees (e.g. cottonwood) evolved biological adaptations, such as high bending stability and strong
anchorage, allowing them to withstand floods, or rapid root
elongation after germination, reducing drought mortality
during flood recession and causing rapid substrate stabilization after germination [26]. Once colonized, these pioneer trees increase surface roughness and favor sediment
accretion, thus actually enhancing the biogeomorphological succession [27]. This in turn eventually leads to the exclusion of this particular pioneer engineer species, which
depends on hydrogeomorphic disturbances for recruitment.
However, the disturbance regime will decrease and eventually disappear because under biotic (here: vegetation)
control, the fluvial corridor tends to raise its topography
and stabilize. If river systems were not naturally dynamic
systems or if the resistance of engineer plants to mechanical constraints were too strong, it could be argued that
this particular engineer species would be excluded from
wide areas or could even become extinct due to its own
ecosystem engineering within fluvial corridors. We suggest that this is not the case as long as river systems
remain dynamic, providing suitable areas for colonization
by new generations of riparian pioneer trees, and as long
as the resistance traits of these riparian plants do not become so strong as to completely stabilize the river system.
Furthermore, certain riparian trees, such as cottonwood,
evolve traits related to the modifications they induce in
the fluvial environment. The stabilization and construction of landforms combined with local accumulation of resources lead to development as a feedback of competitive
adult traits, providing advantages for resource uptake (e.g.
high size and deep root system). Cottonwood currently
combines ‘r’ (ruderal opportunistic) traits at the juvenile
stage with K (competitive) traits at the adult stage when
it reaches its sexual maturity on stabilized surfaces. The
initial abiotic conditions selected the r traits in the long
term, whereas the K traits may correspond to an evolutionary feedback associated with engineer effects [28].
Thus, even if the described engineer effects may seem to
be negative in the short term, they appear to be rather
positive in the long term.
As in the case of riparian trees, the properties of many
landforms do not only adjust to external physical and
chemical forces but are also engineered by life. As a re-
action to the effects and responses of engineer organisms,
modulated landforms become adaptive functional components embedded within the biogeomorphic system (i.e. an
abiotic landform and its associated community, including
engineer species), e.g. by way of positive niche construction sensu [6], become adaptive functional components embedded within the biogeomorphic system (i.e. an abiotic
landform and its associated community, including engineer
species). Ultimately, these modulations can become real
extended phenotypes of the constructing organism, sensu
Dawkins [29] as suggested for ponds created by beavers.
According to [29], ecosystem engineering traits and their
effect on the physical environment and geomorphology can
thus, over evolutionary time, be encoded in the genetics
of the engineer. Biogeomorphic structures (e.g. fluvial islands, galleries in soil, ant or termite mounds, coral reefs)
are not explicitly written in the genotype of the engineer
species, but they emerge from the interactions between
the organisms and the physical environment [2]. In our
opinion, the feedback leading to extended phenotypes of
organisms provides a further argument for suggesting the
term of ‘evolutionary geomorphology’ in the context presented here.
6.
Perspectives
Even if landforms engineered by life are not ’unique’ topographic signatures, the emerging landform properties
adjusted to organisms’ activities must be considered to be
of crucial importance because they contribute to complex
interactions and reciprocal adjustments between ecological and evolutionary processes within the ecosystem. We
therefore suggest that focusing on these geomorphologic
modulations, rather than on the search of a wholly unique
signature of life on topography, will constitute a major
issue for geomorphologists working on the Critical Zone.
We are only just beginning to understand how life affects landform dynamics over geological timescales and
how, in turn, landforms or properties of landforms modify ecosystem structure and function through the modification of organism fitness. Conceptual advances [4] and
recent findings showing that engineered alluvial facies
are the signature of feedback between hydrogeomorphic
processes and plant evolution [12, 16] support the emergence of what we have called ‘evolutionary geomorphology’. Nevertheless, we are aware that further exchanges
and collaborations among geomorphologists, notably biogeomorphologists, ecologists and especially evolutionary
biologists are required to support our as yet limited conceptual framework. As suggested by [9], inter-disciplinary
studies have to be designed to explicitly examine the re-
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Johannes Steiger, Dov Corenblit
cursive influences among genes, phenotypes and geomorphic processes and structures on the Earth.
We suggest that to identify the role of intraspecific genetic variability of organism populations related to engineer traits in order to quantify the variability in abiotic responses and resulting feedbacks on engineer populations and communities will be a future challenge (see
also [2]). This will require a formal examination of how,
and to what degree, genetic variation in engineer species
may influence physical habitat properties and thus community structure and function in the short term, as well
as adaptation and speciation over evolutionary timescales,
especially based on paleontological evidence. Genetically
variable engineer traits that could be associated with crucial biogeomorphologic properties have to be identified by
combining field or laboratory studies. The links among
specific variations of these traits, specific landform modulations, associated community structures and ecological
responses have yet to be established.
We are aware that these suggestions - ranging from
designing inter-disciplinary studies to linking particular
landform modulations with intraspecific genetic variability of engineer ecosystems and biological life traits – are
still in the theoretical stage. We are confident, however,
that more specific and more practical recommendations
will emerge from inter-disciplinary research efforts.
7.
[4]
[5]
[6]
[7]
[8]
[9]
[10]
[11]
Acknowledgments
[12]
The authors wish to thank all four reviewers for their helpful and constructive suggestions and discussions of this
commentary, which helped us advance our own thoughts
and reflections. We also thank S. Lühmann for English
proofreading of the manuscript.
[13]
[14]
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