Hydrological Sciences Journal

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Global hydrological models: a review

Aditya Sood & Vladimir Smakhtin

To cite this article: Aditya Sood & Vladimir Smakhtin (2015) Global hydrological models: a review,
Hydrological Sciences Journal, 60:4, 549-565, DOI: 10.1080/02626667.2014.950580

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 Hydrological Sciences Journal – Journal des Sciences Hydrologiques, 60 (4) 2015 549
http://dx.doi.org/10.1080/02626667.2014.950580

Global hydrological models: a review

Aditya Sood and Vladimir Smakhtin
International Water Management Institute (IWMI), Colombo, Sri Lanka
a.sood@cgiar.org

Received 20 June 2013; accepted 30 April 2014

Editor D. Koutsoyiannis

Abstract Global hydrological models (GHMs) have effectively become a separate research field in the last two
decades. The paper reviews and compares 12 known global modelling efforts since 1989, the year the first GHM
was published. Structure, strengths and weaknesses of individual models are examined, and the objectives of
model development and their initial applications are documented. Issues such as model uncertainty, data scarcity,
integration with remote sensing data and spatial resolution are discussed.
Key words global hydrological models; grids; remote sensing; hydrology

Revue des modèles hydrologiques globaux

Résumé Les modèles hydrologiques globaux (MHG) sont en fait devenus un domaine de recherche distinct au
cours des deux dernières décennies. Cet article examine et compare 12 démarches de modélisation globale
connues depuis 1989, année de publication du premier MHG. La structure, les forces et faiblesses de chaque
modèle ont été examinées et les objectifs de développement des modèles et leurs premières applications ont été
décrits. Des questions telles que l’incertitude du modèle, la rareté des données, l’intégration des données de
télédétection et la résolution spatiale ont été discutées.
Mots clefs modèles hydrologiques globaux ; maillages ; télédétection ; hydrologie

INTRODUCTION and economic implications. Due to globalization,

The continuing debate on climate change (e.g.
Kundzewicz and Stakhiv 2010, Huard 2011) and
other global drivers of change highlights the interde-
pendence of various earth systems and the need for
integration of those systems into global simulation
models (Wilby 2010). There is an impact on regional
and, hence, global climate, from changes in soil moist-
ure and terrestrial evapotranspiration (Munro
et al. 1998, Koster et al. 2003, Koster et al. 2004,
Seneviratne et al. 2010, Dirmeyer 2011), and hence
river discharge which in turn impacts on sea character-
istics (Milly et al. 2010). Land-use changes upstream
affect the hydrology and water quality thousands of
miles downstream (Freeman et al. 2007). Global
hydrology is closely linked with the nutrient cycle
(cause of eutrophication of coastal zones) (Foley
et al. 2005, Rabalais et al. 2009, Fekete et al. 2010)
and the carbon cycle (impacting on the climate). The
changes in these global cycles eventually have social
virtual water trade, for example, has become an impor-
tant factor of both the global water cycle and food
security (Islam et al. 2006). While the scientific com-
munity has long been aware of, and has dealt with,
impacts of climate on hydrology, the feedback has
only recently started to receive attention. All this
leads to the notion of the ‘global water system’
(Alcamo et al. 2008, Alcamo 2009), in which the
global water flow is connected to other systems
through physical relationships, economics and institu-
tions. This system is further complicated by the inter-
ference from humans through water storage and
withdrawals (Rost et al. 2008b).
Models that attempt to simulate global hydrol-
ogy and associated processes are similar to numerous
stand-alone hydrological models and to hydrological
components of the general circulation models
(GCMs). However, they differ in the detail of
description of processes, parameter estimation

© 2015 IAHS
550 Aditya Sood and Vladimir Smakhtin
approaches, time scales, and spatial resolution of
input data and simulations (Haddeland et al. 2011).
The stand-alone models are usually applied at the
basin scale, or a smaller catchment scale, and have
many parameters that need to be calibrated or esti-
mated regionally. Examples of such models include
the Soil and Water Assessment Tool (SWAT; Neitsch
et al. 2002), the Hydrological Simulation Program-
Fortran (HSPF; Bicknell et al. 1997) and the
Hydrologiska Byråns Vattenbalansavdelning (HBV;
Lindström et al. 1997). Some of these models can,
in principle, be applied at the global scale, but, due to
severe information constraints, this never happens in
practice. The hydrological models that are a part of
GCMs are usually land surface schemes (LSSs) that
simulate the energy balance at soil, atmosphere and
vegetation interfaces at finer time scales (often
hours), and do not have a flow routing component.
Examples of such models include the Biosphere–
Atmosphere Transfer Scheme (BATS; Dickinson
et al. 1986), the Simple Biosphere Model (SiB;
Sellers et al. 1986) and the Joint UK Land
Environment Simulator (JULES; http://www.jchmr.
org/jules/index.html).
The actual global hydrological models (GHMs)
have few calibratable parameters and are calibrated
either at eco-region, climatic-region or large river
basin scales (Vörösmarty et al. 1989, Döll
et al. 2003, Widén-Nilsson et al. 2007). Some models,
such as the Water Balance Model – Water Transport
Model (WBM-WTM; Vörösmarty et al. 1998), are not
calibrated per se but have an adjustment factor to tune
them. The model Water – Global Analysis and
Prognosis (WaterGAP; Alcamo et al. 2003, Döll
et al. 2003) is a combination of calibration and tuning.
It is first calibrated with a single parameter against
streamflow. The basins that underestimate or overesti-
mate flows are then tuned by two adjustment factors,
runoff and discharge correction (Döll et al. 2003). The
spatial resolution of GHMs is defined by the resolu-
tion of available global climate input data. GHMs are
relatively new and have emerged in the last two dec-
ades. Lately, there has been increasing activity in this
field and a concerted effort is evolving (Lawford
et al. 2004, Döll et al. 2008). Also, GHMs are becom-
ing more complex and resolute as more functionality
is added to them and finer global spatial datasets are
becoming available. Issues of the sensitivity of models
to varying spatial and temporal scales of input and
output data, measuring modelling uncertainty and cou-
pling GHMs with other models have become promi-
nent (Döll et al. 2008, Voß et al. 2008).
The explosion of global data availability from
satellites in the last two decades (Tang et al. 2009)
has had an influence on the development of GHMs.
Although some GHMs have been applied at a range
of scales, they are, as a rule, built for global-scale
studies. They would not be the preferred choice in
basin-scale applications, due to the coarser resolution
of GHMs at present, and the fact that there is a large
family of hydrological models that have been
designed for this purpose. However, GHMs may
provide valuable spatial and temporal estimates of
global water resources, and help to analyse possible
projections/scenarios of changes of those estimates;
GHMs have been built effectively for this purpose.
Global estimates obtained through GHMs could be
an improvement over those simply based on the
statistical analysis of ground-based observed data,
which, at a global scale, remain limited and, hence,
contain a lot of uncertainty (Rodda 1995). An even
greater use of GHMs is revealed when they are
linked with other models, i.e. describing global econ-
omy, ecology, trade, biodiversity, energy balance,
land-use change, climate change, crop growth and
other development issues/components related to
water (Vörösmarty et al. 2000, Islam et al. 2006,
Alcamo 2009). The requirements from the GHMs
depend upon the demands of such associated models.
To the best of the authors’ knowledge, a single
source/compilation of GHMs does not exist.
Recently, Trambauer et al. (2013) examined GHMs
in the context of drought forecasting in Africa. This
paper extends the discussion to the broader context of
developing simulation tools for global food trade,
agriculture and economy, and examines constraints
and development trends of GHMs, including those of
uncertainties, input data quality and integration with
remote sensing data.
EXISTING GLOBAL HYDROLOGICAL
MODELS—DESCRIPTION
Over the last few decades, many isolated efforts were
made to simulate the global hydrological cycle. A
review of the existing literature suggests that there
are at least 12 GHMs (or tools that can be interpreted
as such) at present. Table 1 lists these GHMs in
chronological order of their development and
includes details of the main developers, their objec-
tives for developing the model and some example
applications. Table 2 summarizes the technical details
of these models.
Table 1 Origin and known applications of existing global hydrological models.
No. Model Developed/maintained by Year Objective(s) Applications

1 HDTM 1.0 (HydroDynamic Model) University of New Hampshire,
/ WBMplus / WBM-WTM (Water
Balance Model - Water Transport
Model)
2 Macro-PDM (Macro Probability
Distribution Model)
3 MPI-HM (Max Planck Institute –
Hydrology Model)
4 GWAVA (Global Water Availability Centre for Ecology & Hydrology 1999
Assessment model)
USA
University of Reading, UK
Max-Planck-Institut für 1998
Meteorologie (Max Planck
Institute for Meteorology),
Germany
(formerly Institute of
1989 To study global biogeochemical 1.
cycles. Linked to Terrestrial
Ecosystem Model and Trace Gas 2.
Model through soil moisture and
evapotranspiration
3.
1998 To build a macro-scale global
model. Among the initial efforts,
only one model existed before:
WBM. This uses the VIC
principle on a global scale
To define lateral flow of water from Validation of the weather forecasting models, European Centre
continents to oceans and to link
with a GCM (ECHAM)
To study progression of global water 1. Estimation of water scarcity for eastern and southern
scarcity due to population
Comparison of PET methods on US watersheds—WBM
(Vörösmarty et al. 1998)
Using GHM in an application of isotope tracers at con-
tinental scale in hydrological modelling (Fekete
et al. 2006)
Calculation of variability and uncertainty in the global
irrigation water demand—WBMplus (Wisser et al. 2008)
4. Analysis of the effect of climate and hydrological altera-
tion in hydrological cycle to study nutrient transport
(Fekete et al. 2010)
5. Using GHM to calibrate remote sensing signal to dis-
charge (Brakenridge et al. 2012)
1. Measurement of uncertainty in an ensemble with 21
GCMs at global scale using a campus-wide computer
grid (Gosling et al. 2010)
2. Study of the impact of climate change on river flow
regimes (Arnell and Gosling 2013)
for Medium-Range Weather Forecasts Re-Analysis (ERA)
and National Center for Environmental Prediction Re-
Analysis (NRA) (Hagemann and Dümenil 2001)
Africa (Meigh et al. 1999)
Global hydrological models: a review

Hydrology), UK
5 VIC (Variable Infiltration Capacity
model)
6 LAD (Land Dynamic model)
University of Washington, 2001
Seattle, USA
National Oceanic and 2002
Atmospheric Administration
(NOAA) / Geophysical Fluid
Dynamics Laboratory,
Princeton, New Jersey, USA
increase and climate change
To improve on the previous model 1. Estimation of global soil moisture content (Nijssen
by incorporating two layers in
soils and introducing sub-grid
heterogeneity for vegetation, soil
moisture storage capacity and
precipitation
To improve the energy and water
balance of the older model
2. Measurement of the impact of climate and land-use
change (Meigh and Tate 2002)
et al. 2001b)
2. Prediction of discharge of the world’s rivers (Nijssen
et al. 2001a)
3. Evaluation of the Atmospheric Model Intercomparison
Project (AMIP II) (Irannejad and Henderson-Sellers 2007)
1. Illustrating an approach of land model evaluation with
precipitation uncertainty (Milly and Shmakin 2002)
2. Measurement of the inter-annual variation in river dis-
charges (Shmakin et al. 2002)
(Continued )
551

Table 1 (Continued).
No. Model
7 WaterGAP (Water – Global Analysis University of Kassel; University 2003 To combine water availability and
and Prognosis model)
8 PCR-GLOBWB (PCRaster GLOBal Utrecht University, The
Water Balance model)
Developed/maintained by
of Frankfurt, Germany
Netherlands
Year Objective(s)
water use based on structural and
technological changes at global
scale
2004 To add advance routine for grid
heterogeneity for surface runoff,
inter-flow and groundwater; also
added routine for lateral transport 2.
of heat by water
552
Applications
1. Identification of regions in which water resources have
higher sensitivity to global change—critical regions
(Alcamo and Henrichs 2002)
2. Global modelling of irrigation water requirements (Döll
and Siebert 2002)
3. Calculation of water availability indicators (Döll
et al. 2003)
4. Integration of Gravity Recovery and Climate Experiment
(GRACE) data into global hydrological models (Werth
et al. 2009)
5. Analysis of river flow alterations due to water withdra-
wals and reservoirs (Döll et al. 2009)
6. Simulation of continental water storage variation at the
global scale (Werth and Güntner 2010)
7. Computing the water mass variations to calculate load-
induced deformation of the Earth’s crust (Fritsche
et al. 2012)
8. Studying the impact of groundwater and surface water
withdrawals on water storage (Döll et al. 2012)
9. Studying large-scale variation characteristics of global
groundwater (Jin and Feng 2013)
10. Calculating seasonal water storage variations as
impacted by water abstractions (Döll et al. 2014)
1. Measurement of the skill of seasonal predictability of river
Aditya Sood and Vladimir Smakhtin
discharge for different European rivers (Bierkens and van
Beek 2009)
Analysis of the depletion of global groundwater resources
(Wada et al. 2010)
3. Modelling methane (CH4) emissions of boreal and arctic
wetlands (Petrescu et al. 2010)
4. Analysis of future global runoff changes (Sperna
Weilanda et al. 2011)
5. Calculation of global water stress (van Beek et al. 2011,
Wada et al. 2011)
6. Study of the impact of human abstraction of surface water
and groundwater on streamflow (de Graaf et al. 2014)
(Continued )
Table 1 (Continued).
No. Model
9 LPJmL (Lund-Potsdam-Jena
managed Land model)
10 WASMOD-M (Water And Snow
balance MODeling system)
11 H08 (H07)
12 ISBA-TRIP (Interactions between
Soil, Biosphere, and Atmosphere
– Total Runoff Integrating
Pathways)
Developed/maintained by
Potsdam Institute for Climate
Impact Research (PIK),
Germany
Department of Earth Sciences,
Air, Water and Landscape
Sciences, Uppsala University,
Sweden
National Institute of
Environmental Studies,
University of Tokyo, Japan
Centre National de Recherchés
Météorologiques, France
Year Objective(s)
2007 To simulate the spatial and temporal 1.
dynamics of global vegetation,
and study its impact on global 2.
hydrological and carbon cycles.
3.
2007 To complement existing global
models and to come up with a set
of minimum parameters for
gauged and ungauged river basins
2008 To assess global water availability at
a sub-annual time-scale
2010 To measure continent-level
terrestrial water storage and to use
GRACE data to validate it.
Applications
Calculation of green water flows—LPJ (Gerten
et al. 2005)
Measurement of global carbon fluxes and pools (Bondeau
et al. 2007)
Checking the uncertainty of the terrestrial carbon and
water cycle at different spatial resolutions using the LPJ-
Dynamic Global Vegetation Model (LPJ-DGVM) (Müller
and Lucht 2007)
4. Studying human alteration of the terrestrial water cycle
through land management (Rost et al. 2008b)
5. Calculation of agricultural green and blue water consump-
tion (Rost et al. 2008a)
6. Calculation of global-scale water withdrawal, allocations
and consumptive use for surface water and groundwater
(Wada et al. 2014)
1. Comparison of two flow network-response functions to
evaluate the improvement in runoff routing (Gong
et al. 2009)
2. Comparison of two precipitation datasets (TRMM and
WATCH Forcing Data) in southern Africa (Li et al. 2013)
Global hydrological models: a review
1. Calculation of national agriculture water withdrawals and
locating water-stressed regions (Hanasaki et al. 2008b)
2. Estimation of global virtual water flows (Hanasaki
et al. 2010)
3. Analysis of scenarios for different socio-economic condi-
tions to calculate water availability and scarcity (Hanasaki
et al. 2013)
Measurement of terrestrial water storage and its seasonal and
inter-annual variability (Alkama et al. 2010)
553
 554

Table 2 Technical features of existing global hydrological models.

No. Model Spatial Temporal PET Reservoirs Irrigation Crop/vegetation Soil layers Number of
resolution resolution / model parameters
simulation time
step1

1 HDTM 1.0 0.5° Monthly / PET, Yes Yes Three vegetation types: forest, grassland and Single layer till >4
month Korzoun et al. (1977) shrubland rooting depth
considered
2 Macro-PDM 0.5° Daily / day Penman, Penman- No No Forest and grassland only 1 8
Monteith and
Priestley-Taylor
3 MPI-HM 0.5° Daily / day Thornthwaite formula No No None 1 3
4 GWAVA 0.5° / Monthly / day Yes Yes Trees, bushes, grass and bare soil
0.1°
5 VIC 2° Daily / day Penman-Monteith No No 13 land cover + water 2–3 Original VIC (not
global) (Liang
et al. 1994): 22
parameters
6 LaD 1° Daily / hour Custom equation as part No No 32 vegetation types of 5
of energy balance Matthews (1983) reduced to 10 and put into a
1° resolution global map
7 WaterGAP 0.5° Monthly / day Priestley-Taylor Yes Yes 16 land cover types 1 36
8 PCR- 0.5° Daily / day Penman-Monteith Yes No Three categories: natural vegetation, rainfed 3 layers: 2
GLOBWB crops and irrigated crops; these are further upper soil
subdivided into tall and short vegetation layers and
one
groundwater
layer
Aditya Sood and Vladimir Smakhtin

9 LPJmL 0.5° Daily / day Priestley-Taylor Yes Yes Natural vegetation: nine plant functional types; 2
agricultural vegetation: 12 crop functional
types
10 WASMOD- 0.5° Monthly / Custom equation based No No None 1 4–6 (based on
M month on air temperature presence of
and relative humidity snow)
11 H08 (H07) 1° / 0.5° Daily / 3-h Custom equation based Yes Yes Crop growth model used similar to SWIM 1
on energy balance
12 ISBA-TRIP 1° Daily / ISBA: Custom equation based No No Coordination of Information on the 3 2 (primary) and 14
20 min, on water balance in Environment (CORINE) secondary that
TRIP: 1 h the soil layer can be derived
from primary
Note: Empty boxes imply missing information.
1
Simulation time step is the smallest time unit at which the model operates, whereas temporal resolution is the time interval at which the output is provided.
 Global hydrological models: a review
An initial effort in the development of GHMs
was made by Vörösmarty et al. (1989), who devel-
oped the Water Balance Model (WBM) and linked it
with the Water Transport Model (WTM). The WBM
predicts soil moisture (SM), evapotranspiration (ET)
and surface runoff at grid level, and the WTM routes
the discharge generated in the grid cell through the
channel using a linear reservoir model (Vörösmarty
et al. 1989). The field capacity (FC) and, hence,
water availability are linked to the vegetation type
and soil at grid-level resolution. The WBM-WTM
model relies on a narrow range of parameters based
on those published in the literature and then iterates
annually until a dynamic steady state is achieved (i.e.
soil moisture, ET, runoff and snowmelt come within
an acceptable level of difference for successive runs).
It has been modified over the years to include irriga-
tion and reservoirs, accounting for the irrigated and
non-irrigated fraction, reservoir operations (WBMplus
class), effect of permafrost (Pan-Arctic Water
Balance Model (PWBM) and HydroDynamic Model
(HDTM 1.0), along with the multi-layer soil moisture
routine (HDTM 1.0; http://www.wsag.unh.edu/wbm.
html). The parameter values in the WBM are
assigned a priori and are not calibrated. In the
Macro Probability Distribution Model (Macro-PDM;
Arnell 1999), developed in 1998, streamflow is simu-
lated as direct runoff and delayed runoff. A similar
vegetation classification as in WBM is used, but the
soil moisture FC is statistically distributed within a
grid to account for sub-grid variations. The ratio
between ET and potential evapotranspiration (PET)
is described by a linear function of average soil
moisture in a grid cell. In this model, the values of
parameters are set from a literature review or pre-
vious model applications. Six out of 13 parameters
are globally uniform. Around the same time, the
Max-Planck Institute (MPI) in Germany developed
a model that coupled the European Centre Hamburg
Model (ECHAM) GCM with a hydrological dis-
charge (HD) model (Hagemann and Dümenil 1997).
This model is known as the Max Planck Institute-
Hydrology Model (MPI-HM). In this model, vegeta-
tion in each grid cell is partitioned based on vegeta-
tion maps, as developed by Matthews (1983), and the
fraction of such vegetation dependent on the avail-
able water content in the root zone (decreases below
40% of FC). The streamflow and surface runoff are
simulated as a cascade of linear reservoirs, whereas
the baseflow is described by a single reservoir. The
Global Water Availability Assessment model
(GWAVA; Meigh et al. 1999), developed to examine
555
issues of water scarcity, also used a probability dis-
tribution to simulate soil moisture storage. This
model includes water demand based on population,
livestock, irrigation schemes and industrial water use.
It uses four simplified land cover categories (trees,
bushes, grass and bare soil). Groundwater is first
explicitly calculated by using potential yield from a
well and estimating borehole density in each grid. It
is then compared to groundwater recharge calculated
through the surface flow model and a conservative
value of the two, based on aquifer classification, is
used.
All the above models treat soil as a single layer.
The Variable Infiltration Capacity model (VIC;
Nijssen et al. 2001b), developed in 2001, considers
two layers of soil and includes sub-grid variability in
land surface vegetation, soil moisture storage capa-
city and precipitation. The VIC model can run either
as an energy and water balance model or just as a
water balance model. Multiple land covers (13)
including bare soil are considered in the VIC
model. It also accounts for the top vegetation cover.
The second layer is modelled as a nonlinear storage.
Snow is simulated by a single layer energy and mass
balance model. A separate river routing model is
used. The Land Dynamics model (LaD; Milly and
Shmakin 2002) was developed to improve an older
water and energy balance model by Manabe (1969)
by adding sensible heat storage on land, groundwater
storage and stomata resistance. Grid cells are treated
as either glaciated or non-glaciated. Water storage is
considered in snow, glacier ice, root zone and
groundwater. Runoff is generated after the soil
reaches its water-holding capacity, and is then passed
through a groundwater reservoir (with some resi-
dence time) before becoming streamflow. The
PCRaster GLOBal Water Balance model (PCR-
GLOBWB; van Beek and Bierkens 2008), developed
in 2004, broke up the soil layers into three storage
‘buckets’, each contributing to the streamflow in the
form of surface runoff, interflow and baseflow,
respectively. The surface vegetation is simplified
into three categories—natural, rainfed and irrigated
—which are further classified as short (retrieving
moisture from the top soil layer) or tall vegetation
(retrieving moisture from the bottom layer).
Evapotranspiration is broken up into transpiration
by vegetation and evaporation from bare soil.
The Water – Global Analysis and Prognosis
model (WaterGAP; Alcamo et al. 2003, Döll
et al. 2003) and the Lund-Potsdam-Jena managed
Land model (LPJmL; Bondeau et al. 2007) are two
556 Aditya Sood and Vladimir Smakhtin
GHMs that were developed in Germany during the
past decade. While the WaterGAP model puts more
emphasis, at least at present, on water demand simu-
lation, the LPJmL model is more detailed in vegeta-
tion and crop simulations. The WaterGAP model is
made up of two modules—the GHM per se and the
global water use model. The latter incorporates
human water consumption (in the form of domestic,
manufacturing, thermal power production, livestock
and irrigation use). The GHM model considers soil as
a single layer and calculates the vertical and lateral
water balance within each grid. Surface and subsur-
face runoff are transported as a fast response, and the
baseflow (i.e. contribution from groundwater) is
transported as a slow response. The lateral movement
of runoff is described through multiple storages
representing local and global lakes, reservoirs, wet-
lands and streams. The LPJmL model was developed
by adding a hydrology component to an existing
dynamic global vegetation model (DGVM). The
vegetation routine of the model simulates photo-
synthesis, evapotranspiration, plant respiration and
the carbon cycle. It allows for nine natural plant
types and 12 crop types (both rainfed and irrigated).
There can be one stand of natural vegetation and
multiple stands of crop types within each grid.
Some of the inputs for the vegetation are dynamically
generated within the model. The hydrological routine
calculates water balance for each stand of the grid
and each grid flows into another via a river routing
module (Rost et al. 2008a). The evapotranspiration is
broken up into productive water consumption (plant
transpiration) and non-productive consumption
(interception loss and evaporation). Soil storage is
represented by two ‘buckets’.
The Water And Snow balance MODeling system
(WASMOD-M; Widén-Nilsson et al. 2007), originat-
ing from WASMOD (Xu 2002), uses a minimum
number (4–6) of calibrating parameters. WASMOD-
M runs at monthly time steps and separates runoff
into fast and slow components for each grid. It con-
siders rain, snowmelt, snow accumulation and evapo-
transpiration for each grid within its routine. The
parameters in the model are regionalized to simplify
model application. It has six parameters, of which
five can be calibrated and one is fixed (Widén-
Nilsson 2007). The H08 model (Hanasaki
et al. 2008a, 2008b) is a more detailed and integrated
system, made up of six modules—land surface
hydrology, river routing, crop growth, reservoir
operation, environmental flow requirements and
anthropogenic water withdrawal. The land-surface
hydrological module calculates both the hydrology
balance and energy balance. It uses the ‘leaky-
bucket’ concept to model soil moisture (Hanasaki
et al. 2008a). The river routing module uses the
Total Runoff Integrating Pathways model (TRIP;
(Oki and Sud 1998). The TRIP model is a network
of integrated pathways providing information on lat-
eral water movement over land. The crop routine
uses heat unit theory (Barnard 1948, Phillips 1950)
and is based on the Soil and Water Integrated Model
(SWIM; Krysanova et al. 1998). The basis of the heat
unit theory is that each plant has its own range of
temperature within which it grows. SWIM is a river
basin model developed by the Potsdam Institute for
Climate Impact Research (PIK), Germany, to simu-
late hydrology, sediments, nutrients and carbon
movement, along with plant growth and yield.
Nineteen crop types are used and large reservoirs
(with a storage capacity greater than 109 m3) are
incorporated.
The ISBA-TRIP model (Alkama et al. 2010)
was developed by combining two models—the mod-
ified land surface model by Noilhan and Planton
(1989) and the surface runoff pathway network
from TRIP, by Oki and Sud (1998). The heterogene-
ity within a grid is described by using a tile approach
that divides a grid into a series of sub-grid patches
based on land use and soil properties; partial grid
saturation is also accounted for. The model
calculates the surface energy and water budget for
each grid with the soil represented by three layers.
TRIP routes the daily discharge simulated by ISBA
into streamflow.
COMPARISION OF MODEL FEATURES
All GHMs run in a grid format. In most GHMs, the
spatial resolution is 0.5 degrees (just over 3100 km2
per grid cell at the Equator). The grid format and
resolution are dictated by the availability of global
meteorological data and computational resources.
The gridded format makes it convenient to manip-
ulate other available input data, which are also
usually available in grid format (e.g. originating
from remote sensing (RS) data). With faster compu-
tational devices and availability of finer resolution
input data, higher resolution GHMs are being
developed, such as a 5-min version of WaterGAP
(http://www.massentransporte.de/index.php?id=303),
6-min version of PCR-GLOBWB (called PCR-
GLOBWB 2.0: http://www.globalhydrology.nl/models/
pcr-globwb-2-0/) and the latest version of WBMplus that
 Global hydrological models: a review
can run at a range of resolutions dictated by the under-
lying gridded network (45″, 90″, 3′, 5′, 6′, 15′ –
B. Fekete, City University of New York (CUNY), pers.
comm.).
Most of the GHMs have a temporal resolution of
1 day despite the fact that the available input data
normally have a monthly time step. Historical data
covering the full 20th century, for example, are only
available at a monthly temporal resolution, which
necessitates statistical downscaling into a daily tem-
poral scale. With improvement of meteorological
reanalysis data, the models have started using more
of these as inputs. The fourth generation of ECMWF
reanalysis (ERA) daily datasets (from 1979 to date)
are being developed; ERA data have already been
used in some GHMs (Wada et al. 2010) and in model
intercomparison studies (Weedon et al. 2011).
Only five models—WBMplus, WaterGAP, PCR-
GLOBWB, LPJmL and H08—explicitly consider
reservoir storage. Out of these, PCR-GLOBWB
does not simulate irrigation. Some models consider
soil layers as a single storage unit (Macro-PDM,
MPI-HM, WASMOD-M, H08 and WaterGAP),
whereas others consider multiple storage layers
(PCR-GLOBWB: two layers, VIC: two layers,
LPJmL: two layers, ISBA-TRIP: three layers, and
LaD: five layers).
Only H08 and LPJmL have an explicit crop
growth model. Although all the models can handle
changes in meteorological data due to climate change
(such as precipitation and ET due to changes in
temperature), only the models that have crop growth
and/or vegetation models can deal with the impact on
plant physiology due to changes in carbon dioxide
concentration and increase in temperature. From that
perspective, LPJmL and H08 may be well suited to
evaluate the impact of climate change scenarios on
crop yield (and hence on hydrology). The LPJmL
modle simulates natural vegetation (along with
agriculture), and the proportion of different vegeta-
tion classes in each grid is based on relative advan-
tage in terms of light, water, space and other
environmental factors (Gerten 2013).
A few models, such as VIC, LaD, WaterGAP
and LPJmL, have a detailed land-use classification.
Other models have used a simplified classification,
for example, WBM-WTM uses only three categories
(i.e. forest, grassland and shrubland); and the newer
version, WBMplus, accounts for multiple categories
(e.g. Wisser et al. 2010 handle four categories). The
model can simulate up to 32 of the vegetation classes
of the Terrestrial Ecosystem model (Melillo
557
et al. 1993). The current version of WBM can read
the parameters related to the vegetation directly as
input from an external source and, hence, the model
can simulate as many classes as formed by the dif-
ferent parameter combinations (B. Fekete, pers.
comm.). Macro-PDM uses only forest and grassland
categories for land use, whereas PCR-GLOBWB
uses three categories: natural vegetation, rainfed
crops and irrigated crops, which are further divided
into short and tall vegetation (to simulate ET from
different layers of the soil profile).
Models also differ in terms of how they account
for energy balance, sub-grid effects and soil stratifi-
cation. About half of the models reviewed (MPI-HM,
PCR-GLOBWB, VIC, LaD, H08 and ISBA-TRIP)
have their own energy balance module. Some models
treat a grid as homogenous with respect to infiltra-
tion, climate data and vegetation, while others
account for sub-grid effects to describe heterogeneity
within a grid cell (VIC, ISBA-TRIP). Some consider
root-zone depth as the height of one of the soil layers
(such as ISBA-TRIP), whereas others have a fixed
depth of the first layer (PCR-GLOBWB: 0.3 m,
LPJmL: 0.5 m).
The main output of all the models is streamflow,
which is derived from partitioning precipitation into
evapotranspiration, soil moisture, fast and slow flow
(although the terms used may be different, e.g. sur-
face runoff, lateral flow and baseflow). The VIC,
WaterGAP, LPJmL and H08 models also look at
crop water requirements. These are the models that
also have a detailed land-use classification (Table 2).
All models provide output at grid level (mostly at
0.5° resolution at present). These grid-level outputs
are aggregated to river basin scale, based on the
number of grids that fall within a river basin bound-
ary. None of the models uses an incremental drainage
sub-basin discretization directly.
Global hydrological models are often developed
to support simulation of certain phenomena which
are not always hydrology per se (Table 1).
Consequently, they have certain areas of preference.
For example, LPJmL has a vegetation model along
with the hydrological model. The LaD, ISBA-TRIP
and VIC models include hydrology, but also focus on
energy balance, and the H08 model focuses on crop
growth and energy balance. WaterGAP, WBMplus
(irrigation only) and GWAVA also consider water
use. Although the use of any PET equation in a
hydrological model may be seen as an energy bal-
ance, in this study, unless the model has an energy
balance built into its framework, it is not considered.
558 Aditya Sood and Vladimir Smakhtin
None of the models reviewed incorporates land-use
change models, i.e. that simulate changes in land use
due to development (for a review of land-use change
models, see Agarwal et al. 2002). Also, the GHMs
do not consider the impact of market forces or eco-
nomics (in terms of human interventions) on
hydrology.
ISSUES AND TRENDS IN GHM
DEVELOPMENT
Attempts are being made to coordinate the develop-
ment of GHMs. The Global Water System Project
(GWSP), along with the Integrated Project Water and
Global Change (WATCH) (http://www.eu-watch.org/
templates/dispatcher.asp?page_id=25222705), invited
teams developing GHMs to participate in a model
intercomparison project (http://www.eu-watch.org/nl/
25222736-Global_Modelling.html). Seven different
GHMs were forced with the same set of climatic
inputs, and their results were compared at both a
continental scale and the scale of major river basins.
The study concluded that there was limited agree-
ment between models on global water projections.
For example, the range of predicted global runoff
from different models was about 45% of the mean
simulated runoff (Haddeland 2011). Similarly,
Gudmundsson et al. (2012) applied an ensemble of
nine large-scale hydrological models in Europe and
found a large spread in model accuracy including
high variation in the low runoff percentile.
Another 4-year initiative, the Inter-Sectoral
Impact Model Intercomparison Project (ISI-MIP;
http://www.pik-potsdam.de/research/climate-impacts-
and-vulnerabilities/research/rd2-cross-cutting-activities/
isi-mip) is currently underway. Several GHMs, includ-
ing VIC, H08, WaterGAP, MacPDM, WBM, MPI-HM
and PCR-GLOBWB, along with other global impact
models, are included in this project, where new climatic
and socio-economic scenarios developed as part of
the Fifth Assessment Report of the IPCC
(Intergovernmental Panel on Climate Change) are used
as forcing, and their possible impact on different sectors
(such as agriculture, water, ecosystems, infrastructure
and health) are being evaluated.
Focusing on uncertainties
In the WATCH study, although, at the global level,
the deviation between the models of the simulated
total annual flow was not very large—a coefficient of
variation (CV) of about 0.09 (Voß et al. 2008), there
were larger deviations in colder (polar, without
Antarctica: CV ≈ 1.0) and drier regions (Africa and
Australia: CV ≈ 0.41). Also, despite the fact that
there was a closer match in the total flow, there was
a very wide variation in the soil moisture (CV ≈ 0.33)
and evaporation products (CV ≈ 0.27) suggesting the
importance of looking into structural differences in
the models. While some models simulated an
increase in total global freshwater flow for the period
2071–2100, others showed a reduction (Voß
et al. 2008), the range being from −40 to +30%.
The study showed high uncertainty in the intermedi-
ate processes. Wada et al. (2013) used seven GHMs
to assess the impact of climate change on future
irrigation water demand. They concluded that high
uncertainty, until at least mid-century, is due to
GHMs rather than to GCMs.
Uncertainties in GHMs have still not been suffi-
ciently analysed or emphasized. Most of the time the
validity of model outputs of GHMs is asserted by
comparing the results with the output of other GHMs.
Sometimes the assessment is made based on anecdo-
tal terms, for example, “... on the whole the selected
parameters produce a reasonably good representa-
tion of monthly hydrological regimes across Europe.”
(Arnell 1999); or “… we see that WBM runoff is
‘well behaved’ in relation to the precipitation data
used ...” (Fekete et al. 2002). There are four types of
uncertainties in hydrological modelling: those asso-
ciated with model inputs, model structure, parameter
values and observed data. Palmer et al. (2008) ana-
lysed the uncertainties associated with WaterGAP.
Based on this and other analyses (such as
Kaspar 2004), it is concluded that the uncertainties
due to inputs are higher than the parameter uncertain-
ties. Using ‘goodness’ of the model calibration as an
indicator, they found total uncertainty in streamflow
to be medium in 43% of large river basins, lower in
32% and higher in 25%. Widén-Nilsson (2007) com-
pared WASMOD-M and five other global models and
found a high volume error (i.e. error in the river
runoff) in all the models, although the regions of
high error were not consistent. The volume error
mostly ranged from −65 to 50%. Biemans et al.
(2009) found a multiplier effect of precipitation
uncertainty in predicting streamflow at the global
scale. An average of 30% uncertainty in the precipi-
tation data, when used as an input with LPJmL,
resulted in an average 90% uncertainty in streamflow.
One of the reasons for high uncertainty in model
inputs is data scarcity, or inconsistency in the input
data (Arnell 1999, Döll and Siebert 2002, McMillan
 Global hydrological models: a review
et al. 2012, Kauffeldt et al. 2013, Mulligan 2013,
Müller Schmied et al. 2014). Data scarcity is greater
in developing regions (Schuol and Abbaspour 2006),
and is due to well-known problems of weak observa-
tional infrastructure, gaps due to missing data, and
unwillingness to share data. Also, low quality ‘disin-
formative data’ (Kauffeldt et al. 2013) often provide
a false sense of data richness, effectively being
responsible for a large part of model uncertainty.
Integrating with remote sensing
Continuously progressing remote sensing (RS) tech-
nologies can help reduce uncertainties associated
with inputs and observations. The future of GHMs
is to a large degree associated with the development
of RS technologies and public availability of RS data.
Most of the GHMs are already using some satellite
products, such as solar radiation, digital elevation
models (DEMs), land use, etc., as their input data.
Although (at least at present) ground-based measure-
ments are assumed or perceived to be more accurate
than satellite-based measurements, it is obviously
impossible to cover the entire world with the former.
All GHMs (other than WaterGAP) are tuned by first
converting flow from the grids into streamflow and
then validating against the observed data from, gen-
erally, large gauged rivers. For example, Macro-PDM
uses 31 large river basins for validation, MPI-HM
uses 35, and PCR-GLOBWB uses annual streamflow
time series from 99 large river basins. VIC, LaD,
WaterGAP (which is also calibrated), GWAVA,
WASMOD-M, H08 and ISBA-TRIP use 26, 82,
724, 96, 663, 37 and 33 river basins, respectively.
However, the number of basins used for validation/
calibration for the same model may vary based on the
study, e.g. Döll et al. (2003) use 724 calibration
stations for WaterGAP, whereas Hunger and
Döll (2008) use 1235 stations for the same model.
Clearly, such limited tuning/calibration introduces
uncertainty in model outputs. There are also inac-
curacies inherent in the land-based streamflow mea-
surements. Also, as most of the large rivers have been
highly modified (Döll et al. 2009), it is imperative for
future GHM modelling activities to include river
alterations and water withdrawals (van Beek
et al. 2011). In addition, there has been a significant
decrease in the number of river gauging stations
worldwide since the early 1990s, and the existing
gauging network monitors streamflow from only
approximately 50% of the land mass (Fekete and
Vörösmarty 2007).
559
The future of hydrological modelling also rests
in using satellite data (directly or derived) for calibra-
tion. Significant progress has been made in satellite-
based data acquisition in the last decade. With the
launch of the European Space Agency’s (ESA) Soil
Moisture and Ocean Salinity (SMOS) satellite in
November 2009, the first global soil moisture maps
have become available. The satellite uses large
microwaves (L-band) to measure ‘brightness tem-
perature’ through which surface soil moisture is cal-
culated every three days. The data are being validated
by a network of ground-based measurements
(International Soil Moisture Network), coordinated
by the Global Energy and Water Cycle Experiment
(GEWEX) in cooperation with the Group on Earth
Observations (GEO) and the Committee on Earth
Observation Satellites (CEOS) and ESA (http://
www.esa.int/esaLP/LPsmos.html). There is a lot of
promise in composite systems comprising active
and passive microwave technologies (Das
et al. 2011). The US National Aeronautics and
Space Administration (NASA) launched a combined
radiometer and synthetic aperture radar operating at
L-band (1.20–1.41 GHz) in January 2015 (SMAP;
http://smap.jpl.nasa.gov/mission/). Although there are
still limitations in terms of measuring soil moisture
beyond a certain depth from the surface, RS soil
moisture can still be used as a boundary condition
for equations of soil moisture dynamics within a
model. Satellite-derived evapotranspiration products
are also becoming easily available (such as the
MODerate-resolution Imaging Spectro-radiometer
(MODIS) product, MOD 16 – (http://modis.gsfc.
nasa.gov/data/dataprod/dataproducts.php?
MOD_NUMBER=16). An algorithm based on a mod-
ified Penman-Monteith equation was used to develop
MOD16 (Mu et al. 2007). There are known attempts
to use satellite-based data to derive ET and calibrate a
model against it (although not a GHM) for a 46 000-
km2catchment in southern India (Immerzeel and
Droogers 2008). The authors used the Surface
Energy Balance Algorithm for Land (SEBAL) to cal-
culate ET from satellite data. Such satellite-derived ET
estimates are in themselves quite complex and para-
meter-intensive, and may hardly be regarded as obser-
vations. However, such attempts point to possible
alternatives for calibration of GHMs in the future,
when and if satellite-based ET estimates become
more accurate and direct. Similarly, satellite-based leaf
area index (LAI) products are now available. They
include MOD15 from MODIS, ECOCLIMAP (http://
www.cnrm.meteo.fr/gmme/PROJETS/ECOCLIMAP/
560 Aditya Sood and Vladimir Smakhtin
page_ecoclimap.htm), GLOBCARBON (http://www.
fao.org/gtos/tcopjs4.html) and CYCLOPES (http://
postel.obs-mip.fr/?CYCLOPES-Project,82;
Garrigues et al. 2008).
Since the launch of the GRACE satellite,
research has focused on measuring terrestrial storage
(Strassberg et al. 2009, Syed et al. 2009, Alkama
et al. 2010, Tang et al. 2010, Döll et al. 2014) and
to incorporate it into hydrology (Werth et al. 2009).
GRACE, however, has a very coarse spatial and
temporal resolution combined with the uncertainties
in partitioning the GRACE signal. GRACE observes
the combined mass changes in the atmosphere, sur-
face waters (rivers, lakes, reservoirs, floodplains and
wetlands), soil and the groundwater. Hence, the only
way to measure any of the above independently
requires some estimates of all the other components.
Also, GRACE only provides infrequent (approx. 10
days) snapshots of changes in continental water sto-
rage, where the actual storage conditions have to be
known at the time of the satellite’s pass over. Its
spatial resolution is approx. 200 km (i.e. 2 × 2° at
the Equator). Overall, at present, the uncertainties
associated with the GRACE data are on a par with,
or even higher than, the uncertainties in GHMs for
small to medium size river basins, which GRACE
measurement can aim to reduce (Alkama et al. 2010).
However, it may be possible that further development
of GRACE or related technologies would reduce
such uncertainties. The handicap of coarser resolution
is being resolved, to some extent, with the Gravity
field and steady-state Ocean Circulation Explorer
(GOCE) mission (2009–2013), which provides data
related to Earth’s gravity at 100-km resolution (http://
www.esa.int/SPECIALS/GOCE/index.html).
Improving precipitation inputs
The performance of any hydrological model depends
on the quality and scale of available input data,
especially precipitation. Traditional (raingauge) pre-
cipitation data remain limited worldwide. Radar tech-
nology, although an improvement over raingauge
data, still has limitations due to obstructions and
range. Satellites can provide uninterrupted, global
meteorological data collection and hence help, in
principle, to improve the primary input to hydrologi-
cal models. Both the geostationary satellites, obser-
ving visible (VIS) and infrared (IR) radiation, and the
low Earth Orbiting (LEO) satellites using microwave
(MW) technology (along with VIS and IR), provide
precipitation information. They have their distinct
strengths and weaknesses (Kidd et al. 2009). While
VIS and IR data are available at higher resolution, the
precipitation derived from it is dependent on the
relationship between cloud top temperature and rain-
fall, i.e. ‘cold-cloud duration’ methods (Kidd
et al. 2009), but the precipitation estimates are indir-
ect measurements and prone to errors. Passive MW
technology addresses these issues, but their existing
spatial and temporal resolution is still a hindrance to
effective use in hydrological modelling. Precipitation
data derived from passive MW techniques still need
to be calibrated. Various efforts are being made to
combine the strengths of the LEO and geostationary
satellites to develop more reliable precipitation data-
sets (Kuligowski 2002). The use of satellite-based
precipitation data in hydrological modelling is overall
still in its early stages and has not yet led to signifi-
cant improvements in the outputs of GHMs as com-
pared to radar- or ground-based precipitation data
(Tobin and Bennett 2009). Meteorological reanalysis
data can help fill the gap for missing long-term
historical rainfall data. Li et al. (2013) compared
hydrological output in southern African river basins
by forcing a regional hydrological model with rainfall
derived from two different sources—satellite-based
Tropical Rainfall Measuring Mission (TRMM) and
reanalysis of ERA-40. According to their analysis,
streamflow simulations were better when the model
was forced with rainfall (after bias correction) from
reanalysis compared to satellite-based rainfall.
Improving spatial resolution
Wood et al. (2011) argue that the uncertainties asso-
ciated with existing GHMs are too high and their
current usual spatial resolution at the scale of 50-km
grids (considering 0.5° grids) is too coarse to simu-
late “water cycle science questions” effectively
(Wood et al. 2011). The authors make a case for
developing future GHMs at “hyper-resolution”
scale, i.e. at the scale of 1 km globally and 100 m
at the continental level. They believe that the compu-
tational constraints that prevented such studies earlier
no longer exist due to easy access to parallel-clus-
tered computing resources. Gosling et al. (2010) used
Campus Grid to run multiple GCMs with climate
change data. They were able to reduce the computa-
tion time from 750 h to 9 h on a single-processer
personal computer. Also, high resolution RS data are
becoming common. Such high-resolution models
would capture heterogeneity and would better repre-
sent surface–subsurface and land–atmosphere
 Global hydrological models: a review
interactions. Although the idea of increasing the reso-
lution of GHMs is feasible with increasing computer
power, it is important to also consider that many local
issues may be better addressed by using basin-speci-
fic models rather than GHMs. Increasing complexity
and resolution of GHMs may require more spatially
diverse observational data at the ground level (which
is already a big problem). One question that can be
raised is that of the scope of application of GHMs:
with increasing spatial resolution, is it the ultimate
intention to address local water resources problems
by means of GHMs?
Modelling at a finer spatial resolution opens yet
another possibility for running GHMs at the small
basin scale (instead of grids). Some countries have a
very detailed system of basin delineation. For exam-
ple, South Africa evaluates and monitors its water
resources on a so-called ‘quaternary’ sub-basin scale.
Quaternary sub-basins are incremental drainage
subdivisions, ‘flowing’ one into another and covering
the entire country. There are almost 2000 of these sub-
basins with an area ranging from some 60 km2 (in
more humid regions of the country) to some 2000 km2
(in arid areas). The recent trend is to address water
management problems at the scale of even finer
subdivisions, the ‘quineries’. Countries like India,
have not yet established such detailed drainage
subdivisions and for many large-scale assessments
only 19 major drainage areas are used. These drainage
areas range from 22 000 km2 (Sabarmati River basin)
to 860 000 km2 (Ganga River basin) (Amarasinghe
et al. 2005). In the United States, watershed delinea-
tion is carried out in detail, with the entire country
delineated to six levels. This is publicly available as
the Watershed Boundary Dataset (http://www.ncgc.nrcs.
usda.gov/products/datasets/watershed/index.html). The
six levels are classified as regions (level 0), sub-regions
(level 1), basins (level 2), sub-basins (level 3), water-
sheds (level 4) and sub-watersheds (level 5). The sixth-
level drainage delineation has a drainage area of up to
160 km2. Such fine-resolution river basin delineations
are still restricted to a few countries, but the recent
trend is to address water management problems at the
scale of even finer subdivisions. The advantage of this
could be closer integration of such models with the
country-specific national drainage divisions.
CONCLUSIONS
The last two decades have seen substantial effort in
the development of GHMs. The models are based on
either energy balance or hydrological balance, or
561
both. The complexities of the models vary based on
their objectives and applications. Structural issues,
such as the number of soil layers, number of land-
use classifications, crop growth models, irrigation
water applications and reservoir storage, are dealt
with differently in the various models. A comparative
study of seven models has shown structural differ-
ences in the models and little agreement in the pro-
jections made by them. The structural differences
also relate to inclusion of anthropogenic distur-
bances, inclusion of vegetation growth models and
the type of bucket model used (leaky bucket versus
non-leaky bucket).
Most of the models use satellite-based or
Climatic Research Unit (CRU) weather input data,
available in a 0.5-degree gridded format. This also
defines the spatial resolution of the models. Most of
the GHMs run on a daily temporal scale. However,
for the long-term assessment of the impacts of cli-
mate change and impact on global water and food
trade, a monthly temporal scale is sufficient and may
simplify the calibration process.
The models also need to move to finer spatial
resolution to address some of the global societal
issues, but they should not aim to replace more
locally focused modelling efforts. Progress in
GHMs is also dependent on improved access to
observed hydrometeorological data, particularly in
developing countries. Access to more observed
data could expand the ‘calibrated area’ of the
globe, thus also improving the overall accuracy
of global hydrological modelling efforts and
providing more accurate, although simulated,
time series of various components of the hydro-
logical cycle. This may be seen as a possible
interim (or even long-term) solution for ground-
based ‘data-poor’ areas, where ground observa-
tions will probably never materialize to required
levels, while RS data still require decades to
ensure that time series are of a reasonable length.
Hence, simulations will always be needed in such
areas. Therefore, improving the accuracy of
GHMs by all possible means, may be effectively
seen as a form of international cooperation and
economic/humanitarian assistance. However,
countries have to be prepared to cooperate by
allowing better access to national archives of
observed hydro-meteorological data.
It is suggested that, to improve the GHMs, there
should be a closer integration of satellite-based pro-
ducts and models that would help reduce the internal
uncertainties within the models.
562 Aditya Sood and Vladimir Smakhtin
Acknowledgements The authors gratefully acknowl-
edge comments provided by Robyn Johnston (IWMI,
Colombo) in reviewing the manuscript. Comments
provided by two anonymous reviewers are also
acknowledged.
Disclosure statement No potential conflict of inter-
est was reported by the author(s).
Funding This study was funded by the CGIAR
Research Program on Water, Land and Ecosystems
(WLE).
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