Advances in Water Resources 51 (2013) 317325

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Advances in Water Resources

journal homepage: www.elsevier.com/locate/advwatres

Hydrology as a driver of biodiversity: Controls on carrying capacity, niche

formation, and dispersal
Megan Konar a,, M. Jason Todd a, Rachata Muneepeerakul b, Andrea Rinaldo c,d, Ignacio Rodriguez-Iturbe a
a

Princeton University, Department of Civil and Environmental Engineering, Princeton, NJ, USA
Arizona State University, School of Sustainability and Mathematical, Computational, and Modeling Science Center, Tempe, AZ, USA
c
cole Polytechnique Fdrale de Lausanne, Laboratorie of cohydrology, Faculte ENAC, CH-1015, Lausanne, Switzerland
d
Universita di Padov, Department IMAGE and International Centre for Hydrology, Dino Tonini, Padua, Italy
b

a r t i c l e

i n f o

Article history:
Available online 3 March 2012
Keywords:
Hydrology
Biodiversity
Dispersal
Carrying capacity
Niches
Climate change

a b s t r a c t
A synthesis is presented highlighting the importance of hydrologic variables and dynamics to biodiversity
patterns. The focus of this paper is the key hydrologic controls crucial towards quantifying the impacts of
climate changes on the distribution of species. Specically, we highlight the hydrologic controls operating
on the carrying capacity, niche formation, and dispersal dynamics. This synthesis will facilitate avenues of
future research and is connected to issues of major practical importance, such as the integration of the
structure of river networks into conservation strategies and the evaluations of the impacts of climate
change on biodiversity.
2012 Elsevier Ltd. All rights reserved.

1. Introduction
Maintenance of biodiversity across multiple scales has been of
basic interest to the biological and ecological sciences for decades
[43,41,31]. The debate over the ultimate controls on biodiversity is
a contentious one and indeed the numerous papers investigating
them rarely reach solid conclusions. However, it is without debate
that patterns exist across taxa, geographical areas, and geological
eras [67]. With the ever growing body of literature detailing the
benets provided by biodiversity [12,14,21,30], increasing attention is being paid towards its fundamental drivers and how
changes to key components could affect specic patterns of biodiversity. Additionally, increased global species loss [58] has made it
critical to better understand the processes that govern biodiversity.
The principal threats to biodiversity vary widely depending on
geographic location and the complicating effects of differences in
spatial and temporal scale. At present, the principal threats to biodiversity are the effects due to land use change and associated habitat loss and fragmentation, as they act on a much shorter time
scale than other processes [28,19,55,56]. Several global modeling
scenarios show that land use changes will continue to be the principal reason for terrestrial biodiversity loss until at least 2050
[34,56,71,72]. However, climate change is likely to be the major
reason for biodiversity loss worldwide after 2050 [48,75,76].

Corresponding author.
E-mail address: mkonar@princeton.edu (M. Konar).
0309-1708/$ - see front matter 2012 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.advwatres.2012.02.009

While the predicted ultimate percentage loss of species due to

climate change varies widely from study to study (e.g. [44,77]),
the IPCC reports that 2030% of animal and plant species are likely
to be at high risk of extinction with a global mean annual temperature rise of 23 C [75]. Indeed, research has shown that despite
the numerous possible explanations for changes in biological patterns and communities, climate change effects are already inuencing biodiversity through range shifts and alteration of phenology
[19,56]. This loss of biodiversity has the ability to produce a multitude of consequences, such as the loss of ecosystem functioning
and reduction or elimination of goods and services [56]. Many of
these responses may be nonlinear and difcult to predict [5], leading to rapid transitions or sudden shifts in ecosystem states
[56,20,74].
One of the major pathways through which climate change will
impact biodiversity patterns is through altered hydrologic patterns
and processes [83]. It is well known that climate change will impact global precipitation patterns [53,75], resulting in increased
variability in rainfall regimes in both time and space [54], which,
in turn, change the hydrologic conditions that regulate ecological
processes [13,67,59,46]. This is one of the reasons why it is important to focus on the specic mechanisms through which hydrology
impacts biodiversity. In this paper, we present a synthesis of some
of the most important hydrologic controls on biodiversity, with an
eye towards understanding the potential impacts of climate
change. Note that we make no attempt to summarize the vast literature that relates to the many and varied interactions between
hydrology and species diversity. Instead, we focus on hydrologic

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controls that can be used to quantify the impacts of climate

changes on biodiversity.
While the role of hydrology on biodiversity within freshwater
ecosystems may seem self evident, hydrologic controls also play
a vital role in structuring and maintaining terrestrial ecosystems
[16,65,73]. Hydrology has been shown to play a vital role in structuring terrestrial vegetation, particularly in water-limited ecosystems. In water-limited ecosystems, soil moisture controls the
availability of nutrients and limits plant transpiration [64,65]. In
humid ecosystems, the interactions between water and energy cycles increases in importance, and the diversity of trees is inuenced
by evapotranspiration [13]. Vegetation communities in riverine
systems are often structured by hydrogeomorphological interactions [2]. Hydrologic disturbances, including droughts and oods,
play an important role in the maintenance of both aquatic and
oodplain ecosystems [38]. Likewise, droughts and oods are
important determinants of vegetation diversity in terrestrial systems [78].
Hydrology is not the only factor impacting biodiversity patterns
and processes. For example, it is well known that the species richness of almost all life forms increases from high to low latitudes
and along elevation gradients. This latitudinal diversity gradient
is thought to be generated by several mechanisms, such as the
availability of energy, historical perturbations, and interactions between species, but may simply be a consequence of more land area
in the tropics [66,67]. The biogeography of plants in mid- to high
latitudes may best be explained by the spacetime patterns of
the shortwave radiative ux [17]. Determining the ultimate controls on biodiversity patterns is complicated by the fact that the
specic underlying mechanisms and their importance may differ
across both spatial and temporal scales. Hydrologic processes act
with considerable variability across multiple spatial and temporal
scales, which is one of the reasons why biodiversity is likely inuenced by some aspect of hydrology at most scales of analysis
[13,63,59].
In this paper we focus on three specic determinants of diversity
for which hydrology may play an important role and which can be
used to quantify potential climate change impacts. In Section 2,
we describe the hydrologic control of the spatial pattern of carrying
capacity in some ecosystems. Section 3 discusses the hydrologic
control of niches favoring or restricting the existence of different
species. Section 4 describes the hydrologic control of dispersal
mechanisms, with a focus on lotic populations in river networks.
We look to the future in Section 5.

2. Hydrologic control of carrying capacity

Carrying capacity is dened as the maximum number of individuals or units of organisms that can be maintained in a given
area on a long-term basis. Some habitats are far more productive
than others and, in general, more productive areas support more
individuals and more species [13]. However, this pattern is often
complicated and may follow a non-monotonic relationship in
many systems [67]. For this reason, we draw examples from natural systems for which an increase in the carrying capacity has been
shown to lead to more species. The focus of this section is the
hydrologic control of the spatial distribution of carrying capacity.
The relationship between hydrology and carrying capacity has
been established for quite some time [81,25,62]. Both terrestrial
and aquatic ecosystems with more freshwater resources tend to
support more individuals. Here, we focus on the controlling inuence of hydrology in both aquatic and terrestrial ecosystems:
namely, the inuence of precipitation on the carrying capacity of
trees and the impact of ow characteristics on the carrying capacity of sh.

Recently, mean annual precipitation was found to be the major

determinant of potential woody cover in African savannas [73].
Sankaran et al. [73] demonstrate that maximum fractional woody
cover, which proxies for the carrying capacity of trees, is primarily
controlled by moisture limitation. They conduct a continentalscale analysis of Africa in an effort to determine whether savannas
are primarily climatically determined or disturbance driven, nding that savannas are predominantly water limited in locations
with less than approximately 650 mm yr 1, while those locations
that receive greater than 650 mm yr 1 are disturbance driven.
Thus, mean annual precipitation controls the upper bound on woody cover, although disturbance regimes and soil characteristics do
impose signicant controls on woody cover below the bound [73].
Recent research builds upon the work of Sankaran et al. [73]
and utilizes mean annual precipitation as a driver of carrying
capacity to model distributions of tree species diversity. Konar
et al. [39] demonstrate that a neutral meta-community model,
coupled with an appropriate representation of tree carrying capacity, effectively reproduces empirical patterns of tree diversity. The
model was not able to reproduce empirical tree diversity patterns
without a spatial representation of tree carrying capacity based on
rainfall [39,11]. This analysis was conducted for the Mississippi
Watershed (shown in grey in Fig. 1A), showing that mean annual
precipitation appropriately characterizes the carrying capacity of
trees in humid ecosystems. Note that forest cover was used as a
proxy of tree carrying capacity in Konar et al. [39], rather than
woody cover as in Sankaran et al. [73], which accounts for differences in functional form. Additionally, Sankaran et al. [73] focus
on savannas, which, by denition are regions with tree-grass coexistence, i.e. tree cover never reaches 100% in savannas. Two
hydrological variables were considered for use as a driver of forest
cover in Konar et al. [39]: evapotranspiration and mean annual
precipitation. The relationship between forest cover and mean annual precipitation exhibited a more well-dened relationship than
that between forest cover and evapotranspiration. Additionally,
projections of mean annual precipitation under climate change
scenarios are readily available, making this variable desirable for
projection purposes.
Importantly, the modeling approach used in Konar et al. [39] has
predictive powers, since it allows for the direct linkage of largescale biodiversity patterns to environmental forcings. Projections
of mean annual precipitation under different climate scenarios
were used to obtain new values of tree carrying capacity for the
Mississippi Watershed. With these resulting new carrying capacities, Konar et al. [39] determine how various climate change scenarios are projected to affect tree diversity patterns in the Mississippi
Watershed. 15 climate change scenarios are implemented in the
model. Here, the spatially-explicit impacts under the most dramatic
species-poor scenario are shown in Fig. 1B. Note that the probability of any particular outcome in large-scale macrobiodiversity patterns is heavily reliant on the probabilities associated with the
projected precipitation patterns provided by the global climate
models. For this reason, the patterns should be interpreted as envelopes of plausible biodiversity scenarios, rather than as predictions
of biodiversity outcomes. Tree diversity patterns are impacted more
under the species-poor scenarios than under the species-rich scenarios, with the exceptions of a few select regions, where impacts
are of comparable magnitudes under both scenarios. Additionally,
rare species are disproportionately impacted under climate change
[39], a nding shared with niche-based models [50].
Recent research indicates that the timing and intensity of rainfall may be a more important driver of carrying capacity than sheer
quantities of rainfall in some systems [70,22,1]. In a continentscale analysis of Africa, Good and Caylor [22] build upon the work
of Sankaran et al. [73] and demonstrate that the quantity and
intensity of rainfall events inuences the upper limit of woody

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319

Fig. 1. (A) Map of the Mississippi Watershed within the continental United States. The Mississippi Watershed is highlighted in grey and the network structure in blue. (B)
Impact of climate change on region-averaged local species richness (LSR) in sub-regions of the Mississippi Watershed. Shades of green indicate the percentage change in the
region-averaged local species richness under climate change, with dark green indicating a higher percentage lost. The general trend is that a higher percentage of species are
lost in the west with a decreasing trend to the east. The change per local community in region-averaged local species richness under climate change is indicated for each
region by the bold numbers. The species-rich regions east of the 100oW meridian lose more species, though these species represent a smaller percentage of species in these
regions. The mean local species richness in the South is anticipated to decrease by 6.3 species under climate change, the largest loss of all sub-regions. Taken and modied
from Konar et al. [39] and Bertuzzo et al. [4]. (For interpretation of the references to colour in this gure legend, the reader is referred to the web version of this article.)

vegetation cover. They show that areas with similar seasonal rainfall totals have higher fractional woody cover if the local rainfall
climatology consists of frequent, less intense precipitation events.
This distinction is crucially important since climate change is projected to lead to more intense, less frequent storm events, with
important repercussions for vegetation. Similarly, Bartholomeus
et al. [1] demonstrate that increased extremes in soil moisture,
similar to those likely to be observed under climate change, will
impact plant functional groups, though this study does not address
the impact of hydrologic variation on species diversity directly. To
our knowledge, there has not yet been a study that quanties the
impact of hydrologic variation on species diversity, so this represents an important area for future research. Thus, although hydrologic averages may drive the carrying capacity and diversity in
some systems, spatial and temporal variability may prove more
important in other ecosystems.
The carrying capacity of organisms in river systems is strongly
impacted by the gradient of a number of physical factors connected
to the drainage network [81]. In fact, the ow of water, which is
strongly correlated with other physical variables, can be considered a master variable that limits the distribution and abundance
of organisms in rivers [63,59]. Recent research on sh biomass
demonstrates the importance of ow characteristics. Halls et al.
[27] developed a ood index that accounts for the extent and duration of ooding during the entire ood period of the Tonle Sap
River, Cambodia. Fish biomass varied by almost a factor of 5 in response to the ood index over the last decade [27]. The Mekong
River is subject to rapid development, with the construction of several large-scale hydropower dams and reservoirs [40]. The related
ow alterations will have signicant impacts on productivity and
numbers of sh in Tonle Sap ecosystem, with signicant repercussions for sh diversity.
Taking advantage of the relationship between the number of
sh and river ow, Xenopoulos et al. [84] analyze the projected impacts of climate change and future water withdrawals on sh
diversity at a global scale. To address this issue, they present the
rst coupling of a global hydrological model with a freshwater biodiversity model. A species-discharge regression model was used to
produce scenarios of sh species loss. Species richness generally
increased with increased mean river discharge, in a similar manner
to the species-area curve of terrestrial ecosystems [84].
Fig. 2 illustrates the 52 watersheds that Xenopoulos et al. [84]
show will lose more than 10% of their sh species. The combination
of increased evapotranspiration and decreased precipitation as a result of climate change (specically, under the IPCC SRES A2 scenario

[75]) is the most important driver of freshwater sh loss. However,

the impact of water withdrawal for human consumption on sh
diversity is particularly important in certain watersheds. For example, consumptive water use in the Euphrates and Cauvery Rivers
was the major driver of sh species losses in these watersheds.
Xenopoulos et al. [84] present an excellent rst approximation to
the projected impacts of climate change and human modication
on sh diversity. However, as with all species-distribution models,
there are inherent problems in making projections based on correlations with current environmental regimes (see e.g. Pearson and
Dawson [57]). For this reason, a model based upon ecological processes would be a most benecial addition to the literature.
In this section, we have shown that hydrologic variables are
important to the carrying capacity of trees and sh. Recent research
demonstrates the inuence of hydrology on carrying capacity
[73,27,22], as well as how to utilize this relationship to quantify
the impact of climate change on biodiversity patterns [84,39,1].

3. Hydrologic control of niches

Niches refer to segments of ecosystems that have been partitioned in both space and time and that are available for utilization
by different organisms. The number of niches that a given ecosystem contains has long been thought to be a major driver of species
diversity [32,67]. The hydrologic control of niches occurs across
both terrestrial and aquatic environments, but may be best displayed within wetlands, as they have been a principal location
for studying the relationship between hydrology and the structuring of vegetation communities [49]. Due to their dynamic hydrology, climate change impacts in these areas could be particularly
severe, as slight changes in water availability may have profound
inuences on the surrounding biodiversity. The focus of this section is the inuence of hydrology on niche structure and the role
climate change may have in altering these relationships.
The hydrology of a wetland environment can be described by
multiple factors, but is most often described by its hydropattern
or hydroperiod, a combination of the periodicity of inundation
events for a given location along with the length and depth of inundation. One such environment where these relationships have been
extensively studied is the Everglades, with multiple studies showing a relationship between the hydropattern and vegetation communities of a given area [15,26,87]. Nevertheless, the role of
hydrology in structuring vegetation communities in wetlands is difcult to determine due to a suite of interacting variables [6,69,86].

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Fig. 2. Impact of change in water discharge from climate change and water consumption on sh diversity at a global scale. Colors display the percentage change in sh
diversity, where red indicates approximately 50% loss of diversity and green indicates roughly 50% gain in diversity. The 52 watersheds that are projected to lose more than
10% of their sh species are numbered. Climate change is projected to decrease sh diversity more than water withdrawals in most watersheds. However, in a few
watersheds, such as the Euphrates and Cauvery, sh species losses due to water withdrawals will be more severe. Taken and modied from Xenopoulos et al. [84]. (For
interpretation of the references to colour in this gure legend, the reader is referred to the web version of this article.)

Additionally, the lack of long term temporal and wide spatial coverage data presents a further complication to understanding the role
of hydrology in structuring vegetation communities.
Todd et al. [79] examine the hydrologic control of niches across
a long temporal and wide spatial scale of the entire Everglades National Park (ENP). This analysis quanties the hydrological niches
of the dominant vegetation communities. Mean hydroperiod depth
and percent time inundated best describe the vegetation niches.
For instance, muhly grass occurs most often at shallower depths
and at locations that are not inundated for long period of time. In
contrast, bay-hardwood scrub favors more hydric locations. However, results for sawgrass, by far the most common vegetation
community across the ENP, supported earlier studies that hydrology is not the only variable structuring sawgrass niches [26,29].
Thus, the work of Todd et al. [79] showed that, while multiple
factors can inuence the landscape distribution of vegetation communities, hydrology plays a principal, if not dominant role, in many
cases. Most importantly, this work indicates that hydrology structures vegetative niches at large spatial and temporal scales, reinforcing the results of many studies conducted previously at
smaller scales.
In an effort to gauge the impacts of projected changes in precipitation on plant diversity, Todd et al. [80] used the relationships between vegetation communities and hydrologic variables developed
in Todd et al. [79]. Specically, Todd et al. [80] utilized projected
precipitation values for this region under various climate change
scenarios to model future hydroperiod characteristics. Projected
values of mean depth and percent time inundated are then used
to determine impacts to vegetation communities, based upon the
relationships described by Todd et al. [79]. Under increasing emissions scenarios, precipitation decreased across the ENP by as much
as 8% from present scenarios, leading to an associated decrease in
mean depth and percent time inundated (refer to Fig. 3). These

changes in hydroperiod were then used to project changes to

vegetation communities. Thus, vegetation communities that favor
xeric conditions increased in abundance and communities that
favor hydric conditions became scarcer (refer to Table 1). For instance, the relative abundance of muhly grass, which favors xeric
conditions, increased by 15% under the high emissions scenario.
In contrast, the relative abundance of bay-hardwood scrub, which
favors hydric environments, decreased by 66%. Thus, Todd et al.
[80] utilizes the previously established relationships between hydroperiods and species distributions [79] to project the potential
impacts of climate change on plant diversity at large spatial scales.
Another wetland region that has undergone intense research
regarding the inuence of changing climate on vegetation community structure and biodiversity is the Prairie Pothole Region of
North America. This region has numerous wetlands covering a
suite of inundation regimes from temporary (one to two months)
to semipermanent (inundated throughout most years). Associated
with these varying inundation regimes, the vegetation found
therein shows complex zonation dynamics. Through the years, various models have been developed to simulate vegetation dynamics
across a range of hydrologic parameters including precipitation,
runoff, potential evapotranspiration, snowpack and subsurface inow [60,61,37]. These models generally did a satisfactory job of
reproducing hydrologic and vegetation conditions of these wetlands. When incorporating climate change scenarios of increases
in temperature as compared to historic levels, these wetlands
showed marked hydrologic changes. In general, the wetlands experienced earlier snowpack melting, decreased water depths and volume, diminished hydroperiods, and reduced peaks from snowmelt
and rainfall unless rainfall across the region increases 57% per
degree of warming [37]. However, the inuence of these changes
was not uniform across wetland type with seasonal wetlands being
the most affected. These seasonal wetlands normally persist into

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321

Fig. 3. The joint probability surface of percent time inundated and mean depth for (A) present conditions and under (B) low (B1), (C) middle (A1B), and (D) high (A2) climate
emissions scenarios. The value of each pixel represents the relative frequency of all pixels across the Everglades National Park meeting both hydrologic conditions. These
hydrologic scenarios were used to determine the impact of climate change on plant communities in the Everglades National Park, provided in Table 1. Taken from Todd et al.
[79].

Table 1
Percent coverage of dominant vegetation types within Everglades National Park under
the present and high emissions scenarios. The percent change of dominant vegetation
types between the present and high emissions scenarios are also provided. Only those
vegetation types constituting more than one percent of the total landscape are listed.
Taken from Todd et al. [79].
Vegetation type

Present

High

% Change

Sawgrass
Mixed Gramminoids
Tall Sawgrass
Muhly Grass
Spike Rush
Red Mangrove Scrub
Bayhead
Pine Savanna
Willow Shrublands
Dwarf Cypress
Bay-Hardwood Scrub
Brazilian Pepper
Cattail Marsh
Slash Pine with Hardwoods
Hardwood Scrub
Subtropical Hardwood Forest

60.68
6.55
5.80
4.07
2.98
2.16
1.72
1.59
1.47
1.45
1.44
1.22
1.09
0.88
0.71
0.75

55.21
8.82
2.24
10.25
1.38
0.92
0.83
5.17
1.36
0.69
0.49
2.50
0.29
2.96
1.57
1.43

9.0
34.7
61.4
152.0
53.5
57.4
51.7
224.3
7.9
52.1
66.1
104.4
73.5
237.2
121.9
91.1

the summertime months, but suffered shortened hydroperiods due

to the enhanced evaporative demand of a warmer climate.
The changes in hydroperiod structure in the Prairie Pothole Region had a marked effect on the habitat available for migratory
waterfowl. Johnson et al. [36] utilized the WETSIM model, a process-oriented, deterministic model that simulates wetland surface
processes and vegetation dynamics, to quantify suitable waterfowl
niche space. Three future scenarios were run in WETSIM to quantify the impacts of climate change on wetland hydrology and corresponding waterfowl habitat availability. These scenarios are (1) a
3 C temperature increase with no change in precipitation, (2) a
3 C temperature increase with a 20% increase in precipitation,
and (3) a 3 C temperature increase with a 20% decrease in precipitation. From Fig. 4, it is clear that all three scenarios lead to

marked geographic shifts in the availability of waterfowl habitat.

The scenario with increased temperature and decreased precipitation led to the most dramatic contraction in suitable habitat
(Fig. 4E).
Many of the studies investigating the role of hydrology in structuring vegetative niches implement climate change as a long, gradual process, with a trend towards elevated temperatures and
associated changes in hydrology. However, it is important to note
that in dynamic environments, such as wetlands, changes in the
occurrence of extreme precipitation events may play an important
role [35]. Additionally, increased variability in rainfall patterns,
leading to ooding and droughts, may have profound inuences
on vegetation [85,47]. As an example, Zedler [85] investigated
the inuences of oods and droughts on wetland vegetation over
thirty years, showing that biodiversity richness and evenness was
reduced abruptly due to sequential, catastrophic oods.
In this section, we have described the important role that
hydrology plays in structuring niches in wetland ecosystems for
both vegetation and migratory birds. Recent research quanties
the hydrologic niches of plants [15,26,79,87] and waterfowl [36]
and has recently been used to quantify the potential ecological impacts of climate change [36,80,85].

4. Hydrologic control of dispersal mechanisms

Dispersal of organisms refers to their movement from one habitat to another and has long been recognized as an important driver
of biodiversity patterns. Dispersal tends to increase the diversity of
local communities [43], but systems with high rates of dispersal
tend to have lower global diversity [31]. The dispersal distance
[8] and shape of the dispersal kernel [68] are important determinants of diversity patterns as well. Empirical evidence suggest that
the dispersal kernels of seeds have fat-tails [9], which indicates
that dispersal is likely to occur if there is an available pathway.
Dispersal of organisms through heterogeneous landscapes also

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M. Konar et al. / Advances in Water Resources 51 (2013) 317325

Fig. 4. A. The Prairie Pothole region of central North America (inset map) and six eco-regions. Yellow symbols represent weather stations (three per eco-region) used in
WETSIM analyses. Simulated waterfowl habitat across the Prairie Pothole region under historic (B) and future scenarios of climate change (C, D, and E). Taken from Johnson
et al. [36]. (For interpretation of the references to colour in this gure legend, the reader is referred to the web version of this article.)

impacts the distribution of organsims. This is because organisms

adapt their dispersal strategies based upon the underlying landscape structure [52].
There are a number of hydrologic variables and hydrologic
dependent landscapes which may impact dispersal. Here, we focus
on systems with hydrologically determined carrying capacities and
habitat connectivity, since organisms adapt their dispersal strategies to both factors [52]. River systems present a unique opportunity to explore how dispersal is inuenced by the combination of
the spatial distribution of carrying capacity and the landscape connectivity structure (i.e. riverine topology). This unique landscape
structure, in turn, makes it possible explore how ecological dispersal might be impacted by human modication. The focus of this
section is recent research that quanties how river topology inuences dispersal and diversity and, subsequently, how human modication to river topology may impact diversity patterns.
With increased variability in rainfall regimes under climate
change and increased demands on freshwater, not only will river
ows be altered, but it is likely that human modication of river
systems will intensify, in an effort to stabilize water supplies for
human consumption, agriculture, and industry [82]. For example,
inter-basin water transfer projects, in which man-made canals
transform the river connectivity to provide water to areas where
demand exceeds supply, are anticipated to become more prevalent
[42], as are water withdrawal projects [84]. River topology is an
important element of dispersal, particularly for aquatic organisms.
In fact, sh and other entirely aquatic organisms are constrained by
the landscape imposed dispersal constraints embedded in river
networks [18].
River networks impose constraints on population processes, such
as spread, growth, and survival [23]. Thus, straight line distances in
2-D space may be very different to distances as measured through
the river network. In other words, as the crow ies may be different to as the trout swims [18]. For this reason, one of the principal
consequences of the riverine structure is to alter the landscape
through which organisms disperse. For example, the river network
structure may inuence population extinction risk, slow the spatial
spreading of exotic organisms, enhance diversity at conuence
points of the network [23], and impact the persistence of species [3].
Muneepeerakul et al. [51] present a neutral model of sh diversity in the Mississippi Watershed that is based upon ecological
processes. In this metacommunity model, the sh disperse based
upon the underlying river network topology. The model produced

excellent ts to the empirical data, once the spatial distribution of

carrying capacity was appropriate characterized and the dispersal
process was constrained to follow the river network. The model
effectively reproduced several key diversity patterns; namely, the
prole of local species richness, the local species richness frequency distribution, the rank-occupancy curve, and the b diversity
as characterized by the Jaccards similarity index [51]. Additionally,
Bertuzzo et al. [4] demonstrate that the model is able to reproduce
spatially explicit biodiversity patterns; namely, the environmental
resistance, which quanties the spatial loss of community similarity and is useful in identifying biogeographic regions. Remarkably,
the model produced a good t to this spatial pattern with no further calibration [4].
The process-based model presented in Muneepeerakul et al.
[51] demonstrates that the dispersal of sh in the Mississippi
Watershed is controlled by the combination of the spatial distribution of carrying capacity and river network structure. Interestingly,
the model results indicate that sh do not exhibit biased dispersal
and the majority travel locally, though an important fraction do
travel very long distances [51,4]. The river network structure is
an essential component of the model; sh dispersal is dramatically
impacted by the drainage structure, since sh offspring are constrained to follow the ecological corridors dened by the network
[51]. Thus, river network structure, a key hydrologically inuenced
landscape, is an important controlling feature of the dispersal of
sh.
Inter-basin water transfer projects restructure river connectivity
and have the potential to alter landscape-scale patterns of species
distributions by introducing new dispersal routes. The landscape
constraints imposed by river topology are essential in determining
the dispersal characteristics and diversity of entirely aquatic organisms (i.e. those constrained to solely within-network movement).
However, the unique topology of river networks does not entirely
control the dispersal of aquatic organisms capable of out-ofnetwork movement [23,24]. For example, salamanders utilize both
out-of-network and within-network movement, which acts to
stabilize their populations [24]. Thus, the network structure is not
as important in the dispersal of organisms that are able to travel
out-of-network, such as salamanders or trees [24,39]. Changing
the connectivity structure of river networks with man-made canals
for inter-basin water transfer projects is an extreme example of
out-of-natural-network movement, most likely to impact entirely
aquatic organisms.

M. Konar et al. / Advances in Water Resources 51 (2013) 317325

323

Fig. 5. (A) Map of the Indian Peninsula. Major rivers are shown in gray, and the 11 proposed links considered in the analysis are shown in black. The four major river basins
involved are shaded and labeled. The links are numbered according to enumeration assigned by Indias National Water Development Agency. (B) Impact of the 11 proposed
interbasin water transfers on local species richness (LSR) for the Indian Peninsula river network. The absolute differences (after minus before) in LSR are shown for each link in
the river network. Taken and modied from Lynch et al. [42].

A neutral metacommunity model of sh in river networks, similar to that of Muneepeerakul et al. [51], can be used to determine
the impacts of inter-basin water transfer projects on sh diversity.
With this approach, Lynch et al. [42] quantify the potential ecological impacts of the proposed inter-basin water transfers in India.
Although the neutral model is not able to predict how the proposed
inter-basin transfer project will impact specic species, it is a suitable framework to evaluate the potential landscape-scale consequences of changes to river connectivity [42].
Changes in species richness due to the 11 proposed inter-basin
water transfers are quantied and shown in Fig. 5. Their study
demonstrates the fragility of reaches in close proximity to the proposed canals. Note that local diversity increases throughout India
with increased connectivity, which is anticipated under the null
scenario of habitat fragmentation [7]. However, global diversity

in the system becomes spatially more homogenous with a notable

decrease in b diversity. Additionally, rare species decrease with the
introduction of the proposed canals [42]. In other words, more species are present in each location, but this is because certain species
become widespread and dominant when otherwise distant basins
are linked through human modication. This increase in common
species comes at the expense of endemic and rare species. The
study by Lynch et al. [42] is the rst to examine the impact of
changing network connectivity on basin-wide patterns of biodiversity and species richness.
We can look to the past to observe how human introduction of
exotic organisms into river networks has exacerbated biological
invasions. Invasive species are known to displace native species,
usually decreasing species diversity in the system as a whole
[10]. Mari et al. [45] show that anthropogenic drivers of dispersal

Fig. 6. Dynamics of the zebra mussel invasion of the Mississippi Watershed. (A) Empirical data on zebra mussel spread over time. (B) Model snapshots of zebra mussel spread.
Note that the model (B) reproduces the data (A) very closely when both the connectivity structure of the river network and human interferences in zebra mussel spread are
taken into account. Human-mediated dispersal impacts the spread of zebra mussels through commercial and recreational boating activities. Taken and modied from Mari
et al. [45].

324

M. Konar et al. / Advances in Water Resources 51 (2013) 317325

(i.e. commercial navigation) in combination with the river network

topology provide the best t to the spatio-temporal dynamics of
invasion of zebra mussels. In Fig. 6, model estimates of zebra mussel invasion in space and time provide excellent ts to the data of
zebra mussel invasion. Dispersal in the model was constrained to
follow the river network, but zebra mussels were transported at
an accelerated rate and to regions of the network that may have
been outside their range by ships. The study by Mari et al. [45]
illustrates that human-mediation of dispersal is of great importance, particularly when temporal dynamics are taken into account
in river network context, where natural ecological distances are
constrained to follow the topology.
In this section, we have presented examples of how network connectivity can be used to hind-cast or fore-cast the potential impacts
of re-routing the connectivity structure of river systems. Examples
of human-mediated dispersal in both India and the Mississipi
Watershed indicate changes to river network topology may have
disastrous consequences for species diversity, largely through the
introduction of invasive species. As increased pressures are placed
on global freshwater resources from population growth and climate
change [82], it is likely that human modication of riverine systems
will intensify. It is imperative to understand how water withdrawals and modications to the connectivity structure of river networks
will impact ecological dispersal, in order to properly evaluate the
projected biodiversity impacts. This is particularly relevant since
policy makers and resource managers are more able to directly impact water withdrawals and river engineering projects than global
climate.

5. Looking forward
It is imperative to understand the key relationships between
hydrologic processes and biodiversity as global freshwater resources are subject to increasing pressures from both population
growth and climate change. There is certainly a vast literature that
relates to the many and varied interactions between hydrology and
species diversity that we do not cover in this paper. Instead, we
have outlined some of the major hydrologic drivers of biodiversity:
carrying capacity, niche formation, and dispersal. In this synthesis,
we have chosen to focus on hydrologic controls that can be used to
quantify the impacts of climate changes on biodiversity.
Forecasting the potential impacts of climate changes on biodiversity is of increasing importance. For this reason, the focus of this
paper is research that uses hydrologic processes to quantify
changes in species compositions. Going forward, there is a clear
need to develop a mechanistic model linking rainfall dynamics
and biodiversity. While the mean annual precipitation may be
the major driver of the distribution of species in some cases, it is
clear that rainfall timing and intensity will play a major role as well
[22]. For this reason, the development of a model that integrates
species diversity processes with rainfall dynamics would make a
valuable contribution to the literature. Similarly, recent research
has highlighted the importance of soil moisture variability to plant
functional groups [64,1]. It is important that future research continue to develop the relationship between soil moisture processes
and plant diversity.
As global water resources are increasingly constrained, water
withdrawals, transfer projects and dams will become increasingly
popular. Understanding the ecological impacts of proposed engineering projects is particularly important and challenging. The impact to sh diversity due to withdrawals and changes in river
network connectivity have only recently been examined quantitatively. In the future, these modeling approaches could be improved
by incorporating a more comprehensive set of relationships between hydrologic variables and the carrying capacity of sh.

Of course, the major hydrologic drivers that we have highlighted

do not act in isolation. For example, changes to global precipitation
patterns will impact the spatial distribution of carrying capacities of
certain organisms. This, in turn, may lead those organisms to adapt
their optimal dispersal strategies accordingly. Similarly, increased
variability in rainfall regimes may lead to enhanced niche spaces,
which may, in turn, affect the carrying capacity and diversity of certain species. Thus, accounting for the interplay between the major
hydrological drivers of biodiversity is an important and challenging
task going forward.
Additionally, the interaction between these hydrologic and other
abiotic variables should be taken into account when appropriate
[33]. In particular, research that mechanistically links hydrologic
processes with other factors, such as nutrients and temperature,
will be of increased importance in the future, particularly as the
scientic community strives to understand the multitude of interactions between a changing climate and the diversity of biological
systems.

Acknowledgements
M.K., R.M., and I.R.-I. acknowledge the support of the James S.
McDonnell Foundation through the Studying Complex Systems
Grant (220020138). M.K., M.J.T., and I.R.-I. acknowledge the support of NASA WaterSCAPES (Science of Coupled Aquatic Processes
in Ecosystems from Space; NASA NNX08BA43A). A.R. acknowledges funding from ERC Advanced Grant RINEC 22761 and SFN
Grant 200021/124930/1.

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