Toward An Extreme World: The Hyper-Arid Ecosystemas A Natural Model

Received: 28 March 2023 Accepted: 6 April 2023
DOI: 10.1002/ecs2.4586
INNOVATIVE VIEWPOINT
Climate Ecology
Toward an extreme world: The hyper-arid ecosystem

as a natural model
Elli Groner 1,2 | Avshalom Babad 1 | Naomi Berda Swiderski 1 | Moshe Shachak 3
1
Ramon Branch, Dead Sea and Arava,
Science Center, Mitzpe Ramon, Israel Abstract
2
Eilat Campus, Ben Gurion University in The forecasted increased frequency and intensity of extreme climatic events may
the Negev, Eilat, Israel strongly affect ecosystem structure and function in the future. It is unclear how
3
Mitrani Department of Desert Ecology, ecosystems will function in the long run over a large spatial scale under a new
Ben-Gurion University of the Negev, Sede
Boqer Campus, Midreshet Ben Gurion, extreme water cycle. This open question calls for a conceptual framework as a
Israel fundamental basis for theoretical and experimental exploration of ecosystem
function on a large scale driven by an extreme climate envelope. To assess the
Correspondence
Elli Groner problem on a large scale, we investigated hyper-arid ecosystems (HAEs) as natu-
Email: elli@adssc.org ral tangible models that already function under an extreme climatic envelope.
Our new assertion is that if extreme climate change drives arid lands to function
Funding information
Israel Science Foundation, Grant/Award under alternate extreme conditions, then arid land ecosystems will function like
Number: 964/14; Ministry of Science and an HAE as an alternative state, rather than progress to desertification. To sup-
Technology, Israel; Open Spaces for
Gvanim LTER
port our assertion, we developed a conceptual framework of HAEs that includes
a geo-hydrological “abiotic engine” that drives HAE function by soil moisture
Handling Editor: Sujith Ravi diversity and plant functional groups. Based on this conceptual framework, we
suggest incorporating two new hypotheses in climate change studies to advance
our understanding of responses of large-scale, water-limited ecosystems:
(1) Hydro-climatic extremes in water-limited ecosystems will reduce the degree
of resource conservation by slope ecosystems due to reduction in plant cover
and soil. The decreased ecosystem function on the slope will be compensated for
by increasing the effect of the abiotic engine on the ephemeral stream, thus
enhancing meta-ecosystem functioning in the ephemeral stream. (2) In
water-limited ecosystems, climate change toward hydro-climatic extremes will
rescale the dominant hydro-ecological processes of pulse–reserve, source–sink,
and connectivity along the semiarid, arid, and HA gradients in two ways:
(i) shrinking of both spatial and temporal dimensions; and (ii) shrinking in the
temporal dimension and expanding in the spatial dimensions. The first rescaling
trajectory is related to biodiversity–ecosystem function and the second to the abi-
otic engine processes.
KEYWORDS
connectivity, desertification, eco-hydrology, source–sink, stream order
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided
the original work is properly cited.
© 2023 The Authors. Ecosphere published by Wiley Periodicals LLC on behalf of The Ecological Society of America.
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2 of 16 GRONER ET AL.
INTRODUCTION worldwide (Martinez-Valderrama et al., 2020). In

hyper-arid areas, vascular plants can only survive in
According to meteorological models, climatic changes are ephemeral streams that receive runoff water from the
expected to force ecosystems to function under increas- slopes. HAEs are driven by long dry phases, representing
ingly extreme climatic conditions. Climate scenarios pro- drought conditions, that are intersected by extremely
ject an increase in the frequency and intensity of extreme short wet phases (flash floods). We suggest that HAEs
climatic events that will directly affect the structure and are natural tangible models that can provide insight on
function of present and future ecosystems (IPCC, 2021; the long-term sustainability of ecosystems’ functioning
Reichstein et al., 2013). Drought and flood events signifi- under extreme water cycling. We propose that investigat-
cantly modulate the water cycle; the degree of their effect ing ecosystems such as HAEs, which have evolved to
depends on both the intensity and duration of the event. cope with extreme hydrological conditions, may increase
Changes in frequency and magnitude of extreme events our conceptual knowledge on ecosystem response and
may force ecosystems to function under two contrasting function under extreme climatic regimes. We assume that
effects of water cycling, causing long-lasting shifts in eco- if extreme climate changes drive arid lands to function
system state and resulting in plant mortality and soil loss under alternating drought and flood conditions with
(Navarro-Cerrillo et al., 2018). The result is that unprece- intensifying frequency and magnitude, then arid land sys-
dented changes in ecosystem functioning and dryland tems will be forced to adapt and function similarly
mechanisms are already occurring in non-water-limited to HAEs.
areas (Grünzweig et al., 2022). It is becoming clear that Some models predict that climate change will cause
more and more ecosystems are facing new water balance desertification in the sense of degradation of arid and
challenges (Allan et al., 2020). As water-limited ecosys- semiarid systems (Martinez-Valderrama et al., 2020). We
tems, deserts react to changes in the water regime and question this notion. Climate change will push the condi-
can serve as models for management of other ecosystems tions toward extreme changes in frequency and magni-
(Grünzweig et al., 2022). Our understanding of ecosystem tude of droughts and floods. As rainfall declines, the
functioning under extreme hydrological regimes is importance of runoff increases, and with it, the implica-
hindered by the lack of a conceptual framework to theo- tions on biota become more significant. We hypothesize
retically and experimentally explore the long-term func- that climate changes will push dryland systems along the
tioning of ecosystems affected by extreme climatic events. climatic gradients toward extremities in which HAEs
Since experimental studies are limited to small spatial already exist. These changes will cause many processes to
and temporal scales, whereas ecosystems respond to switch their operation from the patch level to the land-
extreme climatic events at very large scales, investiga- scape (watershed) level and to rescale the structure and
tions are best carried out on ecosystems that already function of the system. This will lead to a new extremity
function under an extreme climatic envelope. Gaining that can be seen in a different light at the landscape scale.
insights on the function of such ecosystems can help to Following these suggested predictions, we propose a new
construct a conceptual framework for further experimen- specific hypothesis as an alternative to the desertification
tal and theoretical studies, aiming at predicting the effect paradigm. Our alternative hypothesis suggests that when
of extreme climate on dryland ecosystems in general. studying climate change on the watershed scale, it is evi-
If climate changes push ecosystems along a climatic gra- dent that climate change toward aridity may not degrade
dient toward the extreme (dry) end, then studying pres- the entire ecosystem but, instead, “compensation” will
ently extreme ecosystems may assist us in understanding occur. During this process, the functioning of the ecosys-
possible futures of the less extreme ecosystems. The most tem will shift in space but will not necessarily deteriorate
extreme water-limited (dryland) ecosystem is the completely due to the catastrophic shift. In our alterna-
hyper-arid ecosystem (HAE). tive hypothesis, we suggest that extreme climate will
HAEs are defined as areas in which precipitation is rescale the connectivity of runoff flow from short flow
less than 5% of potential evaporation (aridity index <5%; pathways within the slope to long pathways from slope to
Safriel et al., 2006). Although the structure and functions valley. This rescaling process will increase walleye eco-
of HAEs are substantially different from those of arid ones, system functioning that compensates for the decrease in
they are much less studied. A Web of Science search using slope functioning. The compensation hypothesis is an
the keywords ecology and ecosystem with arid provided alternative to the desertification hypothesis that predicts
14,887 papers, compared with only 408 papers using whole ecosystem degradation as the climate shifts toward
hyper-arid (with a maximum of less than 50 a year), which extremeness.
is less than 3% of papers on arid systems. Due to climate To elaborate on our hypothesis, we first explain how
change and anthropogenic impacts, HAEs are expanding HAEs operate differently to arid ones. We show how
ECOSPHERE 3 of 16
HAEs are an extreme ecosystem in many aspects along a Upscaling the hillslope processes to the watershed scale,
climatic gradient and how climate change can drive other that is, slope–ephemeral stream relationships, implies that
systems toward these extremities. We then break down stronger sink functions of the vegetation patches on the
the mechanisms that drive HAEs and explain the struc- slope weaken the source–sink interaction between slope
ture and function of HAEs in respect to abiotic and biotic and ephemeral stream. As such, a trade-off exists between
“engines.” We then elaborate why arid watersheds may water availability for ecosystem functions on the slopes and
experience a state shift that compensates for slope ecosys- the ephemeral streams. Higher water and soil conservation,
tem degradation by valley restoration in response to due to the strong sink function of the slope, implies less run-
predicted climate changes toward extremeness. off and soil flow from the slope to the ephemeral stream.
ARID ECOSYS T E M Arid ecosystem degradation

HYDRO- ECOLOGIC AL FUNC TION
AND DEGRADATION The desertification paradigm proposes that degradation
of arid ecosystem slopes, that is, collapsing of the func-
Arid ecosystem function tion of the two-phase mosaic as a resource conservation
mechanism, will result in desertification throughout the
To understand the expected ecosystem shift under more entire system. The frequency, magnitude, and spatiotem-
extreme climate, we must assess the water flow processes poral distribution of droughts and floods that have
that regulate arid ecosystems’ functioning on the water- already increased around the world (Di Baldassarre et al.,
shed scale prior to the climate changes. Arid watersheds 2010; ISDR, UN, 2017; Winsemius et al., 2016) cause this
(other than flat plains) are composed of two interacting type of degradation. Many ecosystems may experience
ecosystems: slopes and valleys. The function of slopes in periods of prolonged drought conditions and extreme
dryland ecosystems is regulated by feedback between abi- flood events in the future (IPCC, 2021). Arid areas were
otic and biotic processes (von Hardenberg et al., 2001). found to be sensitive to global climate change. Climate
Positive and negative processes induce biotic engineering models and weather trends forecast an increase in
on a small scale, creating a patch mosaic of vegetated drought frequency (Collins et al., 2013; Hill et al., 2018),
patches embedded in a matrix of biocrust and rocks. This conjugated with higher intensity of rainfall events
landscape structure redistributes rainfall pulses via runoff resulting in more frequent floods (Donat et al., 2019).
flows, creating water-deprived patches under the biocrust, A decrease in the length of the rainy season and an
and water-enriched patches under the vegetative canopies. increase in the frequency of large rainfall events have
This redistribution, attributed to the different infiltration been documented (Wittenberg et al., 2007). In arid envi-
rates of the two patch types, drives source–sink dynamics ronments, paleohydrology research found that the second
(Shachak et al., 2008). half of the 20th century was characterized by high-flow
The vegetation patches serve as sinks since their high magnitude events (Greenbaum et al., 2000).
infiltration rates induce belowground engineering by plant According to the desertification paradigm, as the
roots and the associated microbiome that produce magnitude and frequency of extreme events increase, pri-
macropores and improve soil aggregation (Gilad et al., mary productivity will decline, shrubs will die, and the
2007). Hydrophobic biocrusts have low infiltration rates slope ecosystem will collapse to a level at which ecosys-
and thus induce runoff generation (Eldridge et al., 2000; tem productivity is minimal (Whitford & Duval, 2020).
Weber et al., 2016). The runoff produced by the biocrust However, in the coming sections we suggest that arid sys-
areas infiltrates into the vegetated patches, and thus water tems may alternatively shift to a new stable state that is simi-
resources are conserved on the slope and their leakage to lar to the current state of HAEs, which is not a degraded
the ephemeral streams is minimal. This mechanism, state. In order to understand this assertion, we must distin-
which sustains slope productivity and diversity, is guish between the water flow processes of arid and
self-reinforcing as it increases water availability and plant hyper-arid ecosystems along the extreme climate gradient.
growth in the vegetated patches. Rainfall redistribution by
runoff from biocrusts as source areas to vegetation patches
as sinks is therefore the key for maintaining hillslope func- An alternative to the desertification
tionality in arid ecosystems (Shachak et al., 1999). This paradigm: The compensation hypothesis
process occurs on a scale of a few centimeters to very few
meters—the distance runoff water can travel from the Our proposed alternative hypothesis suggests that while
biocrust patches to the shrub patches on the slopes. biological productivity on the slopes may indeed decline
under new extreme climate conditions, the biological responses of drylands to climate change and suggest a
productivity of the entire watershed may not be lower, set of questions for future research.
that is, the loss of productivity on the slopes is compen- Our model is based on a hot, rocky, hilly desert in
sated by higher productivity in the riverbeds (wadis). which topography and geomorphology regulate the water
Our hypothesis states that slope degradation driven by balance, and scant rain falls mostly in the winter. This
extreme hydrological regimes can rescale source–sink model is directly suitable to describe areas that may shift
relationships so that under extreme climatic conditions, to HAEs under climate change. With some modifications,
the entire slope acts as the source of runoff water and the model can describe any ecosystem that is driven by
soil, and the ephemeral dry bed stream becomes the sink. source–sink patch dynamics. However, as deserts include
Our assertion is that in contrast to arid ecosystems, HAEs a large variety of landscapes and conditions, it remains
do not have large water sinks along their slopes, and questionable whether our model can be applied to sandy
therefore, a large proportion of the runoff water reaches deserts with insignificant runoff generation, or deserts
the riverbeds. Accordingly, dry riverbeds of ephemeral with monsoon events and plateaux.
streams serve as water- and soil-enriched landscape units
and compensate for slope degradation by increasing pro-
ductivity and other ecosystem functions. The abiotic engine of HAEs
Thus, under a regime shift induced by climatic
extremes, the former arid landscape may function as an Defining HAEs as the dryland ecosystem with the most
HAE with nonvegetated rocky slopes that are a source of extreme hydrological conditions requires a fresh look at
runoff and ephemeral streams that are greater sinks. the dimensions of extremeness and their control over eco-
Accordingly, the overall functioning of the watershed of system function in relation to source–sink (Merino-Martín
arid ecosystems will resemble that of HAEs and will not et al., 2012) and pulse–reserve (Collins et al., 2014) dynam-
collapse as the desertification paradigm suggests. ics. We suggest five interrelated dimensions that delineate
To support our assertion that the functioning of dry- HAEs as ecosystems that operate under extreme hydrologi-
land ecosystems can shift toward HAEs in response to cli- cal cycle conditions (Figure 1). The first dimension that
mate changes characterized by extreme hydrological drives ecosystem dynamics is the degree to which the
regimes, we present the main properties that define the structure and function of the ecosystem depend on runoff
extremeness drivers of water-limited ecosystems. We then for soil moisture content. Runoff dependency correlates to
suggest a conceptual framework describing the structure rainfall gradient; arid regions with lower rainfall magni-
and function of HAEs as the ecosystem that functions tudes are more runoff-dependent (Shachak et al., 1998). In
under the most extreme climatic envelope. The frame- HAEs, vascular plants cannot survive in patches that only
work is based on core principles of dryland hydro- receive rainwater; therefore, their existence is completely
ecological theories and dry river functions, as reported in dependent on high-magnitude, low-frequency runoff. In
the literature. The conceptual framework includes two contrast, arid ecosystems support vascular plants in plains,
“engines” that drive the system: (1) the geo-hydrological hilltops, and slopes and can function on low-magnitude,
“abiotic engine” that drives biotic dynamics under six high-frequency runoff water. HAEs are at the extreme end
interrelated dimensions that delineate HAEs as systems that of the runoff-dependent gradient (Figure 1A) and the sys-
operate under extreme hydrological conditions; and (2) bio- tem is runoff limited, implying that HAEs represent the
diversity and HAE functioning driven by soil moisture most runoff-dependent ecosystem.
diversity, and plants’ functional and spatial diversity along The dependency of HAEs on high-magnitude runoff
the fluvial network. This “biotic engine,” the final product implies that the system depends on intensive rainfall
of the abiotic engine, affects the entire biodiversity– events and unique surface properties that generate run-
ecosystem function of the HAE. Subsequently, in the off. At low rain intensities, slope–valley runoff connectiv-
Discussion, we refer to the link between the conceptual ity is negligible and does not affect ecosystem functions.
framework and the potential effects of climate change on Only rain above a certain threshold generates runoff that
water-limited systems, emphasizing the rescaling of water reaches the ephemeral streams and drives ecosystem
flow connectivity processes as a main mechanism. We then functions. Since HAEs are entirely runoff-dependent,
emphasize the significance of incorporating HAE stud- they are reliant on a certain rainfall pattern, that is, the
ies as a tangible model for understanding processes of frequency distribution of rainfall events (Figure 1B). Only
dryland transitions in ecosystem functioning under high-magnitude rainfall events effectively contribute to
extreme hydrological regimes. Finally, we summarize the ecosystem (Chen et al., 2019). Generally, rainfall in
the potential contributions of HAE research for drylands follows a gradient of frequent low-magnitude
advancing experimental and theoretical studies on the rainfall events to scarce high-magnitude events. While
ECOSPHERE 5 of 16
F I G U R E 1 Extremeness dimensions of hyper-arid (HA) ecosystems (dotted red lines) as systems under extreme hydrological
conditions. Extremeness implies the high dependency of HAE on: (A) runoff, (B) low frequency, high-magnitude rainfall; (C) high spatial
scale of hydrological process operation and high source to sink ratio; (D) high connectivity pathways; and (E) geological, geomorphological,
and pedological diversity.
semiarid and arid ecosystems utilize the entire range of flows on a small scale from rock patches (source) to soil
the magnitude distribution of rainfall events as water patches (sink; Yair & Shachak, 1982). In HAEs, the
resources, HAEs exploit only the scarce high-magnitude source–sink network functions on a much larger scale;
events (Figure 1B). This property defines HAEs’ second the entire slope is the source of runoff and the stream is
dimension of extremeness, that is, system dependency on the sink. The limited soil cover on the slopes in HAEs
rare, extreme rainfall events. The HAE is unique in that creates negligible sink patches. Although these patches
most rainfall events have little effect on the ecosystem. In may collect runoff water, the soil moisture level in these
dryland ecosystems driven by runoff, landscape configu- “sinks” remains insufficient to support vascular plants,
ration, that is, the runoff source and sink networks of the hence the source–sink impact on productivity at this
watersheds, is essential for their function (Peters scale is negligible. Cyanobacteria, the main component of
et al., 2015). biological soil crust in arid ecosystems, are present in
Figure 1C illustrates the scale at which ecosystem HAEs (Warren-Rhodes et al., 2007) but do not form
processes operate in water-limited ecosystems. In arid biocrusts, hence soil erosion is exacerbated during strong
systems, the source can, for example, be a rock while the floods. The minimum spatial scale in which source–sink
sink may be a soil patch downslope of it, that is, water dynamics becomes relevant in HAEs is the watershed
scale (Figure 1C). Because the entire slope is the source The link between geodiversity and
in HAEs, their source to sink ratio is much higher com- biodiversity
pared with arid systems, in which shrub patches on the
slopes and the streams are sinks. We integrated HAEs’ extreme characteristics into a concep-
Connectivity between different source and sink land- tual model that represents the link between hydrodiversity
scape units plays a key role in dryland ecosystem function- and geodiversity. These properties act as the abiotic engine
ing (Okin et al., 2015). The large spatial scale, as a that drives biodiversity and ecosystem function in HA land-
precondition for HAE operation, requires high water con- scapes (Figure 2). In the model, hydrodiversity is generated
nectivity to link source and sink areas. Source and sink by the transformation of rare, high-magnitude rainfall
areas can be connected in HAEs only if the connectivity pulses (Figure 2i) into runoff pulses (Figure 2ii) that flow
pathway can link runoff generated on the slopes with the various distances along connectivity pathways (Figure 2iii).
entire network of dry riverbeds. Most rain events do not Rare, high-magnitude rainfall pulses are characteristic of
generate runoff that reaches the streams. The fourth HA climate. However, their conversion to runoff pulses of
dimension that drives HAEs is the dependency on long variable lengths is the product of interactions between
connectivity pathways, which are rare because of the low rainfall patterns and geodiversity (Okin et al., 2015). In
frequency of rainfall events that generate high-magnitude the model, these interactions describe the controlling
runoff (Figure 1D). Conversely, in arid and semiarid land- effect of slope geodiversity components (geology and geo-
scapes, low-magnitude rain increases ecosystem function morphology) on runoff generation and length of connec-
because short connectivity runoff supports the growth of tivity pathways (Figure 2iv). The temporal diversity of
shrubs and annuals on the slope. runoff pulses that are generated on the slopes and the
The extreme dimensions that define HAEs demonstrate length of connectivity paths determine the next compo-
the strong link between hydrological diversity and nent of hydrodiversity—the soil moisture diversity within
geodiversity (Figure 1E). In the case of HAEs, hydrological the dry riverbed (Figure 2v). Under relatively low magni-
diversity refers to the variety in magnitude, frequency, dura- tudes of runoff pulses, slope–ephemeral stream connectiv-
tion, and timing of runoff pulses. Geodiversity refers to the ity is limited to first-order streams, and source–sink
heterogeneity of the catchment’s physical environment, such connectivity is restricted to part of the catchment area.
as variable geological formations, geomorphological land- Increasing magnitude of runoff pulses lengthens the con-
forms, and topography. In HAEs, geodiversity and rainfall nectivity pathways, thus linking water flow to higher
pulses are the integral drivers of ecosystem functions that order streams until the entire catchment is connected.
regulate the hydrological diversity, which is crucial for sus- Catchment connectivity and geodiversity throughout the
taining ecosystem processes. Geodiversity provides the sub- stream orders (pedodiversity) regulate the high spatiotem-
strates and landform mosaics that determine the dynamics poral variability in soil moisture (Figure 2vi). Soils with
of runoff pulses; these control primary and secondary pro- high water retention, due to their grain size distribution
duction and soil functions across the catchment riverbed. or thickness, increase primary productivity even at rela-
As suggested by Hjort et al. (2015), ecosystems are the tively low-intensity runoff events. Patches with low water
product of three realms of diversity: geo-, bio-, and cli- retention capacity cannot support productivity at any run-
mate diversity. However, in HAEs, where the system off intensity. Hence, soil moisture varies among the net-
operates under extreme climate variability and depends work of stream orders and creates different hydrological
on runoff pulses, geodiversity predominantly regulates niches along the catchment network. Low stream orders
ecosystem dynamics (Okin et al., 2015). We propose that receive runoff more frequently; since the contributing
in drylands, the relative importance of geodiversity rises area is relatively small, low-intensity rainfall events are
as rainfall magnitude decreases (Figure 1E). Within the sufficient to generate runoff along short connectivity path-
aridity gradient of semiarid, arid, and HA ecosystems, ways. High stream orders receive rare runoff input of high
HAEs are the most dependent on geodiversity. magnitude due to the larger contributing (source) area.
By referring to the hydro- and geodiversity realms as The wide variety of operating scales is common to all
the abiotic engine that drives HAEs, we may conclude that processes related to hydrodiversity. The largest cross-
under extreme conditions, the engine represents large scale relationship is the connectivity between runoff gen-
spatial-scale abiotic processes that regulate small-scale eco- erated in slopes throughout the entire catchment and a
system processes. Thus, the HAE operates as an extreme patch of soil in a high-order stream. The smallest cross-
cross-scale system in which the main abiotic processes scale relationship is the connectivity between runoff gen-
occur on a very large scale (slopes of the entire catchment) erated over a first-order watershed and a patch of sink
and the biotic processes take place on a very small spatial downslope in its ephemeral stream. The dependency of
scale (patches in the dry riverbed). the abiotic engine on a wide range of spatial cross-scale
ECOSPHERE 7 of 16
F I G U R E 2 Conceptual model of the abiotic engine of hyper-arid (HA) ecosystems (dotted red lines) that drives biodiversity and
ecosystem function. The figure integrates water flow (blue) that interacts with various components of geodiversity (orange) and regulates
biodiversity and ecosystem functions (green). The water flow pathway (ii, iii, v) is driven by rare, high-magnitude rainfall events (i).
Transformation of the rainfall pulses to soil moisture reserves among stream orders is a scale-dependent source–sink process that operates
on an extremely large spatial scale of connectivity pathways (iii, v) and is regulated by the various components of geodiversity (iv, vi).
relationships between geo-hydrological processes implies structure in terms of species composition, functional
extremeness in the source–sink ratio, producing water- diversity, and vegetation pattern, that is, the biotic patch
enriched patches that sustain life. mosaic of the dry riverbed. Ephemeral streams sustain a
range of functional plant patches, such as annual and peren-
nial herbaceous plants, shrubs, and trees (Breslau et al.,
Biodiversity and ecosystem function 2019; Fossati et al., 1999; Tooth & Nanson, 2000). Each plant
in HAEs patch is a fertility island that supports aboveground and
belowground secondary production and the interactions
In contrast to HAEs’ abiotic engine that operates on the between them. These functions include maintenance of
catchment scale, biodiversity and ecosystem functioning are diverse invertebrate assemblages (Davidson & Groner,
mostly restricted to the ephemeral streams, where vascular 2021; Sanchez-Montoya, Guerrero-Brotons, et al., 2019;
plants exist. Since the effect of the abiotic engine on the eco- Sanchez-Montoya, Tockner, et al., 2019), composed
system is stream-order dependent, we propose that the tar- mainly of mites, springtails, ants, beetles, and spiders.
get scale for the biodiversity and ecosystem function model This invertebrate community plays a key role in decompo-
is defined by stream order. Stream order defines the hydro- sition and nutrient cycling processes. The plant patches
logical niches (sensu Silvertown et al., 1999) available at a also include debris piles composed of woody and leaf litter
given watershed, which in turn determine plant community (Sanchez-Montoya, Guerrero-Brotons, et al., 2019) that
are sites for organic matter (OM) processing and nutrient network of plants and animals create soil properties with
cycling along the fluvial network. The decomposition pro- a higher rate of litter decomposition and nutrient cycling
cesses are carried out by diverse invertebrate and micro- than that in the inter-patch soil (Lavelle & Spain, 2001).
bial communities that are active after runoff pulses when Hence, most ecosystem functions depend on patterns of
soil moisture rises above a certain threshold (Marcé shrub distribution, which depend on the distribution of
et al., 2019; Marxsen et al., 2010; Zoppini et al., 2014; soil moisture. Although photodegradation also affects
Zoppini & Marxsen, 2011). Debris and detritus maintain decomposition of OM, its impact varies among systems
diverse invertebrate communities (Sanchez-Montoya, (Grünzweig et al., 2022).
Guerrero-Brotons, et al., 2019) that may sustain life and We incorporated HA biodiversity and ecosystem func-
also drive belowground processes. Decomposition and the tion when modeling the abiotic engine. Soil moisture
nutrient cycle link biodiversity to geodiversity; this impor- diversity (heterogeneity) along the various stream orders
tant link operates on both the patch and the fluvial net- of the fluvial network was determined as a product of the
work scales (Grünzweig et al., 2022). On the patch scale, abiotic engine (Figure 3i). We propose (based on observa-
litter deposition under the plant canopy and the plant’s tion on HA catchments in the Negev) that the large scale
species-specific qualities drive soil development and func- of the HAEs’ abiotic engine affects plants’ functional and
tions that create soil-specific properties. These properties spatial pattern diversity (Figure 3ii). We suggest that HA
determine pedodiversity, specifically, the variability in soil ephemeral streams exhibit a gradient of the three main
organic matter (SOM) and nutrient availability (Thibault functional plant types along the stream orders. Slopes are
et al., 2018). On the fluvial network scale, the organic bare of vascular plants. Herbaceous plants are dominant
debris patches under the plant canopies interact with run- in low-order streams, shrubs in medium-order streams,
off pulses that redistribute the organic debris by erosion while trees are associated with high-order streams. The
and deposition processes along the channels of the dry plant type along the stream-order gradient is determined
river (Jacobson et al., 2000). Runoff intensity and the by the interaction between soil moisture distribution, as
length of connectivity pathways, along with the geomor- regulated by catchment geohydrology, and the plants’
phological structure, determine the erosion and deposi- hydrological niche and hydrological trait diversity. We
tion sites along the fluvial network and the redistribution attribute the stream-order gradient to the spatial niche
of OM among the plant patches. Some plant species act as partitioning on a catchment scale; soil water distribution
a source of OM while others act as sinks. The OM redistri- is determined by the abiotic engine. The phenomenon of
bution can also induce OM deposition in new sites that hydrological niche partitioning has been observed in a
contribute to the formation of new vegetative patches variety of ecosystems (Kulmatiski & Beard, 2013; Lipkin,
(Jacobson et al., 2000) that stimulate patch dynamics and 1971; Nippert & Knapp, 2007; Schume et al., 2004;
the creation of vegetation patterns. Schwendenmann et al., 2015). On a small, single
The properties and ecosystem functions driven by stream-order scale, the three functional plant types can
plant patches are also the outcome of ecosystem engi- probably coexist due to the complementarity of the
neering by plants (Gurnell, 2014). Patch modulation by hydrological niche. Hydrological niche complementarity,
plants plays a key role in the development of soils’ physi- or niche partitioning by soil water, can occur due to
cal, chemical, and biological functions. By stabilizing the interspecific differences in root distribution (Li et al.,
sediments that are deposited under their canopy, plants 2018; Ward et al., 2013) or by differential responses to
modify the physical structure of the sediment, supporting shifting water availability (Asbjornsen et al., 2008;
the development of aggregates and macropores that regu- Nippert & Knapp, 2007). In our model, we propose
late both soil moisture regime and the development of a that hydrological niche complementarity may be an
soil microbiome (Batz et al., 2014; Gutierrez & Jones, important mechanism contributing to the positive
2006; Verboom & Pate, 2006). The state of the stabilized biodiversity–ecosystem functioning relationship in HAEs
patches generates a series of changes in physical, micro- on temporal and spatial scales (Figure 3iii). Interspecific
bial, and biogeochemical processes of the plant patches differences among herbaceous plants, shrubs, and trees
(An et al., 2010; Batz et al., 2014; Chotte, 2005; in the timing of water use for productivity pulses
Kay, 1998). (as depicted in Figure 3iv) enhance temporal stability,
Soil patch engineering by plants leads to further engi- while the reliance on variable soil water sources along
neering by macro- and meso-fauna. Through their activi- the fluvial network (Figure 3i) increases the spatial
ties, soil faunae increase the plants’ impacts on soil stability of plants’ function. Based on growing evidence
aggregation and porosity, and hence associated hydrolog- suggesting that functional diversity increases plant pro-
ical properties, and increase SOM availability for micro- ductivity and stability over time and space (Isbell
organisms (Lavelle, 1997). The combined effects of the et al., 2009; Lehman & Tilman, 2000; Zhang et al., 2023),
ECOSPHERE 9 of 16
F I G U R E 3 Conceptual model of hyper-arid ecosystem biodiversity and ecosystem function. The model depicts the role of the end
product of the abiotic engine, soil moisture distribution over stream orders (blue), as the driver of plant (green) and soil (brown) functions in
interaction with dry riverbed geodiversity (orange). Soil moisture distribution over stream orders (i) regulates the abundance of the
functional diversity of herbaceous and woody vegetation, perennials, and trees (ii) along stream order. Plant functional diversity controls
primary production pulses over time (iv). Plants, via litter production (v), induce soil functions (vi) that include decomposition by soil fauna
and soil microbiome. End products of soil functions are soil nutrients (vii) and soil organic matter (SOM; viii) that affect soil moisture and
pedodiversity of the alluvial stream bed by soil ecosystem engineers (EE; ix). Soil positive engineering increases ecosystem productivity.
we argue that hydrological trait diversity among herba- from runoff patterns determined by geodiversity. Thus,
ceous plants, shrubs, and trees regulates productivity and primary productivity increases during the rare events of
stability of HAEs under extreme climatic conditions. In sufficient rain that generates runoff over a large scale.
our framework, we view hydrological trait diversity of The large sink (the streams) sustains relatively high pri-
plants as the biotic response to soil moisture diversity mary productivity and may compensate for the low pro-
generated by the abiotic engine that drives HA primary ductivity at the source (slopes), so that at the watershed
production pulses. Hydrological trait diversity supports scale, productivity is relatively high.
relatively high primary productivity and maintains biodi-
versity, processes that are linked by positive feedback
(Groner & Novoplansky, 2003). We acknowledge that DISCUSSION
microorganisms, such as cyanobacteria and lichens,
inhabit the slopes (Bull & Asenjo, 2013; Vítek et al., 2010) Hyper-arid eco-hydrology
and contribute to watershed productivity; however, this
productivity is much lower than that of vascular plants. Climate change models project a future biosphere domi-
We suggest that although water, the limiting nated by hydro-climatic extremes characterized by high
resource, is so scarce in HAEs (by definition), biodiversity variation in precipitation regime, namely severe droughts
is maintained due to the hydrological diversity created and pulses of floods. In many regions, the effects of
hydro-climatic extremes on ecosystem are projected to Landscape compensation hypothesis

induce state change and even biome shifts (Kulmatiski
et al., 2020). Understanding the direction and magnitude On the community level, hydro-climatic extremes may
of ecosystem changes under these predictions is critical induce substantial spatiotemporal changes in vegetation
for coping with the ecosystem dynamics of an extreme composition and structure (Ummenhofer & Meehl,
world. Living in a world in which ecosystems function 2017). This effect can cascade to the ecosystem level and
under hydro-climatic extremes requires accurate projec- modify primary and secondary productivity and decom-
tions of ecosystem trajectories. We must predict the position processes (Tilman & El Haddi, 1992). In addi-
changes of ecosystem state, from present to future, based tion, mass mortality and widespread dieback events
on experimental data, mathematical models, and, as we driven by extreme drought can modulate landscape struc-
suggest in this paper, tangible models of natural ecosys- ture and function by intensifying runoff and soil erosion
tems that function under hydro-climatic extremes. processes (Weifeng et al., 2012). Plant mortality events
Characterizing ecosystem shift under extreme events, and associated landscape effects are predicted to acceler-
such as intense precipitation, floods, and droughts, is par- ate under increasingly extreme droughts (Momcilo et al.,
ticularly difficult because of the small spatial scale and 2019). The spatiotemporal reorganization of the vegeta-
relatively short observational records of experimental tion cover, driven by hydro-climatic extremes, is funda-
studies. One must consider that hydro-climatic extremes mental for the function of the ecosystems. If extreme
have large-scale effects on ecosystems, including cross-scale climatic events continue to become more frequent, spa-
interactions and feedbacks between physical and biological tial and temporal reorganization of vegetation cover will
processes (Marshall et al., 2008). Models, as a complemen- occur and may trigger reorganization in the spatial distri-
tary tool that indicates ecosystem transformation under bution of ecosystems. In water-limited systems, the
extreme climatic conditions, can overcome the scaling prob- spatial distribution of ecosystems and their interactions
lem (Prentice et al., 2007; Rehfeldt et al., 2006). However, within and across watersheds strongly depend on
their capacity for assessing the magnitude of ecosystem geo-hydrological processes and their modulation by plants
transformation under future extreme climate scenarios is as ecosystem engineers (Gilad et al., 2007). In fact, catch-
limited by the difficulty in evaluating model performance ments are networks of meta-ecosystems defined as a set of
against empirical small-scale records and the limitations of ecosystems connected by spatial flows of water, energy,
extrapolation in cases of nonlinearity and feedback materials, and organisms across ecosystem boundaries
(Jackson & Williams, 2004). Hence, to enhance upscaling (Loreau et al., 2003). The boundaries between ecosystems
and the understanding of experimental results and to pro- in catchments can be delineated in relation to watershed
vide data and insights related to broadscale spatial and tem- (slope–ephemeral stream) and catchment (stream order)
poral patterns and processes, data across a broad range of components. According to our biotic model (Figure 3),
scales must be obtained. We cannot experimentally examine HAEs in a catchment consist of a set of ecosystems
small-scale ecological responses to hydro-climatic extremes connected by water flows where each stream order repre-
and simply upscale them linearly to the catchment scale, due sents a different ecosystem. The connector and the driver of
to inherent biases associated with cross-scale interactions, these meta-ecosystems is the abiotic engine (Figure 2) that
spatial heterogeneity, and nonlinearity (Peters et al., 2007). controls biodiversity and ecosystem function by the
Therefore, we advocate that to understand and pre- strength of slope–ephemeral stream runoff connectivity.
dict the extent and ecological impacts of hydro-climatic HA catchments represent the highest degree of
extremes on ecosystems, naturally functioning HAEs are slope–ephemeral stream runoff connectivity. Since the
a promising area of study. We propose that untangling slopes are bare bedrocks without soil or vegetation, the con-
the properties of HAEs can generate new hypotheses nectivity between slope and ephemeral stream riverbed is
related to the effect of hydro-climatic extremes on high. In water-limited systems, HA catchments exemplify
large-scale landscapes. the extreme case in which the meta-ecosystem networks
From our analysis of HAE properties, specifically, their are restricted to the ephemeral stream, while in arid and
dimensions of extremeness (Figure 1), their abiotic drivers semiarid catchments, the networks are more complex and
(Figure 2), and their biodiversity and ecosystem function include ecosystems on the slopes. We propose that in catch-
(Figure 3), we suggest incorporating two hypotheses in ments whose meta-ecosystem networks include ecosystems
future experimental and modeling studies of climate on the slope, there is a trade-off in water allocation for eco-
change toward extremes. These may advance our under- system functioning on the slope and the ephemeral
standing on the large-scale responses of water-limited eco- stream’s riverbed and between the impact of water on eco-
systems to climate change in terms of hydro-climatic system functioning on slopes and its impact on ecosystem
extremes. functioning in the riverbeds. Water that leaves the slope
ECOSPHERE 11 of 16
(through runoff) can contribute to productivity in the unchanged, still remains. According to Yair et al.
riverbed. In HAEs, the riverbeds support almost all the (2021), runoff in dry riverbeds is higher in HAEs
watershed’s productivity. In arid systems, high water con- because less water is lost on slopes, perhaps resulting
servation on the slopes by vegetation and biotic ecosystem in higher productivity.
engineering lowers the effect of the abiotic engine on the
biodiversity and ecosystem function of the ephemeral
stream. Rescaling of processes hypothesis
We suggest that climate change toward hydro-climatic
extremes modifies this trade-off. In HA catchments, the We propose that the dominant processes that regulate the
trade-off favors the ephemeral streams at the cost of extinc- abiotic engine and biodiversity–ecosystem functions of
tion of slope ecosystems, demonstrating that HAE net- HA, arid, and semiarid ecosystems are identical, but have
works can persist even when slope ecosystems are different rates and spatiotemporal scales of operation. We
eliminated due to hydro-climatic extremes. We propose also suggest that in the three dryland regions, the scale of
that climate change toward extremeness drives a shift in the processes varies in response to the magnitude and fre-
trade-off that decreases slope resources and enriches quency of rainfall pulses and interacts with the specific
ephemeral stream resources, by reducing plant cover on geodiversity of each region. According to the dryland eco-
slopes through widespread mortality and dieback events. logical paradigm, the dominant processes that regulate
The frequency and magnitude of riverbed runoff may dryland ecosystem functions are pulse–reserve (Collins
increase under a decreasing rainfall regime, as vegetation et al., 2014), source–sink (Merino-Martín et al., 2012),
and soil, which retain water on the slopes, are removed and connectivity (Okin et al., 2015) dynamics.
(Belachsen et al., 2017; Cammeraat, 2004; Yair et al., The pulse–reserve framework (Collins et al., 2014)
2021). By studying the function of HAEs, we can infer that refers to the temporal distribution and usage of resources.
higher flows of resources from the slopes to the ephemeral It describes the ecological processes in drylands in which
stream increase ephemeral stream productivity and diver- rain pulses drive a set of abiotic and biotic pulses that
sity, at least up to the (flood intensity) point when water cease when soil moisture is low. When the pulses termi-
becomes a disturbance rather than a limiting resource. nate, they leave abiotic and biotic reserves that drive a
This implies that in order to understand the trajectories of new set of pulses stimulated by precipitation events
water-limited ecosystems under future hydro-climatic within the next growing season. Ephemeral streams are,
extremes, we should refer to the catchment scale and by definition, based on pulses of water that temporarily
examine reorganization of the meta-ecosystems through- flow in the streams. The resultant soil moisture reserves
out the entire catchment. Overall catchment scale predicts, drive productivity and decomposition pulses that repre-
as evident in HA catchments, that resources lost from the sent ecosystem functions.
slopes enrich the ecosystem in the ephemeral stream, thus The source–sink framework relates to the spatial patchy
compensating for loss of productivity and diversity on the distribution of resources. It describes eco-hydrological pro-
slopes. Using HAE as a tangible model of ecosystem func- cesses in which runoff and associated materials flow from
tioning under extreme conditions suggests a general an area of low water infiltration (source) to a high infiltra-
hypothesis on the tendency of water-limited ecosystems tion area (sink), creating resource-enriched patches with
under the expected extreme climatic conditions of the higher productivity and diversity in comparison to the sur-
future. We propose that hydro-climatic extremes in rounding matrix. In HAEs, the rocky slopes are the source
water-limited ecosystems will reduce resource conservation and the ephemeral streams serve as sinks.
by slope ecosystems due to elimination of plants and soil. Connectivity encompasses the processes of movement
The subsequent decrease in ecosystem function on the slope of abiotic and biotic materials (soil, nutrients, and seeds)
will be compensated for by increasing meta-ecosystem func- from one location in the landscape to another as a func-
tioning in the ephemeral stream. In short, the landscape tion of the strength of the transporting factors (wind or
compensation hypothesis proposes that climate change water). In HAEs, the movement is from the slopes to the
toward extremeness should be investigated as a landscape dry riverbeds.
(catchment) phenomenon where climate change shifts The three dominating processes are linked and
plant cover and composition and causes spatial redistribu- together determine dryland ecosystem dynamics. Rainfall
tion of ecosystems. pulse properties regulate the strength of connectivity pro-
This implies that the net effect of extreme hydro- cesses, which in turn control the source–sink relationships
logical climate change is unpredictable. The question, that affect biotic and abiotic pulses and reserves.
whether total ecosystem productivity and diversity Our models of the abiotic engine driving the
in the catchment will decrease, increase, or remain biodiversity–ecosystem functioning of HAEs incorporated
the following components of the three dominant scale of operation of each of the abovementioned
processes: processes varies gradually and regulates the function of
the abiotic engine and biodiversity–ecosystem function.
1. Pulse processes that include transformation of rainfall Figure 4 depicts our interpretation of changes in spatio-
pulses to runoff pulses (Figure 4, arrow i), transforma- temporal scales of the dominant processes along the
tion of runoff to various soil moisture pulses and semiarid, arid, and HA gradient. The figure is based
reserves (Figure 4, arrow vi), and plant productivity on extensive literature that links pulse–reserve and
pulses (Figure 4, arrows ix–xi). source–sink processes to vegetation patterns and engi-
2. Source–sink processes in which hillslope geodiversity neering in semiarid and arid zones and in our models
is a source of water, sediments, and soil for the of HAEs.
ephemeral streams, whereas the soil patches in the In view of our model, we suggest a second hypothesis
various stream orders are sinks for the transported that asserts that climate change toward hydro-climatic
resources (Figure 4, arrow vii). extremes will shift the scales of processes in semiarid and
3. Connectivity processes, including runoff generation arid regimes toward the scales of HAE operation
and the varying length of water connectivity (rescaling; Figure 1). Using the definitions of spatial
pathways along the riverbed network (Figure 4, shrinkage and expansion and temporal acceleration and
arrow vii). deceleration defined by Rose et al. (2017), our hypothesis
on alternating ecosystem function scales along the aridity
In our conceptual scheme, the three processes are gradient driven by hydro-climatic extremes can be formu-
linked. The transformation of rainfall pulses to runoff lated as follows:
pulses is regulated by geodiversity and determines the
length of water flow pathways along the riverbed net- 1. In water-limited ecosystems, climate change toward
work (connectivity). Connectivity determines the distri- hydro-climatic extremes will rescale the dominant
bution of runoff from its source to its destination, where hydro-ecological processes of pulse–reserve, source–sink,
it infiltrates into the riverbed (sink). We propose that and connectivity from arid and semiarid ecosystems to
along the aridity gradient from semiarid, arid, to HA, the HAE along the rainfall gradient.
F I G U R E 4 Possible research topics that emerge from the hyper-arid ecosystem (HAE) paradigm. Water appears in blue, plants in
green, geodiversity in orange, and soil in brown. EE, ecosystem engineers; SOM, soil organic matter.
ECOSPHERE 13 of 16
2. Expansion of the spatial dimensions relates to the (Weiskopf et al., 2020). The factors that limit our ability
abiotic engine processes. The transformation of rain- to forecast ecosystem responses to climate change are the
fall pulses to runoff occurs on a small spatial scale complex interactions among vegetation changes, hydro-
(between patches) in more humid drylands and on logical responses, and the multiscale nature of dominant
larger spatial scale (watershed) as aridity increases. dryland processes. Due to the expected multiple and
3. Hence, as climate changes drive ecosystems at the large-scale responses, as suggested by landscape compen-
wetter end of arid lands toward the drier end, the sation and the rescaling of processes hypotheses, it will
scale of the main processes of ecosystem function will likely be challenging to determine ecosystem reorganiza-
change. We expect that on the spatial scale, abiotic tion in response to hydro-climatic extremes only by
processes will expand. experimental manipulations and mathematical modeling.
Hence, we propose that studying HAEs may improve our
The rescaling of processes hypothesis proposes that ability to predict water-limited ecosystem shifts to a pos-
the effect of climate change toward an extreme hydrologi- sible future ecosystem state characterized by climatic
cal regime on ecosystems should be investigated as a pro- extremes.
cess rescaling phenomenon. Climate change induces a We suggest that studying an ecosystem that already
shift in plant cover and composition that cascades to a functions under hydro-climatic extremes provides unique
shift in the spatiotemporal scales of the dominant abiotic information and insights on ecosystem reorganization to
and biotic ecosystem processes. The rescaling hypothesis cope with climatic extremes. In this paper, we identified
emphasizes that a scale shift due to climate change differs two possibly related reorganization strategies: landscape
between abiotic and biotic processes that together drive compensation and process rescaling. The landscape com-
ecosystem dynamics. pensation strategy is related to resource redistribution in
Our rescaling hypothesis advocates a novel perspec- response to an increasingly extreme hydrological regime.
tive on how the spatial and temporal scales of processes However, compensation processes encompass a scale
and their interactions operate along an aridity gradient transition of source–sink relationships.
and provide a framework to understand the role of cli- Process rescaling provides a theoretical framework on
mate change in process rescaling. It stresses that climate how to assess climate-change-induced regime shifts.
change toward hydro-climatic extremes will modulate Using HAEs as a model emphasizes that it is essential to
not only the magnitude of the dominant processes but incorporate rescaling processes and cross-scale interac-
also their scale of operation. This indicates that drylands tions when projecting ecosystem responses to extreme
driven toward HAE by climate change should be studied hydro-climates. Furthermore, comparing the scales of
using a scientific framework of large-scale and cross-scale operation along the rainfall gradient in semiarid, arid,
case studies. The development of macro-system ecology, and HA aridity landscapes (Figure 1) suggests different
with an emphasis on studying interactions between bio- scaling roles for abiotic and biotic processes.
logical and geophysical components that interact with Our concept of arid ecosystems shifting to hyper-arid
one another at broader and finer scales (Heffernan functioning under future climate change may impact man-
et al., 2014; Rose et al., 2017), together with remote agement practices—degraded ecosystems may receive less
sensing methodologies, could provide the scientific tools conservation efforts as they are pessimistically considered
necessary to understand the rescaling of processes “beyond help.” However, by understanding that state shift
hypothesis. does not necessarily entail degradation, the conservation
potential of the new state may actually be increased.
To conclude, integrating HAEs as a tangible model for
C O N C L U S IO N S projecting ecosystem shifts under extreme hydro-climatic
regimes is valuable, but it raises two future challenges. First,
As a tangible model for a water-limited ecosystem’s the landscape compensation hypothesis must be empirically
response to extreme hydro-climatic change, HAEs pro- tested. Second, our study calls for the development of math-
vide an alternative scenario to desertification scenarios. ematical climate change models and experiments that
By relating ecosystem and landscape processes to the incorporate rescaling and cross-scale interactions of the
whole watershed, we may find that climate change dominant ecosystem processes.
toward extremeness does not lead to desertification, but
rather to a state shift in which rescaling of processes com- ACKNOWLEDGMENTS
pensates for ecosystem degradation. It remains difficult The study was funded by the Israel Science Foundation
to project large-scale ecosystem shifts in water-limited (ISF) grant number 964/14, the grant for Open Spaces for
systems in response to extreme hydro-climatic change Gvanim LTER, and by the Ministry of Science and
Technology. We wish to thank Michelle Finzi for her Stocker, D. Qin, G.-K. Plattner, M. Tignor, S. K. Allen,
editing assistance and all the field assistants: Ariel J. Boschung, A. Nauels, Y. Xia, V. Bex, and P. M. Midgley,
Meroz, Noam Ganz, Tzvia Efrati, and Sofia Galeano. We 1029–136. Cambridge: Cambridge University Press. https://
doi.org/10.1017/CBO9781107415324.024.
also wish to thank Noa Avriel-Avni and Shayli Dor Haim
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617–30. ecs2.4586

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Received: 28 March 2023 Accepted: 6 April 2023

Toward an extreme world: The hyper-arid ecosystem

Ecosphere. 2023;14:e4586. https://onlinelibrary.wiley.com/r/ecs2 1 of 16

INTRODUCTION worldwide (Martinez-Valderrama et al., 2020). In

ARID ECOSYS T E M Arid ecosystem degradation

hydro-climatic extremes on ecosystem are projected to Landscape compensation hypothesis

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