Pond Ecology and Conservation Research Priorities
Pond Ecology and Conservation Research Priorities
Citation: Hill, M. J., H. M. Greaves, C. D. Sayer, C. Hassall, M. Milin, V. S. Milner, L. Marazzi, R. Hall, L. R. Harper, I.
Thornhill, R. Walton, J. Biggs, N. Ewald, A. Law, N. Willby, J. C. White, R. A. Briers, K. L. Mathers, M. J. Jeffries, and P. J.
Wood. 2021. Pond ecology and conservation: research priorities and knowledge gaps. Ecosphere 12(12):e03853. 10.1002/
ecs2.3853
Abstract. Ponds are among the most biodiverse and ecologically important freshwater habitats globally
and may provide a significant opportunity to mitigate anthropogenic pressures and reverse the decline of
aquatic biodiversity. Ponds also provide important contributions to society through the provision of
ecosystem services. Despite the ecological and societal importance of ponds, freshwater research, policy,
and conservation have historically focused on larger water bodies, with significant gaps remaining in our
understanding and conservation of pond ecosystems. In May 2019, pond researchers and practitioners par-
ticipated in a workshop to tackle several pond ecology, conservation, and management issues. Nine
research themes and 30 research questions were identified during and following the workshop to address
knowledge gaps around: (1) pond habitat definition; (2) global and long-term data availability; (3) anthro-
pogenic stressors; (4) aquatic–terrestrial interactions; (5) succession and disturbance; (6) freshwater connec-
tivity; (7) pond monitoring and technological advances; (8) socio-economic factors; and (9) conservation,
management, and policy. Key areas for the future inclusion of ponds in environmental and conservation
policy were also discussed. Addressing gaps in our fundamental understanding of pond ecosystems will
facilitate more effective research-led conservation and management of pondscapes, their inclusion in envi-
ronmental policy, support the sustainability of ecosystem services, and help address many of the global
threats driving the decline in freshwater biodiversity.
Key words: aquatic–terrestrial linkages; biodiversity; connectivity; ecosystem services; management; policy; small
lentic water bodies.
Received 21 April 2021; revised 19 July 2021; accepted 22 July 2021. Corresponding Editor: Samantha K. Oliver.
Copyright: © 2021 The Authors. Ecosphere published by Wiley Periodicals LLC on behalf of The Ecological Society of
America. 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.
E-mail: [email protected]
RESEARCH PRIORITIES
Definition of pond habitats
Since the 19th century there have been numer-
ous attempts to define ponds (see Appendix 1 in
Biggs et al. 2005). The distinction between smal-
ler lakes and ponds has proved particularly diffi-
cult, as well as between ponds and virtually all
other small standing freshwaters into which they
merge (Biggs and Williams 2021), although sev-
eral factors can be used to distinguish larger
lakes from smaller lakes and ponds (e.g., littoral
zone size, catchment size, wind protection, and
environmental heterogeneity; Søndergaard et al.
Fig. 1. Total number of peer-reviewed publications 2005). Despite this, distinguishing between small
based on the search topic (a) “biodiversity” or (b) lakes and large ponds can still be problematic as
“conservation,” with “pond,” “stream,” “lake,” or they share many characteristics (Biggs et al.
“river” between 2000 and 2019, using the Scopus data- 2017). This discrepancy in definition is likely to
base. have implications for our understanding of eco-
logical processes in small water bodies and the
required to advance understanding of pond development of pond management and conser-
ecosystems; (2) develop and/or facilitate more vation strategies.
effective conservation and management strate- In operational terms, ponds have been defined
gies; and (3) establish the required evidence base based on their size as water bodies between 1 m2
to support the future inclusion of pond ecosys- and 2 ha in the UK (Williams et al. 2010a, b),
tems in wider environmental and conservation although the upper size limit used to define a
policy initiatives. Initially, attendees individually pond varies considerably among countries.
and independently listed the “key challenges” Values between 1 ha and 6 ha are often
and “research priorities” for pond ecosystems, employed (Ilg and Oertli 2017), and the Ramsar
based on theoretical (e.g., fundamental under- Convention on Wetlands proposed an area of
standing) and applied (e.g., management, conser- 8 ha to distinguish between ponds and lakes
vation, and wider societal functions) topics. (Ramsar Convention Secretariat 2013). To
Fig. 2. The nine priority research themes and their contribution to the conservation and management of pond
ecosystems.
provide a consistent, scientifically derived defini- and temporary pond types that exist internation-
tion, research is required to examine the signifi- ally, all of which demonstrate significant variabil-
cance of depth, wind action, nutrient transport, ity in environmental conditions and origin (Biggs
and light penetration across a range of water et al. 2017). Given the complexity of this task,
body sizes, as these factors may have a large simple size-based definitions will probably con-
influence on biological communities. Recent tinue to play an important role in practical identi-
research has found that small water bodies may fication and management of small standing
experience strong diurnal stratification and mix- waters. But clearly an overarching, process-based
ing (due to convection), and seasonal stratifica- definition of a pond is required to allow
tion (Sayer et al. 2013), a process which can researchers to undertake more targeted and com-
influence pond environmental conditions, pat- parable research and for practitioners to develop
terns in freshwater biodiversity, and the perfor- effective conservation and management strate-
mance of organisms, particularly sessile taxa gies and policies.
(Andersen et al. 2017, Martinsen et al. 2019). A
threshold in water body size where there is a Key research question.—
change in mixing processes may provide a suit-
able characteristic to distinguish between ponds 1. Are there critical thresholds in physical, bio-
and lakes, although this currently remains inade- logical, and chemical processes, and ecosys-
quately quantified. The wide range of pond defi- tem function, which reflect changing lentic
nitions partly reflects the wide range of perennial water body size?
Table 1. Key research themes, research questions, and future research identified in this paper to advance under-
standing of pond ecology and increase opportunities for pond conservation.
Table 1. Continued.
Pond monitoring eDNA, remotely sensed and The use of remote sensing 1. Can molecular tools be used to assess the
and technological drone-based data have been and drone technologies to distribution of conservation priority and
advances poorly applied to pond identify the distribution invasive species as well as community
monitoring and multi-taxon and physicochemical diversity at the pondscape scale?
assessments. Advances in properties of pond 2. Are molecular tools and remote sensing
biostatistics may increase environments able to identify the effects of
effectiveness of pond Using multiple technologies anthropogenic stressors and climate
conservation (eDNA, UAV) effectively to change on ponds and improve
answer fundamental, management strategies to mitigate
multitrophic ecological stressors?
questions 3. Can UAV-based data collection record
physico-chemical and spatial
characteristics of ponds and pondscapes
more accurately than conventional data
collection?
4. How can the development of new
statistical analyses in biodiversity
assessment contribute to more effective
pond conservation planning?
Socio-economic Limited understanding of the Studies of the human 1. How do ponds contribute to human
factors personal, social, and dimensions of stakeholder physical and mental health and
educational contributions made engagement in pond wellbeing within urban and rural
by ponds, and governance creation and management populations?
conflicts that exist between are needed 2. What are the barriers (including social,
users of ponds. There remain Explore the relationships cultural, institutional, emotional,
gaps in our knowledge of the between ponds, well-being, communicative, and governance) in
contribution of ponds to access, and habitat quality stakeholder–pond conservation
human health and wellbeing Assess the complexities of interactions, and how might these be
engagement, the addressed?
educational value of ponds, 3. What are the short- and long-term effects
and the use of ponds in of environmental education for pond
fostering an environmental conservation?
conscience
Conservation, There is a need to increase Examine the mechanisms 1. Where are the most ecologically
management, fundamental understanding of that affect pond creation important ponds at national and
and policy pond ecosystems to inform and restoration success international scales?
practical pond conservation A need for studies to assess 2. How does pond creation and
and management. Ponds are medium to long-term pond management affect biotic communities at
largely excluded from conservation and local and landscape scales in different
environmental policy and management measures environmental settings?
legislation Identify opportunities for the 3. How can we best conserve pondscapes in
inclusion of ponds in anthropogenically dominated
environmental and landscapes?
conservation policy 4. What are the mechanisms required to
Quantify the distribution of better incorporate ponds into national
rare species in ponds and and international environmental policy
high-quality pond sites for and legislation?
conservation
water bodies; although this database does not stressors may include chemical (e.g., pollution),
currently exist. New large-scale collaborative biological (e.g., invasive species), and physical
studies will be critical to understand how global (e.g., infilling, management, and climatic warm-
pressures (e.g., land-use change and pollution) ing) pressures. In addition, stressors may origi-
are affecting pond diversity (e.g., total biodiver- nate from anthropocentric perceptions of ponds,
sity and rare species distributions) and ecosys- such as the high values placed upon neatness,
tem service provision (e.g., stormwater carefully managed plant communities (removal
collection, water purification, and human well- of “weedy” and aesthetically unpleasing spe-
being). cies), and large areas of open water, perceptions
A further scaling issue that inhibits conserva- which often do not reflect the natural functioning
tion efforts and the development of effective pol- of ponds (Nassauer 2004). Many stressors remain
icy for ponds is the lack of long-term datasets. poorly understood, and in this section, we focus
With a few exceptions (see Jeffries 2011, Williams on the impacts of pollution, non-native species,
et al. 2020), most pond research has been domi- and climate change on pond communities, and
nated by studies covering a single season or year identify knowledge gaps and key areas for future
(Hill et al. 2016), with studies covering time- research.
scales longer than 10 yr largely missing. Conse- Pollution.—One of the most pervasive threats
quently, there are significant knowledge gaps to pond communities is pollution. Nutrient
regarding the long-term dynamics and patterns enrichment, which is known to decrease species
in pond environments, biological diversity, and diversity at a landscape-scale (Rosset et al. 2014),
functioning. As for larger lentic water bodies, is one of the better understood threats to pond
paleoecological approaches could provide critical communities in agricultural and heavily popu-
information for inferring long-term (c.100s–1000s lated landscapes. In urban landscapes, runoff
years) changes in pond ecology in response to from impermeable surfaces can result in an
environmental change and, in turn, inform increase in heavy metals, nutrients, road salts,
restoration strategies (Walton et al. 2021). This is and chloride in ponds (Moss 2017). However,
of particular importance for demonstrating the many natural and anthropogenic ponds in urban
scope for returning communities to those landscapes are designed and managed to hold
recorded prior to major human impacts. As well stormwater runoff and are thus intended to be
as encouraging long-term data collection, the polluted as a means to protect water quality in
comparison of contemporary samples with older the catchment downstream (Gold et al. 2017). As
records (Hassall et al. 2012) and mesocosm such, there is a paradox in urban landscapes,
experimentation may elucidate long-term pro- where ponds have high biodiversity value but
cesses (Yvon-Durocher et al. 2017). are being designed to hold pollutants to improve
downstream water quality of more highly val-
Key research questions.— ued (but not actually more biodiversity rich)
freshwater habitats, such as larger lakes and riv-
1. What are the abiotic and biotic drivers of ers. Given urban areas are predicted to increase
pond environmental and biological patterns in size across the globe (Seto et al. 2012), more
at different spatial and temporal scales? ponds are likely to experience pollution, but the
2. How do biological communities within impacts on urban pond biodiversity are not fully
ponds respond to global environmental understood (Hintz and Relyea 2019). Higher
change along spatial and temporal gradi- numbers of polluted ponds also pose potential
ents? risks from increasing the number of water bodies
which are producing large quantities of climate
heating gases (Peacock et al. 2019, Rosentreter
Anthropogenic stressors et al. 2021).
Ponds are increasingly being subjected to mul- The process of suburbanization also poses a
tiple anthropogenic stressors which can affect the major threat to freshwater biodiversity (Van
diversity and resilience of their communities Acker et al. 2019). Pond habitats frequently sur-
(Ryan et al. 2014). The cocktail of anthropogenic vive the conversion from natural to suburban
land cover, but they are often subject to wastewa- native communities, not all introductions of non-
ter runoff (chemical loading), physical changes native taxa are negative. For example, in Oregon,
(e.g., decreased canopy cover and increased tem- USA, the non-native Reed Canary grass (Phalaris
perature), and isolation which can have large arundinacea) has been beneficial to breeding
effects on food web dynamics, species health, amphibians in many ponds, providing suitable
and taxa richness (Homan et al. 2004, Holgerson sites for oviposition, increasing tadpole survival,
et al. 2018, Van Acker et al. 2019). Pesticide pol- and the abundance of adult amphibians (Holzer
lution in freshwaters also represents a long-term and Lawler 2015) and in Washington State, USA,
persistent issue (Ito et al. 2020), yet quantifying Holgerson et al. (2019) found that non-native
concentrations remains challenging due to high bullfrogs (Lithobates catesbeianus) did not impact
spatiotemporal variability, limitations of instru- pond amphibian occupancy (but non-native fish
ment availability, and difficultly in identifying did). Should ponds be less widely affected than
new/emerging chemicals (Lorenz et al. 2017). In other freshwaters they may represent refugia for
particular, neonicotinoid pesticides (Raby et al. some taxa otherwise threatened by non-native
2018) are contaminants of freshwaters which invasive species in shallow lakes and rivers
have been given little attention in pond environ- (Sayer et al. 2011). Historically, the effects of non-
ments thus far. Moreover, despite a large body of native species on trophic interactions within
literature in lotic systems (Calabrese et al. 2020), ponds have been poorly studied compared to
there has been little research examining interac- their lotic counterparts, and as such, further con-
tive effects of multiple stressors on pond commu- sideration is required to identify the implications
nities, which could be additive, subtractive, of predator–prey interactions between non-
antagonistic, or synergistic (Hassall 2014). native and native species, as well as how non-
Non-native species.—Non-native species are one native species may be driving shifts in
of the greatest threats to many freshwater sys- pond ecosystem functioning. Research is clearly
tems, but are probably less common in ponds required to examine the full range of ecological
than other highly connected freshwater environ- effects (negative, positive, and negligible) that
ments associated with their hydrological isola- non-native taxa may have on native pond com-
tion (Williams et al. 2010a, b), although ponds in munities, and how they can be managed to facili-
urban areas are highly vulnerable to intentional tate conservation of native species.
introductions (Patoka et al. 2017). Climate change.—As small but abundant habi-
Where non-native species become highly inva- tats, ponds are sensitive to the effects of environ-
sive after successfully colonizing and establish- mental change and are therefore essential to
ing in ponds, impacts are invariably detrimental. consider within climate feedbacks. Agricultural
For example, the effects of New Zealand Pigmy- impoundment ponds are among the most signifi-
wort (Crassula helmsii) on pond communities in cant habitats for carbon sequestration due to
Europe are widely acknowledged (e.g., loss of their high nutrient loads and associated high pri-
plant biomass and abundance, plant germination mary productivity (Downing et al. 2008). Even
suppression; Langdon et al. 2004; Smith and accounting for their relatively small size, research
Buckley 2020), and the introduction of the Top- has demonstrated that carbon burial rates in
mouth Gudgeon (Pseudorasbora parva) into Euro- small ponds are at least equivalent to woodland
pean pond habitats has been shown to shift the or grassland habitats (Taylor et al. 2019). How-
trophic position of fishes (Britton et al. 2010). ever, other studies have suggested that small
However, there are a wide range of non-native water bodies can switch between carbon sources
invasive species that are expanding their habitat and sinks, while boreal ponds and small agricul-
range and of which we know little about their tural reservoirs may even act as significant
impacts for pond ecosystem functioning and bio- sources of greenhouse gas emissions (Holgerson
diversity (e.g., Dikerogammarus villosus and Paci- and Raymond 2016; Webb et al. 2019). Given
fastacus leniusculus). recent drives toward afforestation for climate
While it is clear that the establishment of non- mitigation (Brown 2020), determining the net
native invasive species can be detrimental to value of ponds within the carbon cycle may
reveal their potential, helping diversify climate The existing focus has largely been on birds (Rad-
mitigation beyond forestry alone. chuk et al. 2019), and changes in synchrony of
Understanding the different dimensions of cli- interacting taxa have mostly been studied in lar-
mate change—gradual warming, heat waves, ger water bodies (Winder and Schindler 2004).
droughts, and floods—and how they interact and Given the importance of links between aquatic
affect pond communities is vital to the future con- and terrestrial food webs (Stenroth et al. 2015),
servation of pond environments. Extreme climatic more research examining the effect of climate
events may result in homogenized pond communi- change on aquatic–terrestrial networks and the
ties, reducing the biodiversity value of pondscapes flow of nutrients and services between these habi-
(Bertoncin, et al. 2019). Drought is likely to increase tats is required (Soininen et al. 2015).
the proportion of ponds that experience hydrologi-
cal intermittency resulting in changing community Key research questions.—
structures of existing temporary ponds, extirpation
of species unable to adapt to increasing ephemeral- 1. What are the interactive effects of multiple
ity (Ryan et al. 2014, Abney et al. 2019), and poten- stressors on pond ecosystem resilience?
tially influence greenhouse gas emissions from 2. How do pollutants impact pond ecological
ponds (Holgerson and Raymond 2016). These cli- functioning and resilience?
mate shifts typically favor freshwater communities 3. How widely are ponds affected by non-
from warmer regions that can take advantage of native species and what are the dynamics of
higher temperatures to circumvent the increasingly their spread?
ephemeral nature of the ponds. However, a better 4. How do non-native species introductions
understanding of the interaction between short- affect trophic interactions in pond ecosys-
and long-term climate effects on ponds to predict tems?
future effects is required. 5. What are the potential ecological implica-
Climate change has been shown to interact tions for pond communities associated with
with other stressors, for example by exacerbating different climate change scenarios?
the effects of pesticides (Janssens and Stoks 2013) 6. How will climate change affect the interac-
and sedimentation (Piggott et al. 2012) due to cli- tions between aquatic and terrestrial food
matic changes to rainfall and runoff regimes webs?
altering the delivery of fine sediment and its
associated contaminants. In contrast, some stud-
ies suggest that climate change may be antago- Aquatic–terrestrial interactions
nistic to biological invasions, where non-native The importance of freshwater habitats is not
species increase their functional capacity to the limited to the water body itself. Interactions
same level as native species under warming between aquatic and terrestrial ecosystems can be
(Kenna et al. 2017), but climate change has also significant, demonstrated by the importance of ter-
been found to intensify the effects of non-native restrial vegetation shading in structuring freshwa-
introductions (Bellard et al. 2012). While some ter invertebrate assemblages (Suh and Samways
progress has been made in understanding how 2005), the negative effects of land-use change on
co-occurring stressors interact (if at all), a syn- terrestrial insectivores reliant on aquatic insects,
thetic approach to gathering evidence across spa- and the effect of predation (e.g., dragonflies prey-
tial and temporal scales could yield further ing on bees) on aquatic–terrestrial interactions
insights across ponds and the wider freshwater (Knight et al. 2005, Stenroth et al. 2015). There is a
network and inform new research. growing recognition of the importance of interac-
Climate change also potentially mediates tions between ponds and terrestrial species, espe-
interactions among species through driving cially farmland birds (Lewis-Phillips et al. 2020)
spatio-temporal patterns of occurrence. While and insect pollinators (Stewart et al. 2017). The
phenological change has been described for some reliance of terrestrial organisms on ponds may be
freshwater taxa (Hassall 2015), the ecological and largely due to trophic interactions that bridge the
fitness implications of phenological changes in aquatic–terrestrial divide. Farmland ponds
pond communities have received little attention. managed by scrub and sediment removal (to
succession, which may enhance both beta and and genetic resources within and between habi-
gamma diversity (Hill et al. 2017). In contrast, in tats in a landscape or more widely. Published
landscapes where natural disturbances have been studies examining connectivity have accelerated
reduced, uninterrupted succession can result in greatly during the 21st century alongside the
areas entirely dominated by late-succession ponds, growing debate on the relative roles of dispersal
thus reducing biodiversity (Sayer et al. 2012). The limitation and local species sorting across all
influence of succession and disturbance regimes ecosystem types (Heino et al. 2017). A growing
(natural and anthropogenic) on landscape-scale number of studies have reported the importance
species distribution and diversity patterns remains of spatial factors (Juracka et al. 2019), or biologi-
poorly understood, in part due to a lack of long- cal traits linked to dispersal ability (De Bie et al.
term monitoring, but could provide critical infor- 2012) in determining assemblage structure, indi-
mation for the development of landscape-scale cating that dispersal limitation is likely to be an
conservation and management strategies. important influence on pond communities. Con-
Many pond species possess life cycles and adap- trasting active and passive dispersers, both
tive strategies enabling them to persist across the within and across taxon groups, has confirmed
landscape in the face of disturbance, such as dor- expectations that dispersal limitation among
mant and resistant propagules of aquatic inverte- ponds is stronger among less mobile taxa (De Bie
brates and plants (Alderton et al. 2017; Williams et al. 2012; Hill et al. 2017a). The transferability
et al. 1997), and relatively long lifespans for of these patterns across a range of landscape
amphibians and some invertebrates. Efficient dis- types and pond characteristics now needs to be
persal mechanisms by many invertebrates allow determined as most studies on connectivity in
rapid recolonization when favorable conditions freshwater systems focus on longitudinal connec-
return including endozoochary (Kleyheeg and tivity in rivers and tend to examine freshwater
van Leeuwen 2015), aerial dispersal (Bilton et al. habitats in isolation. In the case of ponds, two-
2001), and “hitch-hiking” (Okamura et al. 2019). way connectivity both across the aquatic–terres-
Research focused on the life-history strategies that trial interface and with other freshwater habitats
enable species and communities to persist after a in the wider landscape are likely to be especially
disturbance will be especially valuable to conser- important.
vation managers as manipulating and facilitating Connectivity is complex to quantify and is
disturbance via active management or passively typically estimated using simple indicators,
by rewilding may be key to the survival and pro- including Euclidean distances to similar habitats
tection of many pond species. (Juracka et al. 2019), hydrological pathway
lengths or percentage of surrounding freshwater
Key research questions.— with a buffer zone (Law et al. 2019), waterbird
migration flyways (Viana, et al. 2016), and
1. What role do natural disturbances play human population densities or proximity to
in structuring aquatic and terrestrial biodi- recreational facilities (Chapman et al. 2020).
versity in pond habitats across landscapes Measures of geneflow or similarity in assem-
minimally impacted by human influences? blage composition are often used to infer real-
2. What are the best management strategies ized connectivity (Bilton et al. 2001). Metrics
to reduce detrimental disturbances and based on the structural properties of spatial net-
increase positive disturbance in anthropogeni- works (e.g., centrality and percolation thresh-
cally dominated landscapes? olds) or species-specific dispersal distances and
3. What are the principal trajectories, time- costs to crossing different habitats have also
scales, and outcomes of pond succession been suggested as possible solutions (Thornhill
across different pond types? et al. 2018; Hunter-Ayad and Hassall 2020).
Rapid increases in the quality and availability of
national biological recording datasets are
Freshwater connectivity improving understanding of the role of connec-
Connectivity is important because it facilitates tivity on the large-scale distribution of freshwa-
the movement of energy, materials, organisms, ter taxa. However, among ponds, the lack of
high-resolution mapping may confound such 4. What is the role of spatial processes in
attempts. It is also important to acknowledge assemblage structure and does pond context
that connectivity is not a static property and var- or landscape create regional differences?
ies temporally, for example, with hydroperiod or
hydrological events, or seasonality linked to
water bird migration, and can be fundamentally Pond monitoring and technological advances
altered by anthropogenic activities or ecosystem Recent developments in pond monitoring
engineers (Bilton et al. 2001). Moreover, rare or techniques and biostatistics can advance our
long-distance colonization events may be impor- understanding of the ecology and conservation
tant for connectivity (Jordano 2017), but accu- of pondscapes. There are several challenges fac-
rately modeling or predicting these events will ing pond monitoring including (1) often being
be complex when related to local physicochemi- located difficult to access and remote landscape
cal variables. Freshwater habitats are not dis- settings, (2) identifying representative sites due
crete entities but often exist in networks (e.g., an to their high abundance, (3) their high environ-
interconnected system of ponds, rivers, streams, mental heterogeneity, resulting in multiple
and lakes; Sayer 2014), and ubiquitous species ponds needing to be monitored to capture abi-
are recorded across these freshwater habitats otic and biotic diversity, and (4) the availability
(Davies et al. 2008). As such, it will be mislead- of taxonomic specialists to identify the highly
ing to study ponds in isolation, and more holis- diverse floral and faunal taxa recorded within
tic research is required to understand the ponds. However, the utilization of new tech-
ecological processes operating across a range of nologies may help to overcome some of these
connected freshwater habitats (the “water- challenges. In this section, we first outline the
scape”), which will likely provide more accurate contribution that molecular tools have made to
information for future landscape-scale conserva- pond monitoring (Biggs et al. 2015; Deiner
tion initiatives (Heino et al. 2021). et al. 2017) and then discuss the opportunities
Management strategies to increase habitat that recent technological advances in remote
patch connectedness is a common response to sensing, unmanned aerial vehicles, and bio-
fragmentation; however, the relative isolation of statistics have provided for pond ecology and
some ponds may be an asset in insulating them conservation.
from common stressors such as diffuse nutrient Pond monitoring using molecular tools.—The
loading, pathogens or invasive species that are emergence of environmental DNA (eDNA) anal-
easily transmitted between more connected ysis has the potential to transform freshwater
water body types. Understanding the aspects of biodiversity assessment. eDNA is genetic mate-
connectivity that confer an advantage in terms of rial released by organisms into their environ-
enhanced resilience, and those where it repre- ment, which can be sampled and analyzed to
sents a risk in terms of accelerated transfer of target specific species or passively screen entire
stressors, is therefore critical in terms of future communities (Harper et al. 2019a). Targeted
restoration or the design of pondscapes and eDNA analysis can be used to assess the distribu-
wider waterscape. tion and range of threatened, rare, or non-native
pond species (Biggs et al. 2015; Mauvisseau et al.
Key research questions.— 2018), and estimate relative abundance or bio-
mass, and detection probability (Buxton, et al.
1. How does connectivity between pond habi- 2017). Similarly, eDNA metabarcoding could be
tats influence trophic interactions that employed to assess multi-species distribution,
bridge aquatic–terrestrial divides? reveal species interactions (Harper et al. 2019b),
2. How do species move between ponds and and characterize genetic diversity (Parsons et al.
other freshwaters and what are the domi- 2018), all of which are only beginning to be con-
nant mechanisms? sidered for pond ecosystems. Community DNA
3. How are pond networks (pondscapes) best is distinct from eDNA samples, being sourced
designed or managed to ensure that rarer from biological material such as invertebrate
and less mobile species benefit? blood meals (invertebrate-derived DNA: iDNA),
feces, and collected specimens (Deiner et al. research, using remote sensing, was able to deter-
2017). iDNA analysis uses DNA that was mine the number (>1000 ponds) and distribution
ingested by invertebrates, such as leeches, to of ponds across the Greater Kuala Lumpur
detect biodiversity within freshwater habitats region (˜2950 km2) in Malaysia and quantify par-
(Abrams et al. 2019). iDNA metabarcoding could ticular environmental conditions of each pond
identify vertebrate biodiversity and enable including the surface area, shape, connectivity,
multi-species occupancy modeling (Abrams and surrounding land use (Teo et al. 2021).
et al. 2019), while fecal metabarcoding could be Despite this, it remains a largely unused tool in
used to assess diets of threatened or invasive spe- pond research. However, given the spatial reso-
cies and construct pond food webs (Kaunisto lution of remotely sensed data, it is unclear how
et al. 2017). Although bulk tissue DNA and remote sensing could be used to record small
eDNA metabarcoding have been used for (<10 m2) or intermittent ponds (during the dry
macroinvertebrate assessment in other freshwa- phase), and those located under forest canopy
ter ecosystems (Elbrecht et al. 2017), these tech- (Gallant 2015, Kissel et al. 2020).
niques have rarely been applied to ponds. Using In lotic settings, the recent use of small
bulk tissue DNA and eDNA metabarcoding in unmanned aerial vehicles (UAV) and structure-
pond research may provide more holistic esti- from motion (SfM) photogrammetric processing
mates of alpha and beta-diversity (Harper et al. has provided a significant improvement in the
2020). Similarly, stable isotope analysis can com- objectivity, accuracy, and efficiency of physical
plement metabarcoding to determine trophic habitat data collection (Woodget et al. 2015). The
relationships among pond taxa (Compson et al. parallels between streams and ponds suggest
2019). These tools could more accurately quan- that similar advancements could be made for
tify target species distribution and ranges and surveying surface area (particularly of small
determine the interactive effects of anthro- ponds), substrate composition, aquatic and ripar-
pogenic stressors, such as invasive species on ian vegetation structure, habitat complexity, pol-
communities and food webs. lution events, and the spatial structure of ponds
Technological advances in remote sensing, and pondscapes. UAVs can generate fine spatial
unmanned aerial vehicles, and biostatistics.—Con- resolutions (<10 cm/pixel; Lucieer et al. 2014)
ventional data collection methods for some envi- and, as such, have the potential to significantly
ronmental variables within ponds (e.g., visual increase the accuracy and consistency of pond-
estimation of macrophyte coverage) are typically scape data, elucidating the processes impacting
subjective or at best semi-quantitative and/or their ecology and providing detailed site infor-
lack sufficient detail of wider environmental con- mation to underpin effective monitoring strate-
ditions. Remote sensing has been widely used in gies. However, no empirical studies have thus far
water quality assessment and resource manage- examined the use of UAVs to characterize pond-
ment within lakes and rivers (Gholizadeh et al. scapes.
2016). Remote sensing can accurately measure Recent developments in biostatistics may
certain catchment characteristics (e.g., land cover, also provide considerable advances to our
productivity), physical properties (e.g., surface understanding of pond ecology and their con-
area, turbidity), and biological properties (e.g., servation. Research on the drivers of species
aquatic macrophyte coverage) of freshwater richness and community composition among
environments across large spatiotemporal scales lentic systems has focused almost exclusively on
in a cost-effective and standardized manner local environmental and spatial factors and has
(Giardino et al. 2010). Remote sensing could pro- largely ignored the potential influence of biotic
vide an efficient means to collect large-scale envi- interactions (Heino et al. 2015). Recently, new
ronmental and spatial metadata (e.g., pond statistical analysis has enabled the influence of
numbers, connectivity, pond spatial structure, biotic interactions, environmental conditions,
and physical barriers) for pond research and and spatial factors to be considered together
assistance in the development of effective moni- providing a more realistic understanding of the
toring strategies, particularly for remote ponds factors that govern the spatial and temporal pat-
and pond networks (Rose et al. 2015). Recent terns in pond biodiversity (Garcıa-Giro n et al.
2020). Similarly, Local Contributions to Beta created for a range of purposes, including indus-
Diversity (LCBD; Legendre 2014) analyses goes trial processes such as mineral extraction, provi-
beyond traditional measures of beta-diversity sion of food and water, irrigation, watering
(a single measure of dissimilarly across the land- livestock, and as ornamental features (Gledhill
scape) and calculates the contribution to overall and James 2012). In much of the world, many of
beta-diversity by individual sites (provides a the historical purposes of ponds are now redun-
measure of ecological uniqueness for individual dant and today ponds are often managed as
sites; Heino and Gro €nroos 2017). LCBD may amenity features (e.g., angling) or have been
contribute to pond conservation by identifying abandoned. Angling ponds have been demon-
ponds with high numbers of unique species strated to support limited faunal diversity (Wood
(that may be missed by traditional conservation et al. 2001), but there remains a paucity of
practices that focus on taxonomic richness), research considering social elements associated
whose protection can increase the number of with angling ponds. Research is needed to assess
species that are conserved at a landscape-scale. the personal, social, and educational contribu-
Recent research by Hill et al. (2021) found 70– tions made by angling ponds and to better
97% of the regional species pool was protected understand the practical and emotional gover-
when ecologically unique (sites with high LCBD nance conflicts that exist between anglers and
values) and high taxonomic diversity sites (sites other users of ponds to increase opportunities for
with >50 taxa) were considered together com- sustainable management (Arlinghaus 2005).
pared to 54%–94% when only sites with high Given that the origin of many ponds is a by-
taxonomic diversity were considered. Applying product from industrial processes, their presence
these new statistical techniques to different eco- in rural areas is unwanted by some landowners
logical and geographical settings will facilitate a (Wood et al. 2003). As a result, there are still
greater understanding of pond ecology and numerous barriers to the creation of new ponds
more effective and targeted conservation and the management of existing ponds, despite
strategies. recent evidence of biodiversity gains (Williams
et al. 2020). Studies of the human dimensions (in-
Key research questions.— cluding social, cultural, institutional, emotional,
communicative, governance, and lifestyle toler-
1. Can molecular tools be used to assess the ance factors) of stakeholder engagement with
distribution of conservation priority and pond creation and management are currently lar-
invasive species as well as community gely absent but are required to ensure the success
diversity at the pondscape scale? of any local or landscape-scale pond initiatives.
2. Are molecular tools and remote sensing able There is emerging evidence of the importance
to identify the effects of anthropogenic stres- of blue space for the health and well-being of
sors and climate change on ponds and individuals by promoting psychological restora-
improve management strategies to mitigate tion and providing spaces for physical activity,
stressors? recreation, and social interaction (Gascon et al.
3. Can UAV-based data collection record 2017; Foley and Kistemann 2015). Blue space
physicochemical and spatial characteristics regeneration could also help foster a sense of
of ponds and pondscapes more accurately civic pride and ownership (Higgins et al. 2019).
than conventional data collection? However, existing research has primarily focused
4. How can the development of new statistical on coastal areas or lotic ecosystems, and there
analyses in biodiversity assessment con- remain considerable gaps in our knowledge of
tribute to more effective pond conservation the contribution of ponds to human health and
planning? well-being (Foley and Kistemann 2015). The
immersive benefits of ponds in terms of their
contribution to physical health, imaginative,
Socio-economic factors emotional, and therapeutic aspects, and the
Ponds form an intrinsic component of urban range of meanings for individuals and groups of
and rural landscapes and many were historically pond ecosystems all require a greater
understanding. In addition, research is needed to benefits has developed significantly (Thiere et al.
explore the relationships between ponds, well- 2009). In addition, there is a growing body of
being, access, and habitat quality, particularly in research and expertise centered on the successful
urban landscapes (Higgins et al. 2019). restoration and management of ponds in a variety
With increased urban living, the incorporation of landscape settings, in particular European
of ponds as blue spaces into the aesthetic design of farmland and semi-natural habitats (Sayer et al.
urban areas represents an opportunity to engage 2012; Sayer and Greaves 2020), Mediterranean
the public with freshwater ecosystems (de Bell coastal plains (Sebastian-Gonzalez and Green
et al. 2017). Online packs of pond dipping 2014), and the prairies of Canada and the United
materials and citizen science pond initiatives can States (Bortolotti et al. 2016). Pond creation and
reconnect individuals with nature, while engaging restoration studies are typically short-term in
non-professionals in scientific research and duration, and studies assessing the medium to
practical freshwater conservation (Dickinson et al. long-term success of measures alongside natural
2012). Ponds also provide opportunities for educa- dynamics are needed (Seabloom and van der Valk
tion as the presence of ponds within schools may 2003). These studies will ultimately determine the
provide an important resource for educational need (or not) for subsequent policy and manage-
study and can encourage an initial interaction and ment activities to maintain conservation benefits
familiarization with ponds and wildlife from an (Sayer et al. 2012). Further, research is also
early age (Braund 1997). Research that considers required on the mechanisms that affect pond cre-
the complexities of engagement, the educational ation and restoration success, covering key issues
value of ponds, barriers to environmental such as water quality, grazing regimes, hydrology,
education, and the use of pond ecosystems for the connectivity, and invasive species. In this respect,
development of an environmental conscience in studies that compare and combine pond restora-
individuals and society will be particularly benefi- tion with creation at the landscape-scale will be
cial in addressing freshwater environmental and important to inform pond conservation planning
ecological degradation and increase opportunities and prioritization.
for pond conservation. Policy on the conservation and management of
ponds has generally suffered from the assump-
Key research questions.— tion that small water bodies were not important
due to their size (Biggs and Williams 2021). This
1. How do ponds contribute to human physi- long-standing assumption, noted since the 1950s,
cal and mental health and well-being within has generally led to the absence of small water
urban and rural populations? bodies from environmental and conservation
2. What are the barriers (including social, cul- policy globally (Hill et al. 2018). For example, the
tural, institutional, emotional, communica- European Union’s Water Framework Directive,
tive, and governance) in stakeholder–pond although intended to protect “all” water bod-
conservation interactions, and how might ies specifically excludes those <50 ha from moni-
these be addressed? toring schemes, thereby excluding millions of
3. What are the short- and long-term effects of ponds (although some are specially selected for
environmental education for pond conserva- nature conservation under the EU Habitats
tion? Directive). Similarly, in North America and Asia,
ponds are generally not directly considered in
environmental policy or legislation despite the
Pond conservation, management, and policy establishment of the Clean Water Rule clarifying
While lake and riverine habitats have over- freshwater protection under the Clean Water Act
whelmingly dominated historic freshwater policy, in the United States (Department of Army, Corps
conservation, and management research, more of Engineers and US Environmental Protection
recently there has been a growing focus on ponds. Agency 2015) and various national-scale environ-
Research on strategies for pond creation, pond mental legislation across Asia (e.g., Environment
and pondscape design, and where best to locate Protection Act 1986 and the Wildlife Protection
new ponds to maximize aquatic conservation Act 1972 in India). However, the policy failure
relating to small water bodies is slowly being 4. What are the mechanisms required to bet-
reversed as awareness of the importance of these ter incorporate ponds into national and
water bodies appears to be increasing. For exam- international environmental policy and
ple, the Intergovernmental Science Policy Plat- legislation?
form on Biodiversity and Ecosystem Services
(IPBES) included temporary and perennial ponds CONCLUSION
in its classification of freshwater habitats in 2018
(IPBES 2018). Large knowledge gaps remain in our under-
The future inclusion of ponds in environmental standing of pond ecosystems. Yet, it is clear that
and conservation policy should focus on four ponds can benefit society and wildlife by provid-
areas: (1) stopping further deterioration, especially ing habitats that support significant freshwater
of the most vulnerable high-quality sites. and terrestrial biodiversity across a range of land-
Although evidence is scarce, the quality of the scapes, while also providing ecosystem services
most protected ponds continues to decline required by society. Although ponds have
(Williams 2019); (2) protecting species and com- received less research attention than other fresh-
munities that are a special feature of ponds. water habitats to date, there is an increasing com-
Increasing knowledge of pond communities and munity of researchers and practitioners interested
the distribution of freshwater species will refine in pond ecology and conservation and a rapidly
our understanding of the role of ponds play in increasing awareness of the importance of ponds
protecting endangered freshwater species and by society. This paper has highlighted some of the
inform policy; (3) encouraging the creation of new major themes and provides key questions for
clean water ponds (clean water ponds reflect future pond research, which aim to address exist-
pond water chemistry and biology that is typical ing knowledge gaps and increase fundamental
for a given area in the absence of human activity; and practical understanding of pond ecology.
(Williams et al. 2010a, b)) and ensuring the effec- However, to continue to progress interest, knowl-
tive restoration and management of existing edge and awareness of pond ecosystems, interna-
ponds to maximize opportunities for maintaining tional collaboration, and commitment among
and increasing species diversity at the landscape- researchers and end-users is required. A better
scale; and (4) creating/restoring ponds for ecosys- understanding of pond ecosystems will benefit
tem service measures for land and water manage- society and wildlife by enabling more effective
ment. Emerging evidence suggests that this could research-led conservation and management of
be one of the most effective means of enhancing pondscapes, facilitating their inclusion in environ-
landscape-scale freshwater biodiversity, but to mental policy more clearly while simultaneously
maintain their long-term contribution, manage- addressing many of the threats driving the decline
ment interventions may be required (Williams in global freshwater biodiversity.
et al. 2020). However, it is important to recognize
that ponds for ecosystem service purposes such as ACKNOWLEDGMENTS
sediment retention or to attenuate flood flows
may not always provide suitable conditions for The authors gratefully acknowledge the support of
biodiversity provision (Williams et al. 2020). The British Ecological Society for funding and sup-
porting the workshop. MJH would also like to thank
Key research questions.— to the University of Huddersfield for supporting the
workshop. The authors would like to thank the
reviewers of the draft manuscript for their valuable
1. Where are the most ecologically important comments and input which have greatly improved
ponds at national and international scales? this publication.
2. How does pond creation and management
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