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Plants in aquatic ecosystems: current trends and future directions

Article in Hydrobiologia · May 2018


DOI: 10.1007/s10750-017-3190-7

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Hydrobiologia (2018) 812:1–11
https://doi.org/10.1007/s10750-017-3190-7

PLANTS IN AQUATIC SYSTEMS Review Paper

Plants in aquatic ecosystems: current trends and future


directions
Matthew T. O’Hare . Francisca C. Aguiar . Takashi Asaeda . Elisabeth S. Bakker .
Patricia A. Chambers . John S. Clayton . Arnaud Elger . Teresa M. Ferreira .
Elisabeth M. Gross . Iain D. M. Gunn . Angela M. Gurnell . Seppo Hellsten .
Deborah E. Hofstra . Wei Li . Silvia Mohr . Sara Puijalon . Krzysztof Szoszkiewicz .
Nigel J. Willby . Kevin A. Wood

Received: 20 October 2016 / Revised: 5 April 2017 / Accepted: 8 April 2017 / Published online: 20 April 2017
Ó Springer International Publishing Switzerland 2017

Abstract Aquatic plants fulfil a wide range of this paper, we present an overview of current trends and
ecological roles, and make a substantial contribution to future directions in aquatic plant research in the early
the structure, function and service provision of aquatic twenty first century. Our understanding of aquatic plant
ecosystems. Given their well-documented importance in biology, the range of scientific issues being addressed
aquatic ecosystems, research into aquatic plants contin- and the range of techniques available to researchers have
ues to blossom. The 14th International Symposium on all arguably never been greater; however, substantial
Aquatic Plants, held in Edinburgh in September 2015, challenges exist to the conservation and management of
brought together 120 delegates from 28 countries and six both aquatic plants and the ecosystems in which they are
continents. This special issue of Hydrobiologia includes found. The range of countries and continents represented
a select number of papers on aspects of aquatic plants, by conference delegates and authors of papers in the
covering a wide range of species, systems and issues. In special issue illustrates the global relevance of aquatic
plant research in the early twenty first century but also
the many challenges that this burgeoning scientific
Guest editors: M. T. O’Hare, F. C. Aguiar, E. S. Bakker & discipline must address.
K. A. Wood / Plants in Aquatic Systems – a 21st Century
Perspective

M. T. O’Hare  I. D. M. Gunn (&) P. A. Chambers


Centre for Ecology & Hydrology, Bush Estate, Environment and Climate Change Canada, 867 Lakeshore
Penicuik EH27 0QB, Scotland, UK Road, Burlington, ON L7R 4A6, Canada
e-mail: [email protected]
J. S. Clayton  D. E. Hofstra
F. C. Aguiar  T. M. Ferreira National Institute of Water & Atmospheric Research,
Forest Research Centre, School of Agronomy, University Gate 10 Silverdale Road, Hillcrest, Hamilton, New
of Lisbon, Tapada da Ajuda, 1349-017 Lisbon, Portugal Zealand

T. Asaeda A. Elger
Department of Environmental Science, Saitama EcoLab, Université de Toulouse, CNRS, INPT, UPS,
University, 255 Shimo-okubo, Sakura, Saitama 338-8570, Toulouse, France
Japan
E. M. Gross
E. S. Bakker Université de Lorraine, LIEC UMR 7360 CNRS, Rue
Department of Aquatic Ecology, Netherlands Institute of Général Delestraint, Bâtiment IBISE, 57070 Metz,
Ecology (NIOO-KNAW), Droevendaalsesteeg 10, Lorraine, France
6708 PB Wageningen, The Netherlands

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2 Hydrobiologia (2018) 812:1–11

Keywords Angiosperms  Botany  Herbivory  As a consequence of the growth of aquatic plant


Limnology  Macrophytes  Submerged aquatic research over recent decades, our views on many key
vegetation  Trends in research topics in aquatic botany have shifted (Vermaat &
Gross, 2016; Phillips et al., 2016), and so this
introduction to the special issue on plants in aquatic
Introduction
systems presents an overview of current trends and
future directions in aquatic plant research in the early
In the early twenty first century, researchers recognize
twenty first century. It is a time of newly emerging
the fundamental importance of plants that grow in and
fields and the advancement of long-established
around water to the structure, functioning and service
research areas. The research is set against a back-
provision of aquatic ecosystems (Chambers et al., 2008).
ground of rapid environmental change that has been
Aquatic plants interact with and influence the hydrolog-
on-going for at least the last two centuries. The pace of
ical, geomorphological and physico-chemical environ-
change is unremitting with demands on water
ments, and interact with a wide range of other organisms,
resources set to increase globally (Dudgeon et al.,
from microbes to vertebrates, for example, by providing
2006; Vörösmarty et al., 2010). In the future, the
habitat and food (Brix, 1997; Engelhardt & Ritchie,
response of aquatic plant dominated systems (e.g.
2001; Wood et al., 2017a). The current interest contrasts
shallow lakes and seagrass beds) to global temperature
with the views of earlier limnologists a century ago who
increases and climatic extremes may well become a
considered aquatic plants to be largely unimportant in
focus of research efforts. The in-depth understanding
aquatic ecosystems, for example, Shelford (1918) argued
aquatic botanists possess can only contribute posi-
that ‘‘One could probably remove all the larger plants
tively to our understanding of how climate change will
and substitute glass structures of the same form and
perturb aquatic systems. Trends in aquatic plant
surface texture without greatly affecting the immediate
research reflect the environmental pressures on fresh-
food relations’’. Over the past century the study of
water systems, legislative drivers, technical advances
aquatic plants has expanded considerably, because of the
and developments in the wider fields of ecology and
increased recognition of their importance in fundamental
environmental management.
system processes. Specialist journals have been estab-
Both national and international legislative drivers
lished, such as Aquatic Botany (Den Hartog, 1975) and
have had a clear impact on the direction of aquatic
Journal of Aquatic Plant Management, as well as
plant research. In Europe, the implementation of the
conferences devoted to aquatic plant research.

A. M. Gurnell K. Szoszkiewicz
School of Geography, Queen Mary University of London, Faculty of Environmental Engineering and Spatial
London E1 4NS, UK Management, Poznan University of Life Sciences, Wojska
Polskiego 28, 60-637 Poznan, Poland
S. Hellsten
Finnish Environment Institute SYKE, Freshwater Centre, N. J. Willby
Paavo Havaksen tie 3, 90570 Oulu, Finland Biological & Environmental Science, Faculty of Natural
Science, University of Stirling, Stirling FK9 4LA, UK
W. Li
Key Laboratory of Aquatic Botany and Watershed K. A. Wood
Ecology, Wuhan Botanical Garden, Chinese Academy of Wildfowl & Wetlands Trust, Slimbridge,
Sciences, Wuhan 430074, China Gloucestershire GL2 7BT, UK
S. Mohr
Umweltbundesamt, Schichauweg 58, 12307 Berlin,
Germany

S. Puijalon
Université de Lyon, Université Claude Bernard Lyon 1,
CNRS, ENTPE, UMR 5023 LEHNA,
69622 Villeurbanne, France

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Hydrobiologia (2018) 812:1–11 3

European Union (EU) Water Framework Directive expansion and the intensification of resource use have
(WFD) (European Commission, 2000) led to a mas- increased demands on water supplies and other natural
sive surge in research on monitoring methods, their resources. A striking example is the numerous hydro-
inter-calibration and the analysis of the resulting large power plants constructed in South America that have
multi-site datasets (Hering et al., 2010). As the WFD caused profound changes in aquatic ecosystems,
implementation moves into its second phase, we now including macrophyte community composition and
see a shift in focus to restoration projects. We have patterns of colonization (e.g. Martins et al., 2013). Yet
learnt much during the implementation of this direc- at the same time as these enormous ecological
tive and it is likely that we will see knowledge transfer changes, many developing countries also face reduced
from European scientists to colleagues in countries research funding and weakened environmental legis-
across the globe. We see many countries in Asia and lation, which limits conservation efforts (Azevedo-
Africa now adopting reference-based systems for Santos et al., 2017). The conference attracted dele-
freshwater assessment (e.g. Kennedy et al., 2016). gates from many developing countries and we would
The global financial crash in 2008 has exacerbated strongly encourage their future participation.
the difficulty in obtaining research funding in many While global financial trends and legislative drivers
countries, and immediate output in terms of results have affected the direction of research, technical
reigns over the long-term understanding of complex advances in survey and analytical methodologies have
interactions and processes (Krugmann, 2012). In also been influential. Some established techniques
Europe, we have also seen a reduction in core funding have become increasingly used in aquatic botany, for
for national research organizations and university example, molecular biology and stable isotope anal-
researchers who work on aquatic plant management ysis. Recent reductions in the cost of stable isotope
issues and there are concerns that there will be a slow analysis have facilitated their use. Developments in
erosion of the research base. The United Kingdom’s ecological modelling and computational biology have
decision in 2016 to leave the EU will likely have allowed aquatic plants to be incorporated into models
implications for site-level conservation of aquatic that can predict interactions between macrophytes and
plants under the EU Habitats Directive (Council of the other organisms (e.g. Wood et al., 2014; Stillman
European Communities, 1992), although it is not yet et al., 2015). The continued development of remote
clear what will replace the EU Directives in UK law. sensing, drone technology and the software to interpret
In the USA, the Department of Energy has been aerial photography, now allows new types of spatial
planning to increase hydropower output by retro- analysis. Moreover, the potential for drones to carry
fitting turbines to pre-existing dams that are currently Light Detection and Ranging (LIDAR) equipment
only used for flood control or water supply. While the could facilitate aquatic plant-sediment interaction
election in the USA of President Trump in 2016, who studies. The rise of ‘citizen science’ represents greater
is a climate change sceptic and pro-fossil fuel public participation in scientific research and has the
advocate, makes the implementation of this policy potential to aid data collection (McKinley et al., 2017).
much less certain, it is worth noting that it did have Similarly, the emergence of R (the free statistical
substantial cross-party support. If this work is under- software environment) has encouraged the develop-
taken, it could reduce the USA’s carbon production ment and sharing of new analytical techniques (R Core
and reduce its requirement to buy in fossil fuels from Development Team, 2016).
abroad, but careful assessment of downstream impacts Aquatic botanists work from an especially strong
on aquatic plants and other taxa will need to be position where the physiology of the plants is well
undertaken. In China, the current five-year plan, which described and there is a deep knowledge of the plants’
has significant green policies, has energized the roles in system function. Aquatic plants have many
environmental sector and led to substantial efforts to advantages over other aquatic biota as study organ-
exchange knowledge with western countries. We hope isms: they are sessile, they can be accurately mapped,
this exchange will lead to greater international rapidly surveyed and cultured easily in the laboratory,
collaboration between aquatic botanists in the future. and they are increasingly being used by a wide variety
In developing countries, there is a need too for the of researchers. Although, historically, there was an
services of aquatic botanists where rapid population assumption that publishing aquatic botany studies in

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4 Hydrobiologia (2018) 812:1–11

high impact journals was challenging, there is anec- represented a key session during the conference:
dotal evidence that this is no longer the case. (i) physical habitat interactions, (ii) riparian processes,
Against this background of environmental and (iii) ecological stoichiometry and nutrient cycling, (iv)
societal change, aquatic botanists met recently to take trophic interactions—focused on plant herbivore
stock of their discipline at the 14th International interactions, (v) community responses to environmen-
Symposium on Aquatic Plants, held in Edinburgh in tal change in space and time, (vi) aquatic plant
September 2015. The symposium series originally monitoring, (vii) ecotoxicology, (viii) restoration, (ix)
began as an aquatic weeds meeting, but over time the the future of invasive species management and
focus of the symposia changed as research and (x) fundamental science.
management interests altered. As our understanding
and appreciation of the different roles that macro-
phytes play has increased, so too have the breadth of Overview of current trends and future directions
topics addressed at the symposia. The conference in aquatic plant research
continues to attract delegates involved in the practical
management of aquatic systems and those working Physical habitat interactions and riparian processes
directly in research. The synopsis which follows is
based primarily on the conference output. The 14th The interactions between plants and water flow and
International Symposium was attended by 120 dele- sediments have been championed sporadically for
gates from 28 countries and six continents, and over forty years, but in the last decade work has
featured 79 oral presentations in addition to over 30 accelerated as the importance of the interactions for
poster presentations. Although the 2015 symposium ecology, hydrology and fluvial geomorphology was
and the 13 preceding symposia were held in Europe, fully realized. Plants influence physical processes:
henceforth, every second symposium will be held transport of solutes, sediment deposition/resuspen-
outside Europe to reflect the global nature of the sion, hydraulic conditions and light transmittance
subject and the attendees. Global regions often diverge (O’Hare, 2015; Klančnik et al., 2017). In turn, the
in approaches and attitudes towards macrophytes, for physical environment affects macrophytes. Its effects
instance, weed management with herbicides is well are induced by mean velocity, turbulence and water
accepted in the United States yet largely prohibited in level (O’Hare, 2015). Macrophytes can be affected at
Europe. Therefore, truly international conferences are scales, from individual plants to populations and
vital in order to provide opportunities for global communities. This is exemplified by plant growth
debates on such key issues. The next conference will which is known to be influenced from the microscale,
take place in February 2018 in New Zealand and it will for example, cell ultrastructure (Atapaththu et al.,
be jointly held with the Aquatic Plant Management 2015), to macroscale, for example, biomechanical
Society of North America. The conference will also be traits (Puijalon et al., 2011; Schoelynck et al., 2014).
supported by our colleagues from China, where there Current developments in our understanding of these
has been an upsurge in research interest in aquatic complex two-way interactions between aquatic vege-
plants in recent years. tation and physical factors are tightly linked to fluid
Traditionally, authors of conference presentations dynamics modelling (Marjoribanks et al., 2014; Ver-
elaborated their contributions as full papers published schoren et al., 2016).
in a special issue of Hydrobiologia (e.g. Caffrey et al., While aquatic botanists have tended to focus on
1996, 1999, 2006; Pieterse et al., 2010; Ferreira et al., aquatic macrophytes, geographers have been examin-
2014). Thus, in this special issue of Hydrobiologia, we ing both instream and riparian vegetation. An espe-
present a number of studies of aquatic plants that cially exciting development is the realization that
comprise the peer-reviewed proceedings of the 14th vegetation fringing a river’s edge has a substantial
International Symposium on Aquatic Plants. In the influence on fluvial geomorphological processes. In
remainder of this paper, we present an overview of effect, nearshore plants (emergent and submerged)
current trends and future directions in aquatic plant help engineer river form (Gurnell, 2014; Gurnell et al.,
research in the early twenty first century. We focus on 2016). This has significant practical implications as
the following key areas of study, each of which alterations to hydrology and fluvial geomorphology are

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Hydrobiologia (2018) 812:1–11 5

as widespread as nutrient pollution in Europe, effecting individuals to communities. Despite clear theories
approximately half of all water bodies (Kristensen, (Elser et al., 2000), elemental requirements and the
2012). We speculate that this reflects an unmeasured influence of environmental factors on nutrient uptake
but global trend as evidenced by the contributions from seem more complex for aquatic plant systems. At a
Africa and Asia to this session on impacts of flow global scale, silica is a nutrient which is in surprisingly
disturbance and regulation. Regulation by hydropower a short supply in marine environments requiring
dams influences the colonization rates of aquatic and frequent inputs from freshwater systems. The role of
riparian vegetation, with synergic impacts when rivers macrophytes and other primary producers in influenc-
are subjected to sediment removal or impaired by ing silica delivery is gaining increasing interest and its
storage reservoirs (Aguiar et al., 2016). Such distur- accumulation in macrophytes may be a functional trait
bances create ecosystems prone to alien plant inva- that enables them to adapt to environmental conditions
sions, and regulation alters the growth trajectories, (Schoelynck & Struyf, 2016). At local scales, macro-
composition and complexity of native communities phytes strongly influence their physico-chemical
(Bunn & Arthington, 2002). During the conference, the environment. Aquatic weed mats may constitute
concerning case of Podostemaceae in West-Africa important hotspots for greenhouse gas emissions in
(strictly aquatic angiosperms) was highlighted, where temperate shallow lakes, but wetland vegetation can
six species are critically endangered and four species also assist in nitrogen assimilation (Volkmann et al.,
have become extinct due to altered flows (personal 2016; Ribaudo et al., 2017). Yet, the relation between
communication). Such issues can be overcome: for environmental nutrient availability and macrophyte
example, implementing environmental flows that nutrient content is often less clear. For example,
inundate geomorphological structures and create slack research, presented during the conference, showed that
waters helped with the restoration of regulated rivers upland streams with proliferations of pond water-
by enhancing recruitment and colonization (Souter crowfoot (Ranunculus peltatus L.) tend to have a low
et al., 2014; Rivaes et al., 2015). While most research in N:P ratio at overall very high nitrogen and phosphorus
this field focuses on rivers, data from the UK and concentrations (personal communication). Although
Denmark indicate artificial water-level fluctuations in intra-specific C:N:P stoichiometry of submerged
lakes affect macrophytes (e.g. Baastrup-Spohr et al., macrophytes correlates to sediment and water nutrient
2015; May & Spears, 2012; Smith et al., 1987), and that availability, inorganic carbon availability may also
shoreweed (Littorella uniflora (L.) Asch.) has potential play a strong role in their nitrogen-based metabolism
as a model species in ecological studies of both lake (Hussner et al., 2016). Further research, presented
productivity and morphometry (e.g. Baastrup-Spohr during the conference, found that macrophyte tissue
et al., 2016; Robe & Griffiths, 2000). nutrient concentrations appear more closely related to
In due course, this field of research has the potential plant growth form than to phylogeny (personal
to produce novel tools for management, especially communication).
nature-based solutions to flooding, and fresh insights
into the ecology of aquatic plants. A research effort Trophic interactions: focused on plant herbivore
equivalent to that which elucidated the basic mecha- interactions
nisms of lake eutrophication (Vollenweider, 1968)
will likely be required to resolve these major research Since the seminal paper by Lodge (1991) on herbivory
questions. With this realization will come a far greater of aquatic plants, researchers have been devoting
appreciation of the role of both instream and riparian considerable attention to plant–herbivore interactions
vegetation in engineering physical habitats. Further in aquatic ecosystems. Now, in the early twenty first
collaborative research between geographers and ecol- century, it has now been demonstrated, unequivocally,
ogists will emerge. that herbivores can provide strong top-down regulation
of macrophyte beds (Bakker et al., 2016; Wood et al.,
Ecological stoichiometry and nutrient cycling 2017a). These top-down mechanisms can interact with
recovery from stress; for example, recovery of macro-
Ecological stoichiometry bridges ecology and ecosys- phyte beds after eutrophication attracts herbivorous
tem functions or processes at various levels, from water birds, but the colonization process can be

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hampered by strong vertebrate herbivory. In contrast, the importance of different aquatic habitats to a Scottish
smaller invertebrate grazers may assist recovery of regional flora confirmed the importance of riverine
eutrophic systems. They stimulate submerged macro- backwaters (Keruzore et al., 2013), a habitat that had
phyte growth and establishment by consuming peri- been previously neglected. The study illustrated an
phyton (instead of the tougher macrophytes) that would increasing awareness of spatial processes operating
otherwise reduce light availability for macrophytes beyond individual sites, and the associated issue of
(Bakker et al., 2016; Wood et al., 2017a). scale-dependent responses. Thus, for example, the
Recognizing the importance of herbivory opens effects of land use on macrophyte richness in lakes are
new research avenues by scaling up from macrophyte scale-dependent and are of greater importance at small
beds to aquatic ecosystem functioning, as herbivores spatial scales relative to the influence of hydrological
affect methane emission, carbon cycling and regime connectivity (O’Hare et al., 2012). Looking beyond the
shifts (Hidding et al., 2016). Furthermore, there is an immediate is one of the most powerful approaches of
urgent need to predict how global change will alter space and time analyses, and frequently produces
trophic interactions as a result of exotic species insightful findings. Not only do we see this in relation
invasions (Redekop et al., 2017), temperature rises to hydrological connectivity but also in legacy signals,
(Zhang et al., 2017) or changes in hydrological for example, the lakes of northwest Europe are geolog-
patterns (Wood et al., 2017b). Finally, current and ically young due to their glacial origins, with the signal
future conservation challenges lay in predicting and of glaciation still evident in the composition of their
managing the consequences of recovery of larger flora (Alahuhta et al., 2017).
vertebrate herbivores, through re-introductions such
as the Eurasian beaver (Castor fiber L.) in Europe (e.g. Aquatic plant monitoring
re-wilding), as well as by strong local herbivore
population increases in species such as mute swans Changes in the abundance or composition of an
(Cygnus olor Gmelin). aquatic plant community are often obvious signals of
alteration in the ecological condition of a lake or
Community responses to environmental change stream. In fact, a recent review of assessment methods
in space and time used to implement the EU Water Framework Directive
showed that the majority of methods are based on
The study of the responses of aquatic plant communities macroscopic plants (28% of all methods), followed by
to environmental change in space and time is both a benthic invertebrates (26%) (Birk et al., 2012).
mature field of research and one with critical new Moreover, unlike many other biological indicators,
questions being asked. Current research effort has seen a macrophytes are equally good at detecting eutrophi-
continued focus on the role of bottom-up regulation cation/organic pollution and hydrological/morpholog-
through environmental drivers (e.g. Fernández-Aláez ical changes (Birk et al., 2012). Historically, surveys
et al., 2017) and competitive processes between of abundance and composition were challenging in
macrophyte species (e.g. Gérard & Triest, 2017; Nunes terms of both field effort and taxonomic ability. As
& Camargo, 2017) in shaping aquatic plant community identified at this symposium, improved methods for
composition. Our understanding of how connectivity mapping abundance and composition of aquatic
can influence floodplain macrophyte populations has vegetation are now becoming available: high-resolu-
now matured to the point where scenario modelling is tion aerial images of lake and rivers taken with
feasible, for example, on the Murray-Darling system in unmanned aircraft systems permit identification, map-
Australia where species richness of floodplain plant ping and abundance estimates of non-submerged
communities can be predicted as a function of channel species while near-infrared-sensitive DSLR cameras
connectivity in the watershed (Campbell et al., 2014). can be used to map spatial distribution and depth of
Furthermore, recent studies of aquatic plant responses to submerged species (e.g. Visser et al., 2015).
floods in large floodplains have offered support for the Research is continuing to show that community
flood homogenization hypothesis (Thomaz et al., 2007). metrics (e.g. cover, diversity and richness) and species
Floodplain inundation has received less attention on frequency of occurrence are often related to water
smaller systems; however, comparative assessments of quality, lending support for the development of

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Hydrobiologia (2018) 812:1–11 7

macrophyte-based indices for classification of fresh indirect effects of chemicals on plant populations and
waters and brackish water ecosystems and seagrass communities. On plant-contaminant issues, the interac-
beds (Spears et al., 2016). Although many macrophyte tion between chemical contaminants and other stressors
indices are based only on hydrophytes due to their was evident; for example, evidence presented at the
dependency on the quality of the aquatic environment, conference showed that the stoichiometry (C:N:P) of
the importance of helophytes has been demonstrated Eurasian water milfoil (Myriophyllum spicatum L.) was
as indicators of the eutrophication process, for exam- not only influenced by light and nutrients, but also by
ple, in the bioassessment of lowland lakes (Kolada, herbicides and the metalloid arsenic (personal commu-
2016). Biochemical measurements may also provide a nication). Field monitoring and biomarker assays
new tool for bioassessment: for example, during the revealed a significant relationship between the decline
conference, evidence was presented that 15N and C:N of dwarf eelgrass (Zostera noltei Hornem.) in the
values from caged duck weed (Spirodela sp.) were Vaccarès lagoon in France and its exposure to chemical
found to relate to the proximity and timing of sewage contaminants including metals and pesticides (personal
manure or fertilizer inputs into rivers in South Africa communication).
(personal communication). Despite encouraging
advances in both methods for mapping aquatic veg- Restoration
etation and approaches for assessing water quality,
physical factors such as hydrological modifications to Management of aquatic macrophytes is an essential
water courses or inter-annual variation in water levels part of freshwater restoration projects (Phillips et al.,
can confound the relationship between macrophyte 2016). Macrophyte restoration can have multiple
occurrence and water quality, necessitating caution benefits, for example, supporting endangered water-
when deciding the status of a water body based on fowl and fish species or limiting the spread of invasive
limited (temporal or spatial) macrophyte data. species, such as Nuttall’s waterweed (Elodea nuttallii
(Planch.) H. St. John), in Europe. To successfully
Ecotoxicology restore macrophytes, consideration of the following
factors can be helpful: the genetic background of
The banning of herbicides for use in aquatic systems macrophyte population used, native seed bank viabil-
across the EU resulted in a shift in research away ity, control of herbivores and, in the case of eutrophic
from studies on the efficacious use and impacts of lakes, the use of geo-engineering tools which reduce
pesticides in controlling aquatic plants. A strong internal P loading, (Combroux et al., 2001; Guittonny-
research focus remains, however, on the effects of Philippe et al., 2015; Hussner et al., 2017). Restoration
pesticides and other pollutants derived from terres- science is still under development and new data are
trial systems on aquatic plants (Coutris et al., 2011; desirable; monitoring using macrophyte growth forms
King et al., 2016). can provide a cost-effective tool for evaluating the
This was the first time an ecotoxicology session was effect of individual restoration projects while long-
held at the conference and it focused on linking term records of macrophyte dynamics can provide
ecological studies with chemical risk assessment, with valuable information for assessment of broader, global
the overarching aims to make assessment methods more scale change (Ecke et al., 2016).
realistic and to identify emerging plant-contaminant Throughout the history of this symposium, the loss
issues. The work presented in the session indicated a of lake macrophytes due to eutrophication has been a
continuing shift toward the use of more realistic test core issue. Now, in the twenty first century, research
species. To refine risk assessments, laboratory studies on the mechanisms of eutrophication continues but
used more realistic exposure conditions than standard with a somewhat different emphasis; we now see more
techniques; an example was presented at the conference work presented on systems that are in recovery.
in which pesticide exposure pulses, typical of running Research has turned to drivers that influence the
water bodies, caused less harm to gibbous duckweed recovery trajectory, for example, trophic interactions
(Lemna gibba L.) than standard exposure conditions involving herbivores, which have been somewhat
(personal communication). A higher tier approach, neglected in the past, and issues associated with the
using mesocosms, proved effective when investigating role of invasive species.

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The future of invasive species management In spite of the development of genetic and cyto-
plasmic tools, morphological traits are still relevant as
The spread of invasive species and decline in biodi- well as the role of population traits, for example, for
versity is associated with accelerating globalization, dispersal and survival. Many ecosystem processes are
human migration and increasing pressures on fresh- also driven by vegetation, shaping succession of both
water supplies; however, whilst challenging, success- plant and animal communities, in the short- and long-
ful invasive species management has been terms, in which interspecific competition and envi-
demonstrated using combinations of lake and aquatic ronmental constraints determine the end point. Under-
plant-based approaches matched with appropriate standing such processes is fundamental for
management tools (Havel et al., 2015). In some cases, biomanipulation, ecosystem restoration and the proper
regime shifts amongst aquatic flora, such as floating to management of both constructed and natural wetlands.
submerged vegetation, may follow from the use of
classical biological control (Cuda et al., 2008; Bakker
et al., 2016). Yet in other cases, invasive aquatic plants Conclusions
may not be considered the primary drivers of change,
adding to debate surrounding the anthropocentric Both the conference presentations and this resulting
interpretation of benefits (vs detriments) for many special issue of Hydrobiologia reflect the broad
non-native species in impacted habitats. Increasingly, discipline that aquatic botany has become over the
there is a focus towards, arguably, bigger more last century. Research interest in aquatic plants ranges
‘threatening’ issues such as climate change in the from the use of aquatic plants as model organisms, to
management of invasive species that could result in the roles of aquatic plants within ecosystems and to the
greater impacts from existing nuisance aquatic plants conservation of aquatic plants themselves. Further-
at a global level. For example, alien aquatic species more, the range of countries and continents repre-
can reduce the diversity of native seedbanks, thereby, sented by conference delegates and authors of papers
jeopardizing future restoration. Targeted experimental in this special issue illustrate the global relevance of
work in both field and laboratory conditions is aquatic plant research in the early twenty first century.
allowing researchers to understand competitive inter- Currently, the International Symposia on Aquatic
actions between native and invasive species (Gérard & Plants are dominated by research on freshwater taxa,
Triest, 2017). Continued research investment is and in particular those found in shallow lakes.
required to manage the spread of invasive species. However, greater integration of freshwater macro-
The development of new knowledge and techniques phyte and marine seagrass research efforts, and their
will likely provide new opportunities in the future for associated literatures, would benefit our overall
more effective invasive species management and understanding of aquatic plant biology, management
aquatic restoration (e.g. Lozano & Brundu, 2017). and conservation. Whilst aquatic plant species may
differ across ecotones, the processes that shape aquatic
Fundamental science plant assemblages, such as bottom-up and top-down
control and competitive processes, will share common
Applied aspects dominate much of current aquatic plant elements. For example, recent research into herbivory
research, such as aquatic plant populations’ restoration, on aquatic plants has synthesized information from
monitoring and ecological quality assessment, and freshwater, brackish and marine ecosystems (e.g.
different forms of response of aquatic plants to human Bakker et al., 2016; Wood et al., 2017a).
disturbance or novel ways to control plant overgrowth. Our understanding of aquatic plants, the range of
Nonetheless, fundamental science is often the basis for scientific issues being addressed and the range of
management actions, and indeed many failures relate to techniques available to researchers have all arguably
the lack of taxonomic resolution, the misunderstanding never been greater. This is to be welcomed, as the
of species autecology and role in the ecosystem, or challenges facing researchers and practitioners have
undefined tolerance responses over the disturbance also never been more pressing. Climate change, rising
gradient. Fundamental science, thus, provides, in large human demand for resources including water, pollu-
part, the key to successful plant management. tion of freshwater resources, the spread of invasive

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non-native species, land-use changes and intensifica- Brix, H., 1997. Do macrophytes play a role in constructed treat-
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Acknowledgements We are grateful to André Padial, Baz From biology to management. Hydrobiologia 570: 1–263.
Hughes and two anonymous reviewers for their helpful Campbell, C. J., C. V. Johns & D. L. Nielsen, 2014. The value of
comments on earlier drafts of this manuscript. plant functional groups in demonstrating and communi-
cating vegetation responses to environmental flows.
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