Received: 1 September 2019 Revised: 26 October 2019 Accepted: 2 November 2019

DOI: 10.1002/aqc.3268

INVITED COMMENTARY

Editorial: Conservation of groundwaters and their dependent

ecosystems: Integrating molecular taxonomy, systematic
reserve planning and cultural values

Andrew J. Boulton
School of Environmental and Rural Science, University of New England, Armidale, Australia

Correspondence
Andrew Boulton, School of Environmental and Rural Science, University of New England, Armidale, NSW 2351, Australia.
Email: aboulton@une.edu.au

K E Y W O R D S : cultural groundwater, environmental DNA, groundwater protection, groundwater-dependent ecosystems, rehabilitation,

restoration, socio-hydrogeology, stygofauna, systematic conservation planning

1 | I N T RO DU CT I O N referring to all aspects of groundwater conservation has hovered just

above 20% (Figure 1b; see caption for details of this bibliometric anal-
The fundamental importance of groundwater is indisputable. Compris- ysis). Of course, such analyses severely underestimate the numbers of
ing 97% of Earth's unfrozen fresh water, groundwater is essential to papers that specifically describe examples or strategies of conserving
the global hydrological cycle and provides crucial ecosystem services. groundwater habitats or biodiversity. My analysis of a random subset
Aquifers, particularly shallow ones, are inhabited by diverse and of 100 papers for each of five years (2014–2018) from the
unique invertebrates (stygofauna) and sometimes even fish that have bibliometric data used in Figure 1 indicated this proportion was actu-
adaptations to a lightless environment with limited productivity (Hose, ally only 1–2% of the papers in each year. In Aquatic Conservation:
Asmyhr, Cooper, & Humphreys, 2015). Many of these species are Marine and Freshwater Ecosystems, the proportion of papers describing
‘short-range endemics’ (Harvey, 2002) and may only be known from aspects of groundwater conservation has averaged 2.8% (SE = 0.47%)
one or two sampling bores (Karanovic, Eberhard, Perina, & Callan, over the last 20 years. Adding the phrase ‘Conservation of groundwa-
2013). Groundwater also sustains diverse surface ecosystems, ter habitats’ to the journal's scope in 2016 has not yet led to any
ranging from swamps and baseflow-fed rivers through to large areas appreciable increase.
of groundwater-dependent terrestrial vegetation. These surface- In earlier editorials on this topic (Boulton, 2005, 2009), I
expression groundwater-dependent ecosystems (GDEs) harbour rich expressed concerns about the limited public and political attention
troves of aquatic and terrestrial biodiversity and have significant eco- that groundwater conservation attracts and described some of the
logical, economic, and cultural values (Kløve et al., 2011). In parts of advances in scientific knowledge, legislative protection, and conser-
the world where groundwater has been severely drawn down, con- vation of groundwaters and their dependent ecosystems. It is time
taminated, or both, there have been serious and often irreversible to evaluate progress. As space is limited, I have focused on three
impacts on biodiversity, natural and agricultural productivity, and issues: (a) the feasibility of restoration and rehabilitation of
human well-being (Mammola et al., 2019). degraded groundwaters; (b) integrating molecular tools with system-
Despite the hydrological, ecological, economic, and social impor- atic reserve planning; and (c) the concept of cultural groundwater as
tance of groundwaters, research on their conservation lags far behind part of the socio-hydrogeology of groundwater conservation. I have
that on surface waters. In the last two decades, there has been a steep drawn mainly on Australian examples, because groundwater ecosys-
rise in numbers of papers referring to all aspects of conservation of tems ‘down under’ are most familiar to me. However, all three
fresh waters, whereas the trend is far more modest for the groundwa- issues are global, and parallel examples occur in many other parts of
ter subset (Figure 1a). In the last decade, the proportion of papers the world.

Aquatic Conserv: Mar Freshw Ecosyst. 2020;30:1–7. wileyonlinelibrary.com/journal/aqc © 2020 John Wiley & Sons, Ltd. 1
2 BOULTON

F I G U R E 1 (a) Annual numbers of

papers published between 1991 and May
2019 referring to all aspects of
conservation of all fresh waters (blue line)
and of groundwaters only (black line).
(b) Annual proportions of papers referring
to groundwater conservation expressed
as a percentage of papers referring to all
aspects of conservation of all fresh
waters. As 2019 data are incomplete, the
lines are dotted in (a). Results are from a
Web of Science bibliometric analysis
(27 May 2019) using the search terms:
(groundwater* OR aquifer* OR phreat*
OR spring* OR freshwat* OR aquat* OR
river* OR wetland* OR stream* OR lake*
OR swamp* OR bog* OR fen* OR pond*)
AND (conserv* OR protect* OR preserv*)
and then (groundwater* OR aquifer* OR
phreat*) AND (conservation OR
protection OR preservation). Asterisks
represent ‘wildcards’ to capture plurals
and other suffixes

2 | IS IT FEASIBLE TO RESTORE OR (Costigan, Kennard, Leigh, Datry, & Boulton, 2017). Contamination of
R E H A B I L I T A T E D A M A G E D GR O U ND W A T E R S groundwater with salt, heavy metals, hydrocarbons, or other pollut-
A N D TH E I R DE P E N D E N T E C O S Y S T E M S ? ants is widespread and often devastating to the biota of GDEs
(Brown, Bach, Aldous, Wyers, & DeGagné, 2011). By the time the
The persistence of biota and ecological processes in aquifers and impacts are evident at the surface, the contamination or altered
surface-expression GDEs relies on the maintenance of their natural groundwater regime has often affected large areas of GDEs, and its
groundwater regime and water quality. ‘Groundwater regime’ refers effects may be irreversible.
to the temporal variability in magnitude, frequency, duration, timing, Efforts to restore or rehabilitate the groundwater regime of dam-
and rate of change of groundwater levels, storage, and pressures at aged groundwaters and GDEs have met with limited success. The
given spatial scales (Kath, Boulton, Harrison, & Dyer, 2018). Altering main challenges include obtaining a sustainable supply of sufficient
the groundwater regime can alter the frequency, duration, or timing water of suitable quality to replace lost groundwater, remediating a
of hydrological connectivity between surface-expression GDEs and sufficiently large spatial extent to restore the depleted aquifer and its
vital groundwater, leading to reduced production and even mortality groundwater supply to surface-expression GDEs, and maintaining the
of their biota. For example, sustained drawdown and lowered water amendment into the future until the groundwater regime is restored.
tables have led to changes in composition, condition, and function In some cases, partial restoration is possible (e.g. boreal springs,
of terrestrial vegetation GDEs (Eamus, Zolfaghar, Villalobos-Vega, Lehosmaa et al., 2017), but I was unable to find any published exam-
Cleverly, & Huete, 2015). Lowered rates of groundwater discharge ples where complete recovery of pre-impact species diversity and
into rivers change thermal and chemical dynamics in their hyporheic functional attributes in aquifers or GDEs had been achieved. Further,
zones (Boulton & Hancock, 2006) and reduce flow duration and the as seen in case studies of restoration or rehabilitation of surface
persistence of refugial pools in groundwater-fed intermittent rivers waters (Rubin, Kondolf, & Rios-Touma, 2017), there are seldom
INVITED COMMENTARY 3

adequate pre-impact data or reference sites, clearly stated goals, or vegetation communities, is likely but may need additional manage-
appropriate monitoring of ecological responses to the remedial ment of invasive species and other stressors. In light of this gloomy
activities. prognosis for full restoration of damaged groundwater ecosystems,
Sometimes, efforts to prevent the loss of subterranean species their effective conservation is even more crucial. Fortunately, there
border on the desperate. One poignant example in south-western have been some recent advances in molecular technology and system-
Australia is the effort to address the precipitous declines in waters atic reserve planning that may help — my next point.
and stygofauna of limestone caves underneath the Leeuwin-
Naturaliste Ridge, where climatic drying and other stressors have pro-
gressively lowered groundwater levels (Eberhard & Davies, 2011). The 3 | I N T E G R A T I N G M O L E C U LA R T O O L S
caves' unique stygofaunal communities are associated with mats of W I T H S Y S T E M A T I C R E S E R V E P L A N N I NG TO
submerged rootlets of trees, and several have been listed as Critically C O N S E R V E G R O U N D W A T E RS A N D T H E I R
Endangered under Australia's Environmental Protection and Biodiversity DE P E N D E N T E C O S Y S T E M S
Conservation Act. As reinstating historical groundwater regimes is
impossible, water must be added to the caves to prevent the drying Aquatic conservation of subsurface habitats is severely hampered by
that may lead to extinction of these dwindling communities. In Lake the lack of knowledge about even fundamental attributes, such as
Cave, water captured from winter rainfall is trickled into the cave how many species there are, where they occur, and what influences
during the summer dry season while monitoring water quality to their distribution. Mammola et al. (2019) tabulate these serious knowl-
ensure its suitability for the root-mat communities. Although not ideal, edge shortfalls and the specific problems related to conservation of
these measures appear to have forestalled the loss of Lake Cave's groundwater species. One particular challenge is that most subterra-
stygofauna but are probably too late for those of other caves, such as nean habitats are inaccessible to humans except indirectly (Ficetola,
Jewel Cave (Eberhard, 2014). Canedoli, & Stoch, 2019). Traditionally, stygofauna have either been
Even if groundwater regimes can be restored or rehabilitated, collected from water in accessible caves or by sampling scattered
recovery of aquifer communities to their original composition is far bores and wells (which themselves may not reliably represent adjacent
from certain. Most groundwater animals are intrinsically vulnerable to aquifer communities; Korbel, Chariton, Stephenson, Greenfield, &
periods of drawdown because they lack desiccation-resistant stages Hose, 2017). More recently, these traditional approaches have been
and can be stranded when water levels fall during drawdown complemented by molecular ones, such as the use of environmental
(Stumpp & Hose, 2013). Many have limited tolerance for shifts in DNA (eDNA) shed by organisms into the surrounding environment
abiotic conditions and have low fecundity, so population growth is (e.g. from faeces, chitin, or mucus). Assays for eDNA are claimed to
slow. As there is little redundancy in subterranean communities provide a rapid, non-invasive, and cost-effective option for docu-
(Gibert & Deharveng, 2002), they have low ecological resilience and menting occurrences of species that are otherwise difficult to detect,
are more vulnerable to disturbance (Mammola et al., 2019). Many including rare or threatened aquatic invertebrates (Currier, Morris,
stygofauna species are also ‘short-range endemics’ with extremely Wilson, & Freeland, 2018).
restricted ranges (<1,000 km2; Eberhard et al., 2009), and sometimes Their routine application in groundwaters appears promising. For
much less (<1–100 km2), such as the ‘priority communities’ in Western example, Niemiller et al. (2018) demonstrated the utility of eDNA for
Australian calcrete faunas described in Harvey et al. (2011). These the detection of Stygobromus hayi, a rare and endangered groundwa-
restricted ranges may be exceeded by the footprints of proposed ter amphipod that lives in a unique habitat (the hypotelminorheic,
developments (Halse, 2018) and areas of groundwater drawdown defined as a persistent wet spot fed by subsurface water in a slight
associated with mine dewatering or prolonged irrigation pumping. The depression, rich in organic matter and perched on a clay layer typically
persistence of many groundwater species relies on maintaining or up to 50 cm below the surface; Culver, Pipan, & Gottstein, 2006) that
reconnecting hydrological and ecological connections between is challenging to sample using traditional approaches. This approach is
surface and subsurface systems, and this is often suggested when pro- an efficient way of initially screening potential habitats for rare
posing conservation measures for endangered stygofauna (e.g. Ercoli groundwater invertebrates before investing effort and money into
et al., 2019). traditional sampling approaches, and avoids the ‘take’ of protected
So, the answer to the question posed in this section's title is species (Niemiller et al., 2018). However, there are several methodo-
‘Probably not’. In most cases, addition of water (e.g. aquifer injections, logical constraints (e.g. uncertainty about the quantitative relationship
alluvial recharge schemes) will need to continue for decades until and time frames of continuous eDNA production and degradation in
recovery occurs. Even if the natural groundwater regime and water the environment), and eDNA alone cannot provide quantitative data
quality can be reinstated, it is highly likely that some elements of the on abundance. Nonetheless, combined with DNA metabarcoding
groundwater fauna will become extinct, especially some short-range (a technique based on the extensive parallel DNA sequencing of entire
endemics with ranges that fall completely within the area of impact. communities; Cristescu, 2014) and other molecular approaches (Saccò
Partial restoration or rehabilitation may be possible, resulting in a et al., 2019), there appears to be great potential for considerably
different stygofauna assemblage. More complete recovery of surface- enhancing the quality and coverage of crucial data on the distribution
expression GDEs, such as groundwater-fed swamps and terrestrial of stygofauna to guide planning for effective conservation.
4 BOULTON

Of course, these molecular approaches can be (and are) used to spatially patchy, forcing Linke et al. (2019) to use geological regions
complement traditional sampling of surface-expression GDEs too. and the geophysical template of the aquifers as surrogates for biodi-
Unlike subterranean environments, however, the mapping of these versity. Although this approach is not uncommon in conservation
surface habitats is made easier with drones, satellites, and remote- planning, the authors acknowledged that these surrogates often
sensing technology (Mammola et al., 2019). For some taxa greatly under-represent true biodiversity and short-range endemism,
(e.g. phreatophytic vegetation), these tools can also rapidly collect so that the resulting conservation planning outcomes may be over-
information on groundwater use, condition, and distribution over large simplified for aquifers. Furthermore, fine-scale attributes play a major
areas (Eamus et al., 2015). Far less common is the integrated sampling role in controlling the distribution of stygofauna, yet these attributes
of subsurface and surface-expression GDEs at the same time in a are poorly captured at a regional scale. Nonetheless, Linke et al.
region to obtain a concurrent and collective perspective of the com- (2019) showed that including aquifers in freshwater conservation
position and distribution of groundwater-dependent biota. This allows planning does not necessarily compromise efficiency if it is done in
an assessment of their vulnerability to altered groundwater regimes the planning phase, and only marginally increases the overall land
(Kath et al., 2018) and thus can inform systematic planning to opti- needed compared with considering only surface waters. With more
mize conservation of surface and subsurface habitats and their biota. stygofauna data and finer taxonomic resolution, it will be possible to
Groundwaters have seldom been explicitly included in systematic use a direct biotic signal rather than environmental surrogates for
conservation planning to identify key areas to protect freshwater integrated conservation planning. Importantly, such integrated
organisms. A literature search by Linke, Turak, Asmyhr, and Hose planning can incorporate the crucial connectivity among landscape
(2019) found only three studies since 2007. The first of these used a elements, including among aquifers and between groundwaters and
comprehensive database of 1059 groundwater species collected their associated GDEs — a connection that has been recognized by
across six European regions to prioritize biodiversity hotspots (Michel humans for thousands of years, and which brings me to my final point.
et al., 2009) and advocated an aquifer-selection approach that could
incorporate species representation targets as well as socio-economic
costs associated with the vulnerability of aquifers and the amount of 4 | ‘C U L T U R A L G R O U N D W A T E R ’ A N D
human activity in the catchment. The second study (Asmyhr, Linke, C O N S E R V A T I O N OF CO N N E C T E D W A T E R S
Hose, & Nipperess, 2014) used molecular sequence data and phyloge-
netic diversity as surrogates for stygofauna diversity in aquifers of In Australia, especially the arid and semi-arid zones that comprise
New South Wales, Australia. These data were incorporated as conser- more than 70% of the continent, almost all permanent surface fresh
vation features in the systematic conservation planning software waters are groundwater dependent. The significance of groundwater
Marxan by maximizing phylogenetic diversity instead of species connectivity to maintain these waters and other GDEs has been rec-
diversity. The third study assessed conservation of riverine and non- ognized by Indigenous Australians for thousands of years, forming a
riverine freshwater wetlands on the North China Plain, but it also common theme in many of their narratives (Wooltorton, Collard, &
incorporated five major subsurface aquifers (Li et al., 2017). However, Horwitz, 2019; Yu, 2000). Typically, the Indigenous perspective
this study did not assess groundwater biota (the focal species were emphasizes interdependence among all landscape elements, and inter-
waterbirds), with subsurface aquifers simply included as conservation pretations of the environment do not distinguish landscapes from
features in addressing vertical connectivity (Li et al., 2017). waterscapes or groundwater from surface water (Wooltorton et al.,
None of these three studies integrated aquifers and surface 2019). In Australia, Western recognition of the conservation signifi-
waters spatially in a systematic conservation planning framework that cance of this interdependence among landscape elements appears far
could identify key areas for protection, especially of groundwaters less entrenched and has often resulted in site-specific management
and their associated surface-expression GDEs. The main challenge is strategies and Eurocentric paradigms for perceiving and assigning
obtaining suitable concurrent data sets of taxa and conservation value (Bark et al., 2015).
values for aquifers and surface waters in a large region. Here is where In the last decade, there have been efforts to integrate the rich
the use of molecular tools such as eDNA and DNA metabarcoding to store of Indigenous knowledge, including this understanding of
complement traditional surveys can be integrated with the application interdependence among landscape elements, into water resource
of modern systematic conservation planning methods to address this management and aquatic conservation in Australia (Liedloff,
knowledge gap. Woodward, Harrington, & Jackson, 2013; Woodward, Jackson, Finn, &
Linke et al. (2019) demonstrated the feasibility of this approach in Marrfurra McTaggart, 2012). Over the same period, there have been
their development of a method to link rivers, wetlands, and aquifers reforms in water law and governance to enable more robust participa-
spatially in systematic conservation planning which they applied to tion by Aboriginal peoples and Torres Strait Islanders in water man-
the Hunter Valley in New South Wales, Australia. This region is one of agement across Australian jurisdictions, one component of which is
the few in Australia where stygofauna have been extensively termed ‘cultural flows’ (Nelson, Godden, & Lindsay, 2018).
sampled and there are detailed data sets for fish and aquatic Cultural flows are defined as ‘Water entitlements that are legally
macroinvertebrates in the overlying rivers and wetlands. However, and beneficially owned by Indigenous Nations of a sufficient and
even in this well-studied area, stygofaunal data were limited and adequate quantity and quality to improve the spiritual, cultural,
INVITED COMMENTARY 5

environmental, social and economic conditions of those Indigenous particular, these assessments must conserve hydrological connectivity
Nations’ (Nelson et al., 2018). Although groundwater is included in in all three dimensions so that surface-expression GDEs are explicitly
this definition and in much of the literature on this topic (Australian included in strategies to protect groundwater ecosystems.
Cultural Heritage Management, 2014), the term ‘cultural flows’ may These conservation approaches must be able to monitor and pro-
be misinterpreted to refer solely to surface water flows, analogous to tect the groundwater regimes of diverse GDEs at sufficiently broad
‘environmental flows’ released strategically from dams to meet flow spatial scales (Kath et al., 2018). Where possible, groundwater users
requirements of riverine and floodplain habitats downstream should be actively involved in monitoring water levels and developing
(cf. Finn & Jackson, 2011). However, this narrow interpretation over- collaborative response plans. For example, the potential for cumula-
looks the interdependence among landscape elements inherent to the tive impacts on groundwaters from development of the coal seam gas
Indigenous perspective and the crucial role of groundwater in industry led to the declaration in 2011 of the Surat Cumulative Man-
supporting many surface waters and their associated vegetation. agement Area in the Surat and Southern Bowen Basins, Queensland.
To minimize the risk of this misinterpretation, a term like ‘cultural Under current legislation, where a Cumulative Management Area is
groundwater’ may be useful to supplement the current one. Con- declared, the independent Office of Groundwater Impact Assessment
versely, a collective term like ‘cultural flows and groundwater’ may is required to undertake a cumulative impact assessment on the
better capture the perspective of interdependence. In this way, the groundwater resource, water supply bores, springs, and other GDEs,
integration of Indigenous and Western perspectives in Australian including regional groundwater flow modelling (Office of Groundwa-
water resource management and allocations of water entitlements ter Impact Assessment, 2019). The Office of Groundwater Impact
could be more explicit about acknowledging the fundamental signifi- Assessment also coordinates a Joint Industry Plan where each of the
cance of groundwater. This is the essence of socio-hydrogeology, a main tenure holders is allocated a specific network of monitoring
concept introduced by Re (2015) to specifically incorporate the social bores and responsibility for particular springs that may be affected or
dimension into hydrogeological investigations. Socio-hydrogeology is that may serve as useful reference sites. Every 3 years, the Office of
a rapidly growing field (reviewed in Hynds et al., 2018), but its Groundwater Impact Assessment revises their models and predictions
relevance to conservation of groundwaters and their dependent using data from the joint plan and other sources. This coordinated
ecosystems has not been fully explored. There is scope to use socio- approach involving each tenure holder helps ensure consistent moni-
hydrological approaches to better integrate Indigenous and Western toring protocols and management activities to address cumulative
knowledge and understanding into conservation plans that include impacts and encourages collective responsibility.
cultural groundwater in Australia to protect the biota and ecological Another crucial task in effective groundwater conservation is to
processes in connected waters and their landscapes. communicate to the public more effectively the values of groundwa-
ters and their GDEs, the threats they face, and how we can best pro-
tect them. I have urged this in previous editorials (Boulton, 2005,
5 | C O N CL U S I O N S 2009), and I reiterate it here. In addition to conventional educational
programmes, there is scope for more explicit recognition of Indige-
Given the very limited success of efforts to restore or rehabilitate nous knowledge and understanding of groundwaters and their entitle-
damaged groundwater ecosystems, their protection is essential. ments to ‘cultural groundwater’. The integration of Indigenous and
Groundwater ecosystems harbour numerous unique and often highly Western perspectives in a socio-hydrogeological framework (Hynds
endemic species, perform many essential ecosystem services, and et al., 2018) may be a promising way to enhance communication
support diverse dependent ecosystems, such as baseflow-fed rivers about groundwater conservation and initiate regional protection of,
and particular associations of terrestrial vegetation. However, all these for example, biodiversity hotspots identified by systematic conserva-
are being lost or impaired at an alarming rate. Worldwide, groundwa- tion planning. At the same time, we must address the low level of
ter ecosystems face serious threats, particularly to their groundwater sympathy for subterranean invertebrates evident amongst
regimes and water quality. These threats have prompted many scien- policymakers, the general public, and even many biologists working on
tists to call for urgent improvements to the protection of groundwa- surface fauna (Halse, 2018). Even activities such as recreational
ters (Danielopol, Griebler, Gunatilaka, & Notenboom, 2003; Mammola scuba-diving in groundwater springs can have environmental impacts
et al., 2019). (Thorbjörnsson, Ólafsdóttir, Kristjánsson, Chambers, & Burns, In press)
What can we do? One of the most pressing tasks is to identify and require careful management of diver numbers and behaviour to
the world's subterranean hotspots of diversity, complementing prevent ecological damage to these GDEs.
traditional surveys with molecular methods and then using systematic I echo the recent ‘warning’ by 21 groundwater scientists on the
conservation planning to prioritize areas for protection. Where need for conservation of groundwater ecosystems (Mammola et al.,
species-level taxonomic and distribution data are limited, community- 2019). In the absence of convincing evidence that groundwater eco-
level approaches (Mokany, Harwood, Halse, & Ferrier, 2019) set systems impaired by prolonged groundwater drawdown or extensive
within transdisciplinary research frameworks (Gibson, Humphreys, contamination can be readily restored or rehabilitated, effective pro-
Harvey, Hyder, & Winzer, 2019; Saccò et al., 2019) may be needed tection is imperative. Sustainable stewardship of our groundwaters
to guide identification of groundwater biodiversity hotspots. In and their dependent ecosystems is an ethical duty as well as a
6 BOULTON

practical way to ensure continued provision of their myriad ecosystem freshwater pearly mussel species (Bivalvia: Unionidae). Aquatic Conser-
services and conservation of their unique and diverse biota and eco- vation: Marine and Freshwater Ecosystems, 28, 545–558. https://doi.
org/10.1002/aqc.2869
logical processes.
Danielopol, D. L., Griebler, C., Gunatilaka, A., & Notenboom, J. (2003).
Present state and future prospects for groundwater ecosystems.
ACKNOWLEDGEMEN TS Environmental Conservation, 30, 104–130. https://doi.org/10.1017/
I thank Phil Boon for suggesting I revisit this topic, and I am grateful S0376892903000109
Eamus, D., Zolfaghar, S., Villalobos-Vega, R., Cleverly, J., & Huete, A.
to him, Stefan Eberhard, Steve Flook, Stuart Halse, and Simon Linke
(2015). Groundwater-dependent ecosystems: Recent insights from
for helpful comments on drafts of this editorial. satellite and field-based studies. Hydrology and Earth System Sciences,
19, 4229–4256. https://doi.org/10.5194/hess-19-4229-2015
CONF LICT OF IN TE RE ST Eberhard, S. M. (2014). Climate changes and other stressors on cave
stygofauna. In A. J. Boulton, M. A. Brock, B. J. Robson, D. S. Ryder,
None to declare.
J. M. Chambers, & J. A. Davis (Eds.), Australian freshwater ecology:
Processes and management (p. 200). Chichester, UK: Wiley-Blackwell.
ORCID Eberhard, S. M., & Davies, S. (2011). Impacts of drying climate on aquatic
Andrew J. Boulton https://orcid.org/0000-0001-7393-2800 cave fauna in Jewel Cave and other caves in southwest Western
Australia. Journal of the Australasian Cave & Karst Management Associa-
tion, 83, 6–13.
RE FE R ENC E S
Eberhard, S. M., Halse, S. A., Williams, M. A., Scanlon, M. D., Cocking, J., &
Asmyhr, M. G., Linke, S., Hose, G., & Nipperess, D. A. (2014). Systematic Barron, H. J. (2009). Exploring the relationship between sampling effi-
conservation planning for groundwater ecosystems using phylogenetic ciency and short-range endemism for groundwater fauna in the Pilbara
diversity. PLoS ONE, 9, e115132. https://doi.org/10.1371/journal. region, Western Australia. Freshwater Biology, 54, 885–901. https://
pone.0115132 doi.org/10.1111/j.1365-2427.2007.01863.x
Australian Cultural Heritage Management (2014). Cultural flows literature Ercoli, F., Lefebvre, F., Delangle, M., Godé, N., Caillon, M., Raimond, R., &
review. Brunswick North, VIC, Australia: Murray Lower Darling Rivers Souty-Grosset, C. (2019). Differing trophic niches of three French
Indigenous Nations (MLDRIN), Northern Basin Aboriginal Nations stygobionts and their implications for conservation of endemic
(NBAN) & North Australian Indigenous Land and Sea Management stygofauna. Aquatic Conservation: Marine and Freshwater Ecosystems
Alliance (NAILSMA). https://doi.org/10.1002/aqc.3227
Bark, R. H., Barber, M., Jackson, S., Maclean, K., Pollino, C., & Ficetola, G. F., Canedoli, C., & Stoch, F. (2019). The Racovitzan impediment
Moggridge, B. (2015). Operationalising the ecosystem services and the hidden biodiversity of unexplored environments. Conservation
approach in water planning: A case study of indigenous cultural values Biology, 33, 214–216. https://doi.org/10.1111/cobi.13179
from the Murray–Darling Basin, Australia. International Journal of Biodi- Finn, M., & Jackson, S. (2011). Protecting indigenous values in water man-
versity Science, Ecosystem Services & Management, 11, 239–249. agement: A challenge to conventional environmental flow assess-
https://doi.org/10.1080/21513732.2014.983549 ments. Ecosystems, 14, 1232–1248. https://doi.org/10.1007/s10021-
Boulton, A. J. (2005). Chances and challenges in the conservation of 011-9476-0
groundwaters and their dependent ecosystems. Aquatic Conservation: Gibert, J., & Deharveng, L. (2002). Subterranean ecosystems: A truncated
Marine and Freshwater Ecosystems, 15, 319–323. https://doi.org/10. functional biodiversity. BioScience, 52, 473–481. https://doi.org/10.
1002/aqc.712 1641/0006-3568(2002)052[0473:SEATFB]2.0.CO;2
Boulton, A. J. (2009). Recent progress in the conservation of groundwaters Gibson, L., Humphreys, W. F., Harvey, M., Hyder, B., & Winzer, A. (2019).
and their dependent ecosystems. Aquatic Conservation: Marine and Shedding light on the hidden world of subterranean fauna: A transdis-
Freshwater Ecosystems, 19, 731–735. https://doi.org/10.1002/aqc. ciplinary research approach. Science of the Total Environment, 684,
1073 381–389. https://doi.org/10.1016/j.scitotenv.2019.05.316
Boulton, A. J., & Hancock, P. J. (2006). Rivers as groundwater dependent Halse, S. (2018). Conservation and impact assessment of subterranean
ecosystems: A review of degrees of dependency, riverine processes, fauna in Australia. In O. Moldovan, L. Kováč, & S. Halse (Eds.), Cave
and management implications. Australian Journal of Botany, 54, ecology (pp. 479–494). Cham, Switzerland: Springer International
133–144. https://doi.org/10.1071/BT05074 Publishing. https://doi.org/10.1007/978-3-319-98852-8_23
Brown, J., Bach, L., Aldous, A., Wyers, A., & DeGagné, J. (2011). Ground- Harvey, M. S. (2002). Short-range endemism amongst the Australian
water-dependent ecosystems in Oregon: An assessment of their distri- fauna: Some examples from non-marine environments. Invertebrate
bution and associated threats. Frontiers in Ecology and the Environment, Systematics, 16, 555–570. https://doi.org/10.1071/IS02009
9, 97–102. https://doi.org/10.1890/090108 Harvey, M. S., Rix, M. G., Framenau, V. W., Hamilton, Z. R., Johnson, M. S.,
Costigan, K., Kennard, M., Leigh, C., Datry, T., & Boulton, A. J. (2017). Flow Teale, R. J., … Humphreys, W. F. (2011). Protecting the innocent:
regimes in intermittent rivers and ephemeral streams. In T. Datry, Studying short-range endemic taxa enhances conservation outcomes.
N. Bonada, & A. J. Boulton (Eds.), Intermittent rivers and ephemeral Invertebrate Systematics, 25, 1–10. https://doi.org/10.1071/IS11011
streams: Ecology and management (pp. 51–78). London, UK: Academic Hose, G. C., Asmyhr, M. G., Cooper, S. J., & Humphreys, W. F. (2015).
Press. https://doi.org/10.1016/B978-0-12-803835-2.00003-6 Down under down under: Austral groundwater life. In A. Stow,
Cristescu, M. E. (2014). From barcoding single individuals to N. Maclean, & G. Holwell (Eds.), Austral ark: The state of wildlife in
metabarcoding biological communities: Towards an integrative Australia and New Zealand (pp. 512–536). Cambridge, UK: Cambridge
approach to the study of global biodiversity. Trends in University Press.
Ecology & Evolution, 29, 566–571. https://doi.org/10.1016/j.tree. Hynds, P., Regan, S., Andrade, L., Mooney, S., O'Malley, K., DiPelino, S., &
2014.08.001 O'Dwyer, J. (2018). Muddy waters: Refining the way forward for the
Culver, D. C., Pipan, T., & Gottstein, S. (2006). Hypotelminorheic — A “sustainability science” of socio-hydrogeology. Water, 10, 1111.
unique freshwater habitat. Subterranean Biology, 4, 1–7. https://doi.org/10.3390/w10091111
Currier, C. A., Morris, T. J., Wilson, C. C., & Freeland, J. R. (2018). Valida- Karanovic, T., Eberhard, S. M., Perina, G., & Callan, S. (2013). Two new
tion of environmental DNA (eDNA) as a detection tool for at-risk subterranean ameirids (Crustacea: Copepoda: Harpacticoida) expose
INVITED COMMENTARY 7

weaknesses in the conservation of short-range endemics threatened (MLDRIN), Northern Basin Aboriginal Nations (NBAN) & North
by mining developments in Western Australia. Invertebrate Systemat- Australian Indigenous Land and Sea Management Alliance (NAILSMA).
ics, 27, 540–566. https://doi.org/10.1071/IS12084 Niemiller, M. L., Porter, M. L., Keany, J., Gilbert, H., Fong, D. W.,
Kath, J., Boulton, A. J., Harrison, E., & Dyer, F. (2018). A conceptual frame- Culver, D. C., … Taylor, S. J. (2018). Evaluation of eDNA for groundwa-
work for ecological responses to groundwater regime alteration ter invertebrate detection and monitoring, a case study with endan-
(FERGRA). Ecohydrology, 11, e2010. https://doi.org/10.1002/eco. gered Stygobromus (Amphipoda, Crangonyctidae). Conservation
2010 Genetics Resources, 10, 247–257. https://doi.org/10.1007/s12686-
Kløve, B., Ala-aho, P., Guillaume, B., Boukalova, Z., Ertürk, A., 017-0785-2
Goldscheider, N., … Widerlund, A. (2011). Groundwater dependent Office of Groundwater Impact Assessment. (2019). https://www.business.
ecosystems. Part I. Hydroecological status and trends. Environmental qld.gov.au/industries/mining-energy-water/resources/landholders/-
Science and Policy, 14, 770–781. https://doi.org/10.1016/j.envsci. csg/ogia. [17 July 2019].
2011.04.002 Re, V. (2015). Incorporating the social dimension into hydrogeochemical
Korbel, K., Chariton, A., Stephenson, S., Greenfield, P., & Hose, G. C. investigations for rural development: The Bir Al-Nas approach for
(2017). Wells provide a distorted view of life in the aquifer: Implica- socio-hydrogeology. Hydrogeology Journal, 23, 1293–1304. https://
tions for sampling, monitoring and assessment of groundwater ecosys- doi.org/10.1007/s10040-015-1284-8
tems. Scientific Reports, 7, 40702. https://doi.org/10.1038/srep40702 Rubin, Z., Kondolf, G. M., & Rios-Touma, B. (2017). Evaluating stream res-
Lehosmaa, K., Jyväsjärvi, J., Virtanen, R., Rossi, P. M., Rados, D., toration projects: What do we learn from monitoring? Water, 9, 174.
Chuzhekova, T., … Muotka, T. (2017). Does habitat restoration https://doi.org/10.3390/w9030174
enhance spring biodiversity and ecosystem functions? Hydrobiologia, Saccò, M., Blyth, A., Bateman, P. W., Hua, Q., Mazumder, D., White, N., …
793, 161–173. https://doi.org/10.1007/s10750-016-2760-4 Grice, K. (2019). New light in the dark — A proposed multidisciplinary
Li, X. W., Shi, J. B., Song, X. L., Ma, T. T., Man, Y., & Cui, B. S. (2017). Inte- framework for studying functional ecology of groundwater fauna. Sci-
grating within-catchment and interbasin connectivity in riverine and ence of the Total Environment, 662, 963–977. https://doi.org/10.1016/
nonriverine freshwater conservation planning in the North China Plain. j.scitotenv.2019.01.296
Journal of Environmental Management, 204, 1–11. https://doi.org/10. Stumpp, C., & Hose, G. C. (2013). The impact of water table drawdown
1016/j.jenvman.2017.04.089 and drying on subterranean aquatic fauna in in-vitro experiments. PLoS
Liedloff, A. C., Woodward, E. L., Harrington, G. A., & Jackson, S. (2013). ONE, 8, e78502. https://doi.org/10.1371/journal.pone.0078502
Integrating indigenous ecological and scientific hydro-geological Thorbjörnsson, J. G., Ólafsdóttir, J. H., Kristjánsson, B. K., Chambers, C., &
knowledge using a Bayesian network in the context of water resource Burns, G. L. (In press). Recreational scuba diving in a groundwater eco-
development. Journal of Hydrology, 499, 177–187. https://doi.org/10. system: Disturbance mechanisms, ecological impacts and stakeholder
1016/j.jhydrol.2013.06.051 perceptions. Aquatic Conservation: Marine and Freshwater Ecosystems.
Linke, S., Turak, E., Asmyhr, M. G., & Hose, G. (2019). 3D conservation Woodward, E., Jackson, S., Finn, M., & Marrfurra McTaggart, P. (2012).
planning: Including aquifer protection in freshwater plans refines prior- Utilising Indigenous seasonal knowledge to understand aquatic
ities without much additional effort. Aquatic Conservation: Marine and resource use and inform water resource management in northern
Freshwater Ecosystems, 29, 1063–1072. https://doi.org/10.1002/aqc. Australia. Ecological Management and Restoration, 13, 58–64. https://
3129 doi.org/10.1111/j.1442-8903.2011.00622.x
Mammola, S., Cardoso, P., Culver, D. C., Deharveng, L., Ferreira, R. L., Wooltorton, S., Collard, L., & Horwitz, P. (2019). Living water: Groundwa-
Fišer, C., … Zagmajster, M. (2019). Scientists' warning on the conserva- ter and wetlands in Gnangara, Noongar boodjar. PAN: Philosophy,
tion of subterranean ecosystems. BioScience, 69, 641–650. https://doi. Activism, Nature, 14, 5–23.
org/10.1093/biosci/biz064 Yu, S. (2000). Ngapa kunangkul: Living water. Report on the indigenous cul-
Michel, G., Malard, F., Deharveng, L., Di Lorenzo, T., Sket, B., & De tural values of groundwater in the La Grange sub-basin. Perth, WA,
Broyer, C. (2009). Reserve selection for conserving groundwater biodi- Australia: Water and Rivers Commission of Western Australia.
versity. Freshwater Biology, 54, 861–876. https://doi.org/10.1111/j.
1365-2427.2009.02192.x
Mokany, K., Harwood, T. D., Halse, S. A., & Ferrier, S. (2019). Riddles in
How to cite this article: Boulton AJ. Editorial: Conservation
the dark: Assessing diversity patterns for cryptic subterranean fauna
of groundwaters and their dependent ecosystems: Integrating
of the Pilbara. Diversity and Distributions, 25, 240–254. https://doi.
org/10.1111/ddi.12852 molecular taxonomy, systematic reserve planning and cultural
Nelson, R., Godden, L., & Lindsay, B. (2018). Cultural flows: A multi-layer values. Aquatic Conserv: Mar Freshw Ecosyst.
plan for cultural flows in Australia: Legal & policy design. Brunswick 2020;30:1–7. https://doi.org/10.1002/aqc.3268
North, VIC, Australia: Murray Lower Darling Rivers Indigenous Nations