Marine Pollution Bulletin 163 (2021) 111936

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Marine Pollution Bulletin

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

Assessing intertidal seagrass beds relative to water quality in Vanuatu,

South Pacific
Susana Lincoln a,*, Marta Vannoni a, Lisa Benson a, Georg H. Engelhard a, Dieter Tracey b,
Christina Shaw c, Vatumaraga Molisa d
a
The Centre for Environment, Fisheries and Aquaculture Science (Cefas). Pakefield Road, Lowestoft, Suffolk NR33 0HT, UK
b
Science Graphics, 28 Dalziel Street, Stratford, QLD 4870, Australia
c
Vanuatu Environmental Science Society (VESS), PO Box 1630, Anchor House, Kumal Highway, Port Vila, Shefa Province, Vanuatu
d
Department of Environmental Protection and Conservation, PMB 9063, Port Vila, Shefa Province, Vanuatu

ARTICLEINFO
A B S T R A C T
Keywords:
Seagrass Seagrasses are globally important, but their extent is decreasing due to the impact of human activities and
Water quality changing climatic conditions. Seagrass meadows provide vital services, but their condition and distribution
Vulnerability are not yet well known, particularly in many small tropical Pacific islands. In 2018, observations and samples
Human activities were collected from intertidal seagrasses of Efate Island, Vanuatu (South Pacific). Observations included
Intertidal
canopy height, percentage cover, growth rate and species variety. Water quality samples were also collected
Pacific islands
in some locations. Our seagrass metrics suggested more challenging conditions for the seagrasses near Port
Climate change
Vila, the main urban area in the island, where water quality analyses indicated higher levels of dissolved
inorganic nitrogen and suspended solids.

1. Introduction
Seagrass meadows are globally important coastal habitats. They
provide numerous valuable ecosystem services such as food and
refuge to many marine species of fish, invertebrates and endangered
mega- fauna like dugongs and sea turtles (Bell and Pollard, 1989;
Short et al., 2007; Waycott et al., 2011). Seagrasses are also known to
improve water quality by trapping nutrients and pollutants and
stabilising and accreting fine sediments (Waycott et al., 2011). On a
global scale, sea- grass meadows play a significant role in climate
change mitigation because they are capable of absorbing large
amounts of carbon dioxide and storing it for long periods within the
sediment (Duarte, 2002; Duarte et al., 2013). For some of these
services, seagrasses function together with mangrove forests and
coral reefs (Hemminga et al., 1994; Nordlund et al., 2018).
Disturbances derived from a growing population are causing global
declines in seagrass distribution. Poor water quality and increased
turbidity and siltation are particularly damaging to seagrasses (Abal
and Dennison, 1996; Hemminga and Duarte, 2000; Orth et al., 2006;
Way- cott et al., 2011; Grech et al., 2012; McKenzie et al., 2012).
Future climate change impacts in the form of warming temperatures,
rising sea
levels, and physical damage from intense storms and cyclones are ex-
pected to further exacerbate the stress currently experienced by
seagrass habitats (Waycott et al., 2007; Waycott et al., 2011; Luisetti
et al., 2013). The global annual rate of loss for seagrasses in recent
decades has been estimated to range from 0.4 to 16% (Erftemeijer
and Lewis, 2007; Pendleton et al., 2012).
Efate is the third largest and most populated island of Vanuatu,
South Pacific. The narrow fringing reefs that surround Efate and most
of the other islands of Vanuatu do not provide ideal conditions for
seagrass meadows to grow extensively, although they are reportedly
common (Chambers et al., 1990; Laffoley, 2013; McKenzie and
Yoshida, 2017). There have been some studies but the remoteness of
these islands means that there is a lack of data on extent, species
distribution and condition of the seagrass beds in Vanuatu, making it
difficult to assess their status in relation to the rapid population
expansion, which is taking place not only in Vanuatu (Chambers et al.,
1990; Laffoley, 2013; VNSO, 2016) but also in many other small island
countries in the South Pacific (Bani and Esrom, 1993; Waycott et al.,
2011; McKenzie and Yoshida, 2020). In Efate, most of the population
concentrates in and around the capital Port Vila (VNSO, 2016) which
is also a hub for transport and tourism in Vanuatu. As the urban area
expands and develops, so do the impacts

* Corresponding author.
E-mail address: susana.lincoln@cefas.co.uk (S. Lincoln).

https://doi.org/10.1016/j.marpolbul.2020.111936
Received 21 March 2019; Received in revised form 27 November 2020; Accepted 7 December 2020
Available online 23 December 2020
0025-326X/Crown Copyright © 2020 Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/).
S. Lincoln et al.
from activities such as improper waste disposal, water pollution and
poor coastal management (Bani and Esrom, 1993; Waycott et al.,
2011; Ondiviela et al., 2014; CMEP, 2018). These issues can have
negative consequences for seagrass habitats, potentially undermining
their con- dition and their resilience to other pressures.
Seagrass observations (seagrass canopy height, percentage
cover, growth rate and species presence) as well as water quality
indicators (temperature, salinity, total dissolved inorganic nitrogen,
dissolved inorganic phosphorus, chlorophyll-a and total suspended
solids) were gathered across a range of sites in Efate to assess
whether there is po- tential for seagrass beds nearer to Port Vila to be
impacted due to poor water quality conditions compared to seagrass
beds in more remote areas.
2. Methods
2.1. Study area
This study was carried out during August 2018 in five intertidal lo-
cations in Efate where seagrass beds are found: Fatumaru, Pango,
Era- tap, Epau and Pele Island (Fig. 1).
Locations were chosen so they represented a gradient of
decreasing population density and increasing distance from the urban
area of Port Vila: the nearest to Vila and therefore the most densely
populated location represented by Fatumaru Bay, and the furthest
from Vila and less populated location represented by Pele Island
(VNSO, 2016). Table 1 shows a general description of each survey
location, as well as
the population census for each location’s council area (VNSO, 2016)
and estimated distance from Port Vila.
2.2. Water quality parameters
Three separate water samples were collected at each location on
the
2
Marine Pollution Bulletin 163 (2021)
111936
day of the seagrass survey, coinciding with the ebb, low and flow tide
times, for nutrients, chlorophyll-a and suspended sediments analyses.
Water samples were cooled immediately after collection and stored
frozen (nutrients, chlorophyll) or refrigerated (suspended solids) until
each analysis.
Dissolved nutrients samples were filtered in-situ through a single-use
0.45 μm cellulose acetate syringe filter. Nutrients samples were
analysed for ammonium nitrogen (NH4), and + nitrate nitrite (NOx),
as well as
dissolved inorganic phosphorus or filterable reactive phosphorus (DIP
or FRP) according to methods 4500-Norg D, 4500-NH3, 4500-NO3 and
4500-P (APHA-AWWA Method in APHA, 2005). Detection limits were
as
follows: NOx (1 μg N/L), FRP (1 μg P/L), and NH4 (1 μg N/L).
Analytical uncertainty (95% confidence interval of the standard
deviation) was
±8% for NH4, NOx and FRP.
Chlorophyll a (Chl-a) and phaeophytin samples were filtered
through Whatman glass fibre filters, fixed with 0.01 g magnesium
carbonate preservative, and wrapped in aluminium foil prior to
freezing. Chl-a and phaeophytin concentrations were determined using
a spectrophotom- eter after grinding the glass fibres in 90% acetone
(Method 10200H in APHA, 2005).
Total suspended solids (TSS) samples were analysed for
suspended sediment concentration using the ASTM D3977-97 Method
C (ASTM, 2002) and for TSS using the APHA-AWWA 2005 Method
2540 D (detection= limit 1 mg/L; uncertainty 12%; APHA, 2005).
Water temperature and salinity were also recorded three times a
day at each survey site using a WTW 3430 multi parameter meter
(Xylem Analytics, Germany).
2.3. Field survey of intertidal seagrasses
Surveys of intertidal meadows were carried out on foot, on
seagrass beds that were accessible during the lowest diurnal tide. The
surveys were carried out following a modified version of the
Seagrass Watch
S. Lincoln et al. Marine Pollution Bulletin 163 (2021)
Fig. 1. Locations of seagrass surveys in Efate Island, Vanuatu. In Eratap 111936
(detail map B), two separate sites were surveyed within the same location,
although the scale of the map does not necessarily reflect that.

3
Table 1
Description of chosen seagrass survey locations, including population number and population density per km2 based on the 2016 Census of the respective
council areas (VNSO, 2016). Distance to Port Vila city centre is calculated on a straight line.
Seagrass survey
General description Sediment depth and type Population Population Distance from
location (Council 2
Census density (per km ) Port Vila (km)
Area)
2016

Fatumaru Bay (Port

Sheltered bay within the Vila Bay and Harbour Very shallow (5–11 cm) layer of fine, 51,437 2180 1.7
Vila)
area soft, muddy sediment overlay on hard
coral and rock
Pango Vale Vale
Small bay enclosed by narrow fringe reef but Coral rubble mixed with coarse sand 2359 407 7.4
(Pango)
facing open ocean so relatively exposed to (26–109 cm)
wind and waves
Eratap (Eratap) Sheltered area in a lagoon system surrounded Relatively deep (19–87 cm), mostly 6640 91 8.4
by mangrove forests soft,
muddy sandy sediment
Epau (Eton) Small semi-enclosed shallow lagoon Layer of sandy sediment (89–109 cm) 3518 11 22.4
laid over a hard base of coral and rock
Pele Island (Nguna) Sheltered beach on outer island, facing Efate Shallow (28–42 cm) sandy sediment laid 1728 55 33.5
island over rock and coral

×
method described by McKenzie et al. (2003). Due to the time tape. Four 50 50 cm quadrats were thrown randomly into the area and
available, at each survey location a single 50 m transect was laid secured to the ground where they landed for the duration of the
through the centre of the meadow. This method limits × analysis of experiment. The quadrats were subdivided into 10 10 cm squares using
within-site vari- ance but provides information on the status of the string. At the start of the experiment, every T. hemprichii shoot within
seagrass at each lo- cality. Using a measuring tape and stakes, 10 each quadrat was marked by puncturing through the sheath at the base
randomly spaced points were marked along the transect. At each of of the leaves using a hypodermic syringe needle to create a
×
the 10 points, a 50 50 cm quadrat was fixed to the ground by placing it
alongside the measuring tape with the top-left corner of the quadrat
next to the marking stake. While the survey took place during the
lowest diurnal tide, there were some variations between sites in terms
of water depth: the seagrass beds
along the survey transect in Fatumaru were exposed to air, under 2–13
cm of water in Pele and Pango, and under 15–45 cm of water in
Eratap and Epau.
Within each quadrat, all seagrass species were identified and
anno- tated in order as they appeared to be more or less numerous.
The per- centage of seagrass canopy cover was estimated within each
quadrat using the canopy cover example plates from McKenzie et al.
(2003) as guidance. Canopy height was estimated × within a 10 10 cm
subsection of the quadrat (top-left corner of the quadrat next to the
marking stake) as the height of each leaf measured from tip of the leaf
blade to the base of the rhizome over the sediment. Additional
observations on the overall visual appearance of the seagrass within
each quadrat were also anno- tated, such as: green undamaged
leaves; presence of flowers or fruits; presence of epiphytic growth on
the leaf surface; pale leaves mostly covered with epiphytes; or brown,
fragmented and dying leaves.
The thickness of the sediment layer was measured at each end and
at
the mid-point of the Cefas’ survey transects by pushing a 150 cm metal
rod vertically into the sediment until it reached the hard substrate
underneath.
Waypoint positions of all survey transects were recorded using a
Garmin handheld GPS.

2.4. In-situ seagrass growth measurements

Seagrass growth measurements were carried out in-situ in

Fatumaru Bay and Pango Vale Vale during August 2018 (Fig. 1B). The
experiments followed a simplified version of the plastochrone interval
method described by Short and Duarte (2001). The seagrass
meadows in Fatu- maru and Pango were mostly dominated by
Thalassia hemprichii with
relatively short leaves growing on a thin layer of sandy/muddy
sediment over a hard, shallow underlay of rock and coral. During the
low tide, a 25 m2 area was defined over a section of the meadow using
stakes and
 puncture scar. The growth experiment was terminated after 13 days.
This specific time period agrees with the typical plastochrone interval
for T. hemprichii of 10.9 days (Short and Duarte, 2001) and was
chosen to fit with the diurnal low tides in order to safely access the
locations. Upon the conclusion of the experiment on day 13, a total of
20 shoots of T. hemprichii were collected randomly from each quadrat
on each of the two locations and taken to the laboratory. The total
number of new leaves (with no signs of puncture scar) and old leaves
(bearing the scarring from the needle puncture) were counted and
recorded for each shoot. Leaf length was also measured. T.
hemprichii is characterised by the lack of a ligule. To ensure
consistency of new leaves measurements, length was taken for dark
green tissue and starting at the base of the leaf curvature. The
plastochrone interval was calculated as the duration of the
experiment in days (13) divided by the number of new leaves per
shoot. The total number of new leaves as well as the ratio between
the number of new and old leaves were also calculated. We then
assessed whether there were significant differences in total leaf
length, plasto- chrone interval, number of new leaves, and ratio of
new to old leaves, between Fatumaru Bay and Pango Vale Vale.
This was done using the linear mixed-effects model, where location
was included as fixed effect and the individual quadrats as random
effect, using the statistical package RStudio (Version 1.1.383).
3. Results
3.1. Water quality parameters
The recordings of temperature and salinity are shown in Table 2.
Salinity values were lower in Fatumaru Bay (27.3–33.4 ppt) compared
to the other locations (30.0–35.1 ppt). The water temperature ranged
24.5–30.7 ◦C.
The concentration of dissolved inorganic nitrogen (DIN) was higher
in the seagrass bed in Fatumaru Bay (369 ± 24.88 μg N/L mean
standard deviation) compared to the other seagrass sites sampled (5–
162 μg N/L DIN range). Dissolved inorganic phosphorus (DIP) or
filterable reactive phosphorus (FRP) also appeared to be higher in
Fatumaru Bay (9.33 ± 2.08 μg P/L) compared to the other seagrass
sites
(3–8 μg P/L FRP range). Due to the very high concentration of DIN in
Fatumaru Bay, the N:P ratio was also higher compared to the other sites,
and exceeded the Redfield ratio 16:1. The high value of total
suspended solids (TSS) in Fatumaru Bay coincided with the sample
taken at the end of the survey (48 mg/L) as the tide began to flow into
the bay, which also
showed an elevated concentration of Chl-a (1.4–2.9 μg/L) compared to
the samples taken during the ebb and low tide times. The other four
sites
showed a decreasing trend in Chl-a concentration towards the
seagrass sites further away from Fatumaru Bay and Port Vila with the
lowest value (0.017 μg Chl-a/L) found in the samples collected in Pele
Island.

Table 2 3.3. In-situ seagrass growth measurements

Summary of water quality parameters collected during Cefas surveys including
temperature (T), salinity (S), total dissolved inorganic nitrogen (DIN = nitrites Different endpoints were evaluated during the growth experiment
+ nitrates + ammonium), filterable reactive phosphorus (FRP, also referred to and are shown in Fig. 4. New leaves length showed the highest
as dissolved inorganic phosphorus (DIP)), dissolved inorganic nitrogen to similarity between the two sites±(30.25 2.62 mm, mean standard error
dissolved inorganic phosphorus ratio (N:P), chlorophyll-a concentration (Chl-a)
in Fatumaru and 31.60 ± 2.47 mm in Pango; = t 0.17, p 0.87). Instead,
and total suspended solids (TSS) at the seagrass survey sites. Values indicate = Fatumaru and Pango
for the other endpoints, the differences between
mean (± standard deviation), with sample sizes indicated in brackets [].
were more noticeable: plastochrone interval (Fig. 4b), number of new
Parameters including T, S, DIN, DIP, N:P ratio and TSS were recorded or
sampled up to three times [n = 3] at each location during the survey. In the
leaves (Fig. 4c) and ratio of new:old leaves (Fig. 4d), although these
case of Chl-a, two separate samples were collected up to three times at each differences only approached statistical significance (p ≥ 0.05). Plasto-
location [n = 6]. chrone interval was higher in Fatumaru than Pango (8.91 ± 3.37 days
Seagrass T S DIN FRP N:P Chl-a TSS
survey (◦C) (ppt) (μg N/ or ratio (μg/ (mg/ Fatumaru, 7.79 ± 3.19 days Pango; t = —2.14, p = 0.08) and a
location L) DIP L) higher ratio of new to old leaves was observed in Pango (1.04 ± 0.06)
(μ g L) compared to Fatumaru (0.84 ± 0.43; t = 2.05, p = 0.09). The biggest
P/L) differences
were observed for the number of new leaves which were higher in
Pango
(± (± (± (± (± (± (± = 2.39, p = 0.05).
0.9) 1.3) 24.88) 2.08) 23.74) 1.06) 25.81)
[3] [3] [3] [3] [3] [6] [2]
Pango 29.6 34.8 9.67 5.00 4.71 1.73 7.75 4. Discussion
(± (± (± (± (± (± (±
0.9) 0.2) 1.53) 1.73) 1.91) 0.33) 1.06) To our knowledge, this is the first study that assesses the condition
[3] [3] [3] [3] [3] [4] [2]
of seagrasses in Vanuatu in relation to local water quality conditions,
Eratap 24.7 34.6 6.67 1.90 0.65 12.10
(± (± 5.67 (± (± (± (± using a combination of survey, sampling and in-situ experimental
0.2) 0.1) (± 0.58) 0.32) 0.05) 5.37) techniques. The condition of the seagrass varied highly across the
[3] [3] 0.58) [3] [3] [6] [3] studied sites in
[3] Efate Island in terms of canopy cover and canopy height suggesting that
Epau 27.2 32.5 13.67 7.33 5.96 0.13 7.03
(±
the meadows were generally patchy and composed of an assortment
(± (± (± (± (± (±
0.4) 0.2) 13.05) 0.58) 1.73) 0.06) 1.40)
of species and sizes. No correlation was found in terms of either
[3] [3] [2] [2] [2] [6] [3] distance from Port Vila or population density. Instead, canopy cover
and height
Pele 26.47 34.5 15.33 4.67 8.25 0.09 8.67 appeared related to site-specific characteristics such as sediment type,
Island (± (± (± (± (± (± (± sediment depth and shoreline position relative to tidal level (Waycott
0.4) 0.2) 4.04) 2.08) 3.66) 0.05) 5.05)
et al., 2011; Kilminster et al., 2015). The most compact meadows with
[3] [3] [3] [3] [3] [6] [3]
higher canopies were found in sheltered locations with softer and
deeper sediment, and below the mean low water level, such as Eratap
1.
Five species of seagrass were identified during the surveys: Thalassia

3.2. Results of field survey of intertidal seagrasses

 According to the results of seagrass canopy cover (%) and
seagrass canopy height collected, there was not a clear pattern across
the sites surveyed with high variability within each site. The least
dense meadows occurred in Fatumaru Bay ± (22 11%) and Epau (19
5%)
(Fig. 2).
Based on the canopy height measurements (Fig. 2), larger
±
seagrass shoots were found in Eratap 1 (31 12 cm; mean standard
deviation). The shorter seagrass were found±in Pango (4 2 cm), Pele
(3 0.4 cm) and± Epau (3 0.5 cm).
Table 3 represents the relative dominance of all five seagrass
species recorded at each survey sites, based on ranking their relative
abun- dances within each survey site. Thalassia hemprichii appeared to
be the most common species, as it was present in all 12 survey sites
and was the numerically dominant species in 7 of these. Following on
in order of general abundance was Cymodocea rotundata, which was
likewise pre- sent in all sites and numerically dominant in 4 sites.
Halodule uninervis was present in 6 out of 12 sites, but never dominant.
Enhalus acoroides was the species with the longest leaves of those
observed in Efate during the survey with some shoots measuring over
30 cm long, and was the dominant species in the Eratap 1 site.
Halophila ovalis was only found in Pele Island, were it was relatively
abundant. Across all sites examined, Pele Island was the richest site in
terms of seagrass species.
In Fatumaru Bay, seagrass leaves appeared generally paler and
brownish in colour, almost completely covered by epiphytes (Fig. 3).
The seagrass in the other sites showed some brown, burnt leaves,
particularly in Pango and Pele where some of the seagrass patches
were almost completely exposed during the low tide (Fig. 3). In
general, the seagrass leaves appeared greener in Pango, Eratap,
Epau and Pele Island compared to Fatumaru.
hemprichii, Cymodocea rotundata, Halodule uninervis, Enhalus acoroides
and Halophila ovalis. To date, thirteen species have been recorded in
the Vanuatu Islands; they include in addition to the five species from
this study: Cymodocea serrulata, Halodule pinifolia, Halophila capricorni,
Hal- ophila decipiens, Halophila minor, Ruppia maritima, Syringodium iso-
etifolium and Thalassodendron ciliatum (Chambers et al., 1990; Payri,
2011; McKenzie and Yoshida, 2017). In our study of Efate Island, the
richest sites in terms of number of species found was Pele Island.
Overall, T. hemprichii was found to be the most common species in
Efate Island, in the sense that it was the most numerous species in all
locations except Pango and Pele Island. This agrees with other obser-
vations from elsewhere in the tropical Pacific where either T. hemprichii
or C. rotundata appear to be the most abundant seagrasses in inshore
intertidal coastal areas (Waycott et al., 2011; Kilminster et al., 2015). T.
hemprichii, as well as E. acoroides, are amongst the tallest seagrass
species in the Indo-Pacific region and exhibit traits typical of long-lived,
robust and persistent species capable of tolerating a moderate level of
distur- bance (Kilminster et al., 2015). These two species contributed
to the taller canopy height found in Paonangisu and Eratap. Canopy
height was shorter in Pele Island, Pango and Epau, even though the
sediment layer appeared to be quite deep and therefore optimum for
seagrass growth. However, there was evidence of recent sea turtle
and dugong grazing (dugong sightings, dugong feeding trails and
clipped seagrass leaves) which may also be responsible for the shorter
canopy height at these locations.
In Fatumaru Bay, most of the seagrass patches exhibited brown
fragmented or decaying leaves, more densely covered with epiphytes
compared to the other sites. Nutrient over-enrichment could be an un-
derlying factor as this is known to stimulate overgrowth of epiphytes as
well as macroalgae, inducing a decline in seagrass condition
(Borkholder et al., 2007). Epiphyte growth was more widespread
across the seagrass beds surveyed in Fatumaru and although
macroalgae abundance was
Fig. 2. Summary of seagrass results showing seagrass canopy cover and canopy height. Closed circles represent mean values and error bars indicate standard de-
viation; n = 10 for each site.

Table 3
All five species of seagrass represented across all survey sites ranked by relative abundance, from the
most abundant (darker shade, higher number of *) to the least abundant but still present (lighter shade,
just one *). Blanks denote that a species was not found on a given site.
Seagrass survey Thalassia Cymodocea Halodule Enhalus Halophila
location hemprichii rotundata uninervis acoroides ovalis
Fatumaru ** *
Pango * **
Eratap 1 ** * ***
Eratap 2 *** * **
Epau ** *
Pele Island *** **** * **

not strictly quantified, macroalgae were observed to cover large partly explain the nutrient enrichment, however the nitrate
patches amongst the seagrass in Fatumaru (see Fig. 3) while concentrations
macroalgae pres- ence was much more sporadic in the other sites.
Elevated concentrations of DIN were found in Fatumaru Bay
(exceeding 350 μg/L), together with relatively high concentrations of
inorganic phosphorus (FRP or DIP). Nitrates are usually high in
coastal
waters that receive land drainage from industrial sources (Devlin et al.,
2015). A small industrial estate as well as a number of basic housing
areas and villas surround the shores of Fatumaru Bay, which could
appear exceedingly high compared to other concentration ranges
measured in Efate as well as other comparable regions of the South
Pacific and the Mediterranean (Schaffelke et al., 2012;
Papathanasiou and Orfanidis, 2018; Devlin et al., 2020). The values
of FRP or DIP in Fatumaru Bay appear similar to literature DIP
concentration ranges attributed to land runoff or riverine inputs of
suspended matter, particularly fine particles that are the conduit of
phosphorus (Devlin
et al., 2015). Salinity values were lower in Fatumaru Bay (27.3–33.4
ppt) compared to the other locations (30.0–35.1 ppt), a sign of fresh-
water input into the bay. It is likely that these high nutrient values
were
 Fig. 3. Examples of seagrass quadrats from each of the sampling locations.

Fig. 4. Results from T. hemprichii growth experiments in four quadrats in Fatumaru and four quadrats in Pango. Symbols indicate the means per quadrat, and the bars
represent the standard errors (SE). (a) Total length of new leaves; (b) plastochrone interval; (c) number of new leaves; (d) ratio of new to old leaves. Sample sizes: n =
81 shoots in Fatumaru, n = 78 shoots in Pango.

due to resuspension of the surface sediments and associated pore harbour area have re- ported poor water quality due to a combination of
water, where the nitrates build up from groundwater sources and coastal hydrodynamic
septic efflu- ents originated from the settlements that surround the
bay. Ground-
water inflows from septic leachates and other anthropogenic sources
can contribute up to 800 μM NO3 (Maier and Pregnall, 1990; Harman
et al., 1996; Burkholder et al., 2007), and nutrient concentrations are
often 10-
to 100-fold higher in the sediments than in the water column (e.g.
Sand- Jensen and Borum, 1991; Burkholder et al., 2007). Higher
values of TSS and Chl-a found in the sample taken during the flow tide
in Fatumaru suggest that, as the tide flows into the bay, fine muddy
sediments and benthic microalgae are resuspended causing transient
turbid conditions. Recent studies in Fatumaru Bay and Port Vila
circulation patterns that limits flushing in and out of the bay, and high
nutrient and TSS inputs from the surrounding area, where less than
25% of households have access to public plumbing facilities resulting
in raw domestic waste water being discharged into surrounding
streams or outfalls (Devlin et al., 2019; Devlin et al., 2020; Graham et
al., 2019). Concentration of Chl-a decreased at the sites further away
from Port Vila
(lowest concentration was 0.017 μg Chl-a/L, Pele Island) indicating low
phytoplankton and reduced nutrient load in these areas. Other studies
have previously indicated that in northern Efate, away from populated
coastal areas and particularly in exposed locations, the water quality is
good (Johnson et al., 2017).
The endpoints evaluated during the in-situ growth experiment
showed similar values between Fatumaru and Pango Vale Vale.

Nevertheless, differences for three parameters compared between the et al., 2020). Long-term monitoring and surveying over larger spatial
sites – new leaf production (p = 0.05), plastochron interval (p = 0.08) scales should also be conducted in order to improve understanding of
and ratio of new to old leaves (p 0.09) – approached statistical sig- the condition and distribution of these important habitats across the
nificance (p < 0.1), which was = suggestive albeit not conclusive of Vanuatu islands.
better
growth conditions in the more remote site. We note high variability CRediT authorship contribution statement
between replicates and a fairly limited sample size which may have
influence the statistical power of the growth experiment. Connell et al. Susana Lincoln, 27th November 2020
(2017) exposed seagrass plots to in-situ DIN enrichment and found an (lead author, lead scientist for this research and responsible for
abrupt shift from positive to negative net leaf production (from 0.04 leaf seagrass quadrat surveys, drafted and led the revision and editing of
per day production to 0.02 leaf per day loss). In our experiments, each draft)
growth was equivalent to the formation of 0.11 new leaves per day in Marta Vannoni
Fatumaru (or 1 new leaf in 8.91 days) and 0.13 new leaves per day in
Pango (or 1 new leaf in 7.79 days). The leaf production rate in
Fatumaru in partic-
ular, appears to be higher than the critical “tipping point” rate
suggested by Connell et al. (2017), which coincided with
concentrations of DIN
between 0.13 and 0.15 mg/L in their experimental enrichment. Future
research, including further in-situ growth experiments in Fatumaru Bay
and at other, more remote sites, where possible with enhanced sample
sizes, would help substantiate to what extent seagrass growth in the
area may be challenged.
Poor water quality, along with light limitation due to turbid condi-
tions, is generally considered one of the major drivers of seagrass loss
globally (Erftemeijer and Lewis, 2006; Waycott et al., 2009; Grech et
al., 2012; Unsworth et al., 2015) and in small Pacific islands
particularly (Brodie et al., 2020a, b). The relatively low values in terms
of % canopy cover observed in Fatumaru and Epau might not
exclusively relate to water quality but also to other environmental
characteristics of these sites, such as type of sediment, seagrass
species composition, exposure to waves and tides, and grazing (Short
et al., 2006; McKenzie and Yoshida, 2020). Eutrophication, which may
result from a lack of well-managed coastal development activities, has
the potential to cause dramatic los- ses in terms of coverage as well as
species richness to seagrass com- munities (Short et al., 2006).
The combined impact of warming and cyclones due to climate
change in tropical Pacific island countries such as Vanuatu is likely to
be more damaging in shallow coastal meadows compared to
seagrasses in
deeper areas (Waycott et al., 2011; Brodie and N’Yeurt, 2018). Where
seagrasses are already exposed to challenging local conditions of
water
quality, such as in Fatumaru Bay, this can undermine their resilience
and exacerbate their vulnerability to climate change stressors. The
loss of seagrass also threatens the important natural services they
provide (Hemminga et al., 1994; Duarte, 2002; Duarte et al., 2013;
Potouroglou et al., 2017; Nordlund et al., 2018; Paul, 2018).
This study supports findings of previous reports in terms of water
quality issues in the coastal waters near Port Vila in Efate, and
suggests that those seagrass communities are experiencing
challenging condi- tions due to high nutrient loads. In order to manage
and protect the seagrass habitats in Vanuatu, particularly those
located near populated areas, and preserve the important services
they provide, we recommend that water quality conditions, together
with seagrass condition, should be adequately monitored, and that
action should be taken, for example, to reduce untreated waste inputs
into coastal waters (Graham et al., 2019). The involvement of local
communities will be essential to the success of marine and coastal
management initiatives in small Pacific islands such as Efate (Johnson
 (responsible for seagrass growth experiments, reviewed and edited
this draft)
Lisa Benson
(responsible for overall seagrass survey design, reviewed and
edited this draft)
Georg H. Engelhard
(responsible for amends based on reviewers comments, including
reanalysis of water quality and seagrass data)
Dieter Tracey
(responsible for generating map-based figure used in this article)
Christina Shaw
(responsible for advising on survey areas and survey design)
Vatumaraga Molisa
(responsible for arranging field work and research permit for
this work)
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to
influence the work reported in this paper.
Acknowledgements
The authors thank The Vanuatu Department of Environmental
Pro- tection and Conservation for providing research permits to allow
this study. We thank the Secretariat of the Pacific Regional
Environment Programme, University of the South Pacific, Vanuatu
Department of Water Resources, Vanuatu Environmental Science
Society, Coasts Climate Oceans Consulting and their RESCCUE
marine champions, and the people from the coastal villages in Efate
for their help and support. Our appreciation goes to Len McKenzie,
Rodney Forster and Elisa Capuzzo for their advice. This study was
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Foreign and Commonwealth Office. We thank the two anonymous
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