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Can macroalgae contribute to blue carbon? An Australian perspective

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LIMNOLOGY
and
OCEANOGRAPHY Limnol. Oceanogr. 00, 2015, 00–00
C 2015 Association for the Sciences of Limnology and Oceanography
V
doi: 10.1002/lno.10128

Can macroalgae contribute to blue carbon? An Australian perspective

Ross Hill,*1 Alecia Bellgrove,2 Peter I. Macreadie,3,4 Katherina Petrou,3 John Beardall,5 Andy Steven,6
Peter J. Ralph3
1
Centre for Marine Bio-Innovation and Sydney Institute of Marine Science, School of Biological, Earth and Environmental
Sciences, The University of New South Wales, Sydney, New South Wales, Australia
2
Centre for Integrative Ecology, School of Life & Environmental Sciences, Deakin University, Warrnambool, Victoria,
Australia
3
Plant Functional Biology and Climate Change Cluster, University of Technology Sydney, Broadway, New South Wales,
Australia
4
Centre for Integrative Ecology, Deakin University, Burwood, Victoria, Australia
5
School of Biological Sciences, Monash University, Clayton, Victoria, Australia
6
Wealth from Oceans Flagship, The Commonwealth Scientific and Industrial Research Organisation, Dutton Park,
Queensland, Australia

Abstract
Macroalgal communities in Australia and around the world store vast quantities of carbon in their living
biomass, but their prevalence of growing on hard substrata means that they have limited capacity to act as
long-term carbon sinks. Unlike other coastal blue carbon habitats such as seagrasses, saltmarshes and man-
groves, they do not develop their own organic-rich sediments, but may instead act as a rich carbon source
and make significant contributions in the form of detritus to sedimentary habitats by acting as a “carbon
donor” to “receiver sites” where organic material accumulates. The potential for storage of this donated car-
bon however, is dependent on the decay rate during transport and the burial efficiency at receiver sites. To
better understand the potential contribution of macroalgal communities to coastal blue carbon budgets, a
comprehensive literature search was conducted using key words, including carbon sequestration, macroalgal
distribution, abundance and productivity to provide an estimation of the total amount of carbon stored in
temperate Australian macroalgae. Our most conservative calculations estimate 109.9 Tg C is stored in living
macroalgal biomass of temperate Australia, using a coastal area covering 249,697 km2. Estimates derived for
tropical and subtropical regions contributed an additional 23.2 Tg C. By extending the search to include
global studies we provide a broader context and rationale for the study, contributing to the global aspects of
the review. In addition, we discuss the potential role of calcium carbonate-containing macroalgae, consider
the dynamic nature of macroalgal populations in the context of climate change, and identify the knowledge
gaps that once addressed will enable robust quantification of macroalgae in marine biogeochemical cycling
of carbon. We conclude that macroalgal communities have the potential to make ecologically meaningful
contributions toward global blue carbon sequestration, as donors, but given that the fate of detached macro-
algal biomass remains unclear, further research is needed to quantify this contribution.

Anthropogenic activities since the industrial revolution,
such as the burning of fossil fuels, industrialization, defores-
tation and intensive agricultural practices, have rapidly
raised the atmospheric concentration of greenhouse gases
(Bala 2013). The increased abundance of these heat-trapping
gases has resulted in rising global temperatures and is dis-
rupting climatic processes around the globe. Carbon dioxide
(CO2) is the greatest contributor to greenhouse gas emissions
and is also responsible for causing ocean acidification
(Doney et al. 2009). One third of anthropogenic CO2 emis-
*Correspondence: ross.hill@mq.edu.au
sions dissolve into the oceans which slows the rise in the
atmospheric CO2 concentration, but this process reduces
ocean pH (Sabine et al. 2004). Over the last 250 yr, the CO2
concentration has increased from 280 ppm to over 400 ppm
(Bala 2013). Although this range of concentrations is not
unusual over geological timescales, the rate of CO2 increase
is faster than at any time in the last 300 million years
(Ho€ nisch et al. 2012). As a result, the rapid increase in CO2
concentration is having severe impacts on global climate
patterns (e.g., extreme weather events and loss of habitable
land due to sea level rise), ocean chemistry (e.g., reduction
in carbonate ion availability required for calcification),

1
Hill et al.
biological processes (e.g., hypercapnia, sensing of olfactory
queues), the functioning of ecosystems (e.g., bleaching and
death of corals), as well as human livelihoods (e.g., eco-
nomic loses from agricultural activities affected by drought,
reduced value of ecosystem services) (Guinotte and Fabry
2008; Doney et al. 2009; Hoegh-Guldberg and Bruno 2010;
Hilmi et al. 2013). While the rising CO2 concentration is
causing widespread problems, it is having a positive effect
on the photosynthetic rate of those terrestrial plants that are
otherwise carbon-limited (Long et al. 2004; Cornwall et al.
2012; Koch et al. 2013; Johnson et al. 2014).
During the 20th century, ocean temperatures rose by
0.748C and pH dropped by 0.1 units (equivalent to a 30%
increase in hydrogen ions) (IPCC 2007). Over the next cen-
tury, it is predicted global temperatures may rise by a further
48C and pH decline by a further 0.4 pH units (under the
higher emission A1F1 scenario (IPCC 2007)). Given the
severity of these impacts, mitigation of CO2 emissions is of
great importance. Carbon capture and storage technologies
engineered to sequester CO2 have been developed, but cur-
rently do not represent viable, affordable, long-term or pro-
ven solutions (Gough 2008; Markewitz et al. 2012; Rubin
et al. 2012).
Carbon sequestration through photosynthetic carbon fixa-
tion represents an alternative solution to removing CO2
from the atmosphere, and if the organic carbon is stored
long-term within the system, this represents a viable carbon
sink for mitigating CO2 emissions and climate change
(Duarte et al. 2013; Singh and Ahluwalia 2013). Natural ter-
restrial and aquatic carbon sinks are known to represent a
powerful means of storing carbon, with both mature (e.g.,
ancient forests or seagrass meadows) and immature ecosys-
tems (e.g., forest plantations) capable of building carbon
stocks, where rates of carbon gain through photosynthesis
exceed counteracting carbon losses from respiration (Grace
2004).
Recently, the term “blue carbon” was coined, referring to
carbon sinks in vegetated coastal ecosystems (Nellemann
et al. 2009). Typically, this term refers to the carbon seques-
tered in mangrove forests, seagrass beds, and salt marshes
(Mcleod et al. 2011). The focus of this review, however, is on
macroalgal populations of coastal ecosystems and the poten-
tial role they play as marine carbon sinks. As macroalgae pri-
marily grow on hard substrata, it has been assumed they do
not directly contribute to carbon sequestration due to the
absence of soft sediment that accretes as a result of organic
carbon deposition (Duarte et al. 2013). However, the export
of macroalgal detrital material to other ecosystems, such as
the continental margins (Dierssen et al. 2009), suggest they
may have the potential to act as significant carbon donors to
other long-term sequestration habitats. Australia has approx-
imately 60,000 km2 of coastline (Geoscience Australia 2010)
from tropical to cool temperate latitudes, where large stands
of macroalgae are a dominant feature of many coastal
2
Can macroalgae contribute to blue carbon?
regions. Here, we specifically address the contribution mac-
roalgae could play in carbon sequestration in the Australian
context; a region that possesses the greatest biodiversity of
macroalgae globally and high endemism (Phillips 2001). The
findings of this review, while based on data from the Austra-
lian coastline, are applicable to macroalgal communities
around the world.
Blue carbon definitions, assumptions and
requirements
Carbon sequestration is the capture and long-term storage
of carbon dioxide from the atmosphere through biological,
chemical or physical processes. Carbon accumulates in bio-
logical organisms as living biomass, but only a fraction of
that carbon becomes sequestered. Through photosynthesis,
organisms absorb and fix carbon dioxide incorporating it
into their cells for growth; however, more than 97% of this
fixed carbon is subsequently released back into the atmos-
phere either through plant respiration or the breakdown of
biomass by microbes (Farrelly et al. 2013). Therefore, while
more than 1550 Gt C is stored in the form of biomass, the
loss of carbon via these two pathways results in the net
sequestration of only  5 GtC yr21 (Raven and Falkowski
1999; Sabine et al. 2004; Lal 2005; Farrelly et al. 2013).
The two most significant sinks for long-term carbon stor-
age (after respiration and remineralization are accounted
for) are terrestrial soils and the deep ocean, with the total
stored carbon content estimated at 39,300 Gt (Farrelly
et al. 2013). The oceans are by far the largest carbon sinks,
estimated to hold around 37,000 Gt of inorganic carbon
(Raven and Falkowski 1999), with approximately 2 Gt of
additional carbon absorbed each year via physico-chemical
processes (Behrenfeld et al. 2002).
As with terrestrial soils, sequestered carbon in marine
sediments can remain over geological timescales in the form
of particulate organic carbon (known as blue carbon) unless
disturbed. In particular, disturbance that oxygenates other-
wise anoxic sediment can activate aerobic bacteria, which,
through aerobic metabolic pathways, may convert recalci-
trant carbon back to CO2 (Hartnett et al. 1998). Thus, a
change in the redox state of the sediment (from anoxic to
oxic) could potentially increase the efflux of carbon from
these natural sinks.
Ocean carbonate can also be stored for geological time-
scales in the form of magnesium and calcium salts (Raven
and Falkowski 1999). The oceans are by far the largest car-
bon sinks, estimated to hold around 37,000 Gt of inorganic
carbon (Raven and Falkowski 1999), with approximately 2 Gt
of additional carbon absorbed each year via physico-
chemical processes (Behrenfeld et al. 2002).
All signatories of the Kyoto Protocol are required to
reduce CO2 emissions either by decreasing the consumption
of fossil fuels or by increasing sequestration, particularly by
Hill et al.
terrestrial carbon sinks. More recently, consideration of
coastal systems as carbon sinks has led to increased investi-
gation into understanding carbon cycling and storage in
coastal systems (Pendleton et al. 2012). Carbon stock is
formed from a combination of autochthonous (derived from
primary producers within the habitat) and allochthonous
(derived from external sources) carbon. One measure of dif-
ferentiating sinks from sources is the use of a production :
respiration (P : R) ratio. Having a P : R greater than 1 is said
to be a carbon sink, removing and storing more carbon than
is released, while a ratio of less than 1 is deemed a net car-
bon source. However, in the case of some coastal systems
that have a P : R of less than 1, if significantly large amounts
of allochthonous carbon gets trapped and sequestered, the
system could in fact be a net carbon sink. It has already
been suggested that CO2 drawdown and assimilation by cul-
tured and wild-grown macroalgae represents a significant
sink for anthropogenic emissions (Chung et al. 2011).
Chung et al. (2011) argue that through the correct use of
harvested macroalgae, significant contributions to carbon
sequestration and offsetting of greenhouse gas emissions
would be possible. Their argument is strongly based on the
high productivity rates of macroalgae compared with higher
plants in the context of crops for biofuel production. How-
ever, in order for this offset to be effective, the carbon fixed
through macroalgal photosynthesis (i.e., the harvested bio-
mass) would have to be utilised in biofuels as a replacement
of fossil fuel use to ensure a net gain in the reduction of
total emissions; as no carbon is stored long-term in this sce-
nario. While Chung et al. (2011) demonstrates the potential
for carbon sequestration through macroalgal harvesting, we
focus here on the contribution of wild macroalgal popula-
tions to carbon sequestration in the oceans.
The carbon derived from the primary production of
coastal systems can be transported or donated to other eco-
systems in the form of particulate organic carbon (POC), dis-
solved organic carbon (DOC), and dissolved inorganic
carbon (DIC). Given the complexity involved in measuring
production and export processes, such as the amount of car-
bon that is donated, its impact on the recipient system, and
ultimately its long-term fate, a complete understanding of
macroalgae in the coastal marine system and the importance
in managing global carbon stocks is still lacking. For macro-
algal communities to be considered significant contributors
to blue carbon, they must either have the capacity to
directly store and accumulate carbon within their own habi-
tat, or transport biomass to receiver habitats where carbon
can be effectively buried and the organic material prevented
from undergoing microbial remineralization.
Like all major blue carbon habitats, community produc-
tion (P) by macroalgae generally exceeds respiration (R), thus
making them effective short-term CO2 sinks as carbon is
sequestered into organic matter. However, given that macro-
algae predominantly grow on rocky substrates, long-term
3
Can macroalgae contribute to blue carbon?
sequestration of this fixed carbon is considered less effective
than that of other blue carbon sources as there is less oppor-
tunity for the assimilated carbon to be accreted in anaerobic
sediments (Duarte et al. 2013). Therefore, understanding the
role of macroalgal stands as a donor system and the fate of
the carbon that is stored as biomass is of critical importance.
In this review, we aim to determine whether macroalgae
can contribute to blue carbon. We consider the potential for
direct storage of carbon within macroalgal habitats, as well
as the contribution macroalgae can make as a carbon donor
to receiver sites where allochthonous carbon accumulates. In
our calculations of carbon storage in macroalgal commun-
ities and comparison to other terrestrial and marine carbon
sinks, we specifically focus on the Australian continent,
though our conclusions are equally applicable to similar eco-
systems worldwide. The distribution, biomass and productiv-
ity of macroalgal communities are vulnerable to change and
we consider the impact of anthropogenic factors in influenc-
ing the ability of macroalgal habitats to contribute to blue
carbon into the future.
Methods
This article presents a review of the research literature on
blue carbon habitats, with a focus on Australian coastal mac-
roalgal communities. A literature search was conducted on
the electronic database, Web of Science, to identify publica-
tions on blue carbon, carbon sequestration, macroalgal dis-
tribution, macroalgal abundance, macroalgal productivity,
physiological and structural properties of macroalgae and
threats to marine communities, without restrictions on the
date of publication or geographic location of the study site.
Australian-based studies were targeted to gather data on
the distribution, abundance and productivity of macroalgal
communities to establish a comprehensive dataset on the
abundance, areal extent, depth range, biomass, biomass
turnover, growth rate, rate of maximum net primary produc-
tion, and total carbon stored in living macroalgae in distinct
regions of the Australian coastline. The total amount of car-
bon stored in temperate Australian macroalgae (Table 1) was
estimated from data of the most common assemblages. This
involved (1) summing published values of total carbon
stored in mixed understorey turfs with Ecklonia radiata and
mixed fucoid assemblages and extrapolated values of mean
total carbon stored in Durvillaea potatorum (based on fwt :
dwt ratio of 4.60 and published value of 0.312 gC g21 dwt;
Raven et al. 1989); (2) multiplying the summed value by the
area of coastal temperate waters to 60 m depth (Whiteway
2009); (3) multiplying the product of (2) by two estimates of
the percentage of rocky reef conducive to supporting macro-
algal populations in temperate Australia based on published
data and a more conservative estimate. Similar calculations
were made for tropical Australia based on the scant data
available for the Great Barrier Reef, Queensland. Further
Table 1. Contributions of macroalgae to short-term carbon sequestration throughout Australia, based on available data. Values represent those as published in the literature,
except where otherwise indicated by superscripts and footnotes to explain the derivation of extrapolated values.

Total C
Max net stored
Biomass Growth production in living
Abundance Areal Depth Biomass turnover Growth mg g rate macroalgae
Region Latitude Species % cover extent km2 range m g m22 times yr21 rate % d21 FW21 d21 g C m22 d21 Mg C km22 References
0
Northern GBR 11830 S– Halimeda spp. 20–40 4637 max Drew and
158350 S meadows 525 mean Abe (l 1988)
Northern GBR 148270 S– Halimeda 26% shelf in 2000 (up to 20–>50 Marshall and
158020 S mounds/bioherms northern 19 m thick) Davies (1988),
section Orme and
of GBR Salama (1988)
Great Palm, 188090 S– Sargassum 2–3 190 (AFDW) 4 40 3 Schaffelke and
Fantome 188410 S baccularia Klumpp (1997)
and Brook
islands, GBR
Davies Reef, 18844 Epilithic algal 20 3.9 42.2† Klumpp and
GBR community McKinnon
(crustose (1989, 1992)
corallines)
GBR 158S, 198S, Epilithic 60–80% reef 5–20 45 max 5.6 2.5 25.3† Klumpp and
258S algal substrate 30 mean McKinnon
community on GBR (1989, 1992)
(turfs)
Shark Bay, 268 Drift algae 152 (dwt - in Kendrick
Western (predominantly Amphibilos et al. (1990)
Australia Laurencia spp., antarctica bed)
Vidalia spiralis, 223 (dwt -
Digenia simplex in Posidonia
and Eucheuma australis bed)
speciosum)
Mooloolaba, 268400 S Ecklonia radiata 0 Connell and
Qld and subtidal 57 Irving (2008)
fucoids Turfs
(multispecies)
Houtman 288S–298S Ecklonia radiata 15 20–40 1040 (dwt) 0.3 Hatcher
Abrohlos Red foliose 15 1–50 ND et al. (1987),
Islands, algae Crustose 3 Smale
Western corallines et al. (2010)
Australia
Ballina, NSW 288520 S Ecklonia radiata 84 Connell and
and subtidal 15 Irving (2008)
fucoids Turfs
(multispecies)
TABLE 1. Continued
Total C
Max net stored
Biomass Growth production in living
Abundance Areal Depth Biomass turnover Growth mg g rate macroalgae
Region Latitude Species % cover extent km2 range m g m22 times yr21 rate % d21 FW21 d21 g C m22 d21 Mg C km22 References
0
Rottnest Island 32842 S Ecklonia radiata 30 1–50 Smale
Scytothalia 4 1–50 et al. (2010)
dorycarpa Red 6 1–70 ND
foliose algae 12
Crustose
corallines
Southern 338050 S– Ecklonia radiata 90 (WA and SA) 20m 18,000 6 (SA 0.9 (WA 9 (WA 6 (SA 635 Kirkman (1984, 1989),
Australia 358360 S and subtidal 41 (NSW) (wwt – fucoids) E. radiata) E. radiata) fucoids) (SA)* Cheshire et al. (1996),
Coast fucoids WA Connell and
E. radiata) Irving (2008)
Southern 338050 S– Turfs 6.5–8 (WA and SA) to 4 and 10 m for 88–100 29–79 2.9 (SA) 32–36* Copertino
Australia 358510 S (multispecies) 40 where canopy productivity (AFDW) et al. (2005),
Coast restricted measures Connell and
(SA) 17 (NSW) Irving (2008)
Southern 338050 S– Crustose 0 (WA and SA) Connell and
Australia 358360 S coralline 38 (NSW) Irving (2008)
Coast barrens
South eastern 358S– 438S Durvillaea 0–5 22,500/4891mean 60.3§ 1526*,‡ Cheshire and
Australia potatorum 108,000/23,478 Hallam (1988),
max (wwt/dwt* Raven et al. (1989)
respectively)
Victoria 388110 S– Dominant 34.1 377 10–60 Ierodiaconou
(9% of Vic 388520 S macroalgal et al. (2007, 2011),
coastline) assemblages Rattray et al. (2009),
(predominately Che Hasan
phaeophytes et al. (2012),
and rhodophytes) Monk et al. (2012),
Rattray et al. (2013)
Robins Passage 408420 S– Durvillaea 10–50 0–5 Lucieer
to Swan Island, 418120 S potatorum 60–100 0–25 et al. (2007b,)
north coast Mixed E. radiata 40–60 15–30
Tasmania and fucoids
Mixed turfs
Tasmania – 418S– Macrocystis  11 (1940) 2 mm d21 2400 g Sanderson (1990),
eastern coast 438360 S pyrifera 13.11 (345 beds, 1999) wwt m22 yr21 Edyvane (2003),
< 1 (2010) Johnson et al. (2011)
TABLE 1. Continued
Total C
Max net stored
Biomass Growth production in living
Abundance Areal Depth Biomass turnover Growth mg g rate macroalgae
Region Latitude Species % cover extent km2 range m g m22 times yr21 rate % d21 FW21 d21 g C m22 d21 Mg C km22 References
0
St Helens Point 41817 S– Total algae 70–100 0–25 Ecklonia: Sanderson (1990),
to Schouten 428S (predominantly 4700 g Lucieer
Island, Phyllospora wwt m22 yr21 et al. (2007a)
SE Tasmania comosa and Phyllospora:
E. radiata) 7400 g
wwt m22 yr21
High Yellow 438S– Phaeophyceae 43–48 20–40 Lucieer
Bluff to Cape 438300 S (predominantly 46–10 20–30 et al. (2012)
Hauy, George E. radiata) 7 to 40–50
III Rock, Rhodophyta 20–30
SE Tasmania Chlorophyta

*Values used to estimate carbon storage associated with biomass of south-eastern Australian subtidal macroalgae.
†
Values used to estimate carbon storage associated with biomass of EAC on the Great Barrier Reef.
‡
Extrapolated values from published data based on fwt : dwt ratio of 4.60 gC g21 dwt and 0.312 gC g21 dwt (Raven et al. 1989).
§
Extrapolated values from published data and assuming maximal photosynthetic rate for 12 h d21 (Raven et al. 1989).
Hill et al. Can macroalgae contribute to blue carbon?

Table 2. Organic carbon stored both above and below ground in terrestrial forests and different marine ecosystems in Australia

Carbon stored (Mg C km22) Area in Total C

Australia stored in
Ecosystem Above ground Below ground Total (km2) Australia (Tg C) References
Terrestrial forests 5700–21,700 6200–23,000 11,900–44,700 1,494,000 17,778.6–66,781.8 Malhi et al (1999),
Fourqurean
et al. (2012),
Department of
Agriculture (2013)
Saltmarsh 67–922 207–12,972 274–13,894 13,029 3.6–181.0 Caçador et al. (2004),
Howe et al. (2009),
OzCoasts (2013)
Mangrove 9750–31,724 5731–67,816 16,792–99,540 3976 66.8–395.8 Twilley et al. (1992),
Howe et al. (2009),
Fourqurean
et al. (2012),
OzCoasts (2013)
Seagrass 0.1–555 0.1–83,678 0.2–84,233 296 (temperate 5.92 3 1025–24.9 Fourqurean et al. (2012),
Australia) 4.30 3 1023–1809.5 Waycott et al. (2009),
21,482 (all OzCoasts (2013)
of Australia)
Australian 2200 0 2200 49,939–124,849 109.9–274.7 This study
temperate
macroalgae

details and justification for these choices are given in the
next section.
Studies from around the world were used for all other pur-
poses of the review and informed the findings from the Aus-
tralian continent, in terms of the potential for above- and
below-ground carbon storage across a range of habitats com-
parable to macroalgal communities, which are known for
their efficacy in carbon sequestration (terrestrial forests, salt-
marshes, mangrove forests, and seagrass beds). Total carbon
stored in Australia was calculated for each habitat from data
reported in the literature (Table 2). Minimum and maximum
values were calculated by multiplication of the carbon stor-
age capacity per unit area of each habitat type, by the area
of that habitat found in Australia. Studies from around the
world were also used to provide a rationale for the review
and contributed to the general discussion to provide a global
perspective on calcification, decomposition, and the influ-
ence of anthropogenic impacts on the blue carbon capacity
of macroalgae.
Potential for macroalgae to directly store carbon
Macroalgae are known to have very high rates of primary
productivity and growth and significant biomass on coastal
margins (Mann 1973; Chung et al. 2011), particularly in
temperate regions (Mann 1973; Cheshire et al. 1996; Connell
and Irving 2008). Macroalgae are typically associated with
hard substrata to which they attach and grow, obtaining
their nutritional needs from the surrounding water rather
than via vascular root systems penetrating the sediment. As
such, and unlike other aquatic macrophytes, their biomass is
all “above ground,” with the exception of calcifying algae
that can build up significant calcium carbonate deposits in
the sediments (e.g., Halimeda in Table 1 and below) and sto-
loniferous taxa such as Caulerpa. The capacity for noncalcify-
ing macroalgae to directly store carbon is thus largely
associated with the above ground living biomass and how
long the carbon within living biomass remains fixed before
being remineralized by bacteria.
Australia has an extensive coastline with about 30% in
the temperate coastal zone dominated by large macroalgae
such as Durvillaea potatorum (found 0–5 m deep), Ecklonia
radiata and mixed fucoids (occurring down to 40 m depth)
including Phyllospora comosa, Cystophora spp., Sargassum spp.
(Table 1) and Scytothalia doryocarpa in the west (Wernberg
et al. 2003). However for most of the Australian coast, data
on areal extent, biomass and productivity of macroalgae are
very fragmented and often reported in non-standardised
units (Table 1). Moreover, there are few records of standing
stock of carbon, or carbon contents of particular species that
allow extrapolation from available biomass data. Thus, accu-
rate assessments of the total amount of carbon stored in

7
Hill et al.
living biomass of macroalgae in Australia are, at this stage,
difficult to determine. We estimate, however, that the coastal
waters of southern Australia, with D. potatorum in the upper
subtidal and E. radiata monospecific and/or mixed fucoid
dominated assemblages to 30 m that support an understorey
of turfing algae that may extend to 60 m in depth, could
store in excess of 2200 Mg C km22 (Table 1). If we extrapolate
to the temperate coastal waters of Australia < 60 m depth
(249,697 km2, Whiteway 2009) and assume that approxi-
mately 50% (124,849 km2) of coastal waters are rocky reef
(based on data compiled from Victoria, Australia in Ierodiaco-
nou et al. 2007; Rattray et al. 2009; Ierodiaconou et al. 2011;
Che Hasan et al. 2012; Monk et al. 2012; Rattray et al. 2013)
we estimate 274.7 Tg C may be directly stored in living mac-
roalgal biomass in temperate Australia (Table 2). The above
studies targeted reef areas, but there are large expanses of
sandy sediments in temperate Australia (e.g., Ninety-mile
beach, Victoria); so if we take a more conservative estimate
of 20% (49,939 km2) reef to 60 m, we estimate 109.9 Tg C
stored in living macroalgal biomass in temperate Australia
(Table 2). For tropical and subtropical Australia data are even
sparser, but based on estimates of carbon storage by epilithic
algal communities, the Great Barrier Reef may store 23.2 Tg C
in epilithic algae alone (67.5 g C m22 3 344,400 km2; Table 1).
The capacity for carbon storage per unit area in macroalgae is
generally lower than for other blue carbon habitats and ter-
restrial forests due to the absence of below ground biomass
(Table 2). However the vast extent of temperate macroalgae
communities makes total carbon storage comparable to the
other blue carbon habitats (saltmarsh, mangroves, and sea-
grass beds) which have both above and below ground carbon
storage (Table 2). We of course acknowledge that there are
temporal and spatial variability in productivity, biomass and
carbon content of different species (Table 1) and thus these
values are crude estimates only. New data on aerial extent,
biomass, and carbon storage for algal species and assemblages
from around Australia will improve our estimates. Compara-
ble data for other regions are needed for reliable global esti-
mates of carbon storage in macroalgal living biomass.
Seasonality in alternation of generations, fragmentation
and dislodgment (Thomsen et al. 2004; McKenzie and Bell-
grove 2009; Krumhansl et al. 2011) of macroalgae all con-
tribute to high turnover of macroalgal biomass. High
productivity and growth rates of macroalgae replenish the
biomass. There are however, no comprehensive data on the
spatial and temporal variability in biomass of any macroalgal
species in Australia (references in Table 1). Assuming rela-
tively stable annual macroalgal standing stock, productivity,
growth and turnover do not affect the potential for macroal-
gae to directly store carbon in the above ground biomass.
High productivity will not result in more carbon storage
unless it equates to greater total biomass. However, high
rates of productivity, growth and turnover do become impor-
tant to carbon-storage potential of macroalgae if we consider
8
Can macroalgae contribute to blue carbon?
their possible role as carbon donors to other systems, which
is a function of their decay rate en route to receiver sites, the
burial efficiency of the receiver site, and their structural com-
plexity (i.e., susceptibility to being remineralized). We
explore this role in more detail below (see “Macroalgae as
carbon donors”).
Calcification in macroalgae
All macroalgal groups contain genera that form calcium
carbonate deposits and calcified algae are found in the
photic zone of all marine habitats from polar to tropical seas
(Nelson 2009). In the Phaeophyceae, calcification is most
notable in Padina, though a few other brown algae also cal-
cify. The calcifying Chlorophyte algae are mostly tropical
and contain genera such as Acetabularia, Neomeris, Avrainvil-
lea and, most importantly in terms of its contribution, Hali-
meda. Many red algae are calcifying, both in tropical and
cold waters, but calcification is best known in the Coralli-
nales with both geniculate and nongeniculate or crustose
forms. Free-living nongeniculate corallines are known as rho-
doliths and form extensive beds, which are harvested in
Europe and elsewhere (e.g., Brazil). In most cases the calcium
carbonate is found as the mineral aragonite, either deposited
at the cell surface (e.g., Padina) or in intercellular spaces
(e.g., Halimeda). Calcium carbonate can also be deposited as
calcite, which has a different crystal structure to aragonite,
making it more stable and less prone to dissolution in sea-
water, especially under the lowered carbonate concentrations
likely to be found with increasing ocean acidification in the
future. Calcite is found in external crusts on Chaetomorpha
and within the cell walls of coralline genera such as Litho-
phyllum and Lithothamnion (Lobban and Harrison 1994).
Many calcifying algae also grow epiphytically on seagrass
leaves, as well as on other marine organisms, including
bivalves and molluscs. Indeed, seagrass photosynthesis can
promote calcification rates by other organisms, such as Hali-
meda, growing in seagrass beds (Semesi et al. 2009).
Although many of the algae that calcify are slow growing,
the calcium carbonate (aragonite, and especially calcite) can
persist in the marine environment long after the living cells
have decayed. Iconic calcium carbonate deposits such as the
White Cliffs of Dover are the remains of marine calcifying
microalgae deposited over geological time. The rhodolith
beds in France and Brazil are harvested to the rate of
500 Gg yr21 (Briand 1991) and up to 120 Gg yr21 (Riul et al.
2008), respectively, for industrial processes including agricul-
ture, water purification, mineralization and the manufacture
of cosmetics (Grall and Hall-Spencer 2003). Carbonate depo-
sition (accumulation) in coastal waters (including coral reefs)
globally is in the order of 1.2 Pg CaCO3 yr21 ( 0.17 Pg C yr21)
with approximately the same accumulated in the pelagic
ocean, though production rates in the open ocean are closer
to 1.1 Pg C yr21 (Iglesias-Rodriguez et al. 2002). Decomposed
Halimeda tissues are important sediment elements in tropical
Hill et al.
regions (Table 1), and contribute the major part of the cal-
careous sediment found in the sandy beaches of many tropi-
cal shores. Halimeda bioherms alone have been estimated to
contribute 400 Tg CaCO3 yr21 to CaCO3 accumulation (Hillis
1997). Rhodolith forming species can produce in the region
of 60–1000 g CaCO3 m22 yr21, with CaCO3 deposits off Bra-
zil’s coast alone of 2 Pg (Riul et al. 2008; Nelson 2009).
Importantly, carbonates deposited by oceanic organisms
have not previously been included in blue carbon budgets.
This is because the calcification process (Eq. 1) results in the
release of 1 CO2 for every CaCO3 formed.
21
2 HCO2
3 1Ca ! CaCO3 1CO2 1H2 O (1)
However, the CO2 produced in this way can be rapidly
utilised in photosynthesis and even if it is not (during calcifi-
cation in the dark for instance) there is still the net conver-
sion of one soluble HCO2 3 into one, insoluble, carbonate
(i.e., calcification still results in net drawdown of carbon in
the recalcitrant carbonate minerals aragonite or calcite).
Given the significant contribution of macroalgal calcifica-
tion to the global C budget and the persistence of CaCO3 in
sediments, we would argue that calcification should be con-
sidered in any calculations of blue carbon into the future. If
included, this would constitute a further 0.34 Pg C yr21,
sequestered globally.
Macroalgae as carbon donors
So far we have established that most macroalgal species
have limited capacity to act as long-term carbon sinks in
their own right (Table 2), largely due to their inability to
accumulate below-ground stores of carbon relative to other
vegetated coastal habitats such as seagrasses, saltmarshes,
and mangroves. However, even though macroalgae do not
develop significant carbon deposits themselves, there still
exists the possibility that they could make significant contri-
butions to global carbon sequestration by acting as “carbon
donors.” In this section, we explore this possibility.
What is a “carbon donor”? Here we define a carbon donor
as an autotroph that “donates” carbon to another “receiver”
habitat that ultimately buries that carbon. Therefore, carbon
donors make indirect contributions to carbon sequestration.
Such movement of carbon from one system to another is
referred to as “leakage” (or “spillover”). A well-recognised
example of leakage can be seen in seagrass ecosystems. Sea-
grasses have a high particle-trapping capacity (Agawin and
Duarte 2002; Gruber and Kemp 2010), and due to their
occurrence at the interface between land and sea, they cap-
ture large amounts of terrestrial carbon that runs off the
land following rainfall events (Kennedy et al. 2010; Macrea-
die et al. 2014). This externally derived carbon that leaks
into a receiver habitat is termed “allochthonous” carbon. In
seagrasses, allochthonous carbon can contribute as much as
9
Can macroalgae contribute to blue carbon?
half the total organic carbon that is sequestered in a meadow
(Kennedy et al. 2010).
From a carbon accounting perspective, leakage of carbon
is a complex issue. Can habitats take credit for carbon that
they didn’t produce? What happens if the carbon was
already credited to another system (double-counting)? How
can the origin of carbon be determined? These questions are
at the forefront of many terrestrial carbon-offset programs
(e.g., REDD), but have received comparatively little attention
in the aquatic fields. The matter of carbon accounting of
leakage will, however, become increasingly important with
the rising prominence of blue carbon, but is beyond the
scope of this review. We recommend Murray et al. (2007) for
more information on leakage.
For a system to be a significant carbon donor, there are
three requirements: (1) high rates of biomass production; (2)
effective transfer of biomass to receiver habitats; and (3) the
donor carbon must undergo efficient burial of allochthonous
biomass within receiver habitats, such that it evades micro-
bial attack (the latter is a function of several factors, includ-
ing: the intrinsic stability of the carbon, physical
mechanisms of protection, and environmental conditions
that dictate microbial activity). All three requirements are
necessary; none are redundant or substitutable. The first
requirement has already been discussed earlier in this article,
and can be summarised as follows: macroalgae have excep-
tionally high biomass production; their global extent (2–6.8
million km2) and primary production (0.19–0.64 Pg C yr21)
exceeds that of all the vegetated coastal habitats (seagrass
beds, saltmarshes, mangroves) combined (Duarte et al.
2013). In the following section, we will discuss requirements
2 and 3, which concern the export and fate of this biomass
production.
Export and fate of macroalgal biomass
to receiver habitats
Most macroalgal species spend their early life history
€ ning 1990), but then
attached to benthic (hard) substrata (Lu
become dislodged or fragmented and can undergo a period
of transport in surface or bottom waters by wind and water
movement (potentially as rafts that travel significant distan-
ces), until eventually sinking, disintegrating, or becoming
cast upon land (Thiel 2003; Thiel and Gutow 2005; Macrea-
die et al. 2011). Detachment of macroalgal tissue can arise
from natural senescence (Ang 1985), dislodgement by herbi-
vores (Kingsford 1992; Tegner et al. 1995), and through
hydrodynamic forces such as waves and storms (Duggins
et al. 2003). The supply, quality, persistence, and transport
(trajectory and duration) of macroalgal rafts are all influ-
enced by climate (Macreadie et al. 2011), particularly tem-
perature (Graiff et al. 2013). Thus, whilst there is temporal
variation in the extent of macroalgal rafts related to these
processes, there is continuous export of macroalgal biomass
throughout the year, whereas saltmarsh and seagrass detrital
Hill et al.
export tends to be seasonal (Josselyn and Mathieson 1980).
Furthermore, acknowledging spatial variation in the export
of macroalgal and vascular plant biomass, macroalgae are
often more significant contributors to surf-zone and beach
wrack than seagrasses (Josselyn and Mathieson 1980; Gomez
et al. 2013; Baring et al. 2014; Ruiz-Delgado et al. 2014).
From genetic and tracking studies, we know that exported
macroalgal biomass can be transported long distances (Thiel
and Haye 2006; Fraser et al. 2009, 2010, 2011; Nikula et al.
2010), but there is limited information on what happens to
carbon bound within macroalgal biomass once rafts or frag-
ments sink and disintegrate—i.e., the point at which carbon
bound within macroalgal biomass is broken down into dis-
solved and particulate organic carbon. Wada et al. (2007)
suggested that macroalgae could release 20–40% of their pro-
ductivity as dissolved organic matter (DOM). Much of the
macroalgal DOM is likely to be microbially degraded, but
estimates of the refractory component of this DOM will be
important for understanding how much could be donated to
other habitats. UV is known to degrade DOM; this means
that if this matter can be deposited in regions away from
light such as the deep ocean, the breakdown will be further
diminished.
There is some evidence to suggest that macroalgal carbon
is donated to other receiver habitats that have long-term
sequestration capacity, including the deep sea (Wada et al.
2008; Dierssen et al. 2009) and other vegetated coastal habi-
tats, such as mangrove forests, seagrass beds, and salt
marshes (Hyndes et al. 2012; Macreadie et al. 2012). In tropi-
cal estuaries and bays, it is estimated that about half of the
macroalgal biomass produced is consumed by herbivores and
the other half is exported to other habitats, whereas in the
temperate open coast most of the biomass is decomposed by
detritivores, with a lesser amount being exported, and only
small fractions being consumed or accumulated as detritus
(Hyndes et al. 2013). The fate of macroalgal biomass in tem-
perate estuaries and fjords is largely unknown (Hyndes et al.
2013). Most export of macroalgal biomass occurs via advec-
tion, but there are also complex biotic pathways that trans-
port macroalgal biomass into receiver habitats; such as via
ingestion and concomitant faecal deposition of carbon
within receiver habitats by fish, which can contribute to car-
bon sequestration by fertilising receiver habitats and thereby
increasing their productivity and sequestration capacity
(Valiela 1995; Hyndes et al. 2013). Breakdown of macroalgae
has been shown to be mostly mediated by microbial proc-
esses (Rieper-Kirchner 1990); however protozoans and meio-
fauna also contribute to the decomposition of macroalgal
tissues as well as enhance microbial degradation. Microbial
breakdown rate is also dependent upon the sediment com-
position and oxic status of the water. Sassi et al. (1988)
found that macroalgal decomposition rates over 25 d under
aerobic vs. anaerobic conditions were 35% and 24%, respec-
tively. Therefore, if the macroalgal detritus can be advected
10
Can macroalgae contribute to blue carbon?
to a rapidly sedimenting region where it can be incorporated
into an anaerobic sediment, this will slow the microbial
breakdown and maintain the carbon within the sediment
longer. More research is needed to understand how long this
carbon actually remains within anaerobic sediment.
All macroalgal carbon that enters the sedimentary carbon
pool will eventually be broken down (oxidised and reminer-
alized) by microbes and converted to inorganic forms and
potentially returned to the atmosphere. The rate and effi-
ciency of this remineralization process can dictate whether
macroalgal carbon remains bound within sediments for
months or millennia. Macroalgal carbon entering the sedi-
mentary pool has three possible fates: (1) particulate and dis-
solved organic carbon that is recalcitrant to microbial
degradation is likely to remain stored within the sediment
for prolonged periods, while labile carbon will be recycled by
heterotrophic bacteria and is either (2) assimilated into
microbial biomass or (3) respired as CO2 back to the water
column and possibly the atmosphere. The proportion of
nonrespired carbon represents the amount of macroalgal car-
bon that is sequestered.
It should be recognised that macroalgae lack the struc-
tural polysaccharide complexity of higher plants, and conse-
quently their decomposition rates are faster (Sassi et al.
1988). To better understand the degradation pathways of
macroalgal plant material within aquatic sediments,
researchers have used several techniques. Analyses of ele-
mental composition and stable isotope signatures are rou-
tinely used to “fingerprint” carbon and determine its origin,
although there is difficultly in separating microalgal and
macroalgal organic carbon signatures (Hardison et al. 2011).
Furthermore, isotopic signatures may also be variable in
space and time during the decomposition process (Fenton
and Ritz 1988). Similarly, a problem with identifying sources
of organic carbon based on carbohydrate composition is that
many groups of organisms contain the same carbohydrates
(Vichkovitten and Holmer 2004). Fatty acid signatures have
proven to offer better resolution of the different plant contri-
butions to organic matter (Hardison et al. 2013), although
the level of complexity using this technique is much higher
(Graeve et al. 2002). Manipulative approaches using isotopic
tracer studies (Hee et al. 2001; Hardison et al. 2010, 2011;
Hyndes et al. 2012) and radiolabelling (Chidthaisong et al.
1999) have proved to be robust for studying the incorpora-
tion of macroalgal carbon into sediments.
Reliable data on the fate of decomposed macroalgal
carbon within sediments (e.g., macroalgal carbon that is
“lost” during litter bag mass loss experiments) is scarce.
Hardison et al. (2010) traced carbon from senescing macroal-
gae (Gracilaria spp.) that were pre-labeled with the 13C stable
isotope and found that 2–9% of macroalgal carbon was
incorporated into the sediment. This percentage of macroal-
gal carbon uptake by sediments is relatively high, suggesting
efficient burial; however, the fate of this buried carbon was
Hill et al.
only monitored for 2 weeks, so it is not known whether this
carbon stock was sequestered long-term. Other studies sug-
gest that macroalgal carbon is not sequestered long-term. For
example, Garcia-Robledo et al. (2008) found that macroalgal
detritus (tubular and planar Ulva spp.) that entered sedimen-
tary organic carbon pools vanished within days after its ini-
tial appearance, indicating that carbon from such simple and
ephemeral macroalgae is not retained for durations that
would make them an important component of the long-
term carbon pool within a receiver habitat. However, given
the diversity in the cell walls and carbon chemistry of mac-
roalgae, it is difficult to make conclusions on likely persist-
ence of sequestered macroalgae based on only a handful of
studies.
Kristensen (1994) compared the decomposition rate of six
different plant materials (two macroalgae, two seagrasses,
and two tree leaves) using stepwise thermogravimetry and
found that macroalgae (Fucus vesiculosus and Ulva lactuca)
lost 40–44% of their carbon within 70 d, whereas seagrasses
(Zostera marina and Ruppia maritima) lost 29–55%, and tree
leaves (Fagus sylvatica and Rhizophora apiculata) lost only 0–
8%. Similarly, Josselyn and Mathieson (1980) found that
macroalgae (Ascophyllum nodosum and Fucus vesiculosus)
decomposed 3–10 times faster than a saltmarsh plant (Spar-
tina alterniflora) and seagrass (Zostera marina). Litter bag stud-
ies have reported a lack of change in sediment carbon
following decomposition of buried macroalgae, suggesting
that the organic carbon bound within macroalgal structural
material (e.g., carbohydrates) is highly labile and rapidly
decomposed by microbes (Vichkovitten and Holmer 2004).
Despite this general pattern of high degradability of macroal-
gae compared with aquatic angiosperms (Trevathan-Tackett
et al. in press), there is a high degree of diversity in the
structural complexity of macroalgae, particularly in cell-wall
structure and carbon chemistry, resulting in high variability
in decay rate estimates within the literature (Rice and Han-
son 1984; Twilley et al. 1986; Kristensen 1990; Wada et al.
2008; Trevathan-Tackett et al. in press).
Nedzarek and Rakusa-Suszczewski (2004) examined the
decomposition of 10 Antarctic macroalgal species finding
they released 80% of the nutrients with the first 3 d of a 69
d decomposition experiment; these rates occurred at polar
temperatures (48C), so it would be assumed that tropical
decomposition would be more rapid. Similar results were
found with North Sea macroalgae where 50–80% of the tis-
sue biomass was lost within 14 d at 188C (Rieper-Kirchner
1990) which included the brown leathery kelp Laminaria
saccharina.
Factors influencing the future carbon storage and
donor ability of macroalgae
This review presents a static view on the current state of
carbon cycling, sequestration and donor capacity of macroal-
11
Can macroalgae contribute to blue carbon?
gae, with an emphasis on Australian coastal systems. In real-
ity, the distribution, biomass and productivity of these
macroalgae are vulnerable to change due to anthropogenic
activities and as such, the dynamic nature of these macroal-
gal systems should be taken into consideration. Factors such
as climate change (Duarte et al. 2013) and urbanisation (Air-
oldi and Beck 2007; Airoldi et al. 2008) can greatly alter the
distribution, abundance and productivity of these systems,
although the rate of change in global macroalgae population
size remains unknown.
Climate change can greatly influence the survival, photo-
synthetic capacity, growth and competitive interactions of
macroalgae in coastal habitats (Wernberg et al. 2011b).
Ocean warming on both the east and west coasts of Australia
have caused pole-ward movement of macroalgal species over
the past five decades (Wernberg et al. 2011a). Potentially,
hundreds of species are expected to be driven beyond the
edge of the Australian continent as warming continues, driv-
ing not only local, but global extinctions, as 62% of macro-
algal species found in southern Australia are endemic to the
region (Phillips 2001). On the east coast of Australia, there
has been a southern retreat of three habitat-forming macro-
algal species (Ecklonia radiata, Phyllospora comosa and Durvil-
laea potatorum; Millar 2007), and in eastern Tasmania, the
once common Macrocystis pyrifera has almost completely dis-
appeared with ocean warming (Edyvane 2003; Johnson et al.
2011).
In addition to rising temperatures, climate change is
expected to influence macroalgal distribution and productiv-
ity through ocean acidification, emergence of marine dis-
eases, and shifts in the distribution of consumers. In
terrestrial ecosystems, there have been climate-driven
increases in global net primary production due to the easing
of critical climatic constraints, such as increasing solar radia-
tion from reduced cloud cover (Nemani et al. 2003). In the
oceans, while increased CO2 dissolution is reducing pH and
disrupting carbonate chemistry, this inorganic carbon is a
building block for photosynthesis. The predicted changes in
CO2 in the atmosphere over the next 100 yr will translate to
a 2.5- to 3-fold increase in dissolved CO2 in seawater,
although the subsequent drop in pH will cause shifts in the
equilibria between inorganic carbon forms; HCO2 3 concentra-
tions will only rise by  10% and total Ci available for pho-
tosynthesis (HCO2 3 1 CO2) will thus rise by < 15%. However,
if CO2 availability is a constraint to the rate of photosynthe-
sis, could the more abundant CO2 result in faster macroalgal
growth? Most macroalgae have the capacity to use the
HCO2 3 ion that is dominant in seawater and given that the
HCO2 3 concentrations will not change much in future, an
impact on photosynthesis is unlikely. Furthermore, most
macroalgae and seagrasses (Beer et al. 2014) possess carbon
concentrating mechanisms (CCMs; Giordano et al. 2005)
which actively take up inorganic carbon (as HCO2 3 and/or
CO2) and thereby accumulate CO2 at the active site of the
Hill et al.
CO2-fixing enzyme Rubisco. This effectively saturates
Rubisco for CO2 and limits any increased productivity of
macroalgae from elevated atmospheric CO2 and ocean acidi-
fication. There are few exceptions, mostly subtidal rhodo-
phytes, to the rule that macroalgae possess CCMs, so
increased CO2 per se is unlikely to increase CO2 drawdown
by coastal primary producers. However, there are suggestions
that higher CO2 may increase carbon allocation to seagrass
rhizomes under high light (i.e., in shallow water) and
thereby enhance burial rates (Jiang et al. 2010).
Anthropogenic activities such as climate-change-
associated rising sea temperatures, are expected to increase
the occurrence of macroalgal diseases, through an increased
virulence of opportunistic pathogens, as well as reduced
resistance to disease in thermally stressed hosts (Egan et al.
2013). For example, in the chemically defended macroalgae,
Delisea pulchra, bacterial bleaching of the thallus occurs in
response to elevated temperatures, due to reduced antibacte-
rial chemical defences (Campbell et al. 2011; Case et al.
2011). Such diseases therefore have the capacity to reduce
macroalgal abundance, either through increased consump-
tion (due to reduced herbivory defence) or through direct
host mortality (Goecke et al. 2010).
With the delivery of warmer water further south, the East
Australian Current not only contracts the range of many
macroalgal species, but also extends the range of urchin con-
sumers. Urchin barrens are now extensive throughout south-
eastern Australia and are continuing to expand at the detri-
ment of macroalgal populations (Johnson et al. 2011). Such
altered ecological interactions with herbivores from this tro-
picalisation of high latitudes are therefore causing macroal-
gal populations to decline in Tasmania.
While climate change will have detrimental effects on
macroalgal populations, especially in temperate Australia,
tropical regions may experience increased macroalgal abun-
dance following thermal pulses on coral-dominated reefs. If
temperatures on coral reefs exceed summer averages by as
little as 1–28C, mass coral bleaching events can occur, which
can lead to coral mortality (Hoegh-Guldberg et al. 2007).
Phase shifts toward macroalgal-dominated reefs can follow,
especially if herbivory has been reduced by overfishing
(McManus et al. 2000; Bruno et al. 2009; Hughes et al.
2010). However, not all tropical macroalgal species will bene-
fit from climate change, with species such as Chnoospora
implexa expected to decrease in abundance (Bender et al.
2013).
When considering the donor capacity of macroalgae, algal
rafts have the potential to deliver dislodged macroalgae to
receiver habitats that can bury the carbon and thus enhance
the carbon sink (see above). However, changes in climate
will affect raft supply and persistence (Macreadie et al.
2011). Increased storm intensities (IPCC 2007) will promote
substratum detachment and/or fragmentation and hence
increase the abundance of algal fragments and rafts, and
12
Can macroalgae contribute to blue carbon?
may also increase burial of macroalgal material (Macreadie
et al. 2011). Elevated temperatures however, may counter
this process by reducing raft longevity due to an increase in
herbivore metabolism and consumption (Macreadie et al.
2011). In addition, changes in ocean surface currents may
affect advection routes and potential endpoints of deposition
into more or less favorable receiver environments for
sequestration.
Given the dynamic nature of macroalgal populations and
changes in abundance and productivity, especially from
human activities, the potential for carbon storage and export
may be highly variable. In considering the role macroalgae
may play in climate change mitigation, the factors detailed
above should be taken into account as they are likely to
greatly influence the contribution of macroalgae to blue car-
bon storage.
Conclusions and research priorities
Although macroalgal communities in Australia and
around the world produce massive quantities of plant bio-
mass with high organic carbon content, the presence of a
hard rocky substrate precludes the potential for long-term
storage of carbon within the macroalgal habitat. In isolation,
we therefore conclude that macroalgae do not contribute to
blue carbon.
However, a considerable proportion of macroalgal bio-
mass becomes dislodged from the benthos and is exported
beyond macroalgal habitat boundaries. This donor carbon
may be received by habitats with long-term sequestration
capacity, such as seagrass beds, saltmarshes, mangroves, and
continental shelf regions. Within Australia, and globally,
geographic differences in macroalgal biomass, community
composition and productivity will influence the blue carbon
donor capacity of macroalgae, with the greatest contribu-
tions likely to be in the temperate (and possibly polar)
regions. In addition to a high rate of biomass production, to
be a viable carbon donor, macroalgae must be transferred to
a receiver habitat, undergo burial and avoid microbial remi-
neralization. The fate of detached macroalgal biomass is not
entirely clear, and whilst limited data on the composition of
macroalgal biomass indicates that macroalgal carbon is
highly labile, recent research has highlighted the potentially
significant role of refractory, taxon-specific compounds (car-
bonates, long-chain lipids, alginates, xylans, and sulphated
polysaccharides) from macroalgae in contributing to long-
term blue carbon stores (Trevathan-Tackett et al. in press).
We therefore, conclude that macroalgae have the potential
to make ecologically meaningful contributions toward global
blue carbon sequestration, as significant carbon donors, but
further research is needed to quantify these contributions.
Data on the degradability of macroalgal plant material is
limited to only a handful of taxa, so there is considerable
need for additional testing on other taxa. The capacity for
Hill et al.
any incorporation of organic material from macroalgae into
long-term carbon stores is only possible if the material can
avoid decomposition by microbes, potentially by entering a
storage habitat that is a rapidly accreting zone. Research
therefore needs to focus on determining the burial rates of
macroalgal tissue at receiver sites, which of these sites are
the greatest sink of macroalgal fragments, understanding the
environmental conditions that maximise the burial process,
and assessing whether burial is fast enough to limit micro-
bial remineralization of the organic macroalgal material.
This will also need to include further research identifying
interspecies variation in carbon donor capacity due to rate of
lability and rate of transport to accreting zones. In existing
blue carbon habitats, methods for identifying macroalgal car-
bon in the storage zone also need to be developed to gauge
historical contributions of macroalgae into these zones.
Once these unknowns are addressed, the potential for mac-
roalgae to contribute to blue carbon as a carbon donor will
become less ambiguous. Focusing future research on these
key areas will provide evidence as to whether macroalgae
should be included under the blue carbon umbrella.
Macroalgal aquaculture and the harvesting of natural pop-
ulations are growing industries worldwide and are actions
that can greatly increase the carbon sequestration capacity
afforded by macroalgae (Chung et al. 2011). Furthermore,
the production of macroalgal biochar (a process involving
the pyrolysis of biomass) for addition to soil, may represent
a viable long-term CO2 sink, although research in this field
is in its infancy (Lehmann et al. 2006; Bird et al. 2011).
Given that macroalgal populations have the highest abun-
dance, biomass and productivity of all coastal habitats, they
not only store an immense amount of carbon in their living
biomass, but they also have an outstanding potential to act
as significant carbon donors. However, macroalgae are vul-
nerable to a range of anthropogenic impacts, so it is impor-
tant to protect these ecosystems for their climate-change-
mitigation potential as well as their ecosystem services.
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Acknowledgments
This research is undertaken by the CSIRO Flagship Marine & Coastal
Carbon Biogeochemical Cluster (Coastal Carbon Cluster) with funding
from the CSIRO Flagship Collaboration Fund. The Coastal Carbon Clus-
ter is an Australian research program designed to foster vital scientific
research to strengthen our low carbon economy and prevent future
excessive greenhouse gas emissions and improve methods in estimating
how much carbon is stored in coastal areas. The Cluster is composed of
seven Australian Universities and one Australian Research Institute work-
ing with the CSIRO’s Wealth from Oceans Flagship. Funding support
was also provided by an Australian Research Council DECRA Fellowship
(DE130101084) to PIM.
Submitted 30 November 2014
Revised 7 May 2015
Accepted 1 June 2015
Associate editor: Thomas Anderson