Blunden, Farnham, Jephson, Fenn and Plunkett: The Composition of Maerl from the Glenan Islands of Southern Brittany

121

Botanica Marina
VoLXX.pp. 121-125,1977

The Composition of Maerl from the Glenan Islands of Southern Brittany

G. Biunden, W. F. Famham, N. Jephson, R. H. Fenn.and B. A. Plunkett

Ponsmouth Polytechnic, Portsmouth PO1 2DZ, U.K.

{Received Maich 22, 1976)

Abstract
Around the Glenan Islands of Southern Brittany the depth of the maerl deposits was found to vary with the site, äs
did the Substrate topography. At two out of fwe sites examined, the maerl was present in a ridge and furrow System,
in which there was a greater proportion of living plants in the furrows than in the ridges. The maerl was found to be
composed almost entirely of Phymatolithon calcareum. Other algal species found growing along with the maerl are
listed. Analysis of maerl for calcium and magnesium contents showed no significant difference in the values
obtained for living and dead plant material. The calcium content, calculated äs Ca+*t varied from 25.5% to 33.3% of
the dry weight and the magnesium content, calculated äs Mg**, varied from 1.7% to 3.3%. The major component of
maerl was found to be calcite with magnesium carbonate present äs a solid solution in the calcite structure at a
concentration of about 8%. Aragonite was shown to comprise about 10-15% of the total material.

Introduction the maerl. To sample the maerl quantitatively, a 20 cm2

At present, over 300,000 tonnes a year of certain
calcareous red algae of the family CoraUinaceae, know
collectively äs maerl, are collected commercially from
around the coast of Brittany. The areas and methods of
collection, the processing procedures, chemical con-
stituents and uses of maerl have been reviewed recently
by Blunden, Binns and Perks (1975). The mae'rl from
Brittany has been shown by Cabioch (1966) to be com-
posed mainly of varieties of both Phymatolithon
(Lithotkamnion) calcareum (Fall) Adey et McKibben
and L. corallioides Crouan frat. The two main collection
areas in Brittany are around the Glenan Islands in the
south and in the Gulf of St. Mal o in the north (Blunden,
Binns and Perks 1975). The Composition of some maerl-
beds has been studied, in particular that of the Bay of
Morlaix (Cabioch 1969), but that of the Glenan Islands
has received little attention in recent years. In view of
the commercial exploitation of this area, the maerl-beds
have been studied and this communication records the
findings.
Materials and Methods
Examination of the mae'rl-beds of the Glenan Islands
was made in July, 1975 by SCUBA diving at five
different sites around the Island of St. Nicholas, the
largest of the Islands and the site of commercial
exploitation. General observations were made at each
site äs to the depths of the beds, -Substrate topography,
distribution of living plants and the flora associated with
quadrat/core sampler was used, the top 5 cm of material
being taken. From this a 200 g fresh weight sample was
taken to ascertain the percentage of living and dead
maerl. A particle whose surface colour was 50% or more
pink was counted äs living.
From the material collected at each site, two representa-
tive samples of both living and dead plants were taken
and assayed for total calcium and magnesium content by
EDTA titration using Eriochrome Black T indicator and
for magnesium using Patton and Reader's indicator.
One sample of commercially-collected and processed
maerl from the Glenan Islands was examined by X-ray
diffraction using the Cu wavelength and the powder
diffraction lines were recorded photographically. The
more accurate d-spacings for the major component were
recorded on a powder refractometer, using the Fe
wavelength.
Results and Discussion
EHfferences in the mae'rl-beds were observed at the
various sites examined, äs outlined below. The depths of
the beds have been corrected to depths below Chart
Darum at Penfret.
Site l
At this site, the maerl was found at depths of 3 to 5 m
and the deposits were over l m deep. For this entire area
the mae'rl was present in a ridge and furrow System, the

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 122 Blunden, Farnham, Jephson, Fenn and Plunkett: The Composition of Maerl from the Glenan Islands of Southern Brittany

wavelengths being about 1.5 to 2 m and the wave heights
from 0.3 to 0.6 m. A similar System was described by
Jacquotte (1962) for Mediterranean maerl-beds and by
Keegan (1974) for some Irish beds. The furrows con-
tained a greater amount of living material than the ridges
(Tab. 1) and the average size of the maerl particles was
also larger in the furrows. It appeared that these larger
and more rounded particles had rolled into the furrows
where they had become aggregated. The associated flora
was sparse, but was mainly found growing attached to
the larger maerl particles.
Site 2
Maerl was found at depths of 1.5 to 5 m and the deposits
were over 2 m deep. At this site there was no ridge and
furrow formation, possibly because it was too sheltered,
and the living maerl was distributed evenly in the upper
2 cm of the surface. Grab marks from commercial
collections were observed, the scars being about l m deep
and 2 m square. The material in the scars was generally a
greyish colour and of small particle size, although there
was some indication that larger, living particles had rolled
into the dips.
Site 3
This site appeared to be more sheltered than the earlier
ones studied. The sea floor was almost flat and was
composed of a continuous bed of maerl, found at depths
of 2.5 to 4 m and being over l m thick. The associated
flora was more abundant here than at the earlier sites
examined and was found growing attached to the larger
pieces of maerl and also to shell debris. The most fre-
quent algal species observed were Scinaia turgida Chemin,
Halarachnion ligulatum (Woodw.) Kutz, Dudresnaya
verticillata (With) Le Jo\.,Mesogloia vermiculata (Sm.)
S. F. Gray and Liagora viscida (Forsk.) C. Ag; the most
frequent animals were Buccinum undatum L,fMarthaste-
rias glacialis L. and Echinus esculentus L. Shells of
Teilina sp. and ofPecten maximus L. were abundant in
the maerl.
Site 4
The maerl was found at depths of 4 to 5 m and was over
l m thick. A similar ridge and furrow arrangement äs
that observed at site l was apparent here and the two
sites were generally similar.
Site 5
At this site the maerl-bed, found at depths of 4 to 5 m,
was associated with clumps ofZostera marina L. which
formed scattered patches about 10 m in diameter. Be-
tween the Z marina clumps, the Substrate was fairly flat
and composed of mae'rl to a depth in excess of l m. The
maerl particles at the periphery of the Z marina clumps
tended to be of a greater size than those observed away
from them and the percentage of living plants was high
(Tab. 1). This site may possibly be a significant source
of new spores for the bed at large. This area also Supports
a richer fauna than the more open maerl beds.
From five samples taken at sites l and 4, where the
maerl was observed in ridge and furrow formations, the
average quantity of living maerl present in the surface
layers of the ridges was found to be only 4.5% whereas
the quantity in the furrows averaged 14.3% and the

Tab. l. Percentage of living maerl in surface layers of sea bed at different sites around the Island of St. Nicholas

Site No. Sample Depth (m) Description of site Living maerl

plants (%)

1 A 4.5 Ridge of ridge and furrow System 2.5

6 5 Furrow of ridge and furrow system 14
C 4.5 Ridge of ridge and furrow system 9
D 5 Furrow of ridge and furrow system 7.5
E 4.5 Ridge of ridge and furrow System 2.5
F 5 Furrow of ridge and furrow system 7.5
2 A 5 Bed approximately flat 11.5
B 4.5 Bottom of grab trough - maerl discoloured (grey) 8.5
C 1-5 Bottom of grab trough - maerl discoloured (grey) 8.5
D 4 Bottom flat 7.5
3 A 3 Bottom flat - sheltered site 15
B 2.5 Bottom flat - sheltered site 12.5
C 3 Bottom flat - sheltered site 17.5
4 A 4 Ridge of ridge and furrow system 5
B 4.5 Furrow of ridge and furrow system 11
C 4 Ridge of ridge and furrow system 3.5
D 4.5 Furrow of ridge and furrow system 31.5
5 A 4 In Zostera marina bed - flat bed 25
B 4.5 Away from Z. marina bed — flat bed 8
C 4 Away from Z. marina bed - flat bed 25

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Blunden, Farnham, Jephson, Fenn and Plunkett: The Composition of Maerl from the Glenan Islands of Southern Brittany 123

overall average for all the different sites is given in Table L Tab. 2, Algal species found growing in the maerl beds around the
Island of St. Nicholas
These values for living plante were calculated from
samples taken from only the top 5 cm of the beds and do Chlorophyceae
Codiolum polyrhizum (Lagerh.) Komm., in Shells.
not represent the quantity present in commercially- Ostreobium quekettii Born. et. Flah., in Shells.
collected material, in which the proportion of living Ulva rigida (C. Ag.) Thur.
plants would be very smalL Ptiaeophyceae
Aglaozonia parvula (Grev,) Zanaid. (Sporophyte of Cutleria
At the five different sites studied, a considerable area of parvula), on Shells
ground was examined and only at one site, viz. site 2, Arthrocladia villosa (Huds.) Duby
was there any evidence of commercial exploitation. 1t has Asperococcus turnen (Sm.) Hook.
Chorda filum (L,) Stackh., in Zostera bed
been suggested that maerl-beds represent the accumula- Cladosiphon zosterae (J. AgJ Kylin
tion of material transported by currents from living Gadostephus spongiosus (Huds.) C. Ag.
maerl elsewhere. From the extensive occurrence of Colpomenia peregrina Sauv.
Cutleria multißda (Sm.) Grev. (gametophyte)
living maerl on the surface at the Glenan maerl-beds it Cystoseira nodicaulis (With.) Roberts
seems likely that the dead material originated from Desmarestia aculeata (L.) Lamour.
living plants in situ. D. ligulata (Lightf) Lamour.
D. viridis (O. F, Müll.) Lamour.
Lemoine(1910), at the beginning of this Century, Dictyota dichotoma (Huds.) Lamour,
Halidrys siliquosa (L.) Lyngb.
investigated the distribution of maerl collected by Halopteris filicina (Grat*) Kütz.
dredging around Concarneau and the Glenan Islands. She Laminaria saccharina (L.) Lamour.
identified all her material äs different forms ofPhymato- Liebmannia leveillei J. Ag.
Mesogloia vermiculaia (Sm.) S. F. Gray
lithon calcareum (äs Lithothamniurn calcareum), but at Myrionema strangulans Grev.
that time L· corallioides was not accepted äs a separate Pseudolithoderma extensum (Crouan fratj S. Lund, on Shells
species. Nevertheless, we now consider the maerl beds Sacchorhiza polyschides (Lightf) Batt.
Sauvageaugloia chordariaeformis (Crouan frat.) Kylin
which we surveyed around St. Nicholas to be composed S. griffithsiana (Grev. ex Harv. in Hook.) Hamel ex Kylin
predominantly of P. calcareum* Lemoine (1910) Scytosiphon lomentaria (Lyngb.) Link
suggested that living maerl did not occur immediately Sporochnus pedunculatus (Huds.) C. Ag.
Stilophora rhizodes (Turn.) J. Ag.
around the Glenan Islands, but our in situ observations Taonia atomaria (Woodw.) J. Ag.
do not support this. Possibly the distribution of living
Rhodophyceae
maerl has altered within the past 60 years. Other algal Antithamnion cruciatum (C. Ag,) Nag,
species found growing in the maerl-beds are listed in Asparagopsis armata Harv. (gametophyte)
Table 2. The diversity and distribution of marine algae Bonnemaisonia asparagoides (Woodw.) C. Ag^ (gametophyte)
Brongniartella byssoides (Good ei Woodw.) Schm-
around the Glenan Islands have been discussed by Callocolax neglectus Schm. ex Batt., on Callophyllis
Bouxin (1959), Bouxin and Dizerbo (1971), Castric-Fey, Callophyllis laciniata (Huds.) Kütz.
Girard-Descatoire, Lafargue and L'Hardy-Halos (1973) Ceramium rubrum (Huds*) C. Ag.
Chondria dasyphylla (Woodw.) C. Ag.
and L'Hardy-Halos, Castric-Fey, Girard-Descatoire and Chylocladia verticillata (Lightf.) Bliding
Lafargue (1973). However, these workers paid little Conchocelis rosea Batt. (phase in the life-history of species of
attention to the algae associated with the maerl. None Porphyra and Bangia), in Shells
Cruoria pellita (Lyngb.) Fries
of these is exclusive to maerl-beds and most of the Cryptopleura rarnosa (Huds.) Kylin ex Newton
species recorded in Table 2 had been found previously Dudresnaya verticülata (With,) Le Jol.
around Glenan, with the main exception ofLiagora Gigartina teedü (Roth.) Lamour.
Halarachnion ligulatum (Woodw.) Kütz. (gametophyte)
viscida, which Hamel (1930) recorded elsewhere for the Heterosiphonia plumosa (Ellis) Batt.
French Atlantic* The epiflora of the Glenan maerl-beds Hypoglossum woodwardii Kütz
is generally similar to that of the Bay of Morlaix Kallymenia reniformis (Turn-) J, Ag.
Liagora viscida (Forsk.) C. Ag.
(Cabioch 1969) and for Algerian maerl-beds, äs described Lomentaria articulata (Huds.) Lyngb.
by Feldmann (1943), The dominant species on the Melobesia le/olisii Rosen.
Glenan maerl include Halarachnion ligulatwn, Uudres- M. minutula Fosl.
Naccaria wiggii (Turn.) Endl.
naya verticülata> Arthrocladia villosa and Sporochnus Nitophyllum punctatum (Stackh.) Grev.
pedunculatus. Peyssonelia rubra (Grev.) J, Ag.
Polyneura hittiae (Grev.) Kylin
Analysis of maerl samples from each of the different sites Polysiphonia fruticulosa (Wulf.) Spreng
for calcium and magnesium contents showed that there Rhodophyllis divaricata (Stackh.) Papenf.
Scinaia fwcellata Biv.
was no significant difference in the values obtained for Scinaia turgida Chemin
living and dead plant material (Tab. 3), although differ- Spyridia filamentosa (Wulf.) Harv. in Hook.
ences were found from sample to sample, The calcium Cyanophyceae
content (calculated äs Ca**) varied from 30.0% to Entophysalis deusta (Menegh.) Dr. et D-, in Shells
32.5% of the dry weight of the living plants and from Oscülawria sp., in Liebmannia
Rivularia mesenterica (Thur.) Born et Flah., on Shells
25.5% to 333% of the dead plants, whereas the Schizothrix calcicola (C. Ag.) Gom., in Shells

J
magnesium content (calculated äs Mg++) varied from

10
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 124 Blunden, Farnham, Jephson, Fenn and Piunkett: The Composition of Maerl from the Glenan Islands of Southern Brittany

Tab. 3. Calcium and magnesium contents of living and dead Examination by X-ray diffraction of a sample of com-
maerl from different collection sites around the Island of St
Nicholas mercially-collected and processed maerl from the Glenan
Islands and comparison of the interplanar spacings and
CaJcium and magnesium contents of intensities with JCPDS (Joint Committee for Powder
maerl
% by weight (g metal/100 g maerl) Diffraction Standards) values showed that the main
component of the material was calcite wi#i aragonite
Site no. Sample Living material Dead material present äs a minor component (Tab. 4). The intensity
Mg4 Mg' of the aragonite line at d = l ,98 Ä, given at 65, is equi-
valent to a line with an intensity of about 3 on the calcite
1 A 32.0 2.4 25.5 1.7 scale. Calculation from first principles or using the graph
B 32.2 3.2 32.5 3.1
given by Chave (1954) indicated that aragonite com-
2 A 32.9 2.4 31.9 3.1
B 32.1 3.0 32.1
posed about 10-15% of the material. The presence of
3.0
silicon dioxide is also indicated.
3 A 31.9 2.2 32.6 3.3
B 32.0 3.1 32.3 3.1
The observed d-spacings of the major component are all
4 A 32.5 2.2 31.9 2.9
B 32.3
smaller than those of pure calcite (Tab. 5), which indi-
3.0 33.3 3.2
cates that magnesium is present in the calcite äs a solid
5 A 30.0 3.1 32.9 3.1
B 31.9 2.9 31.7 2.8 solution. Chave (1952) investigated the effect on the
d-spacing of the 100 planes in the rhombohedral lattice
of the weight per cent of magnesium carbonate present
2.2% to 3.2% of the dry weight for the living plant äs a solid solution in the calcite structure. He published a
material and from 1.7% to 3.3% for the dead. The lack graph showing a linear relationship between 2% and
of any noticeable difference between the living and dead 16% w/w of magnesium carbonate. The equation of the
plants is not surprising äs only the surface layer of the line is 3.035-d observed (100) = 0.00298x, where is
maerl particles is made up of living material, whereas the the % w/w of magnesium carbonate. From the powder
bulk of each particle is dead. diffractometer measurements, the value of is 8%.

Tab. 4. CuKa X-ray photographic powder diffraction data of maerl

Line Intensity Interplanar JCPDS JCPDS JCPDS

Spacing Calcite data Aragonite data Silicon Oxide Type 0
(dA> d I/I d l/l d l/l

VW 1.41 1.422 3
W 1.43 1.440 5
ww 1.48 1.510 3
w (doublet) 1.51 1.518 4
1.525 5
w 1.59 1.604 8
VW 1.61 1.626 4
ww 1.75 1.74 25
Spotty 1.82 1.84 60
s 1.862 1.875 17 1.88 25
(doublet) and
32
s 1.898 1.913 17
1.927 5
w 1.99 1.98 65
s 2.083 2.095 18
s 2.272 2.285 18
ww 2.35 2.34 31
2.37 38
m 2.484 2.495 14 2.48 33
ww 2.72 2.70 46
w 2.82 2.845 3
vs 3.016 3.035 100
ww (double) 3.29 3.27 52
Spotty 3.38 3.40 100 3.38 100
m 3.845 3.86 12
Spotty 4.27 4.32 40

All other lines for these materials are weaker than the quoted ones
vs - very strong s = strong m - medium
w = weak vw = very weak ww = very very weak

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Blunden, Farn ha m, Jephson, Fenn and Plunkett: The Composition of Maerl from the Glenan Islands of Southern Brittany 125

Tab. S. Powder diffractometer values for calcite interplanar Our results agree with those of Levy and Strauss (1961)
spacings of maerl using FeKo wavelength who reported that the major component of maerl is a
d d from JCPDS calcite data solid solution of magnesium carbonate in calcite with
OBS
percentages of magnesium carbonate present varying
3.815 3.860 between 9.3 and 18,5 from different sources. They found
3.011 3.035 also a small, unspecified amount of aragonite present.
2.810 2.845
2.477 2.495 Differential thermal analysis of maerl was attempted and
2.265 2.285 one peak corresponding to carbonate decomposition was
2.076 2.095 obtained supporting the idea that the magnesium was an
1.910 .927
1.890 .913 integral part of the calcium carbonate structure and not
1.858 .875 present äs a separate entity.
1.610 .626
1.588 .604
1.578 1.578
1.511 1.525
1.500 1.518 Acknowledgements
1.495 1.510
1.461 1.473 We wish to thank Estabiissements Boennec and Glen-
1.427 1.440
1.406 1.422 organic Ltd- for financing this project, and M. and Mme.
1.327 1.339 1. Boennec and Mr. A. K. R. Spence for their help. We
1.285 1.297
1.257 1.284 are very grateful to Mr. P. MUls-Goodlet for obtaining
1.168 1.180 the powder diffractometer data and to Madame J.
1.141 1.154 Cabioch for verifying the identity of the Phymatolithon
1.034 1.047
calcareum samples.

Literature

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