, published 15 January 2005 , doi: 10.1098/rsta.2004.1488 363 2005 Phil. Trans. R. Soc.

Andrew S. Y. Mackie, P. Graham Oliver, Teresa Darbyshire and Kate Mortimer

Plateau: high diversity in a tropical oligotrophic environment

Shallow marine benthic invertebrates of the Seychelles

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Phil. Trans. R. Soc. A (2005) 363, 203228
doi:10.1098/rsta.2004.1488
Published online 12 November 2004
Shallow marine benthic invertebrates of
the Seychelles Plateau: high diversity in
a tropical oligotrophic environment
By Andrew S. Y. Macki e, P. Graham Oli ver,
Teresa Darbyshi re and Kate Morti mer
Department of Biodiversity and Systematic Biology, National Museum of Wales,
Cathays Park, Cardi CF10 3NP, UK ([email protected])
Soft sedimentary biotopes are extensive in the shallow Western Indian Ocean, espe-
cially on the Seychelles Plateau and Mascarene Ridge, yet pro rata compared with
coral reefs the research eort devoted to them has been minimal. In this study we
examine the benthic mollusc and polychaete worm assemblages of the shallow waters
(1162 m) around Mahe, in the Seychelles, and make direct comparisons with the
temperate Irish Sea area and subtropical waters of Hong Kong, China (using identical
methodology).
Two assemblages were recognized, characterized by depth and sediment type. Of
these, assemblage A (in shallow carbonate sands) was the most diverse, with diversity
and richness measures exceeding those from the Irish Sea or Hong Kong. Hong Kong
generally had the poorest fauna. Considering the Bivalvia alone, estimates of taxo-
nomic distinctness showed this to be least for Seychelles assemblage A. The degree of
conformity of the results to the concept of the latitudinal gradient in species richness
and the possible underlying causes are discussed.
Comparisons with other data suggest that the Seychelles support a benthic fauna
at least as diverse as any other described from the tropics. A tentative examination
of total bivalve species richness suggests a total of 400500 for the Seychelles. This
is in keeping with other Indian Ocean localities, but higher than known gures for
continental east Africa. The ndings of this paper support the case for widespread
ecological and taxonomic studies of the Western Indian Ocean benthic invertebrates.
Keywords: macrobenthos; taxonomy; diversity;
tropics; subtropics; temperate
1. Introduction
The study of the diversity of soft sediment invertebrate communities has an extensive
history in both temperate shelf (less than 200 m) and deep-sea environments (e.g.
Sanders 1968, 1969; Abele & Walters 1979; Grassle 1991; Gray 1997; Gray et al .
1997). However, it was the much discussed paper by Grassle & Maciolek (1992)
One contribution of 24 to a Discussion Meeting Atmosphereoceanecology dynamics in the Western
Indian Ocean.
203
c 2004 The Royal Society
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204 A. S. Y. Mackie and others
that rmly placed the deep sea at the centre of the debate concerning high benthic
diversity. Tropical ecosystems are also regarded as being highly diverse but, in reality,
there are few comparable studies from tropical marine, soft sediment, biotopes (see
Alongi 1990). Many of the studies cited in Alongis review relate to intertidal habitats
and only a few focus on the continental shelf of the Indo-Pacic province.
The latitudinal gradient in species richness is frequently cited in comparative diver-
sity discussions (Krebs 1985; Rosenzweig 1995; Hawkins 2001). In general, the nega-
tive relationship with increasing latitude holds true (Hillebrand 2004), but causation
remains a matter of debate (Levin et al . 2001; Willig et al . 2003). Further, there are
many exceptions, and conicting results exist for the marine macrobenthos (Thorson
1957; Rex et al . 1993; Dauvin et al . 1994; OHara & Poore 2000; Gray 2001).
Thorson (1957) showed that the species richness of several infaunal groups
remained constant between dierent climatic areas, while that of the epifauna
increased in the tropics. Comparative studies of shallow tropical and temperate muds
have lent support to the rst, revealing little dierence in diversity and community
structure (Warwick & Ruswahyuni 1987; Kendall & Aschan 1993). Recently, Frouin
& Hutchings (2001) reported on the macrobenthos of organically enriched sands in
a Tahitian lagoon. Their species-richness estimates were again no higher than those
found in temperate shelf environments.
Nevertheless, the two regions dier in their wider environmental characteristics.
Contrary to the situation in the temperate region, the tropical shelf can be dom-
inated by carbonate sediments and, in the Indo-Pacic, there are large areas of
low-production, oligotrophic, waters (Alongi 1990). Published studies of the mac-
robenthos of shallow carbonate sediments in tropical regions are scant and prelimi-
nary, with that of Lewis & Taylor (1966) relevant to the Seychelles. Notable scien-
tic investigations to the Seychelles include the Percy Sladen Trust (Gardiner 1907,
plates 1418) and the Dutch Oceanic Reefs Expedition (van der Land 1994a, b). Two
unpublished environmental impact reports giving basic macrobenthic data exist for
two localities in the Seychelles, but these are of restricted access, of variable taxo-
nomic resolution, and give no indication of any particular richness. Available studies
have inevitably concentrated on reef-associated communities (van der Land 1994a, b),
and Taylor (1968) reported a diverse molluscan fauna.
In March 2000, in conjunction with the Royal Geographical SocietyRoyal Society
Shoals of Capricorn Programme, the National Museum of Wales carried out marine
biodiversity studies in the Seychelles, with one of its prime aims being the collection of
quantitative benthic samples from carbonate sands. The samples were to be collected
and analysed in line with similar studies carried out by us in the temperate Irish
Sea (Mackie et al . 1995) and subtropical Hong Kong, China (Mackie et al . 1993).
By using identical collection and analytical methods and applying similar taxonomic
rigour, this paper aims to provide empirical comparisons between temperate and
tropical shelf macrobenthic communities.
2. Methodology
For the comparisons made between the three localitiesIrish Sea (51.353.5

N),
Hong Kong (approximately 22.5

N) and the Seychelles (4.54.8

S)data from pre-

viously published studies by Mackie et al . (1993, 1995) were reanalysed to make them
Phil. Trans. R. Soc. A (2005)
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Tropical benthic diversity 205
compatible with the Seychelles results available at the time of writing (January 2004;
see below).
(a) Taxonomy
Measures of ecological diversity and species richness require adequate species recog-
nition but the taxonomic tools available for the Western Indian Ocean are virtually
non-existent and the taxonomic literature is widely dispersed. This taxonomic imped-
iment (Oliver 1993; Kendall et al . 2000) has forced this study to focus on those groups
familiar to the authors, namely the Mollusca and Polychaeta.
Studies of the molluscs of the Seychelles date back to the early nineteenth century
(Dufo 1840) and continue with faunal lists such as Von Martens (1880), Melvill
(1909) and Winckworth (1940). The single identication guide available (Jarrett
2000) covers only the larger molluscs. Current data from other regions in the Western
Indian Ocean (Red Sea, Dekker & Orlin 2000; Arabian Sea, Dance et al . 1995;
Reunion Island, M. Jay 2003, personal communication) suggest that a molluscan
fauna in excess of 2000 species is likely for the Seychelles.
Given that the Seychelles is part of the Indo-Pacic province, accurate species-
level identication is at least lengthy, and often not possible, without undertaking
taxonomic revision. Many of the species identications used here must therefore be
tentative.
This study attempts to test the use of taxonomic distance measures as developed
by the Plymouth Marine Laboratory (Warwick & Clarke 1995, 2001; Clarke & War-
wick 1998, 2001b) and this requires comment on the generic, family and order-level
taxonomy adopted. The genera within the Bivalvia, especially as applied to the Indo-
Pacic fauna, are not yet stable and vary between authors. One of the most common
problems is in the application of subgenera and subfamilies. In the bivalve family
Tellinidae one commonly adopted classication (Millard 1997) recognizes only two
subfamilies but over 30 subgenera. The web-based Indo-Pacic Molluscan Database
(I-PMD) recognizes four subfamilies and raises the subgenera in Millard to generic
status.
Polychaete taxonomy is perhaps less advanced than that for the molluscs, but
nomenclatural issues and practical taxonomic problems are no less common. The lit-
erature is perhaps smaller, though it is equally dispersed. There are no comprehensive
up-to-date taxonomic guides to the group in the Indian Ocean Region, and Fauvel
(1953) and Day (1967a, b) are still commonly used as standard reference works.
Attempts are being made to use the Internet and modern technology (Kendall et
al . 2000) to improve polychaete taxonomic skills. However, much basic descriptive
as well as detailed systematic (Rouse & Pleijel 2003) work remains to be done. The
polychaetes of the Seychelles have been the subject of relatively few historical studies
(e.g. Potts 1909, 1910), though there have been a number of taxonomic and ecological
papers published in recent years (ten Hove et al . 1994; Westheide 2000; Westheide &
Hass-Cordes 2001; Darbyshire & Mackie 2003; Mortimer & Mackie 2003; B oggemann
et al . 2003).
The higher level polychaete designations for the taxonomic distance indices were
approximated to seven orders from the classication outlined in Beesley et al .
(2000). This was based on the cladograms produced by Rouse & Fauchald (1997)
and, as in the recent modern reference (Rouse & Pleijel 2001), did not recognize the
Linnean hierarchy.
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206 A. S. Y. Mackie and others
N
10
11
9
8
7
39
1
2
3
12
6
5
4
Mah
Ste Anne
miles
kilometres 2 4
2 4
Figure 1. Benthic sampling-station positions around Mahe, Seychelles.
In this study, comparison with a European temperate fauna is to be made, and
consequently classications which have comparable levels of denition need to be
adopted. Consequently, the classications used are the I-PMD (recognizing ve
bivalve orders) and the British Species Directory (Heppell et al . 1997; Mackie &
Erseus 1997: lower taxonomic levels).
(b) Sampling
The Seychelles material was all collected from the Seychelles Fisheries Authority
ship, the RV LAmitie, during the National Museum of Waless expedition to the
island of Mahe in March 2000. The aims of the expedition were to collect polychaete
and mollusc specimens for taxonomic research, and benthic samples for biodiversity
assessments.
All samples from the 13 quantitative stations o Mahe (gure 1) considered in
this paper were obtained using the modied 0.1 m
2
Van Veen grabs rst success-
fully deployed in the Irish Sea in 1997 (Mackie & Darbyshire 2001). Samples were
sieved (0.5 mm mesh) and xed in formalin stained with Rose Bengal. Specimens
were subsequently washed and preserved in 80% alcohol with 2% propylene glycol
(Mackie & Oliver 1996). Three replicate samples were taken at each station; two for
macrofauna and one for sediment analysis. The samples were sorted in the Seychelles
by museum sta and identications were made by the authors. At the present time,
all the molluscs but only part of the polychaete fauna have been identied (see Elec-
Phil. Trans. R. Soc. A (2005)
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Tropical benthic diversity 207
tronic Appendix). The polychaetes identied comprised 20 of the 41 families present,
belonging to the Amphinomida, Phyllodocida (except Nereididae, Glyceridae, Gonia-
didae), Eunicida (except Lumbrineridae, Oenonidae, Dorvilleidae), Spionida, and one
family (Orbiniidae) of the Scolecida. Sediments were analysed by Sediment Analysis
Services, Edington, Somerset.
(c) Multivariate and univariate analyses
The polychaete assemblages were initially elucidated by cluster analysis (Bray-
Curtis similarity coecient with group average fusion) using the Plymouth Rou-
tines in Multivariate Ecological Research (Primer version 5) computer program
(Clarke & Gorley 2001; Clarke & Warwick 2001a). The similarity matrix was further
employed to produce a two dimensional non-metric multi-dimensional scaling (MDS)
ordination of the inter-station relationships. Species abundances were log
10
(x + 1)-
transformed to limit the inuence of the most dominant species.
The environmental factors having potential inuence on the mollusc and poly-
chaete distributions were investigated using the BIO-ENV routine (Clarke &
Ainsworth 1993). The following variables were examined: gravel (%), sand (%), silt
clay (%), carbonate (%), depth, latitude and longitude. Following preliminary assess-
ment, gravel and siltclay were log
e
(x + 1)-transformed to reduce skewness in the
data. All other variables were untransformed.
A variety of diversity and evenness measures were calculated for each station
(replicates were combined). These included the diversity measures of Fisher ()
and ShannonWiener (H

), and the Heip evenness measure (E

Heip
; Heip 1974). The
ShannonWiener and Heip indices were calculated using log
2
values. Diversity was
additionally studied with the rarefaction methodology using ES100 values.
Three taxonomic distance measures were calculated for the combined bivalve-
polychaete (part) fauna: the quantitative average taxonomic distinctness (

) using
both raw and log(x + 1)-transformed abundances, its presenceabsence equivalent
(
+
), plus the variance (
+
) of the latter. Branch lengths were equal between each
level: species, genus, family, order, class, phylum. For the bivalve analysis, the super-
family level was also used, since class and phylum had no involvement in the analysis.
Any species (e.g. juvenile) that could not be assigned to a higher category was omit-
ted.
Species-accumulation curves and projected species-richness values were calculated
using the EstimateS program (Colwell 1997) with 50 randomizations.
3. The study areas
The three locations under study dier in geographical, environmental and faunal
characteristics (table 1). The Irish Sea waters can be considered to be mesotrophic,
though an increasing tendency toward eutrophic conditions has been identied in
the northern part (Allen et al . 1998). The southern Irish Sea area is fully marine
with moderate water temperatures typical of temperate European seas. Mackie et
al . (1995) described the benthic faunal assemblages and reviewed the environmental
conditions present there.
Much of the shallow Tolo Channel location in subtropical Hong Kong has been
greatly aected by eutrophication attributed, in part, to organic pollution (Morton
Phil. Trans. R. Soc. A (2005)
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208 A. S. Y. Mackie and others
Table 1. Summary of environmental parameters and macrofaunal attributes
at each geographical location
(Figures in italics relate to the polychaete fraction under study; denotes lack of data.)
southern Irish Sea Seychelles Hong Kong
location
latitude 51.353.5

N 4.54.8

S 22.5

N
longitude 4.26.5

W 55.355.5

E 114.3

E
climate temperate tropical subtropical
water
environment
production status mesotrophic oligotrophic mesotrophiceutrophic
salinity 3435 % % 3536 % % 2437 % %
bottom temperature 813

C 2430

C 1527

C
benthic
environment
sediments muddy sands, mainly mud
sands and gravels calcareous sands
calcium carbonate 868% 10100%
organic matter 0.47.5% 8.7%
depth 7130 m 1363 m 420 m
sampling stations 51 13 10
benthic
macrofauna
assemblages 3 (+2 stations) 2 (+2 stations) 1 (+1 station)
number of taxa
polychaetes and molluscs 469 (295) (317) 117 (76)
polychaetes 341 (167) (145) 95 (54)
molluscs 128 172 22
bivalves 71 82 11
1990; Chan & Wong 1993; Lee & Arega 1999). In addition, elevated temperatures and
increased freshwater inows during the summer months can produce a stratication
of the water column. Dramatic deteriorations in benthic oxygen levels sometimes
occur, killing the invertebrate fauna (Lam 2000). However, there are now some signs
of improvement in the water quality (Broom et al . 2003). Shin (1982, 1990, 2003),
Mackie et al . (1993) and Shin et al . (2003) described the benthic infauna of the
muddy Tolo ChannelHoi Ha Wan area.
Satellite imaging of chlorophyll (Yentsch & Phinney 1992; Banzon et al . 2004;
McClain et al . 2004) and measurement of nutrient (de Sousa et al . 1992; Baars et
al . 1995) concentrations indicate that the oceanic waters in the Seychelles region are
oligotrophic. A recent zooplankton study (Gallienne et al . 2004) shows that increased
productivity can occur south of the Seychelles (914

S). This may be due to nutrient

upwelling associated with the South Equatorial Current and its interaction with the
Mascarene Ridge. Salinity and water temperature and are both high in this region
(de Sousa et al . 1992; Jury et al . 2000), with average values of the former 3536 % %
and the latter 1820

C (at 100 m depth).
Phil. Trans. R. Soc. A (2005)
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Tropical benthic diversity 209
Table 2. Sediment and depth summary for each Seychelles benthic sampling station
station gravel (%) sand (%) siltclay (%) carbonate (%) depth (m)
1 6.25 93.56 0.17 99.9 21.0
2 24.08 75.88 0.04 66.1 30.0
3 5.74 89.19 5.08 41.0 45.0
4 0.88 96.35 2.78 91.8 35.0
5 2.23 97.78 0 93.4 35.0
6 3.46 96.39 0.15 98.8 25.5
7 2.00 86.21 11.80 32.2 62.5
8 10.44 88.84 0.71 22.8 41.5
9 0.27 91.86 7.87 16.4 28.0
10 5.46 69.79 24.75 91.0 56.0
11 5.67 92.97 1.36 10.0 48.0
12 0 56.74 43.26 65.8 27.5
39 1.74 91.37 6.89 99.0 12.5
The sediments of the shallow waters (1235 m) east of Mahe (gure 1) are predom-
inantly calcareous sands (table 2) with over 90% carbonate content. For the stations
sampled, carbonate decreases with increasing depth (2145 m) to 41% in the south-
east (stations 2 and 3). To the northwest, carbonate increases with depth (2863 m)
from 1632%. To the west, station 11 (48 m) had the least carbonate (10%), though
the shallower and deeper stations had higher levels: 66 and 90%, respectively. The
latter were also the muddiest encountered, having 25 and 43% siltclay. The coarser
sand/gravel fractions of the sediments also contained quartz granules. This was noted
in the sampling log for the sieved faunal samples at stations 13, 5, 7, 8 and 11.
4. Can a faunal fraction be used as a surrogate for the whole?
The present study analysed only part of the macrobenthic fauna present, namely all
of the molluscs and a portion the polychaetes. No data are available for the remaining
Seychelles polychaetes, arthropods, echinoderms or other minor groups. Here, half
the 40 polychaete families present have been examined. Collectively, these families
equated to 49% and 57%, respectively, of the total polychaete species present in the
Irish Sea and Hong Kong studies (table 1). Numerically, they accounted for 42% and
90% of the total polychaete individuals present.
These same families corresponded to an average of 48.4 6.8% polychaete species
per station for the Irish Sea and 76.3 15.1% polychaete species per station for
Hong Kong. The relationships between the polychaete fraction and the total number
of polychaete taxa were strong for both the Irish Sea and Hong Kong locations (g-
ure 2). Interestingly, Olsgard et al . (2003) showed that even one polychaete order
the Terebellidacould be used as a surrogate for both total polychaete and total
macrofauna species richness. However, the relationships diered geographically and
were weaker for the whole macrofauna than those derived from using all the poly-
chaetes. The authors suggested that a larger subgroup would probably improve the
correlations.
Previous work in the Irish Sea and elsewhere has shown that the polychaetes
are good surrogates for whole macrobenthic assemblages (Mackie et al . 1995, 1997:
Phil. Trans. R. Soc. A (2005)
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210 A. S. Y. Mackie and others
70
20
30
40
50
60
10
0
120 100 80 60 40 20 0
Polychaete taxa (total)
P
o
l
y
c
h
a
e
t
e

t
a
x
a

(
f
r
a
c
t
i
o
n
)
is
hk
Figure 2. Relationships between polychaete fraction and total polychaete taxa
for Irish Sea (r
2
= 0.901) and Hong Kong (r
2
= 0.983) stations.
Olsgard & Somereld 2003). Comparisons of cluster dendrograms and MDS plots
from the Irish Sea and Hong Kong showed that the polychaete fraction available
for the Seychelles analysis produced the same assemblage patterns as that obtained
using all the polychaetes or all the macrofauna.
While these ndings for the Irish Sea and Hong Kong cannot prove the same holds
for the Seychelles, they do suggest a more general relationship existsat least in
situations where polychaetes are the dominant component of the benthic macrofauna.
(a) Benthic assemblages in the Seychelles
The quantitative cluster analyses for the molluscs (gure 3a) and polychaete frac-
tion (gure 3b) essentially showed the same two assemblages, with the exception of
the anities of station 3. The bivalve analysis (not shown) was very similar to the
overall mollusc analysis, except that station 8 showed a higher anity for stations 3
and 11 than the main assemblage.
The combined analysis (gure 3c) had the same overall structure as the polychaete
analysis, though the internal relationships within the larger group diered. This was
not considered signicant, since the linkages of the majority of stations occurred over
a relatively narrow band of similarity (3646%).
Phil. Trans. R. Soc. A (2005)
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Tropical benthic diversity 211
B
r
a
y

C
u
r
t
i
s

s
i
m
i
l
i
a
r
i
t
y

(
%
)
1
2
3
4
5
6
7
8
9
10
11
12
39
station
1
2
3
4
5
6
7
8
9
10
11
12
39
100
80

60

40

20

0
1
2
3
4
5
6
7
8
9
10
11
12
39
100
80
60
40
20
0
100
80
60
40
20
0
39 6 2 5 4 1 9 8 3 112110 7
39 6 2 9 8 5 4 1111210 3 7
39 6 2 5 4 1 8 9111210 7 3
stress = 0.15
stress = 0.10
stress = 0.08
(a)
(b)
(c)
(d )
(e)
( f )
Figure 3. Cluster and non-metric MDS analyses of Seychelles macrofauna: (a), (d) molluscs;
(b), (e) polychaete fraction; (c), (f) combined mollusc and polychaete fraction.
Overall, the deeper stations (3, 7 and 10) from the southeast to northwest tended to
form a group distinct from the shallower ones in the east/northeast and northwest.
Stations 11 and 12 in the southwest were intermediate, with the former closer to
the larger assemblage (here designated A) and the latter closer to the smaller B
assemblage.
The two dimensional MDS plots of the same data supported this and there was
a high degree of concordance between the patterns. The mollusc plot (gure 3d)
had the greatest degree of stress and there appeared to be more gradation between
Phil. Trans. R. Soc. A (2005)
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212 A. S. Y. Mackie and others
Table 3. Faunaenvironment relationships for the Seychelles benthos
(Highest correlations in bold.)
number of best variable correlation
variables combinations (p
s
)
molluscs and 2 mud, depth 0.659
polychaetes (part) 3 mud, depth, latitude 0.675
5 mud, depth, latitude, sand, carbonate 0.677
molluscs 2 mud, depth 0.682
3 mud, depth, latitude 0.675
5 mud, depth, latitude, sand, carbonate 0.695
polychaetes (part) 2 mud, depth 0.573
3 mud, depth, latitude 0.594
5 mud, depth, latitude, sand, carbonate 0.601
the two (A and B) clusters with stations 3, 11 and 12 occupying the intermediate
position.
Qualitative (presenceabsence) analyses for the above faunal groupings exhibited
a high degree of concordance. Assemblage A was always a coherent group. Assem-
blage B was a little more variable. Stations 7 and 10 always grouped together, but
the anity of station 3 varied, appearing closer to station 11 (for bivalves) or 12
(molluscs). The polychaete and combined analyses produced the same groupings as
in the quantitative analyses (gure 3e, f).
If the Seychelles analyses follow the same pattern as the Irish Sea and Hong Kong
ones, then the total faunal assemblages are likely to agree with the two identied in
the combined analyses carried out here. In the following sections, assemblage A and
B refer to these station groupings (gure 3c, f).
(b) Seychelles faunaenvironment relationship
The BIO-ENV routine compares the similarity matrices between the faunal and
environmental variable plots and attempts to nd the best match. The same ve
variables gave the highest Spearman correlation value in all cases (table 3). The
maximum value, approaching p
s
= 0.70, was similar to that found in the larger Irish
Sea study. It was noteworthy that the two-variable mud-depth combination gave only
slightly lower values in each case. In the Irish Sea study, gravel, depth and silt were
the best combination.
The identication of depth and sediment composition as major factors determining
macrobenthic faunal assemblages is a common (and logically understood) result of
such analyses. Such methods do not, however, prove that the environmental variables
identied cause the animal distributions. Thus some variables identied as causative
may simply be so by chance, or be proxies for other unmeasured ones. In the present
study, the identication of latitude is a case in point. The actual improvement in the
correlation (above that obtained from the lower variable combinations) is small. The
small scale of the latitude range present in the study makes it an unlikely factor in
itself. Perhaps it is related to the inuence of the prevailing wind direction: from the
southeast?
Phil. Trans. R. Soc. A (2005)
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Tropical benthic diversity 213
5. Is benthic diversity really higher in the tropics?
Reference to the summary characteristics of the three locations compared (table 1)
immediately suggested that the Seychelles had a richer fauna than either the Irish
Sea or Hong Kong. There were more mollusc and polychaete taxa in 13 stations
than were obtained for 51 Irish Sea stations. As the Irish Sea assemblage C stations
(shallow to moderate depth gravels) had previously been identied as approaching
those of the deep sea in their richness (Mackie et al . 1995, 1997), this indicated that
a more detailed examination of diversity was called for.
(a) Station level comparisons of species richness, diversity and evenness
The station diversity values for the molluscs and polychaete fraction have been
summarized for each Seychelles assemblage to facilitate comparison with equivalent
data from the Irish Sea and Hong Kong (table 4). While the results for the molluscs
can be viewed in absolute terms (and so can be compared with other studies), the
combined molluscpolychaete and polychaete values are only comparable for the
three locations examined here. Direct comparison with other studies would only be
possible if the polychaete data in those were reduced to the same subset analysed
here.
For the Seychelles, assemblage A stations were generally richer than those of assem-
blage B and the singleton stations 11 and 12. Species-richness estimators (S, ,
ES100) and the ShannonWiener diversity index (H

) were all at a maximum in this

assemblagefor all faunal components. Heips evenness measure (E
Heip
) was more
varied. The molluscs and bivalves exhibited high evenness at all except station 12,
but evenness for the polychaete fraction (and hence also for the combined fauna)
was much lower in assemblage B.
Cross-comparison with data from the other two geographic locations, revealed
Hong Kong to have the poorest fauna and the Seychelles to have the richest. Only
the number of polychaete species at station 40 (Hoi Ha Wan, muddy sand with detri-
tus, 4 m) approached the values for Seychelles assemblage A and the rich Irish Sea
assemblage C (gravelly sediments, 27120 m). The general trend for all the diver-
sity measures was exemplied (gure 4) by the rarefaction species estimates for 100
individuals (ES100). The value for Seychelles station 11 was included for reference,
although there were only 95 individuals present. The molluscan fauna of the Sey-
chelles assemblage stations tended to be richer, more diverse, and have a higher
degree of evenness than the Irish Sea or Hong Kong assemblage stations.
Direct comparison with other studies is dicult at presentpending the com-
plete identication of the Seychelles fauna. However, we know that the Irish Sea
gravelswith an average of 145 species per 0.2 m
2
and ShannonWiener diversity
(H

) of 5.8represent one of the richest shelf habitats currently known. These have
a richer fauna than Tahitian lagoon sediments (Frouin & Hutchings 2001), one of the
few detailed studies available from the tropical Indo-Pacic. The richest assemblage
(using abundance data) in the Tahitian study had an average of 104 species per
0.1 m
2
.
One of the most detailed and methodologically compatible investigations avail-
able from tropical waters is that of Wade (1972) from Kingston Harbour, Jamaica,
Caribbean Sea. These shallow muds and sandy muds yielded 2373 macrobenthic
taxa (excluding sh) per 1 m
2
station. Recalculated diversity (H

) was 3.595.02
Phil. Trans. R. Soc. A (2005)
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214 A. S. Y. Mackie and others
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Phil. Trans. R. Soc. A (2005)
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Tropical benthic diversity 215
60
0
10
20
30
40
50
A A 12 11 B A C B
E
S
1
0
0

(
m
e
a
n

1

s
.
d
.
)

40
Irish Sea Seychelles Hong Kong
Figure 4. Hurlbert rarefaction species estimates (ES100) per station
for combined molluscpolychaete fractions.
(n = 8), ES100 was 22.137.2 (one station incalculable) and Heip evenness 0.37
0.75. Although the evenness values were in the same range as the Irish Sea gravel
fauna, species-richness and diversity values only approached the lower range values of
that assemblage. The Jamaican richness and diversity were more directly comparable
with the muddy oshore Irish Sea assemblage subgroup A1.
The subtropical Hong Kong benthic fauna is not particularly diverse. The richest
benthic assemblage in Hong Kong (Shin et al . 2003) had a two-station average of
50 species per 0.5 m
2
and an H

(converted log
2
value) of 4.43. In comparison, the
Hoi Ha Wan station 40 used in the present analyses had 118 species per 0.2 m
2
and
an H

value of 4.66. It therefore appears that, by any standard, the main Seychelles
assemblage stations are likely to possess faunas as rich as those known for soft-bottom
shelf localities anywhere.
(b) Station-level comparisons of taxonomic distinctness
Taxonomic distinctness measures examine diversity by quantifying the degree of
relatedness between the taxa in each sample or station. A major advantage of such
measures is that they, unlike species-richness measures, have been shown to be inde-
pendent of sample size (Warwick & Clarke 1998). They have been mainly used in
comparisons against expected values to identify dierences in taxonomic structure
attributable to disturbance or other environmental inuence. They can also be used
to compare diversity in dierent geographic locations (e.g. Price et al . 1999).
Contrary to the situation with respect to species richness, patterns in taxonomic
distinctness for the polychaete fraction were found not to be the same as those for
the whole polychaete fauna and they were excluded from the analyses. Only the
Bivalviathe group supported by the most taxonomic hierarchical information
Phil. Trans. R. Soc. A (2005)
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216 A. S. Y. Mackie and others
50
55
60
65
A1 40 12 11 B A C B1
Hong Kong Seychelles Irish Sea
+

(
m
e
a
n

1

s
.
d
.
)
BX A2
0
50
100
150
A1 40 12 11 B A C B1
Hong Kong Seychelles Irish Sea
BX A2
200
250
300
350
50
55
60
65
A1 40 12 11 B A C B1
Hong Kong Seychelles Irish Sea
BX A2

*

(
m
e
a
n

1

s
.
d
.
)

+

(
m
e
a
n

1

s
.
d
.
)

(a)
(b)
(c)
Figure 5. Taxonomic distinctness measures for bivalve molluscs: (a)
+
; (b)
+
; (c)

.
were considered. The Hong Kong assemblage A stations were also excluded as they
possessed too few species. However, the analyses were broadened by including the
four subgroups of the Irish Sea assemblages (see Mackie et al . 1995). Subgroup A1
comprised six deep muddy sand and mud stations and subgroup A2 four deep but
generally more sandy stations. Subgroup B1 comprised eight muddy sand stations
and subgroup Bx the remaining 12, predominantly sandy, stations of the inshore
Irish Sea assemblage B.
The individual station values for each taxonomic distinctness measure exhibited a
high degree of intra-assemblage variability. Hence, to facilitate comparison, assem-
blage station means (1 standard deviation) were graphed (gure 5). The mean
values of

in the untransformed (not shown) and log-transformed analyses (g-

ure 5c) were almost identical for each assemblage/subgroup and Seychelles station 11.
Phil. Trans. R. Soc. A (2005)
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Tropical benthic diversity 217
A combination of low species number and highly abundant tellinids or semelids pro-
duced lower untransformed

values for the other two singleton stations.

The most signicant nding was the lower mean taxonomic distinctness of Sey-
chelles assemblage A (gure 5a, c). The mean variation in taxonomic distinctness
(
+
) exhibited great uniformity across the assemblages/stations, though Seychelles A
and Irish Sea subgroup A1 had higher valuesmarkedly so in the latter case (g-
ure 5b). The higher value for Irish Sea A1 was simply due to marked dierences in
the taxonomic evenness of very few bivalve species (510) per station, for example,
due to a greater prevalence of nuculoid species (Protobranchia). The value for Sey-
chelles station 11 was close to zero, as the ve heterodont species were almost evenly
distributed through the hierarchy (ve genera, ve families, four subfamilies).
A lower average taxonomic distinctness for Seychelles A is perhaps surprising given
that this assemblage contained the most species-rich and diverse fauna. This indicates
that the taxonomic structure of the Seychelles calcareous sand assemblage diers
from the various temperate ones. Reduced taxonomic distinctness has been found
in environmentally degraded habitats (Warwick & Clarke 1995), though this may
not always occur (Somereld et al . 1997). While there is no obvious reason why
the Seychelles calcareous sands should be regarded as degraded, one possibility is
that there are dierences in the trophic structure associated with habitat. Warwick
& Clarke (1998) showed that
+
was explicitly related to the trophic diversity of
benthic nematode assemblages: a reduced trophic diversity being associated with a
reduced taxonomic distinctness.
Close examination of the taxonomic hierarchy shows that Seychelles A and Irish
Sea C have a similar number of entities at each higher level from family to order. Sey-
chelles A diers in having four times the number of Tellinidae species, making up 20%
of the total analysed species complement (60). Furthermore, at the superfamily level
there were over twice the number of species (16) in the Tellinoidea. Hence not only
were there more species, they were more unevenly distributed among the families.
The galeommatoidean species were also about twice as prevalent in Seychelles A. The
remaining species had approximately similar overall distributions within the higher
taxa of both assemblages. The net result of this is a relative reduction in the average
taxonomic distinctness, and a relative increase in its variation, in Seychelles A.
The underlying causes for this dierence in taxonomic distinctiveness are prob-
ably many and, as yet, are speculative. It is known that the taxonomic make-up
of tropical and temperate bivalve faunas is dierent, with the tropics showing a
diverse radiation in the heterodontsparticularly in the Tellinoidea and Veneroidea
(Crame 2000). Although not well documented at this time, current research indi-
cates a similar radiation pattern in the largely commensal Galeommatoidea, with
many undescribed taxa being recognized (Bouchet et al . 2002). The dierences in
distinctiveness could therefore be simply a function of evolutionary history and have
no ecological link. However, the heterodont component of Seychelles A is restricted
mostly to the Tellinoidea, with the Veneroidea not equally prevalent. Tellinoideans
are deposit feeders, whereas veneroids are suspension feeders, and consideration of
the trophic structure of the Seychelles Plateau may be an inuencing parameter.
The sediment type may also be favouring tellinoids over other taxa in that they
are particularly adapted to sandy habitats. Further, Riddle (1988) reported tellinids
as characteristic of the shallow calcareous sands found in Great Barrier Reef lagoons.
In such sediments tellinoids are relatively deep, fast, burrowers compared to the ven-
Phil. Trans. R. Soc. A (2005)
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218 A. S. Y. Mackie and others
eroids. Therefore, consideration also may need to be given to the relative predation
pressures upon bivalves in dierent habitats and latitudes.
(c) Assemblage-level comparisons of species richness
To further investigate the dierences between the three geographic locations,
species-accumulation curves were generated (using randomized stations) for the
major assemblages present at each (gure 6). These quite clearly demonstrated the
higher richness of Seychelles assemblage A for all faunal components. The Seychelles
assemblage B curve, with only three stations, was similar to the initial section of
that for the Irish Sea assemblage C in the combined molluscpolychaete fraction
plot (gure 6b), and to both Irish Sea assemblage B and C curves for the bivalves
(not shown). The remaining Irish Sea and Hong Kong assemblage curves were in the
same relative, and lower, positions on all plots except for the polychaete one. In the
polychaete fraction plot (not shown), the Irish Sea assemblage C curve was inter-
mediate between the two Seychelles curves and the other two Irish Sea assemblage
curves were virtually identical, intermediate between Seychelles assemblage B and
the Hong Kong curve.
6. Concluding discussion
Comparisons of species richness and diversity can be strongly aected by scale (Willig
et al . 2003; Hillebrand 2004). Gray (2000) proposed a unied species-richness ter-
minology of four scales (point, sample, large area and geographical province) and
two categories (habitat and assemblage), the latter being nested in large area species
richness. This paper examines sample (equal to station) and assemblage diversity.
Neither the large area biodiversity of the Seychelles Plateau benthos nor that of the
Western Indian Ocean geographical province can be examined at present due to the
lack of equivalent datasets.
The use of a standardized collecting regime at each of the three locations studied
here allows us to directly compare station diversity from three dierent biogeographic
provinces. The use of multivariate techniques permits us to also regard these in terms
of species assemblages. The Seychelles assemblage A stations clearly possess a very
rich and diverse fauna in comparison woith the temperate Irish Sea or subtropical
Hong Kong, and this holds for both molluscs and polychaetes. Likewise, the same
Seychelles assemblage possesses the highest total species richness.
Interest in determining total species-richness estimates from assemblages (or
regions) has a long history. Many extrapolative techniques have been proposed (e.g.
de Caprariis et al . 1981; Heltsche & Forrester 1983; Chao 1984; Palmer 1990; Sober on
& Llorente 1993; Karakassis 1995; He & Legendre 1996). The paper by Colwell &
Coddington (1994) and the availability of Colwells EstimateS computer program
(Colwell 1997) have increased awareness and use of these methods. Magurran (2003)
provides a very useful overview and analysis of their use.
To investigate the degree of congruence of these extrapolative methods with our
observed species-accumulation curves, we have calculated a variety of estimators for
the bivalve molluscs (table 5) using Colwells EstimateS. No assumptions are made
about which is best. As Magurran (2003) has shown, there are considerable inter-
study dierences in the merits of each. Rather, we have used them to judge the
possible completeness of our assemblage sampling.
Phil. Trans. R. Soc. A (2005)
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Tropical benthic diversity 219
sA
sB
hk
isA
isB
isC
sA
isB
isC
isA
hk
sB
0
50
100
150
200
250
t
a
x
a

(
S
)
t
a
x
a

(
S
)
1 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 20
stations
160
40
60
80
100
120
140
20
(a)
(b)
Figure 6. Randomized species-accumulation curves (error bars omitted for clarity) for Sey-
chelles (s), Irish Sea (is) and Hong Kong (hk) macrofaunal assemblages: (a) molluscs; (b) mol-
luscpolychaete fraction.
Table 5. A comparison of bivalve species-richness estimators for
each benthic assemblage calculated using Colwells EstimateS software
(Bold values indicate where accumulation curves appear to become asymptotic.)
Seychelles Irish Sea Hong Kong

estimator A A B C A
Species (S) observed 67 24 47 52 7
ACE 93 27 49 55 8
ICE 130 32 51 59 10
Chao 1 83 28 51 58 7
Chao 2 125 30 51 61 8
Jacknife 1 99 31 54 62 9
Jacknife 2 118 34 54 66 8
Bootstrap 81 27 51 57 8
MichaelisMenten (mean) 101 32 55 61 10
probable species 100? 32 52 59 8
Phil. Trans. R. Soc. A (2005)
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220 A. S. Y. Mackie and others
For the Irish Sea and Hong Kong, the estimators generally give a narrow range
of values that, from our experience of these faunas, give very reasonable proba-
ble maxima. The situation for the Seychelles assemblage A is quite dierent. The
estimates cover a wide range and only one estimator appears to be becoming asymp-
totic (at a relatively low value). Although factors such as homogeneity of habitat,
species patchiness and species abundance distribution can all inuence the estima-
tors, much of this variation undoubtedly reects the need for additional sampling.
The species-accumulation curve shows little sign of levelling o for an assemblage
of only eight stations. This would indicate a much higher estimate of total species
for that assemblage and we believe a bivalve count of about 100 species is possible.
This postulation may not be excessive as a single grab sample taken by us at 100 m
on Le Constant Bank, Seychelles contained 72 bivalve taxa when both live and dead
shells are included.
Bouchet et al . (2002) reported a very high molluscan diversity from New Caledo-
nia with some 2738 species, of which 519 were bivalves. Using the Jacknife 2 method
in EstimateS they estimated a bivalve fauna of 750 species. Combining data from
Oliver (1992), Dekker & Orlin (2000) and Zuschin & Oliver (2003) gives a bivalve
fauna of 450 for the Red Sea. Maurice Jay (2003, personal communication) estimates
over 400 bivalve taxa for Reunion and up to 500 for Mauritius. This could reason-
ably indicate a total Seychelles fauna of around 400500 species and thus Seychelles
assemblage A would contain 2025% of this total. However, this relatively high pro-
portion may be an overestimate due, in part, to the sampling methodology. Collecting
using a 0.5 mm sieve will reveal many small species rarely included in other studies.
It is likely that a similar strategy used for coral-reef-associated assemblages would
also reveal higher than expected richness compared with published data.
One must be very careful when making species-richness comparisons in the Indo-
Pacic realm as the datasets are rarely comparable. This study, for assemblage A
alone, recognizes 67 bivalve species from a combined sampling area of 1.6 m
2
. Other
datasets cover vast regions and numerous habitats. At this time we cannot assess
dierences in diversity within habitats across a geographic province. Neither can
we assess the contributions made by assemblages to the total faunas. Crame (2000)
postulates that it is the presence of reef systems in the tropics that is a major con-
tribution to bivalve diversityyet few data are available for examining the relative
diversities of reefs versus soft sediments.
Do our ndings of very high species richness in the Seychelles lend support for a
latitudinal species gradient in the marine benthos? A number of studies have reported
such gradients (e.g. Rex et al . 1993, 1997; Roy et al . 2000), though these ndings have
been questioned (Gray 1997, 2001). Taken in isolation, cross-comparisons between
single temperate, subtropical and tropical locations certainly cannot prove such a
hypothesis. The generally lower diversity for the Hong Kong location does not t
any simple pattern, though this can be at least partially attributed to the highly
changeable local hydrographic conditions (see Lee & Arega 1999; Lam 2000) and
cannot be regarded as representative of the subtropics as a whole. High (and low)
benthic species diversities are now known from many dierent latitudes and depths
(Poore & Wilson 1993; Blake & Hilbig 1994; Mackie et al . 1995; Coleman et al . 1997).
Interestingly, Hillebrand (2004) examined 198 marine datasets and concluded that
latitudinal gradients were present, but they were generally weaker for the infauna
and sessile epifauna than for mobile epifauna.
Phil. Trans. R. Soc. A (2005)
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Tropical benthic diversity 221
A recent large-scale coastal assessment of bivalve molluscs (Crame 2000) did
report strong latitudinal (and longitudinal) gradients in diversity. The gradients
were most marked on the coasts of the Americas, but the pattern was much less
apparent in the Indo-Pacicexcept for the steep gradient along the Pacic coast
of China/Japan/Russia and for New Zealand. The Indo-Pacic showed more of a
longitudinal gradient, with a hot spot in the Philippine/Indonesian basin and high
species richness around Australia. Figures for the Indian Ocean were much lower
and on the east African coast an apparent minimum was present on the Equator.
However the data we have on species numbers around the oceanic islands suggest
they are much higher than that given for continental east Africa, giving more even
values across the Western Indian Ocean.
Crames data suggest that a comparison of continental diversity with oceanic island
diversity might throw light on the role of reef communities and the inuence of ter-
rigenous input. Taylor (1997) is unequivocal about the role of nutrients Bivalves,
for instance, are much more diverse and abundant in eutrophic continental envi-
ronments, than on oceanic oligotrophic atolls and reefs. He includes the Seychelles,
as high oceanic islands, in the nutrient-rich category though the terrigenous input
on these cannot compare with that from the rivers of east Africa or the productivity
of the Somalia and Oman upwellings, which, according to Crame (2000), have lower
richness. In general, however, the Seychelles and Mascarene islands lie in an olig-
otrophic region and the extent of inuence of the islands themselves on the extensive
banks around them is unclear. The numbers of individuals in our samples suggests
that productivity is not high on the Seychelles Plateau. The relationship between
diversity and productivity is complex, however, and can change relative to resource
supply and consumer pressure (Worm et al . 2002). Little more can be added regard-
ing water nutrient levels and productivity until detailed studies are made in the
Seychelles and comparative data are obtained from throughout the Western Indian
Ocean. A review of the continental east African and Madagascar faunas would also
be highly relevant.
This study has shown high diversity for tropical soft-sediment assemblages and a
potentially high species richness for the Seychelles benthic invertebrate fauna. We
are currently ignorant of such assemblages anywhere else along the Mascarene Ridge,
especially on the massive Saya de Malha, Nazareth and St Brandon banks. This re-
emphasizes the need for comparative studies across a wide range of habitats and
geographic scale.
Additional comparative data would reveal gradients or patternsperhaps related
to sediment, depth, productivity and latitudenot only along the Mascarene Ridge,
but throughout the Indian Ocean. Such studies are the basis for assessing benthic pro-
ductivity and environmental impacts in the oceans bordering the developed nations.
Perhaps of greatest relevance at this time would be the contribution benthic ecology
could make to the AgulhasSomalia Large Marine Ecosytem (LME) project (Sher-
man et al . 1998). In the Indian Ocean such methodology is rarely adopted because,
as in our study, it is hampered by the taxonomic impediment that is common for
much of the Indo-Pacic (see Oliver 1993). We highlight the need for taxonomic
work to provide congruence across studies and to accurately use taxonomic distance
measures. A high degree of taxonomic suciency is also required to recognize fau-
nistically distinct regions and aid conservation management (Terlizzi et al . 2003).
Phil. Trans. R. Soc. A (2005)
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222 A. S. Y. Mackie and others
This work was funded by the National Museum of Wales and carried out with the help and
support of the Royal Geographical SocietyRoyal Society Shoals of Capricorn Programme. We
acknowledge: the sta of the Shoals of Capricorn programme, including Martin Callow, Jan
Robinson and Caroline Lawton, for their logistical help in the Seychelles; John Collie and the
sta at the Seychelles Marine Parks Authority (MPA) for use of their facilities; the Seychelles
Fisheries Authority, especially Captain Gerrard Ernesta and the crew of the RV LAmitie for
their expertise and enthusiasm in collecting the oshore samples; BioSyB sta for help in col-
lecting and sorting samples; Andy Woolmer (University of Wales, Swansea) and Mairi Best
(University of Chicago) for assistance on LAmitie; and Jon Houghton (University of Wales,
Swansea) for feeding us all. Thanks to Ivor Rees for supplying a copy of an important paper at
short notice. This paper is Shoals Contribution no. PO50.
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Discussion
J. Gerlach (Nature Protection Trust of Seychelles, Cambridge, UK). Can you
identify any notable features in particular samples such as those near the port or
reclaimed areas, or are there only broader geographical patterns?
A. S. Y. Mackie. Our sampling was designed to investigate the benthic invertebrate
infaunal patterns in a broad sense. It is unlikely that we have sucient sampling
stations for the reliable assessment of any specic local inuence.
M. D. Spalding (CCRU, Cambridge University, Cambridge, UK). The deep ocean
benthic studies published to date have shown very high levels of discovery of new
species, previously undescribed, reaching levels of 8090% of all species sampled.
Were you getting such high levels of new species?
A. S. Y. Mackie. That is an interesting question. At present we have only studied
the molluscs and polychaete worms; therefore we must be careful of extrapolation.
Some faunal groups in the Seychelles area may be better known than others. However,
the taxonomic work we have already carried out suggests that many new polychaete
Phil. Trans. R. Soc. A (2005)
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228 A. S. Y. Mackie and others
species are present on the Seychelles Plateau. For example, in the genus Magelona,
we have found some seven speciesof which at least six are new to science. One
thing that is noticeable for both molluscs and polychaetes is that there appears to
be a large species turnover from sample to sample. Thus -diversity of these shallow
water calcareous sands is likely to be high. We will examine this in more detail when
more data are available.
The supplementary Electronic Appendix is available at http://dx.doi.org/rsta.2004.1488
or via http://www.journals.royalsoc.ac.uk.
Phil. Trans. R. Soc. A (2005)
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