CIIMAR Centro Interdisciplinar de Investigacao Marinha e Ambiental, Rua dos Bragas, 123, 4050-123, Porto,
Portugal
2
ICBAS Instituto de Ciencias Biomedicas de Abel Salazar, Universidade do Porto, Lg. Prof. Abel Salazar 2, 4099-003,
Porto, Portugal
(*Author for correspondence: E-mail: ronaldo.sousa@cimar.org)
Received 5 May 2005; in revised form 27 July 2005; accepted 31 July 2005
Key words: Lima estuary, macrobenthic assemblages, abundance, biomass, diversity, environmental factors
Abstract
During the summer of 2002, sampling was carried out in the Lima estuary in order to compare the pattern
of the macrobenthic communitys distribution in relation to physical and chemical variables. A total of 54
macrobenthic taxa were identied. Abundance, biomass and specic diversity varied among the twenty
stations. Abundance ranged from 212 to 9856 ind./m2, with an average of 1581 ind./m2. Abra alba presented the highest density corresponding to 39.1% of the total specimens gathered, followed by Hediste
diversicolor with 31.5%. Biomass ranged from 0.12 to 264.62 g AFDW/m2, with an average of 17.58 g
AFDW/m2. Cerastoderma edule and A. alba were the species with a clear predominance in the total
biomass, contributing 75.3 and 13.8%, respectively. The multivariate techniques used revealed a macrobenthic community with ve distinct groups, particularly related to the sedimentological characteristics and
salinity. These results demonstrated signicant dierences in macrobenthic assemblages composition along
an estuarine gradient. For the rst time the presence of the nonindigenous invasive species Corbicula
uminea was described in this estuary.
Introduction
Estuaries are normally recognized as transitional
areas of distinct biological importance, where
salinity varies between fresh water and seawater
(McLusky, 1989; Little, 2000). These estuarine
areas are used temporarily as a nursery ground for
many commercially and ecologically important
species of marine and migrating shes, shellsh,
crustaceans and shorebirds (Day et al., 1989;
Raaelli, 1999).
At the same time these productive and complex
estuarine ecosystems are frequently located near
large cities and are receptacle of all types of contaminants (Allchin et al., 1999; Baudrimont et al.,
136
benthic macroinvertebrates are one of the most
useful bioindicators of possible environmental
changes (Warwick et al., 2002; Ysebaert et al.,
2002). Since there is a complete lack of quantitative data on the relationship between spatial distribution of macrobenthic assemblages and
environmental variables in the Lima estuary reference work was carried out, during the summer of
2002, on the subtidal area. The objectives of this
work were to compare the pattern of the macrobenthic communitys distribution in relation to
physical and chemical variables, to estimate the
abundance and biomass of subtidal macrozoobenthos and to identify possible key species in the
functioning of the ecosystem.
Figure 1. Map of the Lima estuary showing location of the twenty sampling stations.
137
During eldwork, the following water column
information was collected: temperature, salinity,
dissolved oxygen and pH. This information was
obtained in situ, close to the bottom, using a
multiparametrical sea gauge YSI 820. Particle size
and organic content analyses of the sediment at
every station were carried out. Sediment granulometry was assessed following drying at 60 C for
72 h. Dried sediment was sieved through a column
of sieves corresponding to integer values of the
Wentworth scale and the frequency of each class
was expressed as the percentage of total weight.
The quantity of organic matter contained in the
sediment was determined, after combusting for
24 h at 550 C, in a mue furnace. Values are
expressed in percentage, relatively to the weight
loss on ignition of each sample analysed.
Grab samples were sieved on a 1 mm mesh and
xed with 4% formalin in seawater. Macrofauna
was sorted and, whenever possible, identied to
species level. Biomass was calculated using the Ash
Free Dry Weight Method AFDW (Kramer et al.,
1994).
Data analysis
To compare the similarity between stations (data
pooled over ve biological replicates for each site)
in terms of species composition (abundance and
biomass), univariate measures and multivariate
analyses were applied using the PRIMER package
(Clarke & Warwick, 2001). Univariate measures
included abundance, biomass, number of species,
and diversity (H) and evenness (J) indices. Classication (CLUSTER) and ordination by nonmetric multidimensional scaling (MDS) based on
the BrayCurtis similarity matrix were used to
analyse spatial distribution of benthic assemblages.
In order to establish correlations between biological parameters and abiotic characteristics, indeces of biotic and abiotic similarity were
compared using PCA (Principal Components
Analysis) and BIOENV (using the Spearman
coecient) (Clarke & Ainsworth, 1993).
Biological information was previously transformed (square root transformation; abundance
per unit of area), however, for the abiotic factors
only granulometry and the percentage of organic
matter were transformed )log(1+y).
Results
The physical and chemical parameters are presented in Table 1. The water temperature was
higher in upstream stations and ranged between
17.8 C (station 18) and 13.2 C (stations 5 and 6).
Salinity values varied signicantly with the distance to the river mouth, with the highest value
recorded in station 1 (34.8 psu) and the lowest in
station 20 (0.4 psu). The dissolved oxygen values
varied between 3.7 mg/l (station 14) and 7.8 mg/l
(station 10) and the pH values between 7.0 (station
6) and 8.2 (stations 13 and 14). There was a large
variation in the cumulative curves of distribution
of the sediment throughout the dierent sampling
stations. The stations located in the river mouth
(stations 1, 2, 3 and 4) contained abundant ne
matter and the other stations, located upstream,
contained coarser sediment with a preponderance
of sands and gravels. Organic matter in the sediment ranged from 0.5% (station 19) to 10.8%
(station 3). There were large spatial variations,
related to the type of substrate existent in the
dierent sampling stations and a signicant correlation (r=0.98, p<0.001) between ner sediment
(silt and clay) and organic matter was found.
A total of 7750 individuals corresponding to 54
macrobenthic species were found (Appendix 1).
These species belong to 7 Phyla: Annelida 17,
Mollusca 20, Arthropoda 10, Nemertini 3,
Echinodermata 2, Cnidaria 1 and Spinculida 1.
Abundance per station ranged from 212 ind./
m2 in station 17 to 9856 ind./m2 in station 2 with
an average of 1581 ind./m2 (Table 2 and Appendix
1). Macrofauna abundance was distributed among
Mollusca (56.1%), Annelida (38.8%), Crustaceans
(4.9%) and other faunal groups (0.2%). Abra alba
was the species that presented the greatest number
of individuals per unit of area, which corresponded to 39.1% of the total number of individuals gathered. Hediste diversicolor was the
second most abundant species with 31.5%.
The number of species (Table 2) also had considerable spatial variation. The maximum value of
20 species was registered in station 2 and the
minimum, with only one species, in stations 14, 15
and 17. The ShannonWiener index (Table 2)
values were low with the maximum value at station
1 (H=1.96) and the minimum at stations 14, 15
and 17 (H=0.00). Pielous evenness varied widely
138
Table 1. Physical and chemical parameters of water and sediments
Station T(C) S(psu) DO(mg/l) pH OM(%) G(%) VCS(%) CS(%) MS(%) FS(%) VFS(%) S+C(%) D (m) DST(km)
1
13.5
34.8
6.4
7.6
2.9
0.1
0.1
12.6
27.1
52.1
13.9
33.9
6.4
7.4
8.9
0.6
1.2
5.1
12.3
43.3
33.5
10
0.3
3
4
13.9
14
34.1
33.9
6.2
6.1
7.6 10.8
7.7 7.5
2.4
0.2
1.4
1.2
5.6
5.2
13.4
13.9
17.5
32.6
26.8
27.5
32.9
19.4
10
9
1
1.5
13.2
33.4
5.2
7.4
0.9
27.2
22.9
13.3
9.8
0.5
0.3
2.8
13.2
33.5
6.8
4.3
4.2
14.4
24.1
18.8
23.2
10.6
4.7
13.3
32.4
6.9
7.4
1.9
17.9
19.4
21.3
22.5
15.5
1.6
1.8
13.5
32.2
7.2
7.3
2.2
12.5
33.6
31.8
13.1
2.9
3.6
2.5
14.3
32.3
7.4
7.4
38.4
32.6
18.9
8.6
0.9
0.3
0.3
10
14.3
32.8
7.8
7.5
1.9
21.4
32.6
34.7
8.2
1.2
0.9
11
12
17
14.2
22.9
32.4
5.8
5.4
7.5
7.5
0.8
1
58.9
58.2
17.9
22.4
14.3
16
7.7
2.8
1
0.2
0.1
0.2
0.1
0.2
2
5
7
8
13
15.3
28.9
4.9
8.2
0.7
51.3
28.4
17
2.9
0.2
0.1
0.1
9.5
14
14.7
27.5
3.7
8.2
0.8
19.3
45.1
30.2
4.9
0.3
0.1
0.1
10.5
15
15.2
24.7
3.9
8.1
0.6
66.7
18.4
11.4
3.2
0.1
0.1
0.1
10
12.5
16
17.3
7.4
4.6
2.5
9.1
19.4
36.2
22.2
7.1
3.9
2.1
13.5
17
17.4
4.8
1.9
15.5
18.2
22.9
31.7
7.5
2.8
1.4
14
18
17.8
2.4
5.1
0.6
29.2
35.8
29.3
4.4
0.8
0.4
0.1
14.5
19
20
17.4
17.2
1.1
0.4
5.2
4.7
8.1
8.1
0.5
0.7
54.6
50.5
28.4
19.4
13
16.7
3.8
11.3
0.1
1.9
0.1
0.1
0
0.1
2
3
15
16
26
Temperature (T C), salinity (S psu), dissolved oxygen (DO mg/l), pH, organic matter (OM %), gravel (G %), very coarse sand
(VCS %), coarse sand (CS %), medium sand (MS %), ne sand (FS %), very ne sand (VFS %) and silt+clay (S+C %).
The depth (D m) and distance to the ocean (DST km) of the twenty sampling stations are also given.
139
Table 2. Abundance (A ind./m2), biomass (B g AFDW/m2),
number of species (S), evenness (J) and Shannon/Wiener index
(H) at the stations sampled
Station
420
1.55
13
0.76
1.96
9856
24.53
20
0.21
0.62
1112
8.14
18
0.39
1.13
3265
28.31
0.14
0.27
360
2.13
0.52
1.14
628
0.70
15
0.68
1.85
7
8
3124
1196
264.62
3.58
8
14
0.39
0.35
0.81
0.93
2632
4.76
0.20
0.39
10
2268
2.46
0.19
0.34
11
888
0.73
0.21
0.34
12
1628
1.23
0.17
0.12
13
568
0.37
0.19
0.13
14
516
0.27
0.00
0.00
15
16
900
304
0.24
1.58
1
4
0.00
0.53
0.00
0.74
17
212
0.12
0.00
0.00
18
632
0.44
0.81
0.56
19
444
5.55
0.22
0.16
20
664
0.30
0.09
0.07
Figure 2. A dendogram of the twenty sampling stations present in the Lima estuary.
140
Discussion
The summer temperature values registered in the
Lima estuary increased from the river mouth to
the upstream areas. The stations near the river
mouth are subjected to a great inuence of adjacent marine water, which have lower values than
the water from uvial inux. The upper shallow
stations present higher temperature values due to
atmospheric inuence. Salinity decreased from the
river mouth to upstream areas due to the dilution
of marine water with fresh water derived from
uvial inux. The spatial uctuations of the dissolved oxygen and pH values were never large
(extreme values were never recorded) and the
macrobenthic assemblages present in the estuary
are presumably tolerant to the amplitudes occurTable 3. Summary of results from BIOENV analysis
k
0.545 Silt+Clay
0.703 Salinity
0.681 Salinity
Very ne sand
3
0.731 Salinity
Silt+Clay
0.710 Salinity
Fine sand
Silt+Clay
Fine sand
141
stability of salinity in the lower part of the Lima
estuary may also be responsible for the larger
diversity present, because this area can be easily
colonized by species originating from adjacent
marine areas. There are various examples of marine
species, which generally colonize the lower part of
the estuary: G. convuluta, A. rotundatus, D. pugilator, N. nucleus and A. squamata, amongst others.
These species were present mainly in the river
mouth areas and in the relative deep navigation
channels, where the inuence of the tide is strongest. The decreased diversity in the middle and
upstream areas may be related to the type of habitats available and with environmental stress. The
middle and upper areas of the Lima estuary are
characterized by habitats with clean, coarse sand
and high degree of instability caused by natural
processes (mainly large uctuations of salinity,
high current velocities and high turbidity), resulting in a lower diversity of benthic macroinvertebrates (Marchand & Gascuel, 1988; Attrill et al.,
1996). Furthermore, the middle and upper estuarine areas are poor in phytoplankton (Da Silva,
1990) and organic matter when compared to lower
stations. According to Herman et al. (1999), the
food availability may be a limiting factor for benthic abundances and biomass in estuarine benthic
systems. At the same time, the salinity gradient acts
as a physiological barrier for stenohaline marine
and freshwater species and places environmental
stress on euryhaline organisms (Mannino &
Montagna, 1997) and so great uctuations of
salinity, principally, in the middle estuary can result in a reduction of the number of species.
This kind of macrobenthic distribution, with a
clear dierence in assemblages composition,
abundance and biomass along an estuarine gradient is in conformity with several studies done in
other estuaries and coastal lagoons (Marques
et al., 1993; Mannino & Montagna, 1997; Ysebaert et al. 2002, 2003; Nanami et al., 2005). Other
factors may be responsible for the abundance,
biomass and species diversity observed in the Lima
estuary such as predation, competition, adult-larval interactions and other abiotic factors not
measured in this study. Eects of predation,
competition and adult-larval interactions have
been demonstrated only in dense assemblages of
benthos (Wilson, 1984). They probably are less
important than the abiotic factors in sparse ben-
142
A. alba, H. diversicolor and C. edule suggest that
these species play a key role in the Lima estuary
and possibly serve as important food resources to
the other trophic levels.
Acknowledgements
Special thanks are addressed to Eduardo, Jonathan Wilson, Jose Calvario and Miguel Pardal.
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Actinidae
Sp2
Sp3
3
4
Sp
Sp
11
13
14
15
16
17
18
19
20
21
23
24
25
Bivalvia
22
Streblospio shrubsolii (Buchanan, 1890)
Mollusca
12
x
4
28
72
16
x
x
x
x
S4
16
x
x
60
x
x
15
220 x
x
x
S3
x
x
x
8
28
x
x
S6
96
x
16
x
x
S7
84
24
x
x
S9
28
20
x
40
x
x
S10
x
24
x
x
S11 S12
x
x
x
x
x
x
x
x
x
x
x
x
x
x
32
20
16
x
x
x
x
2420 2116 824 1588 552 516 900 224 212 476 x
x
x
x
x
x
x
x
x
x
x
x
944 4
12
x
x
S8
2180 24
192 x
232 x
x
x
x
x
S5
x
x
44 20
10
x
x
4
x
x
x
S1 S2
Polychaeta
7
Ampharete grubei (Grube, 1860)
Annelida
Oligochaeta
Annelida
Spinculida
Sp1
Nemertini
Cnidaria
Taxa
Appendix 1. List of the taxa identied in the Lima estuary. For each taxa, the average abundance (ind./m2 ) is given (x absent)
Appendix
144
35
36
37
40
41
42
45
46
Diastylis sp
51
48
49
50
Decapoda
Arthropoda
47
Cumacea
Arthropoda
Gammarus sp
44
Amphipoda
Arthropoda
43
Isopoda
Arthropoda
39
38
Gastropoda
Mollusca
33
34
31
32
29
30
26
27
28
4
x
52
x
x
x
8
12
x
x
164
x
x
20
8
4
4
x
x
x
x
116
20
x
28
912
8
x
4
x
x
x
x
x
4
36
x
x
12
x
4
5
x
x
x
x
x
5
50
x
x
25
x
x
x
4
x
x
x
x
x
12
76
x
20
x
x
4
x
x
x
x
4
x
32
4
204
x
20
20
x
x
x
x
x
x
x
x
788
x
12
8
x
x
x
x
x
x
x
x
96
x
x
4
x
x
x
24
40
x
x
x
x
108
x
x
12
x
x
x
x
68
16
x
x
x
36
x
x
x
x
x
x
x
28
x
x
x
x
x
x
x
x
x
x
40
x
x
x
x
x
x
x
x
x
x
16
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
12
x
x
x
x
64
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
428
x
x
x
x
x
x
x
x
x
x
656
x
x
x
x
x
x
x
Continued on p. 146
x
x
x
156
x
x
x
x
x
x
x
145
sp
S3
S4
S5
S6
S7
S8
S9
S10
S11 S12
Actinidae
sp
sp
Annelida
Polychaeta
Oligochaeta
Annelida
0.01
sp3
Spinculida
x
x
sp1
sp2
S1
2
3
Nemertini
Cnidaria
Taxa
S2
0.99
0.36
0.05
0.30
x
0.26
S3
x
x
S4
x
x
0.15
S5
x
x
S6
x
x
S7
0.01
x
x
S8
x
x
S9
x
x
S10
0.01
x
x
S11
x
x
S12
x
x
S13
x
x
S14
x
x
S15
x
x
S16
x
x
S17
x
x
S18
x
x
S19
x
x
S20
420 9856 1112 3265 360 628 3124 1196 2632 2268 888 1628 568 516 900 304 212 632 444 664
S2
Appendix 2. List of the taxa identied in the Lima estuary. For each taxa, the average biomass (g AFDW/m2) is given (x absent)
Total
Echinoidea
54
S1
Echinodermata
53
Echinodermata
Ophuroidea
52
Insecta
Arthropoda
Taxa
Appendix 1. (Continued)
146
16
17
18
19
20
21
22
24
25
26
27
28
29
30
31
32
33
34
35
36
37
Mollusca
Gastropoda
23
Bivalvia
Mollusca
15
14
12
13
11
0.07
x
0.77
0.05
0.04
0.01
x
x
x
x
0.06 x
x
x
0.38 0.88
0.03 x
x
x
0.02 0.04
1.97 x
0.06
0.13
x
x
0.05
0.04
0.05
0.16 0.80
x
x
x
x
x
0.44
0.04 x
0.01 x
x
x
0.03 x
x
x
x
x
0.33 x
x
x
0.13 0.68
0.01 0.01
x
x
0.02 x
0.01 x
x
x
x
0.32 0.05 x
x
x
x
x
x
x
x
x
0.01 x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
0.03 x
0.01 x
0.45 x
0.01 x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
0.07 x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
5.32 x
x
x
x
x
Continued on p. 148
x
x
1.03 x
x
x
0.01 x
x
x
0.02 3.67 2.25 0.69 1.21 0.36 0.27 0.24 0.52 0.12 0.34 x
x
x
0.20 x
0.01 x
0.31 0.01 x
x
0.19 0.01 x
0.02 x
x
x
0.39 x
x
x
0.03
x
0.06
0.01 x
0.35 x
0.03 x
0.01
0.01 0.01
x
x
x
0.02 x
x
x
x
x
10
147
42
46
Diastylis sp
50
51
sp
0.01
0.33 0.03
x
x
0.02 0.01
0.01 x
0.03 x
x
x
0.08 x
0.63 1.72
x
x
S2
Total
54
Echinoidea
Echinodermata
x
S4
x
x
0.14 x
0.11 x
0.01 0.39
x
x
0.04 0.54
0.75 0.70
x
x
0.02 x
S3
x
S6
S7
0.1
x
x
x
x
x
x
x
x
0.01 0.09
x
x
0.62 0.01 x
x
0.01 x
0.08 x
S5
x
x
S9
x
x
0.03 x
x
x
x
x
x
x
x
x
2.36 0.03 0.01 x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
0.01 0.01 0.02 0.01 x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
0.02 x
x
x
x
x
x
x
x
x
x
x
0.02
x
x
x
x
x
0.10 0.23 0.28
x
x
S10 S11 S12 S13 S14 S15 S16 S17 S18 S19 S20
x
x
S8
1.55 24.53 8.14 28.31 2.13 0.70 264.62 3.58 4.76 2.46 0.73 1.23 0.37 0.27 0.24 1.58 0.12 0.44 5.55 0.30
Ophuroidea
53
Amphipholis squamata (Delle Chiaje, 1829) 0.01 x
Echinodermata
52
Insecta
Arthropoda
48
49
Decapoda
Arthropoda
47
Cumacea
Arthropoda
Gammarus sp
Corophium multisetosum (Stock, 1952)
44
45
Amphipoda
Arthropoda
43
Isopoda
Arthropoda
40
41
38
39
S1
Taxa
Appendix 2. (Continued)
148