Major food web properties of two sandy beaches

with contrasting morphodynamics, and effects on the stability

Leandro Bergamino

Julio Gomez

Francisco R. Barboza

Diego Lercari
Received: 8 February 2013 / Accepted: 24 April 2013 / Published online: 1 May 2013
Springer Science+Business Media Dordrecht 2013
Abstract We determined major structural properties
inuencing the food webs of two sandy beaches with
contrasting morphodynamics in the Atlantic coast of
Uruguay: reective (narrow and steep) and dissipative
beaches (wide and at). Furthermore, we evaluated
how these characteristics could inuence the stability
of the local food webs. To this end, we examined the
correlation of several food web properties with
different ecosystem types (including freshwater hab-
itats, estuary, marine, and terrestrial environments)
using a principal components analysis. Sandy beach
food web components included detritus, phytoplank-
ton, zooplankton, benthic invertebrates, shes, and
seabirds. Our results revealed that the dissipative
beach presented higher trophic levels, a higher number
of trophic species, more links per species, as well as a
higher proportion of intermediate trophic species, but
lower connectance and proportion of omnivorous
species than the reective beach. The variation in the
food web properties was explained by two principal
components. Sandy beach food webs contribute
mainly to one dimension of the principal components
analysis that was determined by the number of trophic
species, links per species, the trophic similarity, and
the characteristic path length. We suggest that species
and link characteristics, such as predominance of
scavengers and detritivorous, the relatively high
connectance and the short path length are drivers in
the food web structure and may play a role in the
community dynamic.
Keywords Food web structure Trophic
interactions Binary food webs Sandy beaches
Uruguay
Introduction
Sandy beach ecosystems are physically stressful
habitats where the community structure of the macro-
fauna is assumed to be controlled mainly by physical
factors, such as sediment characteristics and the
intertidal swash climate, while the biological interac-
tions have generally been regarded as negligible
(Defeo and McLachlan 2005; McLachlan and Brown
2006). There are few studies that suggest the effect of
biotic interactions (e.g., competition, food availability)
in the community structure on sandy beaches (Defeo
et al. 1997; Dugan et al. 2004; Rodil et al. 2012).
Exposed sandy beaches are classied based on the
interaction between sand, wave and tidal regimes. This
Handling Editor: Piet Spaak.
L. Bergamino (&)
Department of Zoology and Entomology,
Rhodes University, P.O. Box 94,
Grahamstown 6140, South Africa
e-mail: lbergamino@gmail.com
L. Bergamino J. Gomez F. R. Barboza D. Lercari
UNDECIMAR, Facultad de Ciencias, Igua 4225, 11400
Montevideo, Uruguay
1 3
Aquat Ecol (2013) 47:253261
DOI 10.1007/s10452-013-9440-5
produces a range of beach morphodynamic types,
varying fromreective (narrowand steep) to dissipative
beaches (wide and at), as sand becomes ner and
waves and tides larger (Short 1999). Species richness,
total abundance, and biomass of the resident biota
increase from reective to dissipative beaches, and
biological interactions (e.g., competition, predation) are
overshadowed by physical factors on reective beaches,
but become more inuential on dissipative beaches
(Defeo and McLachlan 2005; Schlacher et al. 2008).
The analysis of the food web structure and trophic
relationships in sandy beach ecosystems has become a
growing area of research (Colombini et al. 2011).
Sandy beach food webs are mainly based on marine
resources, such as phytoplankton, wrack (stranded
algae and sea grasses), and carrion (McLachlan and
Brown 2006). These sources support a macroscopic
food web comprising mainly of scavengers and deposit
feeders as primary consumers, while carnivorous
shes and polychaetes are consumers, which in turn
may be preyed upon by birds (Heymans and McLach-
lan 1996; Lercari et al. 2010; Bergamino et al. 2011).
Recent studies have attempted to elucidate trophic
pathways on sandy beach ecosystems, pointing out that
trophic pathways and food web complexity can be
strongly linked to morphodynamic factors which
inuence the occurrence and abundance of phyto-
plankton (Lercari et al. 2010; Bergamino et al. 2011).
Moreover, recent ndings suggest that the intertidal
fauna is mainly supported by marine resource inputs
(Paetzold et al. 2008; Colombini et al. 2011). Despite
these advances, information describing the food web
structure as such and key network properties that
determine food web dynamics on sandy beaches
remains scarce (but see Lercari et al. 2010).
Food webs describe feeding relationships between
taxa within ecosystems with structural patterns in the
arrangement of feeding links (Camacho et al. 2002;
Dunne et al. 2002; Pimm 2002). Understanding these
patterns is a key aspect of food web ecology, being
crucial for the description of ecosystem functioning
and important for the understanding of the biological
processes that underlie community organization
(Cohen et al. 1990; May 2006). For this reason,
certain topological properties have been studied to
discern food web patterns, including the proportion of
predators and prey, and the number of trophic links
(e.g., Dunne et al. 2004; Stouffer et al. 2005; Romanuk
et al. 2006; Sanchez-Carmona et al. 2012). Previous
work suggests that connectance and the number of
species are important factors in the structure of food
webs in different ecosystems (Vermaat et al. 2009), as
well as being a measure of the robustness of food webs
to species loss (Dunne et al. 2002).
In the present study, we analyzed the structure of
sandy beach food webs and determined major struc-
tural properties of the food web to evaluate their
ecological implications for the functioning of sandy
beach ecosystems. To this end, binary food webs were
used, considering number of species and links per
species, for two sandy beaches with contrasting
morphodynamics. A total of 17 food web properties
were calculated and then examined with published
food web models of other ecosystems to identify
drivers of food web structure.
Methods
Study area
We analyzed the network structure of two exposed
microtidal sandy beaches (tidal range = 0.5 m) with
contrasting morphodynamics located on the Uru-
guayan Atlantic coast: Arachania (reective) and
Barra del Chuy (dissipative) (Fig. 1). The former is
narrow (width approximately 40 m) and contains
coarse sediments (mean grain size = 0.56 mm) and
a steep slope (7.80 %), whereas the dissipative beach
is wider (width approximately 70 m), consisting of
ne sands (mean grain size = 0.20 mm) with a gentle
slope (3.53 %). Among all Uruguayan beaches, this
dissipative beach represents the highest macrofauna
richness, abundance, and biomass, whereas the reec-
tive beach presents relatively low macrofauna species
richness (Lercari and Defeo 2006). A full character-
ization of the main properties of both beaches is
provided in Defeo et al. (1992, 1997) and Gomez and
Defeo (1999).
Data collection and food web construction
The food web structures for these sandy beaches were
previously analyzed by Lercari et al. (2010) using the
ECOPATHII mass balance model software (Polovina,
1984; Christensen and Walters 2004). Both models
254 Aquat Ecol (2013) 47:253261
1 3
considered all main species identied in the ecosys-
tems. The dissipative beach model included 20 trophic
groups and the reective beach model integrated 9
trophic groups, including benthic invertebrate species
as groups, as well as 1 sh group, 1 bird group, 1
plankton group, and 1 detritus group (Table 1).
Functional feeding groups were used to group shes
and birds species according to similar feeding and
habitat characteristics. For the macroinvertebrate
species, the following sample design was followed
on both beaches: three transects perpendicular to the
shoreline and spaced 8 m apart, with sampling units
(SUs) on each transect every 4 mbeginning at the base
of the dunes to the lower limit of the swash zone. At
each SU, a sheet metal cylinder (27 cm in diameter)
was used to remove the sediment up to a depth of
40 cm. Each SU was sieved through a 0.5 mm mesh,
and the organisms retained were xed in 5 % buffered
formalin (see Defeo et al. 2001 for details). The origin
of the trophic links information was extracted from
published information, qualitative records, and stable
isotope analysis determined for some of the consum-
ers. Detailed explanations on the data source for diet
composition of the trophic groups are provided in
Lercari et al. (2010).
Binary networks were constructed to represent the
food web for each sandy beach using Network3D
Software (Williams 2010), developed for previous
food web studies (e.g., Williams and Martinez 2000,
2008; Williams et al. 2002). Input data were set in a
two-column format: a consumers number appears in
the rst column, and one of its resources numbers
appears in the second column.
Food web properties and data analyses
We analyzed seventeen food web properties that
describe species and link characteristics, as well as
food chain properties (see Table 2 for denitions). In
binary food webs, the most accurate trophic level
estimation is called the mean short weighted trophic
level (TL) which is the mean of shortest TL and prey-
averaged TL (Williams and Martinez 2004). For
denitions of the terms used to describe food web
properties, here please refer to Table 2.
In order to investigate the sandy beach food webs in
a global context, we considered 10 of the food webs
described by Dunne et al. (2004). We included two
terrestrial systems: the Coachella Valley Desert
located in California, USA (16637
0
W, 3354
0
N;
area = *740 km
2
; number of trophic species = 29;
Polis 1991) and the Caribbean island of St. Martin in
the northern Lesser Antilles (1804
0
N, 6303
0
W;
number of trophic species = 42; Goldwasser and
Roughgarden 1993); one marine system: the upwell-
ing Benguela current ecosystem (275
0
S, 113
0
E;
number of trophic species = 29; Yodzis 1998); three
freshwater lakes and pond webs: Bridge brook Lake,
South
America
Uruguay
Brazil
South Atlantic Ocean
Arachania
Barra del Chuy
Argentina
Fig. 1 Map of Uruguay
showing the two sandy
beaches analyzed in this
study: Barra del Chuy and
Arachania indicated by a
black circle. Map produced
using SimpleMappr
(Shorthouse 2010)
Aquat Ecol (2013) 47:253261 255
1 3
Upstate New York, USA (442
0
N, 74W; number of
trophic species = 25; Havens, 1992), Little Rock
Lake located in northern Wisconsin, USA (456
0
N
894
0
W; number of trophic species = 92; Martinez
1991), Skipwith Pond in the North Yorkshire, England
(5340
0
N, 059
0
W; number of trophic species = 25;
Warren 1989); and three estuary webs: Chesapeake
Bay in Eastern USA (3650
0
to 3940
0
N; number of
trophic species = 31; Baird and Ulanowicz 1989), St.
Marks Estuary located in Florida, USA (3006
0
N,
8411
0
W; number of trophic species = 48; Christian
and Luczkovich 1999), Ythan Estuary in NE Scotland
(157
0
W, 5720
0
N; number of trophic species = 83;
Hall and Raffaelli 1991).
Since most of the food web characteristics are
correlated (Vermaat et al. 2009), we used principal
components analysis (PCA) to account for the covari-
ance structure of the food web metrics. In this way, we
reduce data dimensionality revealing the similarities
between individual samples and the relationship
between the measured properties. To this end, the R
package for multivariate analysis FactoMineR (Hus-
son et al. 2011) was used. The number of retained
dimensions in the PCA was determined by taking into
account the percentage of variance explained by these,
considering 75 % as a reference. Since we consider
each single beach a unique ecosystem with dynamic
properties, statistical comparison in the food web
properties is not possible.
Results
The dissipative and the reective beach showed
differences in several structural properties of the food
web (Table 2). The mean trophic level and the
maximum trophic level were higher in the dissipative
beach (2.27 and 3.34, respectively) than in the
reective (2.13 and 3.25, respectively). Both food
webs presented a high trophic similarity with a
predominance of intermediate trophic level species
(85 % in the dissipative beach and 67 % in the
reective beach). The dissipative beach presented
higher links per species (2.95) than the reective
beach (2.11), but the reective beach showed a higher
connectance (0.23) than the dissipative beach (0.15).
Moreover, the percentage of omnivorous species was
50 % in the dissipative beach and 55 % in the
reective beach. Functional groups from the dissipa-
tive beach showed higher standard deviation of
vulnerability and generality (1.06 and 1.03, respec-
tively) than those from the reective beach (0.76 and
0.90, respectively). This result indicated that in the
dissipative beach system, a trophic species presented
greater variability in the number of prey organisms
and the number of predators than in the reective
beach system.
As a result of the PCA, only 2 dimensions were
retained, explaining 75.27 % of the variance. Figure 2
shows sandy beaches (dissipative and reective) and
Bridge BrookLakegroupedtogether. These ecosystems,
Table 1 Functional groups considered for the food web
models of the dissipative beach (Barra del Chuy) and the
reective beach (Arachania), located in the Atlantic coast of
Uruguay
Dissipative beach Reective beach
Birds Fishes
Fishes Polychaeta
Polychaeta Hemipodia californiensis
Hemipodia californiensis Amphipoda
Euzonus (Thoracophelia)
furcifera
Atlantorchestoidea
brasiliensis
Spio (Microspio) gaucha Isopoda
Carabide Excirolana braziliensis
Gastropoda Decapoda
Olivancillaria auricularia Emerita brasiliensis
Olivella formicacorsii Bivalvia
Buccinanops duartei Donax hanleyanus
Bivalvia Zooplankton
Amarilladesma mactroides Phytoplankton
Donax hanleyanus Detritus
Amphipoda
Atlantorchestoidea
brasiliensis
Phoxocephalopsis sp.
Isopoda
Chiriscus giambiagiae
Excirolana braziliensis
Excirolana armata
Decapoda
Emerita brasiliensis
Zooplankton
Phytoplankton
Detritus
256 Aquat Ecol (2013) 47:253261
1 3
together with Ythan Estuary, exhibited an important
contribution to the construction of Dimension 2
(Table 3). Dimension 2 was mainly determined by the
number of trophic species, links per species, mean food
chain length, fraction of basal species, and the trophic
similarity (Table 4). In this sense, these variables were
possibly responsible for the cluster formed by the
beaches and the lake systems studied, and the isolated
position of Ythan Estuary.
Similarly, Dimension 1 contained some ecosystems
that particularly contributed to its construction
(Coachella Desert; Skipwith Pond; Ythan Estuary;
Chesapeake Bay; Table 3). Connectance, fraction of
intermediate species, fraction of species that are
cannibalistic, fraction of top species, and normalized
standard deviation of vulnerability (number of con-
sumers per taxon) were the main variables that
determined Dimension 1 (Table 4), being responsible
for the groups described in relation to this dimension.
Skipwith Pond and Coachella Desert formed a sepa-
rate group (Fig. 2).
Discussion
Our analysis revealed that the dissipative and the
reective beaches presented differences in the struc-
tural properties of the food web. The reective beach
had higher degree of connectance and proportion of
omnivorous species, but lower trophic levels, lower
number of trophic species, links per species, and
proportion of intermediate trophic species than the
Table 2 Food web properties of the two contrasting sandy
beaches: dissipative and reective
Food web
properties
Dissipative Reective Denition
Trophic
species (S)
20 9 Number of trophic
species
Links/Species
(L/S)
2.95 2.11 Number of all
trophic links in
the web
(L) divided by S
Connectance
(C)
0.15 0.23 Proportion of all
possible links
that are realized
(L/S
2
)
Percentage of
top predators
(%Top)
5 11 Species with prey
but no predators
Percentage of
intermediate
species
(%Int)
85 67 Species with both
prey and
predators
Percentage of
basal species
(%Bas)
10 22 Species with
predators but no
prey
Percentage of
herbivores
(%Her)
40 22 Species which are
strictly herbivore
Generality
standard
deviation
(GenSD)
1.03 0.9 Number of
resources per
taxon normalized
Vulnerability
standard
deviation
(VulSD)
1.06 0.76 Number of
consumers per
taxon normalized
Link Standard
deviation
(LinkSD)
0.6 0.42 Number of links
per taxon
normalized
Percentage of
omnivores
(%Omn)
50 55 Taxa that feed on
taxa at different
trophic levels
Maximum
trophic
similarity
(MaxSim)
0.81 0.79 Number of
predators and
prey shared in
common divided
by the pairs total
number of
predators and
prey
Percentage of
cannibals
(%Can)
5 11 Taxa that feed on
their own taxa
Trophic level
(TL)
2.27 2.13 Short weighted
trophic level
Table 2 continued
Food web
properties
Dissipative Reective Denition
Maximum
trophic level
(MaxTL)
3.34 3.25 Maximum short
weighted trophic
level
Chain length
(ChaLen)
2.15 2 Mean food chain
length, averaged
over all species
Characteristic
path length
(Path)
1.79 1.55 The mean shortest
food chain length
between species
pairs
The description of the food web properties was taken from
Williams and Martinez (2000) and Dunne (2009)
Aquat Ecol (2013) 47:253261 257
1 3
dissipative beach. Moreover, consumers in the dissi-
pative beach seem to have more generalized diets than
consumers in the reective beach. These results are in
accordance with a previous, more detailed, food web
study for these sandy beaches (Lercari et al. 2010). It
should be noted that trophic species aggregation has
strong inuence on the measurement of %top and
basal species (Martinez 1991). This fact could explain
the over-estimation of these parameters on sandy
beaches since many taxonomic species in our model
were basal resources and top predators.
Our analyses are based on a high-resolution
description following grouping strategy using indi-
vidual species and trophic link information based on
Lercari et al. 2010. In this sense, our results provided a
robust comparison of the food web properties between
the sandy beaches analyzed here.
Sandy beach food webs are dominated by interme-
diate trophic level species, such as lter and deposit
feeders, being food webs characterized by low chain
length (Heymans and McLachlan 1996; Lercari et al.
2010; Colombini et al. 2011). Previous food webs
studied on sandy beaches revealed that the maximum
trophic levels range from 3.82 with 16 compartments
(Heymans and McLachlan, 1996) to 3.14 with 20
compartments (Lercari et al. 2010). Including the dune
system, the maximum trophic levels of the top
predators in the beach-dune system were 3.51 with
51 compartments (Colombini et al. 2011). On dissi-
pative beaches, the presence of a productive surf zone
with diatom accumulation provides large amounts of
food available for lter feeders and could explain the
high trophic similarity with the dominance of inter-
mediate trophic species (Defeo and McLachlan 2005).
On reective beaches, the harsh swash environment
with dynamic and turbulent swashes, and where waves
break directly on the steep beach face, may exclude
organisms without active and rapid burrowing abilities
at low and medium beach levels (Defeo et al. 2001;
Incera et al. 2006). Moreover, it has been suggested
that reective beaches are more stable and safer
environments for the development of supralittoral
species due to the lower risk of immersion and being
washed away (Defeo and Gomez 2005). Supralittoral
species include mostly primary consumers and sec-
ondary consumers (Colombini et al. 2011) such as
insects and talitrid amphipods.
Our study showed that number of species, links per
species, trophic similarity, and characteristics path
length are the major aspects inuencing the food web
structure on sandy beaches. When contrasted with
published information for other food webs, the
proportion of intermediate species on the reective
beach showed similar values to Mediterranean streams
(66 %), while the dissipative beach was close to lake
systems (range 6886 %) (Dunne et al. 2004; San-
chez-Carmona et al. 2012). These values were lower
than those observed in marine systems (range 9295),
but higher than for streams (2227 %) and slightly
higher than in estuarine systems (5669 %) (Dunne
et al. 2004). In our food webs, benthic invertebrates
were mainly scavengers and detritivorous. Moreover,
in both sandy beaches, the production is poorly
-4 -2 0 2 4 6
-
3
-
2
-
1
0
1
2
3
4
Principal component 1 (44.48%)
P
r
i
n
c
i
p
a
l

c
o
m
p
o
n
e
n
t

2

(
3
0
.
7
9
%
)

Ythan Estuary
Bridge Brook Lake
Skipwith Pond
Cheasapeak Bay
Coachella Desert
St. Martins Islands
Reflective Sandy Beach
Dissipative Sandy Beach
Fig. 2 Principal component analysis for aquatic and terrestrial
ecosystems considering the food web properties described in
Table 2. Percentage values represent the proportion of the
variance explained by each principal component
Table 3 Contribution percentages of aquatic and terrestrial
ecosystems to the dimensions considered in the principal
component analysis
Ecosystems Contributions
Dimension 1 Dimension 2
Skipwith Pond 19.44 2.24
Bridge Brook Lake 0.73 13.74
Chesapeake Bay 12.82 0.21
Ythan Estuary 23.38 32.55
Coachella Desert 41.61 10.35
St Martin Island 1.74 2.31
Dissipative sandy beach 0.23 13.45
Reective sandy beach 0.05 25.16
Values in bold indicate contributions higher than 10 %
258 Aquat Ecol (2013) 47:253261
1 3
consumed (4 % on the dissipative 6 % on the reec-
tive), and most of the biomass ows are directed to
exportations and detritus (Lercari et al. 2010). This
suggests that weak interaction effects of primary
consumers on resources are the most frequent inter-
action in these food webs. Moreover, donor control
dynamics are expected, in which the rate of detrital
input is thought to be a major factor inuencing the
interactions within the macrobenthic community
(Pimm 2002). This pattern can enhance the stability
of these food webs, in the sense that the system
recovers faster after a disturbance, by dampening
uctuations of populations densities (May 1973;
McCann et al. 1998; Neutel et al. 2002; Montoya
and Sole, 2003).
We found that the connectance values calculated
for sandy beaches were relatively high compared with
previous works that analyzed 16 food webs and
reported a range of 0.030.32 (Dunne et al. 2002,
2004). The connectance value for the dissipative beach
was close to the Bridge Brook Lake (0.17) and the
value for the reective beach to marine systems (range
of 0.220.24). However, our connectance results were
lower than those of a terrestrial system (the Coachella
Valley, 0.31), and lake/pond (0.32) food webs, while
they were higher than Mediterranean streams (range
0.090.14) and estuarine webs (0.040.1) (Dunne
et al. 2004; Sanchez-Carmona et al. 2012). It has been
suggested that connectance may increase food web
robustness to species extinction and ecosystem stabil-
ity, and that this effect is more important than diversity
(Dunne et al. 2002, 2004; Fussman and Heber 2002;
Kondoh 2003). In this case, robustness of a food web
refers to the propensity for networks to fragment and is
dened in terms of the number of secondary extinc-
tions that result from primary species loss. Moreover,
previous work reported that connectance is a good
predictor of omnivory and that more omnivorous links
increase ecosystem stability (Fussman and Heber
2002). In our results, the reective beach showed a
smaller number of species but higher connectance and
omnivory, which could result in a greater robustness to
species loss (i.e., less secondary extinction occur) than
the dissipative beach which showed intermediate level
of connectance and lower levels of omnivory. In spite
of this, our results showed that sandy beach food webs
present lowmean path length on both beaches (1.79 on
the dissipative and 1.55 on the reective beach),
suggesting that species are highly interconnected
within the ecosystems. This fact has important
ecological implications, suggesting that change in
diversity, by the loss of species (e.g., caused by habitat
loss) or the introduction of new species, can be
propagated through the ecosystem, thereby affecting
the ecosystem structure (Williams et al. 2002). These
results open a new question on sandy beach ecology
concerning the effects of biodiversity loss on the food
web structure.
In summary, our results show new potential effects
of food web interaction patterns in community struc-
ture and dynamic on sandy beaches. This has impor-
tant consequences for conservation issues. The
understanding of species interactions allows predict-
ing the response of ecosystem function to changes in
structural aspects such as the effects of invasive
species and local extinction. Our food web analysis
suggested that species and link characteristics, such as
trophic similarity, number of species, and links per
species, play a critical role structuring the food webs
on sandy beach ecosystems. Moreover, the predom-
inance of weak trophic interactions of primary
consumers and the relatively high connectance could
enhance the stability of these ecosystems and act
together with the strong physical forces in structuring
populations and communities. Although sandy beach
populations are mainly controlled by physical factors,
the effects of interaction patterns on the community
structure and stability remain open. We think that for
Table 4 Contribution percentages of food web properties to
the dimensions considered in the principal component analysis
Variables Contributions
Dimension 1 Dimension 2
S 4.74 17.68
L/S 4.62 16.27
C 17.18 0.26
%Top 11.59 6.36
%Int 14.69 0.16
%Bas 1.43 10.80
GenSD 5.72 0.07
VulSD 11.92 2.52
%Omn 8.30 8.12
MaxSim 2.89 15.18
%Can 14.92 2.12
Path 2.00 20.46
Values in bold indicate contributions higher than 10 %
Aquat Ecol (2013) 47:253261 259
1 3
future works on sandy beaches, an important aspect to
analyze would be the stability of the population
dynamics against species loss by considering the role
of several structural factors of the networks (e.g.,
Dunne et al. 2002). The higher degree of connectance
and omnivory in the reective beach could enhance
the stability and robustness of the food web. In
comparison with the dissipative beach, the food web
may be more fragile to the loss of species with a
greater magnitude of secondary extinctions. This
could be tested by building different congurations
of both food webs simulating the sequential local
extinction of the groups and then comparing the
stability/robustness indicators.
Acknowledgments We thank Omar Defeo (Facultad de
Ciencias, UNDECIMAR, Uruguay) for his mentorship and
friendship through the years. This work was supported by
SANDISA IMWEBU grants (L.B). We thank Katherina Schoo
and Sydney Moyo for the language editing. We are also grateful
to Piet Spaak and an anonymous referee for helpful comments in
the manuscript. DL thanks PEDECIBA and ANII. L.B. thanks
Jesus Orozco (Rhodes University, South Africa) for his
transmission of knowledge and encouragement in a friendly way.
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