Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 November 2021 | 13(13): 20000–20010
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
https://doi.org/10.11609/jott.6340.13.13.20000-20010
#6340 | Received 29 June 2020 | Final
received 20 December 2020 | Finally accepted 12 October 2021
Niche overlap of benthic macrofauna in a tropical estuary: diurnal variation
Mário Herculano
de Oliveira 1, Lidiane Gomes de Lima
2, Caroline Stefani da Silva Lima 3, Jéssica
de Oliveira Lima Gomes 4, Franciely
Ferreira Paiva 5, Graciele de Barros
6, Carlinda Railly
Medeiros 7 &
Joseline Molozzi 8
1–6 Programa de Pós-Graduação
em Ecologia e Conservação, Universidade Estadual da Paraíba, Paraíba, Brazil.
7 Programa de Pós-graduação
em Ecologia, Universidade Federal do Pará, Pará, Brazil.
8 Departamento de Ciências
Biológicas e da Saúde; Programa de Pós-graduação em Ecologia e Conservação,
Universidade Estadual da Paraíba, 58429-500, Campina Grande, Paraíba,
Brazil.
1 mariohecules@hotmail.com
(corresponding author), 2 lidianelimauepb@gmail.com, 3 carollinnestefani@hotmail.com,
4 lima.jessica.bio@gmail.com, 5
franciely0903@gmail.com, 6 gracibarross@gmail.com, 7 carlindarailly@gmail.com,
8 jmolozzi@gmail.com
Editor: Anonymity
requested. Date of
publication: 26 November 2021 (online & print)
Citation: Oliveira, M.H.de., L.G.de. Lima,
C.S.da.S. Lima, J.de.O.L.
Gomes, F.F. Paiva, G.de. Barros, C.R. Medeiros & J. Molozzi
(2021). Niche overlap of benthic macrofauna in a tropical estuary: diurnal variation. Journal of Threatened Taxa 13(13): 20000–20010. https://doi.org/10.11609/jott.6340.13.13.20000-20010
Copyright: © Oliveira et al. 2021. Creative Commons Attribution
4.0 International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: None.
Competing interests: The author
declares no competing interests. The views expressed are those of
the author.
Author details: All the authors were linked to
the Programa de Pós-Graduação
em Ecologia e Conservação (PPGEC-UEPB) during sampling procedures and MS
writing process.
Author contributions: MHO—methodology elaboration, data
sampling, and writing process | LGL—methodology elaboration, data sampling, and
writing process | CSSL—data analyzes and writing
process | JOLG—methodology elaboration, data sampling, species identification,
and writing process | FFP—methodology elaboration, data sampling, and writing
process | GB—species identification, data sampling, methodology elaboration,
and writing process | CRM:—species identification and writing process |
JM—methodology elaboration, data sampling, data analyzes,
and writing process.
Acknowledgements: We thanks the anonymous
reviewers for their comments that improved the manuscript.
Abstract: The complexity of estuaries
allows for the establishment of diverse communities composed of species with
different survival strategies. The vertical migration of animals in the
sediment is linked to competition, escape from predators and adaptations to diurnal
physio-chemical changes related to variations in water levels. The present
study aimed to evaluate niche overlap and amplitude, as well as the composition
and structure, of communities of polychaetes and molluscs between sediment
aliquots during the day and at night. Data sampling was performed in the Tubarão River estuary. The highest individual occurrence
was registered during the diurnal period. Communities of polychaetes varied
significantly between sediment aliquots during the day and at night, while
molluscs did not show diurnal variation. Niche overlap results for
polychaetes showed higher values between aliquots during the night, while
molluscs showed greater overlap during the day. This indicates that polychaetes
and molluscs have different mechanisms of coexistence. This may be related to
different attributes of species allowing for the division of resources among
individuals. Examination of niche overlap provides insights into coexistence of
mechanisms within benthic macroinvertebrate communities.
Keywords: Coexistence, hypersaline
estuary, vertical migration, Mollusca, Polychaeta.
Introduction
Estuaries are considered highly
productive, complex and heterogeneous ecosystems that allow the establishment
of diverse organisms that depend on these habitats for feeding, reproduction,
colonization, and protection against predators (Elliot & MCluscky 2002; Kennish 2002;
Kaiser et al. 2005; Vasconcelos et al. 2007; Potter et al. 2010). Estuarine
dynamics are influenced by seasonal, monthly & daily variations, and
consequently the concentrations of organic matter, nutrients, and water level
can oscillate during these periods (Dyer 1979). These variations come from
chemical, physical and biological alterations, causing changes at spatial and
temporal scales in the functioning of biological communities (Officer &
Lynch 1981; Day et al. 2012; Medeiros et al. 2016).
Among the communities that
inhabit estuarine ecosystems, benthic macroinvertebrates, mainly represented by
Polychaeta and Mollusca, have important roles in the decomposition of organic
matter, nutrient cycles, and energy fluxes (Nunes et al. 2008; Wildsmith et al.
2011; Tweedley et al. 2012). Benthic
macroinvertebrates can migrate vertically within the sediment according to
spatial and temporal scales, reacting to the variations of environmental
conditions (Cardoso et al. 2010). Thus, vertical migration of organisms in the
sediment can occur in function of their tolerance to limiting factors (e.g.,
light availability), life strategy (e.g., protection against predators) or
feeding habits (Cardoso et al. 2010).
This locomotive ability is related to the presence of determined
functional traits of each species (e.g., related to feeding category, larvae
development and body size) (Esselink & Zwarts 1989; Cruz-Motta 2005; Persson & Svensson 2006).
It has been observed in the
sand/mudflats of estuaries, that the vertical locomotion of invertebrates is
influenced according to light availability or tide cycle (Yannicelli
et al. 2001). In this case, the pattern of vertical migration in relation to
tide cycles, determines the density of macrofauna in
the sediment as a form to escape adverse conditions. That is, during low tide,
organisms can migrate to protect themselves from predators, while during higher
tides, organisms, when moving vertically, can maximize foraging opportunities
(Cardoso et al. 2010). Diurnal changes also generate vertical migration,
regulated according to light availability, that alter the visibility, and
consequently influence organisms’ survival in the presence of predators (Estlander et al. 2017). In addition, vertical migrations
can also be enhanced when the organisms of a community share the same
functional categories, this can result in greater niche overlap causing the
development of strategies which, consequently, results in the use of different
resources and organism coexistence (Silva-Camacho et al. 2017).
Hypervolume niche concept was
stated by Hutchinson (1957) who considered a niche as multidimensional space in
which species might be established in accordance with their demands relative to
abiotic factors and resources. Abiotic factors limit species distribution
through their physiological tolerance and space and food as resources limited
through competition among species when they are scarce (Kraft et al.
2015). Some strategies developed by
species viewing coexistence are: the species may use their resources
differently in spatial and temporal scales (Devictor
et al. 2010) or functional traits of species (Violle
2007) which allow them to explore a wider range
or to specialize in a narrow range of resources (Devictor
et al. 2010). In this way, these strategies are not only important to
understand the mechanism underlying community assembly processes but also to
understand how the complexity of species interactions influence their
occurrence and their environment defining ecosystem functioning (Dehling & Stouffer 2018).
Researchers have used different
parameters related to species’ niches, such as amplitude and niche overlap,
associated with other parameters of functional traits that can aid in the
understanding of species coexistence. However, the understanding of how benthic
macrofauna species utilize their habitat resources in
relation to vertical distribution, according to light availability (diurnal
variation), has been neglected. Understanding the pattern of vertical
distribution of species in the sediments of tropical estuaries and coexistence
strategies of benthic macrofauna, is a key factor in
describing the importance of these organisms in the maintenance of ecosystem
functions. As such, the aim of this study was to evaluate amplitude and niche
overlap, as well as the composition and structure of polychaete and mollusc
communities between sediment aliquots during day and night periods. The
questions that directed this study were: i) are there
differences in the composition and taxonomic structure of polychaete and
mollusc communities when comparing sediment aliquots analyzed
during the day and night? and ii) does niche overlap and the overlap of
polychaete and mollusc functional trophic groups occur, when comparing the
sediment aliquots during day and night?
Material
and Methods
Study area
The Tubarão
River is located in the semiarid coast of the state of Rio Grande do Norte, northeastern Brazil (-5.09361111, -36.53916667) (Figure 1),
inserted on the limits of the Sustainable Development Reserve of Ponta do Tubarão, a protected area with sustainable use administered
by the state government. The Tubarão River is an
estuarine system which extends a distance of 10 km, presents preserved mangrove
vegetation and its surroundings have a variety of ecosystems such as marshes,
Caatinga vegetation and dunes (Dias et al. 2007). Based on the precipitation
standards, the climate of the region is characterized by higher evaporation
rates and a lower pluviometric index with the
rainy season occurring between February and May and the dry season
occurring between June and January (INMET 2017). Additionally, the freshwater
input comes from subterranean waters and from the lower precipitation of the
region (Queiroz & Dias 2014, Medeiros et al. 2016). The average rainfall
between January and June is 161 mm (INMET 2010, 2017).
This study was performed
throughout the estuary, where we defined three zones: upstream, intermediate
and downstream. The sampling was carried out covering all the variation of the
environmental variables of the estuarine ecosystem, from the continental
drainage to the sea (Figure 1). The upstream zone presents lower profundity and
higher evaporation rates with salinity varying between 30 and 50 and the
predominant substrate is sandy and muddy. The intermediate zone presents
salinity between 32 and 45, with muddy substrates. The downstream zone is
located near the sea, with salinity
between 35 and 45, and the predominant substrate is sandy with an abundance
of gravel (Sales et al. 2016).
Sampling procedures
Samples were collected in May
2017, during the rainy season. In each zone (upstream, intermediate, and
downstream) three sampling points were determined, totaling
nine points across the estuary. The benthic macrofauna
was sampled once during the day and at night, utilizing a PVC cylindrical core
(16.67 cm²). The core was buried in the substrate (profundity 10 cm) and posteriorly
the sediment was partitioned in two aliquots: A1 (0 to 5 cm) and A2 (5 to 10
cm). The interval between diurnal and nocturnal sampling was 12 hours. The
benthic macroinvertebrates were sampled in the intertidal region, always during
low tide. The samples were washed in a 500 micro mesh sieve for the separation
of benthic macrofauna. Organisms were preserved in
70% alcohol.
Laboratory procedures
In the
laboratory, the organisms were separated and identified to the lowest possible
taxonomic level with specialized keys for Polychaeta (Amaral & Nonato 1996; Amaral et al. 2006), and a specialized
bibliography for Mollusca (Rios 2009; Tunnel Jr. et al. 2010). In addition, the
Conquiliologistas site of Brazil was consulted
(www.conchasbrasil.org.br) and the nomenclature followed the database Malacolog 4.1.1 (Rosenberg 2009).
Measurement of niche breadth and
overlap
Niche breadth and overlap
analyses were performed to quantify how many species and functional guilds
partitioned the spaces as resource between aliquots. The Morisita
simplified index (Krebs 1989) was calculated to analyze
the niche overlap of the functional trophic groups of polychaetes and molluscs
between day and night aliquots based on abundance data. The index value varies
from zero to one: when the index is zero or close to zero, the overlap is low
or absent, while when equal to or close to one, the niche overlap is high or
total. To analyze the taxonomic niche breadth and
functional trophic amplitude of polychaetes and molluscs groups, the
Shannon-Wiener (H’) diversity index was used based on abundance data (Shannon
& Weaver 1949), which corresponds to the niche breadth, considered high when
close to one. These analyses were carried out in the “spaa”
package, in the statistical software program R 1.9.0 (R Development Core Team
2017).
Composition and taxonomic
structure
For the analysis of the
composition and taxonomic structure of polychaetes and molluscs communities,
species richness was analyzed using the
Shannon-Wiener (H’) index and Pielou (J’)
equitability. Species richness reflects the number of different species in an
area, while the Shannon-Wiener index considers the richness and species
equitability, both abundant and rare (Shannon & Weaver 1949). The Pielou (J’) equitability index reflects the uniformity and
species’ distribution (Pielou 1966).
Functional trophic guilds
Macrofauna benthic taxa were classified
within functional trophic guilds according to the food resources consumed,
based not only on the type of resource, but also on the morphological and behavioral mechanisms of acquisition, following Fauchald & Jumars (1979) and Muniz & Pires (1999)
for polychaetes and Linden et al. (2017), Mikkelsen & Bieler (2008) and
Rosenberg et al. (2009) for molluscs (Table 1).
Five guilds were considered for polychaetes: Carnivore (Car), Deposit
feeder: digger (DC) e Deposit feeder: surface (DS), Filter feeder (Fil),
Omnivore (Omn), while six guilds were considered for
molluscs: Carnivore, Filter feeder, Herbivore (Her), Omnivore, Scavenger (Sce), and Suspension feeder (Sus). Functional trophic
guilds were chosen as they reflect the distribution of resources in the
ecosystem, the mechanism of adaptation and coexistence of species, in addition
to influencing ecosystem processes, such as energy flow and nutrient cycle
(Pearson & Rosenberg 1978; Dolbeth et al. 2015).
Data analyzes
To test the differences in the
taxonomic composition and trophic guilds of polychaetes and molluscs between
the sediment aliquots during the day and night, a Permutational Multivariate
Analysis of Variance (PERMANOVA) (with 9999 permutations, p ≤0.05) was
performed, considering a factor (aliquots) with two fixed levels (A1 and A2)
for day and night. For this, the abundance data was transformed into square root values and a
Bray-Curtis matrix was utilized as a dissimilarity measurement. To verify
differences in the taxonomic structure of the polychaetes and molluscs between
sediment aliquots during the day and night, analyzes
of univariate significance were applied to species richness, Shannon-Wiener
index, and Pielou equitability, considering a factor
(aliquots) with two fixed levels (A1 and A2) for day and night (PERMANOVA; 9999
permutations; p ≤0.05). For this analysis the coefficient of euclidean distance was utilized as a measure of
dissimilarity. All the analyzes were performed using
the statistical software PRIMER-6 + PERMANOVA (Anderson et al. 2008).
Results
Composition and taxonomic
structure of polychaete and mollusc communities
A total of 1,329 individuals were
captured and distributed across 19 and 21 polychaete and mollusc taxa,
respectively (Supplementary material: T1; Figure 2). The highest individual
occurrences (705) were registered during the diurnal period, accounting for 53%
of the total abundance. During the day, aliquot A1 obtained the highest
abundance of polychaetes, Cirratulidae (25%), Capitellidae (12%), and the bivalve Anomalocardia
flexuosa (11%), while the polychaetes Orbiniidae (20%), Ampharetidae
(15%), and the bivalve A. flexuosa (10%) had
the highest abundances in aliquot A2 (Figure 2). During the night, 624
individuals were captured with higher abundances registered in aliquot A1 for
the polychaetes Cirratulidae (28%), Spionidae (8%), and the bivalve A. flexuosa
(8%). In aliquot A2, the polychaetes Cirratulidae
(25%) and bivalves A. flexuosa (17%) & Phacoides pectinatus
(15%) were most abundant (Figure 2).
PERMANOVA results show that the
polychaetes group varied significantly between the sediment aliquots during the
day (Pseudo-F1,37= 2.3226; p= 0.027), as well as during the night
(Pseudo-F1,41= 2.8135; p= 0.007). The molluscs did not present
variation between the day and night sediment aliquots, (day: Pseudo-F1,31=
1.4734; p= 0.21; night: Pseudo-F1,37= 1.9075; p= 0.079). The species
richness, Shannon-Wiener index, and Pielou
equitability varied significantly between sediment aliquots during the day and
night for polychaetes and molluscs (Table 2).
Overlap and breadth of
polychaetes and molluscs taxonomic niches
Overlap results showed higher
values between the aliquots for polychaetes during the night and molluscs
during the day (Table 3). For polychaetes niche breadth, the taxa Orbiniidae, Ampharetidae, Pilargidae, and Onuphidae were
found to exhibit higher values when comparing day aliquots, while Ampharetidae, Pilargidae, Onuphidae, Dorvilleidae, and Nereididae demonstrated higher values at night (Table 3).
The molluscs, Veneridae sp., A. succinea, and Macoma
sp.1, exhibited higher niche breadths during the day and P. pectinatus, B. varium,
C. cariabaea, S. fragilis at night
(Table 3).
Overlap and breadth of functional
trophic guilds
The functional trophic guilds of
polychaetes that obtained higher proportions were Deposit feeder: digger,
Deposit feeder: surface, Carnivore and Filter feeder, while for molluscs, the
abundant guilds were Filter feeder, Suspension feeder, and Scavenger (Figure
3). For the polychaetes community, the guilds Deposit feeder: digger (42%),
Carnivore (24%), and Deposit feeder: surface (21%) were most abundant in
aliquot A1, while Deposit feeder: digger (44%), Deposit feeder: surface (33%)
and filter feeder (11%) were most abundant in aliquot A2 during the day. During
the night, Deposit feeder: digger (50%), Carnivore (18%), Deposit feeder:
surface (15%) and filter feeder (14%) presented higher abundance proportions in
aliquot A1, and Deposit feeder: digger (49%), Carnivore (27%), Deposit feeder:
surface (18%) in aliquot A2. Significant differences in the abundance of
trophic guilds were observed only between day and night periods for Molluscs
(Pseudo-F1,68 = 4.05; p= 0.0513) and between aliquots for
polychaetes (Pseudo-F1,76= 2.98; p= 0.0368).
The functional trophic groups of
polychaetes, Carnivore, Deposit feeder: surface and Omnivore exhibited higher
breadth during the night while Filter feeder were more representative during
the day (Table 4). For the molluscs, the only functional trophic group with
higher amplitude was Suspension feeder for both day and night periods (Table
4).
Discussion
The macrobenthic
community of the Tubarão River varied quantitatively
and qualitatively between day and night periods, exhibiting a pattern of
vertical distribution in the sediment, possibly determined by ecological
preferences and as a result of the interactions between species throughout the
estuary which was corroborated with Hutchinson’s niche concept (Hutchinson
1957) who stated that a combination of abiotic and biotic factors influence
species occurrences in given habitats relative to their preferences. These
results are an important contribution to understanding how macrobenthic
communities interact with each other and select their preferential habitats in
tropical estuaries. Other studies (e.g., Poole & Stewart 1976) have
demonstrated that variation in benthic macroinvertebrates communities exists in
function of sediment profundity, which corroborates our results. A study
conducted in an estuary of Portugal found that the vertical migration patterns
of macroinvertebrates depend mainly on the size and feeding behavior
of individuals (Cardoso et al. 2010).
Over short time-scales,
individuals can move within sediments as a result of feeding behavior or to escape from predators (Goeij
et al. 2001; Person & Svensson 2006). This may be
the reason why the polychaete and mollusc presented functional and taxonomic
differences between sediment aliquots. Vertical migration behavior
can also be observed in other communities, such as zooplankton, which migrate
from the water column to submerged macrophytes, avoiding open water predators (Sagrario & Balseiro 2010).
Our results showed that polychaetes and molluscs had a greater abundance in
aliquot A2 at night when compared to the day period. This result may be related
to a survival strategy used by these taxonomic groups to escape from predators
(e.g., fish) at night. In contrast, the considerable increase in nocturnal
carnivorous polychaetes may be related to feeding behavior
and food availability for these species at night. Other factors that can
generate vertical migration are reproductive processes that occur mainly during
seasonal fluctuations (Cardoso et al. 2010), although this was not tested in
this study.
The high niche overlap observed
for the polychaete and mollusc indicates that there are coexistence mechanisms
working in these communities, thus allowing the partitioning of resources
between individuals. However, the mechanisms of coexistence that act on
polychaetes and molluscs, act at opposing times. Different times for daily
foraging are strategies used by individuals to reduce competition, allowing the
partition and coexistence of food (Sánchez-Hernández et al. 2011). The
different types of trophic guilds used by the species in this study may explain
their coexistence. According to Kneitel & Chase
(2004), alternative traits can enable the exploration of the same type of
resource by different strategies, ensuring coexistence between species. This
may explain the fact that in this study we found different types of trophic
guilds, such as Filter feeder, Deposit feeder: digger, Deposit feeder: surface
and Suspension feeder, that utilize the same type of resource, with different
methods of capture.
The significant differences of
the mollusc trophic guilds between the aliquots may be explained by the
reduction in abundance of the three species that demonstrated the highest
abundance during the day. The variation in the contribution of Suspension
feeders occurred mainly by the elevation of Phacoides
pectinatus density during the day. Some species
of the Lucinidae family, such as P. pectinatus have specializations for surviving in
environments with a lower oxygen content, under these conditions these species
are likely to have symbiosis with sulfide oxidant
bacteria (Taylor & Glover 2006), which allows for the displacement of
individuals to areas of deep sediment.
It is possible that the higher
abundance of A. flexuosa in both aliquots was
due to the fact that the sampling period included the start of the rainy
season, which is linked to the greatest reproductive activity of the specie (Boehs et al. 2008). This may also explain the high
proportions of polychaete families, Capitellidae and Cirratulidae, according to Giangrande
& Simonetta (1993) and Gibbs (1971). These authors showed that these
species from both families increase their reproductive activity during the
month of May.
Conclusion
In conclusion, the polychaetes
and mollusks groups occupied different functional
guilds and presented dynamics of vertical migration in the sediment during the
diurnal period, which may be a strategy used by species to obtain food
resources, reproduce and persist in the environment under adverse conditions
(for example, escape predation). These results provide information on the
mechanisms of coexistence of benthic macroinvertebrate communities and may
support other studies aimed at the conservation of tropical estuaries.
Table 1. Functional trophic
guilds of polychaetes and molluscs and their importance in demonstrating the
relationships of organisms with the ecosystem.
|
|
Functional trophic guilds |
Rationale |
|
Polychaeta |
Carnivore |
Reflects the distribution of
resources in the ecosystem and the adaptation of the species to the
habitat. As well as influencing ecosystem processes such asenergy
flux and nutrient cycling (Pearson & Rosenberg 1978; Dolbeth
et al. 2015). |
|
Deposit feeder: digger |
||
|
Deposit feeder: surface |
||
|
Filter feeder |
||
|
Omnivore |
||
|
Mollusca |
Carnivore |
|
|
Filter feeder |
||
|
Herbivore |
||
|
Omnivore |
||
|
Scavenger |
||
|
Suspension feeder |
Table 2. PERMANOVA values for the
taxonomic diversity of polychaetes and molluscs between day and night sediment
aliquots (A1 e A2), in the Tubarão River estuary,
Macau-RN, northeastern Brazil.
|
Polychaeta |
|
|
|
|
|
|
|
|
|
|
|
|
Day |
(A1xA2) |
|
|
|
Night |
(A1xA2) |
|
|
|
|
|
Df |
SS |
MS |
Pseudo-F |
P(perm) |
Df |
SS |
MS |
Pseudo-F |
P(perm) |
|
Richness |
1 |
156.06 |
156.06 |
39.373 |
0.0001 |
1 |
110.01 |
110.01 |
32.225 |
0.0001 |
|
Shannon-Wiener |
1 |
10.542 |
10.542 |
44.901 |
0.0001 |
1 |
8.0869 |
8.0869 |
28.991 |
0.0001 |
|
Pielou |
1 |
4.2535 |
4.2535 |
38.865 |
0.0001 |
1 |
1.9306 |
1.9306 |
12.44 |
0.001 |
|
|
|
|
|
|
|
|
|
|
|
|
|
Mollusca |
|
|
|
|
|
|
|
|
|
|
|
|
Day |
(A1xA2) |
|
|
|
Night |
(A1xA2) |
|
|
|
|
|
Df |
SS |
MS |
Pseudo-F |
P(perm) |
Df |
SS |
MS |
Pseudo-F |
P(perm) |
|
Richness |
1 |
39.014 |
39.014 |
29.061 |
0.0001 |
1 |
66.125 |
66.125 |
15.651 |
0.0001 |
|
Shannon-Wiener |
1 |
4.7689 |
4.7689 |
33.107 |
0.0001 |
1 |
3.9247 |
3.9247 |
16.981 |
0.0002 |
|
Pielou |
1 |
2.6835 |
2.6835 |
22.399 |
0.0001 |
1 |
1.9569 |
1.9569 |
15.462 |
0.0004 |
Table 3. Results of the niche
overlap of polychaetes and molluscs abundance between day and night periods in
the Tubarão River estuary, Macau-RN, northeastern Brazil.
|
|
Polychaeta |
Day |
Night |
|
Niche overlap/Taxonomic Niche breadth |
A1 vs. A2 |
0.47 |
0.88 |
|
Orbiniidae |
0.47 |
0.17 |
|
|
Ampharetidae |
0.47 |
0.57 |
|
|
Pilargidae |
0.51 |
0.56 |
|
|
Onuphidae |
0.45 |
0.56 |
|
|
Dorvilleidae |
- |
0.63 |
|
|
Nereididae |
0.11 |
0.48 |
|
|
Cirratulidae |
0.03 |
0.37 |
|
|
Goniadidae |
- |
0.21 |
|
|
Capitellidae |
0.19 |
0.28 |
|
|
Spionidae |
0.20 |
0.10 |
|
|
Syllidae |
- |
0.24 |
|
|
Niche overlap/Taxonomic Niche breadth |
Mollusca |
Day |
Night |
|
A1 vs. A2 |
0.81 |
0.61 |
|
|
A. flexuosa |
0.20 |
0.54 |
|
|
Macomasp.1 |
0.29 |
0.17 |
|
|
P. pectinatus |
0.27 |
0.68 |
|
|
B. varium |
- |
0.63 |
|
|
Veneridae sp. |
0.33 |
- |
|
|
A. succinea |
0.32 |
- |
|
|
C. cariabaea |
- |
0.63 |
|
|
S. fragilis |
- |
0.63 |
|
|
Macomasp.2 |
- |
0.56 |
|
|
B. striata |
- |
- |
|
|
|
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Supplementary Material 1.
Composition, numerical abundance and functional trophic guilds of the taxa of
Polychaeta and Mollusca captured at day and at night in the Tubarão
River Estuary, Macau-RN, northeastern of Brazil.
Functional trophic guilds: Car—Carnivore | Fil—Filter feeder | Her—Herbivore |
DC—Depositer feeder: digger | DS—Deposite
feeder: surface | Omn—Omnivore | Sce—Scavenger | Sus—Suspension feeder.
|
Benthic macroinvertebrate communities |
|||
|
|
Day (A1/A2) |
Night (A1/A2) |
Trophic functional guilds |
|
Polychaeta |
|
|
|
|
Orbiniidae |
36/8 |
23/1 |
DC |
|
Ampharetidae |
27/6 |
23/8 |
DS |
|
Oweniidae |
1/0 |
0/0 |
DC |
|
Pilargidae |
11/3 |
9/3 |
DC |
|
Onuphidae |
5/1 |
6/2 |
Omn |
|
Dorvilleidae |
9/0 |
4/8 |
Car |
|
Nereididae |
41/1 |
13/3 |
Car |
|
Cirratulidae |
165/1 |
149/21 |
DC |
|
Glyceridae |
1/0 |
0/0 |
Car |
|
Goniadidae |
16/0 |
33/3 |
Car |
|
Capitellidae |
77/4 |
11/1 |
DS |
|
Lumbrineridae |
2/0 |
6/1 |
Omn |
|
Magelonidae |
1/0 |
7/0 |
DS |
|
Sternaspidae |
4/0 |
12/0 |
DS |
|
Spionidae |
55/3 |
45/1 |
Fil |
|
Syllidae |
56/0 |
14/1 |
Car |
|
Sabellidae |
3/0 |
7/0 |
Fil |
|
Paraonidae |
1/0 |
0/0 |
DS |
|
Opheliidae |
0/0 |
1/0 |
DC |
|
Polychaeta total abundance: |
538 |
416 |
|
|
Mollusca |
|
|
|
|
Anomalocardia flexuosa |
75/4 |
45/14 |
Fil |
|
Macomasp.1 |
21/2 |
23/1 |
Fil |
|
Phacoides pectinatus |
12/0 |
16/14 |
Sus |
|
Veneridae |
17/2 |
37/0 |
Fil |
|
Bittiolum varium |
0/3 |
2/1 |
Sus |
|
Tagelus divisus |
1/0 |
3/0 |
Sus |
|
Assiminea succinea |
9/1 |
17/0 |
Sce |
|
Caryocorbula caribaea |
0/0 |
2/1 |
Fil |
|
Acteocina bidentada |
7/0 |
14/0 |
Sus |
|
Sphenia fragilis |
2/0 |
2/1 |
Sus |
|
Neritina virginea |
0/0 |
0/2 |
Her |
|
Ervilia subcancellata |
0/0 |
2/0 |
Fil |
|
Paradentalium sp. |
0/0 |
1/0 |
Omn |
|
Tagelus sp. |
1/0 |
1/0 |
Sus |
|
Haminoea antillarum |
4/0 |
1/0 |
Sus |
|
Volvarina sp. |
1/0 |
0/0 |
Car |
|
Amygdalum papyrium |
0/0 |
5/0 |
Sus |
|
Chione sp. |
0/0 |
1/0 |
Fil |
|
Haminoea sp. |
0/0 |
1/0 |
Sus |
|
Macoma sp.2 |
0/0 |
3/1 |
Sus |
|
Bulla striata |
2/0 |
0/0 |
Her |
|
Mollusca total abundance: |
164 |
211 |
|
|
Macroinvertebrate total
abundance: |
702 |
627 |
|