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Population ecology and secondary production of congeneric bivalves on a sheltered beach in Southeastern Brazil.

ABSTRACT Members of the family Tellinidae are considered major components of trophic chains in sheltered sandy beaches located in temperate and tropical waters, due to their high number of species and abundance, as well as their several ecological functions. The population dynamics and secondary production of two congeneric bivalves Tellina lineata and Tellina versicolor were assessed in a sheltered beach environment. Monthly samplings from December 2006 to February 2009 were carried out at Flexeiras beach in southeastern Brazil. Samplings were conducted according to a systematic design, where six transects (fixed 20 m apart) were established, each with 10 equally spaced sampling levels. From each level, a sampling unit (0.04 [m.sup.2]) was taken. The populations differed in several aspects as follows: (1) T. lineata showed a significant preference for occupying the intermediate levels of the beach, whereas T. versicolor showed higher densities at lower levels, (2) T. lineata had significantly higher density, grew significantly slower, the mortality rate was lower and the biomass (B), secondary production (P) and turnover rates (P/B) were higher than for T. versicolor. The differences in population parameters of the two species may be associated a difference in the species' capacities to use resources available in the marine environment.

KEY WORDS: molluscs, Tellina lineata, Tellina versicolor, depositivores, growth, mortality, Brazil


Molluscs are important ecological macrobenthic components of marine coastal ecosystems, such as estuaries and sandy beaches, and are exposed to different environmental factors such as temperature, salinity, and currents (Degraeret al. 2007), which in turn affect their distribution and influence their population dynamics, demography, and reproduction (Fuiman et al. 1999, Cardoso et al. 2007). In sheltered sandy beaches, molluscs, especially bivalves, stand out for their high abundance, biomass, and dominant taxa (Olafsson et al. 1993, Urban 1994, Vaughn & Spooner 2006). These organisms play an important role in benthic communities, occupying different niches according to their feeding habits (Jaramillo et al. 2007), affecting sediment stability (Hall 1994) and the nutrient flow from the sediment to the water column (Michaud et al. 2006). They are also a compound part of the diet of several aquatic organisms and can affect energy cycling in the food chain (Paine 1966).

Studies on population biology, biomass, and production of highly dominant species, especially bivalves, provide knowledge regarding their role in the communities, as well as allowing for comparisons between ecosystems (Hibbert 1976). On a latitudinal scale, differences in bivalve growth rate have often been associated to latitudinal temperature gradients (Vakily 1992, Heck et al. 2002, Fiori & Morsan 2004), although other factors such as food quality and quantity, tidal level, and type of sediment appear to play a greater role on a local scale (Beukema et al. 2002, Carmichael et al. 2004). Populations from low intertidal and subtidal areas show higher growth rates and reproductive output due to longer submersion periods, allowing for the ingestion of greater amounts of food (Roseberry et al. 1991, Cardoso et al. 2007).

Members of the Tellinidae family are considered major components of trophic chains in sheltered sandy beaches located in temperate and tropical waters, due to the high number of species, abundance, and their ecological functions (Trevallion 1971, Dekker & Beukema 1999, Lizarralde & Cazzaniga 2009). Most of the tellins are distributed in intertidal beach zones and shallow waters (Trevallion 1971, Denadai et al. 2001), but knowledge regarding Tellina species population parameters is scarce. Some studies are available on the distribution of Tellina petitiana (D'Orbigny, 1846) in relation to environmental factors (Lizarralde 2002), predation by migratory birds (Pagnoni 1997), reproductive cycle (Baron & Ciocco 2001), population biology, and mean biomass (Lizarralde & Cazzaniga 2009). There is also information on Tellina tenuis (Da Costa, 1778) (Dekker & Beukema 1999) and Tellina lutea (W. Wood, 1828) (Selin 2010) population dynamics.

In Brazilian coast, Tellina lineata (Turton, 1819) and Tellina versicolor (De Kay, 1843) are widely distributed, occurring from the states of Ceara (05[degrees] 11' S, 39[degrees]17' W) to Santa Catarina (25[degrees] 57' S, 48[degrees]19' W) (Rios 1994). Arruda et al. (2003) classified both species as depositivores, with distribution in the intertidal region of muddy and sandy-muddy sheltered beaches. Cardoso et al. (2012) reported a high abundance of these species in sheltered beaches at Sepetiba Bay, southeastern Brazil. Few studies, however, have focused on the Tellina species of the South American coast.

The aim of this study was to analyze and compare the population ecology of the two congeneric bivalves, Tellina lineata and Tellina versicolor, in a sheltered beach located in southeastern Brazil, assessing their spatial and temporal distribution, population parameters (growth rate, mortality rate, and life span), and secondary production.


Study Area and Sampling

Itacuruca is the largest island inside the Sepetiba Bay (22[degrees]54' S, 43[degrees]34' W), Brazil. It harbors several beaches that provide different environments (protected and exposed) for several marine species that use these areas during all or part of their life cycle (Araujo et al. 1997). Among these beaches, Flexeiras beach (22[degrees]56' S, 43[degrees]53' W), characterized as sheltered according to the classification system proposed by McLachlan (1980), is noteworthy for its high richness and macrofauna diversity, mainly molluscs (Cardoso et al. 2011). The Flexeiras beach has a mean salinity of 33 (Cardoso et al. 2011).

Sampling was carried out monthly within the intertidal zone from December 2006 to February 2009, during the low spring tide. Six transects were set up perpendicularly to the shoreline and spaced 20 m apart. At each transect, sampling units (SU) were taken every 3 m, with a 0.04-[m.sub.2] metal sampler at a depth of 25 cm, from the boulder wall to 9 m below the waterline during low tide. Each sample was sieved through a 0.5-mm mesh and the material was taken to the laboratory (Fig. 1).

Sediment samples for particle size analyses were collected with a 3.5-cm-diameter corer to a depth of 15 cm at the lower, middle, and upper strata of two transects (transects 2 and 5).

Laboratory Procedures

All individuals were preserved in 70% ethanol and shell lengths of Tellina lineata and Tellina versicolor were measured. The anteroposterior length of the valves in all individuals were measured with calipers (0.01 mm precision), with each species grouped into 1.0-mm-size classes. All soft parts were individually removed, dried at 70[degrees]C for 24 h, weighed on a 0.001-g precision balance to obtain dry weight values, subsequently ashed in a muffle furnace at 600[degrees]C for 4 h, and reweighed to obtain the mass of the ash weight. The ash-free dry mass (AFDM) of each individual from both species was obtained by subtracting the dry mass from the ash mass.

The sediment samples were dried for 24 h at 70[degrees]C in an oven and subsequently passed through a series of sieves (-2.5 to 4.0 phi) in order of size to determine the average grain size (Folk & Ward 1957). The mean particle size was then calculated for the three sampled strata. The beach face slope of each transect was measured by the height difference between the drift and water lines (Emery 1961).

Data Analyses

Monthly length-frequency distributions (LFD) for each species were used to estimate growth patterns, following the procedures suggested by Gomez and Defeo (1999) and Defeo et al. (2001): (1) normally distributed components of LFD were separated through the NORMSEP routine of the FISAT program (Gayanilo et al. 1996); (2) absolute ages were assigned to the respective cohorts (lengths) to build an age-length key; and (3) the resulting age-length key for each bivalve was used to fit the generalized von Bertalanffy growth function (VBGF: Gayanilo et al. 1996) by nonlinear least squares:


where [L.sub.t] is length at time t (mm); [L.sub.[infinity]] is the theoretical maximum length attained by the species; K is the curvature parameter; C accounts for the intensity of seasonal growth oscillations; [t.sub.0] is the theoretical age at zero length; and WP is the winter point, that is, the period of growth reduction, expressed as a decimal fraction of the year. To compare the fitted VBGF between the bivalves, an analysis of the residual sum of squares was performed (Chen et al. 1992). The growth index phi prime [PHI]' = 2[1og.sub.10] ([L.sub.[infinity]]) + [log.sub.10]K (Pauly & Munro 1984, Defeo et al. 1992) was used as a measure of overall growth performance.

The instantaneous mortality rate (Z) was calculated for each species using the length-converted catch curve method (Pauly et al. 1995) of the FISAT program (Gayanilo et al. 1996). The Z estimation was given by: ln (n) = g - Z x t, where N is the number of individuals, g is the intercept of the regression, Z is the unbiased mortality estimated, and t is the estimated age in each cohort (Pauly et al. 1995). The annual mortality rate (A) was obtained by the expression: A = 1 - [e.sup.-Z], according to Ricker (1975). Life span ([t.sub.max]) was estimated by the VBGF growth parameters and based on the maximum length to estimate the maximum age ([T.sub.max]) for both species (Cardoso & Veloso 1996).

The relationship between individual shell length and the AFDM of the soft tissues was estimated by a linear regression analysis, following the equation: In W = ln a + b ln L, where W is the mean AFDM (g); L is the individual length (mm); and a and b are the regression parameters. Production was estimated by the weight-specific growth rate method (Crisp 1984), given by equation P = [SIGMA][SIGMA] [f.sub.i][G.sub.i][w.sub.i][DELTA]t, where f is the mean number of individuals from length class i during the time interval [DELTA]t, [G.sub.i] is the specific growth rate in weight within the length class i, and [w.sub.i] is the mean weight of the length class. [G.sub.i] can be obtained by [G.sub.i] = b x K[([L.sub.[infinity]]/[L.sub.i]) - 1], where b is the exponent of the length-weight relationship, K and [L.sub.[infinity]] are VBGF parameters, and [L.sub.i] is the mean length in the length class i. Annual mean biomass was calculated as B = [SIGMA][SIGMA][f.sub.i][w.sub.i][DELTA]t.

Statistical Analyses

A two-way analysis of variance (ANOVA) was applied to test the null hypothesis that Tellina lineata and Tellina versicolor abundance does not differ among months. Cochran's test (Underwood 1997) was used to test for variance homogeneity and then data were log-transformed to remove variance heterogeneity. Tukey's honest significant difference test was used a posteriori to assess significant differences. Nonlinear models were used to assess the relationship between bivalve density and mean grain size. Student's t-test was used to compare the numerical proportions of T. lineata and T. versicolor in each month. The test of slopes was used to compare mortality rates between the species (Zar 1999). In all statistical analyses, a significance level of 5% was adopted (Zar 1999).


Habitat Characterization

The variation in mean grain size ranged from 0.25 mm (media sand) to 1.30 mm (very coarse sand), with a mean value of 0.59 [+ or -] 0.17 mm. The one-way ANOVA indicated significant differences between mean grain size among strata (F = 4.57, df = 2/159, P < 0.05), Tukey's test detected differences between the upper strata in comparison with the other two strata (P < 0.05). Infra and middle strata were composed of coarse sand, whereas medium sand prevailed at the supralittoral levels. The beach presented a gentle slope, ranging from 1/20 to 1/40 m.

Population Structure

Density was significantly higher for Tellina lineata than Tellina versicolor in all months (t = 3.18; P < 0.05). The highest T. lineata densities were recorded in March and May 2008 (autumn) and February 2009 (summer), whereas the highest T. versicolor population peaks were observed in August 2007 and June and July 2008 (winter). The high standard errors in the monthly density of the species suggest a clumped distribution of individuals (Fig. 2).

The between-month abundance variation of the Tellina lineata population density (5.2-123.75 inds/[m.sup.2]) was higher than Tellina versicolor (1.67-36.67 inds/[m.sup.2]). The relationship between tellinid species density and mean grain size exhibited a different pattern, T. lineata density was not significantly correlated with grain size (n = 27, [r.sup.2] = 0.06, P > 0.05), whereas T. versicolor density significantly increased toward coarse grains (n = 27, [r.sup.2] = 0.13, P = 0.059).

The species were distributed in all sampling levels. Tellina Lineata, however, showed a significant preference for occupying the intermediate levels of the beach (SU 5-8) (F = 17.62, df = 9/ 260, P < 0.01), whereas Tellina versicolor showed higher densities at lower levels (SU 1-3) (F = 20.44, df = 9/260, P < 0.01) (Fig. 3). There was a significant inverse correlation between the distance water line and the population densities for T. lineata and T. versicolor (r = -0.321, P < 0.05).

Both species showed a markedly different population structure (Fig. 4), as reflected by the one-way ANOVA (F = 1021.59; P < 0.01): Tellina lineata was dominated by small length classes ranging from 1 to 12 mm (64% of the total population), whereas the dominant size classes in Tellina versicolor ranged from 13 to 32 mm (ca. 73%).


The analyses of the calculated growth parameters revealed significant variations in individual growth between bivalves: Tellina versicolor grew faster than Tellina lineata, as corroborated by the growth index phi prime ([SIGMA]'). The VBGF model with seasonal oscillation through the performance of nonlinear fitting explained more than 99% of the variance for T. lineata and more than 98% for T. versicolor. Growth parameter estimates were statistically significant (P < 0.05; Table 1).

Moderate intra-annual oscillations in growth reflected a minimal growth period in December (summer) for Tellina lineata (C = 0.41, WP = 1.00) and in September (spring) for Tellina versicolor (C = 0.23, WP = 0.70) (Fig. 5). The life span ([t.sub.max]) corresponding to these estimates was more than 3.15 y (31.28 mm) and 2.86 y (32.37 mm) for T. lineata and T. versicolor, respectively. Both sizes represent the largest individuals sampled from each population.


The comparison of the test of slopes indicated that the mortality rate for Tellina lineata (1.96 [y.sup.-1]) was significantly lower than for Tellina versicolor (6.37 [y.sup.-1]) (t-test = 8.13, P < 0.001). For those range sizes where mortality was estimated, the annual mortality rate (A) was 86% for T. lineata and 99.8% for T. versicolor (Fig. 6, Table 2).

Secondary Production

The regression equation between AFDM and length-weight relationship for the bivalve populations were expressed as: [W.sub.T, lineata] = (6 x [10.sup.-6]) [L.sup.2,855] (n = 545, r = 0.97, P < 0.05) and [W.sub.T, versicolor] = (4 x [10.sup.-6]) [L.sup.2,858] (n = 332, r = 0.96, P < 0.05).

The biomass (5), secondary production (P) and turnover rates (P/B) calculated for Tellina lineata (B = 1.391 AFDW g x [m.sup.-2], P = 2.293 AFDW g x [m.sup.-2] x [y.sup.-1], P/B = 1.65 [y.sup.-1]) were higher than for Tellina versicolor (B = 0.577 AFDW g x [m.sup.-2], P = 0.786 AFDW g x [m.sup.-2] x [y.sup.-1], P/B = 1.36 [y.sup.-1]).


The population parameters differed between tellinid species: Tellina lineata showed higher abundance, life expectancy, biomass, secondary production, and turnover rate values, whereas Tellina versicolor showed higher growth and mortality rate. These differences may be caused due to the interaction between both species and their similar lifestyles and food habits.

The distribution of the species is maintained by physical factors associated with biological interactions (McLachlan & Jaramillo 1995). It is unlikely that a single factor will explain patterns of distribution, because a species can occupy distinct habitats. The determination of macrofauna distribution in sandy beaches is difficult, because of the variability of environmental conditions and because most organisms live within the sediment (Amaral et al. 1990). The differences between the density peaks observed for Tellina lineata (autumn and summer) and Tellina versicolor (winter) and the spatial segregation with preference for differentiated levels suggest strategies adopted to avoid possible competition between species, observed in studies on the distribution of macrofauna species in the intertidal zone (Cardoso & Veloso 2003, Mattos & Cardoso 2012). The distribution in these lower beach zones are also related to a higher survival rate due to foraging protection from predators, such as birds and crabs (Zwarts & Wanink 1993). One of the most important factors defining habitat for psammophiles is sediment size. Grain size is considered the most critical factor and exerts direct influence on molluscs burrowing and mobility (Alexander et al. 1993, Ansell & Trevallion 1969). Intertidal or subtidal benthic macrofauna may select for optimal condition. Thus, the correlation positive between T. versicolor and grain size suggests that this species has optimum performance in coarser sands.

The LFD of Tellina versicolor population showed a high percentage of

adults, suggesting a senescent population with a marked absence of recruits (Fig. 4). This population structure could be explained by (1) recruitment failure due to irregular larvae contribution or an adverse effect of hydrodynamic factors (e.g., currents) affecting settlement and (2) low survival of recruits caused by unfavorable habitat conditions. These negative effects of the environment affecting recruitment success have been also documented in other intertidal bivalves inhabiting sandy beaches, tidal flats, and rocky substrates (e.g., Defeo 1996, Caddy & Defeo 2003). These hypotheses should be tested in the future.

The higher growth rate of Tellina lineata in relation to Tellina versicolor can be related to performance of the growth index phi prime, a parameter used to compare the growth intensity between species (Pauly & Munro 1984). Values of WP (point of lowest growth rate during the year) exhibited a different pattern for the two bivalves, T. lineata had its minimal growth in the summer months and T. versicolor during spring. Both periods can be related to higher temperatures and bivalve reproduction (Beukema & Dekker 1999). In subtropical regions, there is an increase in bivalve energy reserves during this period, as well as a reduction in growth and greater energy investment in reproduction (Moura et al. 2008). This pattern is also observed in other species from different taxonomic groups comprising sandy beach macrofauna (Caetano et al. 2006, Petracco et al. 2010). Besides temperature, food availability is also considered as an important parameter controlling growth rate of some Tellina species, because this regulates the beginning/end of the growth period and determines the occurrence of the number of growth periods that will occur during the year (Beukema et al. 1985, Beukema & Desprez 1986). The higher life expectancy recorded for T. lineata in comparison with T. versicolor is related to T. lineata's lower lvalues and lower growth performance rate (phi prime). Organisms with lower growth rates have higher life expectancies (Pauly 1979).

A consistent latitudinal pattern in growth performance was observed for Tellina species, showing higher [SIGMA]' values in low latitudes when compared with higher latitude species (Table 3). Vakily (1992) noted that bivalves grow faster at lower latitudes due to higher temperatures that increase metabolic rates. This pattern, however, may also be related to differences in food availability and plasticity in life habits that allow the species to survive under several temperatures and environmental conditions (Brown 1996, Navarro et al. 2000, Paterson et al. 2003, Cardoso et al. 2007).

The lower mortality rate of Tellina lineata (1.96 [y.sup.-1]), more than three times lower when compared with Tellina versicolor (6.37 [y.sup.-1]), could be related to greater burial depths of this species (Holme 1961). Thus, T. lineata suffers less from predation impacts when compared with T. versicolor, which mainly inhabits shallow waters (Trevallion 1971, Denadai et al. 2001). Other hypothesis would be irregular or spasmodic recruitment pulses that could cause a discontinuity in the arrival of recruits, which could explain the LFD of T. versicolor with the reduction of some year classes, compared with T. lineata (Caddy & Defeo 2003).

The biomass and secondary production values of both species were higher than those found by Warwick et al. (1978) for Tellina fabula (Gmelin, 1791) (B = 0.325 g AFDW [m.sup.-2] and P = 0.292 g AFDW [m.sup.-2] x [y.sup.-1]) in Wales. They suggest that the lower biomass and secondary production values are related to the greater abundance of intermediate-sized individuals in this population, because, in natural populations biomass and secondary production depend on other factors, such as the age structure of the population (Sprung 1993).

The relationship between annual production and mean biomass (P/B) is an adequate method to compare the productivity of different species, populations, and communities (Ansell et al. 1978). Both tellinid species showed higher P/B ratios (Tellina lineata = 1.65/year; Tellina versicolor = 1.36/ year) (Southeastern Brazil, 22[degrees] S) in comparison with those found for other Tellina species, such as T. petitiana (0.97/year) (Argentina, 42[degrees]S) (Lizaralde & Cazzaniga 2009) and T. fabula (0.90/year) (Wales, 51[degrees]N) (Warwick et al. 1978). The higher P/B ratios can be explained by the pattern described by Cardoso and Veloso (2003) for the bivalve Donax hanleyanus, in which higher P/B values are found at lower latitudes. These higher P/B values may be related to greater food availability and individual biomass contributions for energy and nutrient cycling (Souza 1998, Cardoso & Veloso 2003).

The highest abundance of Tellina lineata compared with Tellina versicolor and a spatial distribution with a preference for different levels may be associated with a difference in the species' capacities to use resources available in the marine environment. This can be confirmed by the negative correlation between densities of both species over time. Further experiments are necessary to evaluate interactions between tellinid species, estimating the degree of interactions and effects on population parameters.


We express our deepest gratitude to all fieldwork participants (Bruna Zavarize, Gustavo Mattos, Fabio Sendim, and Carlos Henrique Caetano). We also thank Omar Defeo and anonymous reviewer for their helpful comments that improved the manuscript. Ricardo S. Cardoso was supported by FAPERJ (Fundacao de Amparo a Pesquisa do Estado do Rio de Janeiro) and CT-Infra (Fundo Setorial de Infra-Estrutura) by MCPCNPq (no. E-26/171.164/2006). Tatiana M. B. Cabrini was supported by Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq) and by Universidade Federal do Rio de Janeiro (UFRJ) post-graduate research studentships.


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DOI: 10.2983/035.034.0323


(1) Laboratorio de Ecologia Marinha, Departamento Ecologia e Recursos Marinhos, Universidade Federal do Estado do Rio de Janeiro (UNIRIO), Avenida Pasteur, 458, Sala 407, URCA, Rio de Janeiro, CEP 22290-240, Brazil; (2) Programa de Pos-Graduacao em Ecologia, Universidade Federal do Rio de Janeiro (UFRJ), Rio de Janeiro, CEP 21941-902, Brazil

* Corresponding author. E-mail: rcardoso@!

Growth parameters estimated by nonlinear fitting of the
von Bertalanffv growth function for Tellina lineata and
Tellina versicolor.

                              Tellina lineata

Parameters                  Mean ([+ or -] SE)       P

[L.sub.[infinity]] (mm)    32.33 ([+ or -] 1.62)   0.0000
K[ year                     0.54 ([+ or -] 0.06)   0.0000
C                           0.41 ([+ or -] 0.18)   0.0028
WP                          1.00 ([+ or -] 0.06)   0.0000
[t.sub.o]/year             -0.31 ([+ or -] 0.06)   0.0001
[PHI]'                             2.75
Largest (mm)                      31.28
[t.sub.max]/year                   3.15

                            Tellina versicolor

Parameters                  Mean ([+ or -] SE)       P

[L.sub.[infinity]] (mm)   35.49 ([+ or -] 1.15)   0.0000
K[ year                    0.83 ([+ or -] 0.07)   0.0000
C                          0.23 ([+ or -] 0.09)   0.0034
WP                         0.70 ([+ or -] 0.07)   0.0001
[t.sub.o]/year            -0.03 ([+ or -] 0.02)   0.0050
[PHI]'                             3.02
Largest (mm)                      32.37
[t.sub.max]/year                   2.86

Mortality estimates (Z) for Tellina lineata (n = 20)
and Tellina versicolor (n = 9).

               Tellina lineata             Tellina versicolor

Parameters    Mean ([+ or -] SE)     P     Mean ([+ or -] SE)      P

g            8.86 ([+ or -] 0.15)  0.000  17.18 ([+ or -] 0.20)  0.000
Zj year      1.96 ([+ or -] 0.26)  0.000   6.37 ([+ or -] 0.18)  0.000
[R.sup.2]            0.99          0.000          0.92           0.000
n                   20                            9


Latitudinal variation in growth performance in species
of the genus Tellina.

Species              K/year          (mm)          [PHI]'

Tellina lineata       0.54          32.33           2.75
Tellina versicolor    0.83          35.49           3.02
Tellina petitiana     0.37          50.51           2.97
Tellina foliacea      1.20           7.70           1.85
Tellina tenuis        0.66          23.24           2.66
Tellina tenuis        0.18          21.10           1.90
Tellina tenuis        0.43          15.50           2.01
Tellina tenuis        0.43          16.80           2.08
Tellina tenuis        0.31          19.70           2.08
Tellina tenuis        0.38          22.40           2.28
Tellina lutea         0.10           76.3           2.77

Species                  Latitude                   Source

Tellina lineata      22[degrees]57' S   Present study
Tellina versicolor   22[degrees]57' S   Present study
Tellina petitiana    42[degrees]46' S   Lizaralde and Cazzaniga (2009)
Tellina foliacea     49[degrees]15' S   Negar et al. (2008)
Tellina tenuis       52[degrees]56' N   Dekker and Beukema (1999)
Tellina tenuis       58[degrees]15' N   Evans and Tallmark (1977)
Tellina tenuis       55[degrees]45' N   Stephen (1928)
Tellina tenuis       55[degrees]46' N   Barnett and Watson (1986)
Tellina tenuis       53[degrees]18' N   Wilson (1997)
Tellina tenuis       46[degrees]09' N   Faure (1969)
Tellina lutea        53[degrees]00' N   Selin (2010)

K, curvature parameter; [L.sub.[infinity]] asymptotic
length; [PHI]', growth performance index.
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Author:Cardoso, Ricardo S.; Galhardo, Ludmila B.; Cabrini, Tatiana M.B.
Publication:Journal of Shellfish Research
Article Type:Report
Geographic Code:3BRAZ
Date:Dec 1, 2015
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