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Composition of decapod larvae in a northeastern Brazilian estuarine inlet over a full tidal cycle.

Composicion de larvas de decapodos en la desembocadura de un estuario del noreste brasileno durante un ciclo completo de marea

INTRODUCTION

Many representatives of the estuarine meroplankton use the freshwater and/or marine flows as a means of transport to other locations (mainly to the adjacent continental shelf), where post-embryonic development often occurs. However, a few species have different migratory behaviors and exhibit retention strategies due to circulation and tidal regimes in the estuaries (Christy & Stancyk, 1982; Epifanio, 1988). As a consequence, the distribution of meroplankton in coastal waters is influenced by both behavioral and physical transport mechanisms (Christy & Stancyk, 1982; Butman, 1987; Melo Junior et al., 2012).

In several studies, the effects of tides and photoperiodic variations have focused on the dynamics of estuarine larval decapod, both in temperate areas (Morgado et al., 2006) as well as in tropical (e.g., Silva-Falcao et al., 2007) and subtropical regions (Fernandes et al., 2002; Koettker & Freire, 2006). Usually the synchronization of the larval release with photoperiod seems to be associated with susceptibility to predation (Morgan & Christy, 1995). Most of the time, inconspicuous larvae are released into the estuary regardless of photoperiod, while larvae more attractive to predators are released during the night tide, especially during the spring ebb tides (Morgan & Christy, 1995; Gove & Paula, 2000).

There is extensive information on estuarine and coastal planktonic decapod in neotropical Brazil (Schwamborn & Bonecker, 1996; Schwamborn, 1997; Porto-Neto et al., 1999; Silva-Falcao et al., 2007; Magris & Loureiro-Fernandes, 2011; Melo Junior et al., 2012; Oliveira et al., 2012; Koettker & Lopes, 2013). However, the vertical distribution in relation to tidal cycles is known only for a few taxa. To our knowledge, this is probably the first published study in Brazil that explores the variability of composition of planktonic decapods in an estuarine system, considering its distribution in different layers of the water column. The present study examines the changes of planktonic decapod composition and occurrence of developmental stages over a full tidal and diel cycles in a northeastern Brazilian estuarine inlet.

Study area

The Itamaraca Estuarine System (IES) is located at 7[degrees]34'00"-7[degrees]55'16"S and 34[degrees]48'48"-34[degrees]52'24"W, about 50 km north from Recife City, in the state of Pernambuco, northeast Brazil. It consists of a U-shaped channel (Santa Cruz Channel, 20 km in length) with two connections to the South Atlantic Ocean (Catuama and Orange inlets) and five main tributaries draining into the channel. Mangrove forests (~28 [km.sup.2]) occupy the lowlands along the inner portion of the Santa Cruz Channel and the lower part of its tributaries. The dominant species is the red mangrove (Rhizophora mangle), which its litter is responsible by 7 ton of dry weight of organic matter [ha.sup.-1] [year.sup.-1] (Medeiros et al., 2001). This research was developed in Catuama Inlet (Fig. 1). Sampling was conducted on board of a small vessel during neap tide (August 11-12, 2001) in 3-h intervals over 24 h.

Sampling and laboratory methods

Plankton samples were collected with a pump at three stations along a transect that crosses the inlet. At the central station (Center), three depth levels were sampled (50 cm below the surface, at mid-water, and 50 cm above the bottom, 12-15 m), while at the lateral stations (Continent and Island stations) samples were taken only at the subsurface and above the bottom (8-10 m). Water samples were pumped on board through a conical 300 [micro]m plankton net for 3 to 5 min per sample at approximately 100 L per minute. Immediately after sampling, all samples were fixed in 4% buffered seawater formaline (n = 56).

In the laboratory, planktonic decapods were counted and identified in toto, considering the lowest feasible taxonomic unit by optical analysis. The identification of organisms was made following Gurney (1942), Kurata (1970), Boschi (1981), Calazans (1993), Schwamborn (1997), Pohle et al. (1999), among others. The larval stages for each taxa were determined by characteristics specified mainly by Gurney (1942), Boschi (1981), Calazans (1993) and Pohle et al. (1999). The quantitative data (ind [m.sup.-3]) is not presented here, but is documented elsewhere (Melo Junior, 2005).

For the frequency of occurrence, we used the formula [F.sub.[omicron]] (%) = [Ta.sup.*] 100 * [TA.sup.-1] where Ta: number of samples where the taxon occurs; and TA: total samples. For the interpretation of the result of the frequency of occurrence we used the following scale: >70% very frequent; 70-40% frequent; 40%-10%: infrequent and [less than or equal to]10% sporadic. For the relative abundance, we calculated the percentage of individuals in diurnal and nocturnal photoperiod and at ebb and flood tides. To show the abundance distribution of the individuals in the three depth levels, we used the formula RA (%) = [N.sup.*]100 * [TN.sup.-1] where: N: number of individuals of the taxon in a determinate depth level; and TN: total number of individuals considering the three depth levels.

To detect non-random distributions of decapod larvae between the tidal conditions, photoperiod and depth layers (vertical distribution), the Indicator Species Analysis (Dufrene & Legendre, 1997) was used. This method is based on an indicator value index (IndVal), which combines both the larval relative abundance with its relative frequency of occurrence in defined conditions. The values ranged from 0% (no indication) to 100% (perfect indication). The statistical significance of the IndVal was evaluated using a Monte Carlo test (permutation number = 1,000). The level of significance was set at P < 0.05 for this analysis, and only indicator value indices of more than 70% were considered. Additionally, a clustering analysis was elaborated to establish the main decapod larvae assemblages, based on the Bray-Curtis dissimilarity coefficient and using the density matrix transformed by the UPGMA method. Cophenetic analysis was conducted to measure the fitness of the data. The analyses were run on software R, using the vegan package (Oksanen et al., 2010).

Planktonic decapods in the Catuama Inlet were represented by 27 taxa (Table 1), being distributed in 13 families: Sergestoida (2), Caridea (2), Anomura (2), Thalassinidea (2) and Brachyura (5), besides of non-identified taxa in Hippidea and Stenopodidea. This ecosystem showed a variety of planktonic decapod compared with other coastal and estuarine systems of Brazil (Sankarankutty et al., 1995; Schwamborn et al., 2001; Fernandes et al., 2002; Negreiros-Fransozo et al., 2002; Silva-Falcao et al., 2007). The high number of detected taxa is even more intriguing, since the period examined in this study was inferior than most of surveys conducted at the Brazilian coast. This richness could be even greater if we consider that the pump suction is not one of the most widely used sampling equipments for qualitative analysis of zooplankton, even this technique being adequate for quantitative data at exactly depths (Melo Junior et al., 2015). For example, a study using simultaneous plankton net (300 pm) at the same area (Schwamborn, R. unpublished data), recorded two genera of the complex Penaeus (Litopenaeus sp. and Farfantepenaeus sp.) (Penae-oidea) and the families Galatheidae (Anomura) and Portunidae (Brachyura), which were not collected by pump suction in our sampling.

The number of planktonic decapod species recorded here was below (<10%) the number of species (around 117-all potentially producers of planktonic larvae) inhabiting the IES (Coelho, 2000). This pattern seems to be common in many estuaries and has been observed in many studies of decapod larvae in other Brazilian (Sankarankutty et al., 1995; Schwamborn et al., 2001; Fernandes et al., 2002; Negreiros-Fransozo et al., 2002; Silva-Falcao et al., 2007) and other world estuaries (Dittel & Epifanio, 1990; Dittel et al., 1991; Criales & McGowan, 1994; Paula et al., 2004). Most of the authors attribute this pattern to a lack of knowledge about larvae taxonomy, and is associated with the dominance of a few species in the ecosystems studied or the different reproductive periods of the occurring species.

Among the recorded taxa, 27.8% were very frequent (Table 1), especially the protozoea of Lucifer faxoni and Acetes americanus with 91.7% each, followed by zoea of Pinnixa sp. 1 (78.6%), Uca spp. (76.8%), and Panopeus spp. and the Ocypodidae morphotype A (both with 75%). The zoea of Caridea (others), mysis of Lucifer faxoni, zoea of Petrolisthes armatus, and Alphaeidae were also very frequent. Other decapods were frequent (19.4% of the 27 registered taxa), especially the zoea of Upogebia spp. (64.6%) and Paguridae (62.5%). The 19 remaining decapods were infrequent (25% of them) or sporadic (27.8% of them).

Concerning the quantity of developmental stages, Sergestoida and Brachyura were the two more representative groups of decapod (Fig. 2). However, no evident tidal variation was observed to the distribution of the larval stages in relation to the main groups of decapod larvae. Considering all decapods, we found higher numbers of developmental stages (>40) during flood and high tides, with a maximum of 60 stages at the nocturnal flood tide. Lower numbers were observed during the diurnal low tide (Fig. 2). A greater range among Sergestoida was observed, and a unique adult stage among all planktonic decapod was recorded (Lucifer faxoni). There were occurrences of larvae in early stages of development in all families, and more advanced stages, such as post-larvae, zoea IV or more, megalopa, and glaucothoe were recorded in almost every groups, except in Stenopodidea and Hippidea (Tables 2-3).

We found two main groups of planktonic decapods (Fig. 3): i) an assemblage of mainly infrequent and sporadic larval stages, with variables peaks. Most of these larvae are represented by advanced stages of all decapod taxa; ii) Assemblage of very frequent larval stages, represented by initial and intermediate stages of all registered groups. Twenty larval stages were found to be significant indicators (IndVal > 70%; P < 0.05) of some tidal (mainly, flood) and photoperiod (mainly, night) conditions (Fig. 3). The larvae of Zaops ostreum, Pinnotheridae and Lucifer faxoni were good indicators of flood tides, while the larval stages of Caridea (Alphaeidae and others), Paguridae, Pinnotheridae and Luciferidae, of nocturnal period. These taxa were also associated with other decapod larvae, as is shown in the cluster analysis (Fig. 3).

No difference on planktonic decapod composition or developmental stages was observed in relation to the stations. On the other hand, the vertical distribution of the percentage of the larval stages was variable between the decapod taxa, with evident concentration of individuals in determined layers (Tables 2-3). Even so, no larval stage was indicator of a depth layer (surface, mid-water or bottom), suggesting that the low depth of the inlet (maximum of 15 m) is probably not sufficient to promote patterns of vertical aggregations of the larvae.

Most of the planktonic decapods from Catuama Inlet presented many larval stages (some occurring with four or more developmental stages), suggesting that part of the most larvae develop next to the inlet (Epifanio, 1988; Paula et al., 2004). This pattern is common in euhaline estuarine systems where the development of these species occurs, since the zoea I that remains in the estuary is associated with tidal flooding (Freire, 1998). Since salinity in IES vary from euhaline to mesohaline (Macedo et al., 2000), it is likely that most of the taxa use the region near Catuama inlet as a growth area. This hypothesis was in part confirmed in a study on the transport of decapod larvae by the IES plumes (Schwamborn et al., 2001). In this study, authors observed that larvae dispersion reached only a few kilometers from the coast, suggesting that larvae are retained in estuarine plumes due to the convergence areas formed when the IES's water bodies meet marine water masses.

Considering the classification of Anger (2001), most taxa are larvae of exporting species, while only a small group (formed by Lucifer faxoni, Acetes americanus, and Petrolisthes armatus) seems to have mechanisms for retaining at least one of the larval stages in the IES (Table 2). These three taxa, mainly the initial stages, were grouped in the cluster analysis (Fig. 3). Nevertheless, the high number of larval stages from different taxa indicates that probable mechanisms of retention in the areas adjacent to the inlet should not be discarded. These findings are reinforced by the fact that seven developmental stages were perfect indicators (IndVal > 70%) of flood tide, while just one (Pinnotheridae, zoea II) was indicator of ebb tide.

The role of mangroves in IES as a source of decapod larvae is reinforced by this study. Thus, it is observed that Catuama inlet represents a real corridor for energy exchange in the form of planktonic decapods between the IES and its adjacent coastal shelf. Areas nearby estuaries and mangroves have large densities of decapod larvae at certain times (Neumann-Leitao et al., 1999; Schwamborn et al., 1999, 2001) and one of the main trophic functions of these larvae is to transfer energy to higher links of the food chain (Robertson & Blaber, 1992), including economically important fish. Changes in the tidal cycles and photoperiod, with high numbers of developmental stages during flood and high tides and during the night, promoted a set of variations in the planktonic decapod composition. Besides, the high variability of developmental stages in most of the taxa identified suggests that the development of thesespecies really occurs in the region near to the Catuama inlet.

DOI:10.3856/vol44-issue2-fulltext-21

Received: 20 July 2015; Accepted: 14 January 2016.

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Mauro de Melo Junior (1), Pedro Augusto Mendes de Castro Melo (2) Maryse Nogueira Paranagua (1), Sigrid Neumann-Leitao (3) & Ralf Schwamborn (3)

(1) Laboratorio de Ecologia do Plancton, Departamento de Biologia, Universidade Federal Rural de Pemambuco-DB/UFRPE, 52171-900, Recife, Pernambuco, Brazil

(2) Laboratorio de Ecologia do Plancton, Universidade Estadual de Santa Cruz-UESC Campus Prof. Soane Nazare de Andrade, 45662-900, Ilheus, Bahia, Brazil

(3) Laboratorio de Zooplancton Marinho, Museu de Oceanografia, Centro de Tecnologia e Geociencias Universidade Federal de Pernambuco-MO/CTG/UFPE50670-901, Recife, Pernambuco, Brazil

Corresponding author: Mauro de Melo Junior (mmelojunior@gmail.com)

Paper presented in the 5th Brazilian Congress of Marine Biology, 17-21 May 2015, Porto de Galinhas, Brazil.

Caption: Figure 1. Itamaraca estuarine system (Pernambuco, Brazil) showing the Catuama Inlet and the sampling stations.

Caption: Figure 2. Number of developmental stages of planktonic decapods registered during a 24-h study, in Catuama Inlet (northeastern of Brazil), in August 2011. D: diurnal, N: nocturnal, HT: high tide, ET: ebb tide, LT: low tide, FT: flood tide.

Caption: Figure 3. Cluster analysis based in Bray-Curtis dissimilarity coefficient, using the transformed abundance matrix by the UPGMA method, and significant Indicator Values (IndVal, %) of some taxa in relation to tidal and diel conditions. Car: Caridea (others), Pag: Paguridae, Aa: Acetes americanus, Zo: Zaops ostreum, Bra: Brachyura (others), Uca: Uca spp., OcypA: Ocypodidae A, Pin: Pinnotheridae, Alp: Alphaeidae, Up: Upogebia spp., Lf: Lucifer faxoni, Pi: Pinixa sp. 1, 2 or 3, Pa: Petrolisthes armatus, Pan: Panopeus spp., P: protozoea, M: mysis, J: juvenile, A: adult, Z: zoea. I, II, III, IV, V, VI: stages of development. IndVal legends: N: nocturnal period, F; flood tide, D: diurnal period, E: ebb tide.
Table 1. Composition, description of larval stages and frequency of
occurrence of planktonic decapods registered during a 24 h study, in
Catuama Inlet (northeastern Brazil), in August 2011. P: protozoea, M:
mysis, J: juvenile, A: adult, Z: zoea, PL: post-larvae, G:
glaucothoe, ME: megalopa. I, II, III, IV, V, VI: stages of
development. **** very frequent; *** frequent; ** infrequent;
* sporadic.

Decapod larvae             Larval stages        Frequency (%)

Sergestoida
Family Luciferidae Dana,   PI, PII, PIII, MI,   91.66 ****
  1852 Lucifer faxoni        MII, MIII, J, A
  Borradaile, 1915
Family Sergestidae Dana,
1852
Acetes americanus          PI, PII, PIII, MI,   91.66 ****
  Ortmann, 1893              MII, MIII
Stenopodidea               ZI                   02.08 *

Caridea
Family Hippolytidae        ZI, ZII, ZIII        02.08 *
  Bate, 1888
Family Alpheidae           ZI, ZII, ZIII, PL    68.75 ***
  Rafinesque, 1815
Caridea (others)           ZI, ZII, ZIII, PL    70.83 ****

Anomura
Family Porcellanidae
  Haworth, 1825
Petrolisthes armatus       ZI, ZII              68.75 ***
  Gibbes, 1850
Porcellanidae (others)     ZI, ZII              10.41 **
Family Paguridae           ZI, ZII, ZIII, G     62.50 ***
  Latreille, 1803

Thalassinidea
Family Callianassidae      ZI, ZII              04.16 *
  Dana, 1852
Family Upogebiidae
  Borradaile, 1903
Upogebia spp.              ZI, ZII, ZIII        64.58 ***
Hippidea                   ZI                   02.08 *

Brachyura
Family Dromiidae De        ZI                   10.45 **
  Haan, 1833
Family Leucosiidae         ZI, ZII, ZIII        06.25 *
  Samouelle, 1819
Family Xanthidae
  MacLeay, 1838
Menippe nodifrons          ZI, ZII              06.25 *
  Stimpson, 1859
Panopeus spp.              ZI, ZII, ZIII,       75 00 ****
                             ZIV, ZV
Xanthidae Morphotype A     ZI, ZII, ZIII, ZIV   29.16 **
Xanthidae Morphotype B     ZI                   04.16 *
Family Pinnotheridae De    ZI, ZII, ZIII,       78.57 ****
  Haan, 1833 Pinnixa         ZIV, ZV, ZVI
  sp. 1
Pinnixa sp. 2              ZI, ZII, ZIII, ZIV   27.08 **
Pinnixa sp. 3              ZI, ZII, ZIII,       58.33 ***
                             ZIV, ZV, ZVI
Zaops ostreum              ZI, ZII, ZIII,       39.58 **
  (Say, 1817)                ZIV
Pinnotheridae              ZI, ZII, ZIII,       47.91 ***
  Morphotype A               ZIV, ZV

Family Ocypodidae
  Rafinesque, 1815
Uca maracoani              ZI                   06.25 *
  Latreille, 1802
Uca spp.                   ZI, ZII, ZIII, ZIV   76 79 ****
Ocypodidae Morphotype A    ZI, ZII, ZIII, ZIV   75 00 ****
Ocypodidae Morphotype B    ZI, ZII, ZIII,       16.66 **
                             ZIV, ZV
Brachyura (others)         ZI, ZII, ZIII,       56.25 ***
                             ZIV, ZV, ME

Table 2. Occurrence and relative abundance (%) of larval stages
of planktonic decapods (except Brachyura) registered during a
24 h study, in Catuama Inlet (northeastern Brazil), in August
2011, considering the tides, photoperiod and the depth. S: 50
cm below the surface, m: mid-water, b: 50 cm above the bottom
(8-15 m); -no occurrence; PL: post-larvae; G:
glaucothoe.

Decapod larvae    Diurnal ebb     Nocturnal flood

                 s     m     b     s     m    b
Sergestoida
Lucifer faxoni
Protozoea        53   23    24    61     8    31
Mysis            8    84     8    10    22    68
Juvenile         --   --    --    32    33    35
Adult            --   --    --    11    53    37

Acetes
  americanus
Protozoea        30   53    16    27    38    36
Mysis            --   30    70    --    92    8
Caridea
Hippolytidae
Zoea I           --   --    --    --    --    --
Alpheidae
Zoea I + II      42   50     8    38    57    5
Zoea III + PL    --   --    100   11    79    11

Caridea
  (others)
Zoea I + II      8    18    74    31    23    46
Zoea III + PL    --   --    100   10    69    21

Anomura
Petrolisthes
  armatus
Zoea I           25   40    35    --    97    3
Zoea II + III    5    52    42    --    93    7
Porcellanidae
  (others)
Zoea I           --   --    --     8    83    10
Paguridae
Zoea I           1    90     9    18    44    38
Zoea II +        --   96     4    15    26    59
  III + G

Anomura
  (others)
Glaucothoe       --   --    --    --    100   --
Thalassinidea
Callianassidae
Zoea I           --   --    --    50    50    --
Upogebia spp.
Zoea I           14   81     5    45    32    23
Zoea II + III    --   100   --    26    30    44
Hippidea
Zoea I           --   --    --    100   --    --

Decapod larvae   Nocturnal ebb   Diurnal flood

                 s    m     b     s     m     b
Sergestoida
Lucifer faxoni
Protozoea        53   44    3    17    58    26
Mysis            65   30    6     2    28    71
Juvenile         22   66   13    --    43    57
Adult            6    67   27    21    17    62

Acetes
  americanus
Protozoea        78   14    8     4    78    17
Mysis            59   --   41    --    --    --
Caridea
Hippolytidae
Zoea I           --   --   --    --    72    28
Alpheidae
Zoea I + II      76   19    5    13    26    61
Zoea III + PL    15   80    5    --    47    53

Caridea
  (others)
Zoea I + II      64   28    8    19    47    34
Zoea III + PL    11   28   61    --    --    100

Anomura
Petrolisthes
  armatus
Zoea I           5    26   69    43    48     9
Zoea II + III    3    17   80    74    26    --
Porcellanidae
  (others)
Zoea I           --   --   --    100   --    --
Paguridae
Zoea I           41   43   16     6    38    56
Zoea II +        73   18    9     -    20    80
  III + G

Anomura
  (others)
Glaucothoe       --   --   --    --    100   --
Thalassinidea
Callianassidae
Zoea I           --   --   100   --    --    --
Upogebia spp.
Zoea I           34   19   47     2    84    14
Zoea II + III    --   71   29    --    100   --
Hippidea
Zoea I           --   --   --    --    --    --

Table 3. Occurrence and relative abundance of larval stages
of planktonic Brachyura registered during a 24 h study, in
Catuama inlet (northeastern Brazil), in August 2011,
considering the tides, photoperiod and the depth. Depth: s:
50 cm below the surface, m: mid-water, b: 50 cm above the
bottom (8-15 m), -no occurrence, Morph: morphotype.

Decapod larvae           Diurnal ebb     Nocturnal flood

                         s     m    b     s     m     b
Brachyura
Dromiidae
Zoea I                  --    --    --   53    17    29
Leucosiidae
Zoea I                  --    100   --   15    85    --
Zoea II + III           100   --    --   --    --    --

Menippe nodifrons
Zoea I                  --    --    --   65    --    35
Zoea II                 --    --    --   100   --    --

Panopeus spp.
Zoea I + II              4    60    36    5    70    25
Zoea III + IV + V        3    16    81    5    --    95
Xanthidae Morph A
Zoea I + II             --    38    62   19    37    44
Zoea III + IV           --    --    --    3    61    36

Xanthidae Morph B
Zoea I                  48    --    52   --    --    --
Pinnixa sp. 1
Zoea I + II              9    78    12   19    25    56
Zoea III + IV           --    47    53    2    66    31
Zoea V + VI             --    --    --   --    88    12

Pinnixa sp. 2
Zoea I + II             10    78    13   69    --    31
Zoea III + IV           --    32    68   --    --    100

Pinnixa sp. 3
Zoea I + II              6    82    12    4    55    40
Zoea III + IV + V       --    --    --    1    35    64
Zaops ostreum
Zoea I + II              7    82    11    5    37    58
Zoea III + IV           --    --    --    3    19    77

Pinnotheridae Morph A
Zoea I + II             16    46    38    3    45    51
Zoea III + IV + V       --    27    73   --    100   --

Uca maracoani
Zoea I                  --    --    --   100   --    --
Uca spp.
Zoea I + II             23    60    17   20    38    42
Zoea III + IV            2    37    60    5    62    33

Ocypodidae Morph A
Zoea I + II             11    79    10   59    34     7
Zoea III + IV + V       --    33    67    2    93     6

Ocypodidae Morph B
Zoea I + II             26    59    15   20    18    62
Zoea III + IV           --    94    6    21    29    49

Brachyura (others)
Zoea I + II             18    68    14   91    --     9
Zoea III + IV + V       14    40    46   88    --    12
Megalopa                --    80    20   12    28    60

Decapod larvae          Nocturnal ebb    Diurnal flood

                        s     m     b     s     m     b
Brachyura
Dromiidae
Zoea I                  --   --    --    --    --    --
Leucosiidae
Zoea I                  --   --    --    --    --    100
Zoea II + III           --   --    --    --    100   --

Menippe nodifrons
Zoea I                  --   --    --    --    --    --
Zoea II                 --   --    --    --    --    --

Panopeus spp.
Zoea I + II             20   50    30    13    63    24
Zoea III + IV + V       --   56    44    50    23    27
Xanthidae Morph A
Zoea I + II             26   70     4    10    40    50
Zoea III + IV           50   42     8    100   --    --

Xanthidae Morph B
Zoea I                  --   --    --    --    --    --
Pinnixa sp. 1
Zoea I + II             54   38     8    19    30    51
Zoea III + IV           18   38    44     9    22    69
Zoea V + VI             --   --    --    --    --    100

Pinnixa sp. 2
Zoea I + II             18   21    60    10    84     5
Zoea III + IV           --   --    100   --    89    11

Pinnixa sp. 3
Zoea I + II             23   36    41    14    55    32
Zoea III + IV + V       10    9    81    25    15    59
Zaops ostreum
Zoea I + II             21   79    --    21    39    40
Zoea III + IV           --   --    --    60    30    10

Pinnotheridae Morph A
Zoea I + II             27   25    48    11    47    42
Zoea III + IV + V       9    13    78    100   --    --

Uca maracoani
Zoea I                  --   --    --    --    100   --
Uca spp.
Zoea I + II             54   24    23    23    55    22
Zoea III + IV           13   11    76     9    17    74

Ocypodidae Morph A
Zoea I + II             --   --    --    --    --    --
Zoea III + IV + V       --   --    --    --    --    100

Ocypodidae Morph B
Zoea I + II             73    2    25    19    28    53
Zoea III + IV           --   --    100   23    18    60

Brachyura (others)
Zoea I + II             50   --    50     9    17    74
Zoea III + IV + V       --   100    0    34     3    63
Megalopa                --   --    100   28    --    72
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Title Annotation:Short Communication
Author:de Melo Junior, Mauro; de Castro Melo, Pedro Augusto Mendes; Paranagua, Maryse Nogueira; Neumann-Lei
Publication:Latin American Journal of Aquatic Research
Article Type:Ensayo
Date:May 1, 2016
Words:5020
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