Hydrodynamically-driven distribution of lanternfish larvae in the Southeast Brazilian Bight

Accepted Manuscript Hydrodynamically-driven distribution of lanternfish larvae in the Southeast Brazilian Bight

Cláudia Namiki, Mario Katsuragawa, Dante Campagnoli Napolitano, Maria de Lourdes Zani-Teixeira, Rafael Augusto de Mattos, Ilson Carlos Almeida da Silveira PII: DOI: Reference:

S0924-7963(16)30233-0 doi: 10.1016/j.jmarsys.2017.02.006 MARSYS 2951

To appear in:

Journal of Marine Systems

Received date: Revised date: Accepted date:

1 August 2016 13 February 2017 17 February 2017

Please cite this article as: Cláudia Namiki, Mario Katsuragawa, Dante Campagnoli Napolitano, Maria de Lourdes Zani-Teixeira, Rafael Augusto de Mattos, Ilson Carlos Almeida da Silveira , Hydrodynamically-driven distribution of lanternfish larvae in the Southeast Brazilian Bight. The address for the corresponding author was captured as affiliation for all authors. Please check if appropriate. Marsys(2017), doi: 10.1016/ j.jmarsys.2017.02.006

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ACCEPTED MANUSCRIPT Hydrodynamically-driven distribution of lanternfish larvae in the Southeast Brazilian Bight Cláudia Namiki*1, Mario Katsuragawa1, Dante Campagnoli Napolitano1, Maria de Lourdes Zani-Teixeira1, Rafael Augusto de Mattos2, & Ilson Carlos Almeida da Silveira1 1

Oceanographic Institute, University of São Paulo;

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National Laboratory for Scientific

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Computing, LNCC/MCTI

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*Corresponding author: [email protected], telephone number: +55 11 30916589

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Abstract

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This study analyzes the influence of the Brazil Current and Ekman transport on the distribution of lanternfish larvae in the Southeast Brazilian Bight during summer and winter.

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Larvae of 19 taxa of lanternfish were identified, and Diaphus spp. and M. affine were the most abundant. Three water masses were present in the area: Coastal Water, Tropical Water and

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South Atlantic Central Water. Lanternfish larvae were associated with the Tropical Water in both seasons. During summer, species of Lampanyctinae were associated with the

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shallowest layers and Myctophinae in the deepest layers. In winter most species of both subfamilies were associated with intermediate depths, probably because greater mixing of

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water masses occurred at the surface and 100 m depth, limiting their distribution. During both cruises, the presence of lanternfish larvae in the continental shelf was related to the pattern of

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Tropical Water intrusion, which was mostly driven by the mesoscale activity of the Brazil

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Current and its interaction with the continental shelf.

Keywords: Myctophidae; ichthyoplankton; mesopelagic fish; mesoscale features; vertical distribution; current meandering.

ACCEPTED MANUSCRIPT 1. Introduction

Mesopelagic fishes are among the most abundant group on Earth and they may respire up to 10% of the primary production in deep sea waters (Irigoien et al., 2014). The most speciose mesopelagic family is the Myctophidae, whose members are generally called lanternfish because they have conspicuous luminous organs on their body (Moser and

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Watson, 2006). Lanternfish are composed of approximately 240 species in 32 genera

2014, 2008; Hulley, 1981; Santos and Figueiredo, 2008).

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(Nelson, 2006) and there are at least 81 species in the Southwest Atlantic (Braga et al.

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Lanternfish larvae are dominant in the oceanic larval fish assemblages (Katsuragawa et al., 2014; Muhling et al., 2008; Olivar and Shelton, 1993). The presence of

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mesopelagic larvae in the continental shelf can be associated with meanders and eddies of oceanic currents, because these mesoscale features may transport fish larvae from the

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ocean to the continental shelf and vice-versa (Flierl and Wroblewski, 1985; Franco et al., 2006; Myers and Drinkwater, 1989). In the Brazilian Bight, larvae of mesopelagic fish were

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associated with Tropical Water (TW) (Katsuragawa et al., 2014; Macedo-Soares et al., 2014), which is the warm and salty water mass transported southward by the Brazil Current

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(BC) in the mixing layer (Castro et al., 2006; Silveira et al., 2004, 2000). Two other water masses are present on the Southeast Brazilian Bight (SBB): the less warmer and salty

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South Atlantic Central Water (SACW) of oceanic origin, transported by BC at the pycnocline and Coastal Water (CW), which is formed by mixing between water originating from

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continental discharge and the other two masses. Physical oceanographic studies have demonstrated that coastal and oceanic

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systems interact (Calado et al., 2010; Lee and Pietrafesa, 1987; Palóczy et al., 2014). In the midst of the BC dominated oceanic region and the coastal zone, Ekman transport begins to play a role that might also be important in spreading lanternfish larvae over the SBB continental shelf. In the SBB continental shelf, the northeast winds persist for several days, during the austral summer and promote oceanward transport of the surface Ekman layer, which favors upwelling (Calado et al., 2010; Castelao and Barth, 2006; Palóczy et al., 2014). During austral winter, when atmospheric cold fronts pass more intensely by the region, the surface winds rotate and blow from the southern quadrant, which is favorable to coastward transport of the surface Ekman layer and may cause downwelling (Calado et al., 2010; Castelao and Barth, 2006; Rodrigues and Lorenzzetti, 2001; Stech and Lorenzzetti, 1992).

ACCEPTED MANUSCRIPT These processes lead to a very different vertical condition of the water column during summer and winter that may affect the larval distribution (Muhling and Beckeley, 2007). In Pacific and North Atlantic, larvae of the two lanternfish subfamilies show a different pattern of vertical distribution: Lampanyctinae larvae are located in shallower strata than Myctophinae larvae (Loeb, 1979; Moser and Smith, 1993; Sassa et al., 2002). Although one can assume that the vertical distribution of these larvae could be the same elsewhere, larval

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behavior vary in relation to the environment and local oceanographic processes, which, as

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mentioned before, are very dynamic in the SBB.

Therefore, in the present study we investigate the effects of the Ekman transport and

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BC on the vertical and horizontal distribution of lanternfish larvae in the SBB, during summer

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and winter.

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2. Material and Methods

Two oceanographic surveys were carried out with the R/V “Prof. W. Besnard” during

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austral summer (January 2002) and winter (August 2002) from Cape Frio (23ºS) to São Sebastião Island (25ºS), in the SSB. Each survey comprised a total of 72 stations during

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summer and 66 stations during winter arranged into 14 cross-shelf transects (Figure 1). Ichthyoplankton samples were obtained with Bongo nets in all oceanographic stations and with

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Multi Plankton Sampler (MPS) in each other oceanographic transect (n= 29 in summer; n= 22 in winter) (Figure 1). The Bongo nets (mouth diameter 0.6 m) were fitted with 333 µm and 505

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µm mesh sizes and were towed obliquely according to Smith and Richardson (1977). The maximum sampling depth of the tows was limited to 5.0 m above the bottom in shallow

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stations and to upper 200 m of the water column at offshore stations. Only samples of 333 µm mesh nets were analyzed. The MPS were fitted with 333 µm mesh size and were towed obliquely in five target depth layers (0-20 m, 20-40 m, 40-60m, 60-80 m and 80-100 m). Cable length was limited to 100 m depth, however highest abundance and diversity of lanternfish larvae usually occur within the upper 100 m depth layer (Loeb, 1979; Sassa et al., 2002). Flowmeters in the net mouths measured the volumes of water sampled to estimate the level of larval abundance. All samples were fixed in a buffered 4% formaldehyde-seawater solution. The samples are stored in the Biological Collection of “Prof. Edmundo F. Nonato” Oceanographic Institute, University of São Paulo. Temperature and salinity were measured vertically with a CTD profiler at each

ACCEPTED MANUSCRIPT station. Classical water mass percentage analysis (Mamayev, 1975) was employed to verify the relative contribution of CW, TW and SACW onto the shelf of the study area. This simple approach allows spatial and temporal identification of the water masses, and facilitates analysis of other related oceanographic processes. The characteristic thermohaline indices of CW, TW, and SAWC are inferred directly from the scattered T-S diagram of the summer and winter oceanographic expeditions. The percentage analysis has been successfully

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applied to the investigated region (Castro, 1977, Castro et al., 1977; Foloni Neto, 2010;

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Soares and Möller, 2001).

Daily wind stress maps were plotted from the dataset “erdQSstress3day”, available

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from the National Oceanic and Atmospheric Administration/ National Marine Fisheries Service/ Southwest Fisheries Science Center/ Environment Research. The 3-day composite

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data from the SeaWinds scatterometer of NASA's QuikSCAT satellite is 0.25° gridded from 1999-2009 (http://coastwatch.pfeg.noaa.gov/erddap/griddap/erdQSstress3day.html).

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From the wind stress maps the Ekman transport and Ekman layer depth were calculated following Cushman-Roisin and Beckers (2011). The mean Ekman layer velocity

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was calculated dividing the transport by the layer depth at each grid point. The Ekman velocity maps were plotted for each day of the summer and winter expeditions, following the

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wind stress fields.

Daily-mean geostrophic currents were calculated from the surface streamline function

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field (Kundu, 2008) derived from AVISO+ maps of absolute dynamic topography on a 0.25° grid. This data set is obtained through merging from altimetry sensors from two satellites

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(DT-GLOBAL-TWOSAT-MADT-H). These velocities, obtained for the purpose of a qualitative analysis, were superimposed on Optimum Interpolated Sea Surface Temperature

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(OISST) maps. The NOAA OISST data is an analysis constructed by combining different observations, using optimum interpolation to fill in gaps. OISST for this study uses satellite SST from Advanced High Resolution Radiometer (AVHRR) only, in a regular 0.25° grid (ftp://eclipse.ncdc.noaa.gov/pub/OI-daily-v2/NetCDF/). Fish larvae were identified at the species level, when possible, based on Bonecker and Castro (2006), Moser and Ahlstrom (1996) and Moser and Watson (2006) and they were classified according to their stage of development as preflexion, flexion, postflexion and transformation stages (Richards, 2006). Although descriptions of larval stages of some Diaphus and Lampanyctus species are available (Moser and Ahlstrom, 1996, Moser and Watson, 2006, Sassa et al., 2003) few

ACCEPTED MANUSCRIPT characters are useful to distinguish among 20 species of Diaphus and 10 species of Lampanyctus occurring along the Brazilian coast (Braga et al., 2008; Nafpaktitis et al., 1977; Santos and Figueiredo, 2008), specially the earliest stages. Larvae of Diaphus were separated in "stubby" type (hereafter referred to as Diaphus stubby), with high body and only one or two ventral pigments in the caudal peduncle, and "slender" type (hereafter referred to as Diaphus slender), with elongated body with a series of ventral pigment along

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the caudal peduncle. The larvae of the "stubby" type belong to the group of Diaphus which

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have no supranasal photophores (SNO), while the larvae of "slender" type belong to the group that have one (Moser and Ahsltrom, 1974; Ozawa, 1986; Sassa et al., 2003).

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However, some larvae of Diaphus did not show a very clear pattern of pigmentation and could not be classified in any of the two types. Lampanyctus larvae were separated in

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Lampanyctus type 1 and Lampanyctus type 2, based on unique morphological characteristics, indicating that at least two distinct groups can exist. Lampanyctus type 1 is

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similar to the Lampanyctus sp. described in Moser and Watson (2006). It shows deep, highly compressed body, large head, large dorsal and ventral finfolds and distinctly

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pigments on dorsum and on finfolds. The authors assigned this larva tentatively to L. photonotus, but they highlighted that the general morphology of Lampanyctus sp. is similar

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to L. lepidolychnus (Olivar and Beckley, 1997). The Lampanyctus type 2 larvae are deep bodied, with large head, large rounded eyes and snout blunt. Their general morphology is

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similar to L. alatus and L. tenuiformes described in Moser and Watson (2006). Larval distribution in the area was described using larval abundance (larvae.m‫־‬²) of

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samples obtained with a Bongo net. It was calculated using the number of larvae taken in sampling per volume of water filtered (m³) and multiplied by the depth of haul (m). Volume of

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water filtered was calculated as V = a*n*c, where a = area of net mouth (0.2827 m 2), n = number of rotations of the flowmeter and c = calibration factor of the flowmeter. Larval vertical distribution was described using larval density (larvae.100 m‫־‬³) of MPS samples. It was calculated by the ratio between the number of larvae taken in the sample and the volume of water filtered (m³) multiplied by 100. The non-parametric Mann-Whitney test was used to verify the existence of difference in larval abundance between summer and winter cruises using larval abundance of Bongo net. MPS samples were used to perform univariate PERMANOVA based on Euclidean distance between samples to test vertical variation of the most abundant species. Species

ACCEPTED MANUSCRIPT density was log(x+1) transformed prior to analyses. The factors included in the univariate PERMANOVA design were: sampling depth: 0-20 m, 20-40 m, 40-60m, 60-80 m and 80-100 m (5 levels, fixed) and light condition: sunrise, day, sunset, night (4 levels, random). Detrended correspondence analysis (DCA) was performed using larval density of MPS samples. DCA results showed length of gradients >3 that indicate a unimodal trend in species distribution (ter Braak, 1994), during summer and winter. Based on the DCA results,

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canonical correspondence analysis (CCA) using CANOCO 4.5 was employed to investigate

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the relationships between larval fish and their environment, because it maximizes the separation of their niches (ter Braak, 1986; ter Braak and Prentice, 1988; ter Braak and

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Verdonschot, 1995). Larval densities were log(x+1) transformed to reduce the weighting of dominant taxa, and rare taxa, i.e. the least abundant ones, were down weighted prior to

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analysis (Lepš and Šmilauer, 2003). Six environmental variables (temperature (°C), salinity, plankton biovolume (ml.m-³), sampling layer (m), distance from the coast (km) and transect)

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were used. Temperature, salinity, biovolume and distance from the coast were measured directly as continuous variables. Sampling depth and transect were measured as ordinal

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variables. Temperature and salinity used in CCA analysis were their mean values in each sampling layer, which were calculated using data from CTD vertical profile.

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Monte Carlo permutation test (9999 unrestricted permutations) was used for forward stepwise selection of variables (p<0.05). Statistical significance of the first four axes and of

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3. Results

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the sum of all constrained eigenvalues of the CCA model was also tested.

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3.1. Oceanographic Conditions

During both cruises three water masses were present up to 200 m depth: CW, TW and SACW. Scattered T-S diagrams (Figure 2 a, b) showed a predominantly triangular distribution, indicating mixing processes among the three water masses. Winds from the northeast, which are favorable to coastal upwelling in the region, blew during the summer cruise, except for a four-day period when winds from south/southwest from a cold front passed by the region without modifying the main dynamic scenario in the region. The summer cruise scenario can be depicted by examining the data from January 11 of 2002 (Figure 3). During the transect 6 sampling (red dots on Figure 3c), northeast

ACCEPTED MANUSCRIPT winds (Figure 3a) generated Ekman transport (Figure 3b) offshore, allowing the SACW to enter the continental shelf. This water mass signal can be detected in the SST image (Figure 3c), where cooler temperatures are found in the vicinities of Cape Frio relative to its surroundings and outer portions of the continental shelf. Indeed, the transect 6´s water mass percentage analysis clearly shows SACW over the inner continental shelf (Figure 3d). However, at the surface, the water is still a mixture of the three water types. Geostrophic

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currents (Figure 3c) show the cyclonic Cape Frio Eddy as described by Silveira et al. (2004)

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in the study domain.

Although winter is commonly referred to as a period when coastal downwelling

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prevails, the cruise comprised a period when winds were favoring upwelling (Figure 4 a, b). However, despite the northeasterly winds blowing, the water mass percentage analysis

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results exhibit a wintery scenario and suggesting an impinging TW tongue, a restricted SAWC front and a coastally trapped CW (Figure 4d). As for the geostrophic currents, a BC

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flowing virtually parallel to the isobaths is observed from approximately west of 43°, crossing transect 2 (red dots on Figure 4c).

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Estimated geostrophic velocities were in general one order of magnitude greater than mean Ekman surface layer integrated velocities during both the summer and winter cruises.

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The BC system during the summer cruise accounted for the highest velocity values, reaching ~0.87 m.s-1 (Figure 3c). The winter cruise showed BC maximum velocities of ~0.63

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m.s-1 (Figure 4c). As commented before, in the BC domain, both summer/winter values were one order of magnitude greater when compared to ~0.03 m.s -1 at the Ekman surface

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layer. As we approach shallower areas, however, comparing Figures 3b and 3c (4b and 4c), one can see that Ekman's velocity gradually becomes more significant until it becomes the

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leading mechanism of the dynamics at the inner shelf.

3.2. Taxonomic composition

Myctophidae was the third most abundant family in the samples, after Engraulidae and Sternoptychidae. A total of 2,459 lanternfish larvae were collected with Bongo nets, which represented 11% of the total fish larvae sampled. The MPS captured a total of 1,080 lanternfish larvae. The samples included 19 species or types belonging to 15 genera (Tables 1, 2 and 3). Among these taxa, Centrobranchus nigroocelatus, Myctophum selenops and Lampadena spp. were collected only with Bongo nets and larvae of

ACCEPTED MANUSCRIPT Diogenichthys atlanticus only with MPS.

3.3 Abundance and horizontal distribution

During summer, the mean total abundance of lanternfish larvae was 14.6 ± 20.0 larvae.m-2, and the most abundant taxa were Myctophum affine (27.1%), Diaphus stubby

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(17.7%), Diaphus spp. (12.6%), Lepidophanes spp. (7.7%), Hygophum reinhardtii (2.0%),

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Diaphus slender (1.9%), Myctophum obtusirostre (1.7%) and Benthosema suborbitale (1.2%). Another 13 taxa represented less than 1% each and unidentified larvae accounted

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for 24.3% of the total lanternfish larval abundance (Table 1). In winter, the mean abundance was 10.4 ± 13.2 larvae.m-2, and the most abundant taxa was Diaphus spp. (15.1%),

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Diaphus stubby (13.5%), M. affine (9.9%), Diaphus slender (4.6%), L. guentheri (4.5%), Lepidophanes spp. (2.9%), N. caudispinosus (2.5%), Hygophum hygomii (2.3%), Hygophum

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spp. (2.3%), Lampanyctus type 1 (2.2%), B. suborbitale (2.0%) and Myctophum spp. (1.4%). Another nine taxa represented less than 1% each and unidentified larvae accounted for

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33% of the total lanternfish larval abundance (Table 1). Centrobranchus nigroocelatus, D. atlanticus, Lampadena spp. and M. selenops

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occurred exclusively in the summer (Tables 1 and 2) and Lampanyctus type 1, Lampanyctus type 2, and Notoscopelus causdipinosus occurred only during winter (Table 1

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and 3). Myctophum affine, Diaphus stubby, Lepidophanes spp., H. reinhardtii, M. obtusirostre, Ceratoscopelus spp., N. valdiviae, Nannobrachium sp. and M. asperum were

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more abundant during summer, while L. gemellari, B. suborbitale, Diaphus tipo slender, H. hygomii, L. guentheri were more abundant during winter. However, difference in larval fish

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abundance between seasons were significant only for M. affine (p=0.0007). The majority of taxa were more abundant in the oceanic region (Figure 5), while M. affine and Diaphus stubby were more abundant in the outer shelf (Figure 5). These two species were widely distributed throughout the area and were more frequent and abundant beyond the 100 m isobath (Figures 6 and 7). They were also sparsely distributed in the inner shelf, where Myctophum affine was collected mainly in the north area during summer and at the south during winter (Figure 6). Diaphus stubby occurred in the inner shelf just during winter, in the north (Figure 7). The number of species that occurred in the inner shelf was greater in winter (eight species) than in summer (four species) (Figure 5). The species M. asperum, S. rufinus, Lampadena sp., Lampanyctus type 1, Lampanyctus typ e 2, L.

ACCEPTED MANUSCRIPT gemellari, Nannobrachium sp. and N. valdiviae were restricted to the oceanic region in both seasons (Figure 5). Most taxa were predominantly in preflexion stage in both seasons (Figure 8). The contribution of flexion and postflexion stages of M. affine in both seasons and L. guentheri during winter increase in the shelf region (Figure 8). Myctophum obtusirostre, Lepidophanes sp., H. reinhardtii, during summer, and B. suborbitale, Diaphus spp, showed the opposite

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pattern, with most developed larval stage being more abundant in the oceanic region

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(Figure 8).

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4. Vertical distribution

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The vertical distribution of lanternfish larvae was wider in summer comparing to winter (Figures 9 and 10). During winter less than 20% of lanternfish larval density occurred

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in the 0-20 m depth and less than 4% in the 80-100 m depth, while during summer more than 25% and 15% occurred in the 0-20 and 80-100 m depth respectively. During winter

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Lampanyctinae larvae did not occur in the 80-100 m depth, with exception of one positive sample of Diaphus stubby (Figure 9) and the Myctophinae larvae density were reduced in

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the 80-100 m depth (Figure 10).

Lanternfish larvae were predominantly in preflexion stage in all sampling depth during

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different light conditions, in both summer and winter (Figure 9 and 10). Only Diaphus slender, in both seasons, Diaphus stubby in winter and N. valdiviae in summer, were

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predominantly in flexion and postflexion stages (Figure 9 and 10). Comparing the influence of time of the day on most abundant taxa, M. affine

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concentration was almost equally distributed during day (48%) and at night (47%) in summer, while during winter 51% was found at night and 33% during the sunrise. Myctophum affine was distributed from surface to the 100 m depth during day and at night, with a peak of larval concentration in the 20-40 m depth at night, and highest concentration below 40 m depth during the day and sunrise/sunset. In winter M. affine was more abundant between surface and 40 m depth during the day and between 20-60 m depth at night and sunset (Figure 9), with 82% of its density between 20-60 m, combining the different times of the day. During summer, highest Diaphus stubby density was found above 40 m depth at night and during sunset/sunrise, and below the 40 m depth during the day. In winter highest

ACCEPTED MANUSCRIPT density between was observed between 20-60 m depth at night, above 40 m during the day and below 40 m during the sunrise. Diaphus stubby was more abundant at night in summer (79%) and winter (82%). Diaphus stubby was more abundant in the 40-60 m depth during the day, 20-40 m during sunrise/sunset and in the upper 40 m at night (Figure 10). In winter Diaphus stubby was equally distributed in the upper 80 m during the day, between 40-80 m during the sunrise and most abundant in the 20-60 m depth at night (Figure 10). Although

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the distribution of fish larvae was depth stratified, the univariated Permanova shows that

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differences in vertical distribution of developmental stages of M. affine and Diaphus stubby

condition (day/night/sunrise/sunset) (Tables 4 and 5).

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were neither significant among sampling depths nor among sampling depths within light

Monte Carlo test showed significant associations between species density and

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environmental variables (Table 6). Sampling Depth was significant in both seasons. Temperature, salinity and transect were significant only in the summer, while distance from

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the coast were significant only in winter. The biovolume was not significant in any season. CCA explained about 25% of variation in species matrix in the summer and 12% in winter

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(Table 7). The results were interpreted considering only the first and second axes, as they explained most of the species data and cumulatively accounted for 91% and 100% of the

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species-environment relation variance, in summer and winter respectively (Table 8). In the summer, species were distributed along a clear gradient of salinity and

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temperature, which varied horizontally and vertically (Figure 11). Pair values of temperature and salinity can be used as a proxy of water masses. The species S. rufinus and N.

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valdiviae were related to the deepest layers, TW and northernmost transects. Diaphus stubby, Diaphus slender and L. guentheri, however, were related with the shallowest layers,

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the mixture with CW and the southernmost transects (Figure 11). Myctophum affine was located almost at the center of the diagram, indicating it was distributed along the entire sampled water column, in all transects and associated with TW and its mixture with SACW. Other species were related only to a high percentage of TW (Figure 11). In the winter H. hygomii was associated with the greatest distance from the coast. Myctophum affine, Diaphus stubby, S. rufinus and L. guentheri were associated with intermediate distances. Regarding depth, B. suborbitale was related with the deepest layers and N. caudispinosus and L. guentheri, with the shallowest layers. Most species were situated close to the center of the diagram (Figure 12), indicating that they were related to the intermediate layers.

ACCEPTED MANUSCRIPT These results are mainly related to differences in the vertical and cross-shelf distribution of water masses, as illustrated for the most abundant taxa M. affine (Figures 13 and 14) and Diaphus stubby (Figures 15 and 16).

4. Discussion

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4.1. Taxonomic composition

In Brazilian waters the number of lanternfish identified in the larval stage (Bonecker

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et al., 2012; Castro et al., 2010; Katsuragawa et al., 2014; Nonaka et al., 2000) is much lower than the 81 species that were previously registered during adult and juveniles stages

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(Braga et al. 2014; Santos and Figueiredo, 2008). The low number of lanternfish species identified during the larval stage is also observed in other regions such as Benguela (Olivar

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and Beckley, 1994), probably because the early stages of development of the most myctophid species are unknown. For example, the Diaphus stubby larvae in this study could

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be, among the Diaphus that possess the SNO photophore (Braga et al. 2014; Santos and Figueiredo, 2008) D. garmani or D. dumerilli because the adults are very abundant in the

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area (Braga et al, 2014). However there is no description of the D. dumerilli larvae, and the D. garmani larvae described in the Pacific is very similar to the other Diaphus species. The

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genera Lampadena, Ceratoscopelus and Nannobrachium have fewer species, but their larvae presented few characteristics for species identification. Many lanternfish larvae were

they were

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not identified due to the very early stage of development, lack of pigments and/or because damaged,

preventing the

observation

of

meristic and

morphometric

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characteristics. Most of these larvae may belong to the genera Lepidophanes, Ceratoscoplelus and/or Diaphus, which have very similar body shape during very early stages.

4.2. Horizontal distribution

The horizontal distribution of most lanternfish larvae seems to be related to the adult distribution. The larvae of D. atlanticus were previously recorded in the vicinity of seamounts Vitória and Davis (20° S) (Castro et al., 2010) and it was considered one of the dominant species in the Mid-Atlantic and Walvis Ridges (Kobyliansky et al., 2010). However, in this

ACCEPTED MANUSCRIPT study they were rare, probably because few stations were sampled beyond the 1,500 m isobath. On the other hand, taxa that were previously recorded during the adult stage in the continental shelf, e.g. Diaphus spp. and N. caudispinosus and slope, e.g. M. affine (Figueiredo et al., 2002; Santos, 2003), were more abundant in the SBB. Despite differences in adult distribution, in all cases, lanternfish larvae were associated with the TW domain. The presence of lanternfish and other mesopelagic larvae

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in the shelf was previously associated with TW in the southern (Franco and Muelbert, 2003,

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Macedo-Soares et al., 2014) and southeast coast of Brazil (Katsuragawa et al., 2014) by their surface temperature. Nevertheless, the present study confirmed the lanternfish larval

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association with TW, using stratified vertical sampling and vertical profiles of temperature and salinity.

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Climatologically, the NE winds prevail in the summer, facilitating the intrusion of SACW on the continental shelf. Indeed, during the summer cruise period, the TW was

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limited to the outer shelf, preventing the larval transport to the innermost areas. An exception can be seen at transect 12, where the BC interaction with the shelf promotes

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onshore transport of TW, along with lanternfish larvae (Figure 13 and 15). In the winter climatology of the study region, the SE winds become more frequent, and might favor the

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intrusion of TW in several areas of the inner shelf (Castelao and Barth, 2006). However, during our winter cruise period, the presence of lanternfish larvae collected in the inner shelf

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of the southernmost transects seem still to be related to the BC cyclonic meander observed (Figure 4), which advects particles in a cross-isobath orientation at ~44°W. The BC

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velocities are much greater than those of the Ekman surface drift in summer and winter in the TW domain. Therefore, during both cruises, the presence of lanternfish larvae in the

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continental shelf was related to the pattern of TW intrusion, which occurs majorly through meanders and eddies of BC (Campos et al., 2000; Castelao and Barth, 2006, Palóczy et al., 2014). The Ekman transport influence is likely to be restrained to the innermost areas, where the BC influence is negligible (Castro, 2014; Castro et al., 1987). In the inner portions of the shelf, the BC influence is limited if not absent. Therefore, we speculate that the Ekman transport plays a major role in larvae distribution at depths shallower than 70 m since the wind is one of the main mechanism of generating currents in the inner continental shelf (Castro, 2014; Cerda and Castro, 2014; Dottori and Castro, 2009; Palóczy et al. 2014) and the lanternfish larvae were in the Ekman layer in most cases. Continental shelf productivity is higher than the oceanic area (Lopes et al., 2016),

ACCEPTED MANUSCRIPT thus the spread of mesopelagic larvae over the shelf could enhance their survival because there is more prey available. In general, lanternfish larvae prey upon copepod, both nauplii and adult stages, ostracods, euphasiids and appendicularians (Bernal et al., 2013; Conley and Hopkins, 2003; Sassa, 2010; Sassa and Kawaguchi 2005). Myctophidae larvae feeding was not studied in the SBB to the date, but they exhibit a relatively high feeding incidence elsewhere (Bernal et al., 2013; Conley and Hopkins, 2004; Sassa, 2010) and the nutritional

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condition of M. affine larvae was classified as good in the area (Namiki, 2013).

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During summer, in addition to the agglomeration of zooplankton biomass in the coastal area, due to the enrichment of surface waters by coastal upwelling, patches of

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zooplankton also occur in the outer shelf and close to the slope (Lopes et al., 2006). Therefore, lanternfish with higher larval abundance during summer, e.g. M. affine and

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Diaphus stubby, match the zooplankton production as the epipelagic coastal species Brazilian sardine (Matsuura, 1996). However, other lanternfish species were more abundant

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in the oceanic region and showed no difference in abundance between seasons. The spawning year-round, common among mesopelagic species (Gartner, 1993), should reflect

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the relative stability of an oceanic region (Doyle et al., 1993) that suffers less seasonal variation both in oceanographic conditions and in zooplankton biomass (Lopes et al., 2006).

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In the Benguela Current, larvae of mesopelagic species that occur on the shelf are more abundant during periods of intense upwelling, while those limited to the slope are relatively

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less abundant and occur throughout the year (Olivar and Shelton, 1993). Doyle et al. (1993) proposes that there is a co-evolution of reproductive strategies of fish associated with the

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complexity of the marine environment, which structure the distribution of ichthyoplankton. The spawning year-round and the higher larval abundance in the oceanic area could be a

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behavior to avoid competition with the species that occupy the shelf and spawn intensely during summer.

4.3. Vertical distribution

The lanternfish are subdivided into two subfamilies: Lampanyctinae and Myctophinae that show different patterns of vertical distribution. Myctophinae larvae, which typically occur at greater depths, have elliptical eyes, while the majority of Lampanyctinae, which occur in more surface layers, have rounded eyes (Moser and Ahlstrom, 1974). It is believed that elliptical eyes are an adaptation to deeper regions, because they possess a greater rotation

ACCEPTED MANUSCRIPT capacity, which allows a wider field of vision compared to rounded eyes and, consequently, a greater chance of locating prey (Weihs and Moser, 1981). The pattern of vertical distribution of both subfamilies was clearly observed only during summer, when Myctophinae larvae were more abundant in the deepest layers than Lampanyctinae, with exception of N. valdiviae and L. gemellari. Despite belonging to Lampanyctinae, the larvae of N. valdiviae has elliptical eyes and L. gemellarii has slightly oval eyes (Moser and

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Watson, 2006) and both were collected below 60 m depth also in other studies (Loeb, 1979;

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Sassa et al., 2002). In winter although larvae of the Lampanyctinae L. guentheri and N. caudispinosus occupied the uppermost layers and the larvae of Myctophinae B. suborbitale

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occupied the deepest layers, most species of both subfamilies were more abundant between 20 and 60 m depth. The vertical distribution of larvae varies over time and space

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because it is affected by oceanographic conditions such as temperature and depth of the mixing layer (Muhling and Beckeley, 2007). Therefore, the concentration of larvae of both

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subfamilies in the 20-60 m depth during winter was probably caused by the greater mixing of water masses on the surface and 80-100 m depth.

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The sampling depth of the present study did not cover the entire distribution range of some taxa, especially those that occurred in the deepest sampling layers. In the Kuroshio

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Current, for example, D. atlanticus is located between 75-150 m depth (Sassa et al., 2002), indicating that it must be more abundant at depths higher than those we have sampled.

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Nevertheless, the frequency of occurrence and the center of distribution of different species have been quite consistent in relation to their occurrence at the TW and low occurrence of

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lanternfish larvae at SACW was also verified in Campos Basin (Bonecker et al., 2012). On the California coast lanternfish larvae were distributed above the thermocline, predominantly

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in the upper mixing layer (Ahlstrom, 1959) and most lanternfish larval abundance and highest diversity occur within the upper 100 m layer in the Pacific Central Gyre (Loeb, 1979) and Kuroshio Current (Sassa et al., 2002). This indicates that sampling to 100 m depth is enough to characterize most the vertical distribution of preflexion and flexion lanternfish larvae in the SBB. Some offshore spawning species may use different strategies of vertical distribution to ensure their larvae reach the shelf area by the deep or surface onshore Ekman transport, during upwelling and downwelling events respectively (Rodriguez et al., 2015). The presence of most lanternfish larvae in the first 100 m depth confirm they could be subject to the surface onshore Ekman transport in the inner shelf of the SBB.

ACCEPTED MANUSCRIPT 5. Conclusion

In the present study, larvae of lanternfish species, which during the adult stage usually occur in open ocean beyond the continental slope, were abundant in the SBB continental shelf. The scenarios depicted in both the summer and winter cruises were characterized by

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BC-related currents which are about one order of magnitude larger than those associated

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with the Ekman surface drift. Therefore, we show that lanternfish larval distribution in the SBB is mostly driven by the mesoscale activity of the BC and its interaction with the

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continental shelf.

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Acknowledgments

We thank the CNPq, PRONEX and FINEP for funding the DEPROAS cruises and the CNPq

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for providing the PhD grant to Claudia Namiki (140096/2009-4). Special thanks to Nancy K. Taniguchi for helping with the oceanographic data set and to Gisela M. de Figueiredo, José

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L. de Figueiredo, June F. Dias and Paulo O. Mafalda Junior for great advices. We also

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thank the reviewers that help to improve this work.

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Figure 1. Study area and the location of oceanographic transects, showing oceanographic and biological sampling stations in the SBB during austral summer (January 2002) and winter

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Figure 2. Temperature-salinity (T-S) diagram for summer (a) and winter (b) 2002 in the SBB. The water masses are Coastal Water (CW), Tropical Water (TW) and South Atlantic Central

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Figure 3. Summer scenario during January 11 of 2002 in the SBB. Panel (a): wind stress [Pa];

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water masses percentage and Myctophum affine larval density [larvae.m-3] across the

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transect 6 . Red dots indicate the location of transect 6.

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Figure 4. Winter scenario during August 04 of 2002 in the SBB. Panel (a): wind stress [Pa];

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water masses percentage and Myctophum affine larval density [larvae.m-3] across the

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transect 2 . Red dots indicate the location of transect 2.

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Figure 5. Mean abundance of lanternfish larvae in the SBB during austral summer and winter 2002. BENSU, Benthosema suborbitale; CENIG, Centrobranchus nigroocellatus; HYGHY, Hygophum hygomii; HYGRE, Hygophum reinhardtii; MYCAF, Myctophum affine; MYCAS, Myctophum asperum; MYCOB, Myctophum obtusirostre; MYCSE, Myctophum selenops; SYMRU, Symbolophorus rufinus; CER, Ceratoscopelus sp.; DIA, Diaphus sp.; DIASL, Diaphus slender; DIASTU, Diaphus stubby; LAMPAD, Lampadena sp.; LAM1, Lampanyctus type 1; LAM2, Lampanyctus type 2; LEP, Lepdophanes sp.; LEPGE, Lepidophanes guentheri; LOBGE, Lobianchia gemellarii; NAN, Nannobrachium sp.; NOTVA, Notolychnus valdiviae; NOTCAU, Notoscopelus caudispinosus. Note that species may occur in just one season. Details are shown in Table 1.

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Figure 6. Abundance of Myctophum affine in the SBB during summer (a) and winter (b) 2002.

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Figure 7. Abundance of Diaphus stubby in the SBB during summer (a) and winter (b) 2002.

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Figure 8. Developmental stages distribution of the most abundant lanternfish larvae collected with the Bongo net during austral summer and winter in the SBB. Preflexion stage (light gray bars), flexion stage (dark gray bars), postflexion stage (black bars). Number of positive samples/total samples are indicated in each sampling depth.

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Figure 9. Vertical distribution of developmental stages of most abundant lanternfish larvae collected with MPS during austral summer. Preflexion stage (light gray bars), flexion stage

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Figure 10. Vertical distribution of developmental stages of most abundant lanternfish larvae collected with MPS during austral winter. Preflexion stage (light gray bars), flexion stage (dark

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Details are shown in Table 3.

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Figure 11. CCA ordination diagram with taxa and environmental variables (arrows) in the SBB during summer. BENSU, Benthosema suborbitale; DIASL, Diaphus slender; DIASTU,

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Diaphus stubby; HYGRE, Hygophum reinhardtii; LEPGE, Lepidophanes guentheri; LOBGE, Lobianchia gemellarii; MYCAF, Myctophum affine; MYCOB, Myctophum obtusirostre;

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NOTVA, Notolychnus valdiviae; SYMRU, Symbolophorus rufinus.

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Figure 12. CCA ordination diagram with taxa and environmental variables (arrows) in the SBB during winter. BENSU, Benthosema suborbitale; DIASL, Diaphus slender; DIASTU, Diaphus

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stubby; HYGHY, Hygophum hygomii; HYGRE, Hygophum reinhardtii; LAM1, Lampanyctus type 1; LAM2, Lampanyctus type 2; LEPGE, Lepidophanes guentheri; MYCAF, Myctophum

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affine; MYCNI, M. nitidulum; MYCOB, Myctophum obtusirostre; NOTCAU, Notoscopelus

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caudispinosus; SYMRU, Symbolophorus rufinus.

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Figure 13. Vertical distribution of Myctophum affine larvae and water masses percentage

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between surface and 100 m depth across the sampled transects from inner shelf to oceanic waters during summer. Coastal Water (CW), Tropical Water (TW) and South Atlantic Central Water (SACW).

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Figure 14. Vertical distribution of Myctophum affine larvae and water masses percentage

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between surface and 100 m depth across the sampled transects from inner shelf to oceanic waters during winter. Coastal Water (CW), Tropical Water (TW) and South Atlantic Central Water (SACW).

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Figure 15. Vertical distribution of Diaphus stubby larvae and water masses percentage

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between surface and 100 m depth across the sampled transects from inner shelf to oceanic waters during summer. Coastal Water (CW), Tropical Water (TW) and South Atlantic Central Water (SACW).

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Figure 16. Vertical distribution of Diaphus stubby larvae and water masses percentage between surface and 100 m depth across the sampled transects from inner shelf to oceanic waters during winter. Coastal Water (CW), Tropical Water (TW) and South Atlantic Central Water (SACW).

ACCEPTED MANUSCRIPT Table 1. Number of individuals (N), mean abundance, standard deviation (SD) and percentage (%), and frequence of ocurrence (%) of Myctophidae larvae collected with 333 µm bongo net, between Cape of São Tomé and São Sebastião Island, during the summer (n= 72) and the winter (n=66) of 2002. Abundance expressed in larvae m -2.

AC

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9 179 28 265 1 110 13 1 3 7 355 1444 72

0.10 1.83 0.28 2.58 0.01 1.12 0.13 0.01 0.03 0.08 3.55

0.89 0.7 5.61 12.6 1.60 1.9 7.76 17.7 0.08 0.1 4.10 7.7 0.91 0.9 0.09 0.1 0.26 0.2 0.39 0.6 5.98 24.3

21 24 21 10 18 97 1 4 6

1.4 34.7 8.3 29.2 1.4 18.1 2.8 1.4 1.4 5.6 41.7

6 147 43 160 23 6 29 42 1 1 2 24 329 1015 66

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0.21 0.24 0.24 0.10 0.15 1.03 0.01 0.06 0.06

0.65 0.74 0.88 0.28 0.67 1.82 0.09 0.37 0.26

2.0 2.3 2.3 0.9 1.4 9.9 0.1 0.5 0.5

16.7 16.7 12.1 13.6 7.6 40.9 1.5 3.0 6.1

0.05 1.57 0.47 1.41 0.23 0.07 0.30 0.47 0.01 0.01 0.02 0.26 3.44

0.18 0.5 4.14 15.1 1.48 4.6 3.02 13.5 0.76 2.2 0.29 0.7 1.06 2.9 1.71 4.5 0.12 0.1 0.08 0.1 0.15 0.2 0.55 2.5 6.30 33.1

7.6 31.8 16.7 31.8 13.6 7.6 12.1 18.2 1.5 1.5 1.5 24.2 40.9

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0.69 1.3 8.3 0.09 0.1 1.4 0.10 0.1 1.4 0.22 0.2 2.8 0.70 2.0 23.6 0.23 0.4 5.6 6.78 27.1 62.5 0.23 0.2 1.4 0.10 0.1 1.4 0.88 1.8 15.3 0.14 0.2 2.8 -

Winter Abundance (ind.m -2) F.O. mean sd (%) (%)

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0.18 0.01 0.01 0.03 0.30 0.06 3.95 0.03 0.01 0.26 0.02 -

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16 1 1 3 27 5 391 3 1 23 2 -

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Myctophinae Benthosema suborbitale Centrobranchus Hygophum spp. Hygophum hygomii Hygophum reinhardti Myctophum spp. Myctophum affine Myctophum asperum Myctophum nitidulum Myctophum obtusirostre Myctophm selenops Symbolophorus rufinus Lampanyctinae Ceratoscopelus spp. Diaphus spp. Diaphus slender type Diaphus stubby type Lampadena sp. Lampanyctus type 1 Lampanyctus type 2 Lepidophanes spp. Lepidophanes guentheri Lobianchia gemellari Nannobrachium sp. Notolychnus valdiviae Notoscopelus Larvae not identified Total number of larvae Number of samples

Summer Abundance (ind.m -2) F.O. mean sd (%) (%)

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M

Taxon

ACCEPTED MANUSCRIPT Table 2. Number of individuals (N), mean abundance, standard deviation (SD) and percentage (%), and frequence of ocurrence (%) of Myctophidae larvae collected with the Multi Plankton Sampler between Cape of São Tomé and São Sebastião Island, during the summer of 2002. Density expressed in larvae 100 m -3. Summer Taxon Myctophinae Benthosema suborbitale Diogenichthys atlanticus Hygophum spp. Hygophum hygomii Hygophum reinhardti Myctophum spp. Myctophum affine Myctophum asperum Myctophum nitidulum Myctophum obtusirostre Symbolophorus rufinus Lampanyctinae Ceratoscopelus spp. Diaphus spp. Diaphus slender type Diaphus stubby type Lampadena sp. Lampanyctus type 1 Lampanyctus type 2 Lepidophanes guentheri Lobianchia gemellari Nannobrachium sp. Notolychnus valdiviae Notoscopelus caudispinosus Larvae not identified Total number of larvae Number of samples

0-20 m Density (ind.100 m -3) F.O. mean sd % (%)

N 2 1 24 -

0.20 0.08 3.27 -

0.79 0.45 8.83 -

0.6 6.9 0.2 3.4 9.5 27.6 -

N 1 46 3 -

1 0.13 0.70 0.4 3.4 29 3.38 10.34 9.9 24.1 1 0.08 0.41 0.2 3.4 58 13.75 65.25 40.1 24.1 49 4.62 9.88 13.5 31.0 1 0.06 0.33 0.2 3.4 75 241 29

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-

-

-

C A

26 4.09 9.14 10.7 24.1 1 0.34 1.84 0.9 3.4 39 10.67 35.55 27.9 17.2 13 3.00 8.51 7.8 13.8 8 1.66 8.96 4.4 3.4 -

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-

52 189 29

40-60 m Density (ind.100 m -3) F.O. mean sd % (%)

N

0.21 1.12 0.5 3.4 2 2 1 1 9.03 20.57 23.7 41.4 46 - - 0.60 2.54 1.6 6.9 6 -

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8.72 20.23 25.4 37.9

20-40 m Density (ind.100 m -3) F.O. mean sd % (%)

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-

N 4 2 5 1 37 1 3 1

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4.46 10.37 21.0 29.6 3 0.19 0.98 0.9 3.7 1.86 8.18 8.8 11.1 4 1 - 0.23 1.19 1.1 3.7 1 2

-

-

-

8.58 15.55 22.5 41.4

18 101 27

3.05

-

-

80-100 m Density (ind.100 m -3) F.O. mean sd % (%)

N

1.26 6.04 6.0 4.3 0.36 1.73 1.7 4.3 1.29 3.60 6.2 13.0 0.24 1.17 1.2 4.3 9.62 14.19 46.0 52.2 0.28 1.35 1.3 4.3 0.73 2.56 3.5 8.7 0.28 1.35 1.3 4.3

1 1 2 1 59 1

0.15 0.68 0.7 4.8 0.30 1.35 1.4 4.8 0.59 2.71 2.8 4.8 0.20 0.94 1.0 4.8 8.24 16.27 38.5 38.1 0.30 1.35 1.4 4.8

0.61 0.76 0.24 0.15 0.60

2 2 51 2 1

0.41 1.29 1.9 0.14 0.63 0.6 9.66 44.27 45.2 0.35 1.13 1.6 0.30 1.35 1.4

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A

M

-

14 1 9 1 -

0.41 1.51 1.9 7.4 0.32 1.68 1.5 3.7 0.23 1.19 1.1 3.7 0.25 1.29 1.2 3.7 9.10 16.34 42.8 37.0 1.15 3.07 5.4 14.8 -

60-80 m Density (ind.100 m -3) F.O. mean sd % (%)

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-

-

5.22 14.3 29.6

17 82 23

4.47

2.07 2.78 1.16 0.71 1.98

2.9 3.6 1.2 0.7 2.9

8.7 8.7 4.3 4.3 8.7

-

-

-

-

-

-

-

-

7.21 21.4 43.5

3 126 21

0.76

2.78

3.5

9.5

Table 3. Number of individuals (N), mean abundance, standard deviation (SD) and percentage (%), and frequence of ocurrence (%) of Myctophidae larvae collected with the Multi Plankton Sampler between Cape of São Tomé and São Sebastião Island, during the winter of 2002. Density is expressed in larvae 100 m -3.

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9.5 4.8 4.8 9.5 4.8

ACCEPTED MANUSCRIPT Winter Taxon Myctophinae Benthosema suborbitale Diogenichthys atlanticus Hygophum spp. Hygophum hygomii Hygophum reinhardti Myctophum spp. Myctophum affine Myctophum asperum Myctophum nitidulum Myctophum obtusirostre Symbolophorus rufinus Lampanyctinae Ceratoscopelus spp. Diaphus spp. Diaphus slender type Diaphus stubby type Lampadena sp. Lampanyctus type 1 Lampanyctus type 2 Lepidophanes guentheri Lobianchia gemellari Nannobrachium sp. Notolychnus valdiviae Notoscopelus caudispinosus Larvae not identified Total number of larvae Number of samples

0-20 m Density (ind.100 m -3) F.O. mean sd % (%)

N

20-40 m Density (ind.100 m -3) F.O. mean sd % (%)

N

40-60 m Density (ind.100 m -3) F.O. mean sd % (%)

N

3 3 4 20 -

0.22 0.16 0.11 0.79 -

0.76 0.52 0.39 3.72 -

2.3 1.6 1.1 8.1 -

9.1 9.1 9.1 4.5 -

3 4 1 23 1 2 1

0.41 1.66 0.21 3.98 0.13 0.34 0.27

1.06 5.77 0.98 7.96 0.61 1.14 1.28

2.5 10.2 1.3 24.3 0.8 2.1 1.7

1 13.6 2 13.6 3 4.5 1 - 36.4 20 - 4.5 9.1 1 4.5 1

0.24 0.47 0.73 0.22 4.70 0.23 0.24

1.06 1.40 1.78 0.95 8.54 1.00 1.06

1.4 2.6 4.1 1.2 26.2 1.3 1.4

5.3 10.5 15.8 5.3 - 42.1 - 5.3 5.3

1 4 26 4 1 17 -

0.07 0.36 3.28 0.28 0.07 1.21 -

0.34 0.86 9.75 0.78 0.34 2.75 -

0.7 3.6 33.3 2.8 0.7 12.3 -

4.5 18.2 31.8 18.2 4.5 27.3 -

10 2 29 1 3 -

1.37 0.25 4.90 0.12 0.63 -

3.46 0.82 11.66 0.57 2.17 -

8.4 1.5 29.9 0.7 3.9 -

18.2 9.1 22.7 4.5 9.1 -

8.41 0.40 0.40 -

17.47 1.74 1.74 -

46.8 2.2 2.2 -

36.8 5.3 5.3 -

2

0.55

2.27

5.6

9.1

1

0.13

0.61

0.8

4.5

-

-

-

-

33 118 22

2.74

5.27

27.8

31.8

3.60

12.0

31.8

8 82 19

1.92

3.83

10.7

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12 93 21

1.96

43 1 1 -

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0.53 0.52 0.20 0.26 0.75 0.20 0.60

1.60 2.22 0.85 1.12 1.93 0.87 1.84

6.8 6.6 2.5 3.3 9.5 2.6 7.6

2 3 1 1

1 11 -

0.20 2.03 -

0.87 2.72 -

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-

-

26.3

11 37 18

2.59

4 1 2

80-100 m Density (ind.100 m -3) F.O. mean sd % (%)

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11.1 5.6 5.6 5.6 - 16.7 - 5.6 11.1

1 2 -

2.6 25.7 -

6.64

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U N

60-80 m Density (ind.100 m -3) F.O. mean sd % (%)

N

-

0.18 0.21 0.39 -

0.69 0.81 1.02 -

8.4 9.9 18.3 -

6.7 6.7 13.3 -

5.6 44.4 -

1 1 -

0.18 0.20 -

0.69 0.77 -

8.4 9.5 -

6.7 6.7 -

-

-

-

-

-

-

-

32.7

22.2

4 12 15

0.96

2.52

45.4

20.0

3

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Table 4: Univariate analysis of the influence of sampling depth and light condition on the abundance of M. affine in preflexion, flexion and postflexion stages during summer and winter 2002. Factors: Sa= sampling depth, Li= light condition.

Source

Sa

df 2 1 0

postflexion df MS Pseudo-F p-value 2 2.53E-02 0.58241 0.532 1 4.88E-02 1.1223 0.3503 0 No test

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Li Sa x Li

preflexion df MS Pseudo-F p-value 3 0.67267 1.629 0.5044 1 1.94E-03 2.64E-03 0.9589 1 0.21785 0.29592 0.5984

postflexion df MS Pseudo-F p-value 4 0.30225 2.672 0.1478 1 9.02E-04 1.36E-03 0.9702 4 0.11312 0.17044 0.9383

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Li Sa x Li

df 4 1 1

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Sa

preflexion MS Pseudo-F p-value 0.96743 1.2703 0.4276 0.12124 0.14434 0.8734 0.76157 0.90666 0.4856

df 4 2 4

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Source

Myctophum affine Summer flexion MS Pseudo-F p-value 0.19193 0.36986 0.8179 5.10E-02 0.28156 0.6096 0.55441 3.0626 0.1179 Winter flexion MS Pseudo-F p-value 14.546 98.307 0.1779 12.45 84.14 0.1556 No test

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Table 5: Univariate analysis of the influence of sampling depth and light condition on the abundance of Diaphus stubby in preflexion, flexion and postflexion stages during summer and winter 2002. Factors: Sa= sampling depth, Li= light condition.

Source

Sa Li Sa x Li

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df 4 1 3

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Li Sa x Li

preflexion MS Pseudo-F p-value 1.047 0.50488 0.679 1.9124 0.92218 0.4415 No test

preflexion df MS Pseudo-F p-value 3 1.74E-01 0.2791 0.8232 2 0.74368 1.325 0.3189 2 0.99792 1.778 0.2297

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Sa

df 3 2 0

df 3 2 3

postflexion df MS Pseudo-F p-value 3 9.51E-02 0.17276 0.91 2 1.012 1.8388 0.4421 0 No test

df 4 2 1

postflexion MS Pseudo-F p-value 1.0413 5.3573 0.298 0.18 0.64441 0.6001 0.18969 0.6791 0.5144

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Source

Diaphus stubby Summer flexion MS Pseudo-F p-value 0.89646 1.3225 0.4465 5.09E-03 1.74E-03 0.9675 0.6336 0.21658 0.8795 Winter flexion MS Pseudo-F p-value 0.86571 1.3626 0.2986 0.73656 3.8745 0.0851 0.63535 3.3421 0.0882

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Table 6. Summary of permutation test of Monte Carlo in relation to environmental and spatial variables between Cape of São Tomé and São Sebastião Island, during the summer and winter of 2002. Bold values refer to the significant variables. Analysis based on Multi Plankton Sampler data. Summer Winter Variables p F p F Salinity 0.0001 4.95 0.3070 1.17 Collection depth 0.0009 3.97 0.0145 2.10 Temperature 0.0018 4.00 0.2638 1.24 Transect 0.0031 3.75 0.2376 1.26 Biovolume 0.5069 0.74 0.6674 0.75 Coast distance 0.6836 0.69 0.0002 2.88

5.9% 6.1% 12.7% 75.4%

0.0% 3.8% 8.3% 87.9%

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Environmental - Spatial variation Environmental + Spatial variation Spatial - Environmental variation Unexplained variation

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Summer Winter Environmental Spatial Env - Spat Spat - Env Environmental Spatial Env - Spat Spat - Env 2,495 2,495 2,040 2,173 5,157 5,157 4.589 4.791 0.298 0.455 0.120 0.276 0.194 0.455 0 0.399 11.9% 18.2% 5.9% 12.7% 3.8% 8.8% 0.0% 8.3%

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Sum of all natural eigenvalues Sum of all canonical eigenvalues Explained variation

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Table 7. Summary of analysis of environmental and spatial variables partition in explaining species of Myctophidae larvae distribution between Cape of São Tomé and São Sebastião Island during the summer and winter of 2002. Analysis based on Multi Plankton Sampler data.

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Table 8. Summary of CCA performed on larval fish density of Multi Plankton Sampler, relating species with environmental variables between Cape of São Tomé and São Sebastião Island, during summer and winter of 2002. 1 0.289 0.643

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Eigenvalues Species-environment correlation Percentage of cumulative variance: in the data of species in relation environment species Sum of all natural eigenvalues Sum of all canonical eigenvalues Salinity Temperature Collection depth Transect Distance from the coast

11.6 50.3

Summer Axes 2 3 0.232 0.042 0.643 0.358 20.9 90.6

22.6 98

Total 4 inertia 0.012 2.495 0.329 23 100

1 0.267 0.659 5.2 58.7

Winter Axes Total 2 3 4 inertia 0.188 0.728 0.682 5.157 0.622 0 0 8.8 100

22.9 0

36.2 0

2.495 0.575 -0.7645 -0.5630 -0.3390 -0.0601

0.1906 -0.3993 0.7298 0.5537

-0.0392 0.6514 -0.4615 0.7352

-0.6145 -0.3151 0.3735 -0.3864

5.157 0.455

-0.12

0.993

0

0

0.988

0.156

0

0

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Highlight The mesoscale activity of the Brazil Current influences lanternfish larvae distribution in the SBB. The Ekman dynamic influences the distribution of lanternfish larvae in the innershelf. The vertical distribution of lanternfish larvae changed between summer and winter.

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