Article Details

Neritic Larval Fish Distribution in the Oceanic Area of the Campeche Bay, Gulf of Mexico

Abstract Citation Introduction Materials and Methods Results Discussion Acknowledgment References

Flores-Coto C*, Zavala-García F and Sanvicente-Añorve L

Instituto de Ciencias del Mar y Limnología. Universidad Nacional Autónoma de México, Mexico

Corresponding Author:

Flores-Coto C, Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, CDMX, México, Tel: 55 5622 5811

Keywords: Deep distribution; Oceanic sub-regions; Larval fish

Abstract

Composition and abundance of larval fishes in Campeche Bay were studied during two seasons, winter, 2013 (24 stations) and summer, 2014 (31 stations). Sampling was carried out with open-close nets, mouth 75 cm and 505 µm mesh. The data of salinity and temperature allowed distinguishing three oceanic sub-regions: North, East, and West. There were 236 taxa, belonging to 74 families, 168 species, 154 taxa occurred in winter and 171 in summer; the composition in both cruises was similar with around 70 % of oceanic and 30 % of neritic larvae. The larval density was almost three times larger in summer than winter. The Campeche Bay hydrodynamics fits well with the results; the West sub-region is located where a cyclonic gyre takes place, the North and East sub regions are located in the area of influence of warm currents over the Yucatan shelf. The hydrodynamics also allows understanding the differences in the proportion of neritic larvae among the three oceanic sub-regions, the West and East with the lower and higher number of neritic larvae, respectively. The large difference among regions is related to some neritic taxa occurring exclusively in some of them. Of the total taxa, 55 neritic occurred only once and it means that more than a half of neritic taxa were represented by one organism, 31 from 85 neritic taxa occurred in the layers of 600 to 1000 m depth. Of the neritic larvae, only Syacium papillosum and Apogon sp. appear among the 20 more abundant.

Citation

Flores-Coto C, Zavala-García F and Sanvicente-Añorve L. Neritic Larval Fish Distribution in the Oceanic Area of the Campeche Bay, Gulf of Mexico. Int J Fisheries Sci Res. 2018; 2(1): 1003.

Introduction

The composition, abundance, and distribution of ichthyoplankton in the oceanic waters of Campeche Bay, the southern Gulf of Mexico have not been studied. Most the studies have been carried out on the continental shelf, and few include some oceanic stations but no samples below 200 m. It could mention the papers of Olvera-Limas et al., [1] on some particular species, Flores-Coto and Ordónez-López [2] on the mesopelagic fishes, Flores-Coto and Sánchez-Ramírez [3] on carangids, and Flores et al., [4] with a summary of ichthyoplankton research in the area.

Such papers let us have an idea of the dominant taxa in a general way, for an instant, the dominance in the oceanic area of families of mesopelagic species mainly Myctophidae and Gonostomatidae and those of neritic habitat on the continental shelf, like Sciaenidae, Carangidae, Clupeidae, Engraulidae.

Larval fish distribution around the world depends on the biology of the species and the hydrodynamic regime [5-10]. On the first item, the plankton biomass which means food availability is significant. In the southern Gulf of Mexico, the zooplankton biomass and ichthyoplankton density have a direct relationship, and their distribution patterns are similar, with the lowest values in the oceanic area [11,12]; high biomass densities occurred in the coastal regions of the main rivers [13].

In the continental shelf off Tabasco y Campeche, the hydrodynamics are dominated by a current over the Yucatan shelf (which is a branch of the Yucatán current) and the continental water discharges; environment very different from the oceanic adjacent area in the Campeche Bay where the hydrodynamic is driven by a semi-permanent cyclonic gyre [14] and coastal currents generated by wind force, which change of direction depending on the climatic season [15].

But also the Yucatan shelf current (from now on called YSC) has a significant role because the composition and distribution of larval fish assemblages are determinate firstly by the reproductive habits of the adults, but finally, modulated by the hydrographic stressors that characterize each area.

Richards et al., [16] in their analysis of larvae at the borders of the Loop current in the center of the gulf, recorded among the 25 more abundant families a high proportion of neritic components; however, one would expect in the oceanic waters of Campeche Bay a lower density of larvae and a lower proportion of neritic larvae, as it has been previously reported by Flores-Coto et al., [17] and Espinosa-Fuentes et al., [18].

The present paper analyzes the neritic larval fish composition and distribution in the oceanic waters of Campeche Bay from the surface to 1000m depth, during winter and summer seasons of 2013 and 2014, assuming that the dominant hydrographic features are the controlling factors and to a lesser extent, the larvae biology.

Materials and Methods

The study area was located in the oceanic waters of Campeche Bay, the southern Gulf of Mexico between 18º and 21º 30´N and 93º and 96ºW. The zooplankton samples were obtained, from 24 stations made during the winter (ZOOMEP I, January 23-February 3, 2013) and 31 in the summer (ZOOMEP II, June 4-14, 2014). In the sampling, we used open-close nets 75 cm mouth and 505µm mesh, in five 200 m strata (0 to 1000m) resulting in 270 samples (Figure 1).

Figure 1: Study area and stations located in Campeche Bay, Gulf of Mexico. a - Winter cruise, b - Summer cruise

Samples were initially preserved in 4% formalin neutralized with sodium borate and changed into 70% alcohol 48h later. Fish larvae were all extracted from each sample and identified to the lower taxonomic level as possible. Salinity and temperature data were obtained from a Conductivity, Temperature and Depth profiler (CTD).

For the recognition of larvae assemblages, the Bray-Curtis similarity index was applied [19].

Results

Oceanic regions

Temperature and salinity properties of the sampled water masses allowed the identification of three oceanic sub-regions, here designated as North (NOR), East (EOR), and West (WOR) (Figures 2a and 2b).

Figure 2: Oceanic sub-regions delimited by the average temperature of 0-200 m layer.

<20°C, West sub-region, >21°C North sub-region, between 20 and 21°C East sub-region.

During the winter cruise considering the average temperature of the first level (0 - 200 m) the NOR had values of > 21°C, the WOR of < 19.8°C and the EOR between 20 and 21°C (Figure 2a). Interestingly, during the summer cruise, the same sub-regions were also identified, in similar geographic areas. The NOR with temperatures of > 21°C, the WOR with < 20°C, and the EOR with values between previous sub-regions (Figure 2b). The differences among sub-regions in both cruises were lower but persisted to depth levels of 800 - 1000 m (Figures 3a and 3b).

Figure 3: Temperature profiles (0-1000 m) in three sub-regions of the Campeche Bay, Mexico, during winter and summer cruises.

Salinity differences among sub-regions were small, in the winter cruise. The average values were <36.35 in WOR, >36.4 in the NOR and 36.33 to 36.4 in the EOR. In the summer cruise the values were 34.42 to 36.25, in the NOR, 36.00 to 36.24 in the WOR and 36.17 to 36.26 in the EOR (Figures 4a and 4b).

Figure 4: Salinity profiles (0-1000 m) in three sub-regions of the Campeche Bay, Mexico, during winter and summer cruises.

Common and exclusive taxa in the oceanic sub-regions

A total of 5,612 larvae were collected, only 3,431 larvae were identified corresponding to 236 taxa, 76 families, 148 genera, 161 species. In winter 154 taxa were recorded and 171 in the summer. About 88 taxa were common, 66 only occurred in winter and 83 in summer (Tables 1, 2, 3 and 4).

Table 1: Common taxa among oceanic sub-regions in Campeche Bay, Gulf of Mexico, during winter cruise (January 25 to February 3, 2013).

COMMON TAXA IN THREE OCEANIC SUB-REGIONS
Family Taxa Habitat NOR EOR WOR
Antenaridae Antennarius spp. Neritic 0.013 0.01 0.032
Mugilidae Mugil cephalus Neritic 0.006 0.01 0.004
Congridae Rhynchoconger flavus Neritic 0.011 0.009 0.003
Sternoptychidae Argyropelecus slandeni Oceanic 0.034 0.003 0.012
Sternoptychidae Argyropelecus spp. Oceanic 0.127 0.101 0.077
Myctophidae Benthosema suborbitale Oceanic 0.041 0.144 0.161
Bregmacerotidae Bregmaceros atlanticus Oceanic 0.015 0.055 0.031
Bregmacerotidae Bregmaceros cantori Oceanic 0.023 0.39 0.008
Chiasmodontidae Chiasmodon niger Oceanic 0.015 0.032 0.03
Gonostomatidae Cyclothone acclinidens Oceanic 0.01 0.002 0.03
Gonostomatidae Cyclothone alba Oceanic 0.095 0.144 0.071
Gonostomatidae Cyclothone braueri Oceanic 0.066 0.063 0.043
Gonostomatidae Cyclothone pallida Oceanic 0.113 0.052 0.029
Gonostomatidae Cyclothone pseudopallida Oceanic 0.038 0.06 0.036
Gonostomatidae Cyclothone spp. Oceanic 0.053 0.092 0.043
Myctophidae Diaphus mollis Oceanic 0.017 0.012 0.008
Myctophidae Diaphus spp. Oceanic 0.119 0.158 0.047
Myctophidae Diogenichthys atlanticus Oceanic 0.026 0.043 0.047
Myctophidae Electrona rissoi Oceanic 0.011 0.063 0.041
Gonostomatidae Gonostoma atlanticum Oceanic 0.033 0.071 0.056
Melamphaidae Melampaes simus Oceanic 0.03 0.006 0.035
Myctophidae Myctophum obtusirostre Oceanic 0.042 0.089 0.013
Myctophidae Myctophum spp. Oceanic 0.035 0.02 0.008
Myctophidae Notolychnus valdiviae Oceanic 0.028 0.106 0.138
Gonostomatidae Sigmops elongatum Oceanic 0.059 0.025 0.024
Sternoptychidae Sternoptyx diaphana Oceanic 0.039 0.089 0.063
Sternoptychidae Sternoptyx pseudobscura Oceanic 0.03 0.142 0.155
COMMON TAXA IN NORTH AND WEST SUB-REGIONS  
Family Taxa Habitat NOR WOR  
Bothidae Bothus ocellatus Neritic 0.015 0.007  
Tetraodontidae Canthigaster spp. Neritic 0.005 0.025  
Scaridae Scarus spp. Neritic 0.014 0.015  
Trichiuridae Aphanopus intermedius Oceanic 0.006 0.004  
Sternoptychidae Argyropelecus hemigymnus Oceanic 0.011 0.022  
Gonostomatidae Gonosthoma spp. Oceanic 0.012 0.048  
Myctophidae Myctophum asperum Oceanic 0.006 0.031  
Alepisauridae Omosudis lowii Oceanic 0.021 0.004  
COMMON TAXA IN EAST AND WEST SUB-REGIONS  
Family Taxa Habitat EOR WOR  
Callionymidae Callionymus bairdi Neritic 0.021 0.008  
Synaphobranchidae Dysomma anguillare Neritic 0.05 0.01  
Scorpaenidae Pontinus rathbuni Neritic 0.006 0.003  
Rachycentridae Rachycentrum canadum Neritic 0.023 0.022  
Scorpaenidae Scorpaena spp. Neritic 0.006 0.008  
Sternoptychidae Argyropelecus affinis Oceanic 0.003 0.004  
Gonostomatidae Bonapartia pedaliota Oceanic 0.014 0.006  
Bregmacerotidae Bregmaceros spp. Oceanic 0.06 0.014  
Myctophidae Hygophum taaningi Oceanic 0.016 0.014  
Myctophidae Lampadena spp. Oceanic 0.007 0.004  
Paralepididae Lestidiops affinis Oceanic 0.002 0.006  
Bothidae Monolene sessilicauda Oceanic 0.007 0.005  
Phosichthyidae Pollicththys mauli Oceanic 0.011 0.003  
Scopelarchidae Scopelarchus spp. Oceanic 0.018 0.008  
COMMON TAXA IN NORTH AND EAST SUB-REGIONS  
Family Taxa Habitat NOR EOR  
Apogonidae Apogon spp. Neritic 0.025 0.091  
Bothidae Bothus spp. Neritic 0.01 0.021  
Mugilidae Mugil curema Neritic 0.004 0.019  
Serranidae Serranus spp. Neritic 0.03 0.021  
Myctophidae Ceratoscospelus warmingii Oceanic 0.061 0.007  
Gonostomatidae Cyclothone obscura Oceanic 0.005 0.005  
Gempylidae Diplospinus multistriatus Oceanic 0.01 0.043  
Linophrynidae Haplophryne mollis Oceanic 0.004 0.003  
Myctophidae Lampanyctus nobilis Oceanic 0.005 0.007  
Myctophidae Myctophum affine Oceanic 0.02 0.016  
Myctophidae Myctophum nitidulum Oceanic 0.025 0.044  
Notosudidae Scopelosaurus mauli Oceanic 0.021 0.008  
Myctophidae Symbolophorus rufinus Oceanic 0.004 0.012  
Acropomatidae Synagrops bellus Oceanic 0.004 0.004  
Phosichthyidae Vinciguerria nimbaria Oceanic 0.005 0.011  

Values = average density. NOR = North Oceanic sub-region, EOR = East Oceanic sub-region, WOR = West Oceanic sub-region.

Table 2: Exclusive taxa in each oceanic sub-region in Campeche Bay, Gulf of Mexico, during winter cruise (January 24 to February 3, 2013).

EXCLUSIVE TAXA IN EAST OCEANIC SUB-REGION
Family Taxa Habitat Average density
Mugilidae Agonostoma moticula Neritic 0.002
Serranidae Anthias nicholsi Neritic 0.013
Labridae Bodianus rufus Neritic 0.007
Paralichthydae Citharichthys cornutus Neritic 0.011
Paralichthydae Citharichthys gymnorhinus Neritic 0.004
Paralichthydae Citharichthys macrops Neritic 0.006
Paralichthydae Citharichthys spp. Neritic 0.003
Paralichthydae Citharichthys spilopterus Neritic 0.014
Carangidae Decapterus punctatus Neritic 0.025
Bothidae Engyophrys senta Neritic 0.004
Paralichthydae Etropus microstomus Neritic 0.016
Labridae Halichoeres cyanocephalus Neritic 0.007
Sciaenidae Micropogonias furnieri Neritic 0.013
Ophichthidae Myrophis punctatus Neritic 0.004
Ophichthidae Ophichthus spp. Neritic 0.004
Pomatomidae Pomatomus saltatrix Neritic 0.006
Scorpaemidae scorpaena plumieri Neritic 0.002
Sparidae Sparisoma spp. Neritic 0.017
Paralichthyidae Syacium papillosum Neritic 0.005
Cynoglosidae Symphurus spp. Neritic 0.006
Synodontidae Synodus foetens Neritic 0.011
Synodontidae Synodus spp. Neritic 0.014
Alepisauridae Alepisaurus ferox Oceanic 0.004
Percophidae Bembrops gobioides Oceanic 0.007
Percophidae Bembrops spp. Oceanic 0.007
Trichiuridae Benthodesmus tenuis Oceanic 0.006
Bregmacerotidae Bregmaceros houdei Oceanic 0.007
Ophidiidae Brotula spp. Oceanic 0.016
Gonostomatidae Cyclothone microdon Oceanic 0.018
Myctophidae Diaphus brachycephalus Oceanic 0.011
Directmidae Diretmichthys parini Oceanic 0.005
Microstomatidae Dolicholagus longirostris Oceanic 0.007
Stomiidae Eustomias spp. Oceanic 0.006
Evermanellidae Evermannella melanoderma Oceanic 0.007
Congridae Gnathophis spp. Oceanic 0.007
Myctophidae Hygophum benoiti Oceanic 0.003
Myctophidae Hygophum spp. Oceanic 0.045
Phosichthyidae Icthyococus ovatus Oceanic 0.006
Myctophidae Lampanyctus spp. Oceanic 0.004
Gempylidae Lepidocybium flavobrunneum Oceanic 0.004
Trichiuridae Lepidopus altifrons Oceanic 0.005
Myctophidae Myctophum selenops Oceanic 0.006
Myctophidae Nannobrachium spp. Oceanic 0.015
Gempylidae Nealotus tripes Oceanic 0.014
Polynemidae Polydactilus spp. Oceanic 0.006
Family Taxa Habitat Averaje density
Sternoptychidae Polyipnus spp. Oceanic 0.004
Scorpaenidae Pterois spp. Oceanic 0.006
Paralepididae Sudis atrox Oceanic 0.007
Congridae Uroconger syringinus Oceanic 0.007
Sternoptychidae Valenciennellus tripunctulatus Oceanic 0.016
Phosichthyidae Vinciguerria attenuata Oceanic 0.011
Phosichthyidae Vinciguerria poweriae Oceanic 0.003
EXCLUSIVE TAXA NORTH OCEANIC SUB-REGION
Family Taxa Habitat Average density
Diodontidae Chilomycterus schoepfi Neritic 0.004
Priacanthidae Heteropriacanthus cruentatus Neritic 0.02
Malacantidae Malacanthus plumieri Neritic 0.017
Synodontidae Synodus Neritic 0.013
Sternoptychidae Argyropelecus acuelatus Oceanic 0.007
Myctophidae Ceratoscospelus spp. Oceanic 0.043
Bothidae Chascanopsettalugubris Oceanic 0.005
Myctophidae Diaphus effulgens Oceanic 0.007
Ophidiidae Dicrolene spp. Oceanic 0.007
Gonostomatidae Margrethia obtusirostre Oceanic 0.011
Myctophidae Nannobrachium atrum Oceanic 0.015
Myctophidae Notoscopelus caudispinosus Oceanic 0.006
Gonostomatidae Sigmops spp. Oceanic 0.032
Paralepididae Stemonosudis rothschildi Oceanic 0.011
Cynoglosidae symphuruspiger Oceanic 0.01
Phosichthyidae Vinciguerria spp. Oceanic 0.025
EXCLUSIVE TAXA WEST OCEANIC SUB-REGION
Family Taxa Habitat Average density
Tetraodontidae Canthigaster rostrata Neritic 0.003
Mullidae Mulloidichthys martinicus Neritic 0.013
Nettastomidae Nettenchelys pygmaea Neritic 0.013
Sphyraenidae Sphyraena spp. Neritic 0.013
Microstomatidae Bathylagos spp. Oceanic 0.008
Berycidae Beryx spp. Oceanic 0.01
Stomiidae Chauliodus sloani Oceanic 0.005
Chiasmodontidae Chiasmodon spp. Oceanic 0.008
Nomeidae Cubiceps pauciradiatus Oceanic 0.008
Gempylidae Gempylus serpens Oceanic 0.005
Myctophidae Hygophum reinhardtii Oceanic 0.013
Myctophidae Lampanyctus alatus Oceanic 0.017
Myctophidae Lepidophanes gaussi Oceanic 0.006
Myctophidae Lepidophanes guentheri Oceanic 0.005
Melanocetidae Melanocetus johnsoni Oceanic 0.006
Microstomatidae Melanolagus bericoides Oceanic 0.019
Myctophidae Nannobrachium lineatun Oceanic 0.013
Myctophidae Notoscopelus resplendens Oceanic 0.008
Alepisauridae Omosudis spp. Oceanic 0.003
Nomeidae Psenes pellucidus Oceanic 0.01
Scopelarchidae Scopelarchus analis Oceanic 0.008
Phycidae Urophycis spp. Oceanic 0.013

Table 3: Common taxa among oceanic sub-regions in Campeche Bay, Gulf of Mexico, during summer cruise (4-14 June 2014).

COMMON TAXA IN THREE SUB-REGIONS
Family Taxa Habitat NOR EOR WOR
Bothidae Bothus ocellatus Neritic 0.425 0.781 0.433
Sternoptychidae Maurolicus weitzmani Neritic 0.206 0.318 0.645
Paralichthydae Syacium papillosum Neritic 0.579 1.499 0.705
Sternoptychidae Argyropelecus affinis Oceanic 0.471 0.201 0.304
Sternoptychidae Argyropelecus hemigymnus Oceanic 0.208 0.281 0.171
Sternoptychidae Argyropelecus sladeni Oceanic 0.289 0.369 0.639
Sternoptychidae Argyropelecus spp. Oceanic 0.342 0.423 0.686
Myctophidae Benthosema suborbitale Oceanic 0.506 0.77 0.692
Gonostomatidae Bonapartia pedaliota Oceanic 0.222 0.227 0.154
Bregmacerotidae Bregmaceros atlanticus Oceanic 0.91 1.016 0.76
Bregmacerotidae Bregmaceros cantori Oceanic 1.132 0.589 1.325
Myctophidae Centrobranchus nigroocellatus Oceanic 0.282 0.128 0.143
Myctophidae Ceratoscopelus spp. Oceanic 0.227 0.377 0.638
Gonostomatidae Cyclothone acclinidens Oceanic 0.335 0.598 0.426
Gonostomatidae Cyclothone alba Oceanic 0.403 0.191 0.595
Gonostomatidae Cyclothone braueri Oceanic 0.235 0.388 0.555
Gonostomatidae Cyclothone microdon Oceanic 0.38 0.569 0.286
Gonostomatidae Cyclothone pallida Oceanic 0.255 0.334 0.595
Gonostomatidae Cyclothone pseudopallida Oceanic 0.229 0.418 0.207
Gonostomatidae Cyclothone spp. Oceanic 0.547 0.53 0.535
Myctophidae Diaphus mollis Oceanic 0.482 0.893 0.367
Myctophidae Diaphus spp. Oceanic 0.843 1.264 0.844
Myctophidae Diogenichthys atlanticus Oceanic 0.803 0.636 1.962
Gempylidae Diplospinus multistriatus Oceanic 0.576 0.293 0.276
Microstomatidae Dolicholagus longirostris Oceanic 0.129 0.239 0.255
Myctophidae Electrona risso Oceanic 0.27 0.316 0.113
Gonostomatidae Gonostoma atlanticum Oceanic 0.395 0.443 0.655
Myctophidae Hygophum macrochir Oceanic 0.184 0.341 0.472
Myctophidae Hygophum reinhardtii Oceanic 0.355 0.759 0.637
Myctophidae Hygophum taaningi Oceanic 0.39 0.529 0.558
Myctophidae Lampadena luminosa Oceanic 0.351 0.712 0.402
Myctophidae Lampanyctus alatus Oceanic 0.16 0.756 0.269
Myctophidae Lampanyctus spp. Oceanic 0.351 0.325 0.223
Myctophidae Lepidophanes guentheri Oceanic 0.248 0.128 0.173
Melamphaidae Melamphaes simus Oceanic 0.421 0.562 0.463
Myctophidae Myctophum affine Oceanic 0.328 0.21 0.154
Myctophidae Myctophum asperum Oceanic 0.55 0.518 0.414
Myctophidae Myctophum nitidulum Oceanic 0.361 0.606 0.504
Myctophidae Myctophum obtusirostre Oceanic 0.213 0.544 0.836
Myctophidae Myctophum spp. Oceanic 0.59 0.363 0.559
Myctophidae Notolychnus valdiviae Oceanic 0.88 1.022 0.549
Gonostomatidae Sigmops elongatum Oceanic 0.464 0.902 0.405
Sternoptychidae Sternoptyx diaphana Oceanic 0.335 0.354 0.346
Sternoptychidae Sternoptyx spp. Oceanic 0.333 0.489 0.82
Myctophidae Symbolophorus rufinus Oceanic 0.303 0.293 0.719
Sternoptychidae Valenciennellus tripunctulatus Oceanic 0.329 0.415 0.367
Phosichthyidae Vinciguerria spp Oceanic 0.701 0.34 0.462
COMMON TAXA IN NORTH AND WEST SUB-REGIONS
Family Taxa Habitat NOR WOR  
Exocoetidae Cheilopogon spp. Neritic 0.294 0.679  
Myctophidae Ceratoscopelus warmingii Oceanic 0.65 0.154  
Myctophidae Gonichthys cocco Oceanic 0.229 0.424  
Myctophidae Myctophum selenops Oceanic 0.467 0.272  
Myctophidae Nannobrachium lineatum Oceanic 0.227 0.547  
Myctophidae Nannobrachium spp. Oceanic 0.336 0.594  
Scopelarchidae Scopelarchus spp. Oceanic 0.217 0.154  
Melamphaidae Scopeloberyx spp. Oceanic 0.984 0.173  
Phosichthyidae Vinciguerria poweriae Oceanic 0.183 0.36  
COMMON TAXA IN EAST AND WEST SUB-REGIONS
Family Taxa Habitat EOR WOR  
Microdesmidae Microdesmus lanceolatus Neritic 1.428 0.924  
Congridae Rhynchoconger flavus Neritic 0.285 0.34  
Stomiidae Chauliodus danae Oceanic 0.358 0.499  
Myctophidae Hygophum benoiti Oceanic 0.358 0.164  
Myctophidae Lampanyctus nobilis Oceanic 0.425 0.132  
Myctophidae Notoscopelus spp. Oceanic 0.719 0.376  
Paralepididae Stemonosudis rothschildi Oceanic 0.334 0.173  
Phosichthyidae Vinciguerria attenuata Oceanic 0.734 0.154  
COMMON TAXA IN NORTH AND EAST SUB-REGIONS
Family Taxa Habitat NOR EOR  
Scombridae Auxis thazard Neritic 0.345 0.81  
Carangidae Caranx spp Neritic 0.155 0.332  
Paralichthydae Citharichthys spp. Neritic 0.259 0.152  
Diodontidae Diodon spp. Neritic 0.129 0.274  
Synaphobranchidae Dysomma anguillare Neritic 0.314 0.393  
Scombridae Euthynnus alletteratus Neritic 0.223 0.945  
Howellidae Howella spp. Neritic 0.416 0.369  
Carangidae Selene setapinnis Neritic 0.187 1.735  
Carangidae Selene spp. Neritic 0.323 0.184  
Serranidae Serranus spp. Neritic 0.189 0.573  
Scombridae Thunnus spp Neritic 0.232 0.373  
Bregmacerotidae Bregmaceros spp Oceanic 0.227 0.663  
Stomiidae Chauliodus spp. Oceanic 0.128 0.231  
Coryphaenidae Coryphaena spp. Oceanic 0.259 0.376  
Myctophidae Hygophum hygomii Oceanic 0.195 0.525  
Myctophidae Hygophum spp. Oceanic 0.253 0.673  
Phosichthyidae Ichthyococcus ovatus Oceanic 0.288 0.7  
Myctophidae Lampadena spp. Oceanic 0.187 0.421  
Paralepididae Lestidiops affinis Oceanic 0.339 0.611  
Paralepididae Lestidiops spp. Oceanic 0.642 0.355  
Myctophidae Lobianchia gemellarii Oceanic 0.259 0.661  
Gonostomatidae Margrethia obtusirostre Oceanic 0.256 0.247  
Myctophidae Nannobrachium atrum Oceanic 0.289 0.381  
Alepisauridae Omosudis lowii Oceanic 0.972 0.837  
Chlorophthalmidae Parasudis truculentus Oceanic 0.32 0.332  
Sternoptychidae Polyipnus spp. Oceanic 0.146 0.917  
Scopelarchidae Scopelarchus analis Oceanic 0.266 0.321  
Scopelarchidae Scopelarchus michaelsarsi Oceanic 0.185 0.373  
Sternoptychidae Sternoptyx pseudobscura Oceanic 0.47 0.411  
Paralepididae Sudis atrox Oceanic 0.457 0.258  

Values = average density. NOR = North Oceanic sub-region, EOR = East Oceanic sub-region, WOR = West Oceanic sub-region.

Table 4: Exclusive taxa in each oceanic sub-region in Campeche Bay, Gulf of Mexico during summer cruise (4-14 June 2014).

EXCLUSIVE TAXA IN EAST OCEANIC SUB-REGION
Family Taxa Habitat Average density
Ophichthidae Ahlia egmontis Neritic 0.332
Monacanthidae Aluterus scriptus Neritic 0.296
Antennariidae Antennarius spp Neritic 0.369
Serranidae Anthias woodsi Neritic 0.332
Scombridae Auxis spp Neritic 0.853
Balistidae Balistes capriscus Neritic 0.358
Tetraodontidae Canthigaster spp Neritic 0.425
Carangidae Caranx crysos Neritic 0.358
Paralichthydae Citharichthys arctifrons Neritic 0.126
Paralichthydae Etropus crossotus Neritic 0.332
Paralichthydae Etropus microstomus Neritic 0.332
Gerreidae Eucinostomus spp. Neritic 0.622
Fistulariidae Fistularia spp. Neritic 0.917
Lutjanidae Lutjanus campechanus Neritic 0.425
Lutjanidae Lutjanus spp. Neritic 1.712
Microdesmidae Microdesmus longipinnis Neritic 0.88
Microdesmidae Microdesmus spp. Neritic 0.136
Carangidae Oligoplites saurus Neritic 0.358
Ophichthidae Ophichthus gomesii Neritic 0.358
Ophichthidae Ophichthus spp. Neritic 0.425
Ophidiidae Ophidion nocomis Neritic 0.258
Rachycentridae Rachycentron canadum Neritic 0.332
Scombridae Scomberomorus regalis Neritic 0.229
Scorpaenidae Scorpaena spp. Neritic 1.069
Carangidae Selar crumenophtalmus Neritic 0.167
Carangidae Selene vomer Neritic 0.373
Sphyraenidae Sphyraena guachancho Neritic 1.712
Scombridae Thunnus atlanticus Neritic 1.428
Alepisauridae Alepisaurus spp Oceanic 0.837
Aulopidae Aulopus nanae Oceanic 0.378
Percophidae Bembrops spp Oceanic 0.185
Bregmacerotidae Bregmaceros maclellandii Oceanic 0.185
Chiasmodontidae Chiasmodon niger Oceanic 0.242
Nomeidae Cubiceps pauciradiatus Oceanic 0.332
Gonostomatidae Cyclothone obscura Oceanic 0.56
Stomiidae Eustomias spp. Oceanic 0.358
Nettastomatidae Hoplunnis tenuis Oceanic 0.35
Melamphaidae Melamphaes spp. Oceanic 0.544
Gempylidae Nealotus tripes Oceanic 0.285
Serranidae Pronotogrammus aureorubens Oceanic 0.425
Nomeidae Psenes spp. Oceanic 0.332
Scopelarchidae Scopelarchoides danae Oceanic 0.369
Phosichthyidae Vinciguerria nimbaria Oceanic 0.285
EXCLUSIVE TAXA WEST OCEANIC SUB-REGION
Family Taxa Habitat Average density
Stomiidae Chauliodus sloani Oceanic 0.127
Myctophidae Diaphus brachycephalus Oceanic 0.238
Gonostomatidae Gonostoma spp. Oceanic 0.169
Microstomatidae Melanolagus bericoides Oceanic 0.388
Stomiidae Melanostomias spp. Oceanic 0.34
Stomiidae Photostomias guernei Oceanic 0.499
Scorpelarchidae Scopelarchus guentheri Oceanic 0.216
Dactylopteridae Dactylopterus volitans Neritic 0.34
Mirapinnidae Eutaeniophorus festivus Neritic 0.16
Scombridae Katsuwonus pelamis Neritic 0.679
Syngnathidae Syngnathus louisianae Neritic 0.499
EXCLUSIVE TAXA NORTH OCEANIC SUB-REGION
Family Taxa Habitat Average density
Paralichthyidae Cyclopsetta fimbriata Neritic 0.195
Bothidae Engyophrys senta Neritic 0.172
Muraenidae Gymnothorax ocellatus Neritic 0.259
Nettastomatidae Nettenchelys pygmaea Neritic 0.294
Pomacentridae Stegastes spp. Neritic 0.243
Alepisauridae Alepisaurus brevirostris Oceanic 0.146
Caproidae Antigonia capros Oceanic 0.195
Chlorophthalmidae Chlorophthalmus agassizi Oceanic 0.2
Coryphaenidae Coryphaena equiselis Oceanic 0.328
Opisthoproctidae Dolichopteryx binocularis Oceanic 0.227
Evermannellidae Evermanella balbo Oceanic 0.294
Nettastomatidae Facciolella spp. Oceanic 0.259
Ipnopidae Ipnops murrayi Oceanic 0.745
Paralichthyidae Cyclopsetta fimbriata Neritic 0.195
Myctophidae Lepidophanes gaussi Oceanic 0.452
Trichiuridae Lepidopus spp. Oceanic 0.195
Stomiidae Leptostomias spp. Oceanic 0.984
Macrouridae Mesobius spp. Oceanic 0.16
Myctophidae Nannobrachium cuprarium Oceanic 0.289
Myctophidae Notoscopelus caudispinosus Oceanic 0.648
Paralepididae Paralepis spp. Oceanic 0.119
Nomeidae Psenes arafurensis Oceanic 0.147
Paralepididae Uncisudis advena Oceanic 0.294
Paralepididae Uncisudis spp Oceanic 0.195

The taxa composition in both cruises was similar with around 70% of larvae from an oceanic stock and 30% of the neritic stock. The larval density in summer represents 66.8% of the total of both cruises, and the density of neritic larvae was 13.1% in winter and 26.5% in summer.

The more abundant and frequent species in both cruises were Diogenichthys atlanticus, Notolychnus valdiviae, Benthosema suborbitale, Bregmaceros cantori, B. atlanticus, Gonostoma atlanticum, Sternoptyx diaphana and the genera Diaphus, Cyclothone y Argyropelecus. Of the neritic larvae, only Syacium papillosum and Apogon sp. appear among the twenty-first more abundant.

The composition of the larval community was analyzed with Bray-Curtis dissimilarity index using the total taxa or only neritic larvae. However, the resulting stations groups did not correspond with the oceanic sub-region determined by temperature and salinity.

Considering, firstly that from the 236 total taxa, 151 and 85 were oceanic and neritic larvae, respectively and secondly, that there was no correspondence between the ocean sub-regions and stations groups, we analyzed the differences among oceanic sub-regions, considering common taxa in the three regions, as well as those occurring in two or only one (Tables 1,2,3 and 4).

The larger number of common taxa among three sub-regions was oceanic: 24 and 47 in winter and summer, respectively, with only three and five neritic larvae in each cruise. The number of common taxa among two sub-regions was lower. The low number of common taxa among the WOR with the other two, as well as its low number of neritic taxa, making it different. Therefore, the greatest similarity was among the NOR and EOR particularly in the summer (Table 5).

Table 5: Number of common and exclusive, neritic and oceanic taxa, occurring in each oceanic sub-region and season.

WINTER   SUMMER
Taxa Oceanic Neritic   Oceanic Neritic
109 45   117 54
Total 154   171
WINTER   SUMMER
Common Taxa N-W N-E W-E ALL   Common Taxa N-W N-E W-E ALL
Oceanic 5 11 9 24   Oceanic 8 19 6 47
Neritic 3 4 5 3   Neritic 1 11 2 5
Total 8 15 14 27   Total 9 30 8 52
WINTER   SUMMER
Exclusive Taxa North East West     Exclusive Taxa North East West  
Oceanic 12 30 18     Oceanic 18 15 7  
Neritic 4 22 4     Neritic 5 28 4  
Total 16 52 22     Total 23 43 11  

N = North Oceanic subregion, E = East Oceanic sub-region, W = West Oceanic sub-region.

The large difference among sub-regions was attributed to a few neritic taxa occurring exclusively in some of them, those, in winter there were only four in the NOR and WOR and 22 in the EOR. Similarly, in summer cruise four and five neritic taxa occurred exclusively in the NOR and WOR, and 28 in the EOR (Table 5). The greatest diversity and frequency of neritic taxa corresponded to the EOR.

Vertical distribution

Distribution in the water column of the neritic and oceanic larvae was similar in winter and summer seasons; both groups of larvae presented greater density in the epipelagic layer, strongly declining towards the next level (200-400m); the density continued descending up to 1000 m, except in the summer cruise in which after 800m there was an increase to the next level (Figure 5).

Figure 5: Percentage of larvae density by depth levels.

The transport of larvae from the neritic to oceanic zones let us consider that this kind of larvae could remain in the epipelagic layer, but 31 from 85 neritic taxa occurred in the layers of 600 to 1000m depth.

Most of these taxa were represented by one organism, except Auxis thazard, Syacium papillosum and Bothus ocellatus, occurring several times in such depths (Table 6).

Table 6: Taxa present once.

WINTER   SUMMER
Neritic Oceanic   Neritic Oceanic
Agonostoma moticula Alepisaurus ferox   Ahlia egmontis Alepisaurus brevirostris
Anthias nicholsi Argyropelecus acuelatus   Antennarius spp. Alepisaurus spp.
Bodianus rufus Bathylagos spp.   Anthias woodsi Antigonia capros
Canthigaster rostrata Bembrops gobioides   Auxis spp. Bembrops spp.
Chilomycterus schoepfi Bembrops spp.   Balistes capriscus Bregmaceros maclellandii
Citharichthys gymnorhinus Benthodesmus tenuis   Canthigaster spp. Chauliodus sloani
Citharichthys macros Beryx spp.   Caranx crysos Chiasmodon niger
Citharichthys spp. Ceratoscopelus spp.   Citharichthys arctifrons Chlorophthalmus agassizi
Engyophrys senta Chascanopsetta lugubris   Cyclopsetta fimbriata Coryphaena equiselis
Halichoeres cyanocephalus Chauliodus sloani   Dactylopterus volitans Cubiceps pauciradiatus
Heteropriacanthus cruentatus Chiasmodon spp.   Engyophrys senta Diaphus brachycephalus
Micropogonias furnieri Cubiceps pauciradiatus   Etropus crossotus Dolichopteryx binocularis
Mulloidichthys martinicus Diaphus effulgens   Etropus microstomus Eustomias spp.
Myrophis punctatus Dicrolene spp.   Eutaeniophorus festivus Evermannella balbo
Nettenchelys pygmaea Diretmichthys parini   Fistularia spp. Facciolella spp.
Ophichthus spp. Dolicholagus longirostris   Gymnothorax ocellatus Gonostoma spp.
Pomatomus saltatrix Eustomias spp.   Katsuwonus pelamis Hoplunnis tenuis
Scorpaena plumieri Evermannella melanoderma   Lutjanus campechanus Ipnops murrayi
Sphyraena spp. Gempylus serpens   Lutjanus spp. Lepidopus spp.
Syacium papillosum Gnathophis spp.   Microdesmus spp. Leptostomias spp.
Symphurus spp. Hygophum benoiti   Nettenchelys pygmaea Melanolagusbericoides
Synodus spp. Hygophum reinhardtii   Oligoplites saurus Melanostomias spp.
Synodus synodus Ichthyococcus ovatus   Ophichthus gomesii Mesobius spp.
  Lampanyctus spp.   Ophichthus spp. Nannobrachium cuprarium
  Lepidocybium flavobrunneum   Rachycentron canadum Nealotus tripes
  Lepidophanes gaussi   Scomberomorus regalis Paralepis spp.
  Lepidophanes guentheri   Selar crumenophtalmus Photostomias guernei
  Lepidopus altifrons   Selene vomer Pronotogrammus aureorubens
  Margrethia obtusirostre   Sphyraena guachancho Psenes arafurensis
  Melanocetus johnsoni   Stegastes spp. Psenes spp.
  Myctophum selenops   Syngnathus louisianae Scopelarchoides danae
  Nannobrachium atrum   Thunnus atlanticus Scopelarchus guentheri
  Nannobrachium lineatum     Uncisudis advena
  Nannobrachium spp.     Uncisudis spp.
  Notoscopelus caudispinosus     Vinciguerria nimbaria
  Notoscopelus resplendens      
  Omosudis spp.      
  Polydactilus spp.      
  Polyipnus spp.      
  Psenes pellucidus      
  Pterois spp.      
  Scopelarchus analis      
  Sigmops spp.      
  Stemonosudis rothschildi      
  Sudisatrox      
  Symphurus piger      
  Uroconger syringinus      
  Urophycis spp.      
  Vinciguerria poweriae      
  Vinciguerria spp.      

Of the total taxa, 59% occurred in only once, 85 oceanic and 55 neritic. This means that more than a half of neritic taxa were represented by a single organism and most of them were in the epipelagic layer, mainly in summer.

Discussion

Oceanic sub-regions

Differences in temperature and to a lesser extent of salinity allowed the recognition of three oceanic sub-regions in the Bay of Campeche; in the two seasonal periods studied, these subdivision seem promoted by an almost permanent phenomenon, generated by two hydrographic facts: One is the semi-persistent cyclonic gyre which occurs in the bay [20,21]; and the other, the intrusion of neritic waters from the YSC that flow over the Bay [15,22].

The station groups resulting from the application of the similarity index [19] did not correspond to the oceanic regions determined from temperature and salinity; the differences in temperature and salinity are probably not so severe as to limit the passage of the larvae of a region to another. Richards et al., [16] in their study at the border of the Loop Current, had similar results applying a cluster analysis that might not support their hypothesis of front assemblages; however, in their study of families, they did distinguish coastal and oceanic groups.

The hydrodynamics of Campeche Bay is consistent with the hydrographic sub-regions described in our results. Thus, the WOR is located where the cyclonic gyre takes place [20,21]. It is, the region with the lower temperatures and salinities caused by the upwelling of deep water; while the NOR, which had warmer temperatures and high salinity are formed in the area under the direct influence of the YSC with the east-west direction [15,22,23]. The EOR, on the other hand, with average values of temperatures and salinities strictly intermediate is also influenced by the same current, after flowing through the Campeche-Tabasco shelf.

Neritic and oceanic larvae in the oceanic sub-regions

In the pelagic zone of Campeche Bay predominated larvae from oceanic parents, epipelagic, mesopelagic, bathypelagic or demersal; the families Myctophidae, Gonostomatidae, Sternoptychidae, Bregmacerotidae regularly were the most abundant and frequent [2,18].

The distribution of oceanic species did not appear limited by temperature or salinity differences among the three recorded oceanic sub-regions.

The low densities of neritic larvae 13.1% in winter and 26.5% in summer shows the small influence of neritic communities on the oceanic area as it has already been mentioned by Flores-Coto et al., [11]. This is consistent with its neritic origin [6-10] and their dispersion generated by the warm current that flows over the Yucatan shelf which reaches the oceanic zone.

The higher neritic larvae concentration on the NOR and EOR concerning the WOR fits the trajectories of the surface currents described by Zavala-Sansón et al., [24].

The larval distribution of neritic species confirms the low connectivity that exists between Yucatan and Veracruz reefs [25]. That is to say, those planktonic organisms transported by YSC towards Campeche Bay, do not to reach the Western area off Veracruz. However, possibly some of the neritic larvae identified come from the Veracruz reefs, particularly in the most western stations, since many species of this work also have been recorded in this reef area [26-28].

The hydrodynamics of Campeche Bay explains the differences in the proportion of neritic and oceanic larvae among the three oceanic sub-regions.

In the WOR there were a low number of neritic larvae because it was the area with less influence of the neritic communities swept away by YSC; and also, because in this sub-region the most important physical feature is the cyclonic gyre which, closely occupies the center of the Campeche Bay and generates a boundary in its outer limit for the other two regions [22].

In contrast, the EOR presented the greatest number of neritic larvae, due to the significant influence of the communities coming from the Yucatan shelf, particularly of taxa whose parents are linked to the existing local reefs [26].

On the other hand, in addition to the low number of neritic larvae in the WOR and the low number of common taxa shared with the other two sub-regions, it demonstrated its significant difference with other two sub-regions. In contrast, NOR and EOR were more similar.

In the studied area, larval fish distribution, as observed in another world environment, depend on the parents habit mainly the spawning season and areas and larvae concentration or dispersion by the hydrodynamic regime [4-10,29,30].

Vertical distribution

The high density of larvae both oceanic and neritic in the epipelagic zone seems to be attributed to the increased availability of food in that layer, decreasing to the next depthlevels [31,33]. The increase in density of larvae between 800 and 1000m recorded during the summer cruise corresponded to juvenile and adult mesopelagic and bathypelagic species. However, the increase in larvae of neritic species has no explanation.

On the other hand, the concentration the larvae of neritic species in the surface layer of the water column, usually at depths less than 50 m is a common feature. However, the presence at depths of 600 1000 m larvae from neritic species, coming from areas with depths less than 200 m, suggests a vertical migration process by the larvae, not strictly to a turbulence mechanism. These considerations are based on the fact that 31 of the 85 neritic taxa, more one-third, were at greater depths, in all sub-regions during the two cruises, except in the summer when there were no neritic taxa at those depths in the WOR.

The presence of species represented by a single specimen in the oceanic waters seems a common fact. Richards et al., [18] reported 21.8%, and we recorded 59% in our study: 55 neritic and 85 oceanic taxa.

Many neritic species form schools as adults and larvae, as part of their life strategy, including reproduction and spawning; the shoals of larvae swept away by currents are highly dispersed in oceanic waters. Therefore, the capture of a single specimen of a species in the oceanic area could be considered normal. However, the presence of an individual of an oceanic species, particularly mesopelagic must obey to other causes, such as that not all species form dense schools during the reproduction process, or by a high mortality rate of larvae by inanition or predation.

The presence of the same three oceanic sub-regions in the Campeche Bay, in two different seasonal periods, winter and summer, recorded here, underlines the high relevance of its hydrographic regime upon zooplankton communities. We conclude that the distribution of taxa is not limited by salinity or temperature differences between the ocean’s three sub-regions. However, the distribution of neritic larvae is determined by the mesoscale hydrographic stressors that characterize the area, mainly the YSC and the cyclonic gyre.

The presence of neritic larvae in deep layers seems to obey to a vertical migration process, rather than to an advection mechanism.

Acknowledgment

The authors express their gratitude to the Universidad Nacional Autónoma de México by support the cruises ZOOMEP I and II.

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Received: December 11, 2017

Accepted: January 04, 2018

Published: January 08, 2018