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09-Arceo-Fish assemb.. - Organization for Tropical Studies

1. 89 103 March June 2009 91 highest species richness was recorded 17 67 and the lowest in site 2 10 67 As to even ness the highest corresponded to site 7 0 74 and the lowest in site 1 0 52 On temporal basis higher diversity was registered during norths 2 66 0 69 bits ind while the lowest was during dry season 2 22 0 84 bits ind Likewise the highest species richness happened TABLE 1 Results of ANOVA analysis to test significant differences in environmental variables temperature dissolved oxygen salinity and for some ecological parameters of fish community diversity species richness evenness density and biomass after their transformation log x 1 Significant differences p lt 0 05 Variable analysis result Spatial n 8 Temporal n 3 during norths 17 88 and the lowest during Temperature rains 10 63 and for evenness rainy season F es Pere had the highest 0 68 and the lowest in dry aaa i i Dissolved oxygen season 0 62 F 0 5549 0 6717 p 0 7810 0 5214 Ecological Category Dominating fish Salinity with abundance higher than 70 in each a as Boa of the climate seasons were classified as ee f f f Diversity Shannon H marine euryhaline Among these we found F 0 6190 0 8737 A mitchilli A hepsetus E argenteus E P 0 7330 0 4320 gula Lagodon rhomboides Harengula jag No species daio yasi uana S testudineus and M trichodon Respect 4 0 8777 0 0531 to marine stenohaline c
2. or sometimes to lay eggs F persimilis According to Simier Anchoa hepsetus 3 167 0 408 1 188 et al 2004 the high diversity registered can 1 084 0 572 2 500 be caused by 1 establishment in the estuary of many marine species due to the permanent communication with the ocean and the pres Oligoplites saurus 3 088 0 648 1 064 ence of various freshwater seeps and 2 het erogeneous habitats seagrass meadows prop root mangroves mudflats underground water Opisthonema oglinum 4 381 0 236 1 098 flowerings favoring colonization by different fish species The ichthyofauna in Bocas de Dzilam is Dasyatis americana 0 191 2 940 0 894 dominated by estuarine species characteristic Species Salinity Oxygen Temperature Cynoscion arenarius 2 898 0 096 0 759 Sphoeroides nephelus 2 214 1 199 2 445 Anchoa lamprotaenia 1 837 0 515 3 167 Mugil trichodon Trachinotus falcatus 3 167 0 408 1 188 Harengula jaguana 3 880 0 699 0 848 Ariopsis felis 2 705 0 000 3 613 100 80 60 40 Dissimilarity percentage 20 Sampling stations Fig 5 Dendrogram of dissimilarity of stations based on log x 1 transformed total abundance of fish species using the Bray Curtis index 98 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 Salinity A mitch It ae Temperature E argen PS 12 Oxygen ok polyom 24 42 Sp velif Hj ag ua N Cl 78
3. through their thermal differences but above all because of the joined effect with dissolved oxygen Marshall and Elliott 1998 Environmental variables did not show spatial gradient as in other coastal systems of Yucatan 100 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 Peninsula Rueda 2001 Vega Cendejas and Hern ndez 2004 However there is a clear seasonal pattern influenced by climatic condi tions specially during norths Species found as estuarine residents and most dominating did not show any relation with environmental vari ables but some marine species did so because of their abundance in certain climatic seasons Salinity has a great influence over distribution and abundance of some marine species like in anchovies A hepsetus and A mitchilli and herrings H jaguana and O oglinum We have to take into account that response of many species to salinity fluctuations can vary during their life cycle For instance juvenile herrings can tolerate wider ranges than adults Marshall and Elliott 1998 As for tempera ture it showed certain influence over some species such as mullets M trichodon silver croaker B chrysoura and catfish A felis It is unlikely that thermal tolerance by itself be the main factor to influence the examined species distribution However temperature s influence over fish distribution grows through effects of synergy between temperature and
4. 02 0 05 1 39 0 07 0 01 0 26 0 007 0 009 0 02 2 tH a Eg D 5 mar mar mar mar namas mar Estuar E mar E mar E mar E mar Estuar mar mar mar mar mar mar mar mar aA wv zz zzz nw RND ZZUZ 95 TABLE 2 Continued Relative abundance N and biomass W ecological category E C and seasonal occurrence S O Species Family N W 18 Ce SHO Cyprinodon artifrons Cyprinodontidae 0 05 0 001 Estuar D Chriodorus atherinoides Hemiramphidae 0 05 0 09 E mar D Menidia colei Atherinopsidae 0 05 0 002 E mar Nicholsina usta Scaridae 0 03 0 01 S mar N Gerres cinereus Gerreidae 0 03 0 02 E mar N Haemulon plumierii Haemulidae 0 03 0 02 S mar N Chaetodipterus faber Ephippidae 0 03 0 02 E mar Syngnathus louisianae Syngnathidae 0 03 0 002 E mar Dasyatis americana Dasyatidae 0 02 1 39 E mar R Anchoa lyolepis Engraulidae 0 02 0 002 E mar N Citharichtys spilopterus Paralichtyidae 0 02 0 008 E mar D Eucinostomus melanopterus Gerreidae 0 02 0 001 E mar N Syngnathus floridae Syngnathidae 0 02 0 002 E mar N Lutjanus cyanopterus Lutjanidae 0 01 0 01 E mar R Hyporhamphus meeki Hemirramphidae 0 01 0 01 E mar D Acanthostracion quadricornis Ostraciidae 0 01 0 23 E mar R Diapterus auratus Gerreidae 0 01 0 01 E mar N Eugerres plumieri Gerreidae 0 01 0 07 E mar D Gymnura micrura Gymnuridae 0 01 0 08 S mar R Bathygobius soporator Gobiidae 0 01 0 002 E mar R Ctenogobius stigmaticus Go
5. 1 2 5m with 80 of the bottom covered by macrophytes dominated mainly by Halodule wrightii and Ruppia maritima Herrera Silveira et al 1998 Medina Gomez and Herrera Silveira 2003 It is featured by the presence of subterranean flows and springs giving it a special nature by the presence of ecotones regulated by fresh and salty water MATERIALS AND METHODS Six bimonthly samplings including fish and physical variables were performed from May 2003 to March 2004 during climate sea sons of rain July September northern cold fronts norths November January and dry May March along 8 stations distributed in different environments Fish sampling was realized with a beach seine 15 m x 1 5 m 2 5 cm mesh size hauls by duplication covering an average area of 240 m Prior collection physicochemical variables were measured in situ temperature salinity and dissolved oxy gen using a Yellow Springs Instrument model 85 Fish were preserved in 10 formalin identified counted and measured for standard length to the nearest mm Data Analysis Spatial and temporal differences in salinity temperature and dis solved oxygen were subjected to different variance analysis of two ways variance analy sis ANOVA normality and homogeneity standards of variances were double checked by Kolgomorov Smirnov and Bartlett tests respectively Density and biomass were deter mined as the number and weight of organ isms per unit a
6. F persim 19 L rhomb cil Fig 6 Redundancy analyses RDA diagram of the overall fish abundances versus environmental variables represented by arrows Species abbreviations are the first letter of the genus name and first four letters of the species name TABLE 6 Comparative fish species number and coastal lagoon total area for some systems from Yucatan Peninsula and Gulf of Mexico Lagoon No Spp Bocas de Dzilam 8l Rio Lagartos 81 Celest n lagoon 94 Laguna Madre 84 of other coastal lagoons of the Yucatan Peninsula such as mojarras and anchovies Vega Cendejas et al 1997 Vega Cendejas and Hern ndez 2004 Moreover many commer cial importance species Haemulon plumierii Orthopristis chrysoptera Cynoscion nebulo sus C arenarius Lutjanus griseus A mitchilli A hepsetus E gula and E argenteus used the system for feeding raising and shelter Tough salinity gradients create physiological barriers for most species it has been found that season ally the marine euryhaline component like S notata enter to these coastal systems taking 9 4 km 80 km 28 14 km 200 km Area Authors Present study Vega Cendejas and Hernandez 2004 Vega Cendejas 2004 Raz Guzman and Huidobro 2002 advantage of the diverse food resources Many others use this environment as nursery such as L griseus E argenteus and E gula The small size recorded for most of the individuals indi cates the importance of the seagrass me
7. larity Clarke and Green 1988 Redundancy Analysis RDA was used to evaluate the inter relation among media densities and environ mental parameters This multi varied analysis and IVI were performed with the statistical pro gram ANACOM De la Cruz 1994 Spearman 90 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 correlation coefficient was used to determine the significance of each variable to fish dis tribution Zar 1984 Species ecological clas sification was performed separating fish in marine euryhaline marine stenohaline and estuarine residents species based on their rela tive abundance at each climate season Castro Aguirre et al 1999 Froese and Pauly 2004 RESULTS Hydrologic variables Temperature was significantly different among all climatic sea sons rainy and norths p 0 00014 rainy and dry p 0 002261 norths and dry p 0 00014 with the lowest during norths 24 C and high est in the rainy 31 C Fig 1 Salinity showed the lowest values during norths mainly in November with an average of 29 C with the highest value 37 C in the dry season No significant differences p gt 0 05 were found for dissolved oxygen Table 1 Species composition and spatial tempo ral variation A total of 6 474 fish were cap tured comprising 81 species and 37 families Temp C Salinity O mg l Sampling stations with a total weight of 64 kg Best repre
8. Peel on F polyommus 11 55 S testudineus 10 15 E argenteus 15 99 E gula 10 03 A mitchilli 8 80 S058 oe M trichodon 7 63 H parrai 6 16 F polyommus 6 16 E argenteus 13 64 4 E gula 11 90 78 62 S testudineus 9 67 F polyommus 8 90 M trichodon 8 24 Rains E argenteus 14 61 d 80 91 E gula 14 55 y L rhomboides 11 60 SASA M trichodon 9 93 S testudineus 10 95 E gula 8 59 Norths F polyommus 8 06 dry 83 99 E argenteus 8 04 season L rhomboides 6 20 H jaguana 5 94 F persimilis 5 91 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 TABLE 5 DISCUSSION Correlation matrix of overall means of hydrologic variables with fish species only those species with at least one significant correlation are shown p lt 0 05 Species composition and ecological cate gory Compared with previous studies and with other coastal systems in Yucatan Peninsula and Gulf of Mexico fish species richness of Bocas Symphurus plagiusa 1 344 1 458 2 954 de Dzilam 81 spp can be considered high Vega Cendejas ef al 1997 Raz Guzm n and Huidobro 2002 Vega Cendejas and Hern ndez Bairdiella chrysoura 2 636 0 764 2 636 2004 Table 6 However many of the spe cies are represented by low densities because they are in transit and enter to the system for Achirus lineatus 0 608 2 690 0 388 very short terms usually only to feed them selves Strongylura notata
9. Spatial and temporal characterization of fish assemblages in a tropical coastal system influenced by freshwater inputs northwestern Yucatan peninsula Daniel Arceo Carranza amp Ma Eugenia Vega Cendejas CINVESTAV IPN Unidad M rida Km 6 Antig Carr a Progreso A P 73 Cordemex C P 97310 M rida Yucat n M xico darceo mda cinvestav mx maruvega mda cinvestav mx Received 06 VII 2007 Corrected 10 VII 2008 Accepted 14 VIII 2008 Abstract Coastal lagoons are important systems for freshwater estuarine and marine organisms they are con sidered important zones of reproduction nursery and feeding for many fish species The present study investi gates the fish assemblages of the natural reserve of Dzilam and their relationship with the hydrologic variables A total of 6 474 individuals 81 species were collected contributing with more than 50 considering the Importance Value Index IVI Sphoeroides testudineus Fundulus persimilis Anchoa mitchilli Eucinostomus gula Eucinostomus argenteus and Mugil trichodon Differences in species composition were found between seasons the highest during the cold fronts Spatially differences were related with the presence of freshwater seeps the highest in the ecological characterized eastern part and the lowest with higher difference in specific composition located in the western part of the internal zone due to a higher abundance and dominance of L rhomboides Salinity and temperature were the
10. adows mangroves and mudflat to grow and shelter against predators Similar results were reported by Vega Cendejas and Hernandez 2004 for Rio Lagartos lagoon Temporal and spatial variations among assemblages Temporally during norths the highest species number 59 uses the lagoon as Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 99 protection against strong winds and was char acterized by their abundance and aggregation such as anchovies 4 mitchilli and A hepsetus and jacks Selene vomer High density and biomass percentages are generally associated to the occasional presence of small pelagic fish big schools Simier ef al 2004 Seasonal variability in fish assemblages reflects some species abundance due to reproductive puls es mainly during the first stages in their life cycle Machado and Ara jo 2003 For instance during the dry season 1t was recorded a F persimilis abundance peak in relation to its reproductive stage Similar results were found by Vega Cendejas and Hern ndez 2004 that reported an abundance peak for E grandis simus during dry season in R o Lagartos sug gesting a reproductive season for this species in the Yucatan coast In terms of spatial distribution the inner zone station 1 showed the least number of species 20 and Lagodon rhomboides was the most abundant with more than 60 of the total capture On the other hand stations with the high
11. atistical appraisal Mar Ecol Prog Ser 222 217 226 Simier M L Blanc C Aliaume P S Diouf amp J J Albaret 2004 Spatial and temporal structure of fish assem blages in an inverse estuary the Sine Saloum sys tem Senegal Estuar Coast Shelf Sci 59 69 86 Tsou TS Matheson RE Jr 2002 Seasonal Changes in the Nekton Community of the Suwanee River Estuary and the Potential Impacts of Freshwater Withdrawal Estuaries 25 1372 1381 102 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 Vega Cendejas ME 2004 Ictiofauna de la Reserva de la Biosfera Celest n Yucat n una contribuci n al conocimiento de su diversidad An Inst Biol Univ Nal Aut n M x Ser Zoo 75 193 2006 Vega Cendejas M E 8 M Hern ndez 2004 Fish com munity structure and dynamics in a coastal hypersa line lagoon Rio Lagartos Yucatan Mexico Estuar Coast Shelf Sci 60 285 299 Vega Cendejas ME F Arreguin amp M Hern ndez 1993 Trophic fluxes on the campeche Campeche Bank M xico pp 206 213 Jn Christensen V and Pauly D eds Trophic models of aquatic ecosystems ICLAEM Conf Proc Vega Cendejas M E M Hern ndez amp F Arreguin 1994 Trophic interrelations in a beach seine fishery from the northwestern coast of the Yucatan peninsula M xico J Fish Biol 44 647 659 Vega Cendejas M E M Hern ndez amp G De La Cruz Agiiero G 1997 Los peces de la Reser
12. biidae 0 01 0 001 mar D Garmanella pulchra Cyprinodontidae 0 01 lt 0 001 Estuar D Opsanus phobetron Batrachoididae 0 01 0 12 S mar N Bairdiella sanctaeluciae Sciaenidae 0 01 0 01 S mar N Syngnathus scovelli Syngnathidae 0 01 lt 0 001 E mar D Carangoides ruber Carangidae 0 01 0 007 E mar R Monacanthus ciliatus Monacanthidae 0 01 0 001 S mar N Caranx latus Carangidae 0 01 0 001 E mar N Archosargus probatocephalus Sparidae 0 01 0 004 E mar R Diapterus rhombeus Gerreidae 0 01 0 001 E mar N Syngnathus pelagicus Syngnathidae 0 01 0 001 E mar D Mugil cephalus Mugilidae 0 01 0 03 E mar R Pomatomus saltatrix Pomatomidae 0 01 0 01 S mar N Urobatis jamaicensis Urolophidae 0 01 0 39 E mar N Syngnathus makaxi Syngnathidae 0 01 lt 0 001 E mar D of fish species registered in Dzilam lagoon Yucatan E mar Eurihaline marine Estuar Estuarine S mar Stenohaline marine N norths R rainy D dry season 96 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 TABLE 3 R statistic values and their significance levels for pair wise comparisons of species composition of the sites and seasons using ANOSIM P lt 0 05 Differences inter sites and inter seasons as lvs 4 0 917 l vs 6 1 000 lvs7 0 542 lvs 8 0 750 Svs 7 0 500 6 vs 8 0 417 rains vs norths 0 203 rains vs dry season 0 359 norths vs dry season 0 344 p lt 0 001 p lt 0 05 terms of dissolved oxygen Achirus lin
13. dissolved oxygen Pomfret et al 1991 Blaber 1997 Marshall and Elliott 1998 We have to con sider that biological interactions that influence the composition of estuarine fish assemblages such as habitat preference predator prey rela tion food availability and often reproductive biology of species Therefore estuarine fish patterns of distribution can not be only attribut ed to physicochemical factors but to a synergy between biotic and abiotic factors CONCLUSIONS The main spatial and temporal differ ences between fish assemblages in a coastal system influenced by freshwater seeps was species composition due to the behavior and biology of each species at a given time and place Site 1 located in the inner zone bottom with Halodule wrightii showed the greatest difference amongst all sampling sites In addi tion it can be characterized as having a lower diversity value because of juvenile dominance L rhomboides i e using the system as a nursery Temporarily the greatest diversity and species richness was obtained during the north wind season because several marine species of ecological and economical importance used the system for shelter and feeding Salinity was the hydrologic variable that best related fish distribution patterns with a direct influence on the pelagic species A mitchilli A hepsetus H jaguana and O oglinum ACKNOWLEDGMENTS We are grateful for the help of many col leagues that was indisp
14. eatus and Dasyatis americana showed a signifi cant negative relation Spearman p gt 0 05 Temperature affected significantly Symphurus plagiusa B chrysoura and Sphoeroides neph elus while A lamprotaenia M trichodon and A felis were found in areas with higher temperature values Table 5 RDA analysis indicated that salinity and temperature were the principal hydrologic fac tors that influence composition and species distribution The first two axes explain 41 14 of variance and correlation 0 93 and 0 56 with A mitchilli and A hepsetus occurrence greatly influenced by salinity and temperature Fig 6 Some species distribution such as L rhomboides F persimilis H jaguana and S testudineus were influenced by other variables not measured in this research such as substrate type surrounding vegetation food availability and even specific biological considerations TABLE 4 Species contributing to the dissimilarities between sampling stations and climatic seasons based on relative abundance Groups tee P E a cal Dissimilarity Species L rhomboides 18 95 F persimilis 17 24 OR Sa H jaguana 10 08 E argenteus 9 83 L rhomboides 24 63 S1 S6 86 03 E argenteus 18 56 E gula 9 80 L rhomboides 29 95 E gula 11 06 nee 22 02 S testudineus 8 15 A rhomboidalis 6 97 L rhomboides 19 89 E gula 9 11 S1 S8 90 64 A mitchilli 8 75 M trichodon 8 10 H parrai 6 21 E argenteus 15 39 E gula 14 41
15. ensable to realize the fieldworks In special we want to thank Mirella Hernandez de Santillana for her assistance at fish sampling and in the laboratory thank you to all of you We are indebted to Luis Capurro for corrections to the English versions and to anonymous referees for their comments and suggestions that enrich the manuscript RESUMEN Las lagunas costeras son sistemas importantes para muchas especies de organismos dulceacuicolas estuari nos y marinos ya que son consideradas zonas de repro ducci n refugio y alimentaci n de muchas especies de peces El presente estudio analiz los ensamblajes de la comunidad ctica de la reserva de Dzilam y su relaci n con las variables hidrol gicas Se capturaron un total de 6 474 individuos 81 especies en donde Sphoeroides testudineus Fundulus persimilis Anchoa mitchilli Eucinostomus gula Eucinostomus argenteus and Mugil trichodon contribuyeron con m s del 50 del Indice de Valor de Importancia IVI Las diferencias en compo sici n de especies se encontraron entre temporadas la mayor durante los nortes Espacialmente las diferencias fueron relacionadas con la presencia de afloramientos de agua dulce la mayor en la zona Este de la laguna y la menor con mayor diferencia en composici n espec fica se localiz en la parte Oeste de la zona interna esto debido a la gran abundancia y dominancia de una sola especie L rhomboides La salinidad y la temperatura fueron las variabl
16. es que presentaron la mayor influencia en la distribuci n de algunas especies pel gicas como son A mitchilli y A hepsetus Debido a la abundancia de aflo ramientos de agua dulce caracter sticos de las lagunas costeras de la Pen nsula de Yucat n los ensamblajes de peces muestran diferencias espaciales y temporales en la composici n espec fica Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 101 Palabras clave Ictiofauna composici n de especies variaci n temporal entradas de agua dulce variables ambientales Bocas de Dzilam REFERENCES Blaber S J M 1997 Fish and fisheries of tropical estuar ies Chapman and Hall London England Blaber S J M amp T G Blaber 1980 Factors affecting the distribution of juvenile estuarine and inshore fish J Fish Biol 17 143 162 Brower JE Zar JH 1977 Field and Laboratory Methods for General Ecology W C Brown Co Publishers Dubuque lowa USA Castro Aguirre JP Espinoza H Schmitter Soto J 1999 Ictiofauna estuarino lagunar y vicaria de M xico Colecci n Textos Politecnicos Serie Biotecnologias Ed Limusa M xico Clarke K R amp R H Green 1988 Statistical design and analysis for a biological effects study Mar Ecol Prog Series 46 213 226 Clarke KR Gorley R 2001 PRIMER V5 User Manual Tutorial PRIMER E Plymouth UK Clarke KR Warwick RM 2001 Change in Marine Communities An Approach t
17. est number of species were 6 and 8 In site 6 located in the innermost zone of the lagoon many estuarine type and freshwater species were found 11 such as Cichlasoma urophtalmus C artifrons G pulchra G yucatana P velifera and Mendia colei This site is characterized by freshwater seeps that give hydrologic estuarine conditions and produce an abundance of organic matter personal observation which may explain the presence of these species even if salinity is high furthermore this site is characterized by a high habitat heterogeneity with mangroves and muddy flatland floors that contribute to ichthyofauna diversification even for shelter of important species such as C urophtalmus moreover they provide food to benthopha gous Gerreidae and detritophagous species Mugilidae Simier et al 2004 Station 8 located in the external zone near the inlet is the other site with the highest species rich ness in the lagoon This place has a permanent communication with the sea favoring transit for many interacting species Predominantly open estuaries are species rich their permanent or near permanent connection with the sea allows access into these systems Harrison and Whitfield 2006 According to abundance and species com position sampling sites were distributed in three groups Figure 5 the first one included stations located in the external zone 7 and 8 where specific composition comprises mainly mar
18. eta Orthopristis chrysoptera Strongylura timucu Synodus foetens Trachinotus falcatus Cichlasoma urophthalmus Elops saurus Oligoplites saurus Cynoscion arenarius Cynoscion nebulosus Lucania parva Paralichthys albigutta Scorpaena brasiliensis Prionotus tribulus Sphyraena barracuda Menticirrhus saxatilis Hyporhamphus roberti Bairdiella chrysoura Stephanolepis hispidus Chilomycterus schoepfii Symphurus plagiusa Family Sparidae Mugilidae Clupeidae Haemulidae Sparidae Poeciliidae Belonidae Carangidae Achiridae Haemulidae Tetraodontidae Engraulidae Lutjanidae Ariidae Engraulidae Mugilidae Poeciliidae Batrachoididae Haemulidae Belonidae Synodontidae Carangidae Cichlidae Elopidae Carangidae Sciaenidae Sciaenidae Fundulidae Paralichthyidae Scorpaenidae Triglidae Sphyraenidae Sciaenidae Hemiramphidae Sciaenidae Monacanthidae Diodontidae Cynoglossidae N 4 13 3 02 1 42 1 36 1 28 1 16 1 05 0 71 0 61 0 60 0 43 0 43 0 39 0 39 0 38 0 37 0 30 0 26 0 25 0 21 0 21 0 20 0 20 0 17 0 10 0 08 0 08 0 08 0 08 0 07 0 07 0 07 0 07 0 06 0 06 0 05 0 05 0 05 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 of fish species registered in Dzilam lagoon Yucatan W 2 68 5 39 0 15 2 43 2 68 0 09 4 87 1 84 0 19 0 43 0 43 0 02 0 71 2 53 0 17 0 21 0 007 0 24 0 91 0 87 0 37 1 10 1 79 0 55 0 08 0 04 0 21 0 001 0 03 0
19. il ity of nutritional resources and the protection against predators Blaber and Blaber 1980 Marchand 1993 Blaber 1997 Marshall and Elliot 1998 Estuarine fish assemblages show cyclical variations in abundance and composi tion and since they include species of fresh and marine waters they can show sedentary or migratory behaviors and be present in diverse stages of their development Simier et al 2004 as well as seasonal changes in their habitat types because of coastal processes Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 89 Few studies have been realized in Yucatan Peninsula and specifically the ichthyofauna community in Bocas de Dzilam Reserve has not been assessed Vega Cendejas et al 1993 Vega Cendejas et al 1994 Vega Cendejas et al 220 Because of its reserve status and to its ecological importance for estuarine ichthyo fauna this research aims to investigate fish assemblages distribution related to climatic season and some hydrologic variables Study area Bocas de Dzilam lagoon declared a protected natural area since 1989 Zone of Ecological Conservation Reserve of Dzilam is located in Southeastern Mexico in the central coast of Yucatan State 21 19 21 32 N 88 35 88 58 W with a surface of 9 4 km 12 9 km long a maximum width of 1 65 km with a permanent connection with the sea 375 m wide bordered by mangrove and depth
20. ine species considered only in transit and that occasionally get into the lagoon The use of surf zones by a great number of fish mainly in juvenile stages is likely responsible for the existence of rich sources of food in form of zooplankton and for the protection against predators that those shallow low turbid and low turbulent waters provide Lasiak 1986 The second group is conformed by site 1 located in the inner zone which forms an independent group due to a high representation of small size individuals 3 5 cm of L rhom boides This zone is characterized by a bed of submerged vegetation dominated by Halodule wrigthii Herrera Silveira et al 1998 This shows the ecological role of seagrass meadows as growing and feeding zones for juvenile stages of marine species Jenkins ef al 1997 Gray et al 1998 The third group comprises inner stations with dominating species of wide distribution within the lagoon These sites are also characterized by shallow depths with muddy bottoms sometimes with submerged macrophytes and macroalgae aside from high turbidness and a great quantity of decomposing organic matter which is used by detritopha gous species Relation between species and hydrologic variables The ever changing conditions in hydrology can become a physical barrier for the access of many marine origin species such is the case of salinity gradients while varia tions in temperature can affect ichthyofauna distribution
21. lasoma urophthalmus Lucania parva Cyprinodon artifrons and Garmanella pulchra Fig 4 Table 2 Spatial and temporal differences among assemblages ANOSIM indicated significant differences in fish assemblage composition on the climate season R 0 292 p lt 0 05 and spa tial R 0 335 p lt 0 001 among sites 1 4 1 6 1 7 1 8 5 7 and 6 8 Table 3 Total significant dissimilarity among sampling sites shows a range between 83 09 and 93 62 Table 4 These differences are caused by abundance among dominating species in each sampling site being L rhomboides E argenteus E gula S testudineus H jaguana F persimilis F polyommus A mitchilli M trichodon and H parrai contributing with more than 50 to the spatial dissimilarity Among climate seasons there is a total range of dissimilarity between 78 62 and 83 99 with more than 50 by E argenteus E gula S testudineus F polyom mus M trichodon L rhomboides H jaguana and F persimilis Table 4 According to statistical test SIMPER sta tion 1 is characterized by L rhomboides which was found in 58 40 as well as other species such as S testudineus E gula and E argenteus which showed important percentages in the specific characterization of the sampling sites There are stations where dominancy is shared and characterization is determined by many species that represent percentages lower than 30 such as sites 4 7 and 8 Total similarity for each sa
22. mpling site throughout time was from 5 site 7 to 44 site 2 showing the similarity degree of each site among differ ent seasons Temporally species common for rainy season were E gula E argenteus S testudineus and M trichodon with more than Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 93 120 100 8040 S marine D E 60M co g lt amp 40 Rains Norths Dry season Fig 4 Temporal distribution of ecological categories of the Bocas de Dzilam fish assemblages 60 furthermore norths are characterized by E gula E argenteus S testudineus and E polyommus while dry season shows 8 spe cies E argenteus E gula S testudineus M trichodon L rhomboides H jaguana H bonariense and A lineatus that contributed with more than 85 Considering abundance and species com position sampling sites are conformed by three main groups The first one includes sites 7 and 8 located in the external zone of the lagoon marine zone and characterized by the high est salinity value 35 38 species richness and evenness In the second group the site located in the inner zone No 1 forms an independent entity with the lowest salinity record and the species richness and diversity because of the dominance of L rhomboides Finally the third group is conformed by sites located in the center of the lagoon 2 6 Fig 5 Relation between species a
23. nd hydro logic variables Species that showed a positive correlation Spearman p lt 0 05 with salin ity were Cynoscion arenarius A hepsetus T falcatus Oligoplites saurus H jaguana O oglinum and Ariopsis felis while Bairdiella chrysoura preferred oligohaline waters In TABLE 2 Relative abundance N and biomass W ecological category E C and seasonal occurrence S O of fish species registered in Dzilam lagoon Yucatan Species Family Anchoa mitchilli Engraulidae Fundulus persimilis Fundulidae Anchoa hepsetus Engraulidae Eucinostomus gula Gerreidae Eucinostomus argenteus Gerreidae Sphoeroides testudineus Tetraodontidae Harengula jaguana Clupeidae Floridichthys polyommus Cyprinodontidae N W ETC S O 18 71 1 48 E mar ND 18 18 20 77 Estuar D 8 80 0 37 E mar N 8 18 3 31 E mar RND 8 11 2 54 E mar RND 6 03 30 42 E mar RND 5 67 4 11 E mar RND 4 73 1 28 Estuar RND 94 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 TABLE 2 Continued Relative abundance N and biomass W ecological category E C and seasonal occurrence S O Species Lagodon rhomboides Mugil trichodon Opisthonema oglinum Haemulon bonariense Archosargus rhomboidalis Poecilia velifera Strongylura notata Selene vomer Achirus lineatus Haemulon parra Sphoeroides nephelus Anchoa cubana Lutjanus griseus Ariopsis felis Anchoa lamprotaenia Mugil curema Gambusia yucatana Opsanus b
24. o Statistical Analysis and Interpretation 2 edition Natural Environment Research Council Plymouth Marine Laboratory Plymouth U K De La Cruz AG 1994 ANACOM Sistema para el An lisis de Comunidades en computadoras person ales Versi n 3 0 Manual del usuario CINVESTAV IPN Unidad M rida Yucat n M xico Francis M R Hurst B McArdle N Bagley amp O Anderson 2002 New Zeland demersal fish assem blages Env Biol Fish 65 215 234 Gray C A R C Chick amp D J McElligott 1998 Diel changes in assemblages of fishes associated with shallow seagrass and bare sand Estuar Coast Shelf Sci 46 849 859 Harrison D T amp A K Whitfield 2006 Estuarine typol ogy and the structuring of fish communities in South Africa Env Biol Fish 75 269 293 Herrera Silveira J A R J Ram rez amp A Zaldivar 1998 Overview and characterization of the hydrology and primary producer communities of selected coastal lagoons of Yucatan M xico Aquatic Ecosystem Health and Management 1 353 372 Jenkins GP May HMA Wheatley MJ Holloway MG 1997 Comparison of fish assemblages associated with seagrass and adjacent unvegetated habitats of Port Phillip Bay and Corner Inlet Victoria Australia with emphasis on commercial species Estuar Coast Shelf Sci 44 569 588 Kupschus S amp D Tremain 2001 Associations between fish assemblages and environmental factors in near shore habitats of a subtropical estuar
25. omponent higher per Evenness J centage was shown in norths constituted by F 0 7337 0 3603 Opisthonema oglinum Archosargus rhom P 0 6470 0 7016 boidalis Haemulon bonariense H parrai Log e aid ba i Anchoa cubana A lamprotaenia Trachinotus a 0 0750 0 5955 falcatus and Sphyraena barracuda among Log Biomass others while resident estuarine species were F 2 1177 1 4126 shown with higher incidence in dry season rep P 0 0323 0 2501 resented by F persimilis Floridichthys poly ommus Poecilia velifera Gambusia yucatana pP lt 0 05 A rhomboidalis S notata A hepsetus E polyommus L rhomboides H jaguana M trichodon E argenteus E gula A mitchilli F persimilis i a 4 S testudineus o N gt 6 8 10 o BR a Importance Value index Fig 2 Dominant fish species collected in Bocas de Dzilam lagoon during the study period ranked by the Importance Value measure 92 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 3 8 3 4 J aE 1 6 1 4 1 0 0 6 Bits ind Diversity H 30 24 a Gal No species 0 95 0 85 0 75 eS E yka 0 55 0 45 0 35 0 25 Eyenness J 12 345 678 Sampling stations 12 8 0 75 0 55 0 45 pul Sep y Nov Jan Mar May y Rainy Norths Dry season Fig 3 Diversity index total number of species and evenness of the sampling stations and season variation Cich
26. rea ind m g m extrapolated to 10m Spatial and temporal differences in density and biomass were determined by two ways analysis of variance after transforming data to log x 1 and verifying homocedastic ity For this last analysis extremely abundant species were eliminated Clarke and Warwick 2001 Tsou and Matheson 2002 Community structure was analyzed through richness S evenness J and diversity of species H using Shannon Wiener index Dominance implying hierarchical structuring and evaluation of rela tive specie importance was carried out in each zone and on annual basis with Index of Value of Importance IVI which incorporates informa tion about density DR frequency FR and relative biomass BR for each species IVI DR FR BR Brower and Zar 1977 Spatial and temporal changes in assem blage ecological parameters were performed with statistical program PRIMER 5 Plymouth Routines in Multivariate Ecological Research Clarke and Gorley 2001 Fish assemblage differences among sites and or climatic seasons were tested by Similarity analysis ANOSIM non parametric test that applies the Bray Curtis similarity matrix Clarke and Green 1988 Species contribution by climate season or sampling station was evaluated with similarity percentage SIMPER test which determines percentage contribution of each species in order to classify a group similarity and discrimi nate species among sample groups dissimi
27. sented families by number of species were Gerreidae 4 genus and 7 species Carangidae 4 genus and 5 species Sciaenidae 3 genus and 5 spe cies Engraulidae 1 genus and 5 species and Syngnathidae 1 genus and 5 species The most abundant species being Anchoa mitchilli Fundulus persimilis A hepsetus Eucinostomus gula E argenteus and Sphoeroides testudineus while S testudineus F persimilis and Mugil trichodon contributed with 56 6 of total weigh Results of two way ANOVA P gt 0 05 indicated not differences in density and bio mass among climate seasons and sampling sites Table 1 Considering both ecological parameters and their occurrence frequency 12 species contributed with more than 70 with S testudineus 14 2 being the most representa tive Fig 2 Diversity H evenness and species rich ness did not vary significantly P gt 0 05 among neither climate seasons nor sampling sites Fig 3 Spatially average diversity was 2 40 0 69 bits ind with maximum values recorded in site 5 2 9 bits ind and minimum in site 1 1 75 bits ind In the inlet of the lagoon site 8 the 35 2 30 amp 25 oO W 20 15 10 40 O 35 pete ee So 25 20 15 10 20 15 10 O Jul Sep Nov Jan Mar May Rainy Norths Dry season Fig 1 Salinity temperature and oxygen variations in sampling sites A and bimonthly B 1 STD Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2
28. va de la Biosfera de Celest n CINVESTAV M rida PRONATURA Pen nsula de Yucat n M xico Whitfield A K 1999 Ichthyofaunal assemblages in estu aries A South African case study Rev Fish Biol Fisheries 9 151 186 Zar JH 1984 Biostatistical Analysis 2nd edition Prentice Hall New Jersey USA INTERNET REFERENCES Froese R Pauly D 2004 FishBase version 09 2004 Downloaded Feb 24 2005 www fishbase org Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 57 1 2 89 103 March June 2009 103
29. variables that presented a higher influence in the distribution of some pelagic species such as A mitchilli and A hepsetus Because of the abundant freshwater seeps character istic of the coastal lagoons of Yucatan Peninsula their community structure and fish assemblage display spatial and temporal differences in specific composition Rev Biol Trop 57 1 2 89 103 Epub 2009 June 30 Key words Ichthyofauna species composition temporal variation freshwater inputs environmental variables Bocas de Dzilam Coastal lagoons and estuaries are environ ments featured by being transitional places between land and marine habitats as well as dynamic systems with highly environmental variability in short and long term So that their biological communities are influenced mainly by physical environment Kupschus and Tremain 2001 and their structure diver sity and stability are defined by biologic inter actions intra and interspecific Francis et al 2002 Within these ecosystems fish consti tute the main biological component of marine and estuarine species using them as feeding reproduction growth and protection grounds Raz Guzm n and Huidobro 2002 in terms of their tolerance limits to those hydrologic conditions Whitfield 1999 Variables affect ing their distribution in estuarine waters are salinity temperature turbidity and dissolved oxygen though sediment type heterogeneity and vegetation also influence the availab
30. y J Fish Biol 58 1383 1403 Lasiak TA 1986 Juveniles food and the surfzone habitat implications for teleost nursery areas South African J Zoo 21 52 56 Machado P A L amp F G Ara jo 2003 Spatial temporal and diel variations of fish assemblages at two sandy beaches in the Sepetiba Bay Rio de Janeiro Brazil Estuar Coast Shelf Sci 57 817 828 Marchand J 1993 The influence of seasonal salinity and turbidity maximum variations on the nursery function of the Loire estuary France Netherlands J Aquat Ecol 27 427 436 Marshall S amp M Elliot 1998 Environmental Influences on the Fish Assemblage of the Humber Estuary U K Estuar Coast Shelf Sci 46 175 184 Medina G mez I Herrera Silveira J A 2003 Spatial char acterization of water quality in a karstic coastal lagoon without anthropogenic disturbance a mul tivariate approach Estuar Coast Shelf Sci 58 455 465 Pomfret JR Elliott M O Reilly MG amp Phillips S 1991 Spatial and temporal patterns in the fish communi ties in two U K North Sea estuaries p 277 284 In Elliott M amp Ducrotoy SP eds Estuaries and coasts Spatial and Temporal Intercomparisons Olsen amp Olsen Fredensborg Denmark Raz Guzman A amp L Huidobro 2002 Fish communities in two environmentally different estuarine systems of Mexico J Fish Biol 61 182 195 Rueda M 2001 Spatial distribution of fish species in a tropical estuarine lagoon a geost

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