functional groups and ecological biodiversity, in
Transcripción
functional groups and ecological biodiversity, in
Rev. Soco Mex. Hist. Nat., 49:163-172 FUNCTIONAL GROUPS AND ECOLOGICAL BIODIVERSITY, IN TERMINO S LAGOON, MEXICO* Grnpos FunciofUlies y Biodiversidad Ecológica en la LagUfUl de Ténninos, México A. Yáñez-Arancibia', P. Sánchez-GiI' and A.L. Lara-DomínguezJ• ABSTRACT An intcgratcd approach, as requircd lo comprehend the complexity of tropical coastal biodivcrsity (374 nora specics, and 1468 macro fauna species al the study sitc) is discussed within a frarncwork of lagoon-cstuarinc system gradients of habitats, of forciog fu"clioos (or production mechanisms). and scasonally stable asscmblagcs of biola with similar biological bchavior and ecological strategies (or funcJionalgroups). Thcse assemblages play ao important ecological role by coupling life-history strategics wilh the cnvironmental variability and estuarine gradienls. The complcmentarity of planktonic and maeroOoral primary production and their coupling with secondary produetion are also discusscd and we conclude that scquential pulsing is one of the funclional processcs that sustain hlgh estuarinc production and species diversity. The coneept of seasona/ programming describes how the functional groups of estuarine primary producers interact, and how functional groups of fish population assemblages use the lagooo hahitats in time and space [O reduce the effccts of competition and predation, to enhance food availability and to secure their recruitment. Ke)" words: tropical estuary, functional groups, seasonal programming. RESUMEN Para comprender la complejidad de la biodivcrsidad costera tropical (374 especies de flora y 1468 de especies de macro fauna en el área de estudio) se requiere de una aproximación integral y esto debe ser discutido en el marco de un sistema lagunar-estuarino que presenta diversos gradientes de habitats. diferentes funciones de fuerza (o mecanismos de producción) y numerosos conjuntos bioticos -estacionalmente estables- que tienen comportamiento biológico y estrategias ecológicas similares (o grupos funcionales). Estos conjuntos juegan un importante papel ecol6gico vinculando las estrate gias del ciclo de vida con la variabilidad ambiental y los gradientes estuarinos. Complementariamente se discute la producción planctónica y de la macro flora y su vínculo con la producción secundaria, concluyendo que los pulsos estacionales son el proceso funcional que sostiene la alta productividad estuarina y la diversidad de especies. El concepto de "programación estacional" descriebe como los grupos funcionales de conjuntos de poblaciones de peces usan los hábitats lagunares-estuarinos en tiempo y espacio para reducir los efectos de competencia y predación, favoreciendosc ante la abundante disponibilidad de alimento y asegurando su reclutamiento. Palabras clave: tropical, grupos funcionales, programación estacional. • Los autores dedican este trabajo al Dr. Auslín Ayala 2. ConsuHor en Ecosistemas Costero-Marinos A. P. 206 • Castañares, con profundo reconocimiento por ser el pionero Xalapa 91001 Veracruz, México. de las ciencias marinocosteras en México. 3. Centro de Ecología, Pesquerías y Oceanografía del GolCo l. Departamento de Recursos Costeros, Instituto de de Mexico (EPOMEX). Uninrsidad Autónoma de Campt.-che. Ecololo:ía A.C. Xalapa. Antigua Carretera Coatepec km 2.5, Ap. Postal 520, Campeche 24030 Mexico. A.P. 63, Xalapa 91000 Ver., México. YÁ:';EZ-AR¡\.\lCIB\. A.. e SÁCIlEZ-GIL y A.L. LARA-DOlÍG¡;EZ 164 lnlroduction Biouiversity is usually uefined at three levels, i.e., genetie, spceies/population. anu ecosystem (Ray and MeCormiek, 1992). The hiouiversity 01' eoastal environrnents is generally high in all respcets; however information on tropicallagoon-estuarine ecosystcms is usually no! as detaileu a< required, given the variety 01' n::sponses to cnvironrncntal changcs. lhe divcrse spacc and time scales with which organisms ¡nterael ",ilh their habitats, anu the generallaek ofinformation on these arcas as compared to tempeTate systems (Longhurst and Pauly, 1987). The uiverse biological, eeologieal anu physieal interaetions whieh oeeur is the most conspicuous part orthe Usumacinta Del ta, farmcd by a river with the seoonu highest dischargl.: in the Gulf of Mexieo after the Mississippi. The arca is one of the warlu's best known tropical estuarics (Yáñez-Araneibia and Day, 1988; Yáñez-Arancibia el al. 1988, 1993a; Rojas Galavizel a/., 1992). In tcrms of funetional eharaeterization, this area of2500 km' can be diviueu into three regions (Fig. 1). foHowing Kjerfve (1989,1993) anu Yáñez-Arancibia (1987), as follows: l)A (idal riJ'er Wne. Ruvial-dcltaic zone charactcrizcd by lack oC ocean salinity, but subject to tidal risc and [aH ofwater leve!; within tropical cstuaries and lhe adjacent oecan 2) A mixing zone. Charactcrized by water mass mixing produce a highly dynamie anu variable mosaic 01' anu existenee of strong gradients of physical. eCllsystems (Yáñez-Araneibia el al., 1991). chemical and biotic fcatures, and rcaching from From an ecological stanupoint. the term biodiversity can have several meanings when applieu to tropical the lidal river zone to the seawaru location ofriver mouth bar or ebb-tidal delta; lagoon-cstuarinc ccosystems (Day and Yáñcz 3) A near sllore tllrbid zone. In the near-shore oecan, Araneibia. 1982). It can mean that there is a high bctwcen the mixing zone and the seaward euge of diversity 01' speeies, or that there is a high uiversily 01' the tiual plume al full ebb tide. cnvironrncntal factors, habitats. connections in lhe foou webs. anu a high uiversity 01' eouplings, both internally ano with neighhoring systcms. AIso lhe diversity 01' foreing fllnelions effeetively modulating eCllsystem funetioning is high, anu inelude winu, tiue, rivers tlows, ¡ittoral currcnts, scdiment input, and others. Moreover there is a high uiversity among This subdivision oC Terminos Lagoon diffcrs from those previously proposed in thal it recognizcs and includes nearshore marine eomponents that arc cstuarine in eharacter. and implicitly considcrs the five main habitats in the s)'stem as a wholc, deSlTibcd. by Yáñez-Araneibia anu Day (1982). primaty producers and consumers, manywith different typcs oflife history. Grassle el al. (1991) anu Lasserre (1992) pointeu out that large-seale experiments are neeued on whole system responses anu on earcfuHy seleeteu tropical sites in oruer to identify a new paradigm for understanding coastal-marine diversity. More information is neeucd on tropical estuarics for lheir complexity anu biodiversity to be unuerstood. Habitat diversily The semi-pcrrnanent grauients from the tiual wetlands to the estuarine plume on the inner shelf anu aujaeen! sea leads to the identification of scven main habitats in the study area, as oiscusscd by Yáñcz-Arancihia and Day (1988) and summarizeu in Figure 1: Thus, in this paper, we foeus on the funelional 71fejlw'iaJ.JelJaic sy!J1ems in the southern tittaral zone bimliversity of thcse ecosystems, and describe the divcrsity offorcing functions; using the habitats and the seasonality of funetional groups, i.e., primary of Terminos Lagoon have very Inw salinity. high producen> and consumers, in Terminos lagoon, as rnangrove forests. and in some arcas of c1car walcr, example (se e Fig. 1). turbiuity, high nutrient eoneentrations, silty-c1ay sediments, Crassostrea drginica recfs, rivcrinc subrnergcd fresh water vegetation. Tllecentral basin, which is the transition zone bctwecn marine The lagoon-esluarine system conditions and the zonc intluenccd by the rivers. i characterizcd by mcsohalinc salinity, medium walcr Terminos Lagoon is loeateu in the sOllthern GlIlf of transparene)'. silty-clay to sanuy seuiments and a Mexico at 18" N, were tropical elimatie eonuitions typieal estuarine phytoplankton produetion syslem occur: a rainy season from June lO September, winter storms ("nortes") from Oetober lo February, and a ury season, from February lo May. Terminos Lagoon lOlIe aJ El Canllelllslalld. is uominateu by 1Jwlassia inellluing sorne benthie macroalgae. nI< inller lit/oral te,ftudinul1l ano fringe rnangrove habitat with ncar ECOLOGICAL FüCTIOAL GROLJI'S. el ge [El Puerto Reallnlel TERMINOS LAGOON ca rn Inner LiUoral es lIa 165 ..... "._.. ' . Q Central Basin ns 112 og Fluvial-dettaic '), o El Carmen Inlet ,o \jI.: ..... Net flow 19 .1, 11 " 1, )1' CAMPECHESOUND "1','1'1'.',',',',',',',',',',',','1',',','1',1,11',1t' ""I',',',',I,',',',',',',',',',',I,I,',','¡',!,',','I "I',',',',',',',',',I,',',',','¡',','.',','¡',I,',',' ,'¡',',',',',',',',',',',',',',"',l,',',',',',',',',' '1',',',11',1,',',',',',',',',1,',1,1,',',1,','111',1 ',',',',11'11,','1',1,1,',',',',1,',1,',',1,',1,'11,1 ,',',','1',11',',',1,',',',',',',',1,',',1,',',1.' 1 I 1 I I I 111 III I 1111 I 1 II I I • I ',',I,',I,',I,',',',',"',I,'t'I"'I""'" '",', 1,1,1,',',1,1,',1,'111',','1',','1',1,'1 1,1,1,11 ,',' I 1 I 1 1 I I I I I 1 1 I I 1 I 1. Tran.lbonalun. 111,','1',',',',',',',',1,'1',1,',' 111 1 l' n "il o '1'11111 I I I 1 I I 1 I I 1 1 1 I I III'IIIIIIIIIIIII'IIIII! 1111,1 llllllllllllllllllllllllltlr!.1 1 11 111111111111 11 1111 ,11 11. 1 '1111111'11,1111111.1 1,IIIIIIIIIJIIIII .Ir:: e \- e 1, DZoneA [ili] Zone B .1 T Fluvial discharge Fiure 1. lIabitat dhersity in the estuarine ecosyslem uf Terminos l.agoon (ahuve) and the adjacenl Campeche Snund (bt-Iow), showing the dilTerenl habitats (or ecolugical subsystems). The inner shelf shows Zunes A and B. associaled wilh El Carmen and Puerto Real inlels respecth.ely. The laoon.cstuarine system consists of 'he maio hahitals relaled lo l\ater circulation. rh'er dischal)!:c, salinit)' and water depth. (After Yáñcz.Arancibia and Dn,..• 1988). marine salinity, high waler lransparency, sanoy clear waler 01' full salinity, ealcium carbonate sano sedimenls and a high diversily 01' bOlh benlhie and sedirncnts, ano with seagrass bcds ano malToalgae. nektonic consurncrs. 71le Puet10 Real In/el, whcre there is a nel flow 01' Gulf waler inlo lhe lagoon, is characterized by calciurn carbonate sediments, c1car watcrs ano cxtcnsive scagrass beds. El Carmen In/el, lhe esluarine conneelion wilh the shelf, has a nel transport from the lagoon to the oecan, producing an cxtensive estuarine plumc of meoiurn salinityon the shclf. There are silty-clay sediments ano highly turbio waters without submcrgcd vcgetation in this arca. In Campeche Souno aojaeent lo "'orminos Lagoon, thcrc is a lerrigellolls subs)'slem (7..one A) strongly influcnecd by cstuarinc watcrs with rncúium water transparency, no benthievegelation and wilh a high Divcrsily uf furcing funclions Wilhin lhe physical framework provideo by the wetlands, lhe lagoon-cstuarine cnvironment anO the continental shclf; the living rcsaurces maintain a complex biological organization in thc stuoy arl:a. Figure 2 illustrates the complcxily 01' lhe SystClll (which couples the fish resourees wilh the coaslal ecologieal processes) ano lhe biological proouelion mechanisrns (i.c., the forcing functions modulating thc structurc and functioning of lhe ccosystcm). ennlenl 01' organic maller in lhe silly-clay-sandy Functionally. fish rcsaUTces are rclated lo ecological scoimcnts; and a calcareOllS subS)'stem (lime B) with interactions in thc coastal zanc, anu changes in fish 166 YA:':EZ-ARAXClRlA. A .. P. SAI'CIIEZ-GIL y AL LARA-DOlíI'GUEZ rcsourccs in spacc and time are rcflection of the natu Figure 2 show how these faclors inleraello support ral variabilily of physical and biological processes in lhe various groups forming lhe basis of fisheries in the lhal lOne. However, ehanges within lhe high diversily region, in a highly dynamic interactivc systcm in eeosyslem are generally dimeult lo predi el, beeause various lime/spaee scales. The block of small pclagic of lhe large number of inleraeling groups (e.g., fishes includes sardines (C1upcidae) and anehovies eompclilion, predalion). Slill, quanlilalive models of aspeels of lhe resouree systems can be conslrueted, e.g., for managing lhe coastal fisheries relying on the fish resourcc base (Arreguín-Sánehez el al., 1992). The mosl eonspicuous faelors (or foreing funclions (Engraulidae). The lop predator block comprises sharks, dogfish, grunls (Pomadasyidae), snooks (Centropomidae), groupcrs (Serranidae) and snappcrs (Lutjanidae). The small demersal fish block ( < 15 cm) is comprised offlatfish (Soleidae, Bolhidae), silver pcreh (Gerreidae), and scorpion fish (Scorpcnidae). in lerms of lhis paper) affeeling fishery production, The medium-sizcd demersal fish block (between 15-25 are: 1) physical/chcmical conditions in the water cm) includes gilt heads and yellow jacks (Carangidae). columo Le., transparency. nutrients, salinity, eroakers (Sciaenidae), porgies (Sparidae), puffers tempcralure, 2) lalilude, 3) balhymelry and sedimenl (Telraodonlidae), sea robins (Triglidae) and olhers. lypcS, 4) meleorology and clima te, 5) river discharge, The large demersal fish block (>25 cm) includos 6) lidal range, 7) lhe area of coaslal vegelalion i.e., among Olhers, ribbon fish (Triehiuridae). rays marshes, swamps, lagoons and estuaries, and 8) lhe (Dasyalidae), seaeatfish (Ariidae). lizard fish intcractions bctwcen the estuaries and the sea. (Synodonlidae), croakers (Sciaenidae) and olhers. Fisheries, Coast Zone 25 26287 Southern Gulf of Mexico ( 0.30 to 8.30 Primal)' Production I md ell.a rr.-' eSluanne-lagoo > 3.6 km hol oo Iy d-' t' 8 -0.050-1 10 + 0.0892 m Figure 2. There ¡s 11 high dinrsil)' or species. habita., rorcing runclion. and inlereonneclion in eoasla) systcms, such as Tcrminos Lagoon which is iIIuslrated here. The diaJ:ram show lhe dh'ersily or fishery groups (squares), and thtir intcrrelalionship with eoasta! and biological processes, and produetion mechanisms, (rorcing funclion, indicaled bJ circlesl in Ihe lo,,"er parl (Ir Ihe diagram. The eatehes are in metrie tan s per year. Values af lisheries ealeh are giHn in lons/year and lhe sum of the nows in A,B, and e is 3 x 1()J lonsl)"ear. (Arter lOC-UNESCO, 198). ECOLOGICAL FU!'ICTIO:--lAL GROLPS. Species diversity Because of ilS high biologieal diversity, in lerms of bOlh habilals and speeies, Terminos Lagoon is al presenl being proposed to become an "ecologically 167 2) lhe eenlral basin wilh salinity ranges from 10 lo 25 %0, and where phyloplanklon are lhe dominant primary producers; 3) shallow intertidal and sublidal zones wilh high proleeted coastal ecosyslem" under lhe framework of marine intluence (> 25 %0) dominaled by fringing lhe Mexican "Federal Lawfor Ecolagical Equilibrium mangroves and submcrged seagras.."i beds. ami £1l1,'irOnmenla/ Prolection", (Y áñez-Arancibia el al., 1993b). Resulls oflhe lasl20years ofresearch on ilS valuable wildlife resources led to lhe informalion summarized as follow. The lisl of lolal speeies is induded in lhe above ciled report, consislof374 plant species (of which 3 are endangered and 64 are endemics), and 1468 species ofmacrofauna (ofwhich 510 are inverlebrales and 958 are verlebrales). From lhe land verlebrales, 67 species are endangered, 30 are endemics, 22 are under proteetion and have cconomic valuc. These regions includes lhe habilals in Figure 1. Eaeh of these is dominaled by specifie primary producers forming funelional groups. The importanee of these groups is thal they represent a food source for cstuarine and coastal consurners, bcsidl:s acting, as the case might be, as a critical habitats ror a grcat number of organisms during various life stagcs, and regulating importanl components of lhe eSluarine ehemical cycles (Odum, 1988; Day el al., 1989; Ro jasGalavizetal.,1992). Figure 3 shows lhe clear seasonality of lhe pulses of abundanee and produetivily of differenl primary Diversity of functional groups and seasonality produeers funelional groups. Primary praduetion is generally high during all year, bul each funelional group has a differenl seasonal panem, in relalion 10 Termine" Lagoon has a number of imporlanl functional the environmcntal parameters specific to cach groups, bOlh of primary and sccondary producers (or gradient. The main productivity peak of mangrovcs consurncrs). A fimctional group is a conspicuous oceurs during the rainy season and may he related to assemblage ofbiola with similar biologieal behavior freshwaler inpuI (nulriems and lowered salinity). For and ecological slrategies. These assemblages, whieh submerged grasses, the highesl biomass and contain a numbcr uf diffcrcnt spccies and are produetivity generally occur during lhe dry season, characteristic elements in the structurc ano while the lowcst biomass valucs occur during the rainy functioning of lhe eeosyslem play an important and "nartes" seasons. Both marine and frcshwatcr ecological role by lhe coupling life-history slralegies grasses slarllhcir produelivity pulse allhe end of lhe of their consliluenl species wilh lhe environmenlal "nortes". Maximal produetivily of freshwaler variability and eSluarine gradienls. macrophytes oceur in February and slowly declines during lhe dry and rainy seasons, while peak seagrass biomass and productivity occur during lhe dry season FunClioning o/primary produurs The main eeological role of primary producers is to provide food via pholosynlhesis. In tropical estuarine and lagoon syslems, sorne primary producers also provide imporlanl crilical habilats: mangrove swamps, submerged grass beds. The diversity of fllnclional groups of primary producers can be high in esluarine ecosystems (Rojas Galaviz el al., 1992). Salinity gradicnts, turbidity, scdirncnts, nutricnts, ami tidal range, controllhe distribulion of primary producers, (Mareh-May). Phyloplanklon produclivily and biomass in the mid-lagoon has a scasonal pattcrn opposile thal of lhe aqualie macrophyles. PlanklOnic primary produclivity and ehlorophyll a levels increase lhrough lhe rainy season, reaehing a peak during lhe beginning oflhe "nortes" season from September unlil Dcccmbcr. Aquatic primary productivity in mangrove borde red lidal ehannels is highesl during lhe dry season Dayetal. (1982,1988 a, b) showed lhaldissolved organic matter exported from mangrove stimulated in the three aboye mentionncd regioos a follows: aqualie primary produetivity. 1) lidal freshwater della arcas wilh riverine mangrove Thesc results suggesllhal high year-round produetion swamps, submerged freshwaler aqualievegelalion, and fresh waler marshes. In lhis areas, organie carbon in river input providcs an important addilional food souree; in Terminos Lagoon is maintained by sequemial pul ses of different primary producers, lhis seasonal programming being one of lhe funclional processes su.slaining high estuarine produclion. Y'\:';EZ-AHA:-:CIBL\, A, I sÁ:-:Cm:Z-GIL y A.L. LAHA-DOMí"lGU:Z 168 ! ", / 41 3,. 140 i N 120 100 :;¡ • o. N 35 e "!! e . 2' 25 :. 60 40 iD • :;! o 25 } : 15 15 ¡¡ 2,. 35 e o : 80 ce 1-650 'E '-_._'..¿00!. . O o i,.'" {. .-o 0__0/;' ._\- -/;;/_. tI .--- ¡;; • J J ---a A S O N O J F "'550 o ¡¡ , c: 8 • d. 1-250 ; . e • A 1-350 01 .g u M 450 , a « o M o 30 o o 25 .f 20 15,[Rainy s9a11onL: ..,._ ... _....i-...Nol1es...ea.onILDry 5eaMln Fi¡;un.' 3. Sellsonal patterns uf pl"imary production and p1ant hionmss uf the dilTcrent functional groups of primal)' produ(cr in Tcrminos La¡;oon. The illustration ahove the graph are common rt'presentuth"cs of dilTcrcnt primary producer J.:rollp (1cft to riJ,(ht) hlack (Av;cenn;a germ;nans), and red (Rhizophora manxle) mangroves; t\-\-'o genera of diatoms (Chaelocero.. Skelelonema), an imporlant ropus of cstuarine phytoplankton; Iwn spl'cies of fresh\-\-'ater suhmered aquatic "'cctation, Já1li.fneria aml'ricana and Cabomha palaeformü; and the dominant marine s('arass of the 1a()on. ThalaS.\'ia lesludinum. The decrease uf salinil) (helo\-\-') durin the rlliny and "norte s" seasons corrcsponds to the perilld uf hih riHr no\-\-'. And high input uf riH'r hurne orJ,(anic matter (From Rojas Galaviz el al., 1992). FU1lc/i01li1lg O/macro-l'oflsumers The primary produeers diseussed above and tbe dynamics of the ecologieal systcrn surnmarized in earlier scctions, sustain a high diversity of nektonic speeies. There are clear fish-habital relationships and a sequential use of the gradients in various time/spacc seales (Yáñez-Arancibia and 11ra-Domínguez, 1988; Y,íñez-Araneibia el al., 1988, 1993a). More Ihan ISO fish speeies in different funelional groups occur in Terminos Lagoon (Yáñez-Araneibia el al., 1980) and Ihere arc morc than 250 spccies in Cifferent funetional groups from Campcehc Sound, i.c., the adjaecnt continental shelf (Yáñez-Araneihia el al., 1985; Sánehez-Gil and Yáñcz-Arancibia, 1986). The widesprcad use of estuaries hy larvae and juveni les of so many speeies has Ict 10 thc coneept of Sound hahitats for refuge, reproduetion, fceding, recruitment, nurse!)', and growth for a numbl:r of enastal fishes. In this papcr we consider recruill1u:1ll as tlle key factor that scparatcs thc "estuarinc dependent" from "cstuarine-related" (or cstuarinc opportunistic) lish specics. For cxamplc, for estuarine. dcpcndent spceics, juvcnile habitats arc spatially scparate and struetural\y ditlerCnI from thc habitat 01 thc adults and the early the stagcs (eggs and laryae). Fish asscmblagcs or flme/lonal grollfJs are very important in maintaining the structure and functioning of eonsumcrs. At Ieast three groups 01 fishes o<:<:ur in the lagoon-estuarine system (Fig. 4) as diseussed by Pauly and Yáñcz-Araneihía (1993): 1) Reside1lt species: Those which spend thcir cntirc lifc l)'cle within the systcm; "cstuarine dependencc", implying that the estuary is 2) Seasolla/ migraflls: Those whieh entcr the lagoon required for somc part of the life cycle of thcsc during a morc or Iess wcl\-dclined scason (from organisms, Thcrc is consistcnt cvidcnec of thc cithcr the marinc or thc frcshwater side) and leave eeologieal valuc ofTcrminos Lagoon and Campcehc it duringanother scason; ECOLOGICAL FCCTIOAL GROCPS. 169 3) Occasional 'isitors: Those which cntcr and Icave on the habitats inside the system. Here, the rciativc the systcm without c1car pattero within and among level of rccruitrncnt ioto the lagoon-cstuarint.: s)'stcm is determined locally by the ease with whieh fish can years; sequenfially use the habitats, and between years, hy To Ihesc, two othcr groups maybe added: the over1I number of potential remJiIs along Ihe mas!. ).Itarine. estlliJrine-reÚl/eJ speci£s. Thosc which spcnd Ihcir entire life c)'de on the inncr sea shelf under the estuarinc plurnc ¡ntluence; and For the eoastal fish, randomly spawning on the shell' aojacent to the lagoon-cstuarine systm and Icltiog the juveniles find their way ioto the lagoon would be 5) Fresh water, estuarine.related species. Those which ineffieieot (Yáñez-Araneibia el al., 1993e). RaIher, spcnd Iheir entire life c)'de in Ihe fluvial-deltaic seasonal variations of abiotic pararnctcrs. scasonal rivcrinc zonc, associatcd with thc upper zone of the estuarinc system. One meehanism whieh allows high standing stock to be maintained in the lagoon-cstuarine ecosystcm is small-scak bctween-habitat migration. The funetional groupwhieh mainly exists in the fluvial-deltaie riverine ehangcs of primary prnduetion and of eompclitors must be accommndated, and this is whaI Ieads to the phenorncnon labclcd ht:re as "seasonal prograrnrning" (of eonsumers) which refers to the temporal and spatial scquence oflagoon-habitats used by juveniles and preadult fishes (Fig. 5). Fish migration also allows optimum utilization of prirnary production we havc zone of the estuary (Fig.) utilizcs the estuarine system found that fish tend tovisit diffcTent habiIats during mainly as a feeding and nursery ground. This pallero periods of peak primary produetion (as sho\V in Fig. is similar for lhe funetional group in lhe inncr shelf. 3). This ensures high food availability "nd Ihus high '\be reemitment of these functional groups depend fish growth rates. 'UH(;TIONAL GROUPS " Marln •• "luMin... laI-.cI sp.cin ... . $y..,/um flutlfffl A UPMVu 5p_nlnll """"" z li. Truhuru ytJwmi S"_I c¥"inu OCcilJlonill Vi.llor. c::> E Archo...ll'US rhombold.Jil Orthoprlril chrylOP'HiI HililmuIon iluroNfMilwm ,'J A .. id.nl.ptIci •• lIIJIII!IIn>- SMrdie/lil chrysowil 8i1Írd,.¡y ronchu A"",", ".sllwat.' ... lua.;n .....UUd .p.t<i... ft • 8 . o • i • • .".. ... '....;0.. 1 yo....... DorOI_. Ck¡'¿'l......, '_,fTiI""" \;: ,.",twNiI s¡WndI<b s._rr" m"ll'''ts 8"ll'ft-v..1 eyr,olCIo<> ...........s s,..,., lanceoJ.alus C_lJt""" f'dennn'ul S"i"9 adutts ...,1 !lilbitets fudnor- g<.Q lildults. dry susonl Anus ,"s lildults. _ "ilIOn) fiJ;:ure 4. DisJ:ramstic represenlalion of fundional groups 01 Ilshes "'"llh characleríslic migralion pallerns and hahitat utililalion ","ithin. oulside of. and inlo and oul of Terminos Laoon. The funclional roups are defined in (he lexl (Front IJaul)' and Yáñez.Arancihia. 1993). YASEZ-AR:\:\'CIBIA, A.. P. &\:'\CHEZ-GIL y A.L. LARA-DO\líi\GUEZ 170 100 Diapterus rhombeus SO O .- Sphoeroides testudineus 100 SO O 100 SO O DiO 100L SO O '"> Bairdiella ehrysoura = Jl ==....._ Anchoa mitehil/i _===CL=== 100 '" = Petenia sp/endida SO '" U l: lO O "C l: eieh/asoma urophtha/mus il 100 « SO O Eueinostomus gula 100 SO O s Arius me/anopus 100 [ SO r=J= O I [l SO 1 J F M A M J JASOND Monlhs Figure 5. Scusunalil)' of ahundance of eighl dominant lish species in T... nninos Lagoon. These fishes show a c1e:tf se;lsnnal S('(jUl'nt:c uf nbundancf' and habitat utilizalion, t:allcd herc "s("asona¡ proramming". The sum of the rclalhc ahundance u( (hese .. pecit.", (Iuwcr J.:raph) sUAAesl 3n neo ulilizalion of (he IUJ.:oon.estuarine s)"stem throughoul the )'('ar. Afler Yáñel' Arandhia el al. (l993c). i AII speeies 100 o I 1 f I E n ECOLOGICAL FL:"CTIO:"AL GROLPS. Synthcsis Tropical coastal ccosystcms havc maintaincd Ihemselves over very long periods. The specifie location of the <""035t has changcd a" sea leve) rose and fell, but Ihe eeosystems have a long conlinuity. This 171 Day,.Ir" .I.W. and A, Yáñez-Amncihia. 1982. Coastal lagoons amI cstuarics: ccosystcm approach. Ciencia Illteramericalla Washington DC, 22( 1-2): 11-25. Day, .Ir".I. \\é, R, H, Day, M, T, lIarreirn, F. Le)'-Lou and e..I. Madden, 1982, Primary produclion in the has allowcd the cvaluation of rncchanisms which L.1b'Una de Terminas, a tropical estuar)' in the Southern stabilizc amI refine such seasonal prograrnming, Gulf of Mcxieo. Coastal Lagoons. l Lasserre and Ir. making the fish populalions gradually more Postma. (Eds). Oceall%gica Acta Vol. Spee. "depende nI" on the lagoon s)'stem for the maintenancc 5(4):462 p. of high biomass and sccondary production. \Ve believe that understanding biodiversity from 'In ecological point of view, e.g. the funetional role of estuarine gradients and hahitats, willlead to a belter understanding of tropical coastal zones and is indeed the basis of their sustainahle devclopment. Along tropical coasls, only holistic habitat managcrncnt approaches appcar to he adequate (Fones, 1991; Buttriek, 1992; Ray, 1991). The approaeh for intcrrclating resourccs flows, socioeconomic intcractions and habitat gradicnts on traosect of mast1ines proposed hy Pauly and Lighlfoot (1992) may also contrihutc lo this. Day, .Ir"J,'y', W.ll. Conner, E Ley-Lou, R.H, (Ja) and A_ Machado Na,'arro, 1987. The produclivity and eomposition uf mangrovc forcsts, Laguna dc Terminos, Mexico.Aq/latic BOlaIlJ, 27: 267-284. Day, .Ir., .I.,y., e..I, Madden, E Ley-Lou, R.L. WCI1e1 and A, Machado, 1988a. Aquatie primary productivity in the Terminos Lagoon. In Yáñcz Araneibia, and J.W. Day (Eds.) Ec%,'Y 01 cou.wal ecmystems in Ihe sou/hem GlI/f01Afexico: Ihe '1eJ1lli"os Lagoon regioll, p. 221-236. UNAM Press México, D.!' Da)', .Ir".I.W., W,H. Conner, R, 11, Day, E Le)-lAIlJ, and A.Machado. 1988b. Produetivity and composilion 01' mangrovc forest al Boca Chica and Estero Pargo. ln Yáñez-Arancihia, A and J.w. Day (eds.), Ecu/ol-. oI Acknowlcdgrncnts \Ve acknowlcdgc Prof. P. Lasserre, France, amI Praf. Sean \V Nixon, USA for inviting us to participatc in the lntcrnational Workshop on Ecosystcm Funelioll oC Marine Biodiversity in Estuarics, Lagoons and Near-Shore Coastal EeOS}lemS, he Id in Guaddoupe from Mareh 1410 19, 1993. We also acknowledge the supporl provided by lhe IUllS-SCOPE-UNESCO Programme on Biodiversity, the hOSI facilities ofthe eoas/al ecos)'stems in tire sowhem Gul!of,\ferico: lhe Tenllillo" Laliooll regioll, p. 237-257. UNAM Press Mexien, D.E I>ay .Ir" .I"y., e.A.S.Hall, W.t. Kemp and A. Yáñe1 Arancibia, 1989, ütllarine Ec%gy. Jolm Wiley & Sons,lne. Fortcs, M.O. 1991. Seagrass-mangrovc eeosystcms management: a key to marine eoastal conscrvation in thc ASEAN Region. Marille ['ol/lItion /J/ll/etin, 23: 113 Universite des AntiBes et de la Guyane, Dr. Danid 116. Pauly (ICLARM) and Dr. Jnhn W. Day for many Grassle. J.F.. P. Lasscrrc, A.D.. l\Ic1nt}Tt.' and G.C. helpful enmments on the paper. CONACYT from Ra}'. 1991. Marine hiodi"crsity andccu"lysH:m fUllction. Mexico (Projcet 1'467-19190) also provided lravel A proposal for an international program of rcsean.:h. support. Bi%gy I/l/ematiolla/. Specia/I.u/le, 23. lUIlS, SCOPE, UNESCO. P 1-/9. Litcraturc Citcd IOC-UNESCO. 198. IOC-FAO Guiding group of expcrts nn the pr0,b'Tam of Occan Sciencc in Rl.:lation Arreguín-Sánchel. E, EA. Chá\'ez ,md .lA, Menchaca. lo Living Resourees. Firsl Session. IOC Repons of 1992. A multispeeies stock assessment of pelagic Meeting of Expcns and Equivalent Bodies, Paris, 16 eoastal fishery of the south-west Gulf of Mexico. 20 July, 1984. SC-85/WS/18. p. 1-20. Aquacullure and Fisheries lvtanagemenJ, 23: 1 03 112. Kjerf,'e, B ..I. 1989. Estuarine geomorphology and physical occanography./ll: J. W Day, Jr., C.AS. Hall, lIutlrick, S.e. 1992. 1 lahilal management: a deeision WM. Kemp and A Yáñez-Araneihia (Eds.), E.'t/lllline making process. IUlOdora, 9"«879):258-286. ec%l{}', p. 7-78. John Willey & Sons Nc\V York. YÁ\;EZ-AR,c'\CIBL\. A.. I SACIIEZ-GIL y A.L. LARA-DmlíGUEZ 172 Kjerf\'C, BJ. (Ed.) 1993. COlIItallagoon processeI. Mexieo. In: P. Lasserre and I I. Postma (Eds.) Coastal Elscvicr Scicncc Publishcrs. Amstcrdam, Thc lagoons, Oceanologica ACIa. 5(4):431-440. Nelhcrlands. Yáñez.Arancibia, A .. Sánchez-Gil. 1\1. Tapia and Lasserre,l 1992. Thc role ofbiodívcrsíty in marine M.C. Garcia-Abad. 1985. Eeology, eummunily ccosystcms. In: O.T.. Solbrig. H.M. van Emden and structurc ami evaluation of tropical tlcrnersal fishes P.G.WJ. van Oordt (Eds.). BiodiverIity ami glohal in the soulhern Gulf of Mcxico. Cah. Biol. Mar. n'ance. c/umge. p. 105-\30. International Union of Biological 26(2):137-164. Seicnecs. Monogr. 8. Paris. Yáñez-Arancibia. A. 1987. Lagunas costeras y t:\ Lonhurst. A. and D. I'aul'. 1987_ Ecology 01tropical tuarios: c...Tonología, criterios y conceptos para una oce!llls. Acadcmic Prcss, Ine. San Diego. clasificación ecológica de sistemas costeros. Rt'\: SOCo Odum, W.E. 1988. Comparative ecology uf tidal frcshwatcr ami salt marsht:s. Antlll. Rel'. Ecol. SYS1. Mex. Hist. Nat, 39:35-54. Yáñez-Arancibia, A. and J.W. Day Jr. IEds.) 19XX. Ec%gy of coastal ecosystems in the sow/zem Gulf of /9:147-176. P'IUI. D. and C. Lihtfoot. 1992. A nc\\' approaeh for analyzing and comparing coastal rcsaurce systcms. Mexico: the Terminas Lagoon region. UNA..1 Prcss Méxieo,D.E Yáñez.Ardncibia, A. and A. L. Lara.Domíngucl. :VAGA the ICl..ARM QlIarterly. 15(2):7-10. Pauly, U. and A. Yáílez-Arancibia. 1993. Fisheries in coastallagoons.lnIlJ. KjerlVe (Ed.), CoaIw{ lagoom processes, p. 352-372. Elscvicr Scicncc Puhlishcrs, 1988. Eeology of lhree sea eatfishes (Ariidac) in a tropical coastal ecosystcm-southern Gulfof Mcxico. Mar. Ecol. /'rogr. Ser, 49(3):215-230. Yáñcz.Arancibia, A., A.L. Lara-Domínguez. J.L. Ro Amsterdam, The Netherlands. Ray', G.c. 1991. Coastal-zone biodiversity patlerns. BioScience. .¡ 1 (7):490-498. jas Galaviz, P. Sánchez-GiI, J.\\'. Da; Jr. and C. j. l\Iadden. 1988. Scasonal biomass and diversily of estuarine fishes eoupled with tropical habilal Ra. G.c. and I\I.G. MeCormiek-Ray. 1992. hetcrogcneity (southcrn Gulf of Mexico).1. Fi,h Bio[ Funclionai coastal-marinc biodivcrsity. "uns. 57th 33 (S/lpplement AI:I 91-200. N.A. Wildl. & NiII Res. Conf. p. 384-397. Yáñez.Arancibia. A., P. Sánchez-GiI and A. L. Rojas Galaviz,j.L.. A. Yánez-Arancibia,J.\\: Uay, Jr. LaraDomínguez. 1991. 1 ntcraceioncs ecológicas and E Vera Herrera. 1992. Estuarine primary estuario-mar: Estructura funcional de bocas produccrs: Laguna de Terminas a study case. In: U. cstuarinas y su efecto en la productividad dd Sc.:cligcr (ed.). Coastal plam commlllJities o[ Latin ecosistema. Academia de Ciencias de Sao Pllulo. Pub/. America, pp. 141- 154. Academie Prcss, Ine. New York. AClES/',71(4):49-83 S,ínehez-GiI, P. and A. Yáñez-Arancibia. 1986. Yáñez.Arancibia,A., A.L. Lara.Domíngucl and J.\t Discusión sohre relaciones tle reclutamiento en Day. Jr.• 1993a.lntcractions hctwccn mangrovc ano sistcmas lagunares-cstuarinos de los rx:ccs tlcl sur dcl seagrass hahitats mctliatcd hy cSluarinc Ilckton Golfo de México. 111: Yáñez-Araneibia, A. and D. assemblagcs: coupling uf primary and scconJary Pauly (Eds.), IOC/FAO Wor"-'hop on recmitment in produetion. Hydrobiologia, 26-1: I -12. tropical caastal demersal commlllJities. pp. 215-228. IOC/UNESCO Worksbop Report No. 44 (Supplement), Paris. Yáñez-Arancibia, A.• E Amezeua and J.W. Day, Jr., Yáñez-Arancibia. A_ (and other 20 authors). 1993b. Estudio de declaratoria como área ecológica de protección de flora y fauna silvestre y acuática lit' la región de la Laguna de Términos. Campeche. 1980. Fish eommunity struelure and funetion in Programa EPOMEX, Convenio SEDESOUUAC lcrminos Lagoon, a tropical estuary in the Gulf of p.I-259. Mexico.ln V Kcnnedy (Ed.). ESllla'úJe perspecti\'es, p. 465-482. Academie Press, Ine. New York. Yáñez-Arancibia, A., A.L. Lara-Uomínuez and lJ. Pauly. 1993c. Coastal1agoons as fish habitals.ln B.J. Yáílez-Arancibia, A. and J.W. Day', Jr., 1982. Kjerfve (Ed.). Coastallagoons proceIses, p. 339-351. Eeological cbaraeterization ofTerminos Lagoon a Elsevier Seicnee Publisbers, Amsterdam. Thc tropical estuarine system in the soulhern Gulf of Nethcrlands.