CocMl Lopm Pr'Oeeuu edited by B. KjerfYe(EJ..seyjer Oeeanocraphy Series. 60)C 1m Elsevier Science Publisben B.V. All richu retelTed

Chapter 12

Coastal Lagoons as Fish Habitats

Alejandro Yafiez-Arancibia-, Ana Laura Lara Dominguez- andDaniel Paulyb

-Programa de Ecolog{a, Pesquerlas y Oceanograf£a. del Golfo de Mexico(EPOMEXJ, UniversidadAuttSnoma de Campeche, Apartado Postal 520,Campeche 24000, Campeche, Mexico;blnternational Center for Liuing Aquatic Resources ManagementaCLARM), M.C. P.O. Box 2631, 0718 Makati, Manila, The Philippines

363

'Many fish species use coastal lagoons as habitats during at least some part of theirlife cycles. We discuss these characteristics with emphasis on latitudinal differ­ences. Higher biomass of coastal fishes will occur when lagoon-estuarine habitatspace is available and accessible. The concept of seasonal programming describeshow juveniles use that lagoon habitat space. We also consider the lagoon~stuarine

dependence of subtropical and tropical coastal fishes.

Introduction

Coastal lagoons are defined in geomorphological terms (Phleger, 1969;Lankford, 1977; Kjerfve, 1986; Yanez-Arancibia, 1987). In this, they differgeologically from estuaries, which are defined in terms of the hydrologicalconditions and gradients of salinity and temperature (density) (Cameronand Pritchard, 1963; Pritchard, 1967; Fairbridge, 1980; Yciiiez-Arancibia,1987; Kjerfve and Magill, 1989). Kjerfve (1989) proposed a new classifica­tion for "inland waters which are connected to the sea-, based on six classes(estuaries, coastal lagoons, fjords, bays, tidal rivers and straits), defined byvarious combinations of physical and geomorphological processes.

Kjerfve's assignment of adjacent classes to estuaries and coastal lagoonsis due to their sharing numerous characteristics. This is also the reason whyDay and Y8.fiez-Arancibia (1982) had earlier proposed the (ecological) con­cept of "lagoon-estuarine environments-, Le. shallow semi-enclosed waterbodies ofvariable volume, connected to the sea in a permanent or ephemeralmanner', with variable temperature and salinities, permanent muddy bottoms,high turbidity, irregular topographic characteristics, and biotic elements.

364

In this chapter we present the fish fauna occurring in various lagoon­estuarine environments, with emphasis on those features of lagoons thatare actually 'used' by the fish populations, and on some of the mechanismsinvolved. therein.

We concentrate on fishes because they generally constitute the over­whelming bulk (up to 99%) of the lagoon--estuarine nekton, both in terms ofnumbers and biomass (McHugh, 1967). The few nonpiscine members of thelagoon--estuarine nekton are squids, portunid crabs, penaeid shrimps, rep­tiles, and mammals such as dolphins.

Much information is now available on the life cycles of fishes in relationto lagoon--estuarine environment (McCleave et aI., 1984; Yanez-Arancibia,1985; Dadswell et aI., 1987; Weinstein, 1988; Henderson and Margetts.1988). and various schemes have been proposed to link these cycles withenvironmental parameters and the observed high abundances of fishes inlagoons and estuaries (Day el aL, 1981; Pauly, 1982; Weinstein, 1982;Yanez-Arancibia, 1986).

Two important points have emerged from these studies:- the utilization of the lagoon-estuarine environment is an integral part of

the life cycle ofnumerous fishes, particularly so in the Neotropics and theIndo--Pacific (Longhurst and Pauly, 1987);

- the lagoon--estuarine environment is mainly utilized by juveniles andyoung adults.We shall here consider: (1) the fishes that 'use' coastal lagoons; and (2) the

features of the lagoon habitat(s) that are actually being used by these fishes.

Which Fishes Use Coastal Lagoons?

Although the exact species and dominance ranks change between sites,the dominant fishes of lagoon--estuarine habitats usually belong to only afew taxonomic groups. High latitude estuaries have simple ichthyofaunas,dominated in the northern hemisphere, by SaImonidae (salmon and trout),Osmeridae (smelt and capelin), Gasterosteidae (sticklebacks),Ammodytidae(sandlance), and Cottidae (sculpins) and their relatives (Dadswell et aI.,1987; Day et al., 1989). In the southern hemisphere the SaImonidae arereplaced by their ecological analogs, the Galaxioidae (McDowall and Eldon,1980; McDowall and Frankenberg, 1981). Important families for temperatesystems are theAnguilIidae (freshwater eels), Clupeidtu (herring-like fish),Engraulidae. (anchovies),Ariidae (marine catfish), Cyprirwdonti.dae (killifish),Gadidtu (cod), Gasterosteidae (stickleback), Serranidae (basses), Sciaeni­cku (sea trout, drums, croakers), Sparidae (sea-breams), and Pleuroneetidae(flounders) (Haedrich,1983; McCleave et al., 1984).

In the tropics and subtropics, the important families are the Clupeidae,Engraulidae,Ariidae, Scuumidae,Poecillidae (guppy), Chanidae (milkfish),.

365

Synodontidae (lizardfish). Belon-idae (silver gars), Mugilida.e (mullets), Poly·nemidae (threadfins), Gobiidae (gobies), Cichlidae (tilapia and related fishes),Dasyatidcu (rays), Telraodontidae (puffers), Gerreidae (mojarras), £eiegna­thidae (slipmouth, ponyfish), Pomadasyidae (grunts), and various familiesofflatfishes, such as theBothidae, So/eidae and Cyrwglossidae (Pauly. 1982;yanez·Arancibia, 1985, 1986; Lowe·McConnell, 1987).

Several authors have pointed out that there are a greater number of fishspecies in tropical and subtropical lagoon--estuarine ecosystems than incomparable temperate or boreal systems (Yaftez-Arancibia, 1985; Lowe·McConnell, 1987; Yafiez·Arancibia and Sanchez-Gil, 1988; and Fig. 12.1).However, some large high-latitude estuaries, such as Chesapeake Bay,Narraganset Bay and Delaware Bay, in the northeast USA, also have alarge number of fish species Way et al., 1989). The theory of island bio­geography (MacArthur and Wilson, 1967), which relates species number tothe size of habitats could possibly provide a framework for these apparentexceptions;

'0 A

'0- '"•~, '0z

COlodromousspec:les

B

Marine hoblTOTS

-r~l.000l'.

~E 000

\;~ "'0~

~ ..oo

K~ >00

.~ ~-=~~.g:::~--:;;--;;:~~~ oL 10 20 30 "0 !lO 60 70 eo

Lotitude (ONorS)

Fig. 12.1. Latitudinal trends averaged by 5·'atitude (after Gross n aI. 1988). (A) Number ofanadromotU and eatadromous species occ:urring in COastallagOOllS. (.8) Primary production oflimnic: and marine habitats.

366 Coa.tai Lapotu eu FWi Habitats

As for the general pattern, i.e. that low latitude lagoons have more speciesthan those in high latitude, it could well be explained in terms of the cost ofosmotic regulation relative to total metabolism, which is much lower at hightemperatures.

Various classifications have been proposed to analyze the fish communi­ties in Jagoon-estuarine environments. As they involve some form ofdiadro­mous migration, we briefly define:

Diadromous: fishes migrate between fresh and marine waters.Anadromous: fishes spend most of their lives in the sea and migrate to

freshwater for breeding (e.g. salmon and shad).Semi·aruuiromous: fishes do not travel all the way to freshwater, but

spawn in low-salinity lagoon--estuarine waters.Catadromous: fishes spend mostoftbeir lives in freshwater and migrate to

the sea for breeding. A well-known example are eels oftbe genus Anguilla.Amphidromous: fishes migrate from freshwater to the sea, or vice versa,

not for the purpose of breeding, but regularly at other stages in their lifecycles, often for feeding.

McHugh (1967) identified the following groups:(1) Freshwater fishes that occasionally enter brackish·water.(2) Truly estuarine species that spend their entire lives in estuaries.(3) Estuarine-marine species that use the estuaries and lagoons pri-

marily as a nursery ground, generally spawning and spending muchof their adult life at sea, but often returning seasonally to estuariesand lagoons.

(4) Marine species that pay regular seasonal visit to estuaries andlagoons, usually as adults looking for food.

(5) Anadromous and catadromous species in transit.(6) Occasional visitors, which appear. irregularly.Variants of this scheme exist (pauly, 1975; Yanez·Arancibia and Nugent,

1977) and they tend to emphasize the role ofmigration and other movements.Thus, Miller and Merriner (1985), based on Cronin and Mansueti (1971),pointed out that marine ('sport') fishes exhibit, five distinct life histories orpatterns of migration for habitat utilization: (1) inshore anadromy, referringto species which migrate to spawn in estuarine habitats and move towardcoastal waters as developing juveniles or adults; (2) inshore catadromy, refer­ring to species which migrate to spawn in offshore habitats and toward inshorehabitats as developingjuveniles; (3) inshore nonmigratory, referring to specieswhich utilize estuarine or nearshore coastal habitats at all stages; (4) offshorenonmigratory, referring to species which utilize ocean habitats at all stages;and (5) anadromous, referring to oceanic species which migrate to freshwa­ter to spawn while juveniles migrate to the ocean as smolts. It is understoodthat some of the nonmigratory inshore and offshore species or stages mayactually undertake substantial seasonal migrations or movements alongthe coastline. Fig. 12.2 presents a conceptual model of the stimuli involved.

Light. Temperature,Prey Distribution

Habitat:

physicalProcess:

FIshes'ActivIty:

FIshes'BehavIor:

Stlmull:

, ,r---Plankton----J;--Nearshore---+!-Channel or Mouth-l-Estuary or Lagoon.....

I. ' ...1i! ! ..' ,

Ekman Transport, Ii ..··..Counlefcurronts, ! Alongshore Currenls i Tidal Flux

'''1;'' I. i A/-----------: --: ., -....

j 6oontal~ 10 r--"'~.-.-,Drifl +_ L~90on-EsIUaIY-----...T~~~~~:~m ..... _ -~- ~~~~~:~

r ;oaVertical Movements _l_~ Circadian • __ .1_ ..... : Circa~dal H:lbilat Seloction

f.

I A,,;." rR,,,m. ! A';~."r' rl : Tidal Zeilgeber Habitat Slruclure: )0- Diel ! (Salinity, Temperature. (Covar. Depth.J Zeilgcber ! Turbidity, Pressure. Substrale, Proy.: : OUnclOry Cues. Turbulonce,! i Electric Fields) Conspecilics): : .'.. . .: : '.

I---------AppetitiveBehavior---------.......~

wFig. 12.2. Conceptual model of the slimuli and behaviors involved in fish movements towarns estuaries and coastnllagoons (after Doehlert ~

and Mundy,198B),

368 Coastal LosooTl! as Fish Habitau

Recently, Zijistra (988) based on Louis and Lasserre (1982), presented aclassification consisting of six types, pertaining to different regions rangingfrom temperate to tropical climates and which may therefore have a world­wide application: (1) anadromous and catadromous species; (2) residentspecies spending their entire lives in the coastal areas; (3) marine species,spawning in the coastal region and using it as nursery ground; (4) marinespecies, which use the coastal area as a nursery area, but which spawn andspend most of their adult life in the offshore region; (5) marine species,which pay regular seasonal visits to the coastal zone, usually as adult; and(6) adventitious visitors, which appear irregularly and have no apparentestuarine or coastal requirements.

Real fishes, however, will not adhere strictly to any of these classifica­tions. Rather, a number of populations will belong to several types, inde·pendent of how well-conceived the types are.

More interesting are classifications that are quantitative, even if thisimplies that they become site-specific in the process. Deegan and Thompson(1985) provided such classification for the Mississippi delta. Similarly,Yaiiez-Arancibia et al. (l988b) proposed a classification scheme for Termi­nos Lagoon, Southern GulfofMexico, which provides salinity ranges for thevarious groupings, i.e.:

freshwater spawners, occurring in waters < 10%0.brackish water groups,limited to 10-34%0.marine spawners, occurring in waters ~ 35%0.

Classifications of this type can be constructed, more or less rigorously, forany desired level of resolution, using clu~tering-typetechniques (i.e. usingstation species lists vs. observed environmental parameters; Fig. 12.3)andlor the results of physiological experiments, e.g. on salinity tolerance.

Fish production in lagoon-estuarine systems is generally high (Day et al.,1989), and this generally leads to high fishery catches. One reason for thishigh production is that many of the fish species of importance in lagoon­estuarine environments are either: (1) r-selected sensu (Pianka, 1978), i.e.relatively small, fast-growing animals with high productionJbiomass (PIB)ratio; or (2) K-selected juveniles, i.e. fast-growing, high production stagefishes whose adult form, however, typically occurs outside the lagoon. Thehigh PIB ratios of the constituent species are not the only explanation forthe generally high production of lagoon-estuarine communities. The highbiomasses themselves, i.e., the high carrying capacities of these ecosystemsin terms of food availability to fishes also playa crucial role.

370 Coo.stal LcgOOM Q$ Fish Habitats

Which Features of Lagoon-Estuarine Habitats Do Fishes ActuallyUse?

The two most important processes for living organisms are:(1) energy acquisition, i.e. feeding, as needed to maintain their struc­

ture, and to grow;(2) reproduction: only those organisms are still around whose ancestors

bred and who endowed them with ability to do the same.Except for very few species, fishes do not use lagoons for reproduction

proper, i.e. for late maturation, spawning/fertilization. It is tempting torelate this to the ephemeral nature and the strong environmental variabil­ity of these systems. Rather, it is as feeding grounds that lagoon-estuarineenvironments are important to fishes.

The growth/natural mortality schedules of unexploited fish cohorts gen­erally imply that biomasses are maximized near the mean age at firstmaturity ofthe individual fish, an adaptation obviously related to maximiz­ing the output of reproductive products. However, small fishes always eatmore per unit body weight than large fishes (Pauly, 1986). This results inthe maximum of the absolute food consumption curve, of any stable fishpopulation being shifted to the left, toward the juveniles (Fig. 12.4).

Thus, it is the juveniles of fish populations, and not the adults, whichexert the greatest pressure on the food supply. If a coastal population (B')of adult fishes is to maintain itself, it must place its juveniles in a habitatwith a carrying capacity »B'. This point, apparently not made in theextensive literature on lagoon and estuaries, provides an objective criterionfor referring to these water bodies as critical habitats.

However, for any coastal fish population, randomly spawning on the shelfadjacent to lagoon-estuary systems and letting the juveniles find their wayinto these systems, just does not happen. Rather, seasonal variations ofabiotic parameters and of food supplies and competitors must be accommo~dated, and this is what leads to the phenomenon labeled here as seasonalprogramming, which refers to the temporal and spatial sequence of lagoonhabitat uses by juvenile fishes (Yaiiez-Arancibia et al., 1988a).

Seasonal programming of food resource utilization by various species inTerminos Lagoon is illustrated in Fig. 12.5. Various species use the lagoonat different times, thus reducing their competitive interactions. At the levelof a given species, seasonal programming implies that its representatives(mainly juveniles, see above) feed, in the course of their ontogeny, on asuccession of different food types, often gathered from different subareaswithin a given lagoon (Fig. 12.6). This sequential utilization oHoOO type andfeeding location may be highly repetitive, hence predictable, between seasons.

In areas where lagoon systems have maintained themselves over longperiods, evolutionary mechanisms may emerge which stabilize and refinesuch seasonal programming, making the fish population in question'gradu~

A Ydlln·Atundbio., AI.. Lmu Dominguu and D. Paul, 371

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Fig, 12.4. SchematierepresentatioD oflhe growth and decayofa cohort whose growth (A) canbe depicted by the von Bertalanfl'y growth equation, and whose mortality by a negativeexponential {B}. In this ezample, cohort biomus eC) peaks at age t...u(n= 1 yelU', which is alsothe are at which. flnlt maturity is achieved (W... 0.3 W..). Curve D depicts the decline ofrelative ration. commonly expressed as'" body wei,ht per day' or'J'BWD. The product. ofeurveD and curve C (E) shows the predatory impact ofthe cohort, by age. M might by seen, tlDU(%)

< t-..(n i.e.. it is thejuveniles oia cohort which con.sume most ofthe food nquired by a cohortofflShe!.

372

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Fig. 12.5. Seasonalityofabundance ofeight dominant fish 5pecies inTermiDOfI Lagoon, MWoo.These fishes &how a clear seasonal sequence of abundance and habitat utilization calledseasonal programming. Note that the sum of the relative abundance of these eight speciesOower graph) suggests an even utiliution of Tumin05 Lagoon throughout the year (aft.erYAiiu-Aranc:ibia d oJ., 1988a).

A Ydllu-Arandbia, AI.. L4ra Domingu.a and~. Pau.ly

OnlOQenic chonQu

373

LOQOOn/esluory

SeosonolchonQet

<. '- IDiurnol_ ..,:, chonQu.~ '-

"So.

-----"" Y,," ",de.'-----Y consume.

-'- seC.,nd o,de'-., conSllmo,

-----Irr. Tn;.d ."de,----,. c.,nsllme,

Fig. 12.6. Conceptual diagram illustrating the trophodynarnic eetllogy of a hypotheticallagoon-estuarine fish species. The elements of time and time of the day, season and fish age(size), and space are shown. The spatial elements include transfers between ecosystem, e.g.,from marine areas via lagoons/estuarine to riverine areas, as well as between habitats ofthesame system, e.g. from oyster reefs to seagrass beds within the lagoon/estuarine system (afterYlii\.ez·Arancibia et 01.,1986).

ally more dependent on the lagoon system for the maintenance of highbiomass.

In other areas, where the lagoon systems do not persist or are notregularly open to juvenile immigrations, e.g. off West Africa, use of thelagoons are a matter of random movements along the coasts, and of aninshore movements, i.e. errances. This implies (1) a lower conversion ofprimary and secondary production into fish flesh, and hence (2) lowerbiomasses of coastal fishes,

The difficulty inherent in practical separation of random alongshore!inshore movements, i.e. errances, from evolutionarily fine-tuned, aimedmovements towards and within lagoons, and the difficulties involved inpreoise field eltimatioo ofbIomass and conversion efficiencies, make rigor­ous testing of these hypotheses difficult. This is the main reason for thedebate on the degree of dependence of tropical coastal fishes 00 coastallagoons.

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376 CoaswI Lagoons a. FisA HabitatS

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