Article 2 - Guidini & Santos-Silva

7/31/2019 Article 2 - Guidini & Santos-Silva

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THE JOURNAL OF THE BRAZILIAN CRUSTACEAN SOCIETY

NAUPLIUS | VOLUME 19 | NUMBER 02 | 2011 | ISSN 0104-6497

THE JOURNAL OF THE BRAZILIAN CRUSTACEAN SOCIETYTHE JOURNAL OF THE BRAZILIAN CRUSTACEAN SOCIETY


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Nauplius 19(2): 109-122, 2011 109

Composition, species richness and patterns of nycthemeralvertical distribution of planktonic cladocereans in a black waterAmazonian lake

Andr Ricardo Ghidini and Edinaldo Nelson dos Santos-Silva

Instituto Nacional de Pesquisas da Amaznia. Coordenao de Pesquisas emBiodiversidade. Av. Andr Arajo, 2936. CEP: 69060-001, Manaus, Amazonas, Brasil.E-mails: (ARG) [email protected], (ENSS) [email protected]

Abstract

Te aim o this study was to identiy patterns o vertical distribution o planktoniccladocerean populations throughout the diel cycle, during the low and highwater periods, and its ecological implications or a black water Amazonian lake.up Lake is a black water lake located near the Brazilian city o Manaus. Achannel links the lake with the Negro River and its ood pulse. Tis study wasperormed in a low-water period (November 2005) and in a high-water period(June 2006). Samples were taken on a 24-hour cycle, every 4 hours and at eachmeter o the water column, using a Schindler-Patalas trap equipped with a 55msize mesh. A total o 16 species were registered during the low water period,wherein Bosminopsis deitersi,Moina minuta, and Ceriodaphnia cornutawere themost abundant species. B. deitersimigrated to the bottom during the aternoon,whileM. minuta,Moina reticulata, and Holopedium amazonicum remained at thebottom or the entire diel cycle. During the high-water period, a total o 18 specieswere observed, and B. deitersi, C. cornuta, and Diaphanosoma polyspinawere themost abundant species. During both sampling periods, no pattern was detectedor C. cornuta. Generally, vertical patterns o distribution were less evident in thehigh water period, due to the mixing o the lake.

Key Words: Black water lake, Central Amazon, Cladocera, Vertical migration.

horizontal scale. Te vertical distribution othe organisms is a direct result o their verticalmigration behaviors.

According to Cushing (1951), thecapacity to perorm vertical movementsthrough the water column, vertical migration,was rst observed in marine zooplanktonby Cuvier in 1817. Even though verticalmigration o zooplankton has been requentlystudied in reshwater communities or the

Introduction

Zooplankton populations aredistributed heterogeneously throughout anecosystem and, as previously reported byPinel-Alloul (1995), they usually exploit theenvironment in patches. In tropical lakes, thisheterogeneity generally occurs seasonally in aspatial-temporal scale and daily in a vertical-


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Ghidini, A.R. & Santos-Silva, E.N.: Cladocereans in a black water Amazonian lake110

last two centuries (reviews e.g. Cushing,1951; Hutchinson, 1967; Lampert, 1989;Lampert and Sommer, 1997) the real adaptivesignicance o the phenomenon is still notcompletely understood.

Zaret and Suern (1976) explainedvertical migration as a mechanism ozooplankton to avoid predation, which isrelated to luminosity, body size and visibilityo the prey. Tese actors cause predators likeplanktivorous sh to prey dierently on theCladocera community (Zaret, 1972; Zaret andKeroot, 1975). Later, Lampert (1989) pointedto light-related mortality as the ultimate reasonor the vertical movements.

In the Neotropical region, studies have

associated vertical distribution o planktoniccladocereans with variations in temperatureand, consequently, oxygen (Arcia-Zago, 1978,Matsumura-undisi et al., 1984; Nogueira andPanarelli, 1997). Perticarrari et al. (2003) pointsto a close relationship between the populationso Bosmina tubicen and Chaoborus sp., aninvertebrate predator, reporting predator-preyinteractions as the main inuence on verticalmigration. More recently, a nocturnal verticalmigration pattern was detected or cladocereansin a shallow tropical reservoir in NortheasternBrazil (Silva et al., 2009). Bezerra-Neto etal. (2009) detected the occurrence o dielvertical migration in a Brazilian reservoir withplanktivorous sh and the absence o verticalmovements in a sh absent situation.

Tere are ew existing studies regardingvertical migration in the Brazilian Amazonoodplain, a location where the ood pulseo Amazon River and its tributaries promoteperiodic changes in all aquatic communities. Inwhite-water lakes, Fisher et al. (1983) did notnd clear evidence o vertical movements, butthey discovered Daphnia gessneris preerenceto remain in deeper layers. Te same wasobserved by Keppeler and Hardy (2004) ormost species o the zooplankton community.Rejas et al. (2007) observed the diel verticalmigration o planktonic cladocereans in aBolivian vrzea lake and detected a strongpredator-avoidance relationship between

zooplankton and Chaoboruslarvae. Regarding

black water lakes, Previatelli et al. (2005)detected cladocereans remaining in the deeperlayers o the water or the majority o the 24-hour cycle. Our purpose in this paper wasto determine the diel vertical distribution o

cladocereans and the uncover the compositionand abundance patterns o this community ina black water Amazonian lake, comparing lowand high-water periods, in order to determinepossible shits in the community and explorepossible causes or these attributes.

Materials and Methods

up Lake is a black water lake locatedin the up Sustainable Development Reserve(Reserva de Desenvolvimento Sustentvelup), 30km rom Manaus, in the BrazilianState o Amazonas. Te lake occupies 67hectares, and its water volume ranges between1.44 and 2.57 million m3 throughout the year(Aprile and Darwich, 2005). A 20m channelconnects the lake with the Negro river and,as with many river-connected lakes, mosthydrologic characteristic variations are relatedto the ood pulse o the Negro River. Telakes depth ranges rom 4 to 15 meters in thecentral region o the lake. According to Raiand Hill (1981), the lakes trophic state shitsrom dystrophic to mid-eutrophic during thecourse o the hydrologic cycle.

Te samples were taken in thedeepest region o the lake (030235,4S -601517,5W) (Fig. 1), during both the lowwater period (lake depth = 4.5m) (November11th, 2005) and the high water period (lakedepth =13.5m) (June 13th, 2006). Troughoutthe 24-hour period, sampling was perormedevery 4 hours. Samples were taken at everymeter o the water column (rom the sub-surace to the bottom) using a 12-literSchindler-Patalas trap. A total o 24 liters werecollected, ltered through a plankton net witha 55m mesh size and xed with ormalin toreach a nal concentration o 6%. Dissolvedoxygen (mgO

2L-1), temperature (C),

conductivity (S.cm-1) and light penetration

(m) were measured using an oxymeter (YSI55


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Nauplius 19(2): 109-122, 2011 111

Yellow Springs), a conductivimeter (YSI30 Yellow Springs) and a Secchi disk, respectively.

Te organisms were counted ina Sedgewick-Rater chamber using anoptical microscope. Final abundance was

expressed as organisms per cubic meter. Inorder to establish a relationship between thepredators in the samples, a quantication oChaoboridae populations in the samples wasmade. Identication o the species was basedon previous studies by Elmoor-Loureiro(1997), Smirnov (1996) and Velho et al.(2000). Correlations between physical andchemical parameters and the Cladocera species(abundance and composition) were testedwith a Canonical Correspondence Analysis,

while the correlation between the populationdensities o cladocereans and chaoboridae wasmade using Spearmans index. Kruskal WallisANOVA was perormed to determine i therewas a dierence between the total densitieso cladocereans considering both the hourso the cycle and sample depths. All analyseswere made using PAS (Hammer et al.,2001). Counter plots graphs were made usingSigmaplot 10.0 (Systat, 2006).

Results

Low water period (November/2005)Te transparency o the water column,

measured using a Secchi disk, was 1.2m.Te temperature was lower in the deeperlayer during most o the day (27oC), with avariation o 4oC when compared to the surace(31oC), indicating the existence o thermalstratication in the lake during this period.Regarding the electrical conductivity, it rangedrom 4 to 8 S.cm-1; lower values were registeredat the surace. Dissolved oxygen presentedhigh vertical variation and its distribution wasstratied during the entire day varying rom7 to 2 mgO2/L, rom top to bottom (Fig. 2).Oxygen saturation was higher in the rst 3m othe water column (80 to 100%) and lower inthe bottom (10 to 15%) o the water column.

A total o sixteen species o Cladocerawere identied. Bosminopsis deitersi andMoina minutawere the most abundant species(around 30% each), ollowed byCeriodaphniacornuta (20%) and Diaphanosoma polyspina(15%); these species represented more than90% o the counted organisms. Chydoridae

and Ilyocryptidae were registered in low

Figure 1.Location o the study area, highlighting the Amazonas State, and the up Lake, with the position o the samplingstation.


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TaxaAbundance (%)

Low High

BosminidaeBosmina longirostris(O.F. Mller, 1785) 1.10 1.26

Bosmina hagmanniStingelin, 1904 0.11 4,25

Bosminac. huaroensisDelachaux, 1918 0.02 0,04

Bosmina tubicen Brehm, 1953 0.14 1.08

Bosminopsis deitersiRichard, 1895 29.21 59.22

Bosminopsis brandorRey and Vasquez, 1989 NR 0.15

Bosminopsis negrensisBrandor, 1976 NR 0.75

DaphnidaeCeriodaphnia cornuta Sars, 1886 19.14 20.72

Ceriodaphniac. pulchellaSars, 1862 0.32 0.01

MoinidaeMoina minutaHansen, 1899 29.81 0.23

Moina reticulata(Daday, 1905) 1.01 NR

Moina rostrataMcNair, 1980 0.11 0.72

SididaeDiaphanosoma birgeiKorneck, 1981 0.09 0.08

Diaphanosoma polyspinaKorovchinsky, 1982 14.30 11.42

IlyocryptidaeIlyocryptus spinierHerrick, 1882 0.07 0.02

ChydoridaeAlonella dadayiBirge, 1910 0.05 0.01

Nicsmirnovius incredibilisDaday, 1905 3.22 0.01

Kurzia latissima(Kurz, 1874) NR 0.01

HolopedidaeHolopedium amazonicum Stingelin, 1904 1.30 0.02

Table 1. Cladoceran species and relative abundance, during the high and low water level periods,in up Lake. NR: not registered.

Figure 2. Variation o dissolved oxygen, during the 24-hour cycle in all depths, in up lake during the low water period(November/2005).


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Figure 3. Vertical distribution oB. deitersi(a),M.minuta(b),M. reticulata(c), B. longiostris(d), and H. amazonicum (e) during thelow water period, in up Lake (november/05). Te number represents aproximate density o organisms/m-3.


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abundances (ab. 1).Signicant dierences were observed

in the total vertical density o cladocereans.(Kruskal-Wallis test: H (4, N = 30) =15,02366;p = 0,0047). Mean density o cladocereans in

the low water period was 6141 org/m3, butgreat heterogeneity was observed among thesamples. Te density o organisms was higherat 2:00 am (mean = 15816 org/m3) and at 3mdeep (mean =11591 org/m3). Te lowest meandensities were observed at the surace (1974org/m3) and in the samples taken duringthe daytime (at 6:00 am = 2178 org/m3, at10:00 am = 2970 org/m3). When comparingthe dierent hours o the cycle, no statisticaldierence was observed (Kruskal-Wallis test:

H (5, N = 30) = 9,366452; p =,0953).In the low-water period, M. minuta,

M. reticulata, and H. amazonicum remainedin the deep layers o the water column, whileB. deitersi migrated rom the surace to thebottom during the aternoon and Bosminopsislongirostris showed a typical nocturnalmigration pattern rom the bottom to thesurace (Fig. 3).

A high abundance o B. deitersi (Fig.3a) during the low water period was observedmainly in the deeper layers o the water column,particularly during the aternoon (2:00 pm),when a peak o organisms was registered.

Te vertical distribution oM. minuta(Fig. 3b) was quite peculiar, since higherdensities were registered at 2:00 am, at depthsaround 3 and 4m, ollowed by an intensedecrease at 6:00 am. Te preerence o thisspecies or layers close to the bottom o thelake was evident. Te same was ound orM.reticulata(Fig. 3c), save any intense decrease.

Bosmina longirostris (Fig. 3d) was theonly species with a peak o organisms atthe surace o the water column. Tis peakoccurred at night; during the day, the organismsremained in deeper layers. On the other hand,H. amazonicum was registered only in deepwaters, specically at 3 meters (Fig. 3e).

Te total number o Chaoborus sp.ound in the samples was similar to thedensities o cladocereans. Teir distribution

patterns were very similar (correlation index

Spearman, D = 0,6067; total cladocereansdensities and total Chaoborus densities, logtransormed), and there was a pronounceddecrease o both Cladocera and Chaoborussp.observed during the daytime (Fig. 4).

Te relationship between the speciesdensities o Cladocera and the physical andchemical parameters measured during the24-hour cycle at diering depths was notdetermined, even ater log transormation.Oxygen saturation was not used in the analysis,because it was correlated with dissolvedoxygen. Te Canonical CorrespondenceAnalysis registered very low eigenvalues orspecies and environmental data (around 25%o cumulative variance in both axis), and alsoshowed low correlation between the speciesand environment variables (all r < 0.40) withan exception or electrical conductivity (r= 0.59). Te randomization test results orSpecies-Environment correlation (MonteCarlo test) showed no signicant correlation(p = 0.1872).

High water period (June 2006)During the high water period, it was

not possible to observe a clear pattern overtical variation or the abiotic parametersmeasured. emperature varied rom 27.0 to28.5oC, and higher values were registered atthe surace at 10:00 am. Lower values weremeasured at the bottom, during the night andthe early morning. Conductivity ranged rom9.5 to 11.5 S.cm-1, with the highest valuesregistered at 2:00 pm between 2 and 5 meters.

During the early morning hours, conductivity

Figure 4. Mean density o cladocerans (in the let, representedby bars) and Chaoborussp. (on the right, represent by a black

line), on org/m-3, during the low water period, on up lake.


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Nauplius 19(2): 109-122, 2011 115

Figure 5. Variation o the dissolved oxygen, during the 24-hour cycle in all depths, in up lake during the high water period(June/2006).

Figure 6. Vertical distribution oB. deitersi(a), B. negrensis(b) and D. polyspina(c) during the high water period, in up Lake.


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values were homogeneous in the watercolumn. Furthermore, the early morning hoursbetween 2:00 and 6:00 am registered lowervalues o dissolved oxygen, rom 6m down tothe bottom. In the ollowing hours, the values

ranged rom 2 to 10 mgO2/L, without anyclear vertical pattern, and at 2:00 pm, a valueo 6mgO2/L was ound in the entire watercolumn (Fig. 5). Oxygen saturation was higherat 2:00 pm and 6:00 pm, ranging between80% and 120%, whereas the lowest valueswere recorded at the bottom, during the night/early morning.

A total o 18 species were registeredduring this period. Bosminopsis negrensisand Bosminopsis brandor were registered

exclusively during this period. Tere was asmall increase in the population densitieso Moina rostrata and Bosmina hagmanni.Bosminopsis deitersi was the most abundantspecies, accounting or almost 60% o thecounted individuals, ollowed by C. cornuta,D. polyspina, and B. hagmanni(ab. 1).

Te mean density o cladocereans inthe high water period was 9045 org/m3, buta great heterogeneity was observed amongthe samples. No signicant variation in thecommunity densities was registered withinthe nycthemeral cycle at the high water periodbetween the dierent depths (Kruskal-Wallistest: H (13, N = 84) = 3,74727; p = 0,1311)and hours o the day (Kruskal-Wallis test: H(5, N = 84) =7,211148; p = 0,2054). Meanpopulation densities were higher at 6:00 am(=14894 org/m3); little variation was observedduring the other times o day, when meandensities ranged rom 7000 to 8000 org/m3.When considering vertical distribution, thehighest cladocerean densities were observedbetween 2 and 6 meters deep (maximumo 17336 org/m3 at 5 meters deep) whencompared to other layers. Under 6m deep,densities ranged between 5000 and 9000 org/m3.

During this period, ew species showedpatterns o vertical distribution. Highpopulation densities oB. deitersiwere observedduring all times and all depths, with a peak

o organisms observed closer to the surace at

6:00 am (Fig. 6a), indicating a possible reversein migration movement.

Bosminopsis negrensisreached its peak inthe deeper layers during the early morning andmigrated to the surace at 6:00 pm (Fig. 6b). It

is important to note that although this specieswas registered in low densities, the distributionpattern was strong. A similar pattern to theone registered or H. amazonicum during thelow water period was observed or D. polyspinain the high water period, when higher densitieswere observed between 3 and 6 metersduring most o the day (Fig. 6c). Tis speciespresented a heterogeneous distribution onlyduring the high water period. No clear patterno distribution was detected in the populations

oC. cornutaand B. hagmanniduring the highwater period.

Te Canonical Correspondence Analysisregistered very low eigenvalues or speciesand environmental data (around 13% ocumulative variance in both axis), and alsoshowed low correlation between the speciesand environment variables (all r < 0.44).Te randomization test results or Species-Environment correlation (Monte Carlo test)showed no signicant correlation (p = 0.0621).

Discussion

Seasonal dierences in species richness andabundance

Bosminidae was the most representativeamily, both in terms o species richness andabundance. Bosminopsis negrensis and B.brandorwere registered only during the highwater period and both species are considered tobe endemic to the Amazon basin (Brandor,1976; Rey and Vasquez, 1986).

Numerically, the species richnesswas similar in both periods o sampling, butthere were dierences in species composition,particularly when considering the species thatoccurred only during a specic period o thehydrological cycle. Seasonal changes in thecomposition o the Cladocera communityduring a determined cycle or hydrological

period have been previously registered in


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Nauplius 19(2): 109-122, 2011 117

tropical environments (Robertson and Hardy,1984; Lansac-haet al., 2004).

During the low water period, B. deitersiand M. minuta co-dominated the Cladoceracommunity, with very similar relative

abundances. B. deitersi and M. minuta areknown to be ound in lakes, rivers, and ponds,as these species can respond quickly to changesin the environment (Paggi and Jos de Paggi,1990; Melo and Rocha, 2006). C. cornutaalso occurred in both periods in very highdensities.

Tus ar, the records o M. rostrata,and H. amazonicum have been restricted tothe Amazon Basin (Elmoor-Loureiro, 1997).Holopedium amazonicum has a large body size,and the occurrence o this species in dierentlakes o the Amazon basin has varied romdominant to rare, particularly in acidic waterswith low suspended material (Rowe, 2000).Moina rostratahas also been associated withblack water lakes (McNair, 1980).

Te low abundance o Chydoridaeand Ilyocryptidae species was expected, sincesampling did not include the littoral region othe lake, the preerred habitat o these organisms(Whitesite et al., 1978; Smirnov, 1996). Teexception, Nicsmirnovius incredibilis, is achydorid adapted to environments with littlecurrent water and lentic systems, such as up,during its low-water period (Kotov, 2003).

Te dierences in the populationdensities and species richness between thetwo studied periods could be related tomodications to various lake characteristicsthat happen naturally during the hydrologicalcycle. At the beginning o the ooding, thewater volume increases due to the entrance othe Negro river and the increase in the owo the lakes afuents (Aprile and Darwich,2005).

Higher densities o Cladocera wereexpected during the low water period, whenthe lake is not connected to the Negro River.Te low wind eect caused a stratication othe water column, as demonstrated by thephysical and chemical parameters measured inthis study. On the contrary, such a act was not

observed; the mean population densities were

higher in the high water period.Previously, studies on the zooplankton

community (including cladocereans) detectedlower population densities at the high waterperiod. Hamilton et al. (1990) detected a

decrease in zooplankton densities in lakeslocated in the Orinoco Rivers oodplainduring the high water period; a possibleconsequence o this increased water volumeis a dilution eect. Tis conclusion was alsoound by Brandor and Andrade (1978) inJacaretinga Lake. A dilution eect, however,was not detected in this study.

Carvalho (1983), studied a vrzea lakeand registered higher densities o Copepodaand Rotiera during the low water period;however, cladocereans did not demonstratethis pattern. Detritivourous species occurredduring the low water period, whereasplanktivourous dominated during high waterperiods. Te same study, taking into accountthe entire Cladocera community, ound thatthe dierences throughout the whole yearwere not as pronounced or the individualCopepoda and Rotiera communities.

Comparing both periods, certainspecies, like B. deitersiand C. cornuta, were everrequent. Some species occurred only in oneperiod, like N. incredibilis, H. amazonicum,M. minutaandM. reticulata, which requentlyregistered in the low water period; D. polyspina,M. rostrataand B. negrensis registered mainlyin the high water period.

Patterns o vertical distributionGenerally, patterns o vertical

distribution in Cladocereans species weremore commonly observed during the lowwater period, which could be associated to thestratied state o the lake, as observed in thisstudy. Nogueira and Panarelli (1997) reportedthat the physical instability o the watercolumn inuenced the vertical distribution ozooplankton, and that stratied water bodies,with more stability and less inow o watercould acilitate the vertical movements o theseorganisms.

Te physical and chemical parameters

presented a similar variation when compared


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to a previous study perormed in the lake,in which the higher amplitude o variationo the dissolved oxygen, temperature, andconductivity was detected during the low waterlevel period (Darwich et al., 2005). In the

present study, a dierence 4C was detectedin the temperature between the surace andthe bottom. Tis dierence could constitute aphysical barrier or the organisms distribution,as reported by Arcia-Zago (1978).

Otherwise, Cladoceras speciesdensities were high in the deeper layers whenthe dissolved oxygen concentration was low.Te lowest levels o dissolved oxygen registeredor the low water period (=2 mgO

2/L) did not

seem to be a limiting actor or the organisms.

Tis last statement was also ound by Lansac-ha et al. (1995) when it was detected thatdespite the understanding that Cladocereandensities are related more to oxygenate layers,this element was abundant in the entire system.

Te organisms densities were higher indeeper layers o the water column, but somespecies showed a preerence to the suracelayers, specicallyB. longirostrisduring the lowwater period and B. negrensisduring the highwater period, both o them perorming a dielvertical migration to the surace at night. Tisdistribution pattern oB. longirostriswas alsoreported by Ramos-Jiliberto et al. (2004) in aChilean lake, but they also ound the reversemovement o this species during a certainperiod o the year.

Many Cladocera species eed on detritusand bacteria when they are more abundantthan phytoplankton; this inormation isparticularly pertinent to this study, becausemost o the organisms remained at the bottomwhere phytoplankton densities are not typicallyhigh (Gliwicz, 1969a, b; Monakov, 1972). Inthis sense, higher values o conductivity wereregistered at the bottom, which explains thehigh correlation detected in the CanonicalCorrespondence Analysis. Te correlationcould be simply numerical, since conductivityexpresses the concentration o ions dissolved inthe water, and it has never been directly relatedto Cladocera. It is known that conductivity

is mainly related to decomposition and

productivity, as both processes promote theliberation o ions to the environment, makingthem available to other biological processes(Wetzel, 1983). Higher conductivity at thebottom could be related to a higher abundance

o species like B. longirostris and B. deitersi,since both have detritivorous habitats (Paggi,1979; Kotov, 1997).

Te distribution oM. minuta waspeculiar, as the densities at bottom were veryhigh at night, and then sharply decreased inthe early morning, returning to its maximumonly at the end o the day. A similar patternwas observed with Chaoboridae.

In this study, both Cladocera andChaoborussp. were very abundant during the

night at the bottom and demonstrated a sharpdecrease in the early morning. I we consider thatChaoborussp. usually preys during night, it ispossible to associate the decrease o Cladocera,specically oM. minuta, to Chaoborus sp.predation (deBernardi et al., 1987; Rejas etal., 2007). It is also important to associate thedecrease oChaoborussp. densities in the earlymorning to the burrowing behavior o thislarva, which remains in the mud throughoutthe day (Gosselin and Hare, 2003).

Te Diptera larvae, Chaoborus sp.,is an important zooplankton predator, butluminosity does not play a very importantrole in this case, as these organisms respondto a prey at a distance o 2 to 4mm. Terelationship between these two groupso organisms is more related to chemicalreceptors and the developmental stage o thelarvae than to light itsel (Neill, 1981; ollrianand Dodson, 1999; Monakov, 2003). In thisstudy, the decrease inM. minutas populationscould be a consequence o intense invertebratepredation. In up Lake, sh predation hasalready been previously observed (Previattelliand Santos-Silva, 2011).

Also, it is necessary to consider thatthe intense changes in population densitiesregistered or some species in this study couldbe also related to horizontal migration, sincethis study only sampled a single area o thelake. Migration to the littoral zone or to other

parts o the lake is requently associated with


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predator avoidance, and to the search or abetter ood supply (Lauridsen and Buenki,1996). Ghidini (2011) detected most o theplanktonic cladocereans species o this studyon the littoral zone o up Lake.

Some species, like H. amazonicum,remain at certain layers o the water column,whose densities in the low water period werehigher at 3 and 4 meters o depth. On theother hand, higher densities o D. polyspinawere observed around 5m during the highwater period. H. amazonicums pattern overtical distribution was previously observedby Previatelli et al. (2005). Accordingto DeMeester and Weider (1999), somezooplankton species have a depth selection

behavior, constantly migrating in order toremain in the same layer o the water columnand t into the lakes stratication (light,temperature, ood or predation pressure couldalso be related). Tis could be associated to thevertical distribution pattern registered or H.amazonicum, D. polyspina.

When considering the distribution oB. longisrostris, B. deitersi, M. minutaand M.reticulata it was possible to detect a verticalmigration pattern, since these species switchedtheir density per depth throughout the dielcycle. Broadly, the patterns o distributiono B. longisrostris and B. deitersi could berelated to electrical conductivity (related todecomposition and these species detritivoroushabits), whileM. minutawas inuenced moreby Chaoboridae.

Te absence o a correlation between thedistribution o Cladocera and the physical andchemical parameters is a requently observedresult in natural systems. Usually, morethan a ew actors inuence the organismscomposition and distribution (Lampert andSommer, 1997; Sterzaet al., 2002).

Tis study demonstrated that thedistribution o organisms was not homogeneousand that there were many actors inuencingthe distribution. Tis nding could improvethe data currently available about zooplanktoncommunities o the Amazon Basin. Also, theact that many species distribute themselves

heterogeneously through the diel cycle and

varying depths illustrates that samplingrestricted to only one period o the day oronly at surace level is not suitable or a truecharacterization o the composition andabundance o the Cladoceran community.

Tis nal statement is extremelyimportant, particularly when we reect onprevious studies conducted in Brazil and othercountries, in which sampling was typicallyconducted only during the morning and onlyat a ew dierent depths. Tis study showsthat to properly evaluate the diversity andabundance o the cladocereans planktoniccommunity, one must sample the entirewater column at least twice daily, during themorning and at night. It is also important to

sample other regions o the lake to determinethe role o horizontal migration in this process,consider the littoral cladocereans and monitora higher number o environmental parametersin order to establish better relationships withinthe Cladocera community.

Conclusions

Tere was a dierence in the composition

o planktonic cladocereans between thelow and high water periods. M. minuta, M.reticulata, and H. amazonicum occurred only/mainly in the low water period, whereas B.brandorand B. negrensisoccurred only in thehigh water period. B. deitersiand C. cornutawere the most abundant species throughoutthe entire study. During the low water period,B. deitersimigrated to the bottom o the lake inthe aternoon. B. longirostrisshowed nocturnalmigration to the surace. Most individuals oM. minuta,M. reticulata, and H. amazonicumremained at the bottom at all times. Duringthe high water period, the vertical patterns odistribution were less evident than during thelow water period, with B. deitersi migratingto the surace during the early morning andB. negrensisduring the early aternoon. Mostcladocereans remained between 2 and 6 meterso dept. Patterns o vertical distributon byBlongirostris and B. deitersi were related toelectrical conductivity, whereas the distribution


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oM. minutawas inuenced by invertebratepredation. It became evident that to eectivelysample the cladocereans in the community,the vertical distribution o its members mustbe considered. Sampling should be, as a rule,

at least perormed during the day and at nightand should include the entire water column.

Acknowledgements

We thank Dr. Gerd Oltman Brandorfand Dr. Barbara Ann Robertson or theirconsiderations and suggestions, all thestudents o Planktons Lab or supportduring sampling and data analysis and CNPq

or investing in the Biotup Project and thestudents scholarship.

References

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Submitted 06 May 2011

Accepted 30 November 2011


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Summary Volume 19(2) 2011Printed and Distributed in December 2011

Anala R. Daz, Estela C. LoprettoTe genus ChlamydothecaSaussure (Crustacea: Ostracoda) in northeastern Argentina

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Andr R. Ghidini, Edinaldo N. dos Santos-SilvaComposition, species richness and patterns o nycthemeral vertical distribution o planktoniccladocereans in a black water Amazonian lake

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Carina Appel, Aline F. Quadros, Paula B. AraujoMarsupial extension in terrestrial isopods (Crustacea, Isopoda, Oniscidea) 123-128

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Luis E. A. Bezerra,Alexandre O. AlmeidaTe lie and work o Petrnio Alves Coelho (4 November 1937 28 November2011), with a list o his taxa and publications

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Article 2 - Guidini & Santos-Silva