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Demersal zooplankton communities from tropical habitats in thesouthwestern AtlanticPedro A. M. C. Meloa; Tâmara A. Silvab; Sigrid Neumann-Leitãoa; Ralf Schwambornc; Lucia M. O.Gusmãoa; Fernando Porto Netod

a Federal University of Pernambuco, Department of Oceanography, Recife-PE, Brazil b Bahia StateUniversity, Department of Education, Paulo Afonso, Bahia, Brazil c Federal University of Pernambuco,Department of Zoology, Recife, Pernambuco, Brazil d Federal Rural University of Pernambuco,Department of Zootecny, Recife, Pernambuco, Brazil

First published on: 14 July 2010

To cite this Article Melo, Pedro A. M. C. , Silva, Tâmara A. , Neumann-Leitão, Sigrid , Schwamborn, Ralf , Gusmão, LuciaM. O. and Porto Neto, Fernando(2010) 'Demersal zooplankton communities from tropical habitats in the southwesternAtlantic', Marine Biology Research, 6: 6, 530 — 541, First published on: 14 July 2010 (iFirst)To link to this Article: DOI: 10.1080/17451000903426557URL: http://dx.doi.org/10.1080/17451000903426557

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ORIGINAL ARTICLE

Demersal zooplankton communities from tropical habitats in thesouthwestern Atlantic

PEDRO A. M. C. MELO1, TAMARA A. SILVA2, SIGRID NEUMANN-LEITAO1*,

RALF SCHWAMBORN3, LUCIA M. O. GUSMAO1 & FERNANDO PORTO NETO4

1Federal University of Pernambuco, Department of Oceanography, Recife � PE, Brazil; 2Bahia State University, Department

of Education, Campus VIII, Paulo Afonso, Bahia, Brazil; 3Federal University of Pernambuco, Department of Zoology, Recife,

Pernambuco, Brazil; 4Federal Rural University of Pernambuco, Department of Zootecny, Recife, Pernambuco, Brazil

AbstractDemersal zooplankton were captured with traps from a set of tropical coastal habitats (seagrass bed, coral reef, gravel, andsand bottoms) to allow comparisons among communities. Sampling was carried out during dry and rainy seasons in 2000and 2001. Traps with and without light were placed at 18:00 and removed at 06:00 the next day for three consecutive days.Eighty-eight zooplankton taxa were identified. Copepoda was the most abundant group, outranking in relative abundance inseagrass and in sandy bottoms. Copepoda was mainly represented by Oithona oculata, Pseudodiaptomus acutus, and Acartialilljeborgi. No significant differences were found among substrates (P�0.1464); however, differences were significantbetween light and dark traps communities (P�0.0410). The average density was 7113 (93966) ind m�2 in the light and4759 (94825) ind m�2 in the dark. In the light traps, Amphipoda and O. oculata were more representative. Without light,the main group was Foraminifera (�40%). Cluster analysis presented two main groups, Itamaraca Island and TamandareBay; light and dark traps formed separate groups within these location groups. The results allow us to assess the efficiency ofthe used traps in a set of habitats of the tropical coastal area and gives information on the preference of specific organismgroups in one of the tested substrates.

Key words: Coral reef, demersal, sand, seagrass beds, traps, vertical migration

Introduction

The seagrass, saltmarsh, and coral reef patches

provide a structure with ecological functions that

support high species diversity of many invertebrates

(Sheridan 1997; Beck et al. 2001; Heck et al. 2003;

Touchette 2007). This high biological diversity is

due to important ecological linkages in the hetero-

geneous complex mosaic of ecosystems in tropical

coastal areas (Ogden 1997; Heidelberg et al. 2004).

Among the invertebrates, zooplankton is an im-

portant intermediate component in food webs, acting

as a trophic link between small particles (detritus and

microalgae) and planktivorous fishes (Morgan 1990;

Boltovskoy 1999). It also includes larvae of diverse

organisms; some are commercially important.

Demersal zooplankton are small, active organisms

that reside or hide in or near the substrata, migrating

up into the water column from the bottom at night

and then returning to the substratum before daylight;

therefore, these zooplankton cannot be captured by

daytime tows (Sorokin 1990; Gross & Gross 1996).

Studies have reported high abundances of demersal

zooplankton emerging nightly from coral reefs, kelp

beds, and soft-bottom habitats (e.g. Hammer 1981;

Alldredge & King 1985; Jacoby & Greenwood 1988,

1989; Cahoon & Tronzo 1992; Carleton et al. 2001),

suggesting that demersal zooplankton may play an

important role in the ecology and trophic pathways of

many benthic communities.

Also, numerous experiments in the field and in the

laboratory have shown that vertical migration by

*Correspondence: Sigrid Neumann-Leitao, Federal University of Pernambuco, Department of Oceanography, Av. Arquitetura s/n, Cidade

Universitaria, 50730-540, Recife � PE, Brazil. E-mail: sigrid@terra.com.br

Published in collaboration with the University of Bergen and the Institute of Marine Research, Norway, and the Marine Biological Laboratory,

University of Copenhagen, Denmark

Marine Biology Research, 2010; 6: 530�541

(Accepted 14 October 2009; Published online 15 July 2010; Printed 14 October 2010)

ISSN 1745-1000 print/ISSN 1745-1019 online # 2010 Taylor & Francis

DOI: 10.1080/17451000903426557

Downloaded At: 00:05 20 October 2010

demersal species is an adaptation, in part, to preda-

tion and behaviours that can be induced by endogen-

ous (reproductive events and biological rhythms) and

exogenous factors (light, gravity, dissolved oxygen,

temperature, predator, and prey abundance). How-

ever, light is generally accepted to be the most

significant external factor (Putzeys & Hernandez-

Leon 2005), as well as a limiting factor for the optical

efficiency of visual predators (Ohlhorst 1982; Gliwicz

1986; Ringelberg 1995).

Conventional sampling methods used by zooplank-

ton ecologists are inadequate in most cases for

answering questions regarding demersal plankton

drift ecology (Greene 1990). This difficulty is

explained by the sporadic or cyclic emergence, or

low densities, of demersal drifting biota (Dahms &

Qian 2004). In studies of coral reefs, Emery (1968)

and Sale et al. (1976) found different qualitative

compositions of zooplankton in samples collected by

airlift and light traps as compared with net tows or

even with samples collected by the usual traps at night

or traps without light. The difference in these areas

might be due to the zooplankton being composed of

demersal and meroplanktonic forms, which hide at the

bottom during the day and appear in the water column

mostly at night (Renon 1977). Similar traps were used

by McWilliam et al. (1981) to compare two lagoonal

patch reefs, including an open sand environment that is

1�3 m away from the reef. They distinguished two

types of fauna, ‘coral’ and ‘sand’ fauna.

Emery (1968) reported that different species of

reef zooplankton, including some holoplanktonic

species, prefer different types of shelter at the

bottom. According to Sorokin (1990), these places

include seagrasses, macrophytes, and soft-bottom

substrates (sand or gravel).

Among the marine habitats, the tropical South

Western Atlantic (SWA) is poorly known to many

planktonic dermersal groups and many coastal

habitats are under threat by human activities (e.g.

mariculture, fishing, dumping of waste and pollu-

tion), and until we have a firmer idea of their

biodiversity and what controls the emergence beha-

viour, we have little hope of conserving its ecological

functions and biodiversity.

In our study, due to the successful use of a trap-

sampler in coral reef areas, we used this sampling tool

for the first time on seagrass beds (Halodule wrightii

Aschers) to assess the composition and existence of

vertical migration patterns of demersal zooplankton.

Thus, the aim of the present study was to test the

same trap-sampler in a set of habitats (seagrass bed,

coral reef, gravel, and sand bottoms) that normally

occur together in the same tropical coastal area in a

narrow connectivity to allow for comparisons among

communities sampled with the same method; and to

assess the preference of specific organism groups in

one of the tested substrates.

Materials and methods

Sampling areas

Itamaraca Island and Tamandare Bay are located on

the north and south coasts of Pernambuco, Brazil,

respectively (Figure 1). The area has a humid, tropical

climate with two seasons: dry (September�February)

and rainy (March�August). Itamaraca Island presents

high ecosystem diversity and a strong connectivity

among mangrove forests, small rivers estuaries,

beach rocks, and seagrass beds (Neumann-Leitao &

Schwamborn 2000). These seagrass beds are mainly

composed of Halodule wrightii and occasionally

Halophila decipiens Ostenfeld, which occur all along

the east coast of the island (Cocentino et al. 2004),

being commonly found at a depth of between

2 and 3 m. Tamandare Bay has approximately 9 km

of coastline, enclosed by coastal reefs. These coral reef

formations are parallel to the coastline and resemble

fringing reefs, with tops exposed during low tide

(Maida & Ferreira 1997). Of the 18 species of hard

corals described for the Brazilian coast, nine were

observed in this area (Ferreira & Maida 2006). In this

region, the reefs associated with the mangroves

support intense artisanal fishing (Ferreira et al. 2000).

At Itamaraca and Tamandare, water temperatures

vary from 248C (rainy) to 318C (dry season), and

salinity from 27 psu (rainy) to 34 psu (dry season)

(Moura & Passavante 1995; Medeiros et al. 2001).

Demersal zooplankton sampling and procedures

The demersal zooplankton samples were captured

from two fixed stations at Itamaraca Island, one on a

sandy bottom (A) and another on a seagrass bed (B),

and two fixed stations at Tamandare Bay, one on a

coral reef (C) and another on a gravel bottom (D)

located within the areas shown in Figure 1. Sampling

was carried out at neap tide during the dry (January

and February 2001) and rainy (July 2000 and

August 2001) seasons in both areas.

The samples were taken with emergence net traps

(inverted cones placed over the bottom � according

to Alldredge & King 1980 and Alldredge 1985) with

300 mm mesh size, 1 m mouth diameter and 1.5 m

high (Figure 2). Traps were placed at 18:00 and

removed at 06:00 of the next day during three

consecutive days.

For each type of bottom (A, B, C and D), two net

traps were used: one was left in the light (L) and one in

the dark (D). Triplicates were used for all treatments,

totalling 24 traps. The treatment codes consisted of

type of bottom�trap (e.g. BL). The samples were

Demersal zooplankton communities from tropical habitats 531

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fixed in 4% formaldehyde buffered with borax

(5 g l�1), according to Newell & Newell (1963).

Zooplankton species were identified to the lowest

possible taxonomic unit, as described in specific

literature (Bjornberg 1981; Boltovskoy 1981, 1999),

and taxon abundance (per m2) was counted under

a stereomicroscope, based on 5.0 ml subsamples.

Three subsamples were taken with a Stempel pipette

after sample dilution to 250 ml and homogenization.

The samples with low density were completely

analysed.

Data treatment

Density (ind m�2), relative abundance (%) and

frequency of occurrence (%) were calculated and for

the relative abundance the following scale were used:

dominant (�70%), abundant (70�40%), present

(40�10%), and rare (510%). The results of the

frequency of occurrence were expressed in percen-

tages as very frequent (�70%); frequent (70�40%);

infrequent (40�10%), or sporadic (510%). The

classification ‘others’ used in the figures comprised

all the taxa occurring with low abundance or fre-

quency.

The Shannon diversity index (H’) was applied for

the estimation of community diversity (Shannon

1948), and the evenness was calculated according

to Pielou (1977).

Statistical analyses were based on density data

(ind m�2) of the zooplanktonic community, with

PRIMER 5 for similarity tests among samples.Figure 2. Trap-sampler design.

Figure 1. Demersal zooplankton sampling area at Itamaraca Island (sandy bottom and seagrass) and Tamandare Bay (coral reef and gravel

bottom), Pernambuco, Brazil, with the experimental localization.

532 P. A. M. C. Melo et al.

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One-way Kruskall�Wallis ANOVA (Zar 1996) was

used to test for significant (PB0.05) effects of the

factors ‘bottom types (A, B, C and D)’ and ‘traps

(L and D)’. The Spearman correlation was applied

to determine the association between the traps.

Results

Eighty-eight zooplankton taxa were identified

(Tables I and II), with some occurring in different

life-cycle stages. The frequency of occurrence showed

that 5 in 88 taxa were very frequent (Cumacea,

Amphipoda, Ostracoda, Mysidacea and Foraminifera)

(Tables I and II). Seven taxa were frequent and 10

were infrequent. The remaining taxa were sporadic,

representing 75%, of which 29 occurred in only one

sample (2.94%). As no significant differences were

obtained between replicas, in our results we used the

mean of each treatment.

Copepoda was the most abundant group in nearly

all the treatments, outranking all other groups in

sandy and seagrass substrate, as well as in the gravel

with light treatment; a total of 83.2% were attained in

the seagrass substrate with dark treatment (Figure 3).

This group was represented mainly by Oithona oculata

Farran, 1913, Pseudodiaptomus acutus (F. Dahl,

1894), and Acartia lilljeborgi Giesbrecht, 1889. Be-

sides Copepoda, in coral reef with both light and dark

treatment, as well as in gravel substrate during dark

treatment, Foraminifera and others groups (mainly

Radiolaria and Ostracoda) had larger contributions

besides Amphipoda for CL. In sandy substrate, both

in dark and light, the largest contribution was from

Amphioxus larvae, besides Ostracoda, for AD and

Gastropoda for AL. Among the Copepoda, O. oculata

was dominant in gravel with light and occurring in less

than 10% in the three other substrates. This species

did not occur in sandy and seagrass bottoms.

A. lilljeborgi was present in sandy and seagrass sub-

strates and rare in CL and DD. P. acutus was present in

A and B and rare in D and CL. Another abundant

Copepoda was Calanopia americana F. Dahl, 1894,

which was rare, but comprised almost 10% of the

community in seagrass during dark treatment.

Considering the different substrates (Figure 4),

similarity was observed between the main taxa

(A. lilljeborgi, P. acutus, Mysidacea, Cumacea, and

Amphipoda) in the composition of sandy and

seagrass communities. Calanopia americana, Anom-

ura (Glaucothoea), Teleostei (egg), and Ostracoda

presented greater affinity for seagrass bed, while

Amphioxus (larvae) and Gastropoda (veliger) de-

monstrated a preference on the sandy substrate. The

coral reef and gravel substrates also showed similar-

ity between the main groups.

When comparing sandy and seagrass with coral

reef and gravel substrates, a divergence, caused

mainly by an important decrease on A. lilljeborgi

and P. acutus, was observed. Also, an increase in the

Foraminifera and the appearance of O. oculata in C

and D could be noted.

Analysing the type of trap (light or dark), the

average density was 711393966 ind m�2 in light

and 475994825 ind m�2 in dark. The relative

abundance of some groups had similar importance,

but a difference could be noted (Figure 5). In light-

traps Amphipoda, O. oculata, and other groups were

more representative. Without light, the main group

was Foraminifera at more than 40%. In addition,

there were considerable decreases in the abundances

of Gastropoda (veliger) and O. oculata, from 6.5% to

1.1% and 27% to 1%, respectively (Figure 5).

The affinity observed by different groups for the

type of substrate and trap was very specific for

Amphioxus (larvae) and Gastropoda (veliger) in AL,

Labidocera fluviatilis F. Dahl, 1894 in AD,

C. americana in BD, Anomura in BL, Harpacticoida

Copepoda in CD, Brachyura (zoea and megalopa)

and Oikopleura dioica Fol, 1872 in CL, Foraminifera

in DD, and O. oculata in DL.

The average density observed for the treatments

varied from 10939592 ind m�2 (AD) to 12,2099

15,765 ind m�2 (DL) (Figure 6). Light-traps had

higher density than dark for all substrates, but a

significant difference was registered only for the

sandy substrate (t-test, p�0.0319).

The species diversity ranged from 1.653 bits ind�1

(DL) to 2.642 bits ind�1 (CD), and was classified

between low and medium diversity (Figure 7).

Comparing the treatments, the substrates A, C, and

D had higher diversity in the dark rather than light-

traps. Evenness values ranged from 0.369 (DL) and

0.59 (CD). Generally, the coral reef substrate

showed higher diversity and evenness than other

substrates, followed by the sandy bottom. The

evenness index showed better uniformity for sandy

substrate samples, whereas in the seagrass samples,

only one or two species were dominant. In general,

the a diversity was 2.348 bits ind�1 to Itamaraca

Island and 2.472 bits ind�1 to Tamandare Bay while

the b diversity was 2.725 bits ind�1.

No significant difference was found among sub-

strates (A, B, C, and D with ANOVA, p�0.1464)

and between traps communities (L and D) (t-test,

p�0.4510).

The Cluster analysis grouped the treatments into

two main groups, one containing the samples from

Itamaraca Island and another from Tamandare Bay.

For Itamaraca Island, BD and BL formed a sub-

group (0.25 dissimilarity). In Tamandare Bay, this

Demersal zooplankton communities from tropical habitats 533

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Table I. Composition, relative abundance and frequency of occurrence (for sample) of demersal zooplankton (without Copepoda) captured

with traps at Itamaraca Island and Tamandare Bay, PE, Brazil. Values used in calculation are means.

Substrate Treatment

Taxa A B C D L D FO (%)

Protozoa

Lobosa (Arcella sp.) 0.33 0.13 2.94

Foraminifera 0.38 0.19 19.37 37.49 10.17 43.49 70.59

Radiolaria 9.85 2.79 3.73 3.69 58.82

Tintinnina 0.33 0.37 0.37 0.12 14.71

Cnidaria 0.05 0.01 2.94

Platyhelminthes (Convoluta sp.) 1.84 0.84 0.47 1.42 17.65

Nematoda 0.33 0.19 0.29 8.82

Kinorhyncha 0.28 0.35 5.88

Mollusca

Gastropoda (veliger) 22.85 0.16 1.50 0.47 6.50 1.19 44.12

Bivalvia (veliger) 1.23 2.17 0.19 0.93 0.62 17.65

Polychaeta (larvae) 0.47 0.30 1.17 1.77 0.70 2.13 47.06

Pontodora pelagica Greeff, 1879 0.17 0.10 2.94

Pycnogonida (Nymphon sp.) 0.03 0.83 0.20 0.20 11.76

Crustacea

Crustacea (nauplius) 0.09 0.03 0.03 5.88

Crustacea (others larvae) 0.17 0.37 0.23 0.21 11.76

Ostracoda 0.57 1.46 11.35 4.56 4.46 6.29 91.18

Cirripedia (nauplius) 0.06 0.09 0.01 0.12 8.82

Cirripedia (cypres) 0.24 0.07 0.08 8.82

Leptostraca 0.33 0.19 0.14 0.21 11.76

Stomatopoda (juvenile) 2.94

Decapoda

Penaeidae (mysis) 0.67 0.28 0.07 0.64 11.76

Periclimenes sp. (larvae) 0.33 0.13 5.88

Lucifer faxoni Borradaile, 1915 0.17 0.09 0.07 0.12 5.88

Lucifer faxoni Borradaile, 1915 (protozoea) 0.17 0.10 2.94

Alpheidae (larvae) 1.67 0.19 0.68 0.20 26.47

Acetes americanus Ortmann,1893 0.05 0.02 2.94

Caridea (zoeae) 0.33 1.12 0.24 0.05 23.53

Anomura (zoeae) 0.09 0.03 2.94

Anomura (glaucothoea) 0.61 6.84 1.16 0.17 23.53

Callianassidae 1.09 0.33 0.19 0.53 0.08 17.65

Upogebiidae 0.07 0.01 2.94

Paguridae 0.07 0.17 0.19 0.17 0.10 11.76

Porcellanidae (zoeae) 0.09 0.33 0.03 0.20 8.82

Brachyura (zoeae) 0.99 0.42 1.17 0.19 0.32 0.94 38.24

Brachyura (megalopa) 0.43 1.54 1.50 0.65 1.05 0.68 44.12

Callinectes (juvenile) B0.01 B0.01 2.94

Callinectes (megalopa) B0.01 B0.01 2.94

Decapoda (others) 0.20 0.03 2.94

Mysidacea 6.01 3.80 8.35 4.19 5.26 5.67 88.24

Cumacea 3.93 4.42 4.67 3.26 4.13 3.33 94.12

Isopoda

Isopoda (parasite) 0.83 0.56 0.56 0.35 14.71

Isopoda (larvae) 0.09 0.04 0.17 0.65 0.50 0.22 20.59

Isopoda (others) 0.24 0.62 0.83 0.84 0.93 0.40 50

Amphipoda 12.20 7.43 19.87 3.07 10.59 6.45 91.18

Membranipora sp. (cyphonauta) 0.17 0.10 2.94

Chaetognatha 0.47 0.19 0.08 0.31 11.76

Larvacea (Oikopleura dioica Fol, 1872) 0.83 0.33 2.94

Amphioxus (larvae) 6.86 0.87 1.82 0.32 20.59

Teleostei (egg) 0.14 0.85 1.34 0.19 0.18 1.00 47.06

Teleostei (larvae) 0.17 0.28 0.14 0.23 11.76

Notes. A, sandy bottom; B, seagrass bed; C, coral reef; D, gravel bottom. Net trap left in light (L) and dark (D).

534 P. A. M. C. Melo et al.

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subgroup was between CD and CL (0.43 dissim-

ilarity) (Figure 8).

The Spearman correlation between the set of

treatments presented a significant positive correlation

( pB0.05) for type of trap (0.4740), indicating that

when the number of individuals in one type of trap

increases, the same happens to the other treatment.

Discussion

The constant presence of organisms in early life

stages highlights the importance of seagrass beds

and reefs as nurseries for fish and invertebrates

(Heck et al. 2003). The occurrence of these stages,

including some species of socio-economic interest in

the region, is an essential argument for the conserva-

tion of these habitats (Dorenbosch et al. 2006).

Other studies in the area presented higher richness

than our research, but we observed the importance of

certain groups that normally are not observed in

samples collected by net tows, such as Cumacea and

Amphioxus (larvae), demonstrating the usefulness

of this trap-sampler for studies of zooplankton diver-

sity and temporal variations. Adults and larvae of

Table II. Composition, relative abundance and frequency of occurrence (for sample) of demersal Copepoda captured with traps at

Itamaraca Island and Tamandare Bay, PE, Brazil. Values used in calculation are means.

Substrate Treatment

Taxa A B C D L D FO (%)

Copepoda

Calanoida

Paracalanus indicus Wolfenden, 1905 0.05 0.02 2.94

Paracalanus parvus (Claus, 1863) 0.17 0.10 2.94

Paracalanus quasimodo Bowman, 1971 0.09 0.12 2.94

Parvocalanus crassirostris (F. Dahl, 1894) 0.66 0.11 0.28 0.09 0.51 14.71

Clausocalanus furcatus (Brady, 1883) 1.00 0.37 0.35 0.53 17.65

Lucicutia flavicornis (Claus, 1863) 0.17 0.10 2.94

Temora turbinata (Dana, 1849) 0.24 0.06 5.88

Pseudodiaptomus acutus (F. Dahl, 1894) 18.78 34.27 0.33 0.84 7.87 5.47 44.12

Pseudodiaptomus richardi (F. Dahl, 1894) 0.52 0.13 0.19 0.32 11.76

Pseudodiaptomus marshi Wright, 1936 0.33 0.07 0.01 0.14 5.88

Pseudodiaptomus trihamatus Wright, 1937 0.09 0.07 0.04 5.88

Candacia curta (Dana, 1849) 0.05 0.01 2.94

Paracandacia bispinosa (Claus, 1963) 0.05 0.01 2.94

Labidocera fluviatilis F. Dahl, 1894 1.18 0.09 0.08 0.49 8.82

Pontellopsis brevis (Giesbrecht, 1889) 0.06 0.01 2.94

Calanopia americana F. Dahl, 1894 0.66 4.03 0.28 0.15 1.37 32.35

Acartia lilljeborgi Giesbrecht, 1889 15.14 29.63 0.50 0.09 5.68 5.28 38.24

Cyclopoida

Oithona hebes Giesbrecht, 1891 0.95 0.05 0.09 0.32 0.04 17.65

Oithona oculata Farran, 1913 1.67 31.16 25.98 1.35 29.41

Oithona nana Giesbrecht, 1892 1.32 0.36 2.94

Oithona sp. 0.28 0.23 2.94

Harpacticoida

Darcythompsonia radans Por, 1983 0.17 0.07 2.94

Harpacticus chelifer (O.F. Muller, 1785) 0.09 0.08 2.94

Porcellidiidae 0.17 0.09 0.14 5.88

Porcellidium sarsi Bocquet, 1948 0.17 1.02 1.38 5.88

Tegastidae 0.17 0.10 2.94

Euterpina acutifrons (Dana, 1849) 0,61 0,17 5.88

Tigriopus sp. 0.07 0.01 2.94

Tisbe sp. 0.15 0.33 0.07 0.13 17.65

Eudactylopus sp. 0.01 0.17 0.10 5.88

Longipedia coronata Claus, 1863 0.03 0.33 0.19 0.08 0.31 14.71

Poecilostomatoida

Corycaeus giesbrechti F. Dahl, 1894 0.19 0.16 5.88

Oncaea venusta Philippi, 1892 0.17 0.07 2.94

Corycaeus sp. 0.01 0.17 0.09 0.21 8.82

Farranula gracilis (Dana, 1849) 0.17 0.07 2.94

Asterocheres sp. 0.17 0.07 2.94

Caligus sp. 0.50 0.29 5.88

Copepoda (others) 0.24 0.31 0.09 0.04 20.59

Notes. A, sandy bottom; B, seagrass bed; C, coral reef; D, gravel bottom. Net trap left in light (L) and dark (D).

Demersal zooplankton communities from tropical habitats 535

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Amphioxus normally live in sandy bottoms, present-

ing a diurnal vertical migration immediately after

sunset (Wickstead & Bone 1959), which could

explain its absence in daylight net zooplankton tows.

Copepoda, Mysidacea, Ostracoda, and Amphi-

poda, all with larger contributions, demonstrated

their importance in the demersal plankton, in spite

of being holoplanktonic organisms that live perma-

nently in the water column. During the day, these

organisms are found near the substrate searching

for shelter (Emery 1968). A diverse array of small

invertebrates exhibit this demersal behaviour, which

is particularly prevalent among crustaceans, such as

Copepoda, Ostracoda, and Peracarida (Amphipoda,

Isopoda, Mysidacea, Cumacea, and Tanaidacea)

(Hobson & Chess 1976; Hammer & Zimmerman

1979), as observed in our study.

The dominance of Copepoda, Mysidacea, Ostra-

coda and Amphipoda in our study may be due to

sampling procedures, as compared to demersal zoo-

plankton collected by Youngbluth (1982) in three

structurally different emergence traps, indicating that

density and diversity estimates were affected by design

features and sampling procedures. The smallest mesh

netting (63 mm) contained larger catches than traps

with 202- or 333-mm mesh. Samples from Porter�Porter traps tethered 1 and 10 cm above the bottom

had statistically more zooplankton than all other traps

Figure 4. Composition of demersal zooplankton captured with traps for the substrates at Itamaraca Island and Tamandare Bay,

Pernambuco, Brazil. (A) Sandy bottom, (B) seagrass, (C) coral reef and (D) gravel bottom.

Figure 3. Relative abundance of demersal zooplankton captured with traps at Itamaraca Island and Tamandare Bay, Pernambuco, Brazil.

(A) Sandy bottom, (B) seagrass, (C) coral reef and (D) gravel bottom. (L) light and (D) dark.

536 P. A. M. C. Melo et al.

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set on the substratum. A consistently greater total

number of demersal zooplankton was captured in the

Alldredge�King traps when all three trap types were

sampled at the same time and area. The abundance

and rank order of all but the most numerous animals

(harpacticoid copepods) differed between the traps.

Alldredge�King traps caught more of the larger organ-

isms (polychaetes, cumaceans, and gammaridean

amphipods), whereas the Porter�Porter and Hobson�Chess traps contained larger densities of smaller

zooplankton (copepod nauplii and gastropod veligers).

However, we used the same mesh size and differences

were mainly caused by the different substrates.

The dominance of Copepoda in Brazilian tropical

areas is common (Bjornberg 1981; Neumann-Leitao

1995), but this was not observed for substrates C and

D, the last only in the dark treatment. This taxa did

not exceed 20% in traps completely sealed at

the bottom (Hobson & Chess 1979), indicating the

importance of new methodologies in plankton studies.

Decapoda, Cumacea, and Polychaeta, considered

as true demersal plankton (Sorokin 1990), also

occurred in the samples, although with lesser repre-

sentation. The Mysidacea showed greater fidelity to

seagrass beds, and suffered the strong influence of light

(Cebrian et al. 2001). The light intensity variation is

the main cue controlling vertical migrations of many

Mysidacea (Gal et al. 1999), and they are well known

as nocturnally active, demersal zooplankton (Hobson

& Chess 1976; Hammer & Zimmerman 1979).

Pseudodiaptomus acutus had a high abundance in

light traps in A and B, in agreement with others

Figure 6. Density average (�standard deviation) for the treatments of demersal zooplankton captured with traps at Itamaraca Island and

Tamandare Bay, Pernambuco, Brazil.

Figure 5. Composition of demersal zooplankton captured with light and dark traps at Itamaraca Island and Tamandare Bay, Pernambuco,

Brazil.

Demersal zooplankton communities from tropical habitats 537

Downloaded At: 00:05 20 October 2010

studies which identified Pseudodiaptomus migration.

Rios-Jara & Gonzalez (2000) verified this migration in

seagrass bed, mud and sand in other tropical Bay.

The high abundance of Calanopia americana,

which occurs in the water column at night and

is buried in the mud bottom during the day

(Boltovskoy 1981), in samples of seagrass bed

without light can be justified by the efficient

retention of fine sediments by seagrass bed envir-

onments (Alves 2000). Pseudodiaptomus trihamatus

Wright, 1937 was another species that should be

highlighted. This exotic species, native from Indo-

Pacific coastal waters, was accidentally introduced

in Northeastern Brazil in 1977 (Medeiros et al.

1991) and was not registered to Pernambuco

coastal area until the present study.

Amphioxus larvae and other larvae, such as Gastro-

poda (veliger) and Crustacea (zoea), are included in

the groups that spend the initial stages of ontogenetic

development in the plankton, providing a link with the

substrate.

The large variation in the average density observed

can be related to preference, not only for the

substrate, but also treatment. According to Hobson

& Chess (1979), the results obtained by this kind of

trap in reef areas are difficult to interpret quantita-

tively, being more suitable for studies of taxonomic

composition. This problem is related to the fact that

the substrate is not completely flat and may allow

organisms access through openings between the

bottom and the base of the trap. The same authors

mention that during net tows, Copepoda and

Mysidacea may be overestimated, while Gammar-

idea (Amphipoda) can be underestimated, since this

group swims near the bottom at night.

The high densities found in light-trap samples

were in opposition to that observed by Alldredge &

King (1980). They demonstrated that in shallow

waters, large organisms migrated less frequently into

the water column during moonlit periods than small

forms, suggesting that this behaviour is for avoidance

of visually oriented predators.

The high densities of Oithona oculata in some

samples, especially in gravel bottom and in one coral

reef, achieved densities estimated for swarms

(Ambler 2002). According to Ambler et al. (1991),

swarms occur near seagrass beds, algal beds, and

coral reefs in both tropical and temperate areas. This

species forms swarms in light shafts during the day

and disperses to the water column at dusk (Ueda

et al. 1983; Ambler et al. 1991; Buskey et al. 1996).

Alldredge & King (1977) and Porter & Porter

(1977) suggest that most zooplankton live over

structurally complex coral substrata, but differences

among substrates communities were not found in this

study, as observed by Birkeland & Smalley (1981) and

Ohlhorst (1982). This indicates that the zooplankton

moving along the coral reefs and seagrass become

available in less complex adjacent habitats (sandy and

gravel bottom). Fenchel & Uiblein (2008) say that

Figure 8. Cluster analysis of demersal zooplankton captured with

traps at Itamaraca Island and Tamandare Bay, Pernambuco,

Brazil. Group 1: Itamaraca Island; group 2: Tamandare Bay.

Figure 7. Diversity index (bits ind�1) and evenness of demersal zooplankton captured with traps at Itamaraca Island and Tamandare Bay,

Pernambuco, Brazil.

538 P. A. M. C. Melo et al.

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most people consider soft sediments to be rather

homogeneous material notwithstanding the presence

of burrowing invertebrates; however, their study in

sediment indicate that sandy bottom is a spatially

complex habitat and on a subcentimetre scale this

marine sediment appear as spatially complex and

exciting as do coral reefs. Buhring et al. (2006)

showed that sandy sediments are highly active and

have fast turnover rates.

The diversity ranged from low to medium, being

smaller in BD, which can be attributed to the

predominance of Acartia lilljeborgi and Pseudodiapto-

mus acutus, and DL, which was dominated by

O. oculata and Foraminifera. Studies comparing the

epifauna of seagrasses and adjacent areas without

vegetation have shown higher faunal diversity and

abundance in seagrass areas (Arrivillaga & Baltz

1999; Jackson et al. 2002). However, this was not

observed in our study, mainly due to the intense

degradation of this area caused by intense human

seagrass removal. In gravel bottoms, the low diversity

is probably due to the lack of shelter. The diversity on

coral reef in our study was higher than found by

Suarez-Morales & Gasca (2000) (1.84 bits ind�1)

by horizontal surface hauls, showing the need for

other strategies to study these environments.

The differences observed among the samples, both

in terms of composition, densities, and proportions of

the species, can be explained by the different abiotic

characteristics of these environments (e.g. suspended

materials, light, salinity, temperature, oxygen con-

centration, and pH), and with the fauna, thus

presenting different patterns of diel migration. There-

fore, grouping of samples by treatment (Figure 7)

appears to be the most consistent with this approach.

Divergence in some of these samples can be related to

low density and diversity.

The positive correlation between the set of samples

both with and without light indicated that some other

factor might regulate diel migration. The other factor

that may support this migration could be the tide,

which causes a change in the community composition

through the input of oceanic species or the export of

local species. Another factor may be the season, but

some authors (Heidelberg et al. 2004) found seasonal

differences among only a few of the zooplankton taxa.

In our study significant differences between dry and

rainy seasons were observed only to the gravel bottom

only. This may be caused by the lower atypical rainfall

in the studied period.

Our research proved the effectiveness of trap-

samplers for assessment of demersal zooplankton

diversity in seagrass beds as well as coral reefs. The

differences observed between treatments showed the

importance of this type of trap in studies of com-

munities with diel migration patterns, highlighting

the need for further studies to clarify points about

the other abiotic factors that affect these commu-

nities’ structure, composition, and behaviour.

Acknowledgements

We would like to thank M. F. Costa for comments

and suggestions on the manuscript and to American

Journals Experts for editing the English. We declare

that the experiments comply with the current laws of

Brazil.

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