Population dynamics of cryptogenic calcarean sponges (Porifera, Calcarea) in Southeastern Brazil

ORIGINAL ARTICLE

Population dynamics of cryptogenic calcarean sponges(Porifera, Calcarea) in Southeastern BrazilFernanda F. Cavalcanti1, Luıs Felipe Skinner2 & Michelle Klautau1

1 Universidade Federal do Rio de Janeiro, Instituto de Biologia, Departamento de Zoologia, Rio de Janeiro, RJ, Brazil

2 Universidade do Estado do Rio de Janeiro, Departamento de Ciencias, Patronato, Sao Goncalo, RJ, Brazil

Keywords

Growth; habitat selection; mortality;

Paraleucilla magna; recruitment; Sycettusa

hastifera.

Correspondence

M. Klautau, Universidade Federal do Rio de

Janeiro, Instituto de Biologia, Departamento

de Zoologia, Avenida Carlos Chagas Filho,

373, CCS, Cidade Universitaria, Rio de

Janeiro, RJ 21941-902., Brazil.

E-mail: [email protected]

Accepted: 15 September 2012

doi: 10.1111/maec.12013

Abstract

The calcarean sponge Paraleucilla magna is classified as being an invasive spe-

cies on the Mediterranean Sea, where it causes economic damages to mollusc

farms. On the Brazilian coast, this species is considered to be cryptogenic, and

information on its ecology is scarce. The same is true for Sycettusa hastifera,

another calcarean sponge with a worldwide distribution. Data on the ecology

of these species could help in elucidating their potential to become a threat if

they are found to be exotic species in Brazil. In the present work, we studied

habitat selection, growth and mortality of early juveniles of P. magna and habi-

tat selection of S. hastifera in a Marine Reserve from Southeastern Brazil, where

these species are abundant in the benthic community. Granite plates were used

for habitat selection analysis, varying in substrate inclination (vertical and hori-

zontal) and exposure to light and hydrodynamism (exposed and sheltered). To

analyse the growth and mortality rates, sponges were mapped and then mea-

sured once a week for 10 weeks. If a monitored sponge was not found in the

following week, it was considered to be dead. Our results showed that,

although P. magna and S. hastifera are capable of inhabiting substrates exposed

to different environmental conditions, they showed habitat preferences. Growth

of the juveniles of P. magna seemed not to have damaged any neighbouring

invertebrates. The mortality of juveniles of this species was higher during the

first 2 weeks of life but its causes could not be elucidated.

Introduction

To date, few studies have been carried out on habitat

selection of marine sponges (e.g. Johnson 1980; Vacelet

1981; Zea 1993; Padua et al. 2012). These studies mainly

analysed the influence of light and substrate composition

on the settlement of sponge larvae or the distribution of

adult specimens. Johnson (1980), for example, showed

that the distribution of calcarean sponges along the

length of a cave in Florida was determined by light. Zea

(1993) and Padua et al. (2012) also observed that the

highest settlement of sponge larvae was on substrates pro-

tected from sunlight. The influence of substrate composi-

tion on larvae settlement has only been tested once for

sponges (Vacelet 1981) and, in contrast to the results

observed for several other benthic organisms (e.g. corals;

Creed & Paula 2007), this factor was not relevant to this

group.

Although studies on habitat selection of marine

sponges are scarce, their growth and mortality rates have

been more frequently studied (e.g. Turon et al. 1998;

Tanaka 2002; Blanquer et al. 2008; Koopmans & Wijffels

2008; McMurray et al. 2008; Gunda & Janapala 2009;

Duckworth & Wolff 2011). Sponge growth can be influ-

enced by water motion, light, temperature, substrate, spa-

tial competition and predation, among other factors

(Cerrano et al. 2007; Loh & Pawlik 2009; Mercado-Moli-

na & Yoshioka 2009; Xue & Zhang 2009; Duckworth &

Wolff 2011). The size of the sponge (McMurray et al.

2008; Duckworth & Wolff 2011) and probably its age can

Marine Ecology (2013) 1–9 ª 2013 Blackwell Verlag GmbH 1

Marine Ecology. ISSN 0173-9565

also cause variability in its growth rates, but to date, with

the exception of a few studies (Maldonado & Young

1999; de Caralt et al. 2007; Xue & Zhang 2009), available

data comes only from adult individuals. Moreover, these

studies have focused on species almost exclusively of the

class Demospongiae (Page et al. 2005; Cebrian & Uriz

2006; de Voogd 2007; Koopmans & Wijffels 2008; Ferretti

et al. 2009) and data about Calcarea are rare (Orton

1914; Johnson 1978).

In the present work, we studied the habitat selection,

growth and mortality of the calcarean sponge Paraleucil-

la magna Klautau et al. 2004; and the habitat selection

of the calcarean sponge Sycettusa hastifera (Row 1909)

in the Marine Reserve of Arraial do Cabo, Rio de

Janeiro, Brazil. Data were obtained from juveniles grow-

ing on granite plates placed in situ and therefore the age

of each specimen (measured as weeks of life) could be

estimated. Paraleucilla magna is considered to be an

invasive species on the Mediterranean Sea, where it

causes damage to molluscs farms (Longo et al. 2007). As

both species are considered cryptogenic on the Brazilian

coast, knowledge about these parameters is very impor-

tant to evaluate if they could become a threat to the

marine ecosystem of Arraial do Cabo, should they prove

to be exotic species.

Study area

The city of Arraial do Cabo (23°52′ S 42°01′ W) is

located in Rio de Janeiro State, on the southeast coast of

Brazil (Fig. 1). This region is characterized by a strong

upwelling phenomenon, mainly between September and

March, when the South Atlantic Central Waters rise to

the surface (Yoneshigue-Valentin & Valentin 1992; Flo-

eter et al. 2001). Thus, like other regions under influence

of upwelling, Arraial do Cabo has a high diversity of

marine life. Nevertheless, in the last decades, alterations

in the composition of the sponge fauna have been

observed, with changes in abundance, and appearance

and disappearance of species (M. Klautau, personal obser-

vation).

Methods

Experimental design

The study was performed in Forno Harbour (Arraial do

Cabo) from September 2009 to January 2010 (Figs 1 and

2). Experimental structures for settlement were developed,

based on Marins et al. (2009). Two plastic boxes were

attached to each other side by side, and rugose granite

A B

Fig. 1. Study area. (A) Map of Brazil

showing Rio de Janeiro State (inset). (B) Map

of Arraial do Cabo showing Forno Harbour.

Fig. 2. Schematic representation of the experimental design and main observations over the study period.

2 Marine Ecology (2013) 1–9 ª 2013 Blackwell Verlag GmbH

Population dynamics of cryptogenic sponges Cavalcanti, Skinner & Klautau

plates (22 9 11 cm) were fixed inside and outside these

boxes to be used as substrate for the settlement of larvae

(Fig. 3A). The set, composed of the two boxes plus granite

plates, was suspended at 1 m depth, with the apertures of

the boxes turned towards the sea floor (about 7 m below

the boxes). Each set contained four granite plates, one for

each treatment: (i) inclination preference (horizontal or

vertical); and (ii) exposure preference (exposed or shel-

tered). The term sheltered is used for the plates inside the

boxes, meaning plates protected from sunlight and experi-

encing lower hydrodynamic conditions. Four sets (repli-

cates) were used in the experiment, all of them being

placed next to the artificial rocky shore of the harbour.

Habitat selection

To analyse the preference of the larvae of Paraleucilla

magna and Sycettusa hastifera for the different treatments,

the plates were maintained immersed from September

2009 until January 2010, although some interruptions

occurred due to the analyses of growth and mortality (see

below). After this period, the plates were taken out of the

sea and put inside socks (to avoid dislodgement of the

sponges during transportation), and then fixed and pre-

served in 93% ethanol. The preserved experimental plates

were photographed using a Canon G11 digital camera for

calculating the surface area of the main groups of organ-

isms that colonized the plates. For these calculations, the

software IMAGEJ 1.46 (rsbweb.nih.gov/ij/) was used. The

recruited specimens of P. magna and S. hastifera were

quantified under a stereomicroscope (surface area was not

calculated for the sponges, as juveniles were too small).

The two treatments (inclination and exposure) were con-

fined in a same set of plastic boxes. Therefore, a two-way

blocked analysis of variance (ANOVA) was performed to

compare the recruitment of P. magna and S. hastifera

among treatments [factors: inclination preference (horizon-

tal/vertical) and exposure preference (exposed/sheltered)].

Each set (composed of plastic boxes + granite plates) was

the blocking factor. All analyses were performed using the

software STATISTICA 7.0 (http://www.statsoft.com). Data

did not need to be transformed as they conformed with the

assumptions of normality and homoscedasticity.

Growth

This analysis was made only for Paraleucilla magna as

there were very few recruits of Sycettusa hastifera. After

15 days from the beginning of the experiment, the plates

were analysed weekly until December 2009 (10 weeks;

Fig. 2). For the visualization of the juveniles, the plates

were taken out of the sea and analysed with a hand mag-

nifier for a few minutes. A transparent grid was used to

record the position of each sponge on the plates and the

juveniles were measured with a plastic ruler. When an

individual became visible for the first time under a mag-

nifier, we assumed that it had lived for 1 week.

The shape of the juveniles of P. magna is tubular.

Thus, sponge volume was calculated by the equations for

solid cylinders, as follows:

1 If the sponge had a circular base (Fig. 3B), then

Vsponge = pr2h, where Vsponge = volume of the sponge

(mm3); r = radius of the circular base of the sponge

(mm); and h = height of the sponge (mm).

2 If the sponge had an elliptical base (Fig. 3B), then

Vsponge = pabh, where Vsponge = volume of the sponge

(mm3); a = semimajor axis of the base of the sponge

(mm); b = semiminor axis of the base of the sponge

(mm); and h = height of the sponge (mm).

These calculations allowed us to detect increases and

decreases in rates of growth of the sponges, providing a

better understanding on the growth dynamics, as noted

by Garrabou & Zabala (2001).

Volume dynamics (i.e. changes in volume as the weeks

progressed) was analysed for all monitored specimens of

P. magna (i.e. 72 specimens). The growth rate (i.e. how

much the specimen grew in relation to the previous

week) was calculated only for the sponges that lived for

A B

Fig. 3. (A) Set of plastic boxes and granite

plates used as substrate for the settlement of

larvae of Paraleucilla magna and Sycettusa

hastifera. (B) Measurements taken from P.

magna when the juveniles were circular or

elliptical at their bases. r, radius of the

circular base; a, semimajor axis of the

elliptical base; b, semiminor axis of the

elliptical base; h, height.


Cavalcanti, Skinner & Klautau Population dynamics of cryptogenic sponges

at least 2 weeks. Because of mortality, the number of

specimens decreased over the weeks (see Results section).

Growth rate was calculated according to the following

equation: GRt = (Vt � Vt�1)/Vt�1/d, where GRt = growth

rate of the sponge in the t week; Vt = volume of the

sponge in the t week; Vt�1 = volume of the sponge in the

t�1 week; and d = number of days between the two vol-

ume measurements (6–8 days). This equation has been

widely used in sponge growth studies (de Caralt et al.

2007; McMurray et al. 2008; Ferretti et al. 2009).

Both growth rate and volume dynamics were calculated

considering the age (weeks of life) of each juvenile and

not how long the plates were immersed. Therefore,

growth rate and volume dynamics were not influenced by

differences on the age of the juveniles.

Mortality

It was assumed that the reattachment of dislodged juve-

niles was not possible. Therefore, if a marked sponge was

not found in the following week, it was considered to be

dead. Mortality rate was calculated per week and also for

the entire period of study (referred to as final mortality).

Results

Habitat selection

After 4 months of immersion, the experimental plates

were colonized mainly by encrusting and branched bry-

ozoans, colonial ascidians, serpulid polychaetes and bar-

nacles (Fig. 4). The algae Cladophora sp. was also present

on the plates exposed to light (Fig. 4).

The settlement of Paraleucilla magna was significantly

higher on the sheltered plates than on the exposed ones

(F = 22.43, df = 1, P = 0.018; Fig. 5A) but there was no

significant difference in the number of settlers on hori-

zontal and vertical plates (F = 1.97, df = 1, P = 0.255;

Fig. 5A). As plates were put in a block design, this factor

was also tested and did not show significant differences

(F = 3.73; df = 3; P = 0.154). This indicates that the

recruitment on the structures was not dependent on the

place where each structure was placed. No interaction

was observed between the factors exposure and inclina-

tion (F = 3.72, df = 1, P = 0.149).

In contrast, the juveniles of Sycettusa hastifera did not

show any clear preference for the sheltered or the exposed

Fig. 4. Coverage (%) of the main groups of

organisms colonizing the experimental plates.

A B

Fig. 5. Number of specimens (mean value

and standard deviation) of Paraleucilla magna

(A) and Sycettusa hastifera (B) on the plates

used for the habitat selection experiment.



plates (F = 0.03; df = 1; P = 0.874), either for the vertical

or horizontal plates (F = 0.03; df = 1; P = 0.874)

(Fig. 5B). However, there was a significant interaction

between the factors inclination and exposure (F = 15.71;

df = 1; P = 0.029), meaning that the combination of

these variables influenced S. hastifera settlement. The

blocking factor was also not significant (F = 1.79; df = 3;

P = 0.322) for this species.

Growth

A total of 72 specimens of Paraleucilla magna were moni-

tored over time. At the beginning of the measurements,

the initial volumes ranged from 0.79 to 12.56 mm3. The

volume dynamics varied among individuals. A total of 31

specimens lived for only 1 week, and one specimen lived

for only 2 weeks. The latter did not modify its volume

during the course of its life. Of the 40 remaining speci-

mens, 20 juveniles grew continually, whereas a further 20

alternated between periods of growth and periods of

reduction through the weeks. Ten of the sponges that

presented a volume reduction died, whereas the other 10

continued to live and grow. Two specimens survived for

8 weeks and both presented the same volume in the first

week of life (1.57 mm3; Fig. 6). They grew over the fol-

lowing weeks but, from the seventh to the eighth week,

the largest specimen reduced in size from 2935.9 to

1884.0 mm3, whereas the smallest increased from 2245.1

to 3460.3 mm3 (Fig. 6).

The maximum average growth rate measured for juve-

niles of P. magna was 0.42 (±0.38) day�1, which occurred

in the second week of life (based on 41 specimens;

Fig. 7A). The lowest average growth rate was recorded in

the eighth week of life, 0.01 (±0.09) day�1 (based on two

specimens; Fig. 7A). The growth of the juveniles of

P. magna apparently did not interfere with any neigh-

bouring invertebrates.

Mortality

The mortality of Paraleucilla magna juveniles was high

during the first 2 weeks of life. After the first week, 43%

of the analysed sponges had died and 21% died after the

second week (Fig. 7B). The mortality rate decreased as

the sponges became older, but only two specimens sur-

vived until the end of the experiment, in December 2009,

when both had completed 8 weeks of life. Thus, at the

end of the growth experiment, the final mortality rate

was 97.2% (70 sponges). Neither of the last two surviving

specimens of P. magna was still alive in January 2010.

Discussion

Habitat selection

In Arraial do Cabo, Paraleucilla magna and Sycettusa

hastifera occur in both exposed and sheltered habitats.

Nevertheless, the former appears to be more abundant

inside caves and crevices, whereas the latter appears to be

more abundant when exposed to sunlight (Cavalcanti,

Skinner, Klautau).

In agreement with its natural distribution, P. magna

settled mainly on sheltered plates. The same has already

been found by other authors: Padua et al. (2012)Fig. 6. Volume dynamics of the two specimens of Paraleucilla magna

that completed 8 weeks of life.

A B

Fig. 7. (A) Growth rates (mean value and standard deviation) of Paraleucilla magna juveniles. Numbers within brackets correspond to the

number of specimens used to calculate the growth rate. (B) Mortality rates of the juveniles of P. magna during the experiment.



observed a higher settlement of P. magna on shaded (and

consequently sheltered) plates in Rio de Janeiro and nega-

tive photoresponse of the larva.

The sheltered plates used in the experiment performed

in Arraial do Cabo were attached inside plastic boxes,

and the exposed plates were on the outside. Thus, beyond

light, hydrodynamism is another parameter to be consid-

ered. In Forno Harbour the main hydrodynamic move-

ment is caused by ship propellers, which are turned on

several times a day. The hydrodynamic motions created

by the propellers are so high that they appear to act

simultaneously on the sponges settled inside and outside

the boxes. Nevertheless, small-scale differences between

sheltered and exposed plates cannot be discarded.

Sedimentation is also considered to be an important

factor influencing the recruitment of sponges. Vacelet

(1981) observed the highest settlement of calcarean

sponges on the underside of plates placed at depths from

47 to 130 m. As light was not available at those depths,

the preference for the protected faces was related to the

absence of sedimentation (Vacelet 1981). Hence, the

highest number of individuals inside the boxes (on both

horizontal and vertical sheltered plates) could also be due

to sedimentation. However, as individuals of P. magna

are naturally more frequent in sheltered than in exposed

places, independently of sedimentation exposure, and as

our exposed plates apparently did not present more sedi-

ment than the protected ones, it is more probable that

light (and possibly hydrodynamism) was the most impor-

tant parameter influencing P. magna habitat preferences.

Sycettusa hastifera had no preference for the analysed

parameters (exposure/protection and vertical/horizontal

position) separately. We expected that it would show a

preference for the exposed plates, as in nature this species

is found mainly exposed to the sunlight. However, an

interaction was observed and this species recruited more

in the horizontal/exposed plates and in the vertical/shel-

tered plates. Perhaps S. hastifera was not found in the

vertical and exposed plates because the algae Cladophora sp.

and bryozoans were the main groups of organisms found

on these plates, occupying more than 90% of the avail-

able substrate (Fig. 4). The real reasons for the habitat

preferences of S. hastifera remain unknown and other

parameters, such as competition or symbiosis (this spe-

cies is frequently found associated with a coralline algae),

must be investigated.

Growth

Several studies showed that, among Porifera, calcarean

sponges are pioneers in the colonization of new substrates

(Pansini et al. 1974; Vacelet 1980, 1981; Pansini & Pronz-

ato 1981). Nevertheless, little information is available on

the minimum settling time of those sponges. Some stud-

ies show that only three or four specimens of Calcarea

were present 3 months after plates have been put in the

sea (Pronzato 1972; Pansini et al. 1974) but Padua et al.

(2012) found 100 specimens on plates submerged for the

same period of time. In the present work, P. magna was

the first sponge to settle on the plates. The sponge was

observed for the first time 3 weeks after the plates had

been put in the sea, at which time bryozoans, serpulid

polychaetes and hydroids were also seen on the plates. In

the following weeks, several other sponge species were

also fixed on the plates, all of them belonging to the class

Calcarea. Demosponges, which according to previous

works settle later on the substrate (e.g. Vacelet 1981),

were not observed.

The monitoring of the specimens allowed us to observe

that coalescence of individuals, which had been previ-

ously observed in some calcarean species (Burton 1948;

Johnson 1978), did not occur in P. magna.

Volume dynamics analysis showed that the smallest

specimen is not always the youngest. Similar to several

sponge species (e.g. Crambe crambe, Hemimycale colu-

mella, Oscarella lobularis, Chondrosia reniformis, Scopalina

lophyropoda, Scopalina blanensis; Turon et al. 1998; Garr-

abou & Zabala 2001; Blanquer et al. 2008), reduction in

size was common in P. magna, and might or might not

precede the death of the sponge. Among calcarean

sponges, changes in morphology of species belonging to

the genus Clathrina, including decreases in volume, were

attributed to their reproductive period (Johnson 1978;

Gaino et al. 1996) but reproduction is probably not

related to the volume reduction of P. magna, as the indi-

viduals in our experiment were too young for reproduc-

tion. Histological sections of juveniles 1, 2, 4, 6 and

8 weeks old were prepared and reproductive elements

were not found (data not shown).

Average growth rate was higher in the second week of

life (Fig. 7A). Growth rate varied through the weeks but

was clearly lower after the fifth week of life. A possible

explanation is that, during the first 5 weeks, P. magna

was increasing in volume because it was forming its

aquiferous system (choanocyte chambers, canals and

atrium) and competing to guarantee a place on the sub-

strate; however, data on the development of this species

are lacking.

The current information on growth of calcarean

sponges is based on only a few species, and in those stud-

ies frequently the growth was calculated long after the

sponge had been observed (Dendy 1914; Orton 1914; Coe

1932). Orton (1914), for example, observed that in the

Plymouth Sound, one specimen of Sycon coronatum

reached 28.0 cm height and 2.2 cm width within

10 months, whereas specimens of Grantia compressa



reached 8.0 cm height and 3.5 cm width during the same

period of time. The surface area of Clathrina coriacea and

Guancha blanca was measured over time in California,

with groups of specimens occupying areas ranging from

3.2 to 392.4 mm2, and from 2.4 to 590.3 mm2, respec-

tively (Johnson 1978). In the present work, the volume of

a calcarean sponge was measured for the first time and

was considered a good descriptor of growth, as the juve-

niles were tubular and not encrusting.

Recently, juveniles of the Demospongiae C. crambe and

Dysidea avara showed average growth rates of 0.4 (±0.1)and �0.25 (±0.25) day�1, respectively, 60 days after set-

tlement (de Caralt et al. 2007). In the present study, two

specimens of P. magna survived for 56 days (8 weeks).

Their average growth rate was 0.01 (±0.09) day�1, less

than that observed for C. crambe but greater than the

average observed for D. avara. However, it is important

to note that de Caralt et al. (2007) calculated growth rate

based on sponge area and not volume.

Xue & Zhang (2009) analysed changes in area of juve-

niles of Hymeniacidon perlevis under different conditions

of light, temperature and water flow. At the beginning of

the experiment, juveniles measured 0.14–0.31 mm2

(depending on the treatment), and after 28 days, sizes

varied from 0.05 to 1.3 mm2 (Xue & Zhang 2009). The

same was observed for juveniles of Sigmadocia coerulea

cultivated in vitro, the surface area of which reached 1.0–2.25 mm2 14 days after settlement (Maldonado & Young

1999). Data are lacking on the growth rates of these dem-

osponge species in situ.

Mortality

High mortality rates are frequently observed in early juve-

niles of marine sponges. Studies previously performed

showed that 56.4% of individuals of Haliclona loosanoffi

and 81% of individuals of Halichondria sp. died within

4 weeks of settlement (Hartman 1958; Fell & Lewandrow-

ski 1981). The mortality rates observed in fouling com-

munities of demosponges (unidentified species) from the

Colombian Caribbean were also high, varying from 51 to

89.5% within 8 weeks (Zea 1992, 1993). Thus, although

the mortality of the juveniles of P. magna was the highest

among these studies (97.2%), it was not surprising.

Several factors could be related to the observed mortality

of P. magna. One is the emersion of the experimental

plates. Although it took only few minutes to measure the

sponges, the emersion of the plates could have stressed the

young specimens. However, although it is not common,

specimens of P. magna have already been observed in the

intertidal zone (F. F. Cavalcanti personal observation),

suggesting that P. magna can resist desiccation and air

exposure. Thus, this factor was probably not the main

cause of the high mortality observed. Other possible causes

for the mortality of P. magna are: (i) the high hydrody-

namic motion generated by ship propellers in the study

area, which could have dislodged young specimens of

P. magna; (ii) competition for space with encrusting bry-

ozoans and colonial ascidians found on the experimental

plates; and (iii) predation by sea urchins, which are abun-

dant in the study area, or fishes. Although the authors have

made some in situ observations, new experimental work

should be taken to elucidate the real influence of these

factors on the mortality of juveniles of P. magna.

Conclusions

In this study we monitored several individuals of the

calcarean sponges Paraleucilla magna and Sycettusa hastif-

era. Despite the short duration of the experiments, the

results showed that, although P. magna and S. hastifera

are capable of inhabiting substrates exposed to different

environmental conditions, each species had specific habi-

tat preferences. Paraleucilla magna was the first calcarean

sponge to occupy the available substrate, and quickly

became abundant. Sycettusa hastifera was less numerous

on the settlement plates. Mortality rates of P. magna were

high and similar to those observed in other sponges, but

factors influencing this high mortality could not be eluci-

dated. We also observed that growth of juveniles of

P. magna did not seem to damage other organisms. Nev-

ertheless, the juveniles were small compared with the size

adults can achieve. A study focusing on adult sponges

should be performed to demonstrate whether P. magna

are able to interfere with other organisms.

Our results suggest that, although P. magna and

S. hastifera cannot be classified as either native or exotic

species on the Brazilian coast, both of them appear to be

important for the beginning of the succession process for

the fouling community. More studies are necessary on

the ecological aspects of these species to help to elucidate

whether they could be a threat to the marine ecosystem

of the Marine Reserve of Arraial do Cabo.

Acknowledgements

We are grateful to Fundacao Carlos Chagas Filho de Am-

paro a Pesquisa do Estado do Rio de Janeiro (FAPERJ), to

Conselho Nacional de Desenvolvimento Cientıfico e Tec-

nologico (CNPq), and to Programa de Pos-Graduacao em

Zoologia do Museu Nacional (PPGZoo/MNRJ), for grants

and fellowships; to Lupo S.A. for the socks in which the

plates were placed before fixation; to Companhia Munici-

pal de Administracao Portuaria/Porto do Forno (COMAP)

that administers the harbour where the study was per-

formed; to Instituto Brasileiro de Meio Ambiente e



Recursos Naturais Renovaveis (IBAMA) and Instituto

Chico Mendes de Conservacao da Biodiversidade (ICM-

BIO) for research permission. We are also grateful to Maria

Teresa Szechy for the identification of the algae; to Carla

Zilberberg and Paulo Paiva for their help in the statistical

analyses; to Andre Padua for logistic support; and to Guil-

herme Muricy for critical reading of the manuscript.

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Documents

Population dynamics of cryptogenic calcarean sponges (Porifera, Calcarea) in Southeastern Brazil