succession patterns of polychaetes on algal-dominated rocky cliffs (aegean sea, eastern...

ORIGINAL ARTICLE

Succession patterns of polychaetes on algal-dominatedrocky cliffs (Aegean Sea, Eastern Mediterranean)Chryssanthi Antoniadou

School of Biology, Department of Zoology, Aristotle University of Thessaloniki, Thessaloniki, Greece

Keywords

Colonization; feeding guilds; polychaetes;

rocky cliffs; succession.

Correspondence

Chryssanthi Antoniadou, School of Biology,

Department of Zoology, Aristotle University

of Thessaloniki, Gr-54124, Thessaloniki,

Greece.

E-mail: [email protected]

Accepted: 22 April 2013

doi: 10.1111/maec.12079

Abstract

Ecological succession has been scarcely investigated on sublittoral rocky cliffs.

The few relevant studies deal with the structure of the developing community

and are limited to higher taxa or sessile forms. The objective of the present

study was to examine succession patterns on algal-dominated rocky cliffs both

at the structural (species composition) and functional (feeding guild composi-

tion) level, using Polychaeta, a dominant taxon in this marine habitat, as a ref-

erence group. Cement panels were seasonally installed on the rocky substratum

(25–30 m depth) and sampled every 3 months over a 1-year period. Twenty-

nine polychaete species were recorded, previously reported from the surround-

ing benthic community, and classified into eight feeding guilds. Most species

were assigned as sessile filter-feeders; this guild dominated in abundance and

biomass. A strong effect of the length of immersion and of the seasonal onset

of succession on the developed communities was assessed: species composition

analyses suggested convergence into a similar organization as succession pro-

ceeds, whereas the impact of starting season on succession was stronger when

analysing feeding guilds. In both cases succession was faster on panels installed

in winter. The main emerging patterns were in agreement with relevant surveys

of the entire benthic fauna, thus supporting the efficacy of polychaetes as a sur-

rogate group for studying ecological succession in the benthic marine environ-

ment.

Introduction

Sublittoral rocky cliffs are prominent habitats in the

Mediterranean, sustaining species-rich benthic communi-

ties (Garrabou et al. 1998; Terlizzi et al. 2007) due to

their high structural complexity, i.e. physical structure

(Matias et al. 2010). Rocky cliffs are highly heterogeneous

in topographic relief (Riggs et al. 1996) and are colonized

to a varying extend by algae and sessile invertebrates of

various growth forms, ranging from thin encrusting layers

to massive branching shapes. These interacting organisms

create a biotic complex where sediment is often trapped

(Antoniadou & Chintiroglou 2005). In this way, physical

structure further expands and, accordingly, a greater

diversity of niches or exploitable resources is produced

(Matias et al. 2010). These habitats are considered vulner-

able (Ballesteros 2006), being strongly affected by natural

disturbances and anthropogenic pressure (see Antoniadou

et al. 2011a), which can be also classified under the

recently proposed term ‘affectors’ (Montefalcone et al.

2011). However, disturbances are thought to be the main

causes of increased biodiversity on rocky cliffs; they create

open areas in these space-limited habitats (Jackson 1977;

Sousa 1984; Benedetti-Cecchi & Cinelli 1996; Maughan &

Barnes 2000), triggering colonization and subsequent suc-

cession (Connell & Slatyer 1977; Underwood & Anderson

1994). Both ecological processes are highly variable,

unpredictable, and stochastic (Underwood & Chapman

2006), at least during the early periods of succession

(Antoniadou et al. 2010, 2011a; Pacheco et al. 2011; Rico

et al. 2012); they operate in diverse spatial and temporal

scales and are affected by various factors, such as larval

Marine Ecology 35 (2014) 281–291 ª 2013 Blackwell Verlag GmbH 281

Marine Ecology. ISSN 0173-9565

supply, competition and predation. Accordingly, benthic

communities consist of a puzzling mixture of assemblages

at different successional stages (Sousa 1984; Bulleri &

Benedetti-Cecchi 2006; Chapman 2007), thus enhancing

regional biodiversity.

Colonization and ensuing succession, despite being

fundamental processes in benthic ecology, have been

inadequately studied on sublittoral rocky cliffs (see Anto-

niadou et al. 2010, 2011a). They both induce significant

changes in community structure and function but the rel-

evant mechanisms are highly complex, as large numbers

of species are involved and thus not understood com-

pletely. Biota information usually focuses on higher taxa,

or only sessile forms are taken into consideration,

whereas other aspects of succession related mainly to

community function, such as biomass or life and feeding

traits, have been largely overlooked.

Along the Mediterranean sublittoral zone, rocky cliffs

are occupied by algal or animal-dominated communities

according to the prevailing environmental conditions,

mainly illumination (Garrabou et al. 2002). In these com-

munities, Polychaeta is among the most dominant inver-

tebrate groups of macrobenthos, usually constituting over

one-third of both species richness and numerical abun-

dance (Antoniadou et al. 2004; Chintiroglou et al. 2004;

Giangrande et al. 2005). Furthermore, feeding biology has

been recognized to be one of the primary aspects of com-

munity function (Bremner et al. 2003) for polychaete

communities in particular (Pagliosa 2005; Antoniadou &

Chintiroglou 2006; Dom�ınguez Castanedo et al. 2012).

This animal group has been extensively studied under the

‘feeding guild’ concept, which is much broader than sim-

ple feeding interactions (Fauchald & Jumars 1979). With

this concept, differences in the (i) food particle size and

composition (microphages and macrophages), (ii) mecha-

nism of food intake (herbivores, carnivores, filter feeders,

surface deposit feeders, and burrowers), and (iii) motility

patterns associated with feeding (sessile, discretely motile,

and motile) are integrated. Therefore, polychaetes consti-

tute a representative animal group for thorough studies

of macrobenthic communities on hard-substratum habi-

tats, at both structural and functional levels.

The main task of the present study was to assess the

structure (i.e. species composition) and function (i.e.

feeding guild composition) of polychaete communities,

considering abundance and biomass data as well, during

early succession (1-year period) on a sublittoral rocky

cliff (Aegean Sea), using cement-constructed experimental

panels. Cement panels were selected for conformity rea-

sons, as they have been widely used in succession studies

(see Antoniadou et al. 2010) and also because they con-

sist of a suitable material to construct artificial reefs for

restoring benthic communities (Baine 2001).

Study area

The study was carried out in Porto Koufo Bay located in

the North Aegean Sea, Eastern Mediterranean (Fig. 1).

The sea bottom consists of an almost vertical (80–90°inclination) limestone rocky cliff down to a depth of

60 m. Algal-dominated communities occur down to

35 m depth; beyond that depth, they are replaced by ani-

mal-dominated communities (Antoniadou & Chintirog-

lou 2005). Taking into account the scarcity of data on

benthic succession in the subtidal zone, especially consid-

ering its deeper part (Antoniadou et al. 2010), the depth

zone of 25–30 m was chosen to perform the field experi-

ment. At this depth zone the turf-forming filamentous

Rhodomelacea Womersleyella setacea and Polysiphonia

spp. prevail, forming a dense carpet, sparsely interrupted

by the presence of conspicuous sessile animals such as

sponges, bryozoans, and ascidians (Antoniadou et al.

2010).

Materials and Methods

Experimental square panels (30 9 30 9 2 cm) con-

structed of rough cement were randomly deployed on the

rocky cliff, at the chosen depth zone (25–30 m), by

SCUBA diving. Four seasonal immersion experiments

were carried out during which four sets, each consisting

of 12 panels, were fixed with nails on the substratum in

April 1998, July 1998, October 1998, and January 1999

(Fig. 2). The panels of each set were arranged on the

rock, keeping a distance of 5 m from each other, at a rel-

atively homogeneous area covered by the algal carpet,

avoiding the presence of large animal species. The next

set of panels was installed keeping a distance of 1 m from

the previous one. Each set was surveyed at 3-month

Fig. 1. Map of the study area indicating the sampling field (dot).

282 Marine Ecology 35 (2014) 281–291 ª 2013 Blackwell Verlag GmbH

Succession patterns of polychaetes Antoniadou

intervals. During each sampling, three replicate panels of

each set were randomly collected. Panels were carefully

detached from the rock and kept separately in net bags of

0.25 mm mesh size. Concurrently, the main physical and

chemical parameters (i.e. temperature, salinity, pH, and

dissolved oxygen) were measured in the water column

with an autographic recorder (TOA-DKK WQC-24).

Altogether 48 samples were obtained. In the laboratory

the upper surface of each panel was photographed to esti-

mate cover of sessile organisms (e.g. algae, sponges, bry-

ozoans, ascidians), as percentage of the panel area, and

then carefully scraped using a scalpel. The collected mate-

rial was sieved (mesh opening 0.5 mm) and preserved in

an 8% formaldehyde–seawater solution. After sorting, all

polychaetes were identified to species level, counted, and

the biomass of each individual was estimated as formalin

wet weight (decalcified weight for tube-building species)

using an electronic scale (0.01 mg precision). All speci-

mens have been deposited in the Museum of the Depart-

ment of Zoology in the Aristotle University of

Thessaloniki. Polychaete species were classified to feeding

guilds according to the seminal work of Fauchald & Ju-

mars (1979) taking into account updated information on

this topic (Gambi et al. 1995; Giangrande et al. 2000;

Martin et al. 2000; Box et al. 2010; Castanedo et al. 2012;

Mattos et al. 2013).

Τhe relationships of polychaete species richness, abun-

dance, and biomass with the cover of the sessile compo-

nent of the biota were estimated using a linear regression

analysis to assess possible coactions.

Abundance biomass comparisons (ABC plots) were

constructed per immersion period for each seasonal

experiment to assess relevant accrual rates of the studied

populations. Dominance curves for abundance and

biomass were plotted by ranking species in terms of

importance on x-axis and in terms of cumulative percent-

age of abundance or biomass on y-axis using Primer soft-

ware package (Clarke & Gorley 2006).

Multivariate analyses were used to compare the simi-

larity of polychaete communities developed on the panels

according to immersion period (i.e. among samplings)

for both species and feeding guild compositions.

Non-metric multidimensional scaling (nMDS) via Bray–Curtis distances on root-transformed abundance and

biomass data was used to visualize changes in species

(or feeding guilds) composition across immersion peri-

ods. Permutational analysis of variance, PERMANOVA

(Anderson 2005), was used to test for differences in com-

munity composition among the seasonal installations

(four levels) and across periods of immersion (four lev-

els). SIMPER was used to identify the species (or feeding

guilds) responsible for any differences between the biotic

patterns observed. Matching biotic to environmental

patterns procedure, BIOENV, was used to identify the

environmental parameters that were related to the poly-

chaete community patterns during succession, and the

degree of this relation. A further matching procedure

(RELATE) using Spearman rank correlation was applied

to compare multivariate patterns deriving from the data

matrix of the entire fauna (Antoniadou et al. 2011a) with

that of polychaetes, to assess the surrogacy of the latter

animal group. Multivariate analyses were performed using

the PRIMER software package (Clarke & Gorley 2006).

Results

Species diversity, abundance, and biomass

In all, 29 polychaete species were collected from the four

sets of panels (Table 1). Spirobranchus triqueter was the

most dominant species in terms of abundance and

biomass and it was found settled on panels in all sam-

plings. Five other species –Hydroides norvegicus, Nereis zo-nata, Polyophthalmus pictus, Spirobranchus polytrema, and

Vermiliopsis infundibulum – were routinely collected from

the panels (their frequency of appearance surpassed

65%), and contributed greatly to density and/or biomass.

Filamentous algae and colonial bryozoans constituted

the sessile biota settled on panels; their cover was very

low at first (10% at 3-month immersion) and reached

less than 45% at the end of the experiment (1-year

immersion). Very strong relationships were assessed

between the cover of panels and polychaete species rich-

ness (determination coefficient R2 = 96%), abundance

(R2 = 98%), and biomass (R2 = 99%).

ABC plots produced for each seasonal experiment of

succession (Fig. 3) showed the prevalence of biomass

Fig. 2. Field sampling protocol: installation and collection dates for

each set of panels.


Antoniadou Succession patterns of polychaetes

Table

1.Taxonomic

listan

dfeed

ingguild

classificationofpolychetespeciescollected

from

pan

elsper

immersionperiodat

each

seasonal

installation.

Polychaete

species

Feed

ingguild

Springinstallation

Summer

installation

Autumninstallation

Winterinstallation

36

912

36

912

36

912

36

912

Aphroditidae

Hermionesp.

CMJ

+

Chrysopetalidae

Chrysopetalum

deb

ile(Grube,

1855)

CMX

+

Euphrosinidae

Euphrosinefoliosa

Audouin

&Milne-Ed

wards,

1833

CMX

+

Eunicidae

Eunicevittata(Delle

Chiaje,1828)

CMJ/HMJ/OM

++

++

++

+

Lysidiceninetta

Audouin

&Milne-Ed

wards,

1833

HMJ/CMJ/OM

+

Nem

atonereisunicornis(Grube,

1840)

HMJ/CMJ/OM

++

++

Glyceridae

Glycera

tesselataGrube,

1840

CDJ

++

++

++

++

Hesionidae

Kefersteinia

cirrataKeferstein,1862

CMJ

++

++

Lumbrineridae

Scoletomafunchalen

sis(Kinberg,1865)

HMJ/CMJ/OM

++

+

Nereidae

NereiszonataMalmgren,1867

CDJ

++

++

++

++

++

++

+

Platynereisdumerilli(Audouin

&

Milne-Ed

wards,

1834)

HMJ

++

Opheliidae

Polyophthalmuspictus(Dujardin,1839)

HMX/BMX

++

++

++

++

++

Phyllodocidae

Phyllodoce

mad

eirensisLangerhan

s,1880

CMX

++

++

++

+

Polynoidae

Harmothoeareo

lata

(Grube,

1860)

CMJ

++

++

++

Sabellidae

Amphiglenamed

iterranea

(Leydig,1851)

FST

++

++

++

+

Sabella

pavoninaSavigny,

1822

FST

+

Serpulidae

Hydroides

norveg

icusGunnerus,1768

FST

++

++

++

++

++

Placosteg

uscrystallinus(non

Scacchi,1836)sensu

Zibrowius,

1968

FST

++

Protula

sp.

FST

+

Serpula

concharum

Langerhan

s,1880

FST

++

+

Spirobranchuspolytrem

a(Philippi,1844)

FST

++

++

++

++

++

++

Spirobranchustriqueter

(Linnaeus,

1758)

FST

++

++

++

++

++

++

++

++



curves in most cases, especially at more advanced immer-

sion periods. Seven and nine species constituted the two

most dominant ones in terms of abundance and biomass,

respectively; seven species dominated on panels when

succession started in summer, six in spring, and four in

autumn and winter experiments. In most cases, different

species dominated abundance and biomass.

Feeding guild diversity, abundance, and biomass

Overall seven feeding guilds were detected: three carnivore

guilds (CMJ, CDJ, CMX), two herbivore ones (HMJ and

HMX), one of burrowers (BMX) and one of filter-feeders

(FST) (see Table 1 for list of acronyms). FST was the most

speciose guild (10 species) followed by CMJ (four species).

Six species following diversified feeding modes by con-

suming both algal and animal material, along with

Polyophthalmus pictus, which consumes algae or deposited

material, were classified as omnivorous (OM). FST domi-

nated in abundance and biomass, reaching 14,500 individ-

uals � m�2 and 30812.2 mg � m�2, respectively. OM and

CDJ had a density of 2489 and 1345 individuals � m�2,

respectively; the above guilds contributed significantly to

biomass, reaching 4916.66 and 2214.45 mg � m�2, respec-

tively. The remaining guilds had a very low contribution

to both abundance and biomass (fewer than 650 individu-

als � m�2 and 1825.55 mg � m�2, respectively).

The strongest relationships between the cover of sessile

biota and feeding modes (various guilds were pooled over

main feeding modes) were assessed for herbivores

(R2 = 91%, R2 = 94%, and R2 = 76% for species num-

ber, abundance, and biomass, respectively) and carnivores

(R2 = 82%, R2 = 96%, and R2 = 96%). The abundance

(R2 = 96%, and R2 = 97%) and biomass (R2 = 98%, and

R2 = 75%) of filter and deposit feeders were strongly

related, whereas no significant relationships were found

for species diversity.

Early succession patterns of polychaete communities:

species and feeding guild composition

Multidimensional analysis of species abundance or bio-

mass data over immersion periods (Fig. 4A,B) produced

a rather similar pattern: samples from the succession

experiment that started in spring were separated together

with early stages from the summer (3- and 6-month

immersions) and the winter experiments (3-month

immersion) (Group I), whereas almost all other samples

were arranged in a second group (Group II). When anal-

ysing community structure at a functional level, the same

overall pattern was found; the same single group of sam-

ples diverged (Group II), whereas samples within Group

I were arranged differently into subgroups, as the veryTable

1.Continued

Polychaete

species

Feed

ingguild

Springinstallation

Summer

installation

Autumninstallation

Winterinstallation

36

912

36

912

36

912

36

912

Vermiliopsisinfundibulum

(Philippi,1844)

FST

++

++

++

++

++

Spirorbidae

Spirorbissp.

FST

++

Syllidae

Salvatorialim

bata(Clapar� ed

e,1868)

HMJ

+

SphaerosyllispiriferaClapar� ed

e,1868

ΗΜJ

++

++

SyllishyalinaGrube,

1863

CMJ/HMJ/ΟΜ

++

+

SyllisproliferaKrohn,1852

CMJ/HMJ/ΟΜ

+

SyllisvittataGrube,

1840

CMJ

++

+

Meanden

sity

(individuals�m

�2)

41

189

104

152

104

59

411

533

304

667

537

507

78

389

915

1348

Meanbiomass(m

g�m

�2)

36

1064

270

1685

252

169

2048

2096

744

1734

2557

2112

213

2579

2206

6749

FST,

filter

feed

ers,

sessile,tentaculated;CMX,carnivores,

motile,unarmed

pharynx;

CMJ,

carnivores,

motile,jawed

pharynx;

CDJ,

carnivores,

discretelymotile,jawed

pharynx;

HMJ,

herbivores,

motile,jawed

pharynx;

HMX,herbivores,

motile,unarmed

pharynx;

BMX,burrowers,

motile,unarmed

pharynx;

OM,omnivores;

3–1

2=im

mersionperiodsin

months.



A B

C D

Fig. 3. ABC curves constructed per immersion period at each seasonal onset of succession experiment (3A = spring experiment, 3B = summer

experiment, 3C = autumn experiment, 3D = winter experiment). Bright line = abundance curve, dark line = biomass curve. E.f., Euphrosine

foliosa; E.v., Eunice vittata; H.a., Harmothoe areolata; H.n., Hydroides norvegicus; N.z., Nereis zonata; P.p., Polyophthalmus pictus; S.f.,

Scoletoma funchalensis; Sp.p., Spirobranchus polytrema; Sp.t., Spirobranchus triqueter; V.i., Vermiliopsis infundibulum.



early stage (3-month immersion) was separated when

succession started in winter (Fig. 5A,B). When feeding

guild abundance was analysed, the early stages (3- and 6-

month immersions) from the succession experiment

started in summer were placed together with the more

advanced stages of the relevant spring experiment

(Fig. 5A). When feeding guild biomass was analysed, the

6- and 12-month immersion samples of the spring exper-

iment were placed in a separated sub-group within Group

I (Fig. 5B). SIMPER analysis showed that seven species

(Spirobranchus polytrema, Spirobranchus triqueter, Poly-

ophthalmus pictus, Vermiliopsis infundibulum, Nereis

zonata, Hydroides norvegicus, and Eunice vittata) contrib-

uted 60% of the average dissimilarity of polychaete

assemblages during succession in terms of abundance; the

above species, excluding P. pictus, contributed the same

percentage of average dissimilarity in terms of biomass.

Applying the same analysis on feeding guild data, two

guilds contributed 60% to the average dissimilarity for

both abundance and biomass (i.e. FST and OM). Two-

way PERMANOVA showed that polychaete community

structure was significantly affected by both the immersion

period and the season of installation, considering abun-

dance (pseudo-F = 2.32, P < 0.05 and pseudo-F = 5.67,

P < 0.05) and biomass (pseudo-F = 2.37, P < 0.05 and

pseudo-F = 2.14, P < 0.05) as well. In both cases a signif-

icant interaction between these two factors was detected

(pseudo-F = 2.11, P < 0.05 and pseudo-F = 1.83, P <0.05 for abundance and biomass data, respectively), mak-

ing further comparisons impossible. Applying the same

analysis on feeding guild abundance data, similar results

were found: immersion period (pseudo-F = 3.68, P <0.05) and starting season of succession (pseudo-F = 5.79,

P < 0.05) significantly affected the trophic structure of

polychaete assemblages, with a significant interaction

between the studied factors (pseudo-F = 2.94, P < 0.05).

A

B

Fig. 4. Non-metric multidimensional scaling ordination of polychaete

community structure over immersion periods in seasonal installations,

for species abundance (A) and biomass (B), based on Bray–Curtis

similarity index (Sp = spring, S = summer, A = autumn, W = winter,

3–12 = months of immersion). Triangle marks are centroids of

samples.

A

B

Fig. 5. Non-metric multidimensional scaling ordination of polychaete

community structure over immersion periods in seasonal installations,

for feeding guild abundance (A) and biomass (B), based on Bray–

Curtis similarity index (Sp = spring, S = summer, A = autumn,

W = winter, 3–12 = months of immersion). Triangle marks are

centroids of samples.



Considering biomass data, only the starting season of

succession had a significant effect (pseudo-F = 1.83,

P < 0.05) on feeding guild composition. BIOENV analy-

sis showed that none of the measured environmental

variables was related to the biotic pattern of the develop-

ing polychaete communities, either for species or feeding

guild composition (rs < 0.05). RELATE analysis showed

that the similarity matrices derived from the entire fauna

abundance matrix and from polychaete abundance matrix

were highly correlated (rs = 0.68).

Discussion

Polychaete diversity in algal-dominated Mediterranean

communities ranges from 25 to 152 species with a

decreasing trend as depth or pollution increases, or as

habitat complexity decreases (see Antoniadou & Chinti-

roglou 2005 for a thorough review on sublittoral hard

substrata Mediterranean biodiversity). The field experi-

ments carried out revealed the presence of 29 polychaete

species classified into eight feeding guilds, previously

reported to live in the surrounding algal-dominated com-

munity (Antoniadou & Chintiroglou 2005), confirming

the role of the latter as the basic pool of colonizers (An-

toniadou et al. 2010, 2011a). However, these numbers

correspond to just 37% of the species richness and 57%

of the feeding guild richness reported from the neighbor-

ing community (Antoniadou et al. 2004; Antoniadou &

Chintiroglou 2006). Thus, an impoverished community

was detected at both the structural and functional level,

especially considering each seasonal succession experi-

ment separately. These results can be indicative of the

low ‘adjustment stability’ (Menge 1975) or ‘engineering

resilience’ (Montefalcone et al. 2011), i.e. the low ability

of algal-dominated communities on sublittoral rocky

cliffs to recover after a perturbation. Certainly the current

study covers a relatively short time of succession, as only

early patterns were surveyed; further long-term studies

are crucial to follow the fate of the developing communi-

ties and estimate the time needed for recovery.

Under the recent definition of resilience by Folke

(2006), emphasis is given to the ability of a biotic system

to maintain its original function, through either recovery

of the original state or reorganization in a new context

(see also Montefalcone et al. 2011 for a comprehensive

review of the term). Therefore, functional aspects of suc-

cession are mandatory to understand ecosystem change

for conserving and restoring purposes. According to the

results of the present study, the developing polychaete

community had poor functional organization. Surface

deposit feeders were absent, although highly represented

by four guilds in the surrounding benthic community,

where they exploited resources from the sediment

entrapped among the algal turfs (Kelaher et al. 2001;

Gorgula & Conell 2004; Antoniadou & Chintiroglou

2006). Another difference is the very low abundance of

herbivores, which appeared only at more advanced

immersion periods, despite their increased density in the

algal turfs (Antoniadou & Chintiroglou 2006).

Apart from some tube-building, filter-feeding polychae-

tes, which directly settle on panels after the formation of

biofilm, most species require the development of some

structural elements prior to their settlement. This implies

a strong relation between the development of the sessile

component and the associated motile fauna during suc-

cession (Dean & Connell 1987b; Antoniadou et al. 2010,

2011a). This strong relationship is supported by the

results of the present study. During early succession,

about 10% of the panel surface was covered by sessile

biota (mostly algae and encrusting bryozoan) and this

percentage increased with immersion period; the associ-

ated polychaete fauna responded to this trend by increas-

ing diversity, abundance, and biomass at both structural

(species) and functional levels (feeding guilds). The

strongest relationship considering functional diversity was

assessed for herbivores. So, as algae gradually covered

experimental panels they attracted herbivore polychaetes

on their turfs; the latter probably respond to increasing

habitat complexity taking advantage of the newly avail-

able ecological niche offering food, refuge, and living

space.

The same overall species and feeding guilds have been

reported to colonize experimental panels in another,

organically rich area of the Aegean Sea (Antoniadou et al.

2011b) but with big differences in the abundance and

contribution of carnivores. This implies that pioneer spe-

cies in early succession of sublittoral hard substrata, such

as Spirobranchus triqueter and Spirobranchus polytrema,

are persistent regardless of the studied habitat; the latter,

however, severely influences quantitative aspects of com-

munity structure and function, such as the abundance

and biomass.

Focusing on the parallel examination of species abun-

dance and biomass, the dominance of very few species is

apparent. In most cases S. triqueter prevailed when con-

sidering both parameters and this pattern persisted in all

immersion periods. Serpulid worms were particularly

favoured when succession started in winter. In general,

the biomass curve lay above the abundance curve as suc-

cession proceeded. However, this trend was not consistent

in all seasonal experiments or immersion periods. For

example, the two curves intermingled at the more

advanced 12-month immersion period when succession

started in winter. It seems that successful colonizers expe-

rience different rates of abundance and biomass growth

as succession progresses, according to their unique life



history traits. These traits, especially reproduction output

and recruitment success, are subjected to severe temporal

fluctuations (Fraschetti et al. 2003; Watson & Barnes

2004); for instance, the most dominant polychaete S. tri-

queter has been reported to show great seasonal and

inter-annual variability in recruitment (Castric-Fey 1983;

Cotter et al. 2003a,b).

Polychaetes are one of the three dominant taxa in suc-

cession studies examining vagile fauna, mollusks and

crustaceans being the others (Dean & Connell 1987a;

Kocak et al. 1999; Olabarria 2002). In relevant studies

(Antoniadou et al. 2010, 2011a), although not consider-

ing the functional aspects of the developed community, a

similar pattern of increasing diversity and abundance

through time has been reported. Furthermore, a good

agreement of the ordinations derived from the entire

benthic fauna and the polychaete fauna separately was

found in the present study. This conformity supports the

role of polychaetes as a surrogate group in succession

studies.

Seasonal changes of polychaetes associated with algal-

dominated communities on sublittoral rocky cliffs have

been demonstrated in the composition of taxa (Antonia-

dou et al. 2004) and feeding guilds (Antoniadou & Chin-

tiroglou 2006). The season in which succession starts has

been recognized as among the important factors modulat-

ing its outcome; larval supply is linked to season, deter-

mining in this way the availability of first colonizers

(Anderson & Underwood 1994; Qvafordt et al. 2006),

which after having occupied available space, interact with

new, potential colonizers, facilitating or inhibiting their

settlement (Dean & Hurd 1980). Our results showed that

the starting season severely affected polychaete commu-

nity structure and function. Colonization was favored

when succession started in winter, as more species and

individuals successfully settled on these panels and the

developing communities supported higher functional

diversity. A similar trend was observed when the whole

benthic fauna was analysed (Antoniadou et al. 2011a),

whereas another study reported strong seasonal effects

only on species abundance (Underwood & Anderson

1994).

Whether succession in marine environment follows

predictable, sequential stages remains questionable. Most

studies suggest convergence towards a common structure

as succession proceeds, despite strong initial differences

(Foster et al. 2003; Underwood & Chapman 2006; Anto-

niadou et al. 2010, 2011a; Pacheco et al. 2011; but see

also Brown & Swearingen 1998 reporting the lack of any

unidirectional sequence of succession). Increased diver-

gence of the developed polychaete communities during

very early succession stages has been assessed, in confor-

mity with previous works (Antoniadou et al. 2011a). The

impact of starting season on succession was stronger

when analysing feeding guilds; thus, convergence towards

a common structure cannot be suggested in the latter

case, at least not during the temporal scale of the present

study. Further experiments for much longer periods are

therefore required to address functional issues of succes-

sion in the marine environment.

Conclusions

The results of the present study showed that polychaetes

can be used as a surrogate group in marine benthic suc-

cession studies. Three main patterns emerged: (i) an

increase of diversity, abundance, and biomass of coloniz-

ers as succession proceeded; (ii) a profound effect of the

seasonal onset of colonization on benthic succession at

initial stages with convergence towards a similar structure

as succession progressed; and (iii) a faster rate of succes-

sion when started in winter. These were in agreement

with relevant works in which total benthic fauna was

treated. The analyses of the developed polychaete com-

munities showed strong conformity between structural

and functional levels of organization, with a higher

impact of seasonality in the latter.

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