The macrozoobenthos of the Karavasta lagoon system (Albania): local assemblages and geographical...

ORIGINAL ARTICLE

The macrozoobenthos of the Karavasta lagoon system(Albania): local assemblages and geographical comparisonsCarlotta Nonnis Marzano1, Maria Flavia Gravina2, Alessandra Fianchini2, Giuseppe Portacci3, MyriamGherardi1, Angelo Tursi1 & Giuseppe Corriero1

1 Dipartimento di Zoologia, Universita degli Studi di Bari, Bari, Italy

2 Dipartimento di Biologia, Universita di Roma ‘Tor Vergata’, Roma, Italy

3 Istituto Sperimentale Talassografico ‘Attilio Cerruti’ CNR, Taranto, Italy

Problem

Coastal lagoons are shallow aquatic environments sepa-

rated by sand or shingle bars from the open sea, to which

they remain connected at least intermittently through one

or more restricted inlets (Bird 1994). Due to the intermit-

tent entrance of seawater (Ranasinghe & Pattiaratchi

1999), these environments are generally characterized by

large fluctuations in environmental variables, which cause

changes in the structure and distribution pattern of

organisms (Koutsoubas et al. 2000).

Salinity has long been considered the main ecological

factor able to discriminate composition and distribution

of the aquatic fauna inhabiting coastal lagoons. Within a

given basin, salinity depends on the balance between

fresh- and seawater inputs and evaporation, thus being

subject more than other variables to sudden changes. The

Venice System (1958) mainly classified lagoons on the

basis of their mean values of salinity. The categories range

from less than 5 psu of salinity (oligohaline lagoons, with

consistent freshwater inputs), to more than 40 psu

(hyperhaline lagoons, where evaporation prevails), with

middle values of 30–40 psu (most euhaline lagoons).

D’Ancona (1959) and Sacchi (1959) long ago noted the

role of ‘vivification’ in brackish ecosystems, which refers

to continuous flows of larvae from the adjacent marine

waters. Guelorget & Perthuisot (1983) stressed confine-

ment (the time of renewal of components of marine

Keywords

Coastal lagoons; macrozoobenthos;

Mediterranean Sea; salinity.

Correspondence

C. Nonnis Marzano, Dipartimento di Biologia

Animale ed Ambientale, Universita degli Studi

di Bari, via Orabona 4, 70125, Bari, Italy.

E-mail: [email protected]

Accepted: 6 April 2010

doi:10.1111/j.1439-0485.2010.00381.x

Abstract

The macrozoobenthic assemblage from the Karavasta lagoon system, the main

wetland of Albania, was studied on soft bottoms and artificial hard substrates.

Three different communities were identified: the typical ‘brackish-water com-

munity’, an ‘outlet community’ dominated by filter-feeders, and a typical ‘mar-

ine community’, inhabiting fine sandy bottoms, where polychaetes are

dominant, together with a few species of tunicates and bivalves, which colonize

hard substrates. The occurrence of build-ups of the bryozoan Conopeum seurati,

settled on the upper portion of long submerged wooden poles, was the main

feature of faunistic interest. The faunistic affinity between the zoobenthic

assemblage recorded in the hyperhaline lagoon of Karavasta and the assem-

blages from two Italian lagoons, the brackish Lake of Lesina and the euhaline

Lake of Fogliano, was considered according to the different hydrological condi-

tions and the geographical location. Hydrological confinement and salinity

were likely to be the leading factors influencing the benthic community com-

position in the Karavasta wetland area. Therefore, on a vaster spatial scale, the

isolation and geographical proximity of the ecosystems and colonizing ability

and dispersal changes of the species appear to be the main factors liable to

produce faunal variations.

Marine Ecology. ISSN 0173-9565

622 Marine Ecology 31 (2010) 622–632 ª 2010 Blackwell Verlag GmbH

origin at a given location) as the parameter primarily

determining species distribution and biological zonation

of benthic communities in lagoon ecosystems. Confine-

ment depends on numerous ecological factors, some of

which may locally assume more importance in influenc-

ing species distribution (Koutsoubas et al. 2000).

Among a conspicuous set of other hypotheses (Basset

et al. 2006), the distribution of benthic assemblages in

brackish environments has also been evaluated in terms

of dispersal and differential patterns of colonization and

extinction of animals, together with the random effects

on low diversity and small-locked lagoons (Barnes 1988;

Giangrande & Rubino 1994).

The benthic assemblages inhabiting coastal lagoons

have been deeply studied in the Mediterranean area, and

the literature is extraordinarily rich in comparison with

other coastal marine environments. However, the distri-

bution of study sites almost always coincides with that of

research centres (Basset et al. 2006) and therefore our

knowledge of the Eastern Mediterranean is incomplete

and mainly refers to some Greek lagoons (Koutsoubas

et al. 2000; Reizopoulou & Nicolaidou 2004; Dikou et al.

2008). As a consequence, not all the range of physio-

graphical and hydrological variations of lagoon ecosys-

tems in the Mediterranean region are covered: indeed,

most data on the benthic fauna from Mediterranean

lagoons refer to brackish-water environments, whereas

hyperhaline ecosystems have been neglected.

The coastal wetland of Karavasta is the largest of the

Albanian lagoons. This site, consisting of different basins

showing a wide range of salinity variations, is character-

ized by a large area where hyperhaline conditions prevail

for most of the year, with peaks exceeding 60 psu (Pano

et al. 1975; Pano 1984). Little is known about its benthic

fauna (Peja et al. 1996; Casellato et al. 1997; Casellato

1999; Beqiraj 2004; Beqiraj et al. 2007), and more infor-

mation is needed.

The present research was aimed at studying the macro-

zoobenthos from the coastal wetland of Karavasta to

determine characteristic species of the assemblages from

different basins and possible differences between them. In

addition, the faunistic results were compared with litera-

ture data referred to two Mediterranean lagoon environ-

ments located at the same latitude: the Lake of Lesina,

geographically closer to Karavasta and mostly character-

ized by brackish waters, and the Lake of Fogliano, geo-

graphically separated from Karavasta by the Italian

Peninsula and characterized by euhaline waters. The final

purpose of this comparison was to evaluate the different

role of ecological and geographic factors in determining

the macrozoobenthos distribution.

Study area

The Karavasta lagoon ecosystem (Fig. 1), the main

wetland of the Albanian coast, is located 50 km south of

Durres, between the Shkumbin and Seman river mouths.

It consists of two larger lagoons and a smaller basin

northward. The lagoon of Karavasta (surface 41 km2,

maximum depth 1.3 m) communicates with the open sea

by means of the Vahut Canal (length 700 m, maximum

depth 0.75 m) that needs regular dredging to keep its

outlet open (Chauvelon et al. 2006). An indirect connec-

tion with the sea is also realized through the adjacent

lagoon of Godulla (surface 6.5 km2, maximum depth

1.7 m). The smallest basin of Spiaxho (1.5 km2) is located

a few kilometres to the north, close to the outlet of the

Shkumbin River.

According to the literature (Chauvelon et al. 2006),

throughout the area the sediment is silty sand with clay

and organogenous deposits. Sometimes it is covered by a

thick layer of plant remains. Hard substrates have an arti-

ficial origin, and mainly consist of very numerous woo-

den poles partially submerged, positioned for navigational

purposes. According to local fishermen, most of the poles

may have been in the water for more than 10 years.

Most hydrological data available from the literature for

this site are relative to the largest lagoon of Karavasta,

whereas little information is available regarding the adja-

cent basins of Godulla and Spiaxho. The lagoon of Karav-

asta exhibits a remarkable seasonality in its water regime,Fig. 1. Map of the Albanian lagoon system of Karavasta with

sampling stations.

Nonnis Marzano, Gravina, Fianchini, Portacci, Gherardi, Tursi & Corriero Macrozoobenthos of the Karavasta lagoon system

Marine Ecology 31 (2010) 622–632 ª 2010 Blackwell Verlag GmbH 623

with summers characterized by high evaporation rates

and low precipitation. At present, it is kept isolated from

a significant part of its drainage basin by two artificial

canals that reach the sea to the north and south, respec-

tively. Moreover, most of the surface runoff is diverted to

fill some reservoirs on the surrounding hills at least

during summer, thus resulting in a severe water deficit

for the lagoon (Chauvelon et al. 2006). Due to the limited

communication with the open sea, there is a remarkable

risk of eutrophication, especially during summer when

water temperature is higher. Dystrophic crises with asso-

ciated phenomena of anoxia have already occurred in

some areas of the basin following episodes of strong algal

development (Crivelli et al. 1996; Chauvelon et al. 2006).

As regards salinity, literature data point out a very wide

range (Pano & Hysi 1982; Peja et al. 1996; Casellato 1999;

Chauvelon et al. 2006), with values reaching about 60 psu

during the dry season in a large area of the NE lagoon

(Fig. 1). Water temperature in the basin ranges from

7–8 �C to 26–27 �C, and dissolved oxygen from 3–4 to

7.5 mgÆl)1 (Pano et al. 1975; Pano 1984).

The lagoons of Godulla and Spiaxho, both having an

efficient communication with the sea (Fig. 1), show slight

variations in hydrological parameters (Pano et al. 1975;

Pano 1984).

Material and Methods

Sampling methods

Samples were collected in April 2004 from 14 stations

located in the three lagoons and in the Vahut Canal

(Fig. 1). At each station, two replicates of soft bottoms

were sampled by means of a Van Veen grab; wooden

poles submerged for about 10 years were sampled by

scraping off the substrate from a standard area of

400 cm2 (two replicates).

Samples were preserved in a 4% formaldehyde solution;

in the laboratory the fauna retained on a 1-mm-mesh

sieve was sorted and identified to the species level wher-

ever possible.

To show the spatial distribution of the benthic commu-

nity, a matrix with presence–absence data of the 96

recorded taxa was produced and evaluated by means of the

Sørensen similarity index (Sørensen 1948) and subjected to

non-metric multidimensional scaling (nMDS) ordination.

Comparative analysis with other Mediterranean lagoons

The species composition recorded from the lagoon of

Karavasta was compared with that of two other Mediter-

ranean lagoon environments, Lesina (Nonnis Marzano

et al. 2003) and Fogliano (Gravina 1986; Gravina et al.

1989). The two basins are located in the Southern Adriatic

Sea and Central Tyrrhenian Sea, respectively, on the same

latitude as the Karavasta system (Fig. 1).

The coastal Lake of Lesina (51 km2) is mainly charac-

terized by brackish water, with salinity values decreasing

from west (23–41 psu) to east (4–31 psu); it usually

shows mild water temperatures throughout the year and

moderately high values of dissolved oxygen. Two canals

connect the lake to the sea, but the seawater inflow is

moderate. The zoobenthic assemblage, consisting of 53

species, mostly halolimnobic and typical of brackish

water, colonizes large areas of the lagoon, with only a few

marine species characterizing the zone subject to direct

seawater inflow (Nonnis Marzano et al. 2003).

Fogliano (4 km2) is a shallow coastal lagoon, well con-

nected to the sea, characterized by the absence of freshwa-

ter inputs, which determines eu ⁄ hyperaline conditions. A

total of 62 zoobenthic taxa are reported for this basin,

referred to samples collected in 1983–84, when the salin-

ity ranged from 28.5 to 48.9 psu (mean value 37.4 psu)

(Gravina et al. 1989).

The faunistic affinity between the macrobenthic fauna

from Karavasta, Lesina and Fogliano was measured by

means of the Sørensen similarity index (Sørensen 1948),

considering a matrix of 118 species, with presence–

absence data. This matrix was also submitted to facto-

rial correspondence analysis (FCA) (Legendre & Legendre

1984).

Results

The 96 taxa recorded during this study (64 on soft

bottoms, 74 on hard substrates) included five demospon-

ges, three cnidarians, one turbellarian, one nemertine, 19

molluscs, 40 annelids, 18 crustaceans, 2 insects, 5 bryozo-

ans, 1 echinoderm, 1 tunicate (Table 1). The bivalve My-

tilus galloprovincialis and the amphipods Corophium

acherusicum and Microdeutopus gryllotalpa were recorded

at all the study sites.

The faunal similarity between sampling stations, analy-

sed using the Sørensen index, indicated three major clus-

ters (Fig. 2). The first cluster consisted of two groups of

stations: the stations located in front of the sea inlets (1,

6, 7, 11) in the lagoons of Karavasta and Godulla, with

similarity ranging from about 30 to 55%, and stations 9

and 10 (56% similarity), representing the community of

the outlet canals, with a prevalence of marine species and

the highest species richness values. The second cluster

comprised the Spiaxho stations (12, 13, 14), with similar-

ity values ranging between 20 and 42%. The third large

cluster (48–61% of similarity) included the more confined

stations of Karavasta (2, 3, 4, 5, 8), and was mainly

characterized by typical Mediterranean lagoon fauna. The

Macrozoobenthos of the Karavasta lagoon system Nonnis Marzano, Gravina, Fianchini, Portacci, Gherardi, Tursi & Corriero


lowest number of species was recorded at Spiaxho,

whereas a high number of species and the occurrence of

some marine elements characterized the stations under

sea influence.

The scattering of station-points in the nMDS ordina-

tion (stress = 0.18) showed a clear-cut separation of the

stations located in the canal communicating with the sea

(station 9 at Vahut Canal, station 10 at Godulla), exposed

to sea influence and water renewal, from the cluster com-

prising most of the stations of Karavasta (Fig. 3). Within

the latter, the two stations in front of the mouth are

shifted towards the ‘vivified’ pole. Finally, the station-

points of Spiaxho and station 11 of Godulla were con-

fined at the border of the nMDS plot. The cluster analysis

based on Sørensen similarities revealed three main clus-

ters of species corresponding to the different station-point

assemblages (plot not shown). Many of the species char-

acterizing the stations located at the ‘vivified’ pole in the

MDS ordination were sessile and suspension-feeders

(Sycon ciliatum, Chalinula fertilis, Sarcotragus spinosulus,

Hydroides dianthus, Ostrea edulis, Balanus improvisus, Bu-

gula gracilis, Bugula neritina, Bugula stolonifera, Schizopo-

rella errata, Botryllus schlosseri) usually occurring in areas

subjected to moderate water movement (mainly repre-

sented by tidal currents). Some marine species were con-

nected to the stations bordering the plot (Protodorvillea

kefersteini, Paradoneis lyra, Lumbrineris latreilli, Salvatoria

limbata, Musculus costulatus, Dexamine spinosa, Amphip-

holis squamata) and numerous species characterizing the

central cluster of station-points were common in sheltered

coastal waters (Loripes lacteus, M. galloprovincialis, Cerithium

rupestre, Abra segmentum, Platynereis dumerilii, Perinereis

cultrifera, Nephtys hombergi, Chthamalus montagui,

Chthamalus stellatus, Balanus amphitrite, Lekanesphaera

monodi, Sphaeroma serratum, Leptochelia savignyi, C. ach-

erusicum, Corophium insidiosum, M. gryllotalpa) and in

lagoon environments (Halichondria panicea, Heteromastus

filiformis, Polydora ciliata, Cerastoderma glaucum, Pirenella

conica, Ventrosia ventrosa, Cyclope neritea, Balanus eburn-

eus, Gammarus aequicauda, Corophium orientale, Idotea

baltica, Conopeum seurati).

Lagoon of Karavasta proper

About 31% of the 51 taxa collected in this basin were

exclusively found here. Among the most frequent species

were the gastropods Cerithium rupestre, Cyclope neritea

and Ventrosia ventrosa (this latter locally very abundant),

the polychaete Platynereis dumerilii and the isopod Idotea

baltica. On soft bottoms the bivalves Abra segmentum and

Cerastoderma glaucum were the most common species,

whereas hard substrates were markedly dominated by the

bryozoan Conopeum seurati, which colonized wide

portions of wooden poles. Its massive colonies, forming

Fig. 4. Typical appearance of Conopeum seurati build-up from the

Lagoon of Karavasta and its position around the wooden pole. Scale

bar: 2 cm.

Fig. 2. Dendrogram resulting from Sørensen similarity analysis.

Fig. 3. Multidimensional scaling ordination. Numbers refer to

sampling stations shown in Fig. 1.



bioconstructions up to 10 cm thick (Fig. 4), were formed

by the progressive overlap of laminar layers of skeletal

remains. Constructions extended from the water surface

down to about 50 cm depth, and were commonly inhab-

ited by vagile organisms such as annelids, amphipods and

isopods.

The highest values of species richness were observed

near the main canal inlets (stations 1 and 7), with 30 and

32 species, respectively. The number of species decreased

along the confinement gradient, with 10 species collected

from each of the innermost stations (stations 3 and 8).

Despite having the same number of species, the latter

stations showed a remarkable difference in species com-

position (Table 1).

Vahut Canal and lagoons of Godulla and Spiaxho

About 16% of the 38 taxa recorded in the Vahut Canal, and

32% and 19% of those collected at Godulla (53 taxa) and

Spiaxho (26 taxa), respectively, were exclusive to each site.

In the Vahut Canal (station 9), the zoobenthic assem-

blage inhabiting hard substrates was mainly composed of

sponges (Sycon ciliatum, Chalinula fertilis, Halichondria

panicea and Sarcotragus spinosulus), bryozoans (Bugula

gracilis, Bugula neritina and Bugula stolonifera) and poly-

chaetes (Hydroides dianthus and Pomatoceros lamarckii).

In the lagoon of Godulla, hard substrates (station 10)

were characterized by the dominance of the crustaceans

Leptochelia savignyi (tanaid), Idotea baltica, Sphaeroma ser-

ratum (isopods), Dexamine spinosa, Gammarus aequicauda

(amphipods), and bryozoans (the three species of Bugula

collected at Vahut canal, together with Schizoporella errata,

forming arborescent colonies up to 30 cm high).

In the lagoon of Spiaxho, Mytilus galloprovincialis and

Conopeum seurati were the most frequent species inhabit-

ing hard substrates. Polychaetes (Clymenura clypeata and

Platynereis dumerilii) and crustaceans (the amphipod

Corophium acherusicum and the decapod Carcinus medi-

terraneus) prevailed on soft bottoms. Most of the taxa

were collected near the sea inlet (station 13), whereas only

three taxa were found in the innermost area of the lagoon.

Comparative analysis with other Mediterranean lagoons

The Sørensen index, comparing macrozoobenthos com-

position between the lagoon of Karavasta and the Italian

lakes of Lesina and Fogliano, ranged in value from 16 to

39% (mean value: 27.5%). The highest similarity occurred

between Karavasta and Lesina, the lowest between Karav-

asta and Fogliano.

Only four species (Cerastoderma glaucum, Perinereis

cultrifera, Microdeutopus gryllotalpa, Conopeum seurati)

out of a total of 118 were common to the three basins.

Considering the different benthic components, crusta-

ceans showed the highest similarity values, annelids the

lowest ones.

Benthic data subject to FCA showed the following clus-

tering of sites and species on the plane defined by the first

two principal axes, which explained 58.4% and 41.5% of

the total variance, respectively (Fig. 5). The station point

of Fogliano was located at the positive pole of the first

axis, together with a set of species exclusively found in this

basin, all marine and common in sheltered coastal basins.

Points referring to Karavasta and Lesina were placed at

the opposite pole. Species points were arranged in several

clusters, separated along the second axis, according to

their different distribution in the study sites.

Discussion

The composition of aquatic fauna from the rich and

diversified system of Albanian lagoons is still largely

unknown. In the last decade several scientific papers have

been published, particularly referring to sites with high

naturalistic value like the lagoon of Karavasta (Casellato

1999; Beqiraj 2004; Beqiraj et al. 2007), though the gen-

eral level of knowledge remains low. The large number of

first records reported in this research – 40 out of 96 taxa

recorded (about 42%) – confirms this scenario. From a

faunistic point of view, most of the species found here

are very common and largely reported for other Mediter-

ranean lagoons. A particular feature is represented by the

bryozoan Conopeum seurati. This latter is known as a

truly brackish species, collected in waters with salinity val-

ues ranging between 1 and 40 PSU (Winston 1982, 1995),

where it forms thin encrusting colonies. However, it is

also able to raise its edges, overgrowing parts of the

colony (Pisano 1979; Occhipinti Ambrogi & Ambrogi

1986; Winston 1995). At Karavasta, C. seurati displays a

marked tolerance towards high salinity values (up to 60

PSU), and forms wide and thick (about 10 cm) crusts

built through the progressive overlap of different colonies

with laminar growth. In other words, each new colony

overgrows the skeletal remains of previous colonies. The

large size of such bioconstructions and the high degree of

zoaria calcification observed during the study emphasize

the peculiar hydrological aspects of the lagoon of

Karavasta. Indeed, O’Dea & Okamura (1999) stressed that

temperature, salinity, oxygen and trophic resources

(flagellates) strongly affect zoaria calcification, growth

rate, shape and size of bryozoan colonies.

The ability to build organogenous concretions is well

known for various species of bryozoans (Cocito et al. 1998,

2000); however, the massive constructions here described

for C. seurati are not reported in the literature, and consti-

tute a remarkable feature that should be considered for



Table 1. Spatial distribution of the taxa recorded in the studied lagoon system.

Taxa

Stations

Karavasta Vahut Godulla Spiaxho

1 2 3 4 5 6 7 8 9 10 11 12 13 14

Porifera

Sycon ciliatum (Fabricius) X

Chalinula fertilis Keller X X

Halichondria panicea (Pallas) X X

Haliclona sp. X X

Sarcotragus spinosulus Schmidt X X

Cnidaria

Hydroidea X X X X X X X X

Paranemonia cinerea (Contarini) X X

Actiniaria X X X X

Platyhelminthes

Turbellaria X X X

Nemertea X X X X

Mollusca

Bittium reticulatum Da Costa X

Cerithium rupestre Risso X X X X X X X

Cerithium vulgatum Bruguiere X X

Cyclope neritea (Linne) X X X X X X X X

Haminaea hydatis (Linne) X

Ventrosia ventrosa (Montagu) X X X X X

Pirenella conica (Blainville) X X X

Pusillina lineolata Michaud X

Rissoa labiosa (Montagu) X X

Abra segmentum (Recluz) X X X X X X X X X X

Cerastoderma glaucum (Poiret) X X X X X X X X X X

Loripes lacteus (Linne) X X X X X

Musculus costulatus Risso X

Mytilaster minimus (Poli) X X

Mytilus galloprovincialis Lamarck X X X X X

Ostrea edulis Linne X

Paphia aurea Gmelin X

Parvicardium exiguum Gmelin X

Solen marginatus Pulteney X X

Annelida

Arenicola sp. X

Armandia cirrhosa Philippi X

Capitella capitata (Fabricius) X X

Capitella giardi (Mesnil) X

Capitellidae sp. X X X

Cirratulidae sp. X X

Clymenura clypeata (Saint-Joseph) X X

Dorvillea rudolphii (Delle Chiaje) X

Gyptis sp. X X X

Heteromastus filiformis (Claparede) X

Hydroides dianthus (Verrill) X

Lumbrineris impatiens (Claparede) X

Lumbrineris latreilli Audouin & Milne-Edwards X

Microspio sp. X

Myrianida inermis (Saint Joseph) X X X

Nephtys hombergi Savigny X X

Nereis falsa Quatrefages X

Nereis rava Ehlers X X

Nereis zonata Malmgren X



Table 1. (Continued)

Taxa

Stations

Karavasta Vahut Godulla Spiaxho

1 2 3 4 5 6 7 8 9 10 11 12 13 14

Nereis sp. X X X X X X

Notomastus latericeus Sars X

Paraonidae sp. X

Paradoneis lyra (Southern) X

Parapionosyllis minuta (Pierantoni) X

Perinereis cultrifera (Grube) X X

Phyllodoce macrophtalma Schmarda X

Phyllodocidae sp. X X

Platynereis dumerilii (Audouin & Milne-Edwards) X X X X X X X X

Polydora ciliata (Audouin & Milne-Edwards) X X X

Polyophthalmus pictus (Dujardin) X

Pomatoceros lamarckii (Quatrefages) X X

Protodorvillea kefersteini (Mc Intosh) X

Salvatoria limbata (Claparede) X

Scoletoma tetraura (Schmarda) X X

Simplaria pseudomilitaris (Thiriot-Quievreux) X X X X X

Sphaerosyllis sp. X X

Spionidae sp. X X X X

Syllis gracilis Grube X

Syllis sp. X X X

Oligochaeta X X X X X

Crustacea

Balanus amphitrite Darwin X

Balanus eburneus Gould X X X X X X

Balanus improvisus Darwin X

Chthamalus montagui Southward X

Chthamalus stellatus (Poli) X

Leptochelia savignyi (Kroyer) X X X

Tanais dulongii (Audouin) X

Cyathura carinata (Kroyer) X X X

Idotea baltica (Pallas) X X X X X X

Lekanesphaera monodi (Arcangeli) X

Sphaeroma serratum (Fabricius) X X

Corophium acherusicum Costa X X X X X X X X

Corophium insidiosum Crawford X

Corophium orientale Schellenberg X

Dexamine spinosa (Montagu) X

Gammarus aequicauda (Martynov) X X X X X

Microdeutopus gryllotalpa Costa X X X X X X X

Carcinus mediterraneus (Czerniavsky) X X X

Insecta

Chironomid larvae X X X X X X X X X X

Larvae X X X X X

Bryozoa

Bugula gracilis Busk X X

Bugula neritina Linne X X

Bugula stolonifera Ryland X X

Conopeum seurati Canu X X X X X X

Schizoporella errata Waters X

Echinodermata

Amphipholis squamata (Delle Chiaje) X

Tunicata

Botryllus schlosseri (Pallas) X X



preservation and scientific purposes. Indeed, several works

underline the ecological role of bioconstructions in enhanc-

ing biodiversity, essentially acting as a refuge for many spe-

cies and increasing habitat complexity with solid substrates,

crevices, cavities, mud pockets, etc. (Bianchi 2001; Cocito

2004; Chemello 2009; Cocito 2009). This role, well known

for marine environments, has been recently emphasized

also for lagoon systems (Nonnis Marzano et al. 2003), thus

stressing the ecological importance and the naturalistic

value of these organogenous structures.

Most of the Karavasta lagoon system appears to be

colonized by the typical ‘brackish-water community’

belonging to the Paralic Domain or to the Euryhaline and

Eurythermal Biocoenoses (Peres 1967). However, two

other communities have been recorded: the first referred

to as the ‘outlet community’, dominated by a filter-feeder

epifaunal component, the second corresponding to a typi-

cal ‘marine community’, inhabiting fine sandy bottoms,

where polychaetes are dominant, together with a few

species of tunicates and bivalves, which colonize hard

substrates. Such faunistic differences are confirmed by

the Sørensen index and nMDS analysis, strengthening

Bianchi’s hypotheses (Bianchi 1985, 1988a,b). In general,

a marine benthic fauna typical of semi-enclosed shallow

environments prevails at Vahut Canal and at the sea

mouth of Godulla, whereas several elements usually found

in lagoons or exclusive to these environments characterize

Karavasta and Spiaxho.

The comparison between the macrozoobenthic assem-

blages from the basins of Karavasta, Lesina and Fogliano

emphasizes a marked heterogeneity of benthic fauna, even

when comparing environments geographically close to

each other (Karavasta and Lesina). The average similarity

value was less than 25%. Nevertheless, the faunistic affin-

ity changes considerably relative to different zoobenthic

taxa, being higher for crustaceans (isopods and amphi-

pods) and bryozoans and lower for polychaetes. Such dif-

ferences can be explained in terms of different

adaptability and colonization strategies of the various spe-

cies. Several species of polychaetes are able to inhabit

sheltered coastal marine areas (Giangrande et al. 1983–

1984; Lardicci et al. 1993), showing high diversity of

reproductive traits and life histories (Giangrande 1997).

Therefore, marked changes can occur in the species com-

position of worm fauna, with the functioning of the

brackish community remaining unaltered. Indeed, many

polychaete species are opportunistic, with small size, short

generation times and short larval stages, and can rapidly

increase or decrease in density, responding to biotic and

abiotic disturbances (Giangrande & Fraschetti 1996). In

contrast, among crustaceans, few marine species (Barnes

1989) of isopods and amphipods can colonize brackish

biotopes from the close marine areas, having direct devel-

opment and limited dispersal strategies as adults. It is

likely that they are weak competitors and are driven by

more competitive marine species into the various brackish

Asqu, Cten, Cins, Feni, Gins,Grub, Hdiv, Pcin, Pcil

Aseg, Bebu, Cach, Cori, Ccar,Gaeq, Hpan, Hven, Ibal, Lmon,

Mgal, Sser

Cgla, Cseu, Mgry, Pcul

Acir, Ccap, Pdum

Bamp, Cmon, Cste, Llac, Nhom,Olig, Paur, Tdul

Aova, Bner, Clin, Erap, Gmed,Hhyd, Hfil, Hele, Iche, Lpil, Llut,Mful, Ncau, Ppal, Plam, Sdec, Zp

ri, Zver

Fogliano

Lesina

Karavasta

Bimp, Bsch, Caes, Ccir, Eoli,Hnav, Hdia, Jhop, Mpal, Mlin,

–1.5

–1

–0.5

0

0.5

1

1.5

1.510.50–0.5–1–1.5

F1(58.45%)

F2

(41.

55%

)

Fig. 5. Ordination model of the species and station-points of the Karavasta, Lesina and Fogliano lagoons (based on presence–absence data) on

the plane formed by the first two axes extracted by correspondence analysis.



biotopes, where they can survive thanks to their euryha-

linity (Cognetti & Maltagliati 2000).

The multivariate analysis results suggest that in the

three basins considered, the composition of benthic com-

munities is mainly influenced by biogeographic factors.

Indeed, along the first axis the ordination model clearly

separates Tyrrhenian from Adriatic lagoons. The latter are

arranged along the second axis, in agreement with salinity

differences. According to the literature, Karavasta and

Fogliano are both characterized by salinity values exceed-

ing that of seawater (Pano 1984; Gravina 1986; Gravina

et al. 1989; Technital 1995; Crivelli et al. 1996) and may

be respectively considered as hyperhaline and euhaline

lagoons, whereas brackish conditions prevail at Lesina

(Nonnis Marzano et al. 2003, 2007).

In agreement with Cognetti (1994) and Munari &

Mistri (2008), such results support the hypothesis that

Mediterranean lagoons show remarkable biodiversity lev-

els due to the input of elements from different biogeo-

graphical regions. Most species from the Karavasta

lagoon system are autochthonous, such as Sycon cilia-

tum, Chalinula fertilis, Sarcotragus spinosulus, Pusillina

lineolata, Mytilaster minimus, Pirenella conica and Paphia

aurea. Other species are vicariant of those found in the

Tyrrhenian lagoon of Fogliano, with Idotea baltica

substituting for Idotea chelipes, Sphaeroma serratum and

Lekanesphaera monodi, substituting for Zenobiana prism-

atica among isopods; and Corophium acherusicum and

Corophium orientale substituting for Leptocheirus pilosus

among amphipods. The presence of such species can be

explained in terms of both biogeographical factors and

physiological constraints. Although lagoons represent

suitable sites for the colonization by alien species

(Occhipinti Ambrogi 2000, 2001), non-indigenous spe-

cies were not found in the studied basins, probably due

to their geographic location, far from the main traffic

routes. Such results stress the general integrity of the

Karavasta environment and are consistent with the

increasing role of human impact (commercial traffic by

sea, intensive fishery, deliberate or accidental introduc-

tion of species) on the colonization patterns of benthic

species.

On the other hand, it is well known that brackish envi-

ronments taken as a whole, at least in the same biogeo-

graphical regions, do share many halolimnobic and

cosmopolitan species, forming a typical stock of species

(Bacci 1954; Petit 1962). Clearly, the recent origin of the

majority of brackish biotopes lies at the root of the

paucity of species and uniformity of their fauna from a

macroscopic point of view. At the same time, their high

selectivity and isolation, together with the different life-

history features of the species, their colonizing capability,

dispersal chances and the impact of human activities,

determine the great diversity observed between the indi-

vidual systems.

Summary

The macrobenthos of the main wetland of the Albanian

coast was studied to determine the species composition.

It was compared with literature data on two Italian basins

(Lesina and Fogliano) located at the same latitude on the

Adriatic and Tyrrhenian coasts, respectively, to evaluate

the role of both ecological and geographical factors in

discriminating the macrozoobenthos distribution. The

lagoon ecosystem investigated consisted of two larger

lagoons (Karavasta and Godulla), the canal connecting

the former with the sea (Vahut Canal) and a smaller

basin (Spiaxho), all subject to a wide range of salinity,

with highest values up to 60 PSU. On the whole, the

macrofauna was classified into 96 taxa, 41% of which

were new records for the area. Most of the species were

exclusively found at Karavasta, where a particular feature

was represented by the bryozoan Conopeum seurati, which

is able to build wide and thick crusts on artificial hard

substrates. In the other basins, the zoobenthos inhabiting

hard substrates was mainly composed of sponges, bryozo-

ans, polychaetes, crustaceans and molluscs. Polychaetes

and crustaceans prevailed on soft bottoms. Hydrological

confinement and salinity were the leading factors influ-

encing the benthic community composition in the Karav-

asta wetland area. The comparison between Albanian and

Italian basins showed the great diversity between systems,

and different patterns of similarity according to the vari-

ous zoobenthic taxa considered. The results explain the

remarkable biodiversity of Mediterranean lagoons in

terms of input of elements coming from different biogeo-

graphical regions and different life-history traits of the

species.

Acknowledgements

Thanks to Dr Michele Gristina and Prof. Anna Occhipinti

Ambrogi for their help in bryozoan identification and to

Prof. Giuseppe Baldassarre and Dr Arben Pambuku for

logistical help in Albania. We thank the anonymous review-

ers for their valuable comments on improving this article.

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