Species composition and patterns of diversity of macroalgal ...

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Journal of Natural History 

Vol. 44, Nos. 1–2, January 2010, 1–22 

TNAH 0022-2933 1464-5262 Journal of Natural History History, Vol. 1, No. 1, Oct 2009: pp. 0–0 

Species composition and patterns of diversity of macroalgal coralligenous 

assemblages in the north-western Mediterranean Sea 

Journal L. Piazzi of et Natural al. History 

L. Piazzia , D. Balatab *, E. Cecchic , F. Cinellia and G. Sartonid a Dipartimento di Biologia, Università di Pisa, Via A. Derna 1, 56126, Pisa, Italy; b School of 

Biological and Environmental Sciences, University College Dublin, Science Building West, 

Dublin 4, Ireland; c ARPAT – Agenzia Regionale per la Protezione Ambientale della Toscana, 

Via Marradi 114, 57126 Livorno, Italy; d Dipartimento di Biologia Vegetale, Università di 

Firenze, Via La Pira 4, 50121 Firenze, Italy 

(Received 17 March 2009; final version received 30 September 2009) 

This paper contributes to the knowledge of macroalgal coralligenous systems. The 

species composition of coralligenous assemblages of Tuscany’s coasts and islands 

(north-western Mediterranean Sea) was examined and different aspects of alpha 

and beta diversity were evaluated. The encrusting layer of coralligenous habitat in 

the study areas mostly comprised the Corallinales Mesophyllum alternans. A total 

of 187 epiphytic macroalgal species was observed, among them 29 Ochrophyta, 14 

Chlorophyta and 144 Rhodophyta. In the studied area, coralligenous assemblages 

show a similar structure across large scales, as 41species were common to all the 

studied locations. Multivariate analysis showed that locations along continental 

coasts were distinct from islands. Vertical and horizontal substrata were distinct 

on islands, but not in coastal locations. The studied assemblages showed a high 

diversity, especially in relation to the smaller spatial scales examined. 

Keywords: biodiversity; coralligenous assemblages; macroalgae; Mediterranean Sea 

Introduction 

Coralligenous macroalgal habitats are a peculiar feature of deep subtidal systems in 

the Mediterranean Sea, characterized by calcareous structures produced by Corallinales 

(Rhodophyta) (Ballesteros 2006). 

Coralligenous assemblages can develop on both rocky and soft bottoms in 

relatively constant conditions of temperature, currents, salinity and under reduced 

irradiance (Garrabou and Ballesteros 2000). Corallinales constitute a secondary substrate 

that facilitates the settlement of a large amount of algae and sessile invertebrates. 

The composition of animal and algal species comprising coralligenous assemblages has 

been widely studied (Laborel 1961, 1987; Laubier 1966; Boudouresque 1973; Augier and 

Boudouresque 1975; Hong 1982, 1983; Gili and Ros 1985) and, recently, many 

ecological aspects relating to spatial and temporal variability and the influence of 

human-induced alterations have been investigated (Ferdeghini et al. 2000; Cocito 

et al. 2002; Piazzi, Balata, Perusati et al. 2004; Balata, Piazzi et al. 2005; Balata, 

Acunto et al. 2006; Piazzi et al. 2007). However, coralligenous assemblages are less 

well known in comparison to shallower assemblages and many ecological aspects remain 

to be investigated. A fundamental aspect is biodiversity. Studies of the biodiversity of 

*Corresponding author. Email: david.balata@ucd.ie 

ISSN 0022-2933 print/ISSN 1464-5262 online 

© 2010 Taylor & Francis 

DOI: 10.1080/00222930903377547 

http://www.informaworld.com


2 L. Piazzi et al. 

coralligenous assemblages normally refer to a determinate spatial scale, and are seldom, 

if ever, conducted at scales grater than a few kilometres (Cocito et al. 2002: 

Piazzi, Balata, Perusati et al. 2004; Piazzi et al. 2007; Balata et al. 2005; Balata, Piazzi 

and Cinelli 2007). Biodiversity can vary at different spatial and temporal scales 

(Whittaker 1972; Gray 2000) and the correspondence between values of diversity at 

small and large spatial scales is not self-evident, but depends on the patterns of 

spatial variability of each system (Gray 2000; Gering and Crist 2002). Thus, high 

values of diversity at a small spatial scale may correspond to low values of diversity at 

a larger spatial scale and vice versa, making the patterns dependent on the spatial 

scale examined. 

Moreover, the acknowledgement that environmental changes are expected to 

accelerate in this century makes it crucial to understand how distribution limits are 

likely to be affected by those alterations (Parmesan et al. 2005). In this context, 

understanding of the natural geographical limits and distribution of species is central 

to conserving biodiversity and managing ecosystems for long-term viability and 

sustainability. In fact, floristic lists are often an important source of botanical 

information for a particular area and may serve as a useful starting point for more 

detailed study. Such lists may be used for general comparisons of the vegetation of 

different localities, or that of the same locality at different times. 

Another important aspect of the biodiversity of a system, though rarely considered, 

is the variability in species composition and abundance among habitats or 

along gradients, normally referred to as beta diversity (Gray 1997; Legendre et al. 

2005). In fact, in coralligenous environments, the slope of the substratum plays a 

prominent role in the maintenance of beta diversity (Gaines 1985; Benedetti-Cecchi 

et al. 2000, Balata, Piazzi and Benedetti-Cecchi 2007). Vertical substrata usually support 

distinct assemblages compared with those occurring on horizontal surfaces 

owing to differences in light conditions, susceptibility to colonization, and retention 

of sediment (Witman and Sebens 1991; Baynes 1999). 

The aim of this paper is to contribute to the knowledge of coralligenous systems 

through the assessment of the composition and frequency of macroalgal species in a 

large area of the north-western Mediterranean Sea and the determining of species 

richness over different spatial scales (from a few meters to several kilometres). 

Moreover, we evaluate patterns of alpha and beta diversity comparing assemblages 

of different habitats (vertical vs. horizontal surfaces) and in different environmental 

conditions (continent vs. island). 

Material and methods 

The study area consisted of the coasts and the islands of Tuscany (north-western 

Mediterranean Sea). Twelve locations were considered: eight within the Tuscan 

Archipelago National Park islands, two along the continental coasts (Livorno, 

Argentario) and two on rocky banks (Meloria, Vada) situated a few kilometres off 

the coast (Figure 1). 

The floristic list was compiled by taking into account published (Piazzi and 

Cinelli 2002; Piazzi, Balata and Cinelli 2004; Piazzi, Balata, Perusati et al. 2004; 

Piazzi et al. 2007; Balata, Piazzi and Cinelli 2007) and unpublished data referring to 

samples collected from 1990 to 2005. A total of 250 samples were gathered from 13 

localities. Samples were collected on vertical substrates between 30 and 50 m depth;


Journal of Natural History 3 

Figure 1. Map of the study area. Notes: L, Livorno; A, Argentario; V, Vada; Me, Meloria; 

Go, Gorgona; C, Capraia; E, Elba; P, Pianosa; Mo, Montecristo; F, Formiche di Grosseto; 

Gn, Giannutri; Gi, Giglio. 

horizontal substrate was sampled in four locations (Table 1). Each sample consisted 

of all organisms within 400 cm 2 of surface, removed using a hammer and a chisel. 

Taxonomic nomenclature followed Guiry and Guiry (2009). The frequency of each 

species was evaluated as the percentage of samples in which the species was found. 

Herbarium specimens and slide preparations are deposited in the Herbarium 

Universitatis Florentinae (FI) (Prof. Sartoni collection). 

Alpha diversity was evaluated as species richness at four different spatial scales 

(Gray 2000): plot diversity (each replicate sampling of 400 cm 2 of surface), sample 

diversity (surfaces of a few square metres where three plots were collected), site diversity 

(sites within 1 km of coastline where more sample surfaces were present), location



Table 1. Floristic list of coralligenous macroalgal assemblages. 

L V A Me Go C E Mo Gi P Gn F LH VH GnH GoH F% 

Fucophyceae 

Arthrocladia villosa (Hudson) Duby − − − − − − − + − − − + − − − + 1.6 

Asperococcus bullosus J.V. Lamouroux − − − − − + − + − − + + − + + + 4.0 

Carpomitra costata (Stackhouse) Batters − − − − − − − − − − − − − − − + 1.2 

Cladosiphon irregularis (Sauvageau) Kylin − + − − − + − − − + − − − + − − 0.8 

Cutleria chilosa (Falkenberg) P.C. Silva − + + + + + + + + + + + + + + + 42.8 

(sporophyte “Aglaozonia”) 

Cystoseira spinosa Sauvageau − − − − − + − + − + − − + − + + 4.0 

Cystoseira zosteroides C. Agardh − − − − − − − − − − − − − − + + 2.8 

Dictyopteris lucida M.A. Ribera Siguan, − − − − − − − + − − − − − − − − 0.8 

A. Gómez Garreta, I. Pérez Ruzafa, 

M.C. Barceló Martí and J. Rull Lluch 

Dictyopteris polypodioides (A.P. De − − + − − − + − − + − + − + + + 10.4 

Candolle) J.V. Lamouroux 

Dictyota dichotoma (Hudson) J.V. 

− − − − − − − − − − − − + + + − 1.6 

Lamouroux 

Dictyota linearis (C. Agardh) Greville + + + + + + + + + + + + + + + + 51.6 

Discosporangium mesarthrocarpum 

− − − − − + − + − − − − − − + + 0.8 

(Meneghini) Hauck 

Elachista intermedia P.L. and H.M. − − − − − − − − − − − − − − − + 1.2 

Crouan 

Elachista stellaris Areschough − − − − − − − − − − + − − − − − 0.8 

Gontrania lubrica Sauvageau − − − − − − − − − + − − − − − − 0.4 

Halopteris filicina (Grateloup) Kützing + + + + + + + + + + + + + + + + 73.6 

Nemacystus flexuosus (C. Agardh) Kylin − − − − − − − − − − − − − − − + 0.8 

var. giraudyi (J. Agardh) De Jong 

Lobophora variegata (J.V. Lamouroux) − − − − − + − − − − − − − − − − 0.4 

Womersley ex E.C. Oliveira 

Nereia filiformis (J. Agardh) Zanardini + + + + + + + + + + + + + + + + 22.0



Phyllariopsis brevipes (C. Agardh) Henry − − − − − − − − − − − − − − − + 0.8 

and South 

Sargassum hornschuchii C. Agardh − − − − − − − − − − − − − − + + 1.2 

Spatoglossum solieri (Chauvin ex 

− − − − − − − − − − − − − − − + 0.4 

Montagne) Kützing 

Spermatochnus paradoxus (P.H. Roth) − − − − − − − − − + − − − − + 1.2 

Kützing 

Sphacelaria cirrosa (P.H. Roth) C. Agardh + + + + + + + + + + + + + + + + 48.8 

Sphacelaria plumula Zanardini − + + + + + + + + + + − − + + + 22.8 

Sporochnus pedunculatus (Hudson) C. − − − − − − − + − − + + − − + + 4.8 

Agardh 

Stictyosiphon adriaticus Kützing − − − − − − − − − + − − − − + − 2.4 

Stilophora tenella (Esper) P.C. Silva − − − − − − − + − − − − − − + + 1.6 

Zanardinia typus (Nardo) P.C. Silva − − + + − + + − + + − + + + + + 28.0 

Chlorophyta 

Caulerpa racemosa var. cylindracea − − − − − − + − − − − − + − − − 1.6 

(Sonder) Verlaque, Huisman and 

Boudouresque 

Caulerpa taxifolia (Vahl) C. Agardh − − − − − − + − − − − − − − − − 0.4 

Cladophora coelothrix Kützing − − + + + − + + − − − + − − − − 8.4 

Cladophora echinus (Biasoletto) Kützing + + + + + + + + + + + − + + + + 22.0 

Cladophora pellucida (Hudson) Kützing − − − − − − − + − − − + − + − − 1.2 

Cladophora prolifera (Roth) Kützing + + + + + + + + + + + + + + + + 22.4 

Codium coralloides (Kützing) P.C. Silva − − − − − − + + + − − − − − − − 0.8 

Derbesia tenuissima (Moris and De − − − − − − − + + − − − − − − − 1.2 

Notaris) P.L. Crouan and H.M. Crouan 

Flabellia petiolata (Turra) Nizamuddin + + + + + + + + + + + + + + + + 72.0 

Halimeda tuna (J. Ellis and Solander) J.V. + + + + + + + + + + + + + + + + 52.2 

Lamouroux 

Microdictyon tenuius Decaisne ex J.E. Gray − + − + + + + + + + + + − + + + 6.8 

Palmophyllum crassum (Naccari) 

+ + + + + + + + + + + + + + + + 34.8 

Rabenhorst 

(Continued)



Table 1. (Continued). 


Pseudochlorodesmis furcellata (Zanardini) + + + + + + + + + + + + + + + + 79.2 

Børgesen 

Valonia macrophysa Kützing − − + + + + + + + + + + + + + + 45.2 

Rhodophyta 

Acrodiscus vidovichii (Meneghini) 

− − + + + + + + + + + + + + + + 36.8 

Zanardini 

Acrosorium ciliolatum (Harvey) Kylin + − + + + − − + − − + + − + − − 16.4 

Acrothamnion preissii (Sonder) Wollaston + + + + + + + + − + − + + + − + 49.6 

Aglaothamnion tenuissimum 

+ + + + + + + + + + + + + + − + 22.8 

(Bonnemaison) Feldmann-Mazoyer 

Aglaothamnion tripinnatum (C. Agardh) − + − − − − − − − − + − + + − − 1.2 

Feldmann-Mazoyer 

Amphiroa rubra (Philippi) Woelkerling − − − − − − − + − + − − + − + − 7.6 

Antithamnion cruciatum (C. Agardh) + − − + + − + + + − + + + + + + 15.6 

Nägeli 

Antithamnion heterocladum Funk + − − − + + − + − − − − − + − − 3.2 

Anthithamnion piliferum Cormaci and − − − − − − − + − − − − + − − − 5.6 

G. Furnari 

Antithamnion tenuissimum (Hauck) + + + − + + + − + + + + + + + + 20.4 

Schiffner 

Apoglossum gregarium (Dawson) M.J. − − − − − − − + − − − − − − + − 1.2 

Winne 

Apoglossum ruscifolium (Turner) J. Agardh + + + + + + + + + + + + − + − − 33.6 

Balliella cladoderma (Zanardini) 

− − − − − − − − − − + − − − + + 4.4 

Athanasiadis 

Botryocladia botryoides (Wulfen) 

+ + + + + + + + + + + + + + + + 40.0 

Feldmann 

Botryocladia chiajeana (Meneghini) Kylin − − − − − − + + − − − − − − − − 0.8



Brongniartella byssoides (Goodenough and − − − − − − − + − + − − − − − − 1.2 

Woodward) F. Schmitz 

Ceramium bertholdii Funk + − − − + + + + + + + + + + + + 23.2 

Ceramium bisporum Ballantine − − − − − + − − − − − − − − − + 0.8 

Ceramium cimbricum H.E. Petersen + + − − + + + − − + + − + + − − 14.8 

Ceramium circinatum (Kützing) J. Agardh − − − − − − − − − − − − + − + − 2.4 

Ceramium codii (H. Richards) Feldmann- + + + + + + + + + + + + + + + + 49.6 

Mazoyer 

Ceramium comptum Børgesen − − − − − − + − − − − + − − − 0.8 

Ceramium diaphanum (Lightfoot) P.H. + + + − + + + + + + + + − + + + 14.8 

Roth 

Ceramium flaccidum (Kützing) Ardissone − − + − + + + + + + + + + + − + 17.6 

Ceramium giacconei Cormaci and G. − − − − − − − + − − − − − − − + 0.8 

Furnari 

Ceramium graecum Lazaridou and 

− − − − − − + − − − − − − − − − 1.2 

Boudouresque 

Ceramium tenerrimum (G. Martens) + − + + + − − − + + + + − − − − 9.2 

Okamura 

Champia parvula (C. Agardh) Harvey − + + − + + + + + + + + + + + + 20.4 

Chondria capillaris (Hudson) M.J. Wynne + + + + + + + + + + + + + + + + 26.8 

Chondria dasyphylla (Woodward) C. − − + + − + + + + + − + + − − + 16.8 

Agardh 

Chrysymenia ventricosa (J.V. Lamouroux) − − − − − + − − + + − + − + + + 5.6 

J. Agardh 

Chylocladia verticillata (Lightfoot) Bliding − − − + + − − − − − − − − − − − 2.8 

Contarinia peyssonneliaeformis Zanardini + − + + − + + − − + + − + − − − 12.4 

Contarinia squamariae (Meneghini) + + + + + + + + + + + + + + + + 24.8 

Denizot 

Cottoniella filamentosa var. algeriensis − − − − − − + + − − − − − − + − 1.2 

(Schotter) Cormaci and G. Furnari 

Crouania attenuata (C. Agardh) J. Agardh + + + + + + + + + + + + + + + + 31.2 

(Continued)





Cruoria cruoriaeformis (P.L. and H.M. + + + + − − + + − − − + − − − − 13.6 

Crouan) Denizot 

Cryptonemia lomation (A. Bertoloni) J. − − + − − + + + − − − + − − + + 2.8 

Agardh 

Dasya baillouviana (S.G. Gmelin) 

− − + − − + + − + + + − − + + + 5.6 

Montagne 

Dasya corymbifera J. Agardh − − − − + + − + − + + + − − + + 5.2 

Dasya ocellata (Grateloup) Harvey + + + − + + + + + − + + + − + + 14.8 

Dasya punicea (Zanardini) Meneghini ex − − − − − − − − − − + − − − − − 0.4 

Zanardini 

Dasya rigidula (Kützing) Ardissone + + + + + + + + + + + + + + + + 26.4 

Dasyella gracilis Falkenberg − − + − − − − − − − − − − − − − 0.4 

Dipterosiphonia rigens (Schousboe) + − + − + − − + − − + + − − + + 5.6 

Falkenberg 

Dudresnaya verticillata (Withering) Le − − − − − + − − − + − − − − − + 1.2 

Jolis 

Erythroglossum sandrianum (Kützing) + − + + + − + + − + − + + − − − 11.6 

Kylin 

Eupogodon planus (C. Agardh) Kützing + + + + + + + + + + + + + + + + 54.4 

“Falkenbergia rufolanosa (Harvey) F. + + + − + + − − − − + + + − + + 16.4 

Schmitz” stadium sporophyte of 

Asparagopsis armata Harvey 

Feldmannophycus rayssiae (Feldmann and + + + + + + + + + + − + + + − − 26.4 

Feldmann-Mazoyer) Augier and 

Boudouresque 

Gelidium bipectinatum G. Furnari − − − − − + + + + + + + + + + + 8.0



Gelidium spinosum (S.G. Gmelin) 

− − − − − − + − − − − − − − − − 0.8 

P.C. Silva 

Gloiocladia furcata (C. Agardh) J. Agardh − − − − − − − + − − − + − − − + 2.4 

Gloiocladia repens (C. Agardh) Sánchez − − − − − + − − − + + + − − − − 2.0 

and Rodriguez-Prieto 

Gracilaria bursa-pastoris (S.G. Gmelin) − − − − − − − − − − − − − − + + 1.2 

P.C. Silva 

Gracilaria corallicola Zanardini − − − − − − − + − − − + − − + − 3.2 

Gracilaria dura (C. Agardh) J. Agardh − − − − − − − − − − − − − − − + 0.8 

Griffithsia schousboei Montagne − − + + − − − + − − − + − + + − 6.8 

Gulsonia nodulosa (Ercegovic) Feldmann − − − − − − + − − + + − − + + − 5.2 

and Feldmann-Mazoyer 

Gymnothamnion elegans (Schousboe ex − − − − − − − − + − − − − − − − 0.8 

C. Agardh) J. Agardh 

Halopithys incurva (Hudson) Batters − − + − − − + + − − − − + − − − 3.2 

Halydyction mirabile Zanardini + + + + + + − + − + + − − + + + 10.8 

Halymenia floresia (Clemente and Rubio) − − − − − − − − − + − − + − + − 1.2 

C. Agardh 

Haraldia lenormandii (Derbès and Solier) + − − − + − + + + − + − + − + − 8.4 

J. Feldmann 

Herposiphonia secunda (C. Agardh) + + + + + + − + − − − + + + − + 13.6 

Ambronn 

Heterosiphonia crispella (C. Agardh) − − + + − + + + − − − + + − − − 10.4 

M.J. Wynne 

Hydrolithon farinosum (J.V. Lamouroux) − − − − + + − + + − + − − − + + 6.4 

D. Penrose and Y.M. Chamberlain 

Hypoglossum hypoglossoides (Stackhouse) + + + + + + + + + + + + + + + + 27.6 

Collins and Harvey 

Irvinea boergesenii (Feldmann) 

+ + + + + + + + + + + + + + + + 14.8 

R.J. Wilkes, L.M. McIvor and Guiry 

Jania adhaerens J.V. Lamouroux + + + + + + + + + + + + + + + + 55.2 

(Continued)





Kallymenia feldmannii Codomier − − − − − − − + − − − − − − + + 3.2 

Kallymenia lacerata J. Feldmann − − − − − − − − − − − − − − + − 0.8 

Kallymenia patens (J. Agardh) Parkinson − − − − − − − − − − − − − − + − 0.8 

Kallymenia reniformis (Turner) J. Agardh − − − − − − − + + − − − − − − − 0.4 

Kallymenia requienii J. Agardh − − − − − − − − − − − − − − + − 0.8 

Laurencia chondrioides Børgesen + + + − − + + + + − − + + + − − 7.6 

Lejolisia mediterranea Bornand − + + + − + + + + − + − − + + + 5.2 

Lophocladia lallemandii (Montagne) − − − − − − − + − − − − − − − − 0.8 

F. Schmitz 

Lomentaria chylocladiella Funk − + − − + + + + + + + − + − + − 10.8 

Lomentaria clavaeformis Ercegovic − − − − + − − + − − + − − − + + 4.8 

Lomentaria ercegovicii Verlaque, 

− − − − − + − + − − − − − − − − 0.8 

Boudouresque, Meinesz, Giraud and 

Marcot Coqueugniot 

Lomentaria verticillata Funk − − − − − − − + − − − − − − − − 0.4 

Meredithia microphylla (J. Agardh) + + + + + + + + + + + + + + + + 46.4 

J. Agardh 

Microcladia glandulosa (Turner) Greville − − − − + − − − − − − − − − − − 0.4 

Monosporus pedicellatus (J.E. Smith) Solier + + + + + + + + + + + − + + + + 17.2 

Myriogramme distromatica Rodriguez ex − − − − − − + − + − + + − − − − 2.8 

Boudouresque 

Nemastoma dichotomum J. Agardh − − − − − − − − − + − − − − + − 0.8 

Neurocaulon foliosum (Meneghini) 

− + + + − + + + − + + + − − + + 7.6 

Zanardini 

Nitophyllum micropunctatum Funk + + + + + + + + + + + + + + + + 29.6 

Osmundea pelagosae (Schiffner) F.W. Nam + + + + + + + + + + + + + + + + 65.6 

Osmundaria volubilis (Linnaeus) 

− − + − − − + + − + − + − − + + 4.8 

R.E. Norris



Parviphycus antipai (Celan) B. Santelices − − − − − − − + − − − − − − − − 0.4 

Peyssonnelia armorica (P.L. and 

− − − − + − − − − − − − − − − − 0.4 

H.M. Crouan) Weber van Bosse 

Peyssonnelia bornetii Boudouresque and + + + + + + + + + + + + + + + + 31.6 

Denizot 

Peyssonnelia crispata Boudouresque and − − − − − − + + − − − − − − − − 0.8 

Denizot 

Peyssonnelia dubyi P.L. and H.M. Crouan − − − − − − − − + − + − − − − − 0.8 

Peyssonnelia harveyana P.L. and 

− − − − − − − − − − − + − − − − 0.4 

H.M. Crouan ex J. Agardh 

Peyssonnelia inamoena Pilger − − − − − − − − − − + − − − − − 0.4 

Peyssonnelia orientalis (Weber van Bosse) − − − − − + + − − − − − − − − + 1.2 

Boudouresque and Denizot 

Peyssonnelia polymorpha (Zanardini) − + + + − − + + + − − + − − − − 4.8 

F. Schmitz 

Peyssonnelia rosa-marina Boudouresque − − − − − + − − − − − − − − − + 0.8 

and Denizot 

Peyssonnelia rubra (Greville) J. Agardh + + + + + + + + + + + + + + + + 53.6 

Peyssonnelia squamaria (S.G. Gmelin) + + + + + + + + + + + + + + + + 52.4 

Decaisne 

Peyssonnelia stoechas Boudouresque and − − + + + − + − + + + − − − − − 9.2 

Denizot 

Phyllophora crispa (Hudson) P.S. Dixon − − − − − − + + − + − + − + + + 5.2 

Platoma cyclocolpa (Montagne) F. Schmitz − − − − − − − − − − − − − − + − 0.8 

Plenosporium borreri (J.E. Smith) Nägeli − − − − − − + − − − − − − − − − 0.4 

Plocamium cartilagineum (Linnaeus) + + + + + + + + + + + + + + + + 35.6 

P.S. Dixon 

Polysiphonia elongata (Hudson) Sprengel − − − + + + + + + + + + − − + + 9.2 

Polysiphonia furcellata (C. Agardh) Harvey + + + + + + + + + + + + + + + − 51.6 

Polysiphonia perforans Cormaci, 

+ + + + − + + + + + + + + + + − 29.2 

G. Furnari, Pizzuto and Serio 

(Continued)





Polysiphonia subulifera (C. Agardh) − − + + + + + + + + + + − − + + 15.6 

Harvey 

Predaea ollivieri J. Feldmann − − − − − − − − − − − − − − + − 0.8 

Pterothamnion crispum (Ducluzeau) Nägeli − − − − − − − + − − − − − − − − 0.8 

Pterothamnion plumula (J. Ellis) Nägeli + − + − − − − + − + + + − − + + 8.4 

Ptilocladiopsis horrida Berthold − − − − − − − − − + + − − + + 2.0 

Ptilothamnion pluma (Dillwyn) Thurand + + + + + + + + + + + + + + + + 33.6 

Radicilingua reptans (Kylin) Papenfuss − − − − + + − + − + + + − − + + 7.6 

Rhodophyllis divaricata (Stackhouse) − − + + + + + − + + + + + − + + 20.4 

Papenfuss 

Rhodymenia ardissonei J. Feldmann − − + + − + + + + + + + − − + + 8.4 

Rhodymenia delicatula P.J.L. Dangeard − − − − − − − − − − + − − − − − 0.4 

Rhodymenia holmesii Ardissone − − − − − − − − − − + − − − − − 0.4 

Rodriguezella pinnata Ercegovic − − + + + − + + + − + − − − + + 3.6 

Rodriguezella strafforelloi F. Schmitz + + + + + + + + + + + + + + + − 27.6 

Rytiphlaea tinctoria (Clemente) C. Agardh − − − − + − + + − + − − − − − + 3.2 

Schmitzia neapolitana (Berthold) P.C. Silva − − − − − − − + − − − − − − − − 0.8 

Scinaia furcellata (Turner) J. Agardh − − − − − − − − − − − − − − − + 0.4 

Sebdenia dicotoma Berthold − − − − − − − − − − − − − − + + 1.6 

Sebdenia monardiana (Montagne) Berthold + − − − − − − − − − + − − − + − 1.2 

Sebdenia rodrigueziana (J. Feldmann) − − − − − − − − − − − − − − + − 0.8 

Athanasiasis


Seirospora apiculata (Meneghini) 

− − − − − − − − − − − − − − + − 0.4 

J. Feldmann 

Seirospora interrupta (J.E. Smith) 

− − − − + − + − − − + − − − − + 1.6 

F. Schmitz 

Seirospora sphaerospora Feldmann − − − − + + − + + − − − − − − + 2.0 

Spermothamnion flabellatum Bornet − − − − − − − + − − + − − − − + 2.0 

Spermothamnion johannis Feldmann- − − − − − − − + + − + − − − − − 1.2 

Mazoyer 

Spermothamnion repens (Dillwyn) 

− − − − + − − + + − + − − − − − 4.4 

Rosenvinge 

Sphaerococcus coronopifolius Stackhouse − − + − − − − − − − − − + − − + 1.2 

Sphondylothamnion multifidum (Hudson) − + − − − − − − − − − − − − − − 2.0 

Nägeli 

Spyridia filamentosa (Wulfen) Harvey − − − − − − − + + + + + − − − − 3.2 

Stilonema alsidii (Zanardini) K.M. Drew − + − − + − − + − + − + − + − + 13.6 

Tricleocarpa fragilis (linnaeus) Huisman − − − + − + + − − − − − + − − − 9.2 

and R.A. Towsend 

Womersleyella setacea (Hollenberg) + + + + + + + + + + + + + + + + 79.6 

R.E. Norris 

Wrangelia penicillata (C. Agardh) 

+ + + − + − − + + − + + + + + + 23.2 

C. Agardh 


Notes: L, Livorno; A, Argentario; V, Vada; Me, Meloria; Go, Gorgona; C, Capraia; E, Elba; P, Pianosa; Mo, Montecristo; F, Formiche di Grosseto; 

Gn, Giannutri; Gi, Giglio; H, horizontal substrate; F%, frequency, percentage of the presence of the taxon in the total samples (n = 250).



diversity (each island, bank or trait of continental coast) and regional diversity (the 

whole study zone). 

Species composition between assemblages developing on vertical and horizontal 

substrates was compared for each location where both orientations were 

sampled. 

Permutational multivariate analysis of variance (PERMANOVA) was performed 

in order to discriminate spatial patterns in species composition within the study zone, 

as well as differences between assemblages developing on vertical and horizontal 

substrata (Anderson 2001). Bray-Curtis dissimilarity for this analysis was calculated 

using presence/absence data. The analysis consisted of two crossed factors: Inclination 

(two levels “vertical vs. horizontal surface”) and Condition (two levels “island 

vs. coast”) crossed. Multivariate patterns were visualized using non-metric multidimensional 

scaling (nMDS) inferred from a Bray-Curtis dissimilarity matrix (Clarke 

and Warwick 1994). 

Results 

The encrusting layer of coralligenous habitat in the study areas mostly comprised the 

Corallinales Mesophyllum alternans (Foslie) Cabioch and Mendoza and secondarily 

Lithothamnion philippii Foslie, Mesophyllum macroblastum (Foslie) W.H. Adey, 

Lithophyllum pustulatum (J.V. Lamouroux) Foslie and Lithophyllum stictaeforme 

(Areshough) Hauck. 

In total, 187 macroalgal epiphytes of encrusting Corallinales were observed, 

among them 29 Ochrophyta, 14 Chlorophyta and 144 Rhodophyta (see Table 1 with 

nomenclature authority). 

A total of 45 and 18 species were present at a frequency greater than 20% and 

50%, respectively, while 98 species showed a frequency less than 5%. Among the most 

common species were the Ochrophyta Dictyota linearis, Halopteris filicina the 

Chlorophyta Flabellia petiolata, Halimeda tuna, Pseudochlorodesmis furcellata, the 

Rhodophyta Eupogodon planus, Jania adhaerens, Osmundea pelagosae, Peyssonnelia 

rubra, P. squamaria, and Womersleyella setacea. 

Several species were quite rare or scarcely reported in the study area, including 

Ceramium bisporum, C. graecum, Anthithamnion piliferum, Phyllariopsis brevipes, 

Polysiphonia perforans, Microcladia glandulosa Dictyopteris lucida and Spatoglosum 

solieri. 

The mean number of species per plot was 28.1 ± 5.1 (mean ± SD) on vertical 

substrates and 35.1 ± 3.7 on horizontal substrates. Values of species number 

were higher on horizontal substrates also at sample and site level, while values 

were similar between the two habitats at the larger spatial scales examined 

(Figure 2). 

Multivariate analysis applied to samples collected on vertical substrates showed 

that locations along continental coasts were distinct from islands (Figure 3). 

PERMANOVA on observational data detected significant differences for both 

the factors but not for the interaction between these (Table 2). Multivariate analysis 

applied to samples related to different substrate orientation showed that assemblages 

of vertical and horizontal substrata were distinct, even though this pattern was more 

evident on islands than along continental coasts (Figure 4).



Figure 2. Mean number of species of assemblages developing on vertical and horizontal 

substrate calculated for plot, sample, site, location and region (see the text for explanation of 

these terms). 

Discussion 

The extensive distribution of Mesophyllum alternans in the study area confirms previous 

observations that identified this species as the main algal foundation of coralligenous 

systems in the north-western Mediterranean Sea (Sartoretto 1996; Garrabou 

and Ballesteros 2000). 

Many species found in the present investigation are widely reported by other 

studies on the coralligenous habitat in other Mediterranean areas (Augier et al. 1971; 

Boudouresque 1973, 1984; Augier and Boudouresque 1975; Cinelli et al. 1979; Cormaci 

et al. 1997; Serio et al. 2006), supporting the assumption that in pristine conditions, 

coralligenous assemblages show similar species composition at the largest 

scales. A few species deserve further comments. 

Ceramium bisporum was reported for the first time in the Mediterranean Sea in 

the Tuscan Archipelago, as an epiphyte of Peyssonnelia rubra at a depth ranging 

between 15 and 25 m (Sartoni and Boddi 2002); in the present work, it was also found 

below 30 m depth at Capraia Island. 

Ceramium graecum was not reported for the flora of Tuscany; since its description 

from Greece (Lazaridou and Boudouresque 1992) this species has been reported 

only from Southern Italy (Gómez Garreta et al. 2001). 

Phyllariopsis brevipes is a cold temperate species that is reported in a few localities 

of the western Mediterranean Sea, particularly in deep subtidal habitats characterized 

by strong unidirectional currents (Cinelli 1981; Ribera et al. 1992). In the Tuscan 

Archipelago it was found along the southern coast of Gorgona Island at depths ranging 

between 35 and 50 m (Piazzi, Balata and Cinelli 2004). 

Polysiphonia perforans is rarely reported in the Mediterranean Sea but it was 

present in the majority of the locations studied; this finding suggests that this species 

is much more widespread in the basin than the available records indicate, as also 

asserted by Rindi et al. (2002).



Figure 3. Non-metric multidimensional scaling ordination based on Bray Curtis similarity calculated 

using presence/absence data. Notes: L, Livorno; A, Argentario; V, Vada; Me, Meloria; 

Go, Gorgona; C, Capraia; E, Elba; P, Pianosa; Mo, Montecristo; F, Formiche di Grosseto; 

Gn, Giannutri; Gi, Giglio. 

Dictyopteris lucida has been recently described and reported from Atlantic and 

Mediterranean coasts of Spain (Ribera Siguan et al. 2005). At Montecristo Island, 

the species shows invasive characteristics, colonizing wide areas in shallow subtidal 

and coralligenous habitat. 

Microcladia glandulosa is able to grow on rocks or on other algae or sponges and it is 

considered an endangered species of coralligenous assemblages (Ballesteros 2006). 

Spatoglosum solieri has been reported for several localities of Southern Italy in 

low subtidal habitats (Cossu et al. 1992; Cecere et al. 1996), and it is considered rare 

in the north-western Mediterranean area. 

Among the species that we have identified, Nemastoma dichotonum, Platoma 

cyclocolpa, Ptilocladiopsis horrida, Rodriguezella pinnata, Schmitzia neapolitana and



Table 2. Permutational multivariate analysis of variance examining differences 

between horizontal and vertical surfaces and between island and coastal conditions. 

The mean square (MS) of the residual was used as denominator for each 

source of variability. 

Source df MS Pseudo-F p 

In 1 1016.31 2.81 0.023* 

Co 1 1322.82 3.65 0.011* 

In × Co 1 344.36 0.95 0.455 

Residual 6 361.97 

Total 9 

Notes: In, inclination; Co, condition; *significant effects. 

the above mentioned M. glandulosa are considered endangered species (Ballesteros 

2006). All these species, except R. pinnata, are found only along the smallest and least 

urbanized islands of the study area (Giannutri, Pianosa and Montecristo) supporting 

the hypothesis that these are very sensitive to pollution and increased sedimentation 

rates. 

Five introduced species were found in coralligenous assemblages: the Rhodophyta 

Womersleyella setacea, Acrothamnion preissii and Lophocladia lallemandii and 

the Chlorophyta Caulerpa taxifolia and C. racemosa var. cylindracea. Caulerpa taxifolia 

and L. lallemandii were quite rare, being present only at Elba Island and Montecristo 

Island respectively. The spread of C. racemosa var. cylindracea in coralligenous 

habitat has already been reported and the effects of its colonization on benthic 

assemblages have been described (Piazzi et al. 2007). Although widespread in the 

study area (Piazzi et al. 2005), the alga was found in coralligenous habitat only in 

three localities (Elba island, Capraia Island and Livorno). On the contrary, W. setacea 

and A. preissii were present at all the studied locations. The spread of W. setacea 

and C. racemosa var. cylindracea may affect biodiversity and the natural structure of 

coralligenous assemblages (Piazzi et al. 2007), thus the presence of these two species 

in many localities of the study area represents an important ecological aspect to be 

monitored, especially in the protected islands of the Tuscan Archipelago National 

Park. 

In the studied area, coralligenous assemblages showed a similar structure at large 

scales, as 41 species were common to all the studied locations. Among the most common 

species we can find species considered characteristic of this habitat (Sphacelaria 

plumula, Palmophyllum crassum, Acrodiscus vidovichii, Ceramium bertholdii, Eupogodon 

planus, Osmundea pelagosae, Polysiphonia elongata, P. subulifera, Rodriguezella 

strafforelloi) and generalist species also reported for other Mediterranean habitats 

(Cutleria chilosa, Dictyota linearis, Halopteris filicina, Sphacelaria cirrosa, Flabellia 

petiolata, Halimeda tuna, Valonia macrophysa, Botryocladia botryoides, Ceramium 

codii, Jania adhaerens, Meredithia microphylla, Peyssonnelia rubra, P. squamaria) 

(Boudouresque 1984). The structure of macroalgal assemblages related to coralligenous 

habitat depends on light intensity: shallower assemblages are characterized by 

the dominance of Udoteaceae and deeper assemblages are characterized by the 

dominance of Peyssonneliaceae and laminar Rhodophyta (Ballesteros 2006). The



Figure 4. Non-metric multidimensional scaling ordination based on Bray Curtis similarity calculated 

using presence/absence data. Notes: H, horizontal substrate; L, Livorno; A, Argentario; 

V, Vada; Me, Meloria; Go, Gorgona; C, Capraia; E, Elba; P, Pianosa; Mo, Montecristo; 

F, Formiche di Grosseto; Gn, Giannutri; Gi, Giglio. 

generalist algae reported in the present study may be referred to the shallower assemblage. 

The depth at which we can observe the shift between the two assemblages is 

variable and related to the local characteristics of the water. This aspect was not 

examined in the present study, but it represents an interesting topic that deserves further 

investigations. In fact, the separation between coastal localities and islands 

detected by multivariate analysis may also be linked to different depth-related environmental 

gradients. Coastal locations are subjected to higher sedimentation rates in 

the studied areas (Airoldi, Rindi et al. 1995; Airoldi, Fabiano et al. 1996; Balata et al. 

2005) compared with the small islands of the Tuscan Archipelago that are far from 

sources of sediment flow. Under high levels of water turbidity and sedimentation,



two effects may be possible. First, the shift between shallow and deep macroalgal 

assemblages may occur at a shallower depth (Balata et al. 2006); second, species less 

sensitive to disturbance may be dominant, while species more sensitive may decrease 

(Balata, Piazzi and Benedetti-Cecchi 2007; Balata, Piazzi and Cinelli 2007). Both of 

these effects can determine the structure of assemblages, creating differences between 

assemblages developing at the same depth. 

Analyses detected differences in species composition between assemblages developing 

on horizontal and vertical substrates both on islands and along the continental 

coasts. These results showed that beta diversity related to substrate orientation is 

relevant in coralligenous habitat overall larger spatial scales, highlighting the importance 

of this aspect in ecological surveys. Results of the present study also showed that 

differences between assemblages of different substrate inclination are more evident 

on islands than along the continental coasts. In fact, assemblages of horizontal 

substrates on small islands were characterized by rheophilous species such as the 

Ochrophyta Cystoseira spinosa, C. zosteroides, Dictyopteris polypodioides, Sargassum 

hornschuchii and Zanardinia typus and the Rhodophyta Osmundaria volubilis and 

Phyllophora crispa. This kind of assemblage is considered characteristic of deepwater 

Mediterranean rocky reefs subjected to strong unidirectional currents (Ballesteros 

1990; Ballesteros et al. 1998), in agreement with environmental conditions of 

the small islands of the Tuscan Archipelago. Along the coasts, assemblages developing 

on horizontal and vertical substrates were still separated in the analysis, but were 

closer to each other than their island counterparts. Patterns of coralligenous assemblages 

in relation to the inclination of substrate have already been investigated along 

the coasts of Tuscany. Studies sampling sessile animals (Virgilio et al. 2006) also 

highlighted differences between assemblages that were not evident in the structure of 

macroalgal assemblages (Piazzi, Balata, Perusati et al. 2004). The absence of deepwater 

Fucales along the Tuscan coasts reduced the differences between horizontal 

and vertical assemblages. The lack of canopy species along continental coasts may be 

related to differences of hydrodynamism, but other causes may be possible. In fact, 

C. spinosa and other deep-water Fucales are considered sensitive to disturbance and 

their populations decrease in areas subjected to human pressure (Ballesteros et al. 

1998). It is plausible that under stress conditions the differences in composition 

among assemblages of different habitats (beta diversity) are eroded, even if they are 

still present; moreover, in terms of structure, few species present in both habitats 

become dominant (Balata, Piazzi and Benedetti-Cecchi 2007). 

The studied assemblages showed a high number of species, especially in relation 

to the smaller spatial scales examined (plot, sample), when compared with shallower 

assemblages in the same geographic area (24.5 ± 0.7 in Balata and Piazzi 2008). 

Coralligenous assemblages are characterized by high biodiversity mostly due to a 

higher abundance of sessile animals if compared to other Mediterranean habitats 

(Laubier 1966; Hong 1982). Results of this study confirm previous observations also 

showing high values of diversity for algal species (Balata and Piazzi 2008). This 

pattern is mostly evident at small spatial scales, while a high percentage of common 

species throughout the study area showed moderate levels of diversity at macroscales. 

A high number of species per sampling unit in coralligenous habitat has been attributed 

both to the heterogeneity of the substrate constituted by bioconstruction, which 

creates a high number of microhabitats with different characteristics, and to low 

physical disturbance that enhance the development of biotic interactions (Cocito



2004; Piazzi, Balata, Perusati et al. 2004; Ballesteros 2006). In fact, in subtidal marine 

environments, topographical heterogeneity creates refuges from consumers, offers 

suitable habitat to species with varying physiological requirements, and effects 

patterns of recruitment (Walters and Wethey 1986; Archambault and Bourget 1996). 

The present study, examined many data sampled over a large area, allowing us to 

uncover several floristic features of coralligenous assemblages, which would be difficult 

to pinpoint through small-scale investigations. Results also raise new important questions 

to be addressed in future investigations on one of the key benthic assemblages of 

the Mediterranean Sea. 

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