Spatial and Temporal Variation of Motile Macro-Invertebrate Assemblages Associated with Posidonia Oceanica Meadows
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ISSn: 0001-5113 ACtA AdrIAt., udC: 597.556.333.7: 593.1 (262.16) AAdrAY 52(2): 201 - 214, 2011 Spatial and temporal variation of motile macro-invertebrate assemblages associated with Posidonia oceanica meadows Roberto BEDINI1*, Marco PERTUSATI2, Francesca BATISTINI1 and Luigi PIAZZI1 1Istituto di Biologia ed Ecologia Marina, Piazza G. Bovio 4, 57025 Piombino, Italy 2Centro Interuniversitario di Biologia Marina ed Ecologia Applicata, v.le N. Sauro 4, Livorno, Italy *Corresponding author: e-mail: [email protected] The present study assesses the spatial and temporal distribution of motile invertebrates in Posidonia oceanica meadows located off the coast of Tuscany, in the northwestern Mediterranean Sea. A hierarchical sampling design was used to evaluate changes in composition and abundance of assemblages in relation to depth and to different periods over one year. A total of 136 taxa were determined, among them 34 Mollusca, 94 Arthropoda, 5 Annelida and 1 Echinodermata. Results of both multivariate and univariate analyses showed that the macro-invertebrate assemblage structure varied along a depth gradient but this pattern was not consistent in the two study periods. Results suggest that temporal and spatial differences in the structure of P. oceanica meadows are key driv- ers in determining the structure of associated macro-invertebrate assemblages, as values of richness and abundance decreased with a decrease in leaf length and shoot density. Key words: depth, macro-invertebratemotile assemblages, Mediterranean Sea, Posidonia oceanica, seasonal effects IntroduCtIon a refuge from predators, and allow different species to occupy various ecological niches in Seagrass beds represent one of the most a same habitat (STONER, 1980; ORTH et al., 1984). important coastal ecosystems in terms of extent Moreover, seagrass meadows provide food via and production. Moreover, they play a funda- the high primary production of the plants and mental ecological role as habitat-forming spe- epiphyte algae (VAN MONTFRANS et al., 1984; cies, thus enhancing biodiversity. Seagrass beds DUFFY et al., 2003) and the development of com- allow the development of complex epiphyte plex detritus chains (KLUMPP & VAN DER VALK, assemblages and host more diverse and abun- 1984; EDGAR, 1999). dant animal communities than unvegetated areas Motile macro-invertebrate assemblages (EDGAR, 1990; ARRIVILLAGA & BALTZ, 1999). associated with seagrasses may vary spatially The influence of seagrasses on motile fauna in relation to responses of organisms to environ- is both structural and trophic since the habitats mental gradients and, temporally, in relation to they form enhance habitat complexity, provide the life cycle of organisms and changes in the 202 ACTA ADRIATICA, 52(2): 201 - 214, 2011 structure of seagrass beds (HECK & ORTH, 1980). Effective conservation and management of sea- grass communities requires an understanding of the processes that control patterns of distribution and abundance of organisms. Essential to such an understanding is reliable information on the variation in abundance and diversity of species Figure 1. Map of the northern coast of Tuscany. The ellipse through space and time. indicates the study area. In the Mediterranean Sea, Posidonia oce- dates were chosen in each period and three areas anica (L.) Delile is the most important seagrass separatedFigure 1. by several kms were sampled on each for its vast extent, stability, high biodiversity and date and at each depth. At each site, ten replicate production (MAZZELLA et al., 1992). The motile measures of density were made using a 40x40 macro-invertebrate assemblages of the P. ocean- cm quadrant, and ten shoots were collected to ica leaf stratum have been widely studied (MAZ- assess phonological variables. A hand-towed net ZELLA et al., 1989; 1992; GAMBI et al., 1992; SCIPI- (0.4 mm mesh size) was used to sample motile ONE et al., 1996; BEDINI et al., 1997; BEDINI, 2003; organisms living in the leaf stratum. Three rep- BORG et al., 2010). However, most investigations licate samples were collected for each depth and have focused on particular taxa or have consid- date. Each sample consisted of a series of 60 ered only one aspect of the associated faunal strokes over the seagrass leaf canopy (RUSSO et distribution (SCIPIONE & FRESI, 1984; RUSSO et al., al., 1985). 1984; 1991; 2002; GAMBI et al., 1995; 1998; SCIPIONE, In the laboratory, taxa were identified and 1999; BEDINI & BEDINI, 2008; BORG & SCHEMBRI, the abundance expressed as number of indi- 2000; BEDINI, 2006). Moreover, no information is viduals per sample. Taxa nomenclature fol- available on this topic for the northern Mediter- lowed the WORLD REGISTER OF MARINE SPECIES ranean Sea. (2011). Differences in the species composition The aim of the present study is to describe and abundance were tested using Permutational the spatial and temporal distribution of motile Analysis of Variances - PERMANOVA (ANDER- macro-invertebrate assemblages of P. oceanica SON, 2001). A 4-way model was used with Depth in meadows distributed off the coast of Tuscany (shallow vs. intermediate vs. deep) and Period in the northwestern Mediterranean Sea. In this (2 levels; spring-summer and autumn-winter) context, a hierarchical sampling design was used as fixed and crossed factors, Date (2 levels) as to evaluate changes in composition and abun- random factor nested in Period, Area (3 levels) dance of assemblages in relation to depth and to as random factor nested in the interaction Depth different periods over one year. x Date. The Bray-Curtis index of dissimilar- ity was calculated from untransformed species- MAtErIAL And MEtHodS abundance data. The SIMPER routine was used to establish which taxa contributed most to the 19 The study was carried out off the coast of dissimilarity between samples. Tuscany (northwestern Mediterranean Sea) in Differences in meadow shoot density, mean 2007 (Fig. 1). At the study sites, Posidonia leaf length per shoot, number of taxa and oceanica meadows colonized a rocky bottom. total number of macro-invertebrates per sample Three depth ranges were considered: 8-10 m was tested using Univariate Analyses of Vari- (shallow), 15-17 m (intermediate) and 23-25 m ance - ANOVA (UNDERWOOD, 1997) using the (deep). Two periods were chosen over one year: same factors and levels considered in the PER- spring-summer (s-s) and autumn-winter (a-w); MANOVA test. Homogeneity of variance was these correspond to periods during which P. oce- tested using Cochran’s C test. The SNK test was anica meadows have different structural charac- used to identify the source of difference when teristics (MAZZELLA & OTT, 1984). Two random ANOVA indicated a significant difference. Bedini et al.: Spatial and temporal variation of motile macro-invertebrate assemblages ... 203 rESuLtS brate assemblages. Pair-wise tests showed that differences between depths were significant A total of 136 taxa were identified, among for the spring-summer period but not for the them 34 Mollusca, 94 Arthropoda, 5 Annelida autumn-winter period; moreover, differences and 1 Echinodermata (Table 1). between periods were significant in the shal- PERMANOVA indicated a significant inter- low and intermediate assemblages but not in action between Depth and Period for the species the deep assemblages (Table 2). Differences composition and abundance of macro-inverte- between areas were also significant. Table 1. List of recorded taxa. De = deep, In = intermediate, Sh = shallow, s-s = spring-summer, a-w = autumn-winter Sh Sh In In de de tAXA s-s a-w s-s a-w s-s a-w MOLLUSCA Alvania beani (Hanley in Thorpe, 1844) - - + - + - Alvania discors (Allan, 1818) + + + + + + Alvania lineata (Risso, 1826) + + - - + - Barleeia unifasciata (Montagu, 1804) + - + - + - Bittium reticulatum (Da Costa, 1778) + + + - + + Calliostoma conulus (Linnaeus, 1758) + - + - - - Calliostoma laugieri (Payraudeau,1826) + - - - - - Columbella rustica (Linnaeus, 1758) + - + - - - Cuthona caerulea (Montagu, 1804) - - - + - + Elysia viridis (Montagu, 1804) + + - - + - Eulimella acicula (Philippi, 1836) + - - - - - Gibbula divaricata (Linnaeus, 1758) + - - - - - Gibbula umbilicaris umbiliaris (Linnaeus, 1758) + - - - - - Gibbula varia (Linnaeus, 1758) + - + - + - Jujubinus exasperatus (Pennant, 1777) + + + + + + Monophorus perversus (Linnaeus, 1758) - + - + - - Pusillina marginata (Michaud, 1832) + - - + + + Pusillina philippi (Aradas & Maggiore, 1844) + + + + - + Pisania striata (Gmelin, 1791) - + - - - - Rissoa auriscalpium (Linnaeus, 1758) + + + + + + Rissoa guerinii (Récluz, 1843) + + - + + + Rissoa labiosa (Montagu, 1803) - - - + - - Rissoa membranacea (Adams J., 1800) - + - - - - Rissoa monodonta (Philippi, 1846) + + + + - + Rissoa variabilis (von Muehlfeldt, 1824) + + + + - + Rissoa ventricosa (Desmarest, 1814) + + + + + + Rissoa violacea (Desmarest, 1814) - + + + + + Rissoina bruguieri (Payraudeau, 1826) - - + - - - Smaragdia viridis (Linnaeus, 1758) - - - + - - Tectura virginea (O.F. Muller, 1776) - + - - - - Tricolia pullus pullus (Linnaeus, 1758) + + + + + - Tricolia punctura (Gofas, 1993) + - - + - - 204 ACTA ADRIATICA, 52(2): 201 - 214, 2011 Table 1. cont’ d Tricolia speciosa (von Muehlfeldt, 1824) + - - - - - Tricolia tenuis (Michaud, 1829) + - + - - - CRUSTACEA Acanthonyx lunulatus (Risso, 1816) - - + - - - Acartia clausi (Giesbrecht, 1889) - - - - + + Alpheus dentipes (Guérin-Méneville, 1832) + - - - - - Ampelisca diadema (Costa, 1853) + - - - - - Ampithoe ferox (Cheuvreux, 1902) + - - - - - Ampithoe helleri (Karaman,