Seasonal Grazing and Food Preference of Herbivores in a Posidonia Oceanica Meadow*
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SCI. MAR., 65 (4): 367-374 SCIENTIA MARINA 2001 Seasonal grazing and food preference of herbivores in a Posidonia oceanica meadow* ANDREA PEIRANO, ILARIA NICCOLAI, ROBERTA MAURO and C. NIKE BIANCHI ENEA-Marine Environment Research Centre, S.Teresa, C.P. 316, I-19100, La Spezia, Italy. E-mail: [email protected] SUMMARY: Seasonal grazing of the fish Sarpa salpa (L.), the urchin Paracentrotus lividus Lamarck and the isopods Idotea spp. was compared with the C/N ratio of adult and intermediate leaves and epiphytes of Posidonia oceanica (L.) Delile, collected at three different depths. Despite seasonal differences in grazing, herbivores showed preferences through- out the year for adult leaves with more epiphyte and higher N contents. The maximum grazing on adult and intermediate leaves was observed in September and in June for fish and in March for urchins, whereas it was irregular for isopods. Graz- ing by the three herbivores was not related to their preference for leaves or epiphytes, notwithstanding the seasonal differ- ences in their C and N contents. We concluded that herbivores show no preference for food type throughout the year and that seasonal consumption of P. oceanica is related mainly to herbivore behaviour. Key words: grazing, C/N ratio, seagrass, epiphytes. INTRODUCTION pointed out the difficulty of defining a general strat- egy for grazers, the latter depending both on the Leaf production in the Mediterranean seagrass nutritional value of leaves and epiphytes and on the Posidonia oceanica (L.) Delile begins in August and occurrence of herbivores. attains its maximum in March, followed by a period The quality as a food (palatability) of the sea- of slow growth (Ott, 1980). The leaves and their epi- grass and its epiphytes, which can be expressed in phytes sustain a large number of organisms that either terms of C/N ratios (Mazzella et al., 1992), may actively feed on them or eat their remains, which vary according to season and within sites and enter the food chain (Romero et al., 1992; Mateo and depths, in relation to both environmental factors and Romero, 1997). Among grazers, a major role is the taxa that colonise the leaves (Pirc and Wollen- played by the fish Sarpa salpa (Velimirov, 1984; weber, 1988; Alcoverro et al., 1995; Alcoverro et Francour, 1997), the urchin Paracentrotus lividus al., 1997a). Moreover, a large presence of herbi- (Ott and Maurer, 1977; Kirkman and Young, 1981), vores may exert a controlling influence on epiphyte and the isopods Idotea spp. (Cebrian et al., 1996). biomass and on shoot size and leaf length (Alcover- Reviewing plant-animal trophic relationships in ro et al., 1997b). Data collected along the Spanish the P. oceanica ecosystem, Mazzella et al. (1992) Mediterranean coast were used to evaluate the mag- nitude of herbivore consumption both on a local and *Received February 16, 2000. Accepted May 15, 2001. a large scale (Alcoverro et al., 1997b; Cebrian et al., SEASONAL HERBIVORY AND C-N CONTENTS OF P. OCEANICA AND EPIPHYTES 367 1996) and showed that the percentage of leaf pro- Twenty orthotropic shoots were collected sea- duction removed by herbivores in summer was cor- sonally on 28 September 1995, 11 December 1995, related with both leaf specific growth rates and leaf 21 March 1996 and 10 June 1996 at 5 m (the upper production per shoot (Cebrian and Duarte, 1998). limit of the meadow), 10 m and 17 m (close to the Differences in grazing might also be explained lower limit). At each station, density was estimated by different behaviour of the grazers. Massive by counting shoots in three random quadrats of 1 m2. schools of the fish S. salpa actively feed on P.ocean- In the laboratory leaves were measured and divided ica leaves in Summer to accumulate reserves for into adult and intermediate (Giraud, 1977). Herbi- reproduction. In winter, adults migrate to greater vore consumption was estimated by counting the depths, whereas young fish, in shallow waters, eat number of leaves with grazed tips. Leaf marks were preferentially small crustaceans (Bini, 1968; Tor- attributed to the fish Sarpa salpa, the urchin Para- tonese, 1975, Francour, 1997) and turf epilithic centrotus lividus or the isopods Idotea spp. follow- algae (Havelange et al., 1997). ing Boudouresque and Meisnez (1982). P. lividus forms true populations in P. oceanica Epiphytism on leaves was calculated seasonally meadows, where high densities may have a heavy on five shoots per station using the Index of Epi- impact on the meadow (Kirkman and Young; Ned- phytism, IE (Morri, 1991), which is the portion of elec and Verlaque, 1984). Even though the urchins the leaf surface covered by each category of epi- reproduce throughout the year (Tortonese, 1965; phyte with respect to the total leaf surface. The fol- Fenaux 1968), major spawning is concentrated in lowing epiphyte categories were taken into account: spring and summer (Fenaux 1968; Azzolina Algae (encrusting, erected, algal mat) Hydroids and Willsie, 1987) and is related to the lowest con- (Sertularia perpusilla, Plumularia obliqua, others), sumption of seagrass (Semroud and Kada, 1987). Bryozoa (Electra posidoniae, others), and other ani- While the young urchins feed on both rhizomes and mals. Epiphytes were carefully scraped from leaves leaves, the adults (>2 cm diameter) have a low-spe- with a razor and leaves and epiphytes were desiccat- cialised diet and graze preferentially, on the highly ed for 48 h at 80°C and then weighed (Mazzella and epiphytized distal part of the leaves at nigth (Ned- Ott, 1984). Total C and N contents of leaves and epi- elec and Verlaque, 1984; Zupo and Fresi, 1984). phytes were estimated on three homogenised repli- Little is known about the behaviour of the cates with a CHN-analyser mod. Carlo Erba 1106. isopods. The diet of the isopods I. baltica balsteri The C values were corrected following the proce- was investigated by Lorenti and Fresi (1982), who dures of Nieuwenhuize et al. (1994) to evaluate the observed prolonged survival of the animal feeding calcium carbonate that might still be present on the on living leaves of P. oceanica. blades after scraping. Until now, it is not clear whether seasonal graz- Data were analysed using ANOVA and homo- ing is related to leaf or epiphyte quality of food or geneity of group variances controlled with the depends on other factors. The aim of the present Bartlett test. To satisfy the assumption of work is to assess the relationships between grazing homoscedasticity, the number of grazed leaves was pressure and food availability and C and N contents, log 10 (X + 0.5) transformed. When data did not sat- both in time and space. For this purpose three sta- isfy the assumption of homogeneity of variance, the tions along a depth gradient were observed during Kruskall-Wallis non-parametric one-way analysis of one year. Two levels of food were considered: a) variance was applied. leaves (adult and intermediate) and b) epiphytes Correlations between grazing activity of the fish (divided into major taxa). Seasonal variations in S. salpa, the urchin P. lividus and the isopods Idotea food quality were analysed in terms of C and N con- spp. and C, N contents of leaves and epiphytes and tents and were compared with grazing pressure by the IE indexes were investigated . Sarpa salpa, Paracentrotus lividus and Idotea spp. RESULTS MATERIALS AND METHODS Shoot density showed differences between the The study was carried out in the Bay of Mon- three sampling depths (one-way ANOVA, P = terosso, in the “Cinque Terre” Marine Protected 0.031). Recorded values (293.1 shoots.m-2 at 5 m Area (La Spezia, Ligurian Sea). depth, 233.7 shoots.m-2 at 10 m and 159 shoots.m-2 at 368 A. PEIRANO et al. Adult Leaves Intermediate Leaves Carbon (% DW) Nitrogen (% DW) l(mm) Carbon (% DW) Nitrogen (% DW) 60 4 1200 60 4 1000 3 3 40 800 40 5m 2 600 2 20 400 20 1 1 200 0 0 0 0 0 60 4 1200 60 4 1000 3 3 40 800 40 10 m 2 600 2 20 400 20 1 1 200 0 0 0 0 0 60 4 1200 60 4 1000 3 3 40 800 40 600 17m 2 2 20 400 20 1 1 200 0 0 0 0 0 Sep 95 Dec 95 Mar 96 Jun 96 Sep 95 Dec 95 Mar 96 Jun 96 FIG. 1. – Seasonal variations of carbon (o) and nitrogen (∆) contents in adult and intermediate leaves. Mean leaf length (l) in mm (u). Standard deviations are indicated with bars. 17 m) are characteristic of the present status of the agreement with Romero et al. (1992). The recorded Ligurian Posidonia oceanica meadows, which have values (37.32-49.07%) were similar to those regressed in response to large coastal impacts obtained by Augier et al. (1982) in France (31-45%) (Peirano and Bianchi, 1997). and Enriquez et al. (1995) in Spain (38.12%). The Kruskall-Wallis analysis of variance performed maximum C content was recorded in December and on leaf length showed differences between months March in both adult and intermediate leaves. (P < 0.0001), with the longest leaves in June and Nitrogen showed seasonal differences between September (Fig. 1). Differences were also found intermediate and adult leaves and between depths between depths and between adult and intermediate (Table 1 and Fig. 1). The N content showed higher lev- leaves (P < 0.05) in all months, with the single els in intermediate leaves. The analyses showed a sea- exception of March. Adult leaves, always longer sonal trend at all depths, with minimum values in Sep- than intermediate ones, attained their maximum tember and maximum ones in December and March.