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Grazing Evects of the Periwinkle Echinolittorina Peruviana at a Central Peruvian High Rocky Intertidal

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Grazing Evects of the Periwinkle Echinolittorina Peruviana at a Central Peruvian High Rocky Intertidal Helgol Mar Res (2008) 62 (Suppl 1):S73–S83 DOI 10.1007/s10152-007-0086-3 ORIGINAL ARTICLE Grazing eVects of the periwinkle Echinolittorina peruviana at a central Peruvian high rocky intertidal Fernando J. Hidalgo · Fausto N. Firstater · Eugenia Fanjul · M. Cielo Bazterrica · Betina J. Lomovasky · Juan Tarazona · Oscar O. Iribarne Received: 11 July 2007 / Revised: 6 September 2007 / Accepted: 8 September 2007 / Published online: 13 November 2007 © Springer-Verlag and AWI 2007 Abstract Echinolittorina peruviana is the most common are similar to those reported elsewhere for high shore littor- gastropod in the high intertidal zone of Peru, representing inids and represent baseline data to understand how the role more than 80% of the individuals present at that zone. of intertidal consumers will vary under the cold (La Niña) Experimental removal of snails was used to evaluate their and warm (El Niño) phases of ENSO on these shores. eVects on (a) abundance of epilithic bioWlm, (b) barnacle recruitment, and (c) abundance of macroalgae under “nor- Keywords Echinolittorina peruviana · Grazing · Rocky mal” conditions of the El Niño Southern Oscillation intertidal · Peru · Field experiments (ENSO). Experiments were carried out from October 2005 to April 2007 at two intertidal levels of a semi-protected rocky shore of central Peru. Results demonstrated that Introduction E. peruviana is able to control bioWlm abundance and bar- nacle recruitment at both heights investigated, with marked Grazing by gastropods can inXuence the structure and com- eVects in the lower zone. Erect macroalgae (Ulva spp. and position of assemblages of plants and animals on rocky Gelidium spp.) were less aVected by grazing; but negative shores (e.g., Lubchenco and Gaines 1981; Hawkins and eVects were observed on macroalgal crusts. Season and Hartnoll 1983; Dunmore and Schiel 2003). Grazers can physical stress seem to play a more important role in the have both direct and indirect eVects (e.g., Lubchenco and abundance of macroalgae in the high intertidal. Our results Gaines 1981; Hawkins and Hartnoll 1983; Jara and Moreno 1984; Holmes et al. 2005). Herbivores can control the dis- Communicated by S. Thatje. tribution and abundance of ephemeral and perennial algae directly by consuming adult fronds or removing algal spor- Special Issue: Climate variability and El Niño Southern Oscillation: implications for natural coastal resources and managment. S. Thatje elings (e.g., Jara and Moreno 1984; Cervin and Åberg (ed.) 1997), providing bare space for sessile invertebrates to set- tle (Denley and Underwood 1979; Petraitis 1983). Grazing F. J. Hidalgo (&) · F. N. Firstater · E. Fanjul · M. C. Bazterrica · can also negatively aVect barnacles by crushing newly set- B. J. Lomovasky · O. O. Iribarne tled cyprids and bulldozing juveniles (Hawkins 1983; Laboratorio de Ecología, Departamento de Biología (FCEyN), Universidad Nacional de Mar del Plata, CC 573 Correo Central, Underwood et al. 1983). Moreover, they can indirectly B7600WAG Mar del Plata, Argentina aVect barnacles by grazing algae oV barnacle tests and the e-mail: [email protected] surrounding rock, thus preventing smothering and allowing settlement to occur on the bare rock (Hawkins 1983; B. J. Lomovasky · O. O. Iribarne Consejo Nacional de Investigaciones CientíWcas y Hawkins and Hartnoll 1983; Underwood et al. 1983). Técnicas (CONICET), Buenos Aires, Argentina In high shore habitats, littorinid gastropods are among the most conspicuous inhabitants. Most high shore littori- J. Tarazona nids feed on the epilithic bioWlm of microalgae, cyanobac- Facultad de Ciencias Biológicas, Universidad Nacional Mayor de San Marcos, teria, and bacteria; but also on macroalgae sporelings and Grupo DePSEA, Apartado 1898, Lima 100, Peru adult ephemeral Wlamentous or foliose macroalgae (Norton 123 S74 Helgol Mar Res (2008) 62 (Suppl 1):S73–S83 et al. 1990; McQuaid 1996); thus inXuencing both primary be explored given the general scarcity of baseline data on production and overall community structure (e.g., Castenholz ecological processes in the region. 1961; Underwood 1980; Bertness et al. 1983; Petraitis In this paper we examine the eVects of E. peruviana under 1983; Mak and Williams 1999). On the high shore, grazing “normal” conditions on the high intertidal zone of a rocky eVects are usually swamped by abiotic factors due to lower shore of central Peru. The speciWc aims of this study were to immersion times and increased physical stress (Castenholz (1) assess the eVects of E. peruviana on the abundance of the 1961; Underwood 1980; Cubit 1984), since the relative epilithic bioWlm of microalgae; (2) determine its eVects on importance of biological factors (e.g., predation, competi- the recruitment of the barnacles J. cirratus and N. scabrosus tion) in shaping intertidal communities tends to decrease and (3) evaluate its eVects on the vertical distribution and with the increasing environmental stress (e.g., prolonged air abundance of macroalgae in the upper mid intertidal level. exposure, excessive heat) (Menge and Sutherland 1987). Thus, organism abundance is thought to be determined by biological factors at lower intertidal levels, but physical Methods factors at upper intertidal levels. On Peruvian rocky shores, the high intertidal zone sup- Study site ports high densities of the Littorinid Echinolittorina peruvi- ana. This species ranges from Peru to central Chile The study was conducted from October 2005 to April 2007 (Williams and Reid 2004), reaching up to 2,000 individuals on a moderately protected rocky shore south of Ancón Bay m¡2 in the mid and high intertidal zone, usually in aggre- (11°46ЈS, 77°11ЈW; central Peru). Tides are semi-diurnal gated spatial distributions (Rojas et al. 2000). On moderately with a maximum range of ¡0.17 to 1.21 m respect to Chart protected rocky shores of central Peru, the vertical distribu- Datum (Dirección de Hidrografía y Navegación, Peru 2006). tion of E. peruviana ranges from bare substrate in the high The wave-protected, eastern side of a small island (»1ha, zone (i.e., the supralittoral fringe, sensu Paredes 1974), to a 70 m from the coast) was selected for our experiments. The belt of the chthalamoid barnacles Jehlius cirratus and Not- location was sheltered from severe wave action, but waves in ochthamalus scabrosus lower on the shore (i.e., the upper this site can reach up to 2 m high on stormy days. Samplings midlittoral zone, sensu Paredes 1974). During low tide and and experiments were conducted at two diVerent heights: on on calm days, E. peruviana forages further down the shore the upper limit of the barnacle belt (hereinafter the upper below the barnacle belt to the zone of the mussel Perumyti- high intertidal) and within the barnacle belt (hereinafter the lus purpuratus (F. Hidalgo, personal observation). Other lower high intertidal). The vertical distance between the two accompanying gastropods present in the high intertidal zone zones was on average 0.35 m. The slope at the site was are Austrolittorina araucana and the small limpets Siphona- 40°-60° and the lower and upper limits of the barnacle zone ria lessoni, Scurria variabilis, and Scurria ceciliana (Pare- were »1.07 and 1.69 m above Chart Datum, respectively. des 1974). A mix of many macroalgae species such as Gigartina glomerata, Gelidium spp., Ceramiun spp, Polysi- Gastropods abundance and size phonia spp., Ahnfeltia durvillaei, Chondrachanthus chamis- soi and the green ephemerals Ulva and Enteromorpha spp. Herbivore gastropod densities were quantiWed in 10–20 develops immediately below the barnacle belt (Paredes quadrats (10 £ 10 cm) randomly located at the upper and 1974). Erect mats of the green corticated Ahnfeltia durvillaei lower high intertidal. Sampling was replicated in October occur in the barnacle zone throughout the year. The ecologi- 2005; March, April, May and November 2006; and April cal eVects of E. peruviana in the Peruvian rocky intertidals 2007. To evaluate size diVerences of E. peruviana, the most have not been studied. The region is strongly aVected by the abundant gastropod, 100–200 animals were randomly col- interannual variability of El Niño Southern Oscillation lected in October 2005, April and November 2006, and (ENSO) (e.g., Tarazona and Castillo 1999). During El Niño, April 2007 from both heights, and the shell length (from the the most important eVects on coastal communities are apex to the base) was measured with a caliper (0.01 mm). unusually warm waters, a rise in sea level, greater wave The other gastropod species present in the sampling quad- action, and the depletion of nutrients that reduces primary rats were also measured. Two-way ANOVA was used to production and ultimately aVects all trophic levels (Tarazona compare E. peruviana abundances among sampling dates and Castillo 1999; Vinueza et al. 2006). In this nutrient-rich and heights (Zar 1999). Here and thereafter, when interac- environment (see Tarazona and Arntz 2001), it is predicted tions between two or more factors were signiWcant, multi- that top-down control of community structure during “nor- ple comparisons Tukey test was used to identify how mal” conditions will shift to be controlled by bottom-up pro- factors interacted (Zar 1999). Size diVerences of E. peruvi- cesses due to nutrient scarcity under El Niño events (see ana among heights were explored with separate Kolmogo- Vinueza et al. 2006). However, such predictions need still to rov-Smirnov two-sample test (Zar 1999), since parametric 123 Helgol Mar Res (2008) 62 (Suppl 1):S73–S83 S75 assumptions could not be met. A sequential Bonferroni cor- snails were allowed to enter (controls and experimental con- rection of the signiWcance level was applied (Rice 1989) in trols). Experimental plots (n = 8 per treatment) were estab- comparing sampling dates. Size diVerences of all of the lished at both the upper and lower high intertidal zones in gastropods present in the lower zone (where they were October 2005. Each plot was initially scraped and brushed more abundant, see Results) were pooled for all sampling to bare rock.
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