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the humboldt current system of northern and central chile - figema

the humboldt current system of northern and central chile - figema

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MARTIN THIEL ET AL.<strong>the</strong> winter, <strong>the</strong>se amphipods live higher up on <strong>the</strong> beach, mainly to avoid being washed out byhigher wave activity. During summer, <strong>the</strong> abundances <strong>of</strong> O. tuberculata decrease <strong>and</strong> <strong>the</strong>y are foundcloser to <strong>the</strong> flotsam (Sánchez et al. 1982, McLachlan & Jaramillo 1995), possibly to avoiddesiccation risk in <strong>the</strong> supralittoral zone or to get close to <strong>the</strong>ir food. The cirolanid isopods(Excirolana spp.) have similar abundances in summer <strong>and</strong> winter, but <strong>the</strong>y shift <strong>the</strong>ir position in<strong>the</strong> intertidal zone, moving higher during <strong>the</strong> winter months (Sánchez et al. 1982). On <strong>the</strong> o<strong>the</strong>rh<strong>and</strong>, Emerita analoga in <strong>the</strong> nor<strong>the</strong>rn (22°S) <strong>and</strong> Mesodesma donacium in nor<strong>the</strong>rn-<strong>central</strong> Chile(30°S) reach highest abundances in summer (Sánchez et al. 1982, Contreras et al. 2000). At present,<strong>the</strong> reasons for <strong>the</strong> apparent inverse population dynamics <strong>of</strong> species from <strong>the</strong> upper versus thosefrom <strong>the</strong> lower intertidal zone are speculative, <strong>and</strong> seasonally varying <strong>of</strong>fspring (larval) survival orfood supply could be invoked. One very interesting species from <strong>the</strong> supralittoral zone, Tylosspinulosus, which has a restricted distribution in nor<strong>the</strong>rn-<strong>central</strong> Chile (17°–30°S) (Schmalfuss &Vergara 2000), is little known aside from a few data on its population density (Sánchez et al. 1982,Jaramillo et al. 2003), <strong>and</strong> it appears a promising enterprise to examine its population dynamics.The vicinity to upwelling centres plays an important role in <strong>the</strong> succession <strong>and</strong> structure <strong>of</strong>hard-bottom communities (Broitman et al. 2001, Narváez et al. 2006), but <strong>the</strong>re is no clear indicationthat <strong>the</strong> macr<strong>of</strong>auna composition <strong>of</strong> s<strong>and</strong>y beaches is influenced by <strong>the</strong>ir proximity to upwellingareas (Jaramillo et al. 1998). Contreras et al. (2000) concluded that growth rates <strong>of</strong> Emerita analogafrom a beach near <strong>the</strong> upwelling centre <strong>of</strong> Mejillones (22°S) were within values reported for o<strong>the</strong>rareas, suggesting only limited or no direct effects <strong>of</strong> upwelling on s<strong>and</strong>y beach inhabitants. Communitydynamics <strong>of</strong> s<strong>and</strong>y beaches may be influenced by upwelling to a lesser degree than thosepertaining to hard-bottom communities. For example, higher nutrient availability near upwellingareas positively influences growth rates <strong>of</strong> seaweeds on hard bottoms (Camus & Andrade 1999,Wieters 2005) <strong>and</strong> <strong>the</strong>reby <strong>the</strong> community succession (Nielsen & Navarrete 2004), which clearlyis <strong>of</strong> no importance on exposed s<strong>and</strong>y beaches where algae are usually imported from neighbouring(or distant) hard-bottom habitats. Fur<strong>the</strong>rmore, <strong>the</strong> interplay between upwelling <strong>and</strong> subsequentrelaxation events strongly affects <strong>the</strong> recruitment <strong>of</strong> hard-bottom organisms with planktonic larvae(Narváez et al. 2006), but seems to be <strong>of</strong> minor importance on s<strong>and</strong>y beaches, where <strong>the</strong> mostcommon organisms feature direct development.The effect <strong>of</strong> ENSO has been intensively studied in hard-bottom environments, but its role in<strong>the</strong> dynamics <strong>of</strong> s<strong>and</strong>y beach communities is not well known (Arntz et al. 1987). EN events mayhave deleterious effects on <strong>the</strong> organisms from <strong>the</strong> lower intertidal zone <strong>of</strong> s<strong>and</strong>y beaches (e.g.,Mesodesma donacium; see also Artisanal benthic fisheries, p. 278ff.). On <strong>the</strong> o<strong>the</strong>r h<strong>and</strong>, it is knownthat EN provokes mass mortalities <strong>of</strong> seaweeds <strong>and</strong> animals, many <strong>of</strong> which eventually will str<strong>and</strong>on s<strong>and</strong>y beaches (Arntz 1986). This high supply <strong>of</strong> OM may represent an important food sourcefor scavenging animals <strong>of</strong> <strong>the</strong> supralittoral zone <strong>of</strong> s<strong>and</strong>y beaches (e.g., Orchestoidea tuberculata).In this way, it can be suggested that intertidal organisms are distinctly affected by EN: species from<strong>the</strong> lower shore (suspension feeders with planktonic larvae) may be negatively affected by EN,while those from <strong>the</strong> supralittoral zone (scavenging animals with direct development) might benefitfrom <strong>the</strong> higher food supply <strong>and</strong> more benign climate (lower desiccation risk) during EN.Subtidal s<strong>of</strong>t-bottom communitiesZonation patternsRecent studies suggest that <strong>the</strong> bathymetric distribution <strong>of</strong> subtidal benthic communities <strong>of</strong>f <strong>the</strong>Chilean margin seems to be controlled mainly by bottom water oxygen conditions <strong>and</strong> sedimentorganic loading. OMZs are significant mid-water features in <strong>the</strong> eastern Pacific Ocean (Wyrtki1973, Kamykowski & Zentara 1990) that strongly influence <strong>the</strong> distribution <strong>and</strong> diversity <strong>of</strong>230

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