MARTIN THIEL ET AL.whiplash effect <strong>of</strong> fronds <strong>and</strong> stipes pushes herbivores away from <strong>the</strong> plants, reducing grazingpressure. On <strong>the</strong> o<strong>the</strong>r h<strong>and</strong>, spawning <strong>of</strong> egg cases <strong>of</strong> elasmobranchs on L. trabeculata ties <strong>the</strong>stipes toge<strong>the</strong>r, <strong>the</strong>reby reducing <strong>the</strong> whiplash effect <strong>and</strong> thus permitting grazers to approach kelpplants. Additionally, this ‘tie effect’ modifies plant shape toward <strong>the</strong> tree-like morph, <strong>and</strong> plantsare more easily dislodged by water movement (Vásquez 1992).Longevity <strong>of</strong> kelps from nor<strong>the</strong>rn Chile in <strong>the</strong> field is not well known since <strong>the</strong>y do not showany evident age-related structure. Never<strong>the</strong>less, individual Lessonia plants can survive in <strong>the</strong> fieldfor as long as 5 yr (J.A. Vásquez personal observations), <strong>and</strong> Macrocystis integrifolia has beenreported as a perennial species in nor<strong>the</strong>rn Chile (Buschmann et al. 2004b). Several factors generatesignificant biomass loss in <strong>the</strong> field: grazing pressure, wave impact, <strong>and</strong> spore release, which takesplace mainly during summer (Santelices & Ojeda 1984, Edding et al. 1994).Lessonia <strong>and</strong> Macrocystis populations in nor<strong>the</strong>rn-<strong>central</strong> Chile grow throughout <strong>the</strong> year butexhibit growth peaks during spring–summer (Buschmann et al. 2004b, Tala et al. 2004). Growthpatterns are modified by wave impact, quantity <strong>and</strong> quality <strong>of</strong> light, water temperature <strong>and</strong> nutrientconcentration (Buschmann et al. 2004b, Vega 2005). Local factors such as intraspecific interactions(Santelices & Ojeda 1984), herbivory (Vásquez & Buschmann 1997, Vásquez et al. 2006) <strong>and</strong>coastal upwelling events (González et al. 1998, Vásquez et al. 1998) can modify seasonal patterns<strong>of</strong> abundance <strong>and</strong> distribution (see also Graham et al. 2007). Large-scale phenomena such as ENSOproduce interannual variability in abundance <strong>and</strong> could eventually generate local extinctions, asobserved after <strong>the</strong> EN events <strong>of</strong> 1982–1983 <strong>and</strong> 1997–1998 (Soto 1985, Tomicic 1985, Vega 2005,Vásquez et al. 2006). Major impacts <strong>of</strong> EN were observed in kelp beds from lower latitudes(18–21°S). For example, a kelp bed occupying an area <strong>of</strong> ~40 ha at 18°S during <strong>the</strong> 1970s (IFOP1977) disappeared as a consequence <strong>of</strong> EN 1982–1983 (Soto 1985) <strong>and</strong> has not recovered since.Similarly, during EN 1997–1998, <strong>the</strong> density <strong>of</strong> adult sporophytes on subtidal hard bottoms at 21°Sdecreased rapidly <strong>and</strong> linearly with increasing positive <strong>the</strong>rmal anomalies (Figure 14). Six monthslater <strong>the</strong> site remained completely devoid <strong>of</strong> adult sporophytes, <strong>and</strong> no recolonisation occurred in<strong>the</strong> subtidal zone during <strong>the</strong> study period. In areas south <strong>of</strong> 23°S positive <strong>the</strong>rmal anomaliesregistered during EN 1997–1998 had only limited effects on kelp beds (Figure 14). As a result, <strong>the</strong>spatiotemporal abundance patterns <strong>of</strong> M. integrifolia sporophytes in nor<strong>the</strong>rn-<strong>central</strong> Chile is highlyvariable (Figure 14).Kelp conservation <strong>and</strong> human activitiesMany <strong>of</strong> <strong>the</strong> diverse kelp-associated species have significant socioeconomic importance for humanpopulations along <strong>the</strong> coast in north-<strong>central</strong> Chile <strong>and</strong> have been subject to harvesting by localhuman communities since pre-Columbian times (Jerardino et al. 1992, Vásquez et al. 1996). Spatial<strong>and</strong> temporal dynamics <strong>of</strong> kelp beds are significantly affected by anthropogenic impacts producedby both intense harvesting <strong>and</strong> severe pollution with organic as well as mining waste (Faugeronet al. 2005; Vásquez & Vega 2005). Lessonia nigrescens, L. trabeculata <strong>and</strong> Macrocystis integrifoliaare commercially exploited between 18°S <strong>and</strong> 32°S. These species account for >95% <strong>of</strong> l<strong>and</strong>ings<strong>of</strong> macroalgae <strong>and</strong> basically are used for alginic acid extraction. Until 2002, collected biomass indry weight (dry wt) amounted to ~200,000 t, almost exclusively based on str<strong>and</strong>ed kelps, resultingfrom natural mortality <strong>of</strong> plants with holdfasts that are weakened by grazing <strong>and</strong> <strong>the</strong>n detached bystrong bottom <strong>current</strong>s <strong>and</strong> waves. Since 2003, however, due to international needs for raw (dry)materials <strong>and</strong> also due to increasing dem<strong>and</strong>s for fresh algae (to sustain aquaculture <strong>of</strong> herbivorousinvertebrates in nor<strong>the</strong>rn Chile), harvesting <strong>of</strong> natural kelp increased to ~300,000 t dry wt per year.This has led to <strong>the</strong> recent implementation <strong>of</strong> new administrative rules in order to mitigate <strong>the</strong> impacton natural kelp populations. Regulations aim at <strong>the</strong> establishment <strong>of</strong> a sustainable kelp fishery,applying <strong>the</strong> following strategies: (1) harvest management (Vásquez 1995, 1999, 2006), (2) stock242
THE HUMBOLDT CURRENT SYSTEM OF NORTHERN AND CENTRAL CHILESubtidal kelp 54Intertidal kelp3251San Marcos (21°S)0Abundance <strong>of</strong> adult sporophytes (ind m −2 )43210543210543210543210543210543210Abundance <strong>of</strong> adultsporophytes (ind m −2 )Camarones (18°S)1996 1997 1998 1999 20001−1−3−55Constitución (23°S)31−1−3−55Playa Blanca (28°S)31−1−3−55Los Choros (29°S)31−1−3−55San Lorenzo (30°S)31−1−3−55Los Vilos (32°S)31−1−3−51996 1997 1998 1999 2000Thermal anomalies (°C)531−1−3−5Thermal anomalies (°C)Figure 14 Temporal variation (between 1996 <strong>and</strong> 2000) <strong>of</strong> abundances <strong>of</strong> adult sporophytes <strong>of</strong> M. integrifolia(●) <strong>and</strong> <strong>the</strong>rmal anomalies estimated in situ (line) over a latitudinal gradient in nor<strong>the</strong>rn Chile. Note: At SanMarcos, an intertidal kelp population appeared after <strong>the</strong> El Niño event (▫), while <strong>the</strong> subtidal kelp bed didnot recover. At Camarones (top) no sporophytes were observed during <strong>the</strong> study period. (Modified from Vega2005)243