MARTIN THIEL ET AL.<strong>of</strong> meiobenthic organisms, mostly nematodes, have been recorded on <strong>the</strong> shelf <strong>of</strong>f Concepción(Neira et al. 2001, Sellanes et al. 2003). OMZ macr<strong>of</strong>aunal assemblages thus have low diversity<strong>and</strong> are typically composed <strong>of</strong> organisms with morphological <strong>and</strong> metabolic adaptations <strong>and</strong> feedingstrategies suited to <strong>the</strong>se conditions. A good example is <strong>the</strong> locally abundant polychaete Paraprionospiopinnata, an interface feeder organism, which is also able to switch to suspension feeding(Gutiérrez et al. 2000), with a complete set <strong>of</strong> enzymes for anaerobic metabolism (González &Quiñones 2002), <strong>and</strong> also bearing a set <strong>of</strong> highly branched <strong>and</strong> complex gills. Paraprionospiopinnata comprises about half <strong>of</strong> <strong>the</strong> ~14,000 macr<strong>of</strong>auna individuals m −2 reported from <strong>of</strong>f Concepciónwithin <strong>the</strong> OMZ (Palma et al. 2005).Below <strong>the</strong> OMZ, <strong>the</strong> total abundance <strong>of</strong> macrobenthos decreases while biomass <strong>and</strong> diversityincrease in parallel with increases in oxygen <strong>and</strong> decreases in organic carbon (Levin 2003). In <strong>the</strong>bathyal area <strong>of</strong>f Chile, important crustacean assemblages, mainly <strong>of</strong> commercially valuable gala<strong>the</strong>iddecapods (e.g., Pleuroncodes monodon <strong>and</strong> Cervimunida johni), thrive at <strong>the</strong> slope within <strong>and</strong>below <strong>the</strong> lower boundary <strong>of</strong> <strong>the</strong> OMZ. Below <strong>the</strong> OMZ, groups that are favoured by increasedoxygen concentrations are decapod crustaceans, gastropods <strong>and</strong> at greater depths echinoderms(mainly ophiuroids, asteroids <strong>and</strong> irregular echinoids). The large tubiculous onuphid polychaeteHyalinoecia sp. (tubes 10–20 cm in length) is particularly abundant at mid-slope depths <strong>of</strong>fConcepción <strong>and</strong> Chiloé (J. Sellanes personal observations). Scleractinian corals <strong>and</strong> many species<strong>of</strong> gorgonarians are common on <strong>the</strong> slope <strong>of</strong>f Chiloé, below 500 m water depth.Finally, though <strong>the</strong> three-layer scheme for benthic zonation is <strong>the</strong> general rule, ano<strong>the</strong>r type <strong>of</strong>chemosyn<strong>the</strong>tic community, methane seep areas, has been reported from <strong>of</strong>f Concepción at700–1400 m water depth (Stuardo & Valdovinos 1988, Sellanes et al. 2004, Sellanes & Krylova 2005).In <strong>the</strong>se reducing <strong>system</strong>s, C is fixed locally by both free-living <strong>and</strong> symbiotic chemosyn<strong>the</strong>tic bacteria.Seepage areas thus provide a suitable environment for <strong>the</strong> development <strong>of</strong> singular communitiesconsisting <strong>of</strong> sulphide oxidising bacteria (e.g., Beggiatoa), highly endemic endosymbiont-bearingclams (e.g., Vesicomyidae, Lucinidae, Thyasiridae <strong>and</strong> Solemyidae) <strong>and</strong> tubeworms (e.g., Lamellibrachiasp.). In addition, non-chemosymbiotic megafauna (e.g., crustaceans, gastropods, cephalopods,fish) are massively attracted to <strong>the</strong>se deep-sea hot spots <strong>of</strong> biological activity. The attraction<strong>of</strong> <strong>the</strong>se areas is due to both <strong>the</strong> abundance <strong>of</strong> locally produced organic material <strong>and</strong> <strong>the</strong> presence<strong>of</strong> authigenic carbonate reefs (generated by microbial <strong>and</strong> chemical processes), which are avidlycolonised by a diverse benthic fauna (J. Sellanes unpublished data).Latitudinal <strong>and</strong> bathymetric patternsIn general, for <strong>the</strong> available data <strong>of</strong> macr<strong>of</strong>aunal species richness on <strong>the</strong> Chilean shelf, no clearlatitudinal pattern emerges. Published reports <strong>of</strong> species numbers range from 18 to 56 species atnor<strong>the</strong>rn Chile (~23°S, 15–65 m water depth) (Jaramillo et al. 1998), 42 to 85 species at Huasco(~28°S, 20–50 m water depth) (Lancellotti & Stotz 2004), 10 to 41 species at Concepción (~36°S,15–65 m water depth) (Carrasco et al. 1988), <strong>and</strong> 24 to 34 species fur<strong>the</strong>r south (~45°S)(D. Lancellotti & W. Stotz unpublished data). Highest abundances reported are for <strong>the</strong> shelf <strong>of</strong>fConcepción (36,290–38,590 ind. m −2 ; Carrasco et al. 1988) <strong>and</strong> biomass values <strong>of</strong> almost 900 gwet wt m −2 are reported for south <strong>of</strong> Chiloé (D. Lancellotti & W. Stotz unpublished data).A crude comparison among <strong>the</strong> few available deeper studies (>100 m water depth) suggestshigher macr<strong>of</strong>aunal st<strong>and</strong>ing stocks <strong>and</strong> abundances also at <strong>the</strong> shelf <strong>of</strong>f <strong>central</strong> <strong>and</strong> sou<strong>the</strong>rn areas,both for macro- (body size 0.3–2 cm) <strong>and</strong> megafauna (body size >2 cm), compared with nor<strong>the</strong>rnChile (Figure 11). Palma et al. (2005) reported macr<strong>of</strong>aunal biomass values for three transectscovering a depth range from about 100 m to 2000 m at Ant<strong>of</strong>agasta, Concepción <strong>and</strong> Chiloé (about22°S, 36°S <strong>and</strong> 42°S, respectively). Though biomass trends with depth, beyond <strong>the</strong> OMZ, are ingeneral unimodal at all transects, with higher values at intermediate depths <strong>and</strong> lowest within <strong>the</strong>232
THE HUMBOLDT CURRENT SYSTEM OF NORTHERN AND CENTRAL CHILEMegafaunal abundanceMegafaunal abundance (Log 10 ind 1000 m −2 ), Macr<strong>of</strong>aunal abundance (Log 10 ind m −2 )100000100001000100101100001000010001001011000010000100010010Ant<strong>of</strong>agasta (∼22°S)98Concepción (∼36°S)Chiloé (∼42°S)Macr<strong>of</strong>aunal abundanceMacr<strong>of</strong>aunal biomassMegafaunal species numberMacr<strong>of</strong>aunal species number142 300 520 690 1350 1800120 365 535 800 1290 205070605040302010070605040302010070605040302010Macr<strong>of</strong>aunal biomass (g wet weight m −2 ), Megafaunal <strong>and</strong> macr<strong>of</strong>aunal species number1160 300 480 900Depth (m)1250 19600Figure 11 Macro- <strong>and</strong> megafauna depth-related patterns for three transects (22°S, 36°S <strong>and</strong> 42°S) across <strong>the</strong>shelf <strong>and</strong> upper bathyal zone <strong>of</strong> <strong>the</strong> Chilean margin. Data for macr<strong>of</strong>aunal abundance, biomass <strong>and</strong> speciesnumber from Palma et al. (2005) <strong>and</strong> for megafaunal abundance <strong>and</strong> species number from E. Quirogaunpublished data).OMZ <strong>and</strong> beyond 1350 m, average biomasses are lower <strong>of</strong>f Ant<strong>of</strong>agasta. Maximum values for thistransect (6.9 g wet wt m −2 ) are reported at 518 m water depth, while values about an order <strong>of</strong>magnitude higher (60.7 g wet wt m −2 ) are reported <strong>of</strong>f Concepción at 784 m depth. For sou<strong>the</strong>rnChile (~42°S) intermediate values (39.2 g wet wt m −2 ) are reported for a station located at 1250 mdepth. This also indicates a deepening <strong>of</strong> macr<strong>of</strong>aunal biomass maxima with latitude (Figure 11).For <strong>the</strong> megafauna observed at <strong>the</strong> same three transects, though biomass values are not available,abundances in general exhibited a similar pattern to that previously explained for macr<strong>of</strong>aunal233