Effects of La Nina on recruitment and abundance of juveniles and adults of benthic community-structuring species in northern Chile

evaluated recruitment and abundance of juveniles and adults of the major benthic community-structuring groups (kelps, sea urchins, sea stars, crabs and rockfish) during a period affected by La Nina followed by a period of neutral conditions. Recruitment was several times higher during neutral conditions for sea urchins and sea stars. Lower recruitment during La Nina was probably caused by offshore larval transport caused by increased upwelling. Recruitment of crabs tended to be higher during neutral conditions, but this varied among sites. In contrast to invertebrates, kelp abundance seemed to be controlled by post-settlement processes, as reflected by fewer kelp juveniles after La Nina when the abundance of herbivore fishes strongly increased. ENSO has abnormal warm (El Nino) and cold (La Nina) periods (Fiedler 2002), which strongly impact oceanographic characteristics that affect marine and terrestrial communities (Holmgren et al. 2001; Fiedler 2002). Studying the effects of La Nina on marine environments is gaining importance because this event generates periods of increased primary productivity with a bottom-up effect on the system (Glantz 2002; Thatje et al. 2008). However, few studies have considered the effects of La Nina on marine communities (Arntz et al. 2006; Moreno et al. 2008). An increase in T. niger abundance during La Nina may reflect increased recruitment during the previous El Nino as recruitment in sea urchins is higher during periods of reduced upwelling (Morgan et al. 2000; Menge et al. 2004). During El Nino may also explain the population increase during the subsequent La Nina period. La Nina would reduce recruitment and this should translate into reduced juvenile and adult abundances after a year or more. Moreover, one-way ANOVAs were used to compare mean seawater temperatures between La Nina and neutral periods, both at the surface and the bottom. Negative thermal anomalies, such as those that dominated our system during the 200708 La Nina, represent upwelling conditions, with colder waters on average compared with neutral conditions. At Antofagasta and Coquimbo during La Nina, particularly between September and January of 200708, mean temperatures at the bottom and surface were 18C lower than those a year later. Juvenile kelp abundance was significantly higher during La Nina than under neutral conditions at both Antofagasta and Caldera Sea stars also showed a marked seasonal recruitment pattern, peaking in spring summer. Again, recruitment tended to be greater at the surface than on the bottom and at Caldera and Coquimbo, was higher during neutral conditions than during La Nina. Fish abundance was standardised to density per m2 of bottom surveyed and fish were classified as carnivores (Pinguipes chilensis, Cheilodactylus variegatus), herbivores (Aplodactylus punctatus) and omnivores (Girella laevifrons, Scartichthys viridis). The three groups were more abundant during neutral conditions than during La Nina, except for the herbivores at Coquimbo that had similar abundances across the sampling dates. Our study shows that during La Nina, recruitment of major shallow benthic community-structuring groups in centralnorthern Chile was considerably lower than

under neutral conditions. This was especially evident for the black sea urchin,Tetrapygus niger, and the four sea stars present in the shallow subtidal zone, Heliaster helianthus, Meyenaster gelatinosus, Luidia magellanica and Stichaster striatus. Neutral conditions favoured recruitment of the kelp Lessonia trabeculata, the black sea urchin, sea stars and the rock crab, Paraxanthus barbiger. This variation in recruitment was not merely due to interannual variability (see Palma et al. 2006). Larval behaviour and swimming characteristics seem to strongly influence the effect of ENSO on recruitment. During La Nina, upwelling intensifies and nutrient availability typically increases, with positive impacts on upper trophic levels (Glantz 2002). Augmented productivity could lead to strong settlement of crabs, as shown for the Dungeness crab, Cancer magister (Botsford 2001). In contrast, our study has shown low crab recruitment during La Nina, compared with neutral conditions. Similarly, the mussel (Mytilus spp.) from the intertidal of the west coast of the USA exhibits huge increases in recruitment associated with phytoplankton increases during neutral ENSO conditions (Menge et al. 2009). Our observed high recruitments of T. niger and sea stars during neutral conditions compared with La Nina coincides with the high recruitment of the sea urchin (Strongylocentrotus franciscanus), the sea star (Pisaster ochraceus) and barnacles during reduced upwelling periods occurring during warmer El Nino periods on the west coast of USA Despite the cold conditions that dominated our study, when the mean temperatures between the springsummer under La Nina and neutral conditions were compared, mean minimum and maximum temperatures during the latter were 18C higher, which likely explains the greater valueswe observed under neutral conditions.

perekrutan dievaluasi dan kelimpahan remaja dan orang dewasa dari kelompok masyarakat-strukturisasi besar bentik (kelps, landak laut, bintang laut, kepiting dan rockfish) selama periode dipengaruhi oleh La Nin ~ a diikuti dengan periode kondisi netral. Rekrutmen adalah beberapa kali lebih tinggi selama kondisi netral untuk landak laut dan bintang laut. Perekrutan lebih rendah selama La Nin ~ a mungkin disebabkan oleh transportasi larva lepas pantai yang disebabkan oleh peningkatan upwelling. Perekrutan kepiting cenderung lebih tinggi pada kondisi netral, tapi ini bervariasi antara situs. Berbeda dengan invertebrata, kelimpahan kelp tampaknya dikendalikan oleh proses pasca-penyelesaian, seperti tercermin dari sedikit remaja rumput laut setelah La Nin ~ ketika kelimpahan ikan herbivora sangat meningkat. ENSO memiliki normal hangat (El Nin ~ o) dan dingin (La Nin ~ a) periode (Fiedler 2002), yang sangat berdampak karakteristik oseanografi yang mempengaruhi masyarakat kelautan dan darat (Holmgren et al 2001;. Fiedler 2002). Mempelajari efek dari La Nin ~ pada lingkungan laut semakin penting karena acara ini menghasilkan periode peningkatan produktivitas primer dengan efek bottom-up pada sistem (Glantz 2002;. Thatje et al 2008). Namun, beberapa studi telah menyoroti efek dari La Nin ~ pada masyarakat kelautan (Arntz et al 2006;. Moreno dkk 2008.). Peningkatan T. niger kelimpahan selama La Nin ~ mungkin mencerminkan peningkatan perekrutan selama sebelumnya El Nin ~ o sebagai perekrutan di landak laut lebih tinggi selama periode berkurang upwelling (Morgan et al, 2000;. Menge et al 2004.). Selama El Nin ~ o juga dapat menjelaskan peningkatan populasi selama berikutnya La Nin ~ periode. La Nin ~ akan mengurangi perekrutan dan ini harus diterjemahkan ke dalam mengurangi kelimpahan remaja dan dewasa setelah satu tahun atau lebih. Selain itu, ANOVAs satu arah digunakan untuk membandingkan suhu air laut rata-rata antara La Nin ~ dan netral periode, baik di permukaan dan bagian bawah. Anomali termal negatif, seperti yang mendominasi sistem kami selama 2007-08 La Nin ~ a, merupakan kondisi upwelling, dengan air dingin rata-rata dibandingkan dengan kondisi netral. Pada Antofagasta dan Coquimbo selama La Nin ~ a, terutama antara September dan Januari 2007-08, berarti suhu di

bagian bawah dan permukaan yang 18C lebih rendah dibandingkan setahun kemudian. Kelimpahan rumput laut Juvenile secara signifikan lebih tinggi selama La Nin ~ daripada dalam kondisi netral pada kedua Antofagasta dan bintang laut Caldera juga menunjukkan pola perekrutan musiman ditandai, memuncak pada musim semi-musim panas. Sekali lagi, perekrutan cenderung lebih besar di permukaan dibanding di bagian bawah dan di kaldera dan Coquimbo, lebih tinggi selama kondisi netral daripada selama La Nin ~ a. Ikan kelimpahan adalah standar kepadatan per m2 dari bawah disurvei dan ikan diklasifikasikan sebagai karnivora (Pinguipes chilensis, Cheilodactylus variegatus), herbivora (Aplodactylus punctatus) dan omnivora (Girella laevifrons, Scartichthys viridis). Tiga kelompok yang lebih berlimpah selama kondisi netral daripada selama La Nin ~ a, kecuali untuk herbivora di Coquimbo yang memiliki kelimpahan yang sama di tanggal sampling. Studi kami menunjukkan bahwa selama La Nin ~ a, perekrutan kelompok masyarakat-strukturisasi besar dangkal bentik di Chile tengah-utara adalah lebih rendah daripada kondisi netral. Hal ini khususnya terlihat untuk landak laut hitam, Tetrapygus niger, dan bintang laut empat hadir di zona subtidal dangkal, Heliaster helianthus, Meyenaster gelatinosus, Luidia Magellanica dan Stichaster striatus. Kondisi netral perekrutan disukai dari kelp Lessonia trabeculata, landak laut hitam, bintang laut dan kepiting batu, Paraxanthus barbiger. Variasi dalam perekrutan itu bukan hanya karena variabilitas (lihat Palma et al. 2006). Perilaku larva dan karakteristik renang tampaknya sangat mempengaruhi efek ENSO pada perekrutan. Selama La Nin ~ a, upwelling mengintensifkan dan ketersediaan hara biasanya meningkat, dengan dampak positif pada tingkat trofik atas (Glantz 2002). Augmented produktivitas dapat menyebabkan penyelesaian kuat kepiting, seperti yang ditunjukkan untuk kepiting Dungeness, Kanker magister (Botsford 2001). Sebaliknya, penelitian kami menunjukkan kepiting rekrutmen rendah selama La Nin ~ a, dibandingkan dengan kondisi netral. Demikian pula, kerang (Mytilus spp.) Dari intertidal di pantai barat Amerika Serikat menunjukkan

peningkatan besar dalam perekrutan terkait dengan fitoplankton meningkat selama kondisi ENSO netral (Menge et al. 2009). Kami diamati rekrutmen tinggi T. niger bintang laut selama kondisi netral dibandingkan dengan La Nin ~ a bertepatan dengan perekrutan tinggi, landak laut (Strongylocentrotus franciscanus), bintang laut (Pisaster ochraceus) dan teritip selama periode upwelling berkurang terjadi selama lebih hangat El Nin ~ o periode di pantai barat Amerika Serikat Meskipun kondisi dingin yang mendominasi penelitian kami, ketika suhu rata-rata antara musim semi-musim panas di bawah La Nin ~ a dan netral kondisi dibandingkan, rata-rata suhu minimum dan maksimum selama terakhir adalah 18C tinggi, yang kemungkinan menjelaskan valueswe lebih besar diamati di bawah kondisi netral.