Vol. 454: 147–157, 2012 MARINE ECOLOGY PROGRESS SERIES Published May 21 doi: 10.3354/meps09729 Mar Ecol Prog Ser Contribution to the Theme Section ‘Seabirds and climate change’ OPEN ACCESS Effects of climate variability on breeding phenology and performance of tropical seabirds in the eastern Indian Ocean Christopher A. Surman1,*, Lisa W. Nicholson1, Jarrod A. Santora2 1Halfmoon Biosciences, Ocean Beach, Western Australia 6333, Australia 2Farallon Institute for Advanced Ecosystem Research, PO Box 750756, Petaluma, California 94952, USA ABSTRACT: We compared 20 yr of seabird breeding data with oceanographic variables (sea level, sea-surface temperature) and a climate index (Southern Oscillation Index; SOI) for the lesser noddy Anous tenuirostris, brown noddy A. stolidus, sooty tern Onychoprion fuscata and wedge- tailed shearwater Ardenna pacifica at the Houtman Abrolhos, eastern Indian Ocean. We found that timing of breeding and performance of these species is related to climate variability, mediated by oceanographic conditions within the eastern boundary Leeuwin Current (LC). The 3 migratory species have breeding parameters which are correlated with sea level and the SOI, while the first egg date of the resident lesser noddy is correlated with SST during their pre-breeding phase. There was poor breeding performance for each species during El Niño Southern Oscillation (ENSO) conditions; moreover, poor breeding success was recorded for all species outside of ENSO events in the latter years of the study. In addition, breeding was delayed for all species during the time of the present study. The environmental conditions contributing to this appear to be fewer years of strong LC flow with associated high sea levels, warmer SSTs, stronger eddy energy and increased productivity. Migratory seabird species breeding at this location appear less able to respond to climate variability — and its influence upon prey availability — than the resident spe- cies. It is suggested that the resident lesser noddy may have an advantage due to its response to a localised environmental cue, SST, in the pre-breeding phase, which may be correlated with prey availability in the waters surrounding the Houtman Abrolhos. KEY WORDS: Seabirds · Indian Ocean · ENSO · Climate · Variability · Leeuwin Current · Demography · Lesser noddy Resale or republication not permitted without written consent of the publisher INTRODUCTION ditions such as wind speed and sea-surface tempera- ture (SST; Ramos et al. 2002, Smithers et al. 2003, A diversity of breeding strategies are demonstrated Peck et al. 2004, Frederiksen et al. 2004, Jaquemet et by seabirds to accommodate local trophic connections al. 2007) that may limit the abilities of seabirds to and ensure that reproductive success is maximized adapt to changes in prey availability. Marine ecosys- (Lack 1968). However, seabird breeding parameters tems are changing and are predicted to change sub- (e.g. phenology, participation rates and breeding suc- stantially in response to climate change, with sub - cess) are often affected by large-scale atmospheric sequent impacts to biodiversity and productivity climate phenomena, such as the El Niño Southern (Polo czanska et al. 2007, Burrows et al. 2011). In par- Oscillation (ENSO; Dunlop et al. 2002, Ramos et al. ticular, the timing of food availability is likely to shift 2006, Surman & Nicholson 2009a,b) and the North in most marine environments, and seabirds will need Atlantic Oscillation (NAO; Frederiksen et al. 2004, to adjust their breeding phenology and/ or trophic Wanless et al. 2009), as well as localised weather con- niche if they are to adapt to climate change. *Email: [email protected] © Inter-Research 2012 · www.int-res.com 148 Mar Ecol Prog Ser 454: 147–157, 2012 The ability to respond to climate variability, and fisheries recruitments along the continental slope of variation in food availability (Ainley et al. 1995, the Western Australian coastline (Caputi et al. 2009, Nicholson 2002, Surman & Nicholson 2009a,b), Feng et al. 2009, Thompson et al. 2011). This region is appears to be influenced by life-history traits. Re - one of generally low biological productivity, as the sponse to changing regional cues by seabirds in 2 low-density, warm waters of the LC flow southwards North Sea colonies showed contrasting patterns, with over the top of the cooler, denser, and northward- the breeding phenology of some species becoming flowing Westralia Current, preventing upwellings of earlier, some not changing at all, and still others nutrient-rich waters into the photic zone (Feng et al. becoming later, which appeared to reflect differing 2009). As a consequence, seabird abundance in the life-history traits such as foraging method and diet eastern Indian Ocean is much lower, and tropical spe- (Wanless et al. 2009). Life-history traits such as a pro- cies occur much further south, than at equivalent lati- tracted post-natal growth period can inhibit such tudes off the west coasts of Africa and South America, plasticity in timing of breeding in response to envi- which possess northward-flowing currents and strong ronmental cues, in which case other breeding para- coastal upwellings (Wooller et al. 1991). The strength meters such as participation and success decrease in of flow of the LC and its eddy field varies seasonally, response to low food availability. For example, the mostly due to variation of surface winds, as well as in- wedge-tailed shearwater Ardenna pacifica (previously terannually (with the occurrence of ENSO events) in Puffinus pacificus), a highly synchronous migrant its cycle (Feng et al. 2009), with consequent effects tropical seabird species with protracted post-natal upon marine pelagic production. During the austral growth, experiences high variation in breeding pro- winter and in La Niña (non-El Niño) years, the LC ductivity at 4 colonies located between 20° and 29° S flow becomes more rapid, resulting in higher sea lev- in the eastern Indian Ocean, where poor years have els (by up to 20 cm at Fremantle; Feng et al. 2009) and been correlated with strong ENSO events, as mea- warmer sea temperatures along shelf waters of the sured by the Southern Oscillation Index (SOI; Dunlop Western Australian coastline (Cresswell et al. 1989, et al. 2002, Nicholson 2002). Koslow et al. 2008, Thompson et al. 2011). The degree to which a seabird can adjust their breeding phenology in response to climate variability may be dependent upon the environmental cues that they experience during their non-breeding phase. For example, migratory seabird species in the North Sea had an onset of breeding which was correlated with the NAO, whereas a resident species was more af fected by localised SST (Frederiksen et al. 2004). Responding to more localised cues may confer some advantage for breeding success, as tropical seabird species breeding in the eastern Indian Ocean at 29°S experienced later onset of breeding and reduced productivity during seasons coinciding with ENSO events, although a resident species was less affected than the migratory species (Surman & Nicholson 2009a,b). In the eastern Indian Ocean, interannual variability of the ENSO cycle induces changes in the mesoscale variability (e.g. eddy shedding) of the poleward flow- ing Leeuwin Current (LC; Fig. 1) (Feng et al. 2009); the LC is the primary driver of marine ecosystem pro- ductivity along the Western Australian coastline. The LC is sourced from the Pacific via the Indonesian Through-Flow, and flows between latitudes 22° and Fig. 1. Bathymetry of the Western Australian coastline, de- 34° S before turning eastwards and continuing into picting the Indonesian Through-Flow (ITF), the Leeuwin Current (LC), the Eastern Gyral Current (EGC), the Wes- the Great Australian Bight. Seasonal and interannual tralia Current (WC) and the approximate positioning of the variability of the LC is found to be a key factor influ- seasonal (autumn/winter) meso-scale eddies located near encing coastal SST, sea level, marine productivity and the Houtman Abrolhos Surman et al.: Long-term climate effects upon eastern Indian Ocean seabirds 149 Strong annual and seasonal differences in marine umes of key prey species in regurgitates, particularly pelagic productivity associated with the biophysical in the noddy species (Surman & Nicholson 2009a,b). properties in LC mesoscale eddies have been identi- Over time, increasing years of poor breeding per- fied (Koslow et al. 2008, Feng et al. 2009), particu- formance for all species were recorded outside larly north and south of the Houtman Abrolhos ENSO events, accompanied by a significant seasonal region (29° S; Fig. 1). The Houtman Abrolhos is the delay in the onset of breeding (date of first egg) in the central breeding area for the eastern Indian Ocean 3 tern species (Surman & Nicholson 2009a). Years of metapopulations of the lesser noddy Anous tenui - poor breeding performance were not restricted to the rostris, brown noddy A. stolidus, sooty tern Ony- Houtman Abrolhos seabird colonies, as wedge-tailed choprion fuscata and wedge-tailed shearwater. This shearwater colonies between 20° and 29° S also had region contains one of the southernmost extensions poor productivity in the same seasons (Dunlop et al. of tropical marine flora and fauna in the Indian 2002), suggesting that these seabirds were all re - Ocean, principally due to the southward flow of the sponding to the same large-scale environmental cue LC, which provides breeding seabirds with a mix- (ENSO variability). Therefore, our over-arching hy - ture of tropical and temperate prey sources (Surman pothesis is that timing of breeding and breeding per- & Wooller 2003). In the present paper, we used 20 yr formance by seabirds are related to climate variabil- of data (1991 to 2010) to compare seabird response ity, which is mediated by oceanographic conditions to large-scale, regional and fine-scale climatic sig- within the LC. Specifically, we predict that timing of nals, as indicated by variation in the breeding para- breeding of the noddy and tern species (e.g. egg lay meters of these 4 species.