Survival of microscopic stages of a perennial kelp (<Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>) from the center and the southern extreme of its range in the Northern Hemisphere after exposure to simulated El Niño stress

Many organisms survive stressful conditions through a tolerant life history stage. The life history known as the alternation of generations is typical of temperate kelps, producing diploid macroscopic stages, and both haploid and diploid microscopic stages, with the haploid stages thought to be stress tolerant. The survival of microscopic stages of the giant kelp Macrocystispyrifera during El Niño has been suggested, yet has never been tested. This mechanism could be critical for population persistence, particularly at the southern limit of the range in the Northern Hemisphere, which is greatly impacted by El Niño conditions. The purpose of this study was to determine if microscopic stages of giant kelp could survive and recover from El Niño-type conditions and whether those from a population near its southern limit were more tolerant than a population at the center of its range. Microscopic stages were exposed to a laboratory simulation of potential El Niño conditions (high temperature, with and without light and nitrate) for 8 weeks and then allowed to recover at optimal conditions (low temperature and high nitrate) for 8 weeks, while controls were left at optimal conditions the entire 16 weeks test period. Haploid developmental stages from both populations survived and recovered from stressful conditions with no population level effect, suggesting haploid stress-tolerance may be widespread. The more advanced the developmental stage, and the presence of nitrate, resulted in significantly greater recovery for haploids. Yet, none of these stages were able to go on to produce sporophytes, whereas all controls did. There was a large population-level effect for diploids, however, with only microscopic diploid stages (embryonic sporophytes) from the southern-limit population recovering from El Niño simulated stress, suggesting ecotypic adaptation for microscopic sporophytes. Diploid recovery was significantly greater with light. We propose that the diploid stage is the most likely to survive and recover after El Niño conditions, as it would avoid obligate egg and sperm encounters after the stress period. The survivorship of the microscopic diploid in a seed bank analogue may be how the isolated southern-limit populations are able to recover after mass disappearance during El Niño.     

Reef recovery 20 years after the 1982–1983 El Niño massive mortality

For over 20 years the El Niño-Southern Oscillation (ENSO) has caused damage to the coral reefs of the eastern Pacific and other regions. In the mid-1980s scientists estimated that coral cover was reduced by 50–100% in several countries across the region. Almost 20 years (2002) after the 1982–1983 event, we assessed the recovery of the virtually destroyed reefs at Cocos Island (Costa Rica), previously evaluated in 1987 and reported to have less than 4% live coral cover. We observed up to fivefold increase in live coral cover which varied among reefs surveyed in 1987 and 2002. Most new recruits and adults belonged to the main reef building species from pre-1982 ENSO, Porites lobata, suggesting that a disturbance as outstanding as El Niño was not sufficient to change the role or composition of the dominant species, contrary to phase shifts reported for the Caribbean. During the 1990s, new species were observed growing on the reefs. Notably, Leptoseris scabra, considered to be rare in the entire Pacific, was commonly found in the area. Recovery may have begun with the sexual and asexual recruits of the few surviving colonies of P. lobata and Pavona spp. and with long distance transport of larvae from remote reefs. We found an overall 23% live coral cover by 2002 and with one reef above 58% indicating that Cocos Island coral reefs are recovering.     

Effects of El Niño on distribution and reproductive performance of Black Brant

Climate in low-latitude wintering areas may influence temperate and high-latitude breeding populations of birds, but demonstrations of such relationships have been rare because of difficulties in linking wintering with breeding populations. We used long-term aerial surveys in Mexican wintering areas and breeding areas in Alaska, USA, to assess numbers of Black Brant (Branta bernicla nigricans; hereafter brant) on their principal wintering and breeding area in El Ni?o and non-El Ni?o years. We used Pollock's robust design to directly estimate probability of breeding and apparent annual survival of individually marked brant at the Tutakoke River (TR) colony, Alaska, in each year between 1988 and 2001. Fewer brant wintered in Mexico during every El Ni?o event since 1965. Fewer brant were observed on the principal breeding area following each El Ni?o since surveys began in 1985. Probability of breeding was negatively related to January sea surface temperature along the subtropical coast of North America during the preceding winter. Between 23% (five-year-olds or older) and 30% (three-year-olds) fewer brant nested in 1998 following the strong El Ni?o event in the winter of 1997-1998 than in non-El Ni?o years. This finding is consistent with life history theory, which predicts that longer-lived species preserve adult survival at the expense of reproduction. Oceanographic conditions off Baja California, apparently by their effect on Zostera marina (eelgrass), strongly influence winter distribution of brant geese and their reproduction (but not survival), which in turn affects ecosystem dynamics in Alaska.     

Can ocean acidification affect population dynamics of the barnacle Semibalanus balanoides at its southern range edge?

The global ocean and atmosphere are warming. There is increasing evidence suggesting that, in addition to other environmental factors, climate change is affecting species distributions and local population dynamics. Additionally, as a consequence of the growing levels of atmospheric carbon dioxide (CO2), the oceans are taking up increasing amounts of this CO2, causing ocean pH to decrease (ocean acidification). The relative impacts of ocean acidification on population dynamics have yet to be investigated, despite many studies indicating that there will be at least a sublethal impact on many marine organisms, particularly key calcifying organisms. Using empirical data, we forced a barnacle (Semibalanus balanoides) population model to investigate the relative influence of sea surface temperature (SST) and ocean acidification on a population nearing the southern limit of its geographic distribution. Hindcast models were compared to observational data from Cellar Beach (southwestern United Kingdom). Results indicate that a declining pH trend (-0.0017 unit/yr), indicative of ocean acidification over the past 50 years, does not cause an observable impact on the population abundance relative to changes caused by fluctuations in temperature. Below the critical temperature (here T(crit) = 13.1 degrees C), pH has a more significant affect on population dynamics at this southern range edge. However, above this value, SST has the overriding influence. At lower SST, a decrease in pH (according to the National Bureau of Standards, pHNBs) from 8.2 to 7.8 can significantly decrease the population abundance. The lethal impacts of ocean acidification observed in experiments on early life stages reduce cumulative survival by approximately 25%, which again will significantly alter the population level at this southern limit. Furthermore, forecast predictions from this model suggest that combined acidification and warming cause this local population to die out 10 years earlier than would occur if there was only global warming and no concomitant decrease in pH.     

Distribution of squid paralarvae, <Emphasis Type="Italic">Loligo opalescens</Emphasis> (Cephalopoda: Myopsida), in the Southern California Bight in the three years following the 1997–1998 El Niño

Large numbers of paralarvae of the California market squid, Loligo opalescens (10,560 paralarvae from 422 plankton samples), were collected in the Southern California Bight in 1999, 2000, and 2001 during the spawning season. Paralarval abundance increased dramatically (P<0.0041) from 1.5 squid/1,000m³ in 1999 to 77.9 squid/1,000m³ in 2000, and 73.6 squid/1,000m³ in 2001, following the El Niño of 1997-1998. The effects on the squid fishery of the 1997-1998 El Niño were thus extended for two years, with larval abundance reduced until the 1999-2000 spawning season. Paralarvae were abundant close to shore for up to a month after hatching in 2000 (P<0.003), with tidal surface currents adjacent to shore in the Channel Islands strongly affecting paralarval abundance. Tidally reversing currents within 1-3 km of shore created a boundary layer of "sticky water" within which paralarvae remained entrained inshore immediately after hatching. Neritic currents farther from shore dispersed older paralarvae within the Southern California Bight. The greatest change in paralarval abundance, for all transects, was observed within 1 km of the transition between these two flow regimes. Age of paralarvae (from statolith increments) entrained within the Catalina Island boundary layer averaged 13-16 days, but some individuals remained nearshore for up to a month. Paralarvae in the boundary layer occurred above 80 m depth both day and night, and exhibited a statistically significant pattern of vertical diel migration (P<0.01). Paralarvae at sea were disproportionately abundant adjacent to fronts associated with uplifted isotherms.     

The effects of an extraordinary El Niño / La Niña event on the size and growth of the squid<Emphasis Type="Italic"> Loligo opalescens</Emphasis> off Southern California

The population structure of the California market squid Loligo opalescens was studied for the Channel Islands region off Southern California between June 1998 and March 2000. During this time Californian waters were exposed to an extraordinary El Niño event that was possibly the most dramatic change in oceanographic conditions that occurred last century. There was then a rapid transition to record cool La Niña conditions. Statolith increments were used to determine age parameters and increment periodicity was validated for the first 54 days of life. Based on statolith increment counts, the oldest males and females were 257 and 225 days respectively and individuals matured as young as 129 and 137 days respectively. No distinct hatching period was detected. There was a general trend of increasing body size throughout the study period. Squid that hatched and grew through the El Niño were strikingly smaller and had slower growth rates compared to squid that grew through the La Niña. This was related to oceanography and associated productivity. There was a positive correlation between squid mantle length and upwelling index and a negative correlation between mantle length and sea temperature. The 'live-fast die-young' life history strategy of squid makes them ideal candidates for following the effects of the dramatic changes in oceanographic conditions off California. We propose that squid can serve as ecosystem recorders and productivity integrators over time and space and are useful organisms to tie oceanography to biology.     

El Niño periods increase growth of juvenile white seabass (<Emphasis Type="Italic">Atractoscion nobilis</Emphasis>) in the Southern California Bight

Studies of the impact of El Niño periods on marine species have usually focused on negative, highly visible effects, e.g., decreasing growth rates or increasing mortality due to a decline in primary productivity in typically nutrient rich upwelling zones; but positive effects related to elevated water temperature are also known. This study examined how the growth rate of juvenile white seabass, Atractoscion nobilis, responded to changes in ocean temperature in an El Niño period (1997–1998) in the northern portion of the Southern California Bight, USA. Growth rates of juvenile white seabass during their first 4 years of life were estimated as the slopes of linear relationships between body mass and age (from otoliths) of 800 fish collected at 11 stations throughout the bight. Growth rates differed significantly among cohorts hatched in 1996–2001. Specifically, white seabass that hatched in 1996 and 1997 grew significantly faster than those that hatched in 1998, 1999, and 2001. These differences in growth rates of cohorts appeared to be driven by variation in sea-surface temperature (SST). Growth rates averaged over the first three or 4 years of life were significantly positively correlated to average daily SST during the first 1–4 years of life. Increased growth of juvenile white seabass during the warm El Niño period likely provided a number of benefits to this warm-temperate species. This study demonstrated that some species will benefit from these warm-water periods despite reduced system-wide primary production.     

Spatial and social connectivity of fish-eating “Resident” killer whales (Orcinus orca) in the northern North Pacific

The productive North Pacific waters of the Gulf of Alaska, Aleutian Islands and Bering Sea support a high density of fish-eating “Resident” type killer whales (Orcinus orca), which overlap in distribution with commercial fisheries, producing both direct and indirect interactions. To provide a spatial context for these interactions, we analyzed a 10-year dataset of 3,058 whale photo-identifications from 331 encounters within a large (linear ~4,000 km) coastal study area to investigate the ranging and social patterns of 532 individually identifiable whales photographed in more than one encounter. Although capable of large-scale movements (maximum 1,443 km), we documented ranges generally <200 km, with high site fidelity across summer sampling intervals and also re-sightings during a winter survey. Bayesian analysis of pair-wise associations identified four defined clusters, likely representing groupings of stable matrilines, with distinct ranging patterns, that combined to form a large network of associated whales that ranged across most of the study area. This provides evidence of structure within the Alaska stock of Resident killer whales, important for evaluating ecosystem and fisheries impacts. This network included whales known to depredate groundfish from longline fisheries, and we suggest that such large-scale connectivity has facilitated the spread of depredation.     

Reef coral reproduction in the eastern Pacific: Costa Rica, Panamá and Galápagos Islands (Ecuador). IV. Agariciidae, recruitment and recovery of Pavona varians and Pavona sp.a

The reproductive ecology of two eastern Pacific zooxanthellate coral species was examined as part of a continuing series of studies relating bleaching/mortality events caused by the El Niño–Southern Oscillation disturbance, and is described for study sites in Costa Rica, Panamá, and the Galápagos Islands (Ecuador). This study deals with the sibling agariciid species Pavona varians and Pavona sp.a over a 13?yr period (1985 to 1997). Both Pavona species are broadcast-spawners with some gonochoric, but mostly sequential hermaphroditic colonies. Minimum colony sizes (and ages) at first reproduction were 5?cm (5?yr) and 3?cm (2 to 3?yr), respectively, in P. varians and Pavona sp.a. In the Panamá and Galápagos populations, gonochoric colonies spawn eggs or sperm at least monthly. Six fecundity attributes were not significantly different in the two species, but the eggs of P. varians are white to beige and positively buoyant, and those of Pavona sp.a are dark green and neutrally to negatively buoyant. Eggs of both species lack zooxanthellae. Both species are reproductively active year-round, with maximum activity in the dry season in the nonupwelling Gulf of Chiriquí, and in the wet season in the upwelling Gulf of Panamá. Spawning is predominantly during full moon, and possibly also at new moon at most study sites. Spawning in P. varians and Pavona sp.a is 12?h out of phase, with the former species spawning ～1?h before sunrise and the latter about 1?h after sunset. The fecundity of Pavona spp. at Caño and the Galápagos Islands was much greater (19?900 to 27?900 eggs cm^?2?yr^?1) than at all Panamá sites (14?800 to 19?800 eggs cm^?2?yr^?1). Intraspecific crosses in both species resulted in swimming planula larvae after 25 to 36?h. Recruitment of P. varians was highest in Panamá, moderate in Costa Rica, and nil in the Galápagos Islands, matching, respectively, the contributions of P. varians to the pre-1982/1983 El Niño coral-population abundances in these areas. Recruitment success of P. varians at Uva Island was significantly related to maximum monthly positive sea surface-temperature (SST) anomalies that occurred in the year preceding recruitment over the period 1982 to 1996; recruitment failed when SST anomalies exceeded 1.6 to 1.9?C° during the severe ENSO events of 1982/1983 and 1997/1998.     

Reproductive biology and population structure of the sea cucumber <Emphasis Type="Italic">Isostichopus fuscus</Emphasis> (Ludwig, 1875) (Holothuroidea) in Caamaño,Galápagos Islands,Ecuador

Pre-spawning reproductive biology and population structure of the sea cucumber Isostichopus fuscus were analysed at the Caamaño Island, Galápagos Islands, Ecuador on a monthly basis between January 1995 and June 1996. An average of 25 individuals of all sizes were collected each month. Using microscopic characteristics of the gonads three gonadal stages were described: (1) immature; (2) mature, and (3) spent. Mature individuals showed statistically different morphometric characteristics. I. fuscus was continuously sexually mature throughout the year independent of changes in sea water temperature. Mean oocyte diameter indicated planktotrophic larval development. Mean fecundity for this species was 567,884 (±95,528 SE) eggs per gonad. Size at first maturity was between 161.0 and 170.9 g drained weight and the smallest individual found with mature gonads had a drained weight between 121.0 and 130.9 g. The population sampled ranged between 13 and 31 cm (81.0–400.0 g) with less than 10% of the population of individuals ≥25 cm (≥300.0 g). No juveniles were recorded.