Cephalopod prey of the wandering albatross Diomedea exulans

Cephalopod beaks from the stomach contents of wandering albatross (Diomedea exulans L.) chicks from Bird Island, South Georgia, were sampled between May and September in 1983 and 1984. Lower beaks were identified and measured, and allometric data were used to calculated mantle length and biomass of the species consumed. A total of 3421 lower beaks were examined, representing 35 species in the 1983 sample and 45 species in the 1984 sample. Eight of the twenty families contributed over 95% of the biomass. In 1984 there were less Onychoteuthidae and more Ommastrephidae than in 1983 and a decrease in the number of species known to occur south of the Antarctic Polar Front. There was a difference in the size-frequency distribution of the cephalopod diet in the two years; in 1984 there was a higher frequency of intermediate-sized specimens, reflecting the greater importance of ommastrephids, especially Illex sp. The energy content of cephalopods in 1984 may have been greater than in 1983. Serial sampling of cephalopod beaks during the austral winter did not reveal evidence of growth. By the age of 200 d, wandering albatross chicks have consumed a total of approximately 100 kg wet weight of cephalopods each.     

Seasonal changes in the diet and feeding behaviour of a top predator indicate a flexible response to deteriorating oceanographic conditions

Shifts in the diet of top predators can be linked to changes in environmental conditions. In this study, we tested relationships between environmental variation and seasonal changes in diet of a top predator, the grey-headed albatross Thalassarche chrysostoma, breeding at Bird Island, South Georgia in an austral summer of 1999/2000. Oceanographic conditions in that year around South Georgia were abnormal (i.e. anomalously high sea surface temperature to a relative 19-year long-term mean). The diet of grey-headed albatrosses showed high seasonal variation, shifting from cephalopods (42.9 % by mass) in late February to Antarctic krill Euphausia superba (58.3 %) in late April, and grey-headed albatrosses breeding performance was low (16.8 %). This study shows these albatrosses did not manage to find sufficient alternative prey and highlight the risk to top predators if there is an increase in the frequency or severity of food shortages in Antarctic waters.     

Cephalopod remains from the stomach of a sperm whale stranded off Patagonia

The stomach of a male sperm whale (Physeter macrocephalus) stranded near Comodoro Rivadavia, Argentina, on 21 March 1986, contained 247 cephalopod beaks (128 upper and 119 lower). The lower beaks were identified, their rostral lengths were measured and used to estimate the mantle lengths and wet weight of the representatives of each cephalopod taxon. Fifteen species from nine families were represented. The most numerous species, in order of decreasing abundance, were the large cranchiidMesonychoteuthis hamiltoni (32.8% of total numbers),Mastigoteuthis sp. A (16.0%), andKondakovia longimana andHistioteuthis atlantica (both 13.5%). Almost 85% of the estimated total weight was contributed by two species,Mesonychoteuthis hamiltoni (61.6%) andK. longimana (23.1%). The presence of beaks fromK. longimana, Alluroteuthis antarcticus andM. hamiltoni, which are all thought to be restricted to the Antarctic, suggests that the whale had migrated North prior to stranding.     

Feeding strategies and diets of breeding grey-headed and wandering albatrosses at South Georgia

The foraging areas and diets of the grey-headed albatross Thalassarche chrysostoma and wandering albatross Diomedea exulans were studied in March/April 2000 at Bird Island, South Georgia, during their respective chick-rearing and brood-guard periods. Oceanographically, March/April 2000 was abnormal, with warm conditions close to South Georgia. These conditions affected albatross foraging behaviour, particularly that of grey-headed albatrosses. Both species tended to forage in different areas of the ocean, with significant differences in trip durations. Grey-headed albatrosses (n=9) foraged mainly in Antarctic waters (predominantly shelf waters of the South Shetland Islands and Antarctic Peninsula, and also in oceanic waters around South Georgia), feeding mainly on krill (Euphausia superba; 77% by mass). Foraging trips lasted 13.3 days (range: 5–26 days), far longer than the 1–3 days found in previous studies. Only one grey-headed albatross was associated with the APF (Antarctic Polar Front), a reported foraging area in recent studies. Wandering albatrosses (n=9) foraged in Antarctic (South Georgia Shelf) and Antarctic Polar Frontal Zone (APFZ) waters, with trips of 1–4 days trip duration (usual for this species), feeding on fish (46% by mass) and cephalopods (32%). One bird was associated with the APF, and two birds foraged on the shelf/shelf break over the Patagonian shelf. These findings suggest that sea surface temperature anomalies, produced by movement of the APF closer to South Georgia or by eddies, may have had an effect on the foraging strategy of grey-headed albatrosses that year (the main prey of grey-headed albatrosses in previous studies, the ommastrephid Martialia hyadesi, is known to be associated with the APF). Also, when both albatross breeding periods overlap, their foraging areas were complementary, which reflected the prey taken.     

Offspring sex ratio in relation to parental structural size and body condition in the long-lived wandering albatross (<Emphasis Type="Italic">Diomedea exulans</Emphasis>)

Sex ratio theory is one of the most controversial topics in evolutionary ecology. Many deviations from an equal production of males and females are reported in the literature, but few patterns appear to hold across species or populations. There is clearly a need to identify fitness effects of sex ratio variation. We studied this aspect in a population of a long-lived seabird, the wandering albatross (Diomedea exulans), using molecular sex-identification techniques. We report that parental traits affect both (1) fledgling traits in a sex-dependent way and (2) chick sex: Sons are overproduced when likely to be large at fledging and, to a lesser extent, daughters are overproduced when likely to be in good body condition at fledging. Because for the same population, a previous study reported that post-fledging survival was positively affected by size in males and by body condition in females, our results suggest that wandering albatrosses manipulate offspring sex to increase post-fledging survival.     

Cephalopod prey of the grey-headed albatrossDiomedea chrysostoma

Cephalopod remains were collected, at regular intervals throughout the fledging period, from the stomach contents of chicks of the grey-headed albatrossDiomedea chrysostoma at Bird Island, South Georgia, in 1984 and 1986 and from regurgitations of adults at the nest in 1986. The 1984 sample was taken during a season characterised by abnormal local oceanographic conditions in which the breeding success was very low; in 1986 conditions were normal and breeding success was high. Cephalopod beaks (289 from adults; 5 651 from chicks) were identified, and allometric equations were used to estimate the biomass represented. Five cephalopod species belonging to five families (Gonatidae, Onychoteuthidae, Psychroteuthidae, Ommastrephidae and Cranchiidae) contributed 98% by number and 97% of the biomass fed to chicks. The most important species was the ommastrephidMartialia hyadesi, contributing 68.9 to 77.4% by number and 72.5 to 79.3% of the total biomass fed to chicks. The relative proportions of cephalopod species in the chicks' diet were similar between 1984 and 1986, but the total number and biomass was significantly less in 1984. There is evidence of growth ofM. hyadesi between January and June.     

Trophic ecology of Grey-headed albatrosses from Marion Island, Southern Ocean: insights from stomach contents and diet tracers

During chick-rearing, albatrosses can alternate between long foraging trips that provide the main source of food for the adults and short foraging trips that they use to feed their young. This flexibility in foraging behaviour can lead to differences in diet composition between adults and chicks and implies that they may be vulnerable in different ways to food shortages. The trophic ecology of the Grey-headed albatross Thalassarche chrysostoma was investigated at the sub-Antarctic Prince Edward Islands during the chick-rearing period in April 2006 using a combination of approaches. Diets of adults and chicks were assessed using stable isotope ratios and fatty acid (FA) profiles of blood and/or stomach oils, in addition to stomach contents analysis. Fish from the family Macrouridae and cephalopods (particularly the onychoteuthid Kondakovia longimana) were the primary prey, whereas crustaceans (krill Euphausia superba) represented a smaller proportion of the stomach contents. Stomach oil FA profiles contained more monounsaturated FA than the profiles of plasma, which were richer in saturated FA and arachidonic acid (20:4n-6). There was also a distinct separation of adults from chicks, with higher levels of monounsaturates in chick plasma, and higher saturated FA levels (particularly 16:0) in the adult plasma. Stable carbon isotope ratios of whole blood were similar in adults and chicks, whereas stable nitrogen isotope ratios showed significant enrichment by >1‰ in chicks. The combined FA, stable isotopes and stomach contents analyses suggest clear differences in diet quality between adults and chicks, with chicks feeding at a higher trophic position through feeding more on highly nutritious fish and adults keeping much of the less nutritious zooplankton for themselves.     

Importance of the Japanese anchovy (Engraulis japonicus) to breeding rhinoceros auklets (Cerorhinca monocerata) on Teuri Island, Sea of Japan

Temporal variation in the diet and chick growth of rhinoceros auklets (Cerorhinca monocerata), on Teuri Island, Sea of Japan, was studied to understand how local marine environmental changes affect the reproduction of this piscivorous seabird. The food delivered by parents to chicks was sampled every 1-2 weeks from late May to July, 1994-1998. Overall, the diet of nestling rhinoceros auklets consisted of (by mass) 61% Japanese anchovy (Engraulis japonicus), 18% Japanese sand lance (Ammodytes personatus), 18% Japan Sea greenling (Pleurogrammus azonus), 2% other fish and 1% squid. Among years, the contribution of anchovy ranged from 16% to 93%. Once anchovy occurred in the diet, it dominated (80% on average) thereafter. Accordingly, when anchovy appeared in the diet early in the chick-rearing season (1994, 1998), the contribution of anchovy overall was large. The first appearance of anchovy in the diet of auklets late in the summer of 1997 was possibly related to negatively anomalous sea-surface temperature. Food loads composed of anchovy (34.0 g) were heavier than those of sand lance (22.5 g) and greenling (28.5 g). The energy density of anchovies also was higher: 6.3 kJ g^-1 wet mass compared to 0+ greenling (4.78 kJ g^-1) and 0+ sand lance (3.78 kJ g^-1). Thus, a high proportion of anchovy in the diet resulted in high food load mass, high daily growth rates of chicks and high fledging success. This study highlighted the importance of the time of arrival of migratory high-lipid prey, which is influenced by local oceanographic conditions, to the reproductive performance of a piscivorous seabird.     

Spatial and temporal variation in the diet of a high trophic level predator,the Australian fur seal (<Emphasis Type="Italic">Arctocephalus pusillus doriferus</Emphasis>)

This study quantifies the manner in which Australian fur seals, Arctocephalus pusillus doriferus, use their prey in a spatial and temporal context. We analysed 977 scat and 66 regurgitate samples collected from Tasmanian breeding colonies and haul-outs between 1994 and 2000. Diagnostic prey remains identified in the scats represented 35 fish taxa and 8 cephalopod taxa. The main taxa identified in scats, where frequency of occurrence was 10%, were leatherjacket species (family Monocanthidae), redbait (Emmelichthys nitidus), barracouta (Thyrsites atun), jack mackerel (Trachurus declivis) and red cod (Pseudophysis bachus). Regurgitates were dominated by cephalopods, primarily Goulds squid (Nototodarus gouldi), Octopus maorum, O. berrima/pallidus and Sepia apama. Discriminant function analyses indicated that there were generally no significant differences in the composition of the diet between colonies within a year, suggesting that prey distribution is fairly uniform throughout Bass Strait at those time scales. The diet at breeding colonies, however, exhibited significant inter- and intra-annual variation, determined by the presence of several key taxa, such as barracouta and a species of scorpionfish (family Scorpaenidae). The diet composition also varied regionally, between Bass Strait and southern Tasmania in spring 1999 and autumn 2000, with redbait, barracouta and a species of scorpionfish identified as the main taxa contributing to this difference. Redbait occurred in the diet only in southern Tasmania, whereas barracouta and scorpionfish occurred only in Bass Strait.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-003-1219-0.Communicated by G.F. Humphrey, Sydney     

Diet and nest attendance of incubating and chick-rearing northern fulmars (Fulmarus glacialis) in Shetland

We examined diets and nest attendance patterns of northern fulmars (Fulmarus glacialis) on Foula and Unst, Shetland, UK, during the breeding seasons of 1998 and 1999. Mean foraging trip duration, derived from nest attendance patterns, differed between incubation (32.5 h) and chick-rearing (early stage: 11.2 h, mid-stage: 20.4 h). It was influenced by chick age and obviously also by prey availability. The numbers of fulmars attending the colonies were influenced by wind direction, wind speed, time of day and state of reproduction. Diet samples were collected by regurgitations of adult and young fulmars. Diet was analysed by identifying prey items on the basis of otoliths, vertebrae, premaxillae (fish), cephalopod beaks and fragments of arthropod exoskeletons. Adult fulmars and chicks fed upon a wide range of prey types. Gadoid fish, including Norway pout (Trisopterus esmarkii), were the most common prey in the diet. Fish offal was found in 32% of regurgitates, clupeids in 15%. In contrast, the proportions of sandeels were very low (1%). This is different from previous studies where sandeels formed a large part of fulmar diet at Shetland in summer. Differences in food composition of incubating and chick-rearing fulmars were found for fish, but not for offal, crustaceans or squid. The percentage of non-discard fish (Clupeidae, Ammodytidae, Isospondylae) was significantly different between regurgitates from incubating and chick-rearing fulmars. Samples from chick-rearing adults contained non-discard items more frequently than regurgitates from incubating birds. The results strongly indicate that fulmars select to feed their chicks on energy-rich clupeids.     

Impact of the supplementation of a docosahexaenoic acid–rich emulsion on the reproductive output of oyster broodstock, Crassostrea gigas

Pacific oyster [Crassostrea gigas (Thunberg)] broodstock were hatchery-conditioned with either a mixed algal diet of Dunaliella tertiolecta, Tetraselmis suecica and Rhodomonas sp. [1:1:1, on a dry weight (DW) basis] (Control), a single diet of D. tertiolecta (Duna) or a single diet of D. tertiolecta supplemented with an emulsion rich in eicosapentaenoic (20:5n-3) and particularly docosahexaenoic acid (22:6n-3) (Duna+Em). Oysters were spawned after a conditioning period of 7 and 8 weeks and larvae were reared for 1 week on an Isochrysis galbana (clone T-Iso) diet. The percentage recovery of D-larvae from eggs of Control-fed oysters and Duna+Em-fed oysters was not significantly different (80% and 63% after 7 weeks of conditioning; 87% and 74% after 8 weeks of conditioning, respectively] but was much higher than from eggs produced by Duna-fed oysters (24% and 41% after 7 and 8 weeks of conditioning, respectively). A reduction in the percentage recovery of D-larvae as a result of temperature or salinity stress was most severe in eggs from Duna-fed oysters. Neither the broodstock diet nor the conditioning period affected the size of eggs or D-larvae, larval performance, or lipid content and lipid class distribution of the eggs. The fatty acid composition of the eggs was modified by the fatty acid composition of the algal diet and the lipid emulsion. The supplementation of the emulsion resulted in a pronounced increase in the percentage of 22:6n-3 in the polar (PL) and neutral lipids (NL) of the eggs (4.0-13.4% and 8.5-14.7% in NL and PL, respectively). The egg lipids (7-8 ng egg^-1) contained about 62% NL and were dominated by triglycerides (TAG), which made up 65% of the NL. The major phospholipids were phosphatidylethanolamine (PE), ceramide aminoethylphosphonate and phosphatidylinositol (CAEP+PI), phosphatidylserine (PS) and phosphatidylcholine (PC). Regardless of the broodstock diet treatment, the PL fraction contained high percentages of non-methylene-interrupted dienes (NMID) and plasmalogens (detected as dimethyl acetals).     

The trophic link between squid and the emperor penguin <Emphasis Type="Italic">Aptenodytes forsteri</Emphasis> at Pointe Géologie,Antarctica

Cephalopod beaks retrieved from stomachs of dead emperor penguin chicks at Pointe Géologie, Terre Adélie, provide information on taxonomic and size composition of the penguin’s squid diet, on the trophic range of the squid species preyed upon and on the fractional trophic impact of the penguin on the whole food web. Emperor penguins prey upon four squid species (Psychroteuthis glacialis, Kondakovia longimana, Gonatus antarcticus, Alluroteuthis antarcticus) and do not take squid larger than 480 mm mantle length. Larger squid live either below the penguin’s diving range or are beyond its handling capacity. Nitrogen stable isotope ratios indicate that squids cover a range of about two trophic levels (2.5–8‰ δ¹⁵N). The impact of the emperor penguin, however, concentrates on the upper part of this range, about 68% of its squid prey being >6‰ δ¹⁵N. The principal components of the emperor’s diet, fish, krill and squid, differ distinctly in average trophic level. Consequently the trophic position of the emperor penguin changes accordingly with diet composition and may differ by almost one trophic level between different emperor penguin colonies.     

Fish and squid in the diet of king penguin chicks,Aptenodytes patagonicus,during winter at sub-antarctic Crozet Islands

The diet of king penguins, Aptenodytes patagonicus, rearing chicks was studied during three consecutive austral winters (1990, 1991 and 1992) at Crozet Islands. The mean stomach content mass of the 47 samples was 503 g. Percentages of wet and reconstituted masses showed that both fishes (66 and 36%, respectively) and squid (34 and 64%) are important components of the winter diet. Juveniles of the demersal onychoteuthid squid Moroteuthis ingens form the bulk of the cephalopod diet, and this was the main prey by reconstituted mass (57%). Myctophid fish (lantern-fishes) accounted for most of the fish diet, constituting together 32% by mass. The three main species of myctophids eaten in summer by king penguins were either very rare in winter (Electrona carlsbergi) or accounted for a smaller proportion of the diet (Krefftichthys anderssoni = 1.5% by mass and Protomyctophum tenisoni = 4.6%). Five other myctophids, which are rarely consumed in summer, contributed 24% of the diet by mass in winter (Gymnoscopelus piabilis = 18.1%, Lampichthys procerus = 2.4%, G. nicholsi = 1.3%, and Metelectrona ventralis and Electrona subaspera = 1.0%). The greater diversity of prey in winter suggests a more opportunistic feeding behaviour at a time probably marked by a change in prey availability. Both the known ecology of the fish and squid prey and the barely digested state of some items suggest that in winter breeding adults forage in the outer shelf, upper slope and oceanic areas in the close vicinity of the Crozet Islands to feed their chicks. Finally, using king penguins as biological samplers, the present work provides novel data on the previously unstudied mesopelagic/epibenthic marine community in waters surrounding the Crozet Islands. Seventeen myctophid fish have been identified to species level. These include several poorly known species in the southern Indian Ocean. The occurrence of small, nearly intact, cephalopods in the diet of king penguins suggests that spawning grounds of four squid species may be located near the Crozet Archipelago.     

Cephalopod remains from sperm whales caught off western Canada

Of 379 cephalopod beaks sampled from stomachs of 20 sperm whales caught to the west of Vancouver Island, Canada, 152 beaks were sorted and measured. Thirteen types of beak were identified as belonging to eleven families. The most numerous species represented are Berryteuthis magister and Moroteuthis robusta comprising 28.9 and 24.3% respectively. These species were estimated to comprise 17.6 and 61.9% respectively of the weight of flesh represented by beaks.     

Assessment of the state and management of the cephalopod trawl fishery resources in Greek waters

The cephalopod trawl fishery in Greek waters is reviewed in relation to fishing effort for 1964–1981. The mean annual (1964–81) catch of cephalopods amounted to 1348 tons. Total landings of cephalopods are very well described by the exponential⁹ and linear²⁹ surplus yield models, according to which optimum fishing effort, maximum sustained yield and optimum catch per unit of fishing effort range between 120000–150000HP, 1586–1630 tons and 10–13kgr/HP respectively. In addition, the analysis revealed that cephalopod resources were slightly overfished (according to the linear model) during 1975–1981. Hence, authorities must take immediate measures for the protection and rational management of cephalopod resources in Greek waters. Possible alternative measures are discussed in the text.     

Cephalopod remains from the stomach of a sperm whale stranded at Rio Grande do Sul in Brazil

The stomach of a 10.2 m (34 ft) male sperm whale which became stranded at Rio Grande do Sul near Rio Grande, Brazil contained 227 upper cephalopod beaks and 255 lower cephalopod beaks. The lower beaks were identified and measured. Beaks of 14 species belonging to 9 families are present. Lower, rostral lengths are compared with those of the same species collected from stomachs of whales killed off South Africa and in the Antarctic. Estimates of the weight of flesh of the various species represented by beaks are made. Two ommastrephid species are estimated to contribute 33.1% of the weight of flesh represented by beaks while the octopoteuthids contribute 28.3% and the Onychoteuthids 18.5%. Three Antarctic species in the stomach show that the whale had migrated from south of Lat. 40°S prior to being caught and had not eaten much food during the journey which would probably have taken him more than 5 d.     

Comparison of light- and SST-based geolocation with satellite telemetry in free-ranging albatrosses

Light-based archival tags are increasingly being used on free-ranging marine vertebrates to study their movements using geolocation estimates. These methods use algorithms that incorporate threshold light techniques to determine longitude and latitude. More recently, researchers have begun using sea surface temperature (SST) to determine latitude in temperate regions. The accuracy and application of these algorithms have not been validated on free-ranging birds. Errors in both geolocation methods were quantified by double-tagging Laysan (Phoebastria immutabilis Rothschild) and black-footed (P. nigripes Audubon) albatrosses with both leg-mounted archival tags that measured SST and ambient light, and satellite transmitters. Laysan albatrosses were captured and released from breeding colonies on Tern Island, northwestern Hawaiian Islands (23°52N, 166°17W) and Guadalupe Island, Mexico (28°31N, 118°10W) and black-footed albatrosses from Tern Island. Studies were carried out between December 2002 and March 2003. For all birds combined, the mean ± SD great circle (GC) distance between light-based locations and satellite-derived locations was 400±298 km (n=131). Errors in geolocation positions were reduced to 202±171 km (n=154) when light-based longitude and SST-based latitude (i.e. SST/light) were used to establish locations. The SST/light method produced comparable results for two Laysan albatross populations that traveled within distinctly different oceanic regions (open ocean vs more coastal) whereas light-based methods produced greater errors in the coastal population. Archival tags deployed on black-footed albatrosses returned a significantly higher proportion of lower-quality locations, which was attributed to interference of the light sensor on the tag. Overall, the results demonstrate that combining measures of light-based longitude and SST-based latitude significantly reduces the error in location estimates for albatrosses and can provide valid latitude estimates during the equinoxes, when light-based latitude measurements are indeterminate.Communicated by J.P. Grassle, New Brunswick     

Morphology and sinking velocities of fecal pellets of copepod, molluscan, euphausiid, and salp taxa in the northeastern tropical Atlantic

Morphological characteristics and sinking velocities of naturally occurring fecal pellets of copepods, euphausiids, salps, and pelagic mollusks collected in the northeastern tropical Atlantic were investigated during the period of May-June 1992. The fecal pellets of copepods and euphausiids were cylindrical and distinguished only by their size. Those of salps were, in general, rectangular, and slight differences were noted according to the species. The fecal pellets of the molluscan pteropod Clio sp. were conical, while those of the molluscan heteropod Carinaria sp. were spiral. The sinking velocities ranged from 26.5 to 159.5 m day^-1 for copepod fecal pellets, from 16.1 to 341.1 m day^-1 for euphausiid pellets, from 43.5 to 1167.6 m day^-1 for salps' pellets (Cyclosalpa affinis, Salpa fusiformis, Iasis zonaria, and two unidentified species), from 65 to 205.7 m day^-1 for Clio sp. pellets, and from 120.3 to 646.4 m day^-1 for Carinaria sp. fecal pellets. The measured sinking velocities were compared with estimates predicted using the equations of Komar et al. (1981; Limnol Oceanogr 26:172-180), Stokes' law, and Newton's second law, using either a constant density of fecal pellets (1.22 g cm^-3) or densities estimated with the three different equations. When a constant density was used, the three equations overestimated the sinking velocities; Stokes' law resulted in the largest overestimation, and Newton's second law, the smallest. At the taxa level, the overestimation was greatest for euphausiid 1 fecal pellets and smallest for copepod fecal pellets. When the three equations were used to estimate fecal pellet density, the density estimated using the equation of Komar et al. was the greatest, and that using Stokes' law, the smallest, resulting in over- and underestimation of sinking velocities, respectively. Newton's second law resulted in an intermediate density and gave the closest estimate of sinking velocities. We propose that measurement of sinking velocities of a portion of the fecal pellets might guide in choosing an appropriate equation to be used for a reasonable interpretation of vertical mass flux.     

Seasonal fluctuations in the photosynthetic capacity of photosystem II in symbiotic dinoflagellates in the Caribbean reef-building coral <Emphasis Type="Italic">Montastraea</Emphasis>

The photosynthetic capacity of photosystem II (PS II) in symbiotic dinoflagellates (Symbiodinium sp.), as measured by analysis of chlorophyll fluorescence, was investigated in the primary Caribbean reef-building corals, Montastraea annularis and Montastraea faveolata, for 5 years and Montastraea franksi over 2 years in the Bahamas. Significant seasonal fluctuations in the quantum yield of charge separation (F_v/F_m) of PS II were found in all species at all depths, with the highest photosynthetic capacity consistently recorded between mid-winter and early spring and the lowest photosynthetic capacity occurring in the mid to late summer. Corals residing in shallow depths of 1-2 m showed the greatest fluctuations in F_v/F_m, whereas deeper corals (3-4 and 14 m depths) had consistently higher values of F_v/F_m. Densities of symbiotic dinoflagellates and photosynthetic pigments followed a similar pattern. Fluctuations of photosynthetic capacity showed a strong correlation with seasonal patterns of water temperature and light. Such seasonal shifts in photosynthetic capacity are most likely due to several biochemical processes in the algae that lead to alterations of both photoprotection and photodamage. While symbiont density changed significantly on a seasonal basis, visual signs of coral bleaching were noted only in the fall of 1995 and the spring and summer of 1998. Comparisons of photosynthetic capacity and the decrease in the number of symbionts and their subsequent recovery indicated that symbiont populations in this study had the ability to recover quickly following bleaching events, as long as continued physical perturbation (e.g. thermal stress) did not shorten the recovery phase. Large-scale bleaching events are best viewed as the end points of seasonal physiological variation in which photosynthetic capacity and density of symbiotic dinoflagellates are reduced to a lower level than during "non-bleaching" years.     

Alarm responses of the green sea urchin, Strongylocentrotus droebachiensis, induced by chemically labelled durophagous predators and simulated acts of predation

The green sea urchin, Strongylocentrotus droebachiensis, exhibited immediate behavioural responses to waterborne chemosensory cues from two durophagous predators, the Atlantic wolffish, Anarhichas lupus, and the edible crab, Cancer pagurus; as well as from crushed conspecifics and crushed blue mussels, Mytilus edulis. The response patterns were dose dependent and diet dependent. Strong responses were elicited by water conditioned by echinivorous wolffish (97.5%), undiluted urchin extract (73%), undiluted mussel extract (52.5%), and by water conditioned by echinivorous crab (45%). In contrast, urchin extract diluted to 1%, mussel extract diluted to 10%, and water conditioned by predators on a mussel diet elicited weak responses (~20%). Infection by the endoparasitic nematode Echinomermella matsi had no significant effect on the response pattern of S. droebachiensis. There was no conclusive evidence of an alarm response to the predators per se, as the weak response to stimuli from non-echinivorous wolffish and crab, as well as from the seastar Asterias rubens, may have been caused by chemical cues transmitted from their mussel diet. The diet-dependent response to predators suggests that active predators were labelled by chemical cues from their echinoid prey. The chemical cue from echinivorous wolffish acted as both an arrestant and as a repellent, whereas the response to other cues was predominantly or exclusively repellent. The response to echinivorous wolffish was quantitatively stronger than, and qualitatively different from, the response to other stimuli, including undiluted urchin extract. The wolffish is apparently being labelled by a latent chemical cue which derives its potency from activation by, or interaction with, substances in the digestive system of the wolffish. We interpret this novel phenomenon as evidence of alarm signal magnification. The induced behavioural modifications demonstrate the green sea urchins' ability to detect chemical cues associated with active durophagous predators, and would therefore seem to have adaptive potential as a predator defence mechanism.