Molecular prey identification in wild Octopus vulgaris paralarvae

The trophic ecology of Octopus vulgaris paralarvae collected in 2008 off the Ría de Vigo, NW Spain (42° 12.80′ N–9° 00.00′ W), was approached by both morphological and molecular methods. External digestion of prey and posterior suction of the liquefied contents by wild O. vulgaris paralarvae made the morphological identification of gut contents impossible. Thus, a PCR-based method using group-specific primers was selected to identify prey consumed by O. vulgaris paralarvae in the pelagic realm. The mitochondrial ribosomal 16S gene region was chosen for designing group-specific primers, which targeted a broad range of crustaceans and fishes but avoided the amplification of predator DNA. These primers successfully amplified DNA of prey by using a semi-nested PCR-based approach and posterior cloning. Homology search and phylogenetic analysis were then conducted with the 20 different operational taxonomic units obtained to identify the putative organisms ingested. The phylogenetic analysis clustered ingested prey into 12 families of crustaceans (11 belonging to the order Decapoda and 1 to the order Euphausiacea) and two families of fishes (Gobiidae and Carangidae). According to the Czekanowski’s Index (CI), the trophic niche breadth of O. vulgaris paralarvae is low (CI = 0.13), which means that these paralarvae are specialist predators at least during the first weeks of their life cycle. It is the first time that natural prey has been identified in O. vulgaris paralarvae collected from the wild, and such knowledge may be critical to increasing the survival of O. vulgaris hatchlings in captivity, a goal that has been actively pursued since the 1960s by aquaculture researchers.     

Tetrodotoxin affects survival probability of rough-skinned newts (Taricha granulosa) faced with TTX-resistant garter snake predators (Thamnophis sirtalis)

Lethal chemical defenses in prey species can have profound effects on interactions with predators. The presence of lethal defenses in prey can correct the selective imbalance suggested by the life-dinner principle in which the fitness consequences of an encounter between predator and prey should be much greater for the prey species than the predator. Despite the apparent adaptive advantages of lethality the evolution of deadly prey presents a fundamental dilemma. How might lethal defenses confer an individual fitness advantage if both predators and prey die during interactions? We examined the interaction between the rough-skinned newt (Taricha granulosa), which contains a powerful neurotoxin called tetrodotoxin (TTX), and the common garter snake (Thamnophis sirtalis). In some sympatric populations, Th. sirtalis have evolved physiological resistance to TTX. Whether the newts’ toxin confers protection from snake predators or has been disarmed by the snakes’ physiological resistance has not yet been directly tested. In predator–prey trials, newts that were rejected by snakes had greater concentrations of TTX in their skin (4.52 ± 3.49 mg TTX/g skin) than those that were eaten (1.72 ± 1.53 mg TTX/g skin). Despite the plethora of taxa that appear to use TTX defensively, this is the first direct and quantitative demonstration of the antipredator efficacy of TTX. Because the survival probability of a newt (and thus fitness) is affected by individual TTX concentration, selection can drive the escalation of toxin levels in newts. The variable fitness consequences associated with both TTX levels of newts and resistance to TTX in snakes that may promote a strong and symmetrical coevolutionary relationship have now been demonstrated.     

Predators usurp prey defenses? Toxicokinetics of tetrodotoxin in common garter snakes after consumption of rough-skinned newts

Snakes are common predators of organisms, such as amphibians, with toxic defenses that can be lethal to other predators. Because snakes do not have the option of dissecting prey into edible versus inedible components, they face a full dose of any chemical defenses encountered during attempted predation. This limitation has likely resulted in intense selection favoring the evolution of alternative mechanisms for dealing with prey toxins. These mechanisms can be physiological (e.g., resistance to prey toxins) or behavioral (e.g., toxin sampling and rejection). When physiological resistance arises, the possibility of bioaccumulation of a toxin results. We examined the coevolutionary interaction between the common garter snake (Thamnophis sirtalis) and the rough-skinned newt (Taricha granulosa), which contains a powerful neurotoxin called tetrodotoxin (TTX). In some populations syntopic with newts, individuals of T. sirtalis have evolved resistance to TTX. We examined the persistence of TTX in T. sirtalis after administration of an oral dose of TTX to investigate the possibility that snakes are sequestering TTX. The half-life of TTX in snake liver was estimated at 8.1?days. Accordingly, clearance of 99% of a single dose of TTX averages 61?days. Negative fitness consequences of intoxication during and after newt consumption may be balanced by co-opting the newts?? chemical defense for protection from the snakes?? own predators. Accounting of the coevolutionary dynamic between snakes and newts must incorporate post-consumption affects of lingering TTX.     

Distribution of Todarodes pacificus (Cephalopoda: Ommastrephidae) paralarvae near the Kuroshio off southern Kyushu, Japan

The distribution and abundance of Japanese common squid [Todarodes pacificus (Steenstrup, 1880)] paralarvae off southern Kyushu Island near the Kuroshio were examined in relation to water types in the region. Surveys were conducted in 1996 and 1997 using paired, 70 cm diameter Bongo nets. Temperature–salinity plots from 0, 50 and 100 m depths were used to assign sampling stations to three water types: inshore, mixing and Kuroshio. In total, 4103 T. pacificus paralarvae were collected at 59 of 72 stations. Catch densities at positive stations ranged between 8 and 4282 ind./50 m² sea surface area. Catches during both years were highest in mixing waters, followed by inshore and Kuroshio waters. Total catches were highest just north of the oceanographic front at the inshore edge of the Kuroshio and declined with increasing distance away from the front. Catches of paralarvae ≤1.0 mm mantle length also peaked near the front. In both years, ≥85% of all paralarvae and ≥69% of those ≤1.0 mm mantle length occurred less than 25 km from the front. We conclude that most spawning off southwest Kyushu occurs near the frontal zone, where Kuroshio and inshore waters meet. Received: 18 November 1998 / Accepted: 11 May 1999     

Death in the octopus’ garden: fatal blue-lined octopus envenomations of adult green sea turtles

The blue-lined octopus Hapalochlaena fasciata contains the powerful neuromuscular blocker tetrodotoxin (TTX), which causes muscle weakness and respiratory failure. H. fasciata is regarded as one of the most venomous marine animals in the world, and multiple human fatalities have been attributed to the octopus. To date, there have been no recorded incidents of an envenomation of a wild animal. Here, we present a newly developed, multi-stage tandem mass spectrometry technique that provides unequivocal evidence for two cases of envenomation of two ~110 kg herbivorous green sea turtles by two tiny cryptic blue-lined octopuses (~4 cm body length). These cases of accidental ingestion provide evidence for the first time of the antipredator effect of TTX and highlight a previously unconsidered threat to turtles grazing within seagrass beds.     

Different antipredator behaviour in two anuran tadpoles: effects of predator diet

Recent investigations have indicated that animals are able to use chemical cues of predators to assess the magnitude of predation risk. One possible source of such cues is predator diet. Chemical cues may also be important in the development of antipredator behaviour, especially in animals that possess chemical alarm substances. Tadpoles of the common toad (Bufo bufo) are unpalatable to most vertebrate predators and have an alarm substance. Tadpoles of the common frog (Rana temporaria) lack both these characters. We experimentally studied how predator diet, previous experience of predators and body size affect antipredator behaviour in these two tadpole species. Late-instar larvae of the dragonfly Aeshna juncea were used as predators. The dragonfly larvae were fed a diet exclusively of insects, R. temporaria tadpoles or B. bufo tadpoles. R. temporaria tadpoles modified their behaviour according to the perceived predation risk. Depending on predator diet, the tadpoles responded with weak antipredatory behaviour (triggered by insect-fed predators) or strong behaviour (triggered by tadpole-fed predators) with distinct spatial avoidance and lowered activity level. The behaviour of B. bufo in predator diet treatments was indistinguishable from that in the control treatment. This lack of antipredator behaviour is probably related to the effective post-encounter defenses and more intense competitive regime experienced by B. bufo. The behaviour of both tadpole species was dependent on body size, but this was not related to predator treatments. Our results also indicate that antipredator behaviour is largely innate in tadpoles of both species and is not modified by a brief exposure to predators. Received: 22 August 1996 / Accepted after revision: 31 January 1997     

Vertical distribution of <Emphasis Type="Italic">Todarodes pacificus</Emphasis> (Cephalopoda: Ommastrephidae) paralarvae near the Oki Islands,southwestern Sea of Japan

The diel vertical distribution patterns of Japanese common squid, Todarodes pacificus, paralarvae were examined using a Multiple Opening Closing Net and Environmental Sensing System (MOCNESS) in the southwest Sea of Japan near the Oki Islands (Japan) during five late-autumn surveys in 1998–2002. A total of 1,511 paralarvae ranging in mantle length (ML) from 0.7 to 7.3 mm were collected at 63 of the 68 stations surveyed. Most (84%) were collected above 75 m depth and in the mixed layer. The vertical distribution patterns varied little between day and night. Hatchling-sized (<1.0 mm ML) paralarvae were abundant at 0–25 m depth, and paralarval ML increased with increasing sampling depth. Our results suggest that T. pacificus paralarvae do not exhibit large diel vertical migration patterns, but as they increase in size, paralarvae gradually descend in the water column and the variability in depth increases with ontogeny.     

Intraguild predation by <Emphasis Type="Italic">Harmonia axyridis</Emphasis> on coccinellids revealed by exogenous alkaloid sequestration

Summary.  Under laboratory conditions, the multicolored Asian lady beetle, Harmonia axyridis is well known as an intraguild predator of other ladybirds. However the real impact of this exotic species on native species was poorly investigated in the field. Because many ladybird species produce alkaloids as defensive compounds, we propose here a new method of intraguild predation monitoring in coccinellids based on alkaloid quantification by GC-MS. In laboratory experiments, adaline was unambiguously detected in fourth instar larvae of H. axyridis having ingested one egg or one first instar larva of Adalia bipunctata. Although prey alkaloids in the predator decreased with time, traces were still detected in pupae, exuviae and imagines of H. axyridis having ingested one prey when they were fourth instar larvae. Analysis of H. axyridis larvae collected in two potato fields shows for the first time in Europe the presence of exogenous alkaloids in 9 out of 28 individuals tested. This new method of intraguild predation detection could be used more widely to follow the interactions between predators and potential chemically defended insect preys.     

Improving escape responses of hatchery-reared scallops <Emphasis Type="Italic">Argopecten purpuratus</Emphasis>

Hatchery rearing of the scallop Argopecten purpuratus has resulted in successive generations of scallops not exposed to predators that are less sensitive to and escape more slowly from predators than wild scallops. The present study examined whether conditioning hatchery-reared A. purpuratus to its natural predator, the sea star Meyenaster gelatinosus, improved its escape responses. Both juvenile and adult A. purpuratus from Tongoy Bay, Chile, were exposed for 7 days to different conditions: (1) continuous predator odor, (2) predator contact for 30 min three times a day, (3) a combination of the two previous conditions, and (4) no exposure to the predator (control). After conditioning, we evaluated scallop’s escape responses: reaction time, total clap number, duration of the clapping response, clapping rate, and the time scallops spent closed when exhausted. Conditioning with contact and odor plus contact (i.e., high predation risk) resulted in 25 and 50% shorter reaction times of juveniles and adults, respectively. Further, these stimuli caused juveniles to increase the number of claps and clapping rate. For adults, the time spent closed after exhaustion decreased by 50 and 63% after conditioning with contact and odor plus contact, respectively. Therefore, it is shown for the first time that exposure of scallops to increasing predator stimuli enhances escape responses, evidence of threat-sensitive predator avoidance.     

Occurrence of tetrodotoxin in the gastropodsRapana rapiformis andR. venosa venosa

Paralytic toxicity was detected in four of 17 specimens and six of 28 specimens of the gastropodsRapana rapiformis andR. venosa venosa, respectively, collected from Chiching (Kaohsiung City) and Nanfangao (Ilan County), Taiwan, in 1988 and 1989. The highest toxicities, calculated as tetrodotoxin (TTX) content, were 140 and 13 mouse units (MU) g^–1 digestive gland inR. rapiformis andR. venosa venosa, respectively. The toxins obtained from each species were purified by ultrafiltration and Bio-Gel P-2 column chromatography. The toxin's specific toxicities (as TTX) were 56 MU mg^–1 (R. rapiformis) and 52 MU mg^–1 (R. venosa venosa). Results of analyses by thin-layer chromatography, cellulose acetate membrane electrophoresis and high-performance liquid chromatography showed that TTX and anhydrotetrodotoxin were responsible for the toxicity. The alkali hydrolysate of each toxin showed maximum absorption at ca. 274 nm, which is the unique absorption for the C₉-base of TTX and its related substances (such as anhydrotetrodotoxin, 4-epitetrodotoxin, etc.). Here, we report for the first time the occurrence of TTX in the gastropodsR. rapiformis andR. venosa venosa.     

Evaluating the impact of caribou habitat restoration on predator and prey movement

Fragmentation of the boreal forest by linear features, including seismic lines, has destabilized predator–prey dynamics, resulting in the decline of woodland caribou (Rangifer tarandus caribou) populations. Restoration of human-altered habitat has therefore been identified as a critical management tool for achieving self-sustaining woodland caribou populations. However, only recently has testing of the response of caribou and other wildlife to restoration activities been conducted. Early work has centered around assessing changes in wildlife use of restored seismic lines. We evaluated whether restoration reduces the movement rates of predators and their associated prey, which is expected to decrease predator hunting efficiency and ultimately reduce caribou mortality. We developed a new method for using cameras to measure fine-scale movement by measuring speed as animals traveled between cameras in an array. We used our method to quantify speed of caribou, moose (Alces alces), bears (Ursus americanus), and wolves (Canis lupus) on treated (restored) and untreated seismic lines. Restoration treatments reduced travel speeds along seismic lines of wolves by 1.38 km/h, bears by 0.55 km/h, and caribou by 1.57 km/h, but did not reduce moose travel speeds. Reduced predator and caribou speeds on treated seismic lines are predicted to decrease encounter rates between predators and caribou and thus lower caribou kill rates. However, further work is needed to determine whether reduced movement rates result in reduced encounter rates with prey, and ultimately reduced caribou mortality.     

The interplay between foraging mode, habitat structure, and predator presence in antlions

Antlion larvae are sand-dwelling insect predators, which ambush small arthropod prey while buried in the sand. In some species, the larvae construct conical pits and are considered as sit-and-wait predators which seldom relocate while in other species, they ambush prey without a pit but change their ambush site much more frequently (i.e., sit-and-pursue predators). The ability of antlion larvae to evade some of their predators which hunt them on the sand surface is strongly constrained by the degree of sand stabilization or by sand depth. We studied the effect of predator presence, predator type (active predatory beetle vs. sit-and-pursue wolf spider), and sand depth (shallow vs. deep sand) on the behavioral response of the pit building Myrmeleon hyalinus larvae and the sit-and-pursue Lopezus fedtschenkoi larvae. Predator presence had a negative effect on both antlion species activity. The sit-and-wait M. hyalinus larvae showed reduced pit-building activity, whereas the sit-and-pursue L. fedtschenkoi larvae decreased relocation activity. The proportion of relocating M. hyalinus was negatively affected by sand depth, whereas L. fedtschenkoi was negatively affected also by the predator type. Specifically, the proportion of individual L. fedtschenkoi that relocated in deeper sand was lower when facing the active predator rather than the sit-and-pursue predator. The proportion of M. hyalinus which constructed pits decreased in the presence of a predator, but this pattern was stronger when exposed to the active predator. We suggest that these differences between the two antlion species are strongly linked to their distinct foraging modes and to the foraging mode of their predators. Reut Loria and Inon Scharf contributed equally to the paper.     

Threat-sensitive learning of predators by larval mosquitoes <Emphasis Type="Italic">Culex restuans</Emphasis>

A prerequisite for prey to show adaptive behavioural responses to predators is that the prey has the ability to recognise predators as threats. While predator recognition can be innate in many situations, learning is often essential. For many aquatic species, one common way to learn about predators is through the pairing of a novel predator odour with alarm cues released from injured conspecifics. One study with fish demonstrated that this mode of learning not only allows the prey to recognise the predatory cues as a threat, but also mediates the level of threat associated with the predator cues (i.e. threat-sensitive learning). When the prey is exposed to the novel predator with a high concentration of alarm cues, they subsequently show a high intensity of antipredator response to the predator cues alone. When exposed to the predator with a low concentration of alarm cues, they subsequently show a low-intensity response to the predator cues. Here, we investigated whether larval mosquitoes Culex restuans have the ability to learn to recognise salamanders as a threat through a single pairing of alarm cues and salamander odour and also whether they would learn to respond to salamander cues in a threat-sensitive manner. We conditioned individual mosquitoes with water or a low, medium or high concentration of crushed conspecific cues (alarm cues) paired with salamander odour. Mosquitoes exposed to salamander odour paired with alarm cues and subsequently exposed to salamander odour alone responded to the salamander as a threat. Moreover, the intensity of antipredator response displayed during the conditioning phase matched the response intensity during the testing phase. This is the first demonstration of threat-sensitive learning in an aquatic invertebrate.     

Induction of anti-predator responses in the green-lipped mussel Perna viridis under hypoxia

Hypoxia due to the over enrichment of waters by nutrients is becoming a global problem. In mussels, enhanced byssus thread production is an important adaptation to the presence of crustacean predators and to energetic hydrodynamic regimes. Thread production is an energy-consuming process, so this study used the green mussel Perna viridis (L.) to examine the response to predator exposure combined with hypoxia. Hypoxia is common in sheltered bays in Hong Kong, and the mussels were collected in one such bay, Lok Wo Sha (latitude/longitude: 22o18′ N/114o10′ E) in January, 2009. The predator used in the experiments was the swimming crab Thalamita danae. Oxygen concentrations used in the 48-h experiments ranged from hypoxic to normoxic (1.5 ± 0.3 mg l⁻¹, 3.0 ± 0.3 mg l⁻¹ and 6.0 ± 0.3 mg l⁻¹). Fewer byssus threads which were also shorter and thinner were produced at reduced oxygen levels, no matter if the predator was present or not; the frequency the mussels shed stalks was also lower. Mussels exposed to the predator, however, have enhanced byssus thread production at all oxygen levels when compared with the control. This has highlighted the significance of anti-predator responses for the survival of individuals even under a stressful environment in which energy supply is limited by aerobic metabolism. Interactive effects between oxygen level and predator exposure were observed for the byssus thread production (frequency of shed stalks, mean byssus thread length, cumulative byssus thread volume), with values obtained at 1.5 and 3.0 mg O₂ l⁻¹ being statistically indistinguishable for the control group without predator but not for the predator group. The lack of differences in the byssus thread production at lower oxygen levels in the absence of predator may indicate the minimum amount of byssus that is required for settlement on a substrate.     

Predator inspection behaviour in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>): body size,local predation pressure and cooperation

The apparently maladaptive tendency of fish to approach and inspect potential predators has been explained in terms of useful information gathering or as a signal to the predator that it has been seen. We examined this behaviour in 16 populations of wild-caught stickleback (Gasterosteus aculeatus) from ponds with and without predatory perch (Perca fluviatilis). Three large and three small individuals per population were each exposed to three model predators differing in realism. A final cooperative treatment entailed pairing subjects with a second individual from the same population, but of the alternative size class, during predator presentation. As might be expected, predator inspection behaviour was much greater in the predator-sympatric populations, and only these fish increased their level of inspection as the models became incrementally more realistic. This suggests that reductions occur in the level of costly inspection behaviour in populations without predators. Subject body size had no effect on inspection effort, which suggests a limited role for experience (we assumed larger fish to be older than smaller fish), at least over the relative age differences utilized. However, small predator-sympatric fish were the only subjects to increase inspection significantly when in a cooperative context, perhaps reflecting the inherent value of a relatively larger partner in this context. These results confirm that levels of predator inspection are both population- and situation-dependent, suggesting a trade-off in the potential costs and benefits of this behaviour.Communicated by C. St. Mary     

Inspection of mob-calls as sources of predator information: response of migrant and resident birds in the Neotropics

Migrating animals face numerous mortality risks, such as novel predators with which they may not be accustomed. Most animals can recognize predators innately; however, additional predator information can be collected to enhance familiarity. Because migrating birds rarely participate in mobs, they may seek alternative information sources such as cues provided by other birds that can provide information on predator location, identity, and degree of threat. We predicted that Nearctic–Neotropical migrants (hereafter, “migrants”) would react to vocal antipredator cues (e.g., mob-calls) of species residing in areas through which they migrate. To test this, we conducted experiments in Belize during spring migration, using playbacks of mob-calls of black-capped chickadees (Poecile atricapillus) and blue-gray tanagers (Thraupis episcopus); tanagers are familiar to all birds in Belize; chickadees are novel to residents but familiar to migrants. This also allowed us to assess response to novel and out-of-context antipredator signals. Resident birds did not respond to novel chickadee mob-calls, but did respond to familiar tanager calls. Birds overwintering south of our study area, which were migrating during our study, responded most strongly to chickadee playbacks. Conversely, individuals of species that include our study area in their winter range did not respond to either playback. This is the first evidence that birds react to vocal antipredator cues during migration, which may be a strategy used by migrants to learn about predators. Although residents failed to recognize a foreign cue, migrating birds responded most strongly to the out-of-context chickadee cue, associated with breeding grounds >2,000 km northward. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Local interactions between predators and prey call into question commonly used functional responses

Commonly used functional response models (Holling’s type I and type II models) assume that the encounter rate of a predator increases linearly with prey density, provided that the predator is searching for prey. In other other words, aN (a is the baseline encounter rate and N is prey density) describes the encounter rate. This study examined whether the models are adequate when predators and prey interact locally by using a spatially explicit individual based model because local interactions affect the spatial distribution of predators and prey, which also affects the encounter rate. Predators were assumed to possess a spatial perception range that influenced their foraging behavior (e.g., if a prey is in the perception range, the predator moves towards the prey). The effect of antipredator behavior by prey was also examined. The results suggest that prey and predator densities as well as handling time affect the baseline rate (i.e., parameter a) as opposed to the common assumption that the parameter is constant. The nature of model deviations depended on both the antipredator behavior and the predators’ perception range. Understanding these deviations is important as they qualitatively affect community dynamics.     

Interaction and impacts of two introduced species on a soft-sediment marine assemblage in SE Tasmania

Introduced species are having major impacts in terrestrial, freshwater and marine ecosystems world-wide. It is increasingly recognised that effects of multiple species often cannot be predicted from the effect of each species alone, due to complex interactions, but most investigations of invasion impacts have examined only one non-native species at a time and have not addressed the interactive effects of multiple species. We conducted a field experiment to compare the individual and combined effects of two introduced marine predators, the northern Pacific seastar Asterias amurensis and the European green crab Carcinus maenas, on a soft-sediment invertebrate assemblage in Tasmania. Spatial overlap in the distribution of these invaders is just beginning in Tasmania, and appears imminent as their respective ranges expand, suggesting a strong overlap in food resources will result from the shared proclivity for bivalve prey. A. amurensis and C. maenas provide good models to test the interaction between multiple introduced predators, because they leave clear predator-specific traces of their predatory activity for a number of common prey taxa (bivalves and gastropods). Our experiments demonstrate that both predators had a major effect on the abundance of bivalves, reducing populations of the commercial bivalves Fulvia tenuicostata and Katelysia rhytiphora. The interaction between C. maenas and A. amurensis appears to be one of resource competition, resulting in partitioning of bivalves according to size between predators, with A. amurensis consuming the large and C. maenas the small bivalves. At a large spatial scale, we predict that the combined effect on bivalves may be greater than that due to each predator alone simply because their combined distribution is likely to cover a broader range of habitats. At a smaller scale, in the shallow subtidal, where spatial overlap is expected to be most extensive, our results indicate the individual effects of each predator are likely to be modified in the presence of the other as densities increase. These results further highlight the need to consider the interactive effects of introduced species, especially with continued increases in the number of established invasions.Communicated by M.S. Johnson, Crawley     

Linking predator and prey behaviour: contrasts between Antarctic fur seals and macaroni penguins at South Georgia

Antarctic fur seals Arctocephalus gazella and macaroni penguins Eudyptes chrysolophus are the two main land-based krill Euphausia superba consumers in the northern Scotia Sea. Using a combination of concurrent at-sea (predator observations, net hauls and multi-frequency acoustics), and land-based (animal tracking and diet analysis) techniques, we examined variability in the foraging ecology of these sympatric top predators during the austral summer and autumn of 2004. Krill availability derived from acoustic surveys was low during summer, increasing in autumn. During the breeding season, krill occurred in 80% of fur seal diet samples, with fish remains in 37% of samples. Penguin diets contained the highest proportion of fish in over 20 years of routine monitoring (46% by mass; particularly the myctophid Electrona antarctica), with krill (33%) and amphipods (Themisto gaudichaudii; 21%) also occurring. When constrained by the need to return and feed their offspring both predator species foraged to the northwest of South Georgia, consistent with an area of high macrozooplankton biomass, but fur seals were apparently more successful at exploiting krill. When unconstrained by chick-rearing (during March) penguins foraged close to the Shag Rocks shelf-break, probably exploiting the high daytime biomass of fish in this area. Penguins and seals are able to respond differently to periods of reduced krill abundance (in terms of variability in diet and foraging behaviour), without detriment to the breeding success of either species. This highlights the importance of myctophid fish as an alternative trophic pathway for land-based predators in the Scotia Sea ecosystem.     

Reproduction under predation risk in the sand goby,Pomatoschistus minutes,and the black goby,Gobius niger: the effect of age and longevity

Summary In aquarium experiments, the two marine gobiid fish species Pomatoschistus minutus and Gobius niger were allowed to build nests and to spawn in the presence and absence of a predator (cod, Gadus morhua); behind a glass wall the predator was kept where it could be clearly seen by the gobies and vice versa. P. minutus showed no difference in number of nests or number of spawnings in the different treatments; approximately half of the males built nests, and the females spawned in half of those nests. G. niger, on the other hand, responded differently to the simulated predation risk. No nests were built in sight of the predator, whereas in the absence of predators, half of the males built nests and received eggs. The G. niger individuals in this experiment were 2–3 years old. However, when comparing the reproduction of G. niger of different age in the presence of a predator, older individuals (4–5 years) spawned, whereas younger (2–3 years) did not. No difference in vulnerability towards predators was found between equal-sized P. minutus and G. niger. The optimal behavior during the breeding season must depend on prospects of survival, based both on maximal lifespan and vulnerability to predation.