Look before you leap: is risk of injury a foraging cost?

Theory states that an optimal forager should exploit a patch so long as its harvest rate of resources from the patch exceeds its energetic, predation, and missed opportunity costs for foraging. However, for many foragers, predation is not the only source of danger they face while foraging. Foragers also face the risk of injuring themselves. To test whether risk of injury gives rise to a foraging cost, we offered red foxes pairs of depletable resource patches in which they experienced diminishing returns. The resource patches were identical in all respects, save for the risk of injury. In response, the foxes exploited the safe patches more intensively. They foraged for a longer time and also removed more food (i.e., had lower giving up densities) in the safe patches compared to the risky patches. Although they never sustained injury, video footage revealed that the foxes used greater care while foraging from the risky patches and removed food at a slower rate. Furthermore, an increase in their hunger state led foxes to allocate more time to foraging from the risky patches, thereby exposing themselves to higher risks. Our results suggest that foxes treat risk of injury as a foraging cost and use time allocation and daring—the willingness to risk injury—as tools for managing their risk of injury while foraging. This is the first study, to our knowledge, which explicitly tests and shows that risk of injury is indeed a foraging cost. While nearly all foragers may face an injury cost of foraging, we suggest that this cost will be largest and most important for predators.     

A test of the mismatch hypothesis: How is timing of reproduction related to food abundance in an aerial insectivore?

In seasonal environments, vertebrates are generally thought to time their reproduction so offspring are raised during the peak of food abundance. The mismatch hypothesis predicts that reproductive success is maximized when animals synchronize their reproduction with the food supply. Understanding the mechanisms influencing the timing of reproduction has taken on new urgency as climate change is altering environmental conditions during reproduction, and there is concern that species will not be able to synchronize their reproduction with changing food supplies. Using data from five sites over 24 years (37 site-years), we tested the assumptions of the mismatch hypothesis in the Tree Swallow (Tachycineta bicolor), a widespread aerial insectivore, whose timing of egg-laying has shifted earlier by nine days since the 1950s. Contrary to the mismatch hypothesis, the start of egg-laying was strongly related to food abundance (flying insect biomass) during the laying period and not to timing of the seasonal peak in food supply. In fact, food abundance generally continued to increase throughout the breeding season, and there was no evidence of selection based on the mistiming of laying with the seasonal peak of food abundance. In contrast, there was selection for laying earlier, because birds that lay earlier generally have larger clutches and fledge more young. Overall, initial reproductive decisions in this insectivore appear to be based on the food supply during egg formation and not the nestling period. Thus, the mismatch hypothesis may not apply in environments with relatively constant or abundant food throughout the breeding season. Although climate change is often associated with earlier reproduction, our results caution that it is not necessarily driven by selection for synchronized reproduction.     

Sex ratio of juvenile loggerhead turtles in the Mediterranean Sea: is it really 1:1?

Sex ratios are a crucial parameter for evaluating population viability. In species with complex life history patterns and temperature sex determination mechanisms, such as the loggerhead turtle (Caretta caretta), sex ratios may vary within a population and among populations. In the Mediterranean, juvenile sex ratios appear to not differ significantly from 1:1, although estimates for hatchling sex ratios are highly female biased. The immigration of males from the Atlantic has been suggested as a possible cause of such variation. Here, we present results of a multi-year investigation (2000–2011) on the sex ratios of loggerhead turtles foraging along the south Tyrrhenian coast, Western Mediterranean, with the aim of providing a better understanding of the potentially underlying forces that drive regional and age-dependent differences in sex ratios. Sex was determined through visual examination of the gonads in 271 dead turtles (curved carapace length range 29.5–89 cm). A fragment of the mitochondrial DNA control region was sequenced from 61 specimens to characterise the demographic composition of this foraging assemblage by applying a many-to-many mixed stock analysis approach. No significant association was found between sex ratios and years or size classes, although the largest size was male biased. Juvenile sex ratio was 1.56:1, which was different from an even sex ratio but still less female biased than hatchling sex ratios from Mediterranean beaches. Results of the mixed stock analysis indicate that juvenile sex ratios in the Mediterranean are largely unaffected by immigration of Atlantic individuals into the basin, as previously suggested. Continued long-term monitoring of juvenile sex ratios is necessary to detect biologically significant sex ratio shifts in the Mediterranean loggerhead turtle population.     

Female foraging and male vigilance in white-tailed ptarmigan (Lagopus leucurus): opportunism or behavioural coordination?

For several species of birds, high rates of male vigilance are correlated with high rates of female foraging. This relationship is thought to ultimately result in higher reproductive success for females paired with highly vigilant males. However, previous research has not examined the behavioural mechanism that produces the correlation between male vigilance and rates of female foraging. Foraging females may take advantage of vigilance that males are using for other purposes. Alternatively, the purpose of male vigilance may be to increase females' ability to forage. We examined these alternatives by testing whether vigilance preceded or followed bouts of female foraging more often than would occur by chance alone, using simultaneous behaviour observations of pre-incubation pairs of white-tailed ptarmigan (Lagopus leucurus). Our results indicate that each member of a pair may influence the behaviour of the other. Females were more likely to initiate foraging bouts after males became vigilant than if their mate remained non-vigilant. Moreover, non-vigilant males were more likely to become vigilant if their mate was foraging than if she was engaged in some other activity. Despite the possibility that a sexual conflict exists as each member of a pair attempts to maximize its fitness, both sexes behave as though a major role of male vigilance is to enhance female foraging opportunities. Received: 3 May 1999 / Received in revised form: 14 June 1999 / Accepted: 16 June 1999     

Does interaction between bumblebees (Bombus ignitus) reduce their foraging area?: bee-removal experiments in a net cage

To examine whether the interaction between bumblebees, Bombus ignitus, reduces their foraging area, we conducted bee-removal experiments in a net cage. In the cage, we set potted Salvia farinacea plants, allowed bumblebees to forage freely on those plants, and followed their plant-to-plant movements to identify a bee with a relatively small foraging area. We then removed all the other foraging bees, except for the bee with a small foraging area, and observed the change of the foraging area of the focal bee under conditions of no interaction with other bees. After the removal of the other bees, all five bees tested enlarged their foraging areas, suggesting that the interaction between bees is an important determinant of their foraging areas. The result also means that bumblebees are able to adjust their foraging areas in response to other foragers, indicating the necessity for future studies to clarify what cues bees use to interact with other bees. Moreover, after the removal treatments, all five bees showed temporary increases in the number of flower probes per plant. This can be explained by their optimal foraging according to the old average intake rate for the plant population and by the delayed changes in response to the new high average energy intake rate after the bee-removal treatments.Communicated by M. Giurfa     

Vulnerability of newly settled red drum (Sciaenops ocellatus) to predatory fish: is early-life survival enhanced by seagrass meadows?

We conducted predation experiments to evaluate the vulnerability of red drum Sciaenops ocellatus (Linnaeus) larvae and early juveniles to pinfish Lagodon rhomboides (Linnaeus) predators. Experiments were designed to analyze the effects of habitat complexity, prey size, and rearing condition on prey vulnerability. Three structurally different habitats [unvegetated (substrate only), shoal grass Halodule wrightii (Aschers) and turtle grass Thalassia testudinum (Bank ex König)] were simulated in experimental mesocosms. Instantaneous hourly mortality rates (Z?h^?1 predator^?1) for hatchery-reared red drum were significantly higher in the unvegetated habitat (0.189) than in either shoal grass (0.069) or turtle grass (0.046). A similar trend in predation mortality was observed for wild-caught red drum; instantaneous hourly mortality rates were 0.166, 0.047, and 0.021 in unvegetated, shoal grass, and turtle grass habitats, respectively. Mortality rates (adjusted means) for hatchery red drum were higher than for wild individuals in all three habitats; however, the differences were not significant. Predation mortality decreased with increasing prey size (3 to 9% decrease in Z per mm increase in length), suggesting that small red drum (i.e. new settlers) were most vulnerable to predators. We conclude that habitat complexity is critical to the survival of newly settled red drum, and changes in the complexity or areal coverage of natural seagrass meadows may affect early-life survival and possibly recruitment levels.     

Biofilm cue for larval settlement in Hydroides elegans (Polychaeta): is contact necessary?

Larvae of many sessile marine invertebrates settle in response to surface microbial communities (biofilms), but the effects of soluble compounds from biofilms in affecting larval behavior prior to settlement, attachment, and metamorphosis have been little studied. This question was addressed by videotaping the behavior of competent larvae of the serpulid, Hydroides elegans, above settlement-inducing biofilms. Adult worms were collected in Pearl Harbor, Hawaii, USA in November 2012 and spawned almost immediately. Six-day old larvae were placed in five replicated treatments in small cups: (1) with a natural biofilm; (2) with a natural biofilm on an 8-µm screen, 1 mm above the bottom of a clean cup; (3) with a natural biofilm beneath a clean screen; (4) in a clean cup; and (5) in a clean cup with a clean screen. Using the videotapes, larval swimming speeds and trajectories were quantified within 5 min of the larvae being placed in a treatment. Only larvae that touched a biofilm, i.e., in treatments (1) and (2), slowed their swimming speed and increased the amount of time spent crawling rather than swimming. This shows that under these conditions, any soluble cues emanating from a biofilm do not affect settlement behavior. Furthermore, after 24 h close to 100 % of larva in the two accessible biofilm treatments had metamorphosed and <15 % in treatments that included a biofilm under a clean screen and no biofilm at all, strongly suggesting that soluble cues for settlement were not produced by the biofilms over the longer time period.     

Differential timing of spring migration in northern wheatears Oenanthe oenanthe: hurried males or weak females?

In a field experiment on the island of Helgoland (southeast North Sea), we investigated whether migration strategy or competition between the sexes cause the differential timing of spring migration of male and female northern wheatears (Oenanthe oenanthe) (males migrating earlier). The study included two subspecies, heading towards Greenland/Iceland and Scandinavia, respectively, and is based on colour-ringing and remote weighing of individuals. Despite food offered ad libitum, most Scandinavian birds left the island on the day of arrival or stayed only 1–3 days, whereas more than half of Greenlandic/Icelandic birds stayed for up to 12 days and refuelled rapidly. In the latter subspecies, males showed a positive correlation of departure fuel load and fuel deposition rate, resembling time-minimizers in optimal migration theory. In contrast, females departed irrespective of fuel deposition rate, with an approximately constant level of fuel stores. This level was lower than in males, but sufficient to enable by-passing of stopover sites en route, allowing us to regard females as time-minimizers also. Since females are not able to reach Greenland without additional refuelling elsewhere and males appeared to have a larger potential for by-passing stopover sites, time-selection seems to be more pronounced in males and may be the reason for earlier migration of males. Intraspecific aggressive interactions between colour-ringed birds were predominantly won by the initiator, by males and by larger birds, whereas fuel load and subspecies did not affect the outcome. Although compared to females, males were more often dominant at the feeding stations or held territories, refuelling patterns could not be explained by dominance. Subordinate or non-territorial birds did not refuel at a lower rate or depart with lower fuel loads than dominant or territorial birds. In non-territorial birds, the restricted access to feeding stations was made up with larger doses of food taken per visit, leading to the same energy intake as that of dominant and territorial birds. Therefore, competition during stopover could be eliminated as the reason for differential timing of migration of male and female wheatears, but this result may be species-specific.Communicated by W. Wiltschko     

Changes in the timing of otolith zone formation in North Sea cod from otolith records: an early indicator of climate-induced temperature stress?

We examine the seasonal variation in otolith increment formation in southern North Sea cod as a means of monitoring how changes in sea temperature over the past 20 years have affected cod in the wild. Seasonal opaque zone formation was related to winter and early spring. Timing of opaque zone formation was not influenced by either temperature or fish length, but increasing age led to slightly earlier but slower opaque zone formation. In contrast, there was a clear shift in the timing of translucent zone formation with temperature. In warm years, translucent growth occurs up to 22 days earlier than in colder periods. Increasing age and smaller size-at-age resulted in an earlier transition from opaque to translucent edge formation. Translucent zone formation appears indicative of increasing metabolic stress, and the earlier onset provides direct evidence of the impact of increasing sea temperatures on wild North Sea cod stocks.     

Sexual size dimorphism in fallow deer (Dama dama): do larger, heavier males gain greater mating success?

Sexual size dimorphism may evolve as a result of both natural and sexual selection. In polygynous mammals, the main factor resulting in the evolution of large body size in males is the advantage conferred during competition for mates. In this study, we examined whether sexual selection acts on body size in mature fallow bucks (Dama dama) by examining how the following traits are inter-related: age, body (skeletal) size, body mass, prerut dominance rank, rut dominance rank and mating success. This is the first study to examine how all these factors are together related to the mating success of a large sexually dimorphic and polygynous mammal. We found that male mating success was directly related to body size, but not to body mass. However body mass was related to prerut dominance rank which was in turn strongly related to rut dominance rank, and thus there was an indirect relationship between mating success and body mass. Rut dominance rank was the variable most strongly related to mating success. Mating success among mature males was unrelated to age. We conclude that larger mature fallow bucks have advantages over other males when competing for matings, and sexual selection therefore continues to act on sexual size dimorphism in this species. Heavier fallow bucks also have advantages, but these are mediated through the dominance ranks attained by males before the rut.     

In-depth studies of Magellanic penguin (<Emphasis Type="Italic">Spheniscus magellanicus</Emphasis>) foraging: can we estimate prey consumption by perturbations in the dive profile?

A new concept based on analysis of dive depth data was developed to help estimate prey consumption in ten free-ranging Magellanic penguins (Spheniscus magellanicus) that were brooding chicks. By simultaneously analysing the undulations in the dive depth profile (measured by time-depth recorders, TDRs) and beak opening (obtained from the recently developed intra-mandibular angle sensors, IMASEN), it was possible to determine the proportions of the undulations in the dive profile that resulted (or not) in prey capture. This methodology allowed the number of prey consumed to be estimated with a mean error of 10±6% using TDR data alone. If the mean mass of prey is known, then the overall mass of prey consumed per unit time can be determined. Additionally, the method allows estimation of the depth at which prey is taken and thus indicates how penguins exploit the water column. Due to its simplicity, the proposed methodology has applications for other Spheniscus penguin species and should be considered for other marine endotherm divers that show undulations in the dive depth profile.Communicated by O. Kinne, Oldendorf/Luhe     

Shell-boring versus tube-dwelling: is the mode of life fixed or flexible? Two cases in spionid polychaetes (Annelida, Spionidae)

Free crawling was an original life style, and the ability to build tubes and bore into shells evolved independently in various annelids and more than once within the family Spionidae. Absence of morphological differences between the shell-boring (SB) and tube-dwelling (TD) spionid worms and the innate ability of borers to build tubes raised a question whether the mode of life is fixed or flexible. Sequence data of three gene fragments of the mitochondrial 16S rDNA and nuclear 18S and 28S rDNA (in total 1,677 bp) have shown that SB and TD individuals of Dipolydora carunculata from the Sea of Japan were genetically identical, whereas those of Polydora triglanda from Taiwan were genetically distant and not conspecific. These data indicate that some species are constrained to a certain mode of life, while others are flexible and individuals can be either SB or TD depending on the place of settlement in the end of their larval development.     

The foraging ecology of Arctic cod (Boreogadus saida) during open water (July–August) in Allen Bay, Arctic Canada

Arctic cod (Boreogadus saida) is a schooling fish providing a critical link between lower and upper trophic levels in the Arctic. This study examined foraging of Arctic cod collected from Allen Bay, Cornwallis Island, Canada (~75 N 95 W), during summer 2010 using temporal indicators of diet including stomach content, and carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotopes of liver and muscle. Foraging at the time of capture reflected sympagic and epi-benthic habitats indicated by the prevalence of cyclopoid and harpacticoid copepods in stomachs, whereas stable isotope data, which provide an estimate of feeding over a longer period, indicated pelagic prey as important. Prey selection of juveniles differed from adults based on stable isotopes, while large adults showed the most separation based on stomach contents, suggesting size-related diet shifts. Compared to studies near Resolute in the 1970s, 1980s, and 1990s, growth and pre-spawning gonadal conditions of Arctic cod have not changed.     

Does cannibalism in Spodoptera frugiperda (Lepidoptera: Noctuidae) reduce the risk of predation?

The incidence of cannibalism of larval Spodoptera frugiperda (Lepidoptera: Noctuidae) on maize under field conditions was investigated using field cages. Cannibalism was found to account for approximately 40% mortality when maize plants were infested with two or four fourth-instar larvae over a 3-day period. Field trials examined the effect of larval density on the prevalence of natural enemies of S. frugiperda. The abundance of predators (earwigs, staphylinids, other predatory beetles, and Chrysoperla spp.) was significantly greater on maize plants with higher levels of larval feeding damage, while the relationship between predator abundance and number of S. frugiperda larvae per plant was less clear. As larval damage is probably a more reliable indicator of previous larval density than numbers collected at an evaluation, this indicates that predation risk will be greater for larvae living in large groups. Parasitism accounted for 7.1% mortality of larvae in sorghum, and involved six species of Hymenoptera and Tachinidae. There was no effect of larval density or within-plant distribution on the probability of larval attack by parasitoids. The selective benefits of cannibalism, in relation to the risk of predation and parasitism, are discussed. Received: 23 March 2000 / Accepted: 24 June 2000     

Short retention times of stomach temperature loggers in free-living seabirds: is there hope in the spring?

Stomach temperature loggers have proved useful for the study of feeding activity in free-living seabirds, but their usage has been restricted because they are frequently dislodged and lost during the seabirds' regular regurgitation of indigestible prey remains. In the present study we examine the incidence of spontaneous regurgitation (pellet production) in free-living seabirds, consider the effect this has on the likely retrieval of stomach temperature loggers and present a structural modification of the logger housings which leads to a much lower incidence of regurgitation. Systems were tested on albatrosses, cormorants, gannets and penguins. Received: 15 July 1997 / Accepted: 31 October 1997     

Who goes first? Condition and danger dependent pioneering in a group-living bark beetle (Dendroctonus ponderosae)

Among group-living organisms, some individuals initiate groups by being the first to attack a prey item or the first to colonize a new settlement site. In the group-living mountain pine beetle (Dendroctonus ponderosae), first attackers (known as pioneers) on live trees suffer higher mortality due to tree defenses than do beetles that join aggregations. This study examined factors that affect an individual’s propensity to initiate an aggregation. When placed on an unoccupied tree, the probability of successfully entering the tree was positively correlated with body condition (residual of mass versus length regression). However, beetles in better condition took longer to initiate tunnel construction than those in poorer condition, suggesting that pioneering is a “desperation” strategy used when low energy reserves preclude further dispersal or when potential trees are rare. These contrasting patterns suggest pioneering is a nonlinear behavioral response, such that beetles with the smallest energy reserves and beetles with the greatest energy reserves both avoid pioneering. We further found that pioneering was more likely when the environment favored success, such as in smaller diameter trees (which may have weaker defenses) and earlier in the season (when the probability of recruiting conspecifics is higher). Our results suggest that pioneers incorporate both internal and external variables in their decision to attack an uncolonized tree.     

Mitochondrial genomic divergence in coelacanths (Latimeria): slow rate of evolution or recent speciation?

Dating the divergence between the two known living species of coelacanths has remained a difficult issue because of the very ancient origin of this lineage of fish, which is more closely related to tetrapods than to other fishes. We sequenced the complete mitochondrial genome of a recently captured individual of the Indonesian coelacanth in order to solve this issue. Using an approach based on loglinear models, we studied the molecular divergence between the two species of coelacanths and three other pairs of species, one that has diverged recently (Pan) and two that have diverged more distantly in the past. The loglinear models showed that the divergence between the two species of coelacanths is not significantly different from the two species of Pan. A detailed gene by gene analysis of the patterns of nucleotide and amino acid substitutions between these two pairs of species further supports the similarity of these divergences. On the other hand, a molecular dating analysis suggested a much older origin of the two coelacanth species (between 20 and 30 million years ago). We discuss the potential reasons for this discrepancy. The analysis of new individuals of the Indonesian coelacanth will certainly help to solve this issue.     

Assessing the antipredatory defensive strategies of Caribbean non-scleractinian zoantharians (Cnidaria): is the sting the only thing?

The relative importance of chemical, nematocyst, and nutritional defenses was examined for 18 species of Caribbean sea anemones (actinarians), zoanthids, and mushroom polyps (corallimorpharians) from the Florida Keys and the Bahamas Islands, 2008–2010. Feeding assays were performed using the fish Thalassoma bifasciatum with artificial foods containing crude organic extracts of cnidarian tissues. A novel behavioral assay using brine shrimp nauplii was used to assess nematocyst defenses. The nutritional quality of cnidarian tissues was examined using bomb calorimetry and soluble protein assays. In general, actinarians invested in nematocyst defenses, zoanthids in either nematocyst or chemical defenses, and corallimorpharians lacked both, except for 1 of 3 species that was chemically defended. Relative to other coral reef invertebrates, cnidarian tissues had similar caloric values but lower soluble protein concentrations. Trade-offs between chemical and nematocyst defenses were observed for 65% of species, while habitat and behavior provided a likely explanation for undefended species.     

Determination of carazolol in pharmaceutical formulations by gas‐liquid‐chromatography (GLC)

A gas‐liquid‐chromatographic procedure for the quantitative determination of authentic carazolol and carazolol in tablets (Conducton^(R)?) has been described. Carazolol in tablets was extracted with absolute alcohol before injecting onto the gas Chromatograph. The concentration range adopted varied between 0.2 mg to 1.0 mg/ml of carazolol in alcohol solution. The results obtained were 97.6%±3.2 and 98.4%±2.6 for the stated and added amounts respectively.     

Spatial segregation of calving and nursing Risso’s dolphins (Grampus griseus) in the Azores,and its conservation implications

Calving can be a critical period for cetaceans. Areas providing security for vulnerable calves, and high food availability for lactating females can be critical habitats requiring specific conservation measures. Here, we test the hypothesis that calving and nursing habitat could be defined for Risso’s dolphins (Grampus griseus). We investigated the spatial and temporal preferences of this species around Pico Island, Azores, using data gathered from land-based surveys and dedicated at-sea observations between 2004 and 2007. We divided observed pods into three groups: (1) those with newborn and young calves, (2) those with older calves or juveniles and (3) those consisting only of adults or sub-adults. We analysed eco-geographical variables and incorporated them into a presence-only spatial distribution model to evaluate differences in habitat suitability among the groups. We identified 694 pods of Risso’s dolphins overall. On the 267 pods observed in the first and second groups, 136 calves or juveniles were identified, of which 22 were newborns. The peak of the calving season was between June and August. The pods with newborn calves were larger and closer to shore, whereas the other groups were more widely dispersed offshore. Our results support the definition of critical habitat areas for this species, but we suggest widening the geographical coverage for better mapping around the island and throughout the archipelago generally. Conservation measures can, nevertheless, be implemented immediately, in order to reduce human impacts on a vulnerable component of the Risso’s dolphin population