The effects of wing loading and gender on the escape flights of least sandpipers (<Emphasis Type="Italic">Calidris minutilla</Emphasis>) and western sandpipers (<Emphasis Type="Italic">Calidris mauri</Emphasis>)

High body mass caused by fat storage during migration is believed to increase a bird's risk of predation by decreasing its ability to escape predators. We demonstrate the negative effect of wing loading (mass/wing area) on escape speed and angle of two migrating species of shorebird. We also show significant differences in escape performance between the species and genders. To help explain these differences, we test two potential proximate causes, wing shape and leg bone length. Wing shape is correlated with differences in escape performance between the species, but we found no correlation of wing shape or leg bone length with gender. Ultimately, greater predation risk due to habitat use or larger body size, for the species and genders respectively, may have resulted in evolution of enhanced escape ability.     

Intra-colonial foraging specialism by honey bees (Apis mellifera) (Hymenoptera:Apidae)

Summary Colonies of honey bees with two identifiable subfamilies were established. Returning foragers were captured and killed at two different sampling times. The mean volume and per cent soluble solids of crop contents were determined for each subfamily, as was the mean weight of the pollen pellets. No significant differences in nectar volume or concentration were detected between subfamilies within colonies. However, in a few colonies, significant subfamily by sampling-time interactions were present, suggesting that in these colonies subfamilies differed in their nectar and pollen collecting behavior at different times of day. The plant genera worked by pollen foragers were also determined. In four of six colonies, bees of different subfamilies were found to be majoring on different plant species (Fig. 1). Implications of this intra-colonial variance in foraging behavior for colony fitness are discussed. Offprint requests to: B.P. Oldroyd     

Predation on walleye pollock (Theragra chalcogramma) eggs in the western Gulf of Alaska: the roles of vertebrate and invertebrate predators

Estimates were made of the predation rate upon eggs of walleye pollock (Theragra chalcogramma) in Shelikof Strait in the western Gulf of Alaska by midwater and near-bottom fish and invertebrate predators during April 1990. Adult and juvenile walleye pollock were the dominant (99% of total abundance) planktivores collected in midwater samples. Based on visual inspection of stomach contents, a high percentage of the sampled fish were found to have consumed pollock eggs. Daily egg consumption by older age groups of walleye pollock was estimated to be <1% of the eggs available at all sampling locations. The only other fishes found to consume pollock eggs were flatfishes collected in bottom trawls but their abundances and egg consumption were very low. Gammarid and hyperiid amphipods were important invertebrate predators on eggs in the water column, as determined by immunoassays using antibodies developed specifically to ascertain the presence of pollock egg-yolk protein. Decapod shrimp showed a high proportion of positive assays in near-bottom collections. Invertebrate predators may have consumed up to 4% of the total number of eggs available in the water column, but <1% of the total near the bottom on a daily basis. Although we were not able to account for the entire daily egg mortality estimated for this stock, our method of using a combination of techniques is promising in terms of future attempts at estimating total predation mortality.     

Shifts in foraging behavior by a Battus philenor population: field evidence for switching by individual butterflies

Summary A bivoltine east Texas population of the pipevine swallowtail butterfly (Battus philenor) exhibits a seasonal shift in the shapes of non-host leaves upon which ovipositing females land. Field mark-recapture studies and analysis of the behavior of wild females of different ages were used to distinguish between two alternative mechanisms for the shift in the populations's predominant leaf-shape search mode: seasonal differences in the outcome of learning for successively emerging naive foragers which exhibit one preference for most of their lives vs. synchronous switching by experienced foragers from one learned preference to another. Results supported each hypothesis: (1) Since wild butterflies are short-lived, the seasonal shift in searching behavior must reflect at least partly the successive emergence of naive females that learn to prefer host species with different leaf shapes. The leaf-shape preferences of older females were, in fact, stronger and less variable than those of younger individuals. About 80% of 51 marked individuals maintained stable leaf-shape search modes over recapture events, exhibiting slightly stronger preferences in later observation periods. (2) Almost 16% of marked females switched search modes across recapture events. Unmarked females sometimes switched search modes within an observation period; switching occurred only after the discovery of the host species with a leaf shape differing from that originally preferred. Switching by individual butterflies was generally more frequent at the time an adult brood shifted from one dominant search mode to another. Individual switching within an adult brood was more common in those years in which the population's shift in predominant search mode occurred during that brood.The evolution of rapid learning by naive females and conservative switching by experienced females is discussed in relation to a quantitative model of switching dynamics.     

Foraging in a tidally structured environment by Red Knots (Calidris canutus): ideal, but not free

Besides the "normal" challenge of obtaining adequate intake rates in a patchy and dangerous world, shorebirds foraging in intertidal habitats face additional environmental hurdles. The tide forces them to commute between a roosting site and feeding grounds, twice a day. Moreover, because intertidal food patches are not all available at the same time, shorebirds should follow itineraries along the best patches available at a given time. Finally, shorebirds need additional energy stores in order to survive unpredictable periods of bad weather, during which food patches are covered by extreme tides. In order to model such tide-specific decisions, we applied stochastic dynamic programming in a spatially explicit context. Two assumptions were varied, leading to four models. First, birds had either perfect (ideal) or no (non-ideal) information about the intake rate at each site. Second, traveling between sites was either for free or incurred time and energy costs (non-free). Predictions were generated for three aspects of foraging: area use, foraging routines, and energy stores. In general, non-ideal foragers should feed most intensely and should maintain low energy stores. If traveling for such birds is free, they should feed at a random site; otherwise, they should feed close to their roost. Ideal foragers should concentrate their feeding around low tide (especially when free) and should maintain larger energy stores (especially when non-free). If traveling for such birds is free, they should feed at the site offering the highest intake rate; otherwise, they should trade off travel costs and intake rate. Models were parameterized for Red Knots (Calidris canutus) living in the Dutch Wadden Sea in late summer, an area for which detailed, spatially explicit data on prey densities and tidal heights are available. Observations of radio-marked knots (area use) and unmarked knots (foraging routines, energy stores) showed the closest match with the ideal/non-free model. We conclude that knots make state-dependent decisions by trading off starvation against foraging-associated risks, including predation. Presumably, knots share public information about resource quality that enables them to behave in a more or less ideal manner. We suggest that our modeling approach may be applicable in other systems where resources fluctuate in space and time.     

Are meiofaunal species cosmopolitan? Morphological and molecular analysis of Xenotrichula intermedia (Gastrotricha: Chaetonotida)

Many meiofaunal species are reported to be cosmopolitan, but due to uncertainties of identification, the affiliation of specimens from geographically distant areas to the same species-taxon is problematic. In this study, we examined morphological and molecular variation in samples of Xenotrichula intermedia Remane (Gastrotricha: Chaetonotida) from the Mediterranean Sea, the northwestern Atlantic and the northern Gulf of Mexico. Univariate analysis of 16 morphological traits was unable to detect differences among populations, except for the length of the pharynx, which was significantly shorter in the Gulf of Mexico specimens. Canonical discriminant analysis separated the Gulf of Mexico specimens from the other two populations, with pharynx length contributing about half of the total discrimination. Molecular analysis based on restriction-fragment length polymorphisms (RFLPs) in a 710-base pair polymerase chain-reaction (PCR) produet representing roughly half of the mitochondrial cytochrome oxidase I (COI) gene detected four haplotypes: one each from the Mediterranean and the Gulf of Mexico populations and two coexisting within the Atlantic population. The estimated nucleotide-sequence divergence calculated for each pairwise combination of haplotypes (based on the proportion of shared fragments) ranged from 5.3 to 11.5%. The high genetic divergence and the inability to clearly separate populations based on morphology suggest that individuals characterized by different haplotypes are genetically isolated sibling species.     

The influence of reproductive status on foraging by hoary marmots (Marmota caligata)

Summary Predictions were made and tested, comparing the foraging behavior of reproductive (rf) and nonreproductive (nrf) female hoary marmots, with the following findings: In June, no differences occur between rf's and nrf's, regarding daytime foraging. However, rf's forage more in the evening and during inclement weather, suggesting that greater nutritional needs and higher reproductive value select for the assumption of more risk while foraging. These differences disappear by August, when nutritional needs and reproductive values of rf's and nrf's are comparable. Finally, rf's reduce their near-burrow foraging relative to nrf's, which minimizes competition with their own young.     

Predation on zooplankton by the benthic anthozoans Alcyonium siderium (Alcyonacea) and Metridium senile (Actiniaria) in the New England subtidal

The Alcyonacean octocoral Alcyonium siderium Verrill and the sea anemone Metridium senile (L.), the only common perennial zooplanktivores on shallow (16-m depth) subtidal rock walls in much of northern New England, USA, are of similar heights and overlap in their habitat and microhabit distributions. The coelenteron contents of both species were sampled at four-hour intervals over a diel cycle and were compared to zooplankton available in the water at 1 to 5 cm from the rock wall, the height at which the cnidarians held their feeding tentacles. Prey in coelenterons of A. siderium were significantly smaller (means of 256 to 345 m), and those in coelenterons of M. senile were equal to or slightly greater in length (means of 415 to 1006 m) than the available zooplankton. The diets of A. siderium and M. senile differed significantly from each other and from the available zooplankton. A. siderium showed strong positive electivites for ascidian larvae and for foraminiferans, and strongly negative electivities for most crustaceans. M. senile had strong positive electivities for barnacle cyprids, ascidian larvae, and gammarid amphipods, and strong negative electivities for invertebrate eggs, foraminiferans, calanoid and harpacticoid copepods, and ostracods. Electivities may reflect tentacle avoidance or escape by motile prey as well as predator preference. Substratum-associated organisms (e.g. demersal crustaceans, larvae of benthic invertebrates) were the most common items in the diets of both species, suggesting a tight benthic food web, similar to the situation for coral reef anthrozoans which rely on reef-generated zooplankton. A. siderium ate large numbers of ascidian larvae which, as benthic adults, compete for space with A. siderium and can overgrow small colonies. Predation on the larvae of a competing species may alleviate competition by decreasing the competitor's recruitment.     

Worker size variability and foraging efficiency in Veromessor pergandei (Hymenoptera: Formicidae)

Summary As foragers of the harvester ant, Veromessor pergandei, travel further from their nest they spend significantly more time sampling seeds in experimental patches. Although accepted seeds are heavier than offered seeds, mass of accepted seed is not correlated with sampling time. Variably sized V. pergandei workers do not size-match; little, if any, variance in size of seed selected can be attributed to body size of forager. The lack of size-matching in V. pergandei suggests individual performance may be an inadequate measure of colony foraging success.     

The foraging ecology of hoverflies (Diptera,Syrphidae): circular movements on composite flowers

Summary An energetic analysis of the foraging behaviour for nectar of Eristalis tenax L. is presented. The rate of energy gain while foraging on Aster is low (0.01 W) relative to similar calculations for bees, but the flies can fill their crop in about 75–220 min. Flies visit the nectar-bearing ring of florets on a capitulum systematically, leaving when they have circled it once (Fig. 1). A simple decision-making rule appears to be used to decide when to leave.     

Adaptive peaks and alternative foraging tactics in brook charr: evidence of short-term divergent selection for sitting-and-waiting and actively searching

Some recently emerged brook charr (Salvelinus fontinalis) inhabiting still-water pools along the sides of streams are sedentary and eat crustaceans from the lower portion of the water column. Others are more active and eat insects from the upper portion of the water column. We provide evidence that this divergent foraging behavior reflects short-term divergent selection brought about by intraspecific competition in the presence of alternative food sources. Rates of encounters and interactions between individuals were density dependent, and encounter and interaction events were closely timed with prey capture attempts. In addition, aggressive fish made more foraging attempts per minute than nonaggressive fish. Aggressive fish were also either inactive or very active, while nonaggressive fish exhibited intermediate levels of activity. Growth rate potential, an important component of fitness during the early life stages of brook charr, was assessed using tissue concentrations of RNA and found to be highest for sedentary fish and for active fish making frequent foraging attempts, and lower for fish exhibiting intermediate levels of activity. Our findings support contentions that individual behavior plays an important role during initial steps in the evolution of resource polymorphisms. Received: 27 July 1998 / Accepted after revision: 16 November 1998     

Feeding by larvae of two different developmental modes in Streblospio benedicti (Polychaeta: Spionidae)

Females of the spionid polychaete Streblospio benedicti (Webster) produce either small eggs (60–70 μm diameter) and planktotrophic larvae, or large eggs (100–200 μm) and lecithotrophic larvae that reportedly do not feed. This intraspecific polymorphism, a form of poecilogony, is potentially useful in studies of larval ecology and evolution, but necessary data on larval form and function are lacking. This study describes the morphology and nutritional biology of larvae obtained from Atlantic (South Carolina) and Pacific (California and Washington) populations from 2003 to 2005. The two types of larvae produced by Atlantic S. benedicti differed greatly in length (229±22 μm SD for planktotrophs vs. 638±40 μm for lecithotrophs) and chaetiger number (2–5 vs. 10–11) at release from the female’s brood pouch. Planktotrophic larvae bore long provisional chaetae on their first chaetiger; provisional chaetae were absent in lecithotrophic larvae. Larvae from Pacific populations were all of the lecithotrophic form, and were similar to their Atlantic counterparts in all respects. High-speed video microscopy revealed that both types of larvae used opposed bands of cilia to capture suspended particles and transport them to the mouth, where they were often ingested. Lecithotrophic larvae reared with suspended phytoplankton (Rhodomonas sp., 10⁴ cells ml⁻¹) for 2 days grew significantly faster than sibling larvae reared without added food, indicating that these larvae can digest and assimilate ingested food. Larvae of S. benedicti that develop from large eggs are thus facultative planktotrophs instead of obligately non-feeding lecithotrophs, a result that affects the interpretation of comparative studies of the ecology and evolution of larvae in S. benedicti and certain other marine invertebrates.     

Effects of interactions among genotypically diverse nestmates on task specialization by foraging honey bees (Apis mellifera)

Summary Recent studies have shown that differences in patterns of task specialization among nestmate honeybee workers (Apis mellifera) can be explained, in part, as a consequence of genotypic variability. Here, we present evidence supporting the hypothesis that an individual's pattern of task specialization is affected not only by her own genotype, but, indirectly, by the genotypes of her nestmates. Workers from two strains of honey bees, one selected for high pollen hoarding, the other for low pollen hoarding, were observed in colonies of their respective parent strains and in colonies of the other strain. Worker genotype and host-colony type affected foraging activity. Workers from the high strain fostered in low-strain colonies returned with pollen on 75.6% of total foraging trips, while workers from the high strain fostered in high-strain colonies returned with pollen on 53.5% of total trips. Workers from the low strain fostered in low-strain colonies returned with pollen on 34.8% of total foraging trips while workers from the low strain fostered in high-strain colonies returned with pollen on 2.6% of total trips. Similar results were obtained in a second experiment. We suggest that workers influence the behavior of their nestmates indirectly through their effects on the shared colony environment. The asymmetry seen in the response of workers from these strains to the two types of colony environments also suggests that these genotypes exhibit different norms of reaction. Offprint requests to: N.W. Calderone     

Twilight migrations and foraging activities of the copper sweeper Pempheris schomburgki (Teleostei: Pempheridae)

During 1976 and 1977, movements and foraging activities of Pempheris schomburgki were studied on the shallow coral reefs of northeastern St. Croix, U.S. Virgin Islands. P. schomburgki emerge from daytime refuges on the backreef at about local sunset. Fifteen to 20 min after sunset, local milling groups move beyond the confines of the reef and assemble into larger groups. Twenty-five to 30 min after sunset they move in several waves along complex routes through the reef to forereef feeding grounds. Migrations of up to 1 km occur along the forereef. The reverse sequence occurs in the morning, but is earlier with respect to ambient light levels. The principal stages of these activities appear to be triggered by a combination of absolute light level, rate of change of light and state of adaptation of the eye. Migrating aggregations gradually split up into small, well-dispersed feeding groups, relatively evenly spaced along the forereef. Few individuals feed on the backreef. The principal available food consists of meroplanktonic crustaceans not available during the daytime. P. schomburgki mainly select the larger-sized individuals (mean length 5 to 6 mm), although some particles less than 1 mm are taken. These events probably represent adaptations to optimize diurnal sheltering sites, feeding grounds and the avoidance of predator activity.     

Predation on Euphausia pacifica by demersal fishes: predation impact and influence of physical variability

The importance of Euphausia pacifica Hansen to the demersal fish community off Sendai Bay, northern Japan, was studied based on a total of 256 trawl samples collected during May, October and November, from 1989 to 1992. The samples were classified into 16 assemblages according to year, month and depth. The importance of E. pacifica to the total diet of each of the assemblages was measured using an index considering both fish diet and species composition. A total of 24 fish species out of 87 were found to ingest E. pacifica. Of these, Gadus macrocephaus and Theragra chalcogramma had the highest predation impact due to their dominance in the fish assemblages and the high proportion of E. pacifica in their diets. The contribution of E. pacifica to the total diet of the fish assemblage was highest in the shallow (≤300 m) regions during May, accounting for an average of 38.5% for the 4 years. However, during May 1990, when the warm Kuroshio Extension prevailed, the contribution was higher in the deep (>300 m; 22.6%) region, reflecting active and/or passive movements of E. pacifica. The average predation impact was maximal in the May/shallow assemblage (mean ± SE; 4.6 ± 1.4 kg wet wt km² d⁻¹) and was minimal in the November/shallow assemblage (0.4 ± 0.3). Annual estimates of consumption by demersal fish fishes ranged from 43 to 128 metric tons, representing 15 to 64% of the annual commercial catch of E. pacifica by local fisheries. It is therefore suggested that the fisheries on E. pacifica have the potential to considerably impact the demersal fish assemblages. Received: 3 December 1997 / Accepted: 10 June 1998     

Preliminary evidence for the metabolism of benzo(a) pyrene by Plantago lanceolata

Polycyclic aromatic hydrocarbons (PAHs) have long been recognised as potential carcinogens in animals in which biotransformation into reactive metabolites can lead to DNA damage. In animals PAHs metabolism mainly occurs in hepatic microsomes and is associated with the cytochrome p-450 mediated mixed functional oxidase (MFO) system. PAH metabolism in plants has been shown to occur via a similar enzyme system, but has received relatively little attention. This study is looking at how the plant species Plantago lanceolata metabolises benzo(a)pyrene (B(a)P), which is one of the PAHs whose metabolism has been studied extensively in animals. The aim of the work is to establish firstly that the B(a)P is taken up and secondly that it is biotransformed by the plant to products possibly similar to those found in animals. This work is achieved by using C-14-B(a)P along with whole body autoradiography, scintillation analysis and chromatography techniques to locate the B(a)P and its metabolites.     

Life-history evidence for sibling species in Axiothella rubrocincta (Polychaeta: Maldanidae)

Individual Axiothella rubrocincta Johnson on the Californian coast live in U-shaped tubes, feed on the surface of the sediment, attain densities of 100 m^-2, attain lengths of 140 mm, have maximum oocyte diameters of 220 μm and produce demersal larvae. Individuals from Washington populations live in vertical tubes, feed 10–15 cm below the sediment surface, reach densities of 5000 m^-2, attain lengths of 60 mm, have maximum oocyte diameters of 385 μm and brood their young in their tubes to at least the 7-setiger stage. Individuals from California have a scalloped posterior margin of the cephalic plate, while individuals from Washington have entire margins. There are no other consistent morphological differences between populations. The population from Eagle Cove, Washington has individuals similar in size and oocyte diameter to individuals from California, but which maintain vertical tubes. A. rubrocincta represents a sibling species complex.     

Molecular and morphological evidence of a single species, Boccardia proboscidea (Polychaeta: Spionidae), with multiple development modes

Most species of benthic marine invertebrates have a single mode of larval development. Poecilogonous species are those that produce more than one type of larval offspring. Reports of variable development within one species, especially in combination with widely differing ecological habitat, are frequently attributed to cryptic species. The spionid polychaete Boccardia proboscidea Hartman, 1940 exhibits development that varies both within a single brood and among broods produced by different females. Some females have planktotrophic development and produce many small larvae with a 2 week planktonic period before metamorphosis. Other females produce broods containing both planktotrophic larvae as well as nurse-egg-ingesting (adelphophagic) offspring that hatch as juveniles. Molecular analysis (RAPD-PCR) showed that a significant proportion of genetic variance is attributable to geographic origin, and not to developmental type. Adults of both developmental types showed no consistent differences in taxonomically important features (e.g. type and arrangement of chaetae, modified fifth setiger, caruncle, branchiae, pygidium) when examined with SEM. These data support the hypothesis that developmental variability in this species is a case of poecilogony, and is not attributable to cryptic species. Received: 21 April 1998 / Accepted: 20 April 1999     

Effects of parasites on foraging and defense behavior of a termitophagous ant,Pheidole titanis Wheeler (Hymenoptera: Formicidae)

Summary Pheidole titanis Wheeler, an ant that occurs in desert and deciduous thorn forest in the southwestern United States and western Mexico, is a predator on termites. In the dry season well-coordinated raids against termite foraging parties occur early in the morning or late in the afternoon, whereas in the wet season most raids occur at night. This seasonal shift in the timing of raids is due to the increased activity of a fly (Diptera: Phoridae) that is a specialist parasitoid on P. titanis workers and soldiers. When parasitic flies discover P. titanis nest entrances or raiding columns, workers stop foraging and defend themselves against oviposition attacks. Flies are only active during the day and never interfere with foraging at night. However, P. titanis does not increase the frequency of raids at night and, as a result, colonies collect less food in the wet season compared to the dry season. Presence of parasitic flies also interferes with normal defense behavior of P. titanis against conspecific and heterospecific enemy ants. Dissections of P. titanis workers and soldiers suggest that the parasitism rate by flies is less than 2% and observations indicate that parasitic flies are much rarer than their host workers and soldiers. Nonetheless, these parasites exert a strong ecological impact on their host.