Relationship between body size, provisioning rate, longevity and reproductive success in females of the solitary bee Osmia cornuta

Body size has often been related to reproductive success in bees and wasps. The objective of this 3-year study was to analyze the relationship between nesting female body size, provisioning rate and longevity and their effect on several traits related to parental investment and reproductive success in the solitary bee Osmia cornuta. Body size was not correlated to longevity, and it was only correlated to provisioning rate in the third year (with poor weather conditions during nesting). Variation in fecundity, offspring size and offspring mortality was not well explained by nesting female body size in any of the 3 years. However, in the third year, small females biased their investment toward males, the sex requiring smaller pollen–nectar provisions. Large females were more successful usurpers of other females' nests, but fecundity of usurpers was no higher than fecundity of nonusurpers. Large females were more likely to establish at the release site, probably in relation to size-dependent vigor at emergence. A review of the literature on parental investment in solitary aculeate Hymenoptera showed a stronger relationship between body size and reproductive success in wasps than in bees. In O. cornuta, fecundity was strongly related to longevity and provisioning rate in all 3 years. Offspring size was associated with provisioning rate in 1 year, when females with higher provisioning rates tended to produce larger sons and daughters. Both longevity and provisioning rate appeared to be strongly conditioned by stochastic events.     

Interference by wood ant influences size selection and retrieval rate of prey by Formica fusca

Summary Dominance relations between the territorial wood ant Formica polyctena and the submissive F. fusca generate testable predictions on the mechanisms of their coexistence. Here I tested the influence of interference competition by the dominant F. polyctena on the foraging of F. fusca. In the presence of F. polyctena, the activity and the retrieval rate of items of F. fusca decreased significantly. When F. fusca were given a choice between small chironomids and flies a hundred times heavier they selected nearly always flies in the absence of wood ants; when disturbed F. fusca took proportionately more chironomids. In nature, irrespective of distance from the wood-ant mound the size distribution of potentially available food items was the same. F. fusca collected smaller items close to the wood-ant mound, where the density of interfering wood ants was high, than far from it. Also, F. polyctena carried larger protein items from the outskirts of its territory than from the center. The items of F. fusca were on average smaller than those of F. polyctena although the item-size overlap was substantial. When encountering a F. polyctena, those F. fusca workers carrying a fly always lost their booty to the dominant but always managed to bring the chironomids to the nest. F. fusca and F. polyctena were equally efficient in detecting single chironomids placed on the surface of ground, but the presence of either species decreased the discovery rate of the other. This implies mutual exploitation competition between the species for the locally most abundant protein resource. F. polyctena found single flies faster than F. fusca. In an earlier study I showed that close to the wood-ant mound the colony size and production of sexual offspring of F. fusca were reduced. I suggest that the suppressed colony success of F. fusca is attributable to the diminished size and decreased retrieval rate of prey items close to the woodant mound abounding with interfering wood-ant foragers.     

Intragroup dynamics of a cooperative breeder: An anlysis of reproductive roles in the acorn woodpecker

Summary The acorn woodpecker is a cooperative breeder generally thought to breed promiscuously within groups, but alternative patterns of reproductive investment can be expected and remain to be examined. With this feature of mating system in mind, we studied the reproductive roles of individuals in a single group of this species in central coastal California over a three-year period. Particularly detailed observations were made in 1979, when the group consisted of two potentially breeding sibling males and a single breeding female. One male (298) contributed significantly more than 297 in virtually all aspects of nest and fledgling care. These include (1) feeding nestlings, (2) nest sanitation, (3) nocturnal incubation and brooding, (4) feeding fledglings, and (5) escorting fledglings. In the few observed and rarely occurring dominance interactions, 298 prevailed. These results suggest that 298 had a higher probability of fathering the group's two 1979 offspring than did his brother. Thus, the two males differed substantially in their parental roles within the group, and very likely in reproductive roles, also. At present stage of knowledge, monogamy in groups with one female and more than one potential reproductive male is no less likely than promiscuity. These findings are discussed with respect to the evolution of cooperative breeding in this species.     

Clam predation by whelks (Busycon spp.): Experimental tests of the importance of prey size,prey density,and seagrass cover

In 57 l-m² samples within a meadow of Halodule wrightii in Bogue Sound, North Carolina, USA, densities of the clams Mercenaria mercenaria and Chione cancellata were positively associated with seagrass cover. Where seagrass was experimentally removed, marked individuals of both clam species exhibited high rates of mortality in fine sand sediments during two successive experiments spanning 13 months. In the unaltered (control) seagrass meadow, M. mercenaria density remained constant over 13 months and C. cancellata density declined at a slower rate than in the unvegetated plots. Seagrass provides these clams with a refuge from whelk (Busycon carica, B. contrarium, and B. canaliculatum) predation, the major cause of mortality and population decline in experimentally unvegetated plots. In 2 factorial field experiments in unvegetated substratum in which densities of M. mercenaria and C. cancellata were varied independently, first over 5 levels (0 X, 1/2X, 1 X, 2 X, 4 X) and subsequently over 4 levels (0 X, 1/4 X, 1 X, 4 X), there was no repeatable intra- or interspecific effect of density on percent survival, or on the rate of any mortality type. Whelk predation fell preferentially on larger size classes of both species, whereas factors which contribute to clam disappearance usually acted more intensely on smaller sizes. Experimental exclusion of large predators by caging demonstrated that even in unvegetated substratum survivorship of both clam species was high in the absence of whelks and other predators. Individuals of C. cancellata live closer to the sediment surface than those of M. mercenaria, which may explain why seagrass does not serve as effectively to protect them from whelk predation. The mechanism of whelk inhibition may depend upon sediment binding by the H. wrightii root mat, which produces a demonstrable decrease in the physical penetrability of surface sediments.     

Predator biomass,prey density,and species composition effects on group size in recruit coral reef fishes

Group incidence and size are described for recruit parrotfishes, wrasses, and damselfishes on Hawaiian reefs over 3 years (2006–2008) at sites spanning the archipelago (20–28°N, 155–177°W). Coral-poor and coral-rich areas were surveyed at sites with both low (Hawaii Island) and high (Midway Atoll) predator densities, facilitating examination of relations among predator and recruit densities, habitat, and group metrics. Predator and recruit densities varied spatially and temporally, with a sixfold range in total recruit densities among years. Group (≥2 recruits) metrics varied with time and tracked predator and recruit densities and the proportion of schooling species. Groups often included heterospecifics whose proportion increased with group size. A non-saturating relationship between group size and recruit density suggests that the anti-predator benefits of aggregation exceeded competitive costs. Grouping behavior may have overarching importance for recruit survival–even at high recruit densities–and merits further study on Hawaiian reefs and elsewhere.     

Predator and prey space use: dragonflies and tadpoles in an interactive game

Predator and prey spatial distributions have important population and community level consequences. However, little is known either theoretically or empirically about behavioral mechanisms that underlie the spatial patterns that emerge when predators and prey freely interact. We examined the joint space use and behavioral rules governing movement of freely interacting groups of odonate (dragonfly) predators and two size classes of anuran (tadpole) prey in arenas containing two patches with different levels of the prey's resource. Predator and prey movement and space use was quantified both when they were apart and together. When apart from predators, large tadpoles strongly preferred the high resource patch. When apart from prey, dragonflies weakly preferred the high resource patch. When together, large prey shifted to a uniform distribution, while predators strongly preferred the high resource patch. These patterns qualitatively fit the predictions of several three trophic level, ideal free distribution models. In contrast, the space use of small prey and predators did not deviate from uniform. Three measures of joint space use (spatial correlations, overlap, and co-occurrence) concurred in suggesting that prey avoidance of predators was more important than predator attraction to prey in determining overall spatial patterns. To gain additional insight into behavioral mechanisms, we used a model selection approach to identify behavioral movement rules that can potentially explain the observed, emergent patterns of space use. Prey were more likely to leave patches with more predators and more conspecific competitors; resources had relatively weak effects on prey movements. In contrast, predators were more likely to leave patches with low resources (that they do not consume) and more competing predators; prey had relatively little effect on predator movements. These results highlight the importance of investigating freely interacting predators and prey, the potential for simple game theory models to predict joint spatial distributions, and the utility of using model choice methods to identify potential key factors that govern movement.     

Paternal care in the European starling,Sturnus vulgaris: nestling provisioning

The extent to which male birds in polygynous species with biparental care assist in nestling feeding often varies considerably between nests of different mating status. Both how much polygynous males assist and how they divide their effort between nests may have a profound effect on the evolution of mating systems. In this study we investigated how males in the facultatively polygynous European starling Sturnus vulgaris invested in their different nests. The amount of male assistance affected the quality of the offspring. Polygynous males invested as much as monogamous males, but divided their effort asymmetrically between nests, predominantly feeding nestlings of first-mated (primary) females. Although females partly compensated for loss of male assistance, total feeding frequency was lower at primary females’ nests than at monogamous females nests. Secondary females received even less assistance with nestling rearing, and the extent to which males assisted decreased with the length of the interval between the hatching of the primary and secondary clutches. These results are contrasted with those from a Belgian populations of starlings with a much more protracted breeding season and thus greater opportunities for males to attract additional mates during the nestling rearing period. The results show that both the “defence of male parental investment model” and the “asynchronous settlement model” have explanatory power, but that their validity depends on the potential length of the breeding season. Received: 21 July 1995/Accepted after revision: 13 July 1996     

Relationship between phytoplankton cell size and the rate of orthophosphate uptake: in situ observations of an estuarine population

Orthophosphate uptake by a natural estuarine phytoplankton population was estimated using two methods: (1) ³²P uptake experiments in which filters of different pore sizes were used to separate plankton size-fractions; (2) ³³P autoradiography of phytoplankton cells. Results of the first method showed that plankton cells larger than 5 m were responsible for 2% of the total orthophosphate uptake rate. 98% of the total uptake rate occurred in plankton composed mostly of bacteria, which passed the 5 m screen and were retained by the 0.45 m pore-size filter. There was no orthophosphate absorption by particulates in a biologically inhibited control containing iodoacetic acid. Orthophosphate uptake rates of individual phytoplankton species were obtained using ³³P autoradiography. The sum of these individual rates was very close to the estimated rate of uptake by particulates larger than 5 m in the ³²P labelling experiment. Generally, smaller cells were found to have a faster uptake rate per m³ biomass than larger cells. Although the nannoplankton constituted only about 21% of the total algal biomass, the rate of phosphate uptake by the nannoplankton was 75% of the total phytoplankton uptake rate. Results of the plankton autoradiography showed that the phosphate uptake rate per unit biomass is a power function of the surface: volume ratio of a cell; the relationship is expressed by the equation Y=2x10^-11 X ^1.7, where Y is gP m^-3 h^-1 and X is the surface: volume ratio. These results lend support to the hypothesis that smaller cells have a competitive advantage by having faster nutrient uptake rates.     

Trade-offs between maternal foraging and fawn predation risk in an income breeder

The choice of neonatal hiding place is critical for ungulates adopting hiding anti-predator strategies, but the consequences of different decisions have rarely been evaluated with respect to offspring survival. First, we investigated how landscape-scale choices made by roe deer fawns and their mothers affected predation risk by red foxes in a forest–farmland mosaic in southeastern Norway. After, we examined the effect of site-specific characteristics and behaviour (i.e. visibility, mother–fawn distance and abundance of the predator’s main prey item—small rodents) on predation risk. The study of habitat use, selection and habitat-specific mortality revealed that roe deer utilised the landscape matrix in a functional way, with different habitats used for feeding, providing maternal care and as refugia from predation. Mothers faced a trade-off between foraging and offspring survival. At the landscape-scale decisions were primarily determined by maternal energetic constraints and only secondarily by risk avoidance. Indeed, forage-rich habitats were strongly selected notwithstanding the exceptionally high densities of rodents which increased fawn predation. At fine spatial scales, a high visibility of the mother was the major factor determining predation risk; however, mothers adjusted their behaviour to the level of risk at the bed site to minimise predation. Fawns selected both landscape-scale refugia and concealed bed sites, but failure to segregate from the main prey of red foxes led to higher predation. This study provides evidence for the occurrence of spatial heterogeneity in predation risk and shows that energetically stressed individuals can tackle the foraging-safety trade-off by adopting scale-dependent anti-predator responses.     

Feeding behaviour and prey size selection of gilthead seabream,Sparus aurata,larvae fed on inert and live food

Prey selection shortly after the onset of feeding by laboratory-reared gilthead seabream, Sparus aurata L., larvae was studied using larvae fed on two types of microcapsule (hard- and soft-walled) having diameters ranging from 25 to 300 m. Preferences between inert food and live prey (rotifers and Artemia sp. nauplii) were also studied. Seabream larvae were able to ingest inert food from first feeding. Larvae of all size classes ingested hard microcapsules with diameters in the range 25 to 250 m. However, larvae with a total length (TL) below 4 mm preferentially selected particles 25 to 50 m in diameter, larvae of TL 4 and 5 mm preferred particles 51 to 100 m in diameter, while larvae above 5 mm TL preferred particles 101 to 150 m in diameter. With soft microcapsules, larvae always preferred particles larger than in the previous case, and above 4.5 mm TL they preferentially selected particles 201 to 250 m in diameter. In addition, the gradual increase of preferred diameters with increasing TL was more pronounced when larvae were increasing TL was more pronounced when larvae were fed on soft particles. Mean values for prey width/mouth width ratios were approximately 0.24 and 0.30 when larvae were fed on hard-walled and soft-walled microcapsules, respectively, irrespective of the absolute value of larval length. When a mixed diet of live and inert food items was offered, live prey were always preferentially selected, even if the prey width/mouth width ratio was apparently not favourable. Only a physical constraint such as excessive prey width could counter this preference for living prey vs inert microcapsules. These results contribute to our knowledge in larval feeding behaviour, especially in the presence of inert food, and represent a fundamental step in developing prepared food for marine fish larvae.     

Chinstrap penguins alter foraging and diving behavior in response to the size of their principle prey, Antarctic krill

Penguins may exhibit plasticity in their diving and foraging behaviors in response to changes in prey availability. Chinstrap penguins are dependent predators of Antarctic krill in the Scotia Sea region, but krill populations have fluctuated in recent years. We examined the diet of chinstrap penguins at Livingston Island, South Shetland Islands, in relation to their diving and foraging behavior using time-depth recorders over six breeding seasons: 2002–2007. When krill were smaller, more chinstrap penguins consumed fish. In these years, chinstrap penguins often exhibited a shift to deep dives after sundown, and then resumed a shallower pattern at sunrise. These night dives were unexpectedly deep (up to 110 m) and mean night dive depths sometimes exceeded those from the daytime. The average size of krill in each year was negatively correlated to mean night dive depths and the proportion of foraging trips taken overnight. Based on these patterns, we suggest that when krill were small, penguins increasingly targeted myctophid fish. The average krill size was negatively correlated to the time chinstrap penguins spent foraging which suggests that foraging on smaller krill and fish incurred a cost: more time was spent at sea foraging.     

Vigilance behaviour and fitness consequences: comparing a solitary foraging and an obligate group-foraging mammal

Vigilance behaviour in gregarious species has been studied extensively, especially the relationship between individual vigilance and group size, which is often negative. Relatively little is known about the effect of conspecifics on vigilance in non-obligate social species or the influence of sociality itself on antipredator tactics. We investigated predator avoidance behaviour in the yellow mongoose, Cynictis penicillata, a group-living solitary forager, and compared it with a sympatric group-living, group-foraging herpestid, the meerkat, Suricata suricatta. In yellow mongooses, the presence of conspecifics during foraging—an infrequent occurrence—reduced their foraging time and success and increased individual vigilance, contrary to the classical group-size effect. Comparing the two herpestids, sociality did not appear to affect overt vigilance or survival rates but influenced general patterns of predator avoidance. Whereas meerkats relied on communal vigilance, costly vigilance postures, and auditory warnings against danger, yellow mongooses avoided predator detection by remaining close to safe refuges and increasing “low-cost” vigilance, which did not interfere with foraging. We suggest that foraging group size in herpestids is constrained by species-distinct vigilance patterns, in addition to habitat and prey preference.     

Energetics,reproductive suppression and obligate communal breeding in carnivores

Summary Leptogenys processionalis Jerdon forages on termites and other arthropods by raiding in branched trails. Growth and topology of these search trails were studied using Horton's (1945) technique orginally developed to analyze the branching pattern of river systems. Branching was always a bifurcating process and branches emerged symmetrically on either side of the main trail. Branching coefficients (R _b ) were similar to those of a few biological branching systems, such as lungs, that are considered to be non-random in their branching pattern. The R _b values indicated that the rate of branching and growth of trails remained constant within each foraging bout. The length of trails became shorter as they grew out and branched. The branching process was a function of the spatial separation of food patches in the terminal search field. Ants in the terminal search field send signals on encountering prey. The recruits cannot discriminate between these signals if they arise from two food patches situated <40 cm from each other, and hence converge on them in a single trail. However, discrimination is possible when food patches are >40 cm apart and hence recruits congregate on them separately in two trails, resulting in branching. Thus, the branching process is a result of independent decision by the ants conforming to certain simple rules and not a collective decision of the whole colony. We argue that mass recruiting ants selected to forage by branching pattern of trails because of its efficiency over other topologies (Stevens 1973) in minimizing the cost of travel, both from the nest to the food patches and between food patches. Further, the branch angles appear to be a trade-off to minimize travel cost and the resistance to the flow of ants comprising the column.Offprint requests to: T. Veena     

Hatching asynchrony and survival of insurance offspring in an obligate brood reducing species,the American white pelican

American white pelicans (Pelecanus erythrorhynchos) lay two eggs but typically rear only one young owing to siblicidal brood reduction affecting the later-hatched, or B-chick. When the A-offspring fails at an early age, the B-chick may survive as a replacement (insurance) offspring. Using a combination of nests with natural and artificially manipulated hatching asynchrony, I examined the hypothesis that hatching asynchrony in this species is adaptively tuned to permit B-chicks to survive during the time they are most likely to be needed as replacements, with brood reduction following when they become redundant. Hatching asynchrony over the natural range of 0–4 days significantly increased within-brood mass differentials and reduced B-chick lifespan. Full synchrony had a marginally negative effect on A-chick mass. Greater asynchrony did not significantly affect the number of days B-chicks survived after hatching of the A-chick, owing to a corresponding extension of time B-offspring were protected from harassment while still within the egg. This resulted in a high probability (> 0.8) of B-chicks surviving through the initial period (5–7 days) of maximum early A-chick loss. Redundant B-chicks were subject to heavy brood reduction, with both chicks likely to have survived at only one each of 94 natural and 84 manipulated (0, 2, and 4 days asynchrony) nests. Hatching asynchrony in American white pelicans, in combination with a rapid development of senior chick siblicidal competence, appears to result in a time course of brood reduction appropriate for an effective insurance reproductive strategy. Received: 2 February 1996 /Accepted after revision: 18 May 1996     

Temporal variability in functional responses and prey selectivity of the pelagic mysid, Mysis mixta, in natural prey assemblages

Predation rates and prey selection of the pelagic mysid shrimp, Mysis mixta, were studied experimentally in the northern Baltic Sea in 1998 during their most intensive growth period, from June to October. Functional responses during 5 months were determined by providing the mysids with a natural zooplankton assemblage, diluted to several different concentrations. The results show that ingestion rate increased, along with mysid growth, from early summer to autumn and that saturation level was reached between 400 and 500 μg C l⁻¹. Ingestion rates increased with increasing prey concentration, and sigmoidal curves explained mostly the variation in ingestion rates (explanatory levels of 86–97%). Prey selection was evident in June, July and August, though weaker during the latter 2 months. Selection differed between the studied months but, generally, copepods were more positively selected than cladocerans. Rotifers were the main prey during June and July, when mysids were small, while larger mysids fed on copepods and cladocerans. Of the copepods, Eurytemora affinis was a truly selected species. This study shows that mysids feed on many zooplankton taxa and that they undergo ontogenetic diet shifts. Received: 19 July 2000 / Accepted: 19 October 2000     

Echolocation call design and limits on prey size: a case study using the aerial-hawking bat Nyctalus leisleri

The echolocation calls used by Nyctalus leisleri during search phase in open air space are between 9 and 14 ms long, with the peak energy between 24 and 28 kHz. The pulses are shallowly frequency-modulated with or without an initial steep frequency-modulated component. The diet consists primarily of small flies (Diptera), including many chironomids (wingspan 9–12 mm) and yellow dung flies (Scatophaga; wingspan 24 mm), but also of some larger insects such as dung beetles (Coleoptera; Scarabaeoidea), caddis-flies (Trichoptera) and moths (Lepidoptera). The echo target strength of some prey items was measured. Contrary to models based on standard targets such as spheres or disks, the echo strength of real insects was found to be virtually independent of the emitted frequency within the 20–100 kHz frequency range. A model was used to calculate probable detection distances of the prey by the bat. Using narrow-band calls of 13.7 ± 2.7 ms duration, a bat would detect the two smallest size classes of insect at greatest range using calls of 20 kHz. The results may therefore explain why many species of large and medium sized aerial-hawking bats use low-frequency calls and still eat mostly relatively small insects. The data and model challenges the assumption that small prey are unavailable to bats using low-frequency calls.     

Complexity in an obligate mutualism: do fungus-gardening ants know what makes their garden grow?

Fungus gardening ants make clear choices among fungal substrates (food for their fungus). It has been proposed, but never demonstrated, that these ants are collecting the best for their symbiotic fungus and the production of ant biomass (fitness). The goal of this study was to determine whether preferred substrates lead to higher fitness in the attine, Trachymyrmex septentrionalis. Preferences exhibited by foragers were established. Colonies were fed a single substrate or a mixture of substrates during the entire course of the experiment, which ended when sexual offspring appeared in the nest. The response variables were numbers and weights of ant offspring and the chitin content of fungus gardens. Preference was not strongly related to fitness. The preferred oak catkins produced the highest amounts of ant and fungal biomass, but the ants collected much more material than needed, which indicates that forager activity is decoupled from fitness. The preferred caterpillar feces were rejected shortly after the feedings began. The unpreferred oak leaves were just as effective at producing ant and fungal biomass as catkins. Leaves are possibly unpreferred because they are expensive to cut. The unpreferred huckleberry flowers were inferior but did not cause rejection behavior. The mixed diet was just as productive as catkins or leaves. This study indicates that foragers possess a default mechanism to prefer catkins and frass, which can be quickly changed if substrates are bad. In contrast, there does not appear to be a similar mechanism causing substrates to become preferred quickly.     

Graded recruitment and hunting strategies linked to prey weight and size in the ponerine ant Ectatomma ruidum

According to the weight and size of their prey, Ectatomma ruidum workers can employ different recruitment systems (solitary hunting, cooperative hunting and group hunting with recruitment) when mastering and retrieving prey items from short distances from the nest. Prey size determined the backwards entry typically adopted by this species, while prey weight determined the predatory strategy selected. After a common initial sequence (search for prey, detection, localization), predatory sequences varied in terms of the type of approach, the site of seizure, the reaction after stinging and the type of transport. Nevertheless, irrespective of prey weight and size, seizure was preferentially oriented towards the head and prey were always stung. Short-range recruitment and mass recruitment without trail laying were elicited by a large range of heavy prey (> 2.5 times the weight of an individual worker). According to the mortality risk associated with each prey, hunters exhibited a “prudent” stinging posture associated with an increase in the duration of the subsequent phase of waiting for prey immobilization. The overall time of capture was positively correlated with the weight of the prey. When collective hunting strategies were involved, E. ruidum colonies matched the number of recruited hunters to the size and weight of the prey. Compared to solitary hunting strategies, for short food–nest distances, this graded recruitment appeared to enhance the energetic benefits derived by this species from the use of recruitment systems: the higher the number of workers involved in the recruitment process, the greater the energetic benefits obtained. The exhibition or absence of trail laying behavior in the recruitment responses displayed by E. ruidum workers is discussed in relation to their involvement in scavenging or predatory behavior. Received: 27 June 1996 / Accepted after revision: 3 March 1997     

Relative importance of prey size and concentration in determining the feeding behaviour of the herringClupea harengus

The feeding behaviour ofClupea harengus L. in the light is dependent primarily on prey concentration. In the laboratory the fish feed by biting at low prey concentrations and by filtering at high concentrations. With the brine shrimpArtemia sp. as prey, the concentration required for the onset of filter-feeding was directly dependent on prey size, but the concentration at which 50% of feeding fish were filtering differed little between three sizes of brine shrimp (nauplii, and 2 and 4 mm larvae). When fed onCalanus finmarchicus, however, 50% of fish fed by filtering at concentrations at least six times lower than on any size of brine shrimp. Filter-feeding thresholds forC. finmarchicus were six to ten times lower than for any size ofArtemia sp. and, on the basis of biomass, approximately eight times lower than for equivalent sizedArtemia sp.