Host location and exploitation by the cleptoparasitic wasp Argochrysis armilla: the role of learning (Hymenoptera: Chrysididae)

Summary The nesting behaviors of many solitary ground-nesting wasps incorporate temporal barriers against would-be cleptoparasites. Nests being excavated are conspicuous but relatively invulnerable to parasites, while nests being provisioned, often several hours to days later, are inconspicuous but highly vulnerable, Argochrysis armilla, a cleptoparasite of solitary ground-nesting wasps, Ammophila spp., bridges the temporal gap between nest excavation and provisioning by (i) visually locating digging hosts, (ii) learning the locations of associated nests, (iii) maintaining surveillance on a series of nests during the hosts' absence, and (iv) ovipositing in nests when the host returns with provisions. Patterns of surveillance and parasitism of Ammophila dysmica nests were generated by the number of cleptoparasites discovering and learning the nest's location during excavation. These results support recent suggestions that learning may play an important role in shaping foraging strategies of insect parasites.     

Relatedness among nestmates in a primitively social wasp,Cerceris antipodes (Hymenoptera: Sphecidae)

Summary Kin selection, acting through high levels of relatedness, may be an important promoter in the evolution of nest sharing. Cerceris antipodes is a sphecid wasp that shares nests in contrast to the majority of sphecids where only one female occupies a nest. Nest sharing results from females remaining in their natal nests and females moving to already occupied nests. Average relatedness among nestmates of C. antipodes was calculated from allele frequencies of phosphoglucomutase to determine whether nests were usually shared by close relatives. Relatedness among nestmates was high (0.5 to 0.6) at one aggregation in two consecutive years. Preferential association of relatives away from the natal nest can be inferred from these high values combined with the frequency of nest switching observed. Estimates of relatedness were lower (about 0.3) and associated with large standard errors at 3 other aggregations. Inbreeding and relatedness between neighbouring nests were only significant at the aggregation with the fewest females. This may be a result of the small effective population size. The levels of relatedness observed are consistent with kin selection through relatedness being an important factor in the evolution of nest sharing.     

Sex ratios and life-history patterns of a solitary wasp,Trypoxylon (Trypargilum) politum (Hymenoptera : Sphecidae)

Summary Werren and Charnov (1978) and Seger (1983) proposed a model to explain a fairly common pattern of alternating sex ratio biases between generations in partially bivoltine insects. When first-generation males overlap and mate with females of the second generation, then females should bias sex ratios in favor of sons for the first generation and daughters for the second generation. In an intensive, 7-year study at four sites in northern Florida, pipe-organ mud-daubing wasps (Trypoxylon (Trypargilum) politum; Hymenoptera: Sphecidae) were found to have strongly male-biased sex ratios in the first or overwintering generation and 1:1 or female-biased sex ratios in the second or summer generation. These differences were not due to differences in mortality of the two sexes but rather resulted from changing female sex-allocation decisions. In some respects the mud dauber results fit Seger's model well: alternating sex ratios in partially bivoltine populations, first-generation males overlapping second-generation females and perhaps most convincingly, northern, univoltine populations do not have a male bias. Despite this qualitative fit, however, our data do not meet the quantitative predictions of the model. This could result from the fact that some assumptions of the model are not met by the life history of T. politum. Alternative explanations for alternating sex ratios include split sex ratios, seasonal differences in cost ratios, facultative maternal investment rules and facultative overwintering decisions by offspring. Despite the position that sex ratios have achieved in the modern study of evolution, it is clear that accurate, quantitative predictions on sex-allocation patterns demand the same detailed understanding of the biology of the organism that is required for the study of other adaptations.     

Predatory behavior of Polybia sericea (Olivier), a tropical social wasp (Hymenoptera: Vespidae)

Summary Polybia sericea (Olivier) (Hymenoptera: Vespidae) foragers were trained to visit experimental foraging plots and tests were conducted to determine the role of visual, olfactory, and chemotactile cues in prey location. Foragers approached prey from downwind and hovered downwind of visual and olfactory stimuli. Olfactory cues were more likely to elicit landing than were visual cues. Biting of potential prey was most consistently elicited by a combination of visual, tactile, and chemotactile cues. Foragers encountering large prey carried a piece back to the nest and returned for prey remains. These returning foragers used visual cues to direct intensive aerial search; olfactory prey cues elicited landing.     

Gregarious nesting of a digger wasp as a “selfish herd” response to a parasitic fly (Hymenoptera: Sphecidae; Diptera: Sacrophagidae)

Summary Females of a digger wasp Crabro cribrellifer nested in a 234 nest aggregation covering 2x65 m in area. Variability in 3 edaphic factors (soil temperature, hardness, and per cent moisture) had no significant influence on the non-random spatial distribution of nests within the aggregation. A significant positive correlation between wasp nest density and the mean number of parasitic flies, Metopia campestris, present was shown (Fig. 1). Yet, based on 20 excavated nests, the probability of cell parasitism decreased as nest density increased (Fig. 2). This may give females nesting within a dense aggregation a selective advantage, and supports the selfish herd model developed by Hamilton (1971).     

Protandry in western cicada killer wasps, (Sphecius grandis,Hymenoptera: Sphecidae): an empirical study of emergence time and mating opportunity

Summary Sphecius grandis is a univoltine, colonial wasp. Females mate once and are sexually receptive when they emerge in July and early August. Males generally emerge earlier in the summer than females. The opportunity for each male to acquire mates is a function of the number of females emerging during his lifetime and the number of competitiors that are active when the females emerge. I determined a mating opportunity index (MOI) for each male in an aggregation of wasps for three separate summers, and correlated the MOI of individual males with their emergence date. The MOI of a male estimates the potential contribution that timing of emergence makes to his reproductive success. In 1984 males emerging near the mean of the male emergence distribution had the highest MOI. These males emerged between one and two weeks prior to the mean female emergence date. However, in 1981 late emerging males had the highest MOI. In 1983 there was no significant difference in MOI among males. As a result of between-year variation in female emergence schedules and in the duration of male lifetime, the selection pressures influencing male emergence time vary between years.     

Size-dependent, spatial and temporal genetic variation at a leucine aminopeptidase (LAP) locus among blue mussel (Mytilus galloprovincialis) populations along a salinity gradient

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus is related to salinity variation in several marine bivalve molluscs. This paper details an investigation of the Long Island Sound model of LAP selection (LAP genotype-dependent mortality occurring among newly settled Mytilus edulis mussels) carried out in 1997 among three M. galloprovincialis mussel populations along the salinity gradient of Wellington Harbour, New Zealand. Significant LAP genotypic heterogeneity was observed at the LAP locus between small (<25 mm shell length) and large (>25 mm shell length) M. galloprovincialis from Petone and Eastbourne (the two sites experiencing the greatest salinity variation), whereas genotypic heterogeneity was not significantly different between small and large mussels from Seatoun (the site experiencing the least salinity variation). The Lap ³ allele decreased in frequency and the Lap ⁴ allele increased in frequency at Petone and Eastbourne, whereas the Lap ³ and Lap ⁴ allele frequencies remained effectively constant at Seatoun. Both these findings are consistent with the Long Island Sound model of selection. At all three locations, the Lap ^3,3 genotype decreased in frequency from small to large mussels, whereas the Lap ^3,4 genotype increased in frequency from small to large mussels. All other LAP genotypes occurred at low frequencies (<0.10) at all three locations and showed no evidence of frequency change from small to large-size mussels nor evidence of clinal change among the three locations. These genotype frequency data possibly indicate that the Lap ^3,3 genotype is at a selective disadvantage compared to the Lap ^3,4 genotype at all three locations, and that this selective disadvantage is related to the extent of salinity variation which exists at each location. Further investigation is required before it can be determined if the Long Island Sound model of selection best describes the size-dependent and location-specific changes in LAP allele and genotype frequencies along this salinity gradient. Comparison of the population genetic structure at the LAP locus in 1995 and in 1997 revealed a profound change from heterozygote excesses to heterozygote deficiencies for all three M. galloprovincialis populations. The reason for the change is unknown, but the change indicates that population genetic structure at the LAP locus is highly variable in time, but consistent in space, among these M.␣galloprovincialis populations. Received: 5 February 1998 / Accepted: 27 May 1998     

Prey dynamics affect foraging by a pelagic predator (Stenella longirostris) over a range of spatial and temporal scales

Studies have shown that pelagic predators do not overlap with their prey at small scales. However, we hypothesized that spinner dolphin foraging would be affected by the spatio-temporal dynamics of their prey at both small and large scales. A modified echosounder was used to simultaneously measure the abundance of dolphins and their prey as a function of space and time off three Hawaiian islands. Spinner dolphin abundance closely matched the abundance patterns in the boundary community both horizontally and vertically. As hypothesized, spinner dolphins followed the diel horizontal migration of their prey, rather than feeding offshore the entire night. Spinner dolphins also followed the vertical migrations of their prey and exploited the vertical areas within the boundary layer that had the highest prey density. Cooperative foraging by pairs of dolphins within large groups was evident. The geometric and density characteristics of prey patches containing dolphins indicate that dolphins may alter the characteristics of prey patches through this cooperative foraging. The overlap of Hawaiian spinner dolphins and their prey at many temporal and spatial scales, ranging from several minutes to an entire night and 20 m to several kilometers, indicates that the availability of truly synoptic data may fundamentally alter our conclusions about pelagic predator-prey interactions.     

Age-related repertoire expansion and division of labor in Pheidole dentata (Hymenoptera: Formicidae): a new perspective on temporal polyethism and behavioral plasticity in ants

The controversy concerning the extent to which the organization of division of labor in social insects is a developmental process or is based on task allocation dynamics that emerge from colony need independent of worker age and endocrine or neural state has yet to be resolved. We present a novel analysis of temporal polyethism in the ant Pheidole dentata, demonstrating that task attendance by minor workers does not shift among spatially associated sets of behaviors that minimally overlap but rather expands with age. Our results show that the number of tasks performed by older minors increases through the addition and retention of behaviors, with up to a sixfold increase in repertoire size from day 1 to day 20 of adult life. We also show that older minors respond to colony needs by performing significantly more brood care as its demand increases, indicating that they can quickly upregulate nursing according to labor requirements. This level of plasticity was absent in younger siblings. The breadth of responsiveness to task-related olfactory stimuli increased with age. In a binary choice test in which young and old minor workers could orient toward odorants from brood or food, older workers responded to both brood and food, whereas young workers responded only to brood. These dissimilar responses to stimuli associated with nursing and foraging indicate age-related differences in sensory ability and provide a physiological basis for the age-related repertoire expansion model. We discuss repertoire expansion in P. dentata in light of behavioral development and caste flexibility in ants.     

Provisioning behavior and the estimation of investment ratios in a solitary bee,Calliopsis (Hypomacrotera) persimilis (Cockerell) (Hymenoptera: Andrenidae)

Summary One aspect of behavioral ecology that has received considerable attention, especially by students of social insects, is the relative amount of energy invested by parents in the rearing of male versus female offspring. Sexual selection theory makes predictions about how individuals should allocate their total investment in the sexes. To test these predictions we must accurately quantify the relative cost incurred by a parent in the production of an average individual of either sex. Body weight ratios are the most common estimate of cost ratio, but the correspondence between offspring body weight and energetic investment on the part of the parent has rarely been determined. Calliopsis (Hypomacrotera) persimilis is a solitary, ground-nesting bee whose natural history makes it particularly convenient for studies of investment patterns and foraging behavior. Each day females construct and provision from 1 to 6 cells in linear, closely-spaced series. Each cell is provisioned with pollen from Physalis Wrightii flowers, which is collected on two or three foraging trips. However, the temporal sequence in which two- and three-trip foraging bouts occur is not random. Females invariably begin each day provisioning cells with three trips worth of pollen and usually switch to provisioning the latter cells of the day with just two trips worth of pollen. The sex of the offspring within the same co-linear series of cells also varies non-randomly — female offspring predominate in the first cells of each series and male offspring in the latter cells. The correspondence between the number of foraging trips to provision a cell, the total time spent foraging, and offspring sex was determined for 36 cells. The data indicate a close, though not absolute, relationship between the number of foraging trips and the sex of the offspring: males usually received two trips of pollen, though some received three, whereas female offspring invariably received three trips worth of pollen. A number of potential estimates of the relative cost of female and male offspring production were calculated. Estimates of the cost ratio based on the amount of time spent foraging, adult dry body weight, and pollen ball dry weight all give similar values. Female offspring receive an energetic investment of from 1.3 to 1.5 times that of males. These results support the use of adult dry body weight ratios in the estimation of cost ratios.     

Local mate competition and precise sex ratios in Telenomus fariai (Hymenoptera: Scelionidae), a parasitoid of triatomine eggs

Telenomus fariai is a gregarious endoparasitoid of the eggs of several species of Triatominae (Hemiptera) with a high degree of sibmating: males fertilize their sisters inside the host egg before emergence or emerge first and copulate with their sisters as these emerge. Our results show that, when laying alone, T. fariai behaves adaptively, minimizing offspring mortality and conforming to the prediction of local mate competition (LMC) theory by laying a single male, which is sufficient to fertilize all the sisters. When more than one wasp was placed with one host, sex ratios still conformed to LMC predictions but, despite the decreasing number of eggs laid per wasp, clutch size could not be completely adjusted to avoid mortality. This is not surprising, as superparasitism is rare in the field. Offspring production was independent of the contacts between conspecifics but was affected by the number of mothers laying on a single host egg. The sex of the progeny was precisely determined: a female produced one male per clutch when laying on both unparasitized or previously parasitized hosts. On the other hand, a mother produced less daughters when superparasitizing. Under crowded conditions, the number of eggs laid per female wasp and per host decreased as the number of mothers increased. Developmental mortality also increased with the number of T. fariai eggs per host, determining a maximum of approximately 14 emerged adults. Host resources per individual affected male and female adult size with similar intensity, and male adult mortality was slightly higher than that for females. These results, and previous findings, suggest that T. fariai attains Hamiltonian sex ratios by laying one male and a variable number of females, and that the detection of chemical marks left by conspecifics provides information on the number of foundresses sharing a patch. Received: 4 February 2000 / Received in revised form: 19 April 2000 / Accepted: 20 May 2000     

Assured fitness returns favor sociality in a mass-provisioning sweat bee,Megalopta genalis (Hymenoptera: Halictidae)

Assured fitness returns models for the evolution of sociality emphasize the selective value of ensuring that offspring receive adequate parental care to reach maturity. If a member of a social group dies, it can accrue returns on investment in offspring through the efforts of surviving social partners. We provide evidence that in the mass-provisioning, facultatively social sweat bee Megalopta genalis, adult presence in the nest throughout brood development provides protection from ant predation. Nests with adults present were well protected, and brood in nests with adults removed suffered higher predation. Females in observation nests showed effective defensive behavior against experimentally introduced ants, and bees in natural nests repulsed naturally occurring ant raids. Megalopta nest architecture and behavior are such that the brood of several cooperating females can be defended with little additional cost relative to solitary nesting. The benefits of cooperative defense may favor group living in mass provisioning bees. Our observations and experiments suggest that parental care throughout brood development can be adaptive in mass provisioning species, supporting the predictions of assured fitness returns models.     

Feeding opportunities of larval and juvenile cod (Gadus morhua) in a Greenlandic fjord: temporal and spatial linkages between cod and their preferred prey

Feeding of fish depends on a spatial and temporal match with prey, and since larval and juvenile feeding can be highly selective, their preferences for given prey sizes and taxa should be considered when quantifying the actual availability of potential prey. We investigated the diet and prey preferences of the early-life stages of Atlantic cod (Gadus morhua) to quantify the availability of prey during a spring-summer season in a West Greenlandic fjord. We hypothesized that abundances of larval and juvenile cod at size were synchronized to optimal availability of preferred prey in space and time. The present analysis is based on nine cruises each covering 5 stations visited between 24 May and 5 August 2010 comparing zooplankton abundance, cod gut content and distribution patterns. Cod 4–25 mm in length preferred prey of about 5 % of their own length. During ontogeny, their preferences changed from calanoid nauplii towards Pseudocalanus spp. and Calanus spp. copepodites. The larvae/juvenile had an exceptionally high dietary contribution from cladocerans, which were highly preferred by cod larger than 9 mm, while the abundant Metridia longa and the non-calanoid copepods contributed less. These findings stress the importance of focusing on abundance of preferred prey when assessing the actual prey availability to young fish. We found a spatio-temporal overlap between cod and their preferred prey, and observations suggest that advection of both zooplankton and cod contributed to this overlap. Hence, the larval feeding opportunities might be sensitive to climate-related changes affecting the circulation patterns in this fjord.     

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.     

Discrimination between nestmate and non-nestmate kin by social wasps (Polistes fuscatus,Hymenoptera: Vespidae)

Summary The threespine stickleback,Gasterosteus aculeatus is a voracious cannibal of both its own eggs and those of conspecific neighbours. Females, but not males, can distinguish their progeny from those of other fish and attacked alien eggs more frequently. In experiments to examine nest raiding, females that initiated raids on nests resulting in cannibalism were the first females to spawn in the reconstructed nest. These results support the hypothesis that cannibalism by females may be adaptive in situations where intense female competition for male guardians occurs.     

Attraction and learning in mate-finding by solitary bees,Lasioglossum (Dialictus) figueresi Wcislo and Nomia triangulifera Vachal (Hymenoptera : Halictidae)

Summary Males of the solitary sweat bee, Nomia triangulifera, patrol over large areas where thousands of females emerge, searching for receptive females. The daily operational sex ratio is strongly male-biased. Males contact dead, frozen (untreated) females more frequently than they contact females which were washed in hexane, showing that olfactory cues are utilized in mate-finding. A major source of female sex pheromone is in the head. Male pouncing on females is temporally non-random, indicative of group stimulation. Bioassays show that newly emerged females are more attractive to males than are older pollen-collecting females. Female odors are individually distinctive, based on male responses, and there is much variation among females in their attractive properties. Male responses to female odors suggest that learning is important for mating in natural populations. In contrast, the following hypotheses are unlikely to account for the observed behavior: (1) dissipation of female odors; (2) site learning and avoidance behavior by males; (3) decay of male motivation; or (4) male-produced repellents effective against other males. Laboratory and field studies show that female Lasioglossum figueresi produce individually-distinctive odors, which are attractive to males. There is considerable inter-individual variation among females in their attractiveness to males among sexually immature females. Male responses to female odors decay over the course of the presentation, suggesting the importance of learning in natural populations, although several alternatives could not be tested.     

Arsenic (As), antimony (Sb), and lead (Pb) availability from Au-mine Technosols: a case study of transfer to natural vegetation cover in temperate climates

Soils from old Au-mine tailings (La Petite Faye, France) were investigated in relation to the natural vegetation cover to evaluate the risk of metals and metalloids (Pb, As, Sb) mobilizing and their potential transfer to native plants (Graminea, Betula pendula, Pteridium aquilinum, Equisetum telmateia). The soils are classified as Technosols with high contamination levels of As, Pb, and Sb. The single selective extractions tested to evaluate available fraction (CaCl₂, acetic acid, A-Rhizo, and DTPA) showed low labile fractions (<5 % of bulk soil contents), but still significant levels were observed (up to 342.6 and 391.9 mg/kg for As and Pb, respectively) due to the high contamination levels of soils. Even at high soil contaminations (considered as phytotoxic levels for plants), translocation factors for native plants studied are very low resulting in low concentrations of As, Sb, and Pb in their aerial part tissues. This study demonstrates the important role of (1) native plant cover in terms of “stabilization” of these contaminants, and (2) the poor effectiveness of extraction procedures used for this type of soil assemblages, i.e., rich in specific mineral phases.