Division of labour in a crisis: task allocation during colony emigration in the ant Leptothorax unifasciatus (Latr.)

Division of labour during colony emigration is widespread in ants. An important problem is how tasks are allocated during colony movement from one nest site to another. The generally favoured view is that emigrations are organised by a minority group of individuals, which either work unusually hard at tasks (elites) or have the exclusive task of carrying out the emigration (moving specialists). Five consecutive emigrations of a Leptothorax unifasciatus (Latr.) colony showed that the number of transporters, i.e. the individuals that took an active part in the emigration by transporting brood and ants, was smaller than it would have been if allocation of this task was random during each emigration. However, single emigrations of another three colonies, for which the spatial distribution and behaviour of the workers had been observed for a week prior to the emigration, demonstrated that the transporters did not form a homogeneous group. They differed in their spatial positions and tasks before the emigration. There was also no evidence that transporters worked harder or less hard than their nestmates before the emigration. Therefore, the individuals which carry out emigrations in L. unifasciatus colonies appear to be neither moving specialists nor elites. We propose that task allocation during emigrations of L. unifasciatus colonies is based on a feedback mechanism that involves learning.     

Effects of worker genotypic diversity on honey bee colony development and behavior (Apis mellifera L.)

There have been numerous reports of genetic influences on division of labor in honey bee colonies, but the effects of worker genotypic diversity on colony behavior are unclear. We analyzed the effects of worker genotypic diversity on the phenotypes of honey bee colonies during a critical phase of colony development, the nest initiation phase. Five groups of colonies were studied (n = 5–11 per group); four groups had relatively low genotypic diversity compared to the fifth group. Colonies were derived from queens that were instrumentally inseminated with the semen of four different drones according to one of the following mating schemes: group A, 4 A-source drones; group B, 4 B-source drones; group C, 4 C-source drones; group D, 4 D-source drones; and group E, 1 drone of each of the A-D drone sources. There were significant differences between colonies in groups A-D for 8 out of 19 colony traits. Because the queens in all of these colonies were super sisters, the observed differences between groups were primarily a consequence of differences in worker genotypes. There were very few differences (2 out of 19 traits) between colonies with high worker genotypic diversity (group E) and those with low diversity (groups A-D combined). This is because colonies with greater diversity tended to have phenotypes that were average relative to colonies with low genotypic diversity. We hypothesize that the averaging effect of genotypic variability on colony phenotypes may have selective advantages, making colonies less likely to fail because of inappropriate colony responses to changing environmental conditions.     

Interspecific aggression in colonies of the slave-making ant Harpagoxenus sublaevis

Colonies of the slave-making ant, Harpagoxenus sublaevis, may simultaneously contain workers of several Leptothorax slave species. We observed aggressive interactions among slave-makers, between slavemakers and slaves, and among slaves in 11 mixed colonies. The first two types of aggression appear to be correlated with reproductive competition for the production of males. Aggressive interactions among slaves, however, occurred mainly between slaves belonging to different species. In two colonies, in which one slave species clearly outnumbered the other, the majority attacked and finally expelled all nestmates belonging to the minority species. Our observations thus suggest that in Harpagoxenus colonies a homogeneous colony odor is not always achieved and that heterospecific slaves may occasionally be mistaken for alien ants. Gas chromatographic analyses of ants from mixed colonies similarly show that cuticular hydrocarbon profiles may differ strongly between heterospecific nestmate slaves.     

Salinity tolerance of benthic estuarine diatoms as tested with a rapid polarographic measurement of photosynthesis

Measurements of net photosynthesis of benthic estuarine diatoms were made by polarographic registration of oxygen saturation. A measuring cell was constructed in which media with salinities of 2.0 to 100.7 were pumped over the algae between measurements. Diatoms from unialgal cultures and mixed populations from intertidal flats appeared to be highly tolerant of extreme salinities. During short-term exposures (20 min) the net photosynthesis of the algae did not drop below 70% of the initial values, within the salinity range 4.0 to 60.0. Prolonged exposure (up to 6 h) gave essentially the same results. Populations of benthic diatoms, sampled from field stations with mean salinities of about 30, 12, and below 5, showed only gradual differences in their tolerance of salinities between 2.0 and 33.7. Two species, Navicula arenaria and Nitzschia sigma, were cultured in media ranging in salinity from 8.0 to 45.0 and from 2.0 to 45, respectively. The tolerance to changing salinity was only slightly affected by the salinity of the medium in the preculture. The role of salinity in the production and distribution of intertidal diatoms is discussed.     

Nutrition studies in the nauplius larva of Calanus pacificus (Copepoda: Calanoida)

Nauplii of Calanus pacificus were raised on a mixture of algae. Details of the mouth-parts, such as denticles, labial palps and lobes, setations and structure of the masticatory teeth were examined by scanning electron microscopy (SEM). Under the experimental conditions (15°C and 300 gC l^-1), exponential growth coefficients for the period Nauplii II–VI were 0.179 for carbon and 0.228 for nitrogen. C:N ratios dropping from 5.1 to 4.7. Growth was isochronous, each stage lasting 1.5 days. Respiratory losses were 15 to 19.6% of body carbon daily. Nauplii raised on a given alga showed higher rates of ingestion in the presence of this food, compared to nauplii switched to other algae. Minimal threshold concentrations for feeding were found, depending on the size of the food offered and ranging from 5.8 gC I^-1 for Lauderia borealis (28.7 m spherical diameter) to 47.1 gC 1^-1 for Chlamydomonas sp. (11.0 m). Unlike the Copepodite I stage, Nauplii II–VI larvae were not able to ingest small cells such as Isochrysis galbana (4.3 m), or very large ones such as Ditylum brightwellii (47.5 m) at more than maintenance rations. Below the critical concentration for maximal feeding, ingestion was clearly dependent on size of the cells offered, but the size-dependent relationship was different for diatoms and non-diatoms. Filtering rates increased from a threshold concentration to a maximal rate at about 50 gC 1^-1, and decreased at higher concentrations. Critical concentrations ranged from 125 gC 1^-1 for L. borealis to 1000 gC 1^-1 for Chlamydomonas sp. Maximal daily rations ranged between 100 and 150% of body carbon.     

Population characteristics of the deep-sea lobsters Polycheles typhlops and Stereomastis sculpta (Decapoda: Polychelidae) in a bathyal mud community of the Mediterranean Sea

Distribution and abundance as a function of depth, and population parameters such as sex and size structure of the population and aspects of reproductive biology have been studied in the deep-sea polychelid lobsters Polycheles typhlops and Stereomastis sculpta. Samples were taken by otter trawl in the Balearic Sea, a deepsea basin in the northwestern Mediterranean Sea from 1985 to 1989. In both species the largest size classes were dominated by females. In S. sculpta, male and female sizes were very similar. In P. typhlops, ovigerous females and those with external spermatophores were 23 and 25 mm in carapace length (CL), respectively; males with external spermatophores were 17 mm in CL. In S. sculpta, ovigerous females and those with external spermatophores were 24 and 19 mm CL, respectively, and males with external spermatophores 19 mm CL. Highest densities of P. typhlops occurred along the middle slope at depths between 500 and 1000 m. Only small-sized individuals were found at the deepest depths sampled; some recruitment must therefore occur in waters much deeper than those usually inhabited by the adult population. The sex-ratio was 1:1 in most samples, but in some of the shallowest samples females predominated. The depth distribution range of S. sculpta was 981 to 2253 m: densities clearly increased with increasing depth. There were no apparent variations in size distribution as a function of depth. Since very few adult males and females were captured, the population in the survey area would seem to be comprised mainly of juveniles. As a function of depth, females were numerically dominant only in some of the shallowest samples taken in the distribution range of this species. There is strict habitat partitioning between the two species. In both species, the variation in the sex-ratio as a function of depth suggests differential migration between the sexes, probably related to egg incubation and hatching.     

On the evolution of male workers in the Hymenoptera

Summary The effects of brood sex-ratio (investment ratio) and the presence of laying workers on relatedness in the Hymenoptera are analyzed. It is shown that the conditions of brood composition that generate degrees of relatedness favorable to the evolution of eusocial type helping behavior among females will select against such helping behavior among males, and vice versa (see Fig. 1). Thus, societies in the Hymenoptera can be expected to have male workers or female workers, but not both. It is argued that the conditions leading to degrees of relatedness favorable to male helping are quite restrictive and unlikely to be met in haplodiploid species.The presence of laying workers is shown to lead to biasses in relatedness such that females may be selected to be workers even when the sex-ratio is male-biassed. This result sheds new light on a possible pathway to eusociality in the Hymenoptera. It is argued that offspring parasitism of the natal nest may have been important in the evolution of eusociality.     

Female defence polygyny in the bumblebee wolf,Philanthus bicinctus (Hymenoptera: Sphecidae)

Summary In the digger wasp Philanthus bicinctus the first adult males emerge about a week before the first females. The sex ratio of emerging wasps is near unity. Females appear to mate once and do so when they first start to nest, not when they first emerge.Males defend mating territories in a large area of female ground nests. Territoriality begins when females first start to nest and throughout the season coincides with the digging activities of females initiating rests. During the day territoriality occurs only when females are engaged in digging activities.Males compete for certain territory sites within the resting area; the most frequently occupied male territories are in areas where the density of nests is highest. About half of the males are excluded from nesting area territories. Some of these males defend areas among the flowers of snakeweed where females occasionally forage; this appears to be a loser tactic of males unable to obtain nesting area territories.Although there was no significant difference in the sizes of males on nesting area and snakeweed territories, within the nesting area larger males usually occupy territories with the higher nest densities.     

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).     

On the integration of individual foraging strategies with colony ergonomics in social insects: nectar-collection in honeybees

Summary We experimentally tested whether foraging strategies of nectar-collecting workers of the honeybee (Apis mellifera) vary with colony state. In particular, we tested the prediction that bees from small, fast growing colonies should adopt higher workloads than those from large, mature colonies. Queenright small colonies were set up by assembling 10 000 worker bees with approximately 4100 brood cells. Queenright large colonies contained 35 000 bees and some 14 500 brood cells. Thus, treatments differed in colony size but not in worker/brood ratios. Differences in workload were tested in the context of single foraging cycles. Individuals could forage on a patch of artificial flowers offering given quantities and qualities of nectar rewards. Workers of small colonies took significantly less nectar in an average foraging excursion (small: 40.1 ± 1.1 SE flowers; large: 44.8 ± 1.1), but spent significantly more time handling a flower (small: 7.3 ± 0.4 s ; large: 5.8 ± 0.4 s). When the energy budgets for an average foraging trip were calculated, individuals from all colonies showed a behavior close to maximization of net energetic efficiency (i.e., the ratio of net energetic gains to energetic costs). However, bees from small colonies, while incurring only marginally smaller costs, gained less net energy per foraging trip than those from large colonies, primarily as a result of prolonged handling times. The differences between treatments were largest during the initial phases of the experimental period when also colony development was maximally different. Our results are at variance with simple models that assume natural selection to have shaped behavior in a single foraging trip only so as to maximize colony growth. Offprint requests to: P. Schmid-Hempel     

Analysis of two genetic models for the innate components of colony odor in social Hymenoptera

Summary We propose two models for the inheritance of the innate components of colony odor in social Hymenoptera. Under the Individualistic model, individuals are hostile unless they share at least one allele at all colony-odor loci. Under the Gestalt model, colony-odor pheromones are transferred between individuals, resulting in a gestalt colony odor; colonies will not fuse unless they have the same genetic mix of workers. We analyze these models for the case of colonies founded by single, once-mated queens. The Gestalt model seems generally favored for most species, although some evidence suggests the Individualistic model may operate in some primitive ants. A single colony-odor locus is possible in the Gestalt model, but seems improbable in the Individualistic case. We propose a general inbreeding test to estimate the number of loci involved in colony odor.     

Functional reference in an alarm signal given during nest defence: seet calls of yellow warblers denote brood-parasitic brown-headed cowbirds

Field observations and model-presentation experiments have shown that yellow warblers (Dendroica petechia) produce seet calls preferentially in response to brood-parasitic brown-headed cowbirds (Molothrus ater). In this study, we investigated whether seet calls are functionally referential alarm calls denoting cowbirds by determining whether female warblers responded appropriately to seet calls in the absence of a cowbird, whether alarm calling by warblers varied with response urgency, and how warblers in a population allopatric with cowbirds responded to cowbird and avian predator models and seet playbacks. As a control, we presented chip calls, which are elicited by nest predators as well as by non-threatening intruders, but are not strongly associated with cowbirds. Yellow warblers responded differently to playbacks of seet than chip calls. To seet playbacks, almost 60% of females gave seet calls and rushed to sit in their nests, responses typically elicited by cowbirds, whereas these responses were given infrequently in response to chip calls. Yellow warblers seet called equally in situations that simulated low, medium and high risk of parasitism, which suggests that these calls did not vary with response urgency. In a population allopatric with cowbirds, seet calls were rarely produced in response to cowbird or avian nest predator models and never to seet playbacks. These results suggest that seet calls are functionally referential signals denoting cowbirds and that cowbird parasitism was a strong selective pressure in the evolution of functional referentiality in the seet call of yellow warblers.Communicated by W.A. Searcy     

The kin recognition system of carpenter ants (Camponotus spp.)

Summary The phenotypic cues that provide for rejection of non-nestmates by workers of the ant genus Camponotus could derive from any or all of four sources: (1) environmental odors; (2) the individual's own genetically-determined recognition pheromones or discriminators; (3) a gestalt or mixture of transferable discriminators, produced by each nestmate and absorbed by all; and/or (4) the discriminators of the queen applied to all nestmates. To test these hypotheses, four series of small experimental colonies were created: inter- and intraspecific mixed colonies containing queens, queenless worker groups, and pairs of worker groups between which a single queen was repeatedly switched. Intraspecific mixed colonies and queenless groups were further divided into groups receiving different diets. Aggression of workers in 165 experimental colonies was assayed in a total of 4064 neutral arena tests. Workers adopted into inter- and intraspecific mixed colonies with queens were highly aggressive to unfamiliar kin from pure colonies, independent of diet and of the proportion of different kin groups in the colony. However, queenless workers exhibited less aggression to unfamiliar kin than to non-kin, demonstrating the existence of worker discriminators. Diet differences slightly enhanced aggression among unfamiliar queenless kin. Non-kin sharing a switched queen were as unaggressive to one another as were sisters. The ability to adopt queenless workers between colonies gradually declined over 1–2 wks following their emergence from pupae. We propose a hierarchy of importance of cue sources in determining nestmate discrimination in small Camponotus colonies: Queen discriminators> worker discriminators>environmental cues. A flow-diagram model of social insect kin recognition, based on the phenotype matching concept of Holmes and Sherman (1983), is discussed.     

Effects of nitrogen and phosphorus enrichement on in vivo symbiotic zooxanthellae of Pocillopora damicornis

The reef coral Pocillopora damicornis (Linnaeus) was grown for 8 wk in four nutrient treatments: control, consisting of ambient, unfiltered Kaneohe Bay seawater [dissolved inorganic nitrogen (DIN, 1.0 M) and dissolved inorganic phosphate (DIP, 0.3 M)]; nitrogen enrichment (15 M DIN as ammonium); phosphorus enrichment (1.2 M DIP as inorganic phosphate); and 15 M DIN+1.2 M DIP. Analyses of zooxanthellae for C, N, P and chlorophyll a after the 8 wk experiment indicated that DIN enrichment increased the cellular chlorophyll a and excess nitrogen fraction of the algae, but did not affect C cell^-1. DIP enrichment decreased both C and P cell^-1, but the decrease was proportionally less for C cell^-1. the response of cellular P to both DIN and DIP enrichment appeared to be in the same direction and could not be explained as a primary effect of external nutrient enrichment. The observed response of cellular P might be a consequence of in situ CO₂ limitation. DIN enrichment could increase the CO₂ (aq) demand by increasing the net production per unit area. DIP enrichment could slow down calcification, thus decreasing the availability of CO₂ (aq) in the coral tissue.Hawaii Institute of Marine Biology Contribution No. 920     

The effect of male calling on female locomotor activity of Teleogryllus commodus

Summary The calling song of the -cricket, Teleogryllus commodus, attracts for mating. Both calling and -locomotor activity are under circadian control and the behaviors occur approximately at the same time. Daily playback of calling songs for 3–12 h in constant light failed to entrain -running. Instead, calling acts as an external stimulus which can release -locomotion at any time of the 24-h period, while the clock-induced activity continues to run freely. The stimulatory effect of the calling song causes flexible extension of -locomotor activity, but a subsequent restorative period is necessary. The ecological consequences of acoustical stimulation are discussed.     

Risk-taking by female ducks: intra- and interspecific tests of nest defense theory

We tested several predictions of nest defense theory by observing variation in flushing distance and probability of nest abandonment within and between six species of waterfowl. In these species, only the females incubate eggs and attend offspring. First, we examined whether flushing distance by females varied in relation to clutch size, stage of incubation, and time of season, after controlling for the number of visits made to nests by observers. Revisits by observers appeared to affect flushing distance by females for reasons unrelated to the relative value of the current clutch. We found that as incubation progressed, females allowed observers to approach more closely before flushing from the nest. In some species, females with larger clutches allowed closer approaches to nests before flushing which was also consistent with nest defense theory. In contrast, time of season (Julian date) did not relate to flushing distance for any species. When species were compared, we found that species with moderate to high yearly mortality and high reproductive output per breeding attempt (e.g., northern shoveler and blue-winged teal) were less likely to abandon nesting attempts and exhibited riskier behavior (remained at nests when approached closely by observers) than species that had lower yearly mortality (e.g., mallard). Our results show that flushing distance and patterns of nest abandonment by female ducks conform to several predictions of nest defense theory.Correspondence to: M.R.L. Forbes     

Sexual conflict in the house sparrow: interference between polygynously mated females versus asymmetric male investment

Summary The reproductive success of female house sparrows mated with polygynous males depended to some extent on the aid received from their mates. Polygynous males fed nestlings at the same rate as monogamous males although polygynous males gave aid almost exclusively to one of their mates (the preferred). As a consequence, the number and quality of young raised by preferred females were similar to those of monogamous females, whereas nonaided females experienced a reduction of clutch size, hatching success, and fledgling quality. Males aided the harem female that hatched her clutch earliest (in 11 out of 12 cases). However, harem females that laid eggs earliest (i.e., the potential preferred) lost eggs and/or nestlings more frequently than harem females that delayed laying. This resulted from competing females' infanticide suited for redirecting males' aid. All polygynously mated females delayed the start of a new clutch and reproduced fewer times per season than monogamous females. Consequently, monogamy seems to be the optimum mating situation for females; whereas polygynous males raised more young per year than monogamous males, interference between harem females reduced their maximum expected success.     

13C/12C ratios and the trophic importance of algae in Florida Syringodium filiforme seagrass meadows

Over 380 stable carbon isotope (¹³C) analyses made during 1981–82 showed that Syringodium filiforme Kutz seagrass meadows in the Indian River lagoon of eastern Florida have food webs based on algal rather than seagrass carbon. Seagrasses averaging approximately-8 were isotopically distinct from algae epiphytic on seagrass blades (X=-19.3) and particulate organic matter in the water column X=-21.6. ¹³C values of most fauna ranged between-16 and-22, as would be expected if food web carbon were derived solely from algal sources. These results counter the idea that seagrass detritus is the dominant carbon source in seagrass ecosystems. Two factors that may contribute to the low apparent importance of seagrass in the study area are high algal productivities that equal or exceed S. filiforme productivity and the high rates of seagrass leaf export from meadows.     

The evolution of cheating and selfish behavior

Summary We examine the necessary conditions for the spread of genes that determine selfish and cheating behaviors and the rate of spread of these genes through structured populations, in order to address the question of the invadability of altruistic systems by anti-social mutations. We find that, although cheaters always have a higher relative fitness than altruists within groups, population structures which permit the evolution of altruism also preclude invasion by anti-social mutations. These results are related to a discussion by Hamilton (1971) concerning the limits to the evolution of altruistic and selfish behaviors.     

Indoor and outdoor household dust contamination in Cincinnati, Ohio, USA

Despite some progress in reducing the average lead level in the USA, the streets of Cincinnati, Ohio, are still contaminated by heavy metals. High levels of heavy metals will have significant unequivocal ecological impacts and pose a potential health hazard. This study evaluates the level of heavy- metal contamination in household dust and examines its relationships with the external environment. Samples of outdoor and indoor dust were collected from middle-income residential homes in the Greater Cincinnati Metropolitan District and the metal content was analysed by inductively coupled plasma atomic emission spectrophotometry. Results showed that the mean levels of lead in outdoor and indoor dusts are 650 g g-1 and 377 g g-1 and the copper levels are 253 g g-1 and 510 g g-1, respectively. The median levels are 156 g g-1 and 139 g g-1 for outdoor and indoor lead dusts and 35 g g-1 and 124 g g-1 for outdoor and indoor copper dusts. The degree of contamination may be ascribable to the age of the dwelling unit and the neighbourhood, the time when the unit was last painted, the presence of pets and the type of space heating. In addition, car exhausts seems to be a possible source of contamination.