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.     

Reproductive competition in queenless honey bee colonies (Apis mellifera L.)

Previously we reported that there are subfamily differences in drone production in queenless honey bee colonies, but these biases are not always explained by subfamily differences in oviposition behavior. Here we determine whether these puzzling results are best explained by either inadequate sampling of the laying worker population or reproductive conflict among workers resulting in differential treatment of eggs and larvae. Using colonies composed of workers from electrophoretically distinct subfamilies, we collected samples of adult bees engaged in the following behavior: true egg laying, false egg laying, indeterminate egg laying, egg cannibalism, or nursing (contact with larvae). We also collected samples of drone brood at four different ages: 0 to 2.5-h-old eggs, 0 to 24-h-old eggs, 3 to 8-day-old larvae, and 9 to 14-day-old larvae and pupae. Allozyme analyses revealed significant subfamily differences in the likelihood of exhibiting egg laying, egg cannibalism, and nursing behavior, as well as significant subfamily differences in drone production. There were no subfamily differences among the different types of laying workers collected from each colony, suggesting that discrepancies between subfamily biases in egg-laying behavior and drone production are not due to inadequate sampling of the laying worker population. Subfamily biases in drone brood production within a colony changed significantly with brood age. Laying workers had significantly more developed ovaries than either egg cannibals or nurses, establishing a physiological correlate for the observed behavioral genetic differences. These results suggest there is reproductive conflict among subfamilies and individuals within queenless colonies of honey bees. The implications of these results for the evolution of reproductive conflict, in both queenright and queenless contexts, are discussed.     

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     

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.     

Sperm utilization pattern in the honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis>)

Queen honeybees (Apis mellifera) mate with a large number of drones on their nuptial flights. Not all drones contribute equally to the queens offspring and the queens utilization pattern of spermatozoa from different drones has an important impact on the genetic composition of the colony. Here we study the consequences of sperm use for the fitness of the queens mates with microsatellite DNA-fingerprinting. Eight queens were instrumentally inseminated with semen of six or seven drones. Each drone contributed either 0.5 µl or 1.0 µl semen, respectively, and we analyzed both the impact of the insemination sequence and the amount of semen on the sperm utilization. Our data show no significant effect of the insemination sequence but a strong impact of the semen volume of a drone on the frequency of his worker offspring in the colony. This effect was not linear and the patriline frequencies of the drones contributing larger semen volumes are disproportionately enhanced. If these observations are also valid for natural matings, drone honeybees should maximize the number of sperm but not apply specific mating tactics to be first or last male in a mating sequence.Communicated by R. PageAn erratum to this article can be found at     

Environmental Accumulation of Airborne Fluorides in Romania

The nature and extent of pollution from an aluminium smelter and a fertiliser factory in Romania were studied. These are large industrial complexes, and both types of industry are known to release fluorides into the atmosphere. In grass samples collected from around the aluminium smelter, the maximum fluoride levels were found to be 4023mgkg^–1 and 162mgkg^–1 in unwashed and washed grass samples respectively, and 89mgkg^–1 in soils. For the fertiliser factory, the maximum levels in washed grasses were found to be 207mgkg^–1, and 11mgkg^–1 in the soils. In both locations, these maximum values were obtained in samples collected from within 200m of the factory limits, and compare with regional background levels of less than 10mgkg^–1 for grasses and 2mgkg^–1 for soils. The high fluoride levels of fluoride in the grasses are sufficient to give cause for concern for the effects that these could have on the local population and on grazing animals.     

Temporal variation and distribution of trace metals in freshwater and fish from Calabar River, S.E. Nigeria

An investigation on the abundance and distribution of trace metals (Fe, Cu, Zn, Mn, Cr, Cd and Pb) in water, and nine species of fish samples from Calabar river was carried out in 1992. The concentrations of iron (6000–7240gl^–1), zinc (4910–7230gl^–1), and cadmium (3–7gl^–1) showed moderate pollution while those of copper (420–630gl^–1), manganese (23–48gl^–1), chromium (<10–20gl^–1) and lead (<1–10gl^–1) in water were well below WHO permissible levels. Significant seasonal changes (0.001p0.25) were obtained for iron, copper, zinc, manganese and cadmium in water. Furthermore, iron, zinc and cadmium showed statistically significant spatial changes (0.005p0.10). Of the nine fish species studied, no statistically significant relationship between body weight and the concentrations of the metals was observed. The concentrations of the metals per mean total body weight apparently decreases in the order Fe>Zn>Cu>Mn>Pb>Cd=Cr and were within the limits that were safe for consumption.     

Arsenic tissue concentration of immature mice one hour after oral exposure to gold mine tailings

A potentially high bioavailability of arsenic in gold mine tailings from a site in northern California has been suggested by solubility studies. To help address this issue, an in vivo dosing study was conducted using 12dayold Swiss Webster mouse pups (n=8/group). A sample of sizefractionated mine tailings from the site (<20m particle size, 691g g^–1 arsenic) was prepared as an aqueous suspension and administered by gavage in a volume that provided 4mg As/kg body weight. The control group received the same volume of a commercial soil (1g g^–1 As) of similar particle size (<60m). No mortality or toxic signs were noted in either group. Tissue samples were collected 1h after gavage, freezedried, microwavedigested and analysed for arsenic by ICP/MS (detection limit 2ng As g^–1 dry weight). Arsenic concentrations (ng As g^–1 dry weight) in tissues from the pups who received mine tailings were significantly higher than in control tissues. The mean elevation in arsenic concentration was highest in the liver (3364% of control, p<0.0001), followed by blood (818 of control, p<0.0001), skin (207% of control, p=0.07), and brain (143% of control, p<0.0001). The carcass arsenic concentration (excluding the GI tract, liver, brain and skin) was 138 of control (p=0.02). The data indicate uptake of arsenic from weathered mine tailings by the immature mouse pups after oral exposure.     

Environmental Impacts of a Secondary Lead Smelter in Landskrona, Southern Sweden

Scandinavia has one secondary lead smelter that recycles lead from approximately 85% of used car batteries in Scandinavia and which has been active since the 1940s. The smelter, situated in Landskrona, has undergone a comprehensive clean up programme during the last decade, during which time production has doubled, while at the same time discharges of dust and lead to the atmosphere have decreased.Top and depth soil samples were taken on a 0.5km×0.5Km grid throughout the city of Landskrona, which covers an area of approximately 15km². Samples were analysed by ICPAES for a number of elements including Pb, Zn, Cu, Cd, As, Sb and Hg. Road dust samples from selected sites were collected and similarly analysed. Blood samples were taken from 37 volunteer schoolchildren (aged 8–11) from two schools in Landskrona. House dust samples were taken from each child's home. Soil samples were taken from homes which had gardens, public and school play areas. Elevated heavy metal concentrations were found in close proximity to the secondary lead smelter, and this soil enrichment influences the whole of the town, modified to some extent by the prevailing wind. The smelter does not influence the soil lead concentration at distances greater than 3.5km, where the soil reflects the background value for the area.Road dust samples also show decreases in lead concentrations with distance from the smelter. The average level of lead in house dust was considerably lower than that found in Birmingham, UK. Blood lead levels in the child population ranged from 1.5–5.1gdl^–1, with a mean of 3.05gdl^–1, showing a distinct decrease from those measured in 1978–82. No significant difference in blood lead concentrations with distance of the home from the smelter, nor between attenders at the two schools was revealed in the limited number of children studied.     

Aqua regia extractable trace elements in surface soils of venezuela

Surface soils (0–15 cm) were sampled at 10–20 km intervals along two transects in Venezuela. One (1162 km, 70 samples) ran west to east parallel with the Caribbean coastline, the other (920 km, 92 samples) ran south to north from the frontier with Brazil to the Caribbean shore. Sampling took place in both a wet and a dry season. Trace metals were extracted from dried, sieved (<2 mm) soil with boiling aqua regia followed by analysis by ICP or flame AAS. Metal values did not differ significantly between the two seasons and dates were averaged. Geometric mean values for the west–east transect were: Cr=41.5, Cu 17.9, Cs 3.6, Li=13.9, Mn=294, Ni=21.3, Pb=17.4, Sr=39.4, V=60.4 and Zn = 83.7g g^–1, respectively. Similarly, for the south–north transect Cr=21.3, Cu=4.3, Cs=1.1, Li=2.0, Mn=55.7, Ni=4.4, Pb=6.1, Sr=13.3, V=28.2 and Zn=16.7g g^–1, respectively. A classification of samples by lithology showed surface soil composition to be related to rock composition. Metal values were low in the soils in the south of the country, in the Guyana highlands (Gran Sabana). Low Zn contents were prevalent. Lead contents were affected by roadside fallout from vehicles using leaded petrol except that high Pb contents of soils in the Gran Sabana were of more complex origin.     

Growth and survival of Mytilus edulis larvae exposed to low levels of dibutyltin and tributyltin

Two studies were conducted to observe effects of dibutyltin (DBT) and tributyltin (TBT) on larvae of Mytilus edulis for an exposure period of 25 d. Endpoints for evaluation were shell growth and mortality measured at 33 d. Larvae were cultured in a new laboratory assay chamber in a recirculating static test. The control, 2, 20, and 200 g/l DBT-treated populations had mean shell lengths of 527, 523, 417, and 180 m, respectively. Survival was 1% for the 200 g/l DBT-treated population, but ranged from 73 to 83% for controls, 2, and 20 g/l treatments. The no-observed-effect concentration (NOEC) was 2 g/l for DBT, while the lowest-observed-effect concentration (LOEC) was 20 g/l. The chronic toxicity value was 6.3 g/l. In the TBT bioassay, mean shell lengths for the control, 0.006, 0.050, and 0.130 g/l-treated populations were 565, 437, 385, and 292 m, respectively. Control survival was 74%, whereas TBT-treated populations survival ranged from 52 to 58%. The NOEC for TBT was 0.006 g/l TBT and the LOEC was 0.050. A chronic toxicity value of 0.017 g/l was calculated. The results of this study indicated that the toxicity of DBT was less than that of TBT. It was concluded that shell length was inversely related to exposure level in both DBT and TBT bioassays. In this study, we have observed TBT effects at lower exposure levels in the laboratory than previously reported, and also report the first data for DBT effects on mussel larvae.     

Environmental Geochemistry of Soils and Waters of Susaki Area, Korinthos, Greece

Soil and water samples were collected from the Susaki area of Korinthos and analysed for heavy metals in order to evaluate their environmental impact. The geology of the studied area includes ultrabasic rocks and Neogene and Quaternary deposits whereas magnesite veins are found within the ultrabasic rocks. In the north part of the studied area post volcanic emissions of H₂S, CO₂ and H₂O vapor continue to the present day. All the samples were analysed for heavy metals by the ICP method. The element ranges (in g g^–1) for soil samples are: Cu 11–63, Pb 5–256, Zn 21–604, Ni 183–2665, Co 12–124, Mn 456–1434, As 5–104, Sr 44–730, V 21–84, Cr 163–2346, Ba 48–218, Zr 3–41, Y 3–13. The metals Pb, Zn, Ni, Co, Cr, Fe, Cu, Mn, As and Sr are enriched in the Susaki soils. The element ranges for water samples are: Cu 65–103ppb, Pb<10ppb, Zn<5ppb, Ni 21–163ppb, Co 2–12ppb, As<30ppb, Cr<20ppb, Ba 36–785ppb, Sb<10ppb, W<10ppb, Bi<30ppb, Mn 0.0–0.9 g g^–1, Fe 0.01–0.22 g g^–1, Na 843–3076 g g^–1, K 98–278 g g^–1, Si 39–65 g g^–1, P 0.1–0.2 g g^–1. There is a natural pollution of soils with elevated concentrations of Ni, Co, Mn, Fe and Cr due to the presence of ultrabasic rocks. Another natural case of As pollution of soils is due to the volcanic activity and the geothermal field in the area. The geochemical data of ground waters and also the ¹⁸, D data showed a mixing in different proportions between sea water and meteoric water.     

Annual cycles in worker size of the seed-harvester antVeromessor pergandei (Hymenoptera: Formicidae)

Summary The seed-harvester antVeromessor pergandei Mayr is primitively polymorphic; workers are monophasically allometric. There is a distinct annual cycle in mean worker body size that replicates across colonies and habitats (Fig. 1); this cycle occurs through alteration of the worker size distribution (Fig. 2). There is little, if any, morphspecific task specialization by workers suggesting worker size variance is a colony-level adaptation permitting maintenance of a large and constant worker force during periods of resource fluctuation. Smaller workers appear in the foraging force following the triple crunch of reduced seed availability, reduced favorable times to forage, and alate production during winter months. Adult and startingV. pergandei colonies exhibit strong intraspecific territoriality, suggesting the selective advantage for maintenance of a large and constant worker force. Such selective pressures may have provided the initial variance in worker size distributions that led subsequently to specialized castespecific task performance in more distinctly polymorphic ant species.     

Circadian rhythmicity of locomotor behavior and oviposition in female Teleogryllus commodus

Summary Locomotor activity of the Teleogryllus commodus is under circadian control, with _LL=25.3 h and _DD=23.4 h. In LD 12:12 h, running occurs either exclusively during darkness (57%), mainly in the dark phase (35%), or with substantial activity peaks in both phases (8%). For oviposition, _LL>24 h and _DD<24 h; in LD 12:12 h, 80% of the studied deposited their eggs mainly during the light phase and 20% primarily during darkness. In the discussion, a temporal correlation between stridulation, spermatophore formation, and locomotion, and oviposition is established, which serves to make an encounter and mating between the sexes as likely as possible.Dedicated to Prof. Dr. M. Gersch on the occasion of his 70th birthday     

Metabolic diversity in epibiotic microflora associated with the Pompeii wormsAlvinella pompejana andA. caudata (Polychaetae: Annelida) from deep-sea hydrothermal vents

Specimens of alvinellid polychaetes (Alvinella pompejana Desbruyères and Laubier, 1980 andA. caudata Desbruyères and Laubier, 1986) and their tubes were sampled from deep-sea hydrothermal vents at 13°N from the manned submersible Nautile during the Hydronaut cruise (October to November 1987) on the East Pacific Rise. Samples were subjected to bacterial analysis aboard the mother ship Nadir to detect bacteria involved in the nitrogen and sulphur cycles, in non-specific heterotrophic processes, and displaying resistance to selected heavy metals. Cultures were incubated at different temperatures under atmospheric and in situ (250 atm) pressures. Bacterial growth was observed in enrichment cultures for most metabolic types screened. Heavy-metal-resistant bacteria were also detected in many samples. No filamentous bacterial form was observed in the cultures. The results demonstrate a high metabolic diversity in episymbiotic flora, indicating that the worm (A. pompejana orA. caudata), its tube and its epiflora represent a complex micro-ecosystem.     

Genetic variation and biochemical systematics in the marine bryozoan Alcyonidium mytili

For many years the taxonomic position of the marine ctenostome bryozoan Alcyonidium mytili Dalyell (1848) has been in doubt, with some authorities regarding it as being conspecific with A. polyoum (Hassall). We have set out to resolve this problem using enzyme electrophoresis to examine the genetics of the population structure in the two species. The results show beyond reasonable doubt that sympatric populations of the two species are indeed from quite separate and non-interbreeding gene pools. An examination of various allopatric populations shows a new species of A. mytili to exist sympatrically with the other at Langstone Harbour, Hampshire, England. This species can be readily distinguished on morphological as well as genetic ground. Samples of A. mytili from Guernsey also show substantial genetic differences from other populations examined. It is tentatively concluded that this population also represents a separate gene pool and consequently should be recognised as a third species of A. mytili.     

Investigation of potential gene flow limitation of behavioral adaptation in an aridlands spider

Summary This study investigates the possibility that gene flow underlies the apparently maladaptive behavior of a riparian woodland population of the desert spider Agelenopsis aperta with respect to territorial, foraging, and antipredatory behaviors. I found that other local populations of A. aperta in the vicinity of the riparian woodland habitat are prey-limited and exhibit an aridlands phenotype (high aggressiveness in competitive interactions over energy-based territories and a lack of discrimination among potential prey types). The riparian woodland population deviates from surrounding populations in the area in that prey are abundant and this population shows a mixture of aridlands and riparian (low aggressiveness towards conspecifics and discrimination of prey profitability) phenotypes. Electrophoretic analyses of population subdivision in the area indicate that significant levels of gene flow have occurred, at least, sometime in the past. Drift fence analyses of spider movement futher indicate that there is marked unidirectional movement of spiders each year from the more arid habitats into the riparian woodland. Experimental manipulation of gene flow and predation pressure demonstrates that gene flow restricts adaptation in this habitat: one generation of predation pressure in the absence of gene flow is sufficient to cause a marked shift in spider behavior towards the expected riparian phenotype.     

Effects of light intensity on nitrate and nitrite uptake and excretion by Chaetoceros curvisetus

Michaelis-Menten uptake kinetics were observed at all light intensities. With constant illumination, the V_max and K₁ in nitrate uptake over the natural light intensity range of 0 to 2000 E were 0.343 g-at NO₃–N(g)^-1 at protein-N h^-1 and 26 E, respectively. Nitrate uptake was inhibited at higher light intensities. The K_s for nitrate uptake did not vary as a function of light intensity remaining relatively constant at 0.62 g-at NO₃–N 1^-1. With intermittent illumination, the V_mzx for light intensity in nitrate uptake over a light intensity range of 0 to 5000 E was 0.341 g-at NO₃–N(g)^-1-at protein-N h^-1. No inhibition of nitrate uptake was observed at higher than natural light intensities. Chaetoceros curvisetus will probably never experience light inhibition of nitrate uptake under natural conditions.     

Primary production in Dumbell Bay in the Arctic Ocean

Photosynthesis, chlorophyll, inorganic nutrients, and related factors were measured throughout a productive season in a small coastal inlet of the Arctic Ocean. Significant production was confined to 5 m for a very limited time period beacause of both light and nutrient limitations. Maximum chlorophyll was 8.2 mg Chl a m^-3 and maximum gross and net photosynthesis rates were 830 and 550 mg C m^-2 d^-1. Annual gross and net photosynthesis is estimated at about 12 and 9 g C m^-2 respectively. The effect of light stimulation on assimilation is described, sources of coastal zone nutrient enrichment are considered, and discrepancies between gross and net photosynthesis are discussed with reference to the growth characteristics of the phytoplankton population.Bigelow Laboratory for Ocean Sciences. Contribution No. 76008     

Alternative mating tactics and the law of Diminishing Returns: The Satellite Threshold Model

Summary The mating systems of many species comprise at least 2 alternative male mating tactics — broadly termed Dominant and Satellite tactics here. But, there exists no satisfactory conceptual framework in which to explain their co-occurrence and persistence.A Dominant male's ability to copulate is a positive, but decelerating, function of either his territory's attractiveness or his own attractiveness to females. This Diminishing Returns Effect affords mating opportunities for Satellite males associated with the most attractive sites (or males). A graphical Satellite Threshold Model, analogous to the Polygyny Threshold Model (Orians 1969), is developed to predict quantitatively the presence or absence of Satellite male tactics.Five predictions of the Model are supported quantitatively by data from dragonfly and anuran social systems and, more generally, by evidence from a wide variety of species.The Model should apply both to circumstances in which males are able to alternate between tactics and the instances in which males make an irrevocable tactical decision.