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.     

Social foraging in honey bees: how nectar foragers assess their colony's nutritional status

Summary A honey bee colony operates as a tightly integrated unit of behavioral action. One manifestation of this in the context of foraging is a colony's ability to adjust its selectivity among nectar sources in relation to its nutritional status. When a colony's food situation is good, it exploits only highly profitable patches of flowers, but when its situation is poor, a colony's foragers will exploit both highly profitable and less profitable flower patches. The nectar foragers in a colony acquire information about their colony's nutritional status by noting the difficulty of finding food storer bees to receive their nectar, rather than by evaluating directly the variables determining their colony's food situation: rate of nectar intake and amount of empty storage comb. (The food storer bees in a colony are the bees that collect nectar from returning foragers and store it in the honey combs. They are the age group (generally 12–18 day old bees) that is older than the nurse bees but younger than the foragers. Food storers make up approximately 20% of a colony members.) The mathematical theory for the behavior of queues indicates that the waiting time experienced by nectar foragers before unloading to food storers (queue length) is a reliable and sensitive indicator of a colony's nutritional status. Queue length is automatically determined by the ratio of two rates which are directly related to a colony's nutritional condition: the rate of arrival of loaded nectar foragers at the hive (arrival rate) and the rate of arrival of empty food storers at the nectar delivery area (service rate). These two rates are a function of the colony's nectar intake rate and its empty comb area, respectively. Although waiting time conveys crucial information about the colony's nutritional status, it has not been molded by natural selection to serve this purpose. Unlike signals, which are evolved specifically to convey information, this cue conveys information as an automatic by-product. Such cues may prove more important than signals in colony integration.     

Influence of three sediment types on copper toxicity to the polychaete Neanthes arenaceodentata

Adult male Neanthes arenaceodentata were exposed to 0.10±0.015 mg l^-1 copper in the seawater of a continuous-flow bioassay system in the presence of a sand, a mud, a mixture of the sand and mud, and no sediment, to assess the influence of sediment type on Cu-induced mortality. The sediment type did influence mortality. The time to 50% mortality was 7.8 days without sediment, 36.5 days with sand, 54.5 days with the mixture, and 50.0 days with mud. There was no threshold Cu body burden that caused death. The mean Cu concentration per gram of N. arenaceodentata (24 h after death) without sediment was 270 g, in sand 994 g, in the mixture 1047 g, and in mud 1464 g. The differences in the toxic responses are discussed.     

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.     

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

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

Grazing and ingestion rates of nauplii,copepodids and adults of the marine planktonic copepod Calanus helgolandicus

The average grazing and ingestion rates of all stages of the marine planktonic copepod Calanus helgolandicus (Calanoida) from nauplius stage IV to adults were measured experimentally at 15°C in agitated cultures. The chain-forming diatom Lauderia borealis and the unarmoured dinoflagellate Gymnodinium splendens were offered as food. The food concentrations were close to natural conditions and ranged from 36 to 101 g of organic carbon per liter. The medium body weights expressed in g of organic carbon of almost all larval stages raised at 49 g C/1 were identical with the weight of the same stages caught in the Pacific Ocean off La Jolla, California, USA. In a log-log system, grazing and ingestion rates increased almost linearly with increasing body weight. Grazing rates ranged from 4 to 21 ml/day/nauplius stage IV to 286 ml to 773 ml/day/female. Ingestion rates increased from 0.2 g to 0.8 g C/day/nauplius stage IV to 18 g to 69 g C/day/female. Grazing and ingestion rates per unit body weight decreased gradually with increasing body weight. The daily ingested amount of food decreased from 292 to 481% of the body weight (g C) of nauplius stage V to 28–85% of the body weight of adult females. Grazing and ingestion performances of all stages increased with increasing particle size. Grazing rates decreased and ingestion rates increased with increasing food concentrations. The published data on food intake of the different age groups of C. helgolandicus show that the young stages of herbivorous planktonic copepods can play a major part in the consumption of phytoplankton in the sea due to their high grazing and ingestion rates.     

Why are bumble bees risk-sensitive foragers?

Summary In a controlled laboratory experiment, we re-examined the question of bumble bee risk-sensitivity. Harder and Real's (1987) analysis of previous work on bumble bee risk aversion suggests that risk-sensitivity in these organisms is a result of their maximizing the net rate of energy return (calculated as the average of expected per flower rates). Whether bees are risk-sensitive foragers with respect to minimizing the probability of energetic shortfall is therefore still an open question. We examined how the foraging preferences of bumble bees for nectar reward variation were affected by colony energy reserves, which we manipulated by draining or adding sucrose solution to colony honey pots. Nine workers from four confined colonies of Bombus occidentalis foraged for sucrose solution in two patches of artificial flowers. These patches yielded the same expected rate of net energy intake, but floral volumes were variable in one patch and constant in the other. Our results show that bumble bees can be both risk-averse (preferring constant flowers) and risk-prone (preferring variable flowers), depending on the status of their colony energy reserves. Diet choice in bumble bees appears to be sensitive to the target value a colony-level energetic requirement. Offprint requests to: R.V. Cartar     

Division of labor during honey bee colony defense

Summary Some worker honey bees respond to major disturbances of the colony by flying around the assailant and possibly stinging; they are a subset of the bees involved in colony defense. These defenders have an open-ended age distribution similar to that of foragers, but defensive behavior is initiated at a younger age than foraging is. Behavioral and genetic evidence shows that defenders and foragers are distinct groups of older workers. Behaviorally, defenders have less worn wings than foragers, suggesting less flight activity. Genetically, defenders differ in allozyme frequencies, demonstrating different subfamily composition from foragers in the same colony. They also differ in allozyme frequencies from guards in the same colony, providing further evidence for division of labor associated with colony defense. We use this information to develop a model for honey bee colony defense involving at least two distinct groups of workers and we propose that the non-guard defenders be called soldiers, due to their important role in colony defense.Offprint requests to: M.D. Breed     

Dominance and fertility in a functionally monogynous ant

Summary Colonies of some leptothoracine ants may contain several inseminated but sterile females in addition to a single, fertile queen (functional monogyny). We here report the first observations on the behavior of these supernumerary females in the nearctic ant Leptothorax sp.A, a species belonging to the L. muscorum complex.In four colonies, each with up to eight intermorphic females, ritualized or openly aggressive interactions between individuals were observed, similar to those among workers of some other leptothoracine ants. The responses of individual females during encounters with nestmates apparently reflect the existence of linear dominance hierarchies. In each colony, the highest ranking individual was fed and groomed significantly more often than other females, and was the only one to oviposit after hibernation and to become fully physogastric during the first weeks of spring. When these -females were removed from three colonies, several other females started to lay eggs. However, in each colony only the highest ranking individual remained fertile; the others either were pushed out of the nest and finally killed by the workers, or their ovaries degenerated again.Aggressive interactions among females may also play an important role in the foundation of new colonies either by inducing intermorphs to leave the maternal nest to found new colonies solitarily or by inducing colony fission. Offprint requests to: J. Heinze at his new address     

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.     

Lack of social entrainment of free-running circadian activity rhythms in the Australian sugar glider (Petaurus breviceps: Marsupialia)

Summary Social influence on circadian activity rhythms was investigated in the nocturnal Australian marsupial Petaurus breviceps. The activity of two and two was recorded electroacoustically and observed by an IR television camera in LD 12:12 (10¹:10^-1lx) and in LL (10^-1lx) when housed isolated and in pairs (+). In LD-entrained animals the average duration of locomotion, orientation movements, grooming and total activity is influenced by social housing, and individually different activity patterns are harmonized. In constant illumination conditions, however, the members of a pair are not mutually synchronized and free-run with different circadian periodlengths. Therefore social influence on the LD-entrained activity rhythm in Petaurus should be interpreted as social masking rather than direct influence on the circadian system.     

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     

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.     

The role of male vs male interactions in maintaining population dialect structure

Summary Nonmigratory populations of Whitecrowned Sparrows in coastal California exhibit dialects in territorial male songs that are stable in space and time. By field playback experiments, we tested a prediction from the hypothesis that male aggressive interactions prohibit mixing of song dialects. Playback of the home dialect Clear song to territorial males singing the Clear dialect resulted in less response than that given to playback of the neighboring Buzzy dialect. Response to both Buzzy and Clear dialect songs by target Clear males, however, was greater than that given to the Bodega dialect recorded 55 km away (Figs. 2 and 3). We conclude that these results are consistent with the hypothesis that male-male aggressive interactions play an important role in reducing dialect mixing.     

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.     

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.     

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.     

Behavioral and life-history components of division of labor in honey bees (Apis mellifera L.)

Variability exists among worker honey bees for components of division of labor. These components are of two types, those that affect foraging behavior and those that affect life-history characteristics of workers. Variable foraging behavior components are: the probability that foraging workers collect (1) pollen only; (2) nectar only; and (3) pollen and nectar on the same trip. Life history components are: (1) the age the workers initiate foraging behavior; (2) the length of the foraging life of a worker; and (3) worker length of life. We show how these components may interact to change the social organization of honey bee colonies and the lifetime foraging productivity of individual workers. Selection acting on foraging behavior components may result in changes in the proportion of workers collecting pollen and nectar. Selection acting on life-history components may affect the size of the foraging population and the distribution of workers between within nest and foraging activities. We suggest that these components define possible sociogenic pathways through which colony-level natural selection can change social organization. These pathways may be analogous to developmental pathways in the morphogenesis of individual organisms because small changes in behavioral or life history components of individual workers may lead to major changes in the organizational structure of colonies. Correspondence to: R.E. Page, Jr.     

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.     

Carotenoids in the tiger prawn Penaeus esculentus during ovarian maturation

Female Penaeus esculentus Haswell were collected by 15 to 20 min duration trawls during 1990. Carotenoids were analysed in the digestive gland, abdominal muscle, the remainder of the body (hereafter called integument) and ovary of prawns in Stage 2 through Stage 4 (fully mature) of maturation. The only oxycarotenoids (xanthophylls) identified were astaxanthins or astaxanthin esters; occasionally low levels of -carotene were detected in the digestive gland. The concentrations of astaxanthin monoesters (AM) and diesters (AD) were highest, with only minor amounts of free astaxanthins (Ast), except in the maturing ovaries, where free astaxanthins predominated (up to 80% of the total carotenoid). Of the total carotenoid, 82 to 94% was in the integument, but at maturity the digestive gland contained 10.7±3.4% and the ovary 5.6±0.9% of the total carotenoid. Only the ovary increased in mass during maturation, reaching up to 5.2% of total prawn mass. During this period, digestive gland concentrations of AM, AD and Ast all increased (tota 20 to 120 g g^-1); levels in the muscle and integument varied little throughout maturation (total 0.4 and 100 g g^-1, respectively); ovary AM levels remained low throughout (1.5 to 1.2 g g^-1), AD increased from only 2 to 5 g g^-1, but Ast increased from 2 to 34 g g^-1. Apart from the ovary, AM concentrations were the most variable. In common with other decapod Crustacea, the maturing ovary of P. esculentus contained high levels of carotenoids, indicating that these may have an important role in early development. The natural diet of P. esculentus includes a variety of carotenoids, but except for a little -carotene, the digestive gland, where absorption occurs, contained astaxanthins, with only an occasional trace of -carotene. This suggests that the conversion of dietary carotenoids to astaxanthin occurs soon after ingestion.