Self-organizing pattern formation on the combs of honey bee colonies

Summary A characteristic pattern of brood, pollen, and honey develops on the combs of a honey bee colony, consisting of three distinct concentric regions — a central brood area, a surrounding rim of pollen, and a large peripheral region of honey. That the pattern is consistent and well-organized suggests its adaptive value for the colony, yet the mechanism of pattern formation has not been elucidated. Two hypotheses are presented. The blueprint (or template) hypothesis suggests that there are particular locations specified for the deposition of eggs, pollen and honey, i.e., the pattern develops as a consequence of the bees filling in the comb according to the orderly arrangement latent in the blueprint. An alternative is the self-organization hypothesis: pattern emerges spontaneously from dynamic interactions among the processes of depositing and removing brood, pollen and honey, without a plan specifying spatial relationships. Computer simulation of the self-organization hypothesis demonstrates how the colony-level pattern can emerge and how, using only local cues and simple behavioral rules, the bees can create an overall, global pattern of which they have no concept.     

Environmental policy with collective waste disposal

Centralized collection and disposal is an integral component of waste management strategies for many solid and liquid wastes, and carbon capture and storage is currently being considered for gaseous waste. In this paper we show how collective waste disposal systems introduce essential changes in the design of optimal environmental policy. Absent collective disposal, an optimal environmental policy imposes relatively stringent regulations on polluters in regions where local environmental damage functions are “high”; however, under collective waste disposal, the optimal environmental policy level increases monotonically over distance from the disposal site, and this is true irrespective of the degree of spatial heterogeneity in local environmental damage functions. We characterize the optimal spatial pattern of environmental policy levels under collective waste disposal and identify optimal membership size for waste disposal networks comprised of homogeneous producers.     

Brood sorting by ants: distributing the workload over the work-surface

Summary Leptothorax unifasciatus ant colonies occupy flat crevices in rocks in which their brood is kept in a single cluster. In artificial nests made from two glass plates sandwiched together, designed to mimic the general proportions of their nest sites in the field, such colonies arrange their brood in a distinct pattern. These patterns may influence the priority with which different brood are tended, and may therefore influence both the division of labour and colony demography. Different brood stages are arranged in concentric rings in a single cluster centred around the eggs and micro-larvae. Successively larger larvae are arranged in progressive bands away from the centre of the brood cluster. However, the largest and oldest brood items, the prepupae and pupae, are placed in an intermediate position between the largest and most peripheral larvae and the larvae of medium size. Dirichlet tessellations are used to analyze these patterns and show that the tile areas, the area closer to each item than its neighbours, allocated to each type of item increase with distance from the centre of the brood cluster. There is a significant positive correlation between such tile areas and the estimated metabolic rates of each type of brood item. The ants may be creating a domain of care around each brood item proportional to that item's needs. If nurse workers tend to move to the brood item whose tile they happen to be within when they have care to donate, they may apportion such care according to the needs of each type of brood. When colonies emigrate to new nests they rapidly recreate these characteristic brood patterns.     

Why do Varroa mites invade worker brood cells of the honey bee despite lower reproductive success?

Varroa jacobsoni reproduces both in drone and worker brood cells of honey bees, but in drone cells reproductive success is higher than in worker cells. A simple model using clonal population growth as a fitness measure has been developed to study the circumstances under which specialization on drone brood would be a better strategy than reproduction in both types of cell. For European Apis mellifera, the model suggests that if mites have to wait less than 7 days on average before they can invade a drone cell, specialization on drone brood would be a better strategy. This is close to the estimated waiting time of 6 days. Hence, small differences in reproductive success in drone and worker cells and in the rate of mortality may determine whether specialization on drone brood will be promoted or not. In European A. mellifera colonies, Varroa mites invade both drone and worker cells, but specialization on drone brood cells seems to occur to some extent because drone cells are more frequently invaded than worker cells. In the parasite-host association of V. jacobsoni with African or Africanized A. mellifera or with A. cerana, the mites also invade both drone and worker cells, but the mites specialize on drone brood for reproduction since a large percentage of the mites in worker brood do not reproduce. Only in the parasite-host association of Euvarroa sinhai, a mite closely resembling V. jacobsoni, and A. forea is specialization complete, because these mites only invade drone brood.     

The behaviour of ant transporters at the old and new nests during successive colony emigrations

Colonies of the ant Temnothorax albipennis improve their collective performance over successive emigrations (Langridge et al. Behav Ecol Sociobiol 56:523–529, 2004, Behav Ecol Sociobiol 62:447–456, 2008). Here, by analysing the performance of individual transporters (workers that carry the brood, queen and a proportion of adults), we investigate whether they spend less time at the old and new nests during repeated emigrations. Transporters expedited choosing and picking up brood items at the old nest and depositing them in the new nest. Such improvements were not associated with adult transport. Generally, when carrying brood items, but not when carrying adults, transporters visited several locations in the new nest before depositing them. Transporters did not interact with other adults when depositing brood items. Consequently, reductions in depositing times are the sum of time savings made by individual transporters. By contrast, transporters spent most time interacting with other adults before picking up brood items at the old nest. As the frequency of these interactions did not decline, we suggest the behaviours of interacting adults were modified in a way that hastened their completion. Thus, reductions in picking-up times probably occur because of time saved during interactions.     

Patterns of brood division and an absence of behavioral plasticity in a neotropical passerine

Studies of parental behavior in various habitats provide an opportunity to gain insight into how different environments may mold strategies of parental care. Brood division by parents has been hypothesized to occur facultatively within and among species. Brood division occurs when each parent cares for specific offspring within a brood. We studied brood division in a neotropical passerine, the western slaty antshrike (Thamnophilus atrinucha). Our results present a unique picture of a highly specialized example of avian brood division. Division was a fixed behavioral pattern in the population studied: all broods divided by fledging and remained divided during the entire post-fledging period. Brood division before fledging, a previously unreported phenomenon, occurred in 40% of nests observed. Parents that preferentially fed a certain offspring (defined as their focal offspring) in the nest fed the same individual after fledging. Each parent fed only its focal offspring in broods of one and two. The male parent cared for the heavier offspring and the first offspring to leave the nest. Siblings were segregated spatially during the time of highest predation risk. These observations suggest that a consistently high risk of predation on offspring has favored initial spatial segregation and inflexibility of brood division behavior in this species. Factors other than predation risk alone may explain the observed patterns of long-term, perfect brood division. Because high predation is common and relatively predictable in the tropics, selection for fixed brood division may be stronger in tropical birds than in the temperate zone.     

Manipulation of nest-box density affects extra-pair paternity in a population of blue tits (<Emphasis Type="Italic">Parus caeruleus</Emphasis>)

We tested the effect of manipulation of breeding density on the occurrence of extra-pair paternity in a blue tit (Parus caeruleus) population during two consecutive years. In a homogeneous oak forest, nest-box manipulation provided a high density plot (plot A, 1.10 and 1.32 pairs/ha) and a low density plot (plot B, 0.43 and 0.46 pairs/ha). Microsatellite analysis on 91 broods revealed a higher proportion of extra-pair paternity in broods in plot A (mean of 17.2%) than in plot B (mean of 11.4%). A correlative approach showed that the proportion of extra-pair young in broods was affected by the number of breeding neighbours within 100 m around the nest-box, by the distance to the nearest breeding neighbour and by an additional plot effect. However, the nearest neighbours accounted for only 39.3% of extra-pair paternities and distance to extra-pair fathers was significantly higher than the nearest neighbour distance in both plots. This implies that the effect of density on the occurrence of extra-pair paternities is associated with active female choice to enhance the brood fitness. Although there were more extra-pair young in broods when density was high, the number of extra-pair fathers did not increase and stayed close to one. We suggest that density increases the cost of mate guarding by males, thereby increasing the possibility for females to solicit extra-pair paternities to the cuckolding male they have chosen. Finally, we discuss why correlatives approaches do not always show evidence for an effect of breeding density on extra-pair paternity occurrence.Communicated by M. Soler     

Learning,retention and coding of nest-associated visual cues by the Australian desert ant, <Emphasis Type="Italic">Melophorus bagoti</Emphasis>

A variety of social insects use visual cues for homing. In this study, we examine the possible factors affecting the learning and retention of nest-associated visual cues by the Australian desert ant Melophorus bagoti and the manner in which such cues are encoded by foraging ants. We placed four prominent cylindrical landmarks around a nest and trained foragers from that nest to a food source. Ants were tested with the landmark array in a distant testing field after (1) a known number of exposures to the landmarks (1, 3, 7 or 15 trials, spread over a period of 1 day, 2 days or ≥3 days) and (2) after a known period of delay (0, 24, 48, 96 or 192 h). The results show that a combination of an increase in training trials and an increase in number of training days affected the acquisition of landmark memory. Moreover, once the landmarks were learnt, they became a part of long-term memory and lasted throughout the ants’ foraging lifetime. To examine visual cue encoding behaviour, ants trained under similar conditions for 4 days were tested with (1) an identical landmark array, (2) landmarks of the same size used in training, but placed at twice the distance from each other, and (3) landmarks whose dimensions were doubled and placed at twice the distance from each other. In conditions (1) and (3), the ants searched extensively at the centre of the four landmarks, suggesting that, similar to the Saharan ant (genus Cataglyphis) and the honeybee, M. bagoti too uses a snapshot to match the view of the landmarks around the nest. But contrary to the snapshot model, in condition (2), the ants did not search extensively at the centre of the landmarks, but searched primarily 0.5 m from the landmark, the distance from each landmark to the nest during training. We discuss how various search models fare in accounting for these findings.     

Dispersion of Tellina tenuis from Kames Bay,Millport, Scotland

The dispersion of Tellina tenuis da Costa in the laboratory was analysed by the Clark and Evans nearest-neighbour test and by the Kolmogorov-Smirnov one-sample test. The dispersion was random, with a slight tendency toward aggregation that was independent of density. In the field, the dispersion was analysed by the Clark and Evans nearest-neighbour test and by the X² approximation to Fisher's coefficient of dispersion. The dispersion was again random, this time with a slight tendency toward spacing out that was independent of density. The tendency toward aggregation displayed in the laboratory was independent of the dispersion pattern shown at the start of the experiment, and also unaffected by the edge of the container. The apparent randomness suggests that T. tenuis is primarily a suspension feeder, but may be a deposit feeder under certain environmental conditions.     

Differences in olfactory sensitivity and behavioral responses among honey bees bred for hygienic behavior

Honey bees, Apis mellifera L., bred for hygienic behavior uncap and remove diseased and mite-infested brood. We hypothesized that within a colony bred for hygienic behavior, there would be differences in olfactory sensitivity among bees of the same age. We predicted that bees that initiate the behavior by perforating and uncapping brood would have greater olfactory sensitivity to the odor of the diseased brood, and would be better able to discriminate between odors of healthy and diseased brood, compared to bees that complete the behavior by removing the uncapped brood from the cells. Electroantennogram recordings of 15- to 21-day-old bees from three colonies demonstrated that bees collected while uncapping dead brood had significantly greater olfactory sensitivity to all concentrations of diseased brood odor compared to bees collected while removing brood. Proboscis-extension reflex discrimination conditioning demonstrated that 15- to 21-day-old bees collected while uncapping discriminated significantly better and generalized significantly less between the odors of diseased and healthy brood compared to bees collected while removing, when the odor of diseased brood was rewarded, but not when the odor of healthy brood was rewarded. Bees collected while uncapping brood that had been pierced with a pin had significantly less olfactory sensitivity than bees collected while uncapping freeze-killed brood, most likely because the pierced brood had greater stimulus intensity. Initiation of hygienic behavior depends on the olfactory sensitivity of the bee and stimulus intensity of the abnormal brood. Differential olfactory sensitivity and responsiveness among hygienic bees could lead to the apparent partitioning of the behavior into uncapping and removing components.Communicated by R.F.A. Moritz     

The effect of helpers on artificially increased brood size in sociable weavers (Philetairus socius)

In cooperatively breeding birds, the presence of helpers is expected to increase the reproductive success of the breeding pair. However, some studies fail to find this effect. A positive effect of helpers may be restricted to cases in which a breeding pair has a poor likelihood of raising the entire brood on its own, as would be the case under stressful environmental conditions or with enlarged brood sizes. We conducted brood size manipulations in a cooperative breeder, the sociable weaver, Philetairus socius, to investigate the relationship between the difficulty of raising nestlings and the effort and impact of helpers. Overall, sociable weavers did not work harder to raise the enlarged broods. However, the presence of helpers significantly increased the feeding rates at enlarged nests, but not controls. This was insufficient to prevent generalised brood reduction in enlarged broods, whether attended by pairs alone or with helpers. Nonetheless, the presence of helpers was associated with decreased nestling mortality and an increase in the numbers of young fledged. Our results suggest that groups are better able to respond to the needs of enlarged broods than pairs alone and that the presence of helpers has a beneficial effect on overall reproductive success.Communicated by J. Dickinson     

Costs and benefits of surplus offspring in the lesser kestrel (Falco naumanni )

Lesser kestrels (Falco naumanni) lay clutches which appear excessive as only 3% of them yield as many young as eggs laid. Four hypotheses may explain the adaptive value of producing surplus eggs: (1) the bet-hedging hypothesis assumes that the environment varies unpredictably and surplus eggs serve to track uncertain resources; (2) the ice-box hypothesis suggests that surplus offspring serve as a reserve food during a period of shortage; (3) the progeny choice hypothesis says that parents produce surplus offspring in order to choose these with higher fitness; and (4) the insurance-egg hypothesis proposes that extra eggs are an insurance against the failure of any egg. To test the significance of this strategy in the lesser kestrel, an experiment manipu-lating brood size at hatching was carried out over 2 years, with good and bad feeding conditions. The experiment consisted of adding a chick to experimental broods where one egg failed to hatch or removing a randomly selected chick from experimental broods where all eggs had hatched. Independently of annual food availability, pairs with brood sizes reduced by one chick fledged more nestlings than pairs with brood size equalling their clutch sizes. Body condition of young was also better in the former group, but only in 1993 (a high-food year). Independently of year, mean local survival of parents with complete broods at hatching was lower than for parents raising reduced broods. These results supported only the insurance-egg hypothesis which says that surplus eggs may be an insurance against the failure of any egg, but parents may suffer reproductive costs when all eggs hatch. Received: 17 January 1997 / Accepted after revision: 27 April 1997     

Family structure and variation in reproductive success in blackbirds

In avian families, some offspring are rendered unequal by parental fiat. By imposing phenotypic handicaps (e.g., via asynchronous hatching) upon certain of their offspring and not others, parents structure the sibship into castes of advantaged “core” offspring and disadvantaged “marginal” offspring that results in an asymmetric sibling rivalry. Here, I show how this family structure scales up to population level reproductive consequences. In a 17-year study of red-winged blackbirds (Agelaius phoeniceus), I show that year-to-year variation in the number of surviving offspring is driven primarily by variation in the number of marginal offspring at hatching and their posthatching survival. Clutch size, core brood at hatching, and fledging varied little from year to year and had little direct effect on year-to-year variation in total brood size at fledging; conversely, variation in the size of the marginal brood at hatching and at fledging was much greater. Marginal but not core brood size at hatching rose with mean clutch size; in years where parents laid larger average clutches they did so by adding marginal progeny. The mean posthatching survival of marginal offspring was always lower than that of core offspring in a given year, and there was no overlap in the distributions. The highest mean survival of marginal offspring across years fell below the lowest mean survival of core offspring; broods were deeply structured. There was an overall female bias among fledglings, and the sex ratio varied across years, with a higher proportion of the smaller female nestlings in years of below average reproductive success. Such variation was especially pronounced in the marginal brood where a higher incidence of brood reduction allowed greater potential for sex-biased nestling mortality. In years of the highest average reproductive success, the sex ratio in the marginal brood approached equality, whereas in years of the lowest average reproductive success, more than two thirds of 8-day-old nestlings were female. Structuring the brood into core and marginal elements allowed parents to modulate both offspring number and sex under ecological uncertainty with direct consequences for population-level reproductive success. They produced fewer and less expensive fledglings in below average years and more and more expensive fledglings in above average years.     

Brood dispersal and multiple central place foraging by Lapland longspur parents

Summary Lapland longspur chicks continued to be fed by their parents for 2 weeks after nest departure. Shortly after they left the nest, broods were divided evenly into two units each tended by a single parent. Female-tended brood units dispersed away from the nest at a faster average distance per day than those tended by males. The distance between offspring within a brood unit also increased as chicks got older. For the first 8–10 d after nest departure, parents were multiple central place foragers, making 1–8 foraging trips away from each dispersed chick (i.e. the central place) before moving on to feed another chick in a similar fashion. During the final 5–7 d before independence, chicks were quite mobile and followed parents on their foraging bouts. A simulation model shows that brood dispersal reduced total parental travel time per chick, primarily because each chick moved closer to a foraging site. By comparing models for a variety of brood division and foraging itinerary scenarios, we also show that multiple central place foraging by parents visiting different, separated brood units is less energetically expensive than other reasonable alternatives.Although brood dispersal is usually considered to be an adaptation for avoiding predation, it can also help parents reduce the energetic costs of parental care.     

Reproductive patterns of three intertidal salt-marsh gammaridean amphipods

Among three sympatric species of epibenthic amphipods found at different tide marks at Jamaica Bay, New York (USA), the length of time juveniles spend with the mother increases and the number of juveniles per brood decreases as tidal height increases. Each brood has two developmental periods: (1) the embryonic period, from ovulation to hatching; (2) the juvenile period, from hatching to emergence from the marsupium. Gammarus palustris, found at the high-tide mark, has a mean juvenile period of 1.7 days and a mean brood size of 12.4 offspring; G. mucronatus, found at mean-tide mark, has a mean juvenile period of 0.8 days and a mean brood size of 27.4 offspring; Melita nitida, found at low-tide mark, has a mean juvenile period of 0.5 days and a mean brood size of 30.0 offspring. Further, the range of days that a juvenile may emerge is widest for G. palustris (0 to 8 days after hatching) and narrowest for M. nitida (0 to 2 days).This work was submitted in partial fulfillment of the requirements for the Ph.D. at the City University of New York.     

Caste determination and differential diapause within the first brood of Halictus rubicundus in New York (Hymenoptera: Halictidae)

Summary The partially bivoltine, primitively eusocial sweat bee Halictus rubicundus produces two female castes, gynes and non-gynes, in its first brood in New York. Castes in this brood differentiate within the first few days of adult life, with gynes leaving the population to overwinter as early as mid-June (non-gynes further differentiate into replacement queen and worker subcastes, with older females typically dominant). Analysis of possible mechanisms of caste determination reveals that although gynes average significantly larger than non-gynes within the first brood, this appears largely due to a late mean emergence data coupled with an increase in the size of emerging females over the course of brood emergence, rather than a causal relationship. A strong correspondence between male abundance (relative to newly-emerged females) and the pattern of gyne production, along with data from dissections, suggests that females that mate when young become diapausing gynes, while those that do not mate promptly become non-gynes and do not diapause even though many mate later. Although alternatives to this simple mechanism cannot be ruled out entirely, it nevertheless offers profound implications for theoretical and empirical understanding of the evolutionary origins of the worker caste.     

Competition with flies promotes communal breeding in the burying beetle,Nicrophorus tomentosus

Communal breeding through nest-sharing may benefit cooperating individuals indirectly, in increased inclusive fitness, or directly, when environmental constraints reduce the fitness of solitary breeders. Burying beetles provide extensive parental care and can breed either in pairs or in larger groups of unrelated males and females. Parentage of communally-reared broods is usually shared but is skewed in favor of the individuals of each sex that provide longer care. Females provide care longer than males, and two females are more likely to remain together in the brood chamber than two males are. Flies and other burying beetles are the major competitors for carcasses and this study suggests that it is competition with flies that promotes communal breeding inNicrophorus tomentosus On medium-size carcasses (35–40 g) the presence or absence of oviposition by flies had a significant effect on the size of the brood reared, and on large carcasses (55–60 g) the number of beetles present, two or four, had a significant effect on brood size. On both medium and large carcasses, pairs rearing broods on flyblown carcasses had fewer young than pairs on clean carcasses or foursomes on flyblown carcasses. There was a strong trend for an interaction effect between number of beetles and competition with flies (Table 1). Duration of parental care was not affected by competition with flies except for that of the first male to depart, which provided care longer on flyblown carcasses (Table 2). Pairs and foursomes were equally able to defend the carcass and brood from conspecific intruders and from larger intrudingNicrophorus orbicollis (Table 3).     

Evidence of kin-selected tolerance by nestlings in a siblicidal bird

Behaviorally dominant members of blue-footed booby (Sula nebouxii) broods can effect siblicide by restricting access of subordinate siblings to parents providing food. In spite of their capacity for siblicide, dominant chicks permit subordinates to feed during short-term food shortage; in fact, the proportion of the food that the dominant takes is independent of the total amount delivered in older chicks. A model of optimal food distribution suggests that dominant chicks maximize their inclusive fitness with this pattern, rather than by satisfying their own food requirements and leaving what remains for the subordinate sibling. The indirect reproductive potential represented by a chick's sibling appears to have influenced the evolution of siblicidal brood reduction in this species.     

A GIS-based moving window analysis of landscape pattern in the Beijing metropolitan area,China

Quantifying landscape pattern and its change is essential for monitoring and assessment of ecological consequences of urbanization. Using the GIS-based land-use data for 2002, we combined moving window analysis along an urban-rural gradient and around the urban centre with landscape metrics to quantify the spatial pattern of urbanization in Beijing. The results of moving window analysis along the urban-rural gradient indicated that, for class-level metrics, the spatial pattern of urbanization could be quantified using landscape metrics, different land-use types exhibited distinctive spatial signatures, and, for landscape-level metrics, the increase in urbanization in the metropolitan Beijing region has resulted in dramatic increases in patch density (PD), edge density (ED), and patch and landscape shape complexity, and sharp decreases in the largest and mean patch size (MPS), agriculture land-use type, and landscape connectivity. The results of moving window analysis around the urban centre showed that the direction of urbanization could be quantified using the class-level metrics, and landscape-level metrics indicated similar results to gradient analysis. In general, moving window analysis showed that the increasingly urbanized landscape became compositionally more diverse, geometrically more complex, and ecologically more fragmented.