Regulation of worker reproduction in bumblebees (Bombus terrestris): workers eavesdrop on a queen signal

In the annual bumblebee Bombus terrestris, the onset of queen-worker conflict over male production is seasonally and socially constrained. Workers will do better if they start to reproduce (the so-called competition phase) only after ascertaining that larvae are committed to gyne development but before the season ends because they gain more by rearing sister-gynes than their own sons. Here, we tested two nonmutually exclusive hypotheses as to what triggers the onset of worker reproduction: Workers can directly monitor larval development and/or workers eavesdrop on the queen signal that directs gyne development. Exposing workers to gyne larvae through a double mesh did not advance the competition phase compared to control colonies. However, when workers, but not the queen, were allowed contact with gyne larvae, both the competition phase and gyne production were advanced. Thus, while larvae do not emit a volatile pheromone that discloses their developmental route, the physical contact of workers with such larvae triggers early competition phase. However, workers exclusively exposed to worker larvae (colonies prevented from producing gyne larvae) started to reproduce at the same time as control colonies. Replacing the resident queen with an older queen (from gyne-rearing colonies) advanced the competition phase, irrespective of worker age. The results are consistent with the hypothesis that workers eavesdrop on the queen pheromones. This is adaptive because it allows workers a broader time-window for reproduction and thus to gain fitness from rearing both sister-gynes and sons before the season ends without affecting colony development.     

Chemistry of the postpharyngeal gland secretion and its implication for the phylogeny of IberianCataglyphis species (Hymenoptera: Formicidae)

Summary A comparative morphological and chemical study of six endemic species ofCataglyphis from the Iberian Peninsula:C. ibericus, C. rosenhaueri, C. hispanicus, C. humeya, C. velox andC. floricola and the Moroccan speciesC. bombycinus is described. The morphological study relied primarily on genitalia characteristics, whereas the chemical study concentrated on the postpharyngeal gland constituents. Cladograms based on the morphological and chemical data were performed using Ward's method. The dendrogram based on morphological features revealed that the IberianCataglyphis can be classified into three species groupsalbicans, altisquamis andemmae. The same pattern occurred when the dimethylalkanes constituents of the postpharyngeal gland were utilised as character states, with a slight displacement of species within thealtisquamis group. However, when the complete hydrocarbon blend was utilised major discrepancies in the dendrograms occurred.Cataglyphis velox proved to be very similar toC. bombycinus, whereasC. floricola clustered with the other two species of thealtisquamis group. Based on the geographical distribution and paleontological data (Tinaut 1993) it is assumed thatC. floricola recently invaded the Iberian Peninsula. Based on the chemical findings we postulate that chemical character displacement occurred inC. floricola as a result of its sympatry withC. velox after the former colonized the Iberian Peninsula. We further discuss the possible reason for the different dendrograms obtained when only the dimethylalkanes are considered and its implication for the communicative role of the postpharyngeal gland secretion in these ants.     

Global warming and flowering times in Thoreau's Concord: a community perspective

As a result of climate change, many plants are now flowering measurably earlier than they did in the past. However, some species' flowering times have changed much more than others. Data at the community level can clarify the variation in flowering responses to climate change. In order to determine how North American species' flowering times respond to climate, we analyzed a series of previously unstudied records of the dates of first flowering for over 500 plant taxa in Concord, Massachusetts, USA. These records began with six years of observations by the famous naturalist Henry David Thoreau from 1852 to 1858, continued with 16 years of observations by the botanist Alfred Hosmer in 1878 and 1888-1902, and concluded with our own observations in 2004, 2005, and 2006. From 1852 through 2006, Concord warmed by 2.4 degrees C due to global climate change and urbanization. Using a subset of 43 common species, we determined that plants are now flowering seven days earlier on average than they did in Thoreau's times. Plant flowering times were most correlated with mean temperatures in the one or two months just before flowering and were also correlated with January temperatures. Summer-flowering species showed more interannual variation in flowering time than did spring-flowering species, but the flowering times of spring-flowering species correlated more strongly to mean monthly temperatures. In many cases, such as within the genera Betula and Solidago, closely related, co-occurring species responded to climate very differently from one another. The differences in flowering responses to warming could affect relationships in plant communities as warming continues. Common St. John's wort (Hypericum perforatum) and highbush blueberry (Vaccinium corymbosum) are particularly responsive to changes in climate, are common across much of the United States, and could serve as indicators of biological responses to climate change. We discuss the need for researchers to be aware, when using data sets involving multiple observers, of how varying methodologies, sample sizes, and sampling intensities affect the results. Finally, we emphasize the importance of using historical observations, like those of Thoreau and Hosmer, as sources of long-term data and to increase public awareness of biological responses to climate change.     

Park isolation in anthropogenic landscapes: land change and livelihoods at park boundaries in the African Albertine Rift

Regional Environmental Change - Landscapes are changing rapidly in regions where rural people live adjacent to protected parks and reserves. This is the case in highland East Africa, where many...     

In-nest environment modulates nestmate recognition in the ant<Emphasis Type="Italic"> Camponotus fellah</Emphasis>

Multiple behavioral and chemical studies indicate that ant nestmate recognition cues are low-volatile substances, in particular hydrocarbons (HCs) located on the cuticular surface. We tested the hypothesis that in the ant Camponotus fellah, nest environment, in particular nest volatile odors, can modulate nestmate-recognition-mediated aggression. Workers were individually confined within their own nest in small cages having either a single mesh (SM = limited physical contact permitted) or a double mesh (DM = exposed to nest volatiles only) screen. Individual workers completely isolated outside their nest (CI) served as control. When reintroduced into a group of 50 nestmates, the CI workers were attacked as alien ants after only 2 weeks of separation, whereas the SM workers were treated as nestmates even after 2 months of separation. Aggression towards DM ants depended on the period of isolation. Only DM workers isolated for over 2 months were aggressed by their nestmates, which did not significantly differ from the CI nestmates. Cuticular HC analyses revealed that the profile of the non-isolated ants (NI) was clearly distinct from that of CI, SM and DM ants. Profile differences matched the aggressive response in the case of CI ants but were uncorrelated in the case of SM or DM ants. This suggests that keeping the ants within the nest environment affected nestmate recognition in additional ways than merely altering their HC profile. Nest environment thus appears to affect label–template mismatch by modulating aggressive behavior, as well as the direction at which cuticular HCs diverged during the separation period.     

Dufour's gland secretion of the queen honeybee (Apis mellifera): an egg discriminator pheromone or a queen signal?

The role of Dufour's gland secretion as an egg discriminator pheromone was reevaluated by simultaneously exposing workers to two combs, one containing queen- or worker-laid eggs and the second containing treated or untreated worker-laid eggs. Treatments included extracts of Dufour's gland secretion as well as the synthetic esters that were identified in the secretion. Policing was clearly detected both in queenright and queenless colonies by the swift removal of worker, but not of queen eggs. However, neither the glandular secretion nor its synthetic ester constituents were able to protect worker-born eggs from policing. Treated worker eggs were removed significantly faster than queen eggs, and at the same rate as non-treated worker eggs. These results are not consistent with the hypothesis that the secretion serves as an egg-marking pheromone. Chemical analyses of the queen abdominal tips revealed the presence of Dufour's esters, indicating that the glandular secretion oozes out and spreads over the cuticle around the genital chamber. However, contamination while ovipositing may also explain the minute amounts of these esters that were detected on the egg surface. Dufour's gland caste-specific composition suggests that in queens it may constitute a signal that plays a role in queen-worker interactions. Attraction bioassays revealed that the queen secretion, but not that of workers, is very attractive to workers. When applied either on a glass slide or on another worker, a retinue formed around the "surrogate queen". We conclude that Dufour's gland secretion constitutes part of a complex queen signal that is the basis for the social integrity of the honeybee colony.     

Real-time ozone mapping using a regression-interpolation hybrid approach, applied to Tucson, Arizona

Real-time ozone (O3) maps, intended for public access and mass media, are generated from spatially interpolating (i.e., kriging) sparse monitoring data and are typically characterized by over-smoothed surfaces that inadequately represent local-scale spatial patterns (e.g., averaged over 1 km2). In this paper, a hybrid regression-interpolation methodology is developed to enhance the representation of local-scale spatiotemporal patterns with an application to Tucson, Arizona. The mapping of local patterns is enhanced with pre-interpolation regression modeling of local-scale deviation-from-mean variability, preserving variation in the monitor data that is ubiquitous across the modeling domain (i.e., the areal mean). The model is trained on several years of deviation-from-mean hourly O3 data, and predictor variables are developed using theoretically and empirically derived proxy regression variables. The regression model explains a significant proportion of the variation in the data (r2 = 0.54), with an average error of 7.1 ppb. When augmented with the areal mean, the r2 of the pre-interpolation model increases to 0.847. Model residuals are then spatially interpolated to the extents of the modeling domain. Final concentration estimate maps are the summation of areal mean, regression, and spatially interpolated surfaces, preserving absolute values at monitor locations.     

Reproductive plasticity in bumblebee workers (<Emphasis Type="Italic">Bombus terrestris</Emphasis>)—reversion from fertility to sterility under queen influence

Worker sterility in the bumblebee Bombus terrestris is conditional and is linked to the social development of the colony. Workers refrain from reproducing or overtly challenging the queen until gyne production has initiated, at the so-called competition point (CP). It is not known whether this behavior is hard-wired or workers show reproductive plasticity. It also remains unclear whether worker reproductive decision is under queen and/or worker control. In this study, we tested worker reproductive plasticity in an attempt to assess whether and under which conditions worker sterility/fertility are reversible. We introduced egg-laying workers into colonies with different social structures for 1 week then monitored their reproductive status. We revealed a remarkable reproductive plasticity in the introduced workers that was social-condition-dependent. In the presence of a pre-CP queen, the introduced workers reverted to sterility, whereas in the presence of a post-CP queen, such workers remained egg-layer. Reversion to sterility does not occur when direct contact with the queen is prevented, as the introduced workers remained egg-layer in the queenright colonies with a confined queen. Egg-laying workers that were introduced into queenless colonies mostly maintained their fertility regardless of colony social phase. This shows that worker transition from cooperative to selfish behavior is reversible depending on the social context.     

Dufour’s gland pheromone as a reliable fertility signal among honeybee (Apis mellifera) workers

When a colony becomes queenless and without the possibility of requeening, honeybee workers initiate reproduction and lay male eggs about a week later. Assays in which two bees were confined in a small arena revealed that they establish a reproductive dominance hierarchy, i.e., one worker demonstrates greater ovarian development than her paired bee. Reproductive dominance is independent of relatedness, and can be established between full sisters, cousins, or random nestmates. A social environment, however, is obligatory, as singly housed bees fail to develop ovaries on the same time scale. Allowing varying degrees of social interactions between the paired bees revealed that olfaction of volatile bee compounds, as well as tactile communication, seem to provide the necessary social environment. Ovarian development was accompanied by the production of queen-like Dufours gland secretion in these workers. Especially notable was the increase in the queen-like esters. This increase was tightly linked to ovarian development and not necessarily to the dominance status of the bees in the pair. Thus, the occurrence of queen-like esters can serve as a reliable fertility signal. Advertising ovarian status may recruit helper workers with less developed ovaries (and which are less likely to successfully reproduce before colony breakdown) to assist their nestmates and thereby gain inclusive fitness. Revealing the role of Dufours gland secretion as a fertility signal adds another dimension to our understanding of how queen pheromones operate. The mandibular-gland secretion is a good predictor of dominance hierarchy, being correlated with false-queen characteristics but not fertility, whereas Dufours gland secretion is a good predictor of fertility but not dominance hierarchy.Communicated by R.F.A. Moritz