Social evolution in the genusHalictus: a phylogenetic approach

Summary Halictine bees exhibit an enormous diversity of solitary and social colony structures. To investigate social evolution in the genusHalictus, phylogenies of 15 species of the subgeneraH. (Halictus) andH. (Seladonia) were constructed based on protein electrophoretic data. Solitary, social, and socially polymorphic species were included.Halictus (Seladonia) apparently rendersH. (Halictus) paraphyletic. The common ancestor ofH. (Halictus) andH. (Seladonia) was probably social or socially polymorphic. This implies that some solitary and socially polymorphic species, such asH. confusus andH. tumulorum, represent evolutionary reversals from a completely eusocial condition to the solitary condition that is thought to be primitive for the subfamily as a whole.     

Survival time and resistance to desiccation of diapause and non-diapause eggs of temperate Aedes (Stegomyia) mosquitoes

Time-related changes in the patterns of host discrimination may have a complex form depending on whether or not hosts are marked following first parasitisation, the persistence of chemicals in these marks, and changes in the host as a result of being parasitised. Here we present an experimental design capable of detecting temporal patterns in host discrimination of solitary parasitoids. This method is tested against experimental data from Aphidius ervi attacking the aphid, Acyrthosiphon kondoi. The results demonstrate that there are statistically significant changes in the patterns of egg distribution in aphid hosts held for intervals between 1 and 48 h between parasitisation. These changes appear to be attributable to increasing oviposition restraint by the superparasitising female as the time interval between attacks is increased. The form of this relationship is not significantly different to a sigmoid function and the rate of change of restraint was greatest between 6–8 h after the first oviposition.     

Ant predation ofEriocrania miners in a polluted area

We tested the hypothesis that air pollution may affect population densities of birch-feeding leafminers via changes in ant predation. Foraging activity of three ant species (Formica rufa, F. fusca andF. lemani), predation rates and population densities of both solitary and gregariousEriocrania (Lepidoptera: Eriocraniidae) miners were investigated at 13 sites around the Harjavalta copper-nickel smelter, SW Finland. Ant species differed in their distribution patterns relative to pollution. However, the total percentage of birch trees foraged by ants (all species combined) showed no correlation with the distance from the factory complex. As a result, no clear trends in predation rates were apparent in relation to the distance from the pollution source for either solitary or gregariousEriocrania species. Densities of the solitaryEriocrania species tended to increase with the distance from the pollution source whereas densities of the gregariousE. haworthi peaked close to the factory complex. No corresponding differences in predation rates between solitary and gregarious miners were found. Ant predation, thus, did not explain density patterns ofEriocrania miners in the polluted area.     

Biology of <Emphasis Type="Italic">Lasioglossum</Emphasis> (<Emphasis Type="Italic">L.</Emphasis>) <Emphasis Type="Italic">majus</Emphasis> (Hymenoptera: Halictidae), a largely solitary sweat bee with behavioural adaptations to communality

Within the bee family Halictidae there have been three origins of sociality. Although detection of origins and reversal from sociality requires phylogenetic studies, at a behavioural level a predisposition to sociality can be detected by analysis of intra-specific interactions. We studied aspects of nesting biology and behavioural interactions in Lasioglossum (Lasioglossum) majus, a poorly known halictine inhabiting temperate regions of Europe, which is suspected to be solitary. Nests were found to be largely used by one female, but some were shared by more than one individual. These few nests, whose entrances were very close to each other, were found to be connected underground. A few individuals were observed to enter in a nest where a female was waiting, behaving as a guard and allowing the incoming bee to enter the nest. By use of circle-tube experiments, the behavioural repertoire exhibited by females during encounters was assessed. Levels of withdrawal and cooperative events were comparable with those observed in other solitary nesting species, but aggressive events were very rare, as in several observed communal species. We conclude that L. (L.) majus females, despite general solitary nesting, possess behavioural components enabling them to adopt, probably in high nest-density areas, nest-sharing strategies. A similar kind of local social polymorphism has been observed in two other species of the subgenus Lasioglossum, but these are the first data available on a European species and the first record of subterranean connections among nests of halictid bees.     

Social polymorphism in the sweat bee <Emphasis Type="Italic">Lasioglossum (Evylaeus) baleicum</Emphasis> (Cockerell) (Hymenoptera,Halictidae) in Hokkaido,northern Japan

Summary The life cycle and social behaviour of the sweat bee Lasioglossum (Evylaeus) baleicum (Cockerell) was investigated in two geographically separate populations in Hokkaido, northern Japan. Colonies were excavated throughout the brood rearing season from an aggregation in Nishioka forest park, Sapporo, and near Kawakita in eastern Hokkaido during 2000 and 2001. The Nishioka population produced two discrete broods during the year and was weakly eusocial; 57% of workers were mated and 28% exhibited some ovarian development, 12–16% of the first brood was male, and workers were on average 4.5% smaller than their respective queen. In contrast, the population at Kawakita was solitary, and produced a single brood per year with an unbiased sex ratio. In addition however, there were some solitary colonies in the Nishioka population and evidence of a partial second brood in some colonies at Kawakita, suggesting differences between the populations are not fixed and that this species is truly socially polymorphic. L. (E.) baleicum is a member of the fulvicorne species group, which includes other examples of social polymorphic species as well as solitary and eusocial species, though this is the only species of this group so far known to exhibit a solitary/non-delayed eusocial polymorphism. Recent studies suggest that social polymorphism has both genetic and environmental influences, raising questions as to the relative import of each.     

Gregariousness,field distribution and defence in the sawfly larvae Croesus varus and C. septentrionalis (Hymenoptera,Tenthredinidae)

Summary The larvae of Croesus varus are cryptic and those of C. septentrionalis aposematic. In both species, females lay eggs in groups, but the groups formed by young larvae are smaller in C. varus than in C. septentrionalis. In both species, group size decreases during successive larval instars, and the cryptic species becomes solitary, while the aposematic species remains gregarious during all larval instars. On Alnus shrubs in the field, the distribution of the groups ofC. septentrionalis larvae is more aggregative. They are found mainly on mediumsized shrubs (1.5–2.5 m high), whereas groups of C. varus larvae are also found on larger shrubs. Moreover, during the 2 years of observation, C. septentrionalis was present for a shorter time than C. varus, even when the former was much more abundant than the latter. Thus, conspicuousness of the aposematic species should be amplified by this spatial and temporal aggregation. The efficiency of both defensive strategies is compared and discussed, taking into account these particular field distributions, predation and some parasitism factors, and the chemical defences of both species.     

Copepods in spring annual sea ice at Terra Nova Bay (Ross Sea,Antarctica)

The aim of this study was to investigate patterns of abundance, distribution, temporal changes and species composition of the dominant ice-associated copepods in the spring annual pack ice, platelet ice and water column at Terra Nova Bay, Ross Sea, during late spring 1997. Ice cores were drilled for temporal and spatial scales. Stephos longipes and Harpacticus furcifer dominated the sea ice meiofauna in terms of numbers in the lower few centimeters of the bottom ice associated with high chlorophyll a and phaeopigment levels. Nauplii dominated the S. longipes population (91.6%) and occurred in extremely high concentrations. In contrast, copepodids were the dominant stages in H. furcifer. How H. furcifer carries out its entire life cycle and how it differs from ecologically similar species such as Drescheriella glacialis should be examined in more detail.     

Comparing infant and juvenile behavior in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes): a preliminary study

The dichotomy between the two Pan species, the bonobo (Pan paniscus) and chimpanzee (Pan troglodytes) has been strongly emphasized until very recently. Given that most studies were primarily based on adult individuals, we shifted the “continuity versus discontinuity” discussion to the infant and juvenile stage. Our aim was to test quantitatively, some conflicting statements made in literature considering species differences between immature bonobos and chimpanzees. On one hand it is suggested that infant bonobos show retardation in motor and social development when compared with chimpanzees. Additionally it is expected that the weaning process is more traumatic to chimpanzee than bonobo infants. But on the other hand the development of behaviors is expected to be very similar in both species. We observed eight mother–infant pairs of each species in several European zoos. Our preliminary research partially confirms that immature chimpanzees seem spatially more independent, spending more time at a larger distance from their mother than immature bonobos. However, the other data do not seem to support the hypothesis that bonobo infants show retardation of motor or social development. The development of solitary play, environmental exploration, social play, non-copulatory mounts and aggressive interactions do not differ between the species. Bonobo infants in general even groom other group members more than chimpanzee infants. We also found that older bonobo infants have more nipple contact than same aged chimpanzees and that the weaning process seems to end later for bonobos than for immature chimpanzee. Additionally, although immature bonobos show in general more signs of distress, our data suggest that the weaning period itself is more traumatic for chimpanzees.     

The influence of marking upon bee behaviour in circle tube experiments with a methodological comparison among studies

Summary. Circle tubes were used to investigate the influence of marking upon the behavioural profiles of two halictid species; one, Penapis toroi Rozen, putatively solitary, the other, Corynura chloris (Spinola) thought to be semisocial. Previous studies have shown that solitary bees tend to exhibit avoidance behaviour and those with a reproductive division of labour tend to behave aggressively within circle tube arenas. Marked pairs of C. chloris were significantly more cooperative than unmarked pairs and marked P. toroi showed significantly increased aggression and decreased cooperation. These findings suggest that marking bees may reduce behavioural differences among taxa with different types of social organization. Recommendations are made on how to use the circle tube apparatus to detect bee populations that might be worth more detailed studies of social organization and also for standardization in the way results of these studies are presented. Further experiments are suggested to clarify the effects of certain variables upon bee behaviour using this apparatus. The data presented here are the first for any member of the subfamily Rophitinae and the first for any bee thought to be primarily semisocial. A narrower range of behaviours were observed in the Rophitine, Penapis toroi Rozen, in comparison to species from other subfamilies of Halictidae and Corynura chloris Spinola had a behavioural profile similar to that of eusocial species.Received 15 December 2003; revised 12 September 2004; accepted 15 October 2004.     

Signalling through acoustic windows: nightingales avoid interspecific competition by short-term adjustment of song timing

The function of bird song is closely linked to sexual selection through female choice and male–male competition, and thus variation in communication success is likely to have major fitness consequences for a singing male. A crucial constraint on signal transmission is imposed by background noise, which may include songs from other species. I investigated whether singing nightingales (Luscinia megarhynchos) avoid temporal overlap with the songs of other bird species in a playback experiment. I analysed the temporal song patterns of six males, each of which were exposed to songs of other species. The nightingales significantly avoided overlapping their songs with the playback songs, and started singing preferentially during the silent intervals between the heterospecific songs. This timing of song onset produced a greater variability in pause duration compared to the nightingales’ undisturbed solo singing. These findings suggest that birds adjust their song timing to avoid acoustic interference on short temporal scales, and thus are able to improve the efficiency of acoustic communication in complex sonic environments. Moreover, the results indicate that temporal song patterns can be affected by the songs of other species, and thus such influences should be taken into account when studying bird song in the field.     

Effects of herbivory by chrysomelid beetles on the growth and survival ofRumex plants

The effects of herbivory by the guild of chrysomelid beetles on the growth and survival ofRumex plants were examined in relation to the distribution and size of the plants. Gastrophysa atrocyanea never appeared on solitary plants whereasMantula clavareaui showed even utilization of solitary and clumped plants.Galerucella vittaticollis utilized large solitary plants most frequently. In patches of the host plants, the frequency of withering of the aerial parts was higher on small plants. The regrowth rate was higher when the aerial parts withered in spring than in summer. In patches of the host plants, the regrowth rate was higher on small plants. On the other hand, it was lower on small solitary plants than on large solitary plants and small plants in patches. In large plants, no difference in survival rate was recognized between solitary and clumped plants, because both the frequency of withering of the aerial parts and the regrowth rate were lower in clumped than solitary plants. The high vigor against intense herbivory by the chrysomelid beetles brought on the delay in the phenology ofRumex plants. This shift permits in parts the existence ofG. vittaticollis in mid summer. The temporal and spatial interactions between herbivore guild and the host plants were discussed.     

Chilean Bromeliaceae: diversity, distribution and evaluation of conservation status

Chile is home to 23 species of Bromeliaceae, including 2 subspecies and 4 varieties. Twenty species are endemic to the country. We examined 883 herbarium specimens from 27 herbaria for our treatment of the Bromeliaceae for the “Flora de Chile”. These data and field observations resulted in a comprehensive database that we used to generate distribution maps for each species. We applied ecological niche modelling to reveal distribution areas and centers of Bromeliaceae diversity. We further analysed the collecting dates of the herbarium specimens to assess possible changes in species abundance. In this study we assess the conservation status of the bromeliad species in Chile. IUCN categories were assigned to the 27 bromeliad taxa as follows: Critically endangered: 4, Endangered: 6, Vulnerable: 11, Near threatened: 2, Least concern: 4. No species has become “Extinct” up to now. We also put forth a hypothesis about their biogeographic history.     

An overview of proximate factors affecting the nesting behavior of solitary wasps and bees (Hymenoptera: Aculeata) in preexisting cavities in wood

Guilds of Aculeate solitary wasps and bees that nest in preexisting cavities in wood are important components of terrestrial ecosystems because they engage in several ecological interactions (e.g. predation and pollination) with other species of plants and animals. Spatial and temporal variations in richness and abundance of solitary wasps and bees can be related to changes in environmental structure and in the diversity of other groups of organisms. The nesting period of these Aculeata is their most critical life cycle stage. Females of solitary wasp and bee species invest relatively more time constructing and provisioning their nests than do females of social species. Differently from species that nest in the soil or construct exposed nests, the main factors affecting the reproductive success of solitary species nesting in preexisting wood holes are still unknown. Our objective is to provide an overview of the role of proximate causes of nesting failure or success among solitary wasps and bees (Aculeata), for designing effective conservation and management strategies for these Hymenoptera.     

Rates of cross-pollination inLycopersicon pimpinellifolium: Impact of genetic variation in floral characters

Cross-pollination was tested between different genotypes ofLycopersicon pimpinellifolium in an experimental plot situated in the southerly range of the species. Rates of cross-pollination were found to be highly correlated with flower size and were related to a lesser extent with degree of stigma exsertion. Comparisons of fertility suggest that exsertion of 1.5 mm or more from the anther tube orifice interferes with automatic self-pollination, thereby increasing rates of cross-pollination. A single species ofExomalopsis (Anthophoridae), observed to visit all tested genotypes ofL. pimpinellifolium, was responsible for most of the pollination. Other species of solitary bees and other bee families participated in cross-pollination.     

The effect of fire on flower visitation rate and fruit set in four core-species in east Mediterranean scrubland

The recovery of vegetation following fire has been studied intensively in Mediterranean-type ecosystems. Little attention, however, has been given to floral traits, and almost no data have been collected on the effects of fire on pollinator activity and fruit-set. This paper reports the effects of fire on flower visitation rates and the possibly related fruit-set. We compared visitation rates of the main pollinators on four plant core-species in burned and adjacent unburned areas. Measurements were performed at an unburned phrygana (scrub lands), and at a burned area (5–7 years post-fire). Bumble bees and solitary bees were the main taxa of visitors, while few honeybees were recorded. Solitary bees were almost absent from the burned area. Fruit-set was significantly higher in the unburned area for three out of the four plant species. The lower fruit-set in the burned area was possibly the result of low activity of solitary bees which are the main effective pollinators of the examined species. We hypothesize that the populations of the solitary bees were diminished or extirpated either directly by the fire, or indirectly by the scarcity of nectar in the early post-fire years due to dominance of young pine and Cistus spp. seedlings. The short foraging range of the solitary bees and their slow invasion rate into the burned area may explain our results.     

High-resolution spatio-temporal distribution of a coastal phytoplankton bloom using laser in situ scattering and transmissometry (LISST)

Development of optical observation technologies provides new insights into harmful algal bloom (HAB) detection and assessment of HAB species dynamics. Based on preliminary laboratory tests, a laser in situ scattering and transmissometry instrument (LISST-100X) was used to monitor a high-biomass phytoplankton proliferation in the field. Short-term spatial and temporal changes in particle size distribution were measured during a recurrent Alexandrium taylori outbreak. Since the bloom was not monospecific, a size-fraction method to discriminate particular species from LISST-100X measurements was proposed. The results were validated to simultaneous microscopic counts of phytoplankton, and the significantly positive correlation obtained between the two methodologies confirmed the instrument's ability to discriminate phytoplankton at the group and species level. The LISST-100X obtains high-resolution in situ data, and is therefore a better alternative than the traditional microscope for assessing temporal and spatial evolution of HABs. Field observations showed high variability over a short time scale associated with diel vertical migration of A. taylori and the whole phytoplankton population (nanoplankton and microplankton). A numerical circulation model was used to investigate the influence of beach hydrodynamics in the observed horizontal variability. Simulations of the model suggested an important role of daily coastal circulation in determining the distribution of A. taylori in coastal environments.     

The allometry of echolocation call frequencies of insectivorous bats: why do some species deviate from the pattern?

The peak echolocation frequency of insectivorous bats generally declines as body size increases. However, there are notable exceptions to this rule, with some species, such as Rhinolophus clivosus, having a higher than expected peak frequency for their body size. Such deviations from allometry may be associated with partitioning of foraging habitat (the foraging habitat hypothesis) or insect prey (the prey detection hypothesis). Alternatively, the deviations may be associated with the partitioning of sonar frequency bands to allow effective communication in a social context (the acoustic communication hypothesis). We tested the predictions of these hypotheses through comparisons at the family, clade and species level, using species of rhinolophids in general and R. clivosus, a species with a wide distribution, as a specific test case. We compared the wing parameters, echolocation frequency and ecology of R. clivosus to those of the sympatric R. capensis. Rhinolophus clivosus has a much higher echolocation frequency than predicted from its wing loading or body mass. Furthermore, contrary to the predictions of the foraging habitat hypothesis, we found no difference in foraging habitat between R. clivosus and R. capensis. The size range of insect prey taken by the two species also overlapped almost completely, contrary to the prey detection hypothesis. On the other hand, the variation of echolocation frequencies around the allometric relationship for rhinolophids was smaller than that for Myotis spp., supporting the prediction of the acoustic communication hypothesis. We thus propose that the relatively high peak frequency of R. clivosus is the result of partitioning of sonar frequency bands to minimize the ambiguity of echolocation calls during social interactions.     

Temporal analysis of tone pulses within the courtship songs of two siblingDrosophila species,their interspecific hybrid,and behavioral mutants ofD. melanogaster (Diptera: Drosophilidae)

Temporal analyses were applied to the tone pulses within the courtship songs of Drosophila melanogaster, D. simulans,their interspecific hybrid, and behavioral mutants of D. melanogaster.Linear regression was performed on various parameters of the song pulses (cycles per pulse, absolute peak amplitude, intrapulse frequency, number of peaks in fast Fourier transform, width of the primary frequency peak, and interpulse interval), as a function of their positions within pulse trains. Significant differences in the slope values of these two species and of the mutant genotypes allowed for discriminative quantification of temporal changes within trains. These results are discussed in relation to previous kinds of temporal analyses of Drosophilacourtship songs and also with regard to the mechanisms of song production.     

Is there a reproductive basis to solitary living versus pair-formation in coral reef fishes?

Many species of coral reef fishes form pairs. While it is assumed that pairs represent the breeding unit of these species, the reproductive status of paired versus solitary individuals, and changes in status associated with pair-formation have seldom been investigated. In order to assess whether pairing is related to reproduction we examined whether the ontogenetic timing of pair formation coincided with the onset of maturation in four species of fishes: Chaetodon lunulatus and Chaetodon melannotus (family Chaetodontidae), and Valenciennea muralis and Valenciennea strigata (family Gobiidae). 65–78% of all fishes occurred in pairs. In C. lunulatus and V. muralis, pair-formation coincided with maturation, suggesting that these species form pairs for breeding. Further, C. lunulatus and V. muralis exhibited significant positive size-assortative pairing, which is often associated with monogamous mating. In contrast, pair formation in C. melannotus and V. strigata did not coincide with maturation. In both these species many solitary individuals were reproductive, and same sex pairs were common. While reproduction may be the basis for pairing in some species, both solitary and paired individuals are capable of breeding in others. We propose that non-reproductive mechanisms, such as predator vigilance, may explain pair-formation in coral reef fishes with non-monogamous breeding systems.