Pollinator visitation closely tracks diurnal patterns in pollen release.

PREMISE: Animal-pollinated plants face a high risk of pollen loss during its transfer. To limit the negative effect of pollen losses by pollen consumption and heterospecific transfer, plant species may adjust and stratify their pollen availability during the day (i.e., "schedule" their pollen presentation) and attract pollinators in specific time frames. METHODS: We investigated diurnal patterns of pollen availability and pollinator visitation in three coflowering plant species: Succisa pratensis with open flowers and accessible pollen, pollinated mainly by pollen-feeding hoverflies; Centaurea jacea with open flowers and less accessible pollen, pollinated mainly by pollen-collecting bees; and Trifolium hybridum with closed flowers and pollen accessible only after the active opening of the flower, pollinated exclusively by bees. RESULTS: The three plant species differed in the peak pollen availability, tracked by the visitation activity of their pollinators. Succisa pratensis released all pollen in the morning, while pollinator activity was still low and peaked with a slight delay. In contrast, C. jacea and T. hybridum had distinct pollen presentation schedules, peaking in the early afternoon. The pollinator visitation to both of these species closely matched their pollen availability. CONCLUSIONS: Stratifying pollen availability to pollinators during the day may be one of several mechanisms that allow coflowering plants to share their pollinators and decrease the probability of heterospecific pollen transfer.

Polyandrous bee provides extended offspring care biparentally as an alternative to monandry based eusociality.

Parental care behavior evolves to increase the survival of offspring. When offspring care becomes complicated for ecological reasons, cooperation of multiple individuals can be beneficial. There are two types of cooperative care: biparental care and worker (helper)-based care (e.g., eusociality). Although biparental care is common in several groups of vertebrates, it is generally rare in arthropods. Conversely, eusociality is widespread in insects, especially the aculeate Hymenoptera. Here, we present a case of biparental care in bees, in Ceratina nigrolabiata (Apidae, Xylocopinae). Similar to eusocial behavior, biparental care leads to greater brood protection in this species. Male guarding increases provisioning of nests because females are liberated from the tradeoff between provisioning and nest protection. The main benefit of parental care for males should be increased paternity. Interestingly though, we found that paternity of offspring by guard males is extraordinarily low (10% of offspring). Generally, we found that nests were not guarded by the same male for the whole provisioning season, meaning that males arrive to nests as stepfathers. However, we show that long-term guarding performed by a single male does increase paternity. We suggest that the multiple-mating strategy of these bees increased the amount of time for interactions between the sexes, and this longer period of potential interaction supported the origin of biparental care. Eusociality based on monandry was thought to be the main type of extended brood protection in bees. We show that biparental care based on polyandry provides an interesting evolutionary alternative.

Nuclear Genome Size in Contrast to Sex Chromosome Number Variability in the Human Bed Bug, Cimex lectularius (Heteroptera: Cimicidae).

The human bed bug Cimex lectularius is one of the most prevalent human ectoparasites in temperate climate zones. The cytogenetic features of this resilient pest include holokinetic chromosomes, special chromosome behavior in meiosis, and numerical variation of chromosomes, where the diploid number ranges from 26 + X1 X2 Y to 26 + X1-20 Y. It is desirable to assess the nuclear DNA content of various cytotypes for a further detailed study of the C. lectularius genome. Detailed knowledge of the DNA content of this parasite could also clarify the origin of additional chromosomes. The average nuclear genome size C. lectularius with 2n = 26 + X1 X2 Y is 2C = 1.94 pg for males and 1.95 pg for females. There is a significant correlation between genome size and the number of chromosomes, but in some specimens with additional chromosomes, nuclear genome size decreases or remains average. Several species used as the internal reference standard were tested for further investigations of genome size in C. lectularius, and the plant Solanum pseudocaspicum turned out to be the most suitable. © 2019 International Society for Advancement of Cytometry.

Autumnal migration patterns of hoverflies (Diptera: Syrphidae): interannual variability in timing and sex ratio.

Background: The migration of hoverflies (Diptera: Syrphidae) is a well-known phenomenon, with growing interest due to the ecosystem services provided by migrants. However, we still lack fundamental data on species composition, timing of migration, or sex ratio of migrants. To address this gap, we focused on the southward autumnal migration of hoverflies through central Europe. Methods: To recognize migrating individuals from resident ones, we used a pair of one-side-blocked Malaise traps, exposed in a mountain pass in the Jeseníky mountains, Czech Republic, where a mass migration of hoverflies takes place annually. Traps were set for 4 years, from August to October. Results: In total, we recorded 31 species of migrating hoverflies. The timing of migration differed between the years, taking place from the beginning of September to the end of October. Differences in phenology were observed in the four most common migrant species, where larger species seemed to migrate earlier or at the same time compared to the smaller ones. The sex ratio was strongly asymmetrical in most common species Episyrphus balteatus, Eupeodes corollae, and Sphaerophoria scripta, and varied between years for each species. Weather conditions strongly influenced the migration intensity at ground-level: hoverflies migrate mainly during days with south wind, high temperature, high atmospheric pressure, and low precipitation.

An intergenerational approach to parasitoid fitness determined using clutch size.

Parasitoids, as important natural enemies, occur in high numbers and help maintain balance in natural ecosystems. Their fitness is traditionally studied as fertility based on the number of offspring in the F1 generation. Here, using gregarious parasitoids as models, we show that this traditional approach omits one important parameter: the clutch size-body size-fertility correlation among offspring. As a result of this correlation, when females adjust the number of offspring laid in a host, they determine not only the number of offspring produced but also the body size and reproductive potential of those offspring. Although parasitoid fertility has been determined several times from clutch size, here we use Anaphes flavipes to demonstrate the use of this relationship in an upgraded intergenerational approach to parasitoid fitness. We show that with a range of hosts simultaneously utilized by female parasitoids, identical fertility in the F1 generation can lead to distinctly different fertility values in the F2 generation. Even with the same number of hosts, lower fertility in the F1 generation can generate higher fertility in the F2 generation. Our approach provides an intergenerational perspective for determining individual fitness of gregarious parasitoids and new possibilities for the modelling of parasitoid population density.

Individual-level specialisation and interspecific resource partitioning in bees revealed by pollen DNA metabarcoding.

It is increasingly recognised that intraspecific variation in traits, such as morphology, behaviour, or diet is both ubiquitous and ecologically important. While many species of predators and herbivores are known to display high levels of between-individual diet variation, there is a lack of studies on pollinators. It is important to fill in this gap because individual-level specialisation of flower-visiting insects is expected to affect their efficiency as pollinators with consequences for plant reproduction. Accordingly, the aim of our study was to quantify the level of individual-level specialisation and foraging preferences, as well as interspecific resource partitioning, in three co-occurring species of bees of the genus Ceratina (Hymenoptera: Apidae: Xylocopinae), C. chalybea, C. nigrolabiata, and C. cucurbitina. We conducted a field experiment where we provided artificial nesting opportunities for the bees and combined a short-term mark-recapture study with the dissection of the bees' nests to obtain repeated samples from individual foraging females and complete pollen provisions from their nests. We used DNA metabarcoding based on the ITS2 locus to identify the composition of the pollen samples. We found that the composition of pollen carried on the bodies of female bees and stored in the brood provisions in their nests significantly differed among the three co-occurring species. At the intraspecific level, individual females consistently differed in their level of specialisation and in the composition of pollen carried on their bodies and stored in their nests. We also demonstrate that higher generalisation at the species level stemmed from larger among-individual variation in diets, as observed in other types of consumers, such as predators. Our study thus reveals how specialisation and foraging preferences of bees change from the scale of individual foraging bouts to complete pollen provisions accumulated in their nests over many days. Such a multi-scale view of foraging behaviour is necessary to improve our understanding of the functioning of plant-flower visitor communities.

Effect of host switching simulation on the fitness of the gregarious parasitoid Anaphes flavipes from a novel two-generation approach.

Herbivorous insects can escape the strong pressure of parasitoids by switching to feeding on new host plants. Parasitoids can adapt to this change but at the cost of changing their preferences and performance. For gregarious parasitoids, fitness changes are not always observable in the F1 generation but only in the F2 generation. Here, with the model species and gregarious parasitoid Anaphes flavipes, we examined fitness changes in the F1 generation under pressure from the simulation of host switching, and by a new two-generation approach, we determined the impact of these changes on fitness in the F2 generation. We showed that the parasitoid preference for host plants depends on hatched or oviposited learning in relation to the possibility of parasitoid decisions between different host plants. Interestingly, we showed that after simulation of parasitoids following host switching, in the new environment of a fictitious host plant, parasitoids reduced the fictitious host. At the same time, parasitoids also reduced fertility because in fictitious hosts, they are not able to complete larval development. However, from a two-generation approach, the distribution of parasitoid offspring into both native and fictitious hosts caused lower parasitoid clutch size in native hosts and higher individual offspring fertility in the F2 generation.

Host Specificity of the Parasitic Wasp Anaphes flavipes (Hymenoptera: Mymaridae) and a New Defence in Its Hosts (Coleoptera: Chrysomelidae: Oulema spp.).

The parasitic wasp Anaphes flavipes (Förster, 1841) (Hymenoptera: Mymaridae) is an important egg parasitoid of cereal leaf beetles. Some species of cereal leaf beetle co-occur in the same localities, but the host specificity of the wasp to these crop pests has not yet been examined in detail. A lack of knowledge of host specificity can have a negative effect on the use of this wasps in biological control programs addressed to specific pest species or genus. In this study, laboratory experiments were conducted to assess the host specificity of A. flavipes for three species of cereal leaf beetles (Oulema duftschmidi Redtenbacher, 1874, Oulema gallaeciana Heyden, 1879 and Oulema melanopus Linnaeus, 1758) in central Europe. For the first time, a new host defence against egg parasitoids occurring in O. gallaeciana from localities in the Czech Republic, a strong dark sticky layer on the egg surface, was found and described. The host specificity of A. flavipes was studied in the locality with the presence of this defence on O. gallaeciana eggs (the dark sticky layer) (Czech Republic) and in a control locality (Germany), where no such host defence was observed. Contrary to the idea that a host defence mechanism can change the host specificity of parasitoids, the wasps from these two localities did not display any differences in that. Respectively, even though it has been observed that eggs with sticky dark layer can prevent parasitization, the overall rate of parasitization of the three species of cereal beetles has not been affected. However, in our view, new host defence can influence the effects of biological control, as eggs of all Oulema spp. in the locality are protected against parasitization from the wasps stuck on the sticky layer of the host eggs of O. gallaeciana.

Host population density and presence of predators as key factors influencing the number of gregarious parasitoid Anaphes flavipes offspring.

The number of parasitoids developed per host is one of the major factors that influences future adult body size and reproductive success. Here, we examined four external factors (host species, heritability, host population density, and presence of predators) that can affect the number of the gregarious parasitoid Anaphes flavipes (Förster, 1841) (Hymenoptera: Mymaridae) wasps developing in one host. The effect of host population density on the number of parasitoid offspring developed per host was confirmed, and for the first time, we also showed that the number of offspring per host is influenced by the presence of predators. Low host density and presence of predators increases the number of wasps developed in one host egg. However, a higher number of A. flavipes in one host reduces A. flavipes body size and hence its future individual fertility and fitness. Our results highlighted the importance of some external factors that distinctly affect the number of wasp offspring. Therefore, in this context, we suggest that in comparison to solitary parasitoids, the gregarious parasitoid A. flavipes can better respond to various external factors and can more flexibly change its population density.

European bees of the genera Epeolus Latreille, 1802 and Triepeolus Robertson, 1901 (Hymenoptera: Apidae: Nomadinae: Epeolini): taxonomy, identification key, distribution, and ecology.

Seventeen species of Epeolus Latreille, 1802 and one species of Triepeolus Robertson, 1901 have been recorded from Europe. Members of these genera are cuckoo bees, cleptoparasites that appropriate provisions of other bee genera (Colletes Latreille [hosts of Epeolus] and Tetraloniella Ashmead [hosts of Palaearctic Triepeolus]) for their own offspring. We present an identification key to all European species plus four species known from neighbouring regions (the Middle East and North Africa) that might also occur in Europe. The taxonomy of all species is reviewed, with new synonymies proposed, one subspecies elevated to a species (Epeolus eriwanensis Bischoff, 1930, stat. nov.), and a new species (Epeolus ibericus Bogusch, sp. nov.) described. Five new synonymies are proposed: E. giannelli Gribodo, 1894 syn. nov. and Epeolus speculifer Pérez, 1895 syn. nov. under E. fallax Morawitz, 1872, E. berlandi Benoist, 1943 syn. nov. and E. laevigatus Bischoff, 1930 syn. nov. under E. flavociliatus Friese, 1899, and E. julliani Pérez, 1884 syn. nov. under E. transitorius Eversmann, 1852. The following species are redescribed because their identity is less clear due to type specimens and/or vague original descriptions: Epeolus aureovestitus Dours, 1873, Epeolus cruciger (Panzer, 1799), Epeolus fallax Morawitz, 1872, Epeolus schummeli Schilling, 1849, Epeolus siculus Giordani Soika, 1944, and Epeolus variegatus (Linnaeus, 1758). Males are described for the first time for the following species: Epeolus compar Alfken, 1938, Epeolus eriwanensis Bischoff, 1930, Epeolus intermedius Pérez, 1884, Epeolus laevifrons Bischoff, 1930, and Epeolus subrufescens Saunders, 1908. Distribution maps created using records from the literature and data from private and museum collections are also provided, which show changes in the distributions of epeoline bees before and after 1990. Several species are considered endangered, especially Epeolus fasciatus Friese, 1895, which is nearly extinct, and Epeolus tarsalis Morawitz, 1874, which was recently reported only from the coastal parts of the Netherlands and Belgium. Epeolus variegatus is the only common and widespread species still encountered in most of Europe. Comprehensive information on the ecology of these cuckoo bees is also included.

Vertical stratification of plant-pollinator interactions in a temperate grassland.

Visitation of plants by different pollinators depends on individual plant traits, spatial context, and other factors. A neglected aspect of small-scale variation of plant-pollinator interactions is the role of vertical position of flowers. We conducted a series of experiments to study vertical stratification of plant-pollinator interactions in a dry grassland. We observed flower visitors on cut inflorescences of Centaurea scabiosa and Inula salicina placed at different heights above ground in two types of surrounding vegetation: short and tall. Even at such a small-scale, we detected significant shift in total visitation rate of inflorescences in response to their vertical position. In short vegetation, inflorescences close to the ground were visited more frequently, while in tall vegetation, inflorescences placed higher received more visits. Moreover, we found major differences in the composition of the pollinator community on flowers at different heights. In a second experiment, we measured flower visitation rate in inflorescences of Salvia verticillata of variable height. Total flower visitation rate increased markedly with inflorescence height in this case. Data on seed set of individual plants provide evidence for a corresponding positive pollinator-mediated selection on increased inflorescence height. Overall, our results demonstrate strong vertical stratification of plant-pollinator interactions at the scale of mere decimetres. This may have important ecological as well as evolutionary implications.

Large-scale assessment of commensalistic-mutualistic associations between African birds and herbivorous mammals using internet photos.

Birds sitting or feeding on live large African herbivorous mammals are a visible, yet quite neglected, type of commensalistic-mutualistic association. Here, we investigate general patterns in such relationships at large spatial and taxonomic scales. To obtain large-scale data, an extensive internet-based search for photos was carried out on Google Images. To characterize patterns of the structural organization of commensalistic-mutualistic associations between African birds and herbivorous mammals, we used a network analysis approach. We then employed phylogenetically-informed comparative analysis to explore whether features of bird visitation of mammals, i.e., their mean number, mass and species richness per mammal species, are shaped by a combination of host mammal (body mass and herd size) and environmental (habitat openness) characteristics. We found that the association web structure was only weakly nested for commensalistic as well as for mutualistic birds (oxpeckers Buphagus spp.) and African mammals. Moreover, except for oxpeckers, nestedness did not differ significantly from a null model indicating that birds do not prefer mammal species which are visited by a large number of bird species. In oxpeckers, however, a nested structure suggests a non-random assignment of birds to their mammal hosts. We also identified some new or rare associations between birds and mammals, but we failed to find several previously described associations. Furthermore, we found that mammal body mass positively influenced the number and mass of birds observed sitting on them in the full set of species (i.e., taking oxpeckers together with other bird species). We also found a positive correlation between mammal body mass and mass of non-oxpecker species as well as oxpeckers. Mammal herd size was associated with a higher mass of birds in the full set of species as well as in non-oxpecker species, and mammal species living in larger herds also attracted more bird species in the full set of species. Habitat openness influenced the mass of birds sitting on mammals as well as the number of species recorded sitting on mammals in the full set of species. In non-oxpecker species habitat openness was correlated with the bird number, mass and species richness. Our results provide evidence that patterns of bird-mammal associations can be linked to mammal and environmental characteristics and highlight the potential role of information technologies and new media in further studies of ecology and evolution. However, further study is needed to get a proper insight into the biological and methodological processes underlying the observed patterns.

Flower visitation by hoverflies (Diptera: Syrphidae) in a temperate plant-pollinator network.

Hoverflies (Diptera: Syrphidae) are among the most important pollinators, although they attract less attention than bees. They are usually thought to be rather opportunistic flower visitors, although previous studied demonstrated that they show colour preferences and their nectar feeding is affected by morphological constraints related to flower morphology. Despite the growing appreciation of hoverflies and other non-bee insects as pollinators, there is a lack of community-wide studies of flower visitation by syrphids. The aim of this paper is to provide a detailed analysis of flower visitation patterns in a species rich community of syrphids in a Central European grassland and to evaluate how species traits shape the structure of the plant-hoverfly flower visitation network. We found that different species varied in the level of specialisation, and while some species visited a similar spectre of flowers, others partitioned resources more strongly. There was a consistent difference in both specialisation and flower preferences between three syrphid subfamilies. Eristalinae and Pipizinae were more specialised than Syrphinae. Trait-based analyses showed that relative flower visitation (i) increased with plant height, but most strongly in Eristalinae; (ii) increased with inflorescence size in small species from all three subfamilies, but was independent of inflorescence size in large species of Eristalinae and Syrphinae; and (iii) depended on flower colour, but in a subfamily-specific way. Eristalinae showed the strongest flower colour preferences for white flowers, Pipizinae visited mostly white and yellow flowers, while Syrphinae were less affected by flower colour. Exploration of the structure of the plant-hoverfly flower visitation network showed that the network was both modular and nested. We also found that there were almost no differences in specialisation and relative visitation frequency between males and females. Overall, we showed that flower visitation in syrphids was affected by phylogenetic relatedness, body size of syrphids and several plant traits.

Conspecific and heterospecific plant densities at small-scale can drive plant-pollinator interactions.

Generalist pollinators are important in many habitats, but little research has been done on small-scale spatial variation in interactions between them and the plants that they visit. Here, using a spatially explicit approach, we examined whether multiple species of flowering plants occurring within a single meadow showed spatial structure in their generalist pollinator assemblages. We report the results for eight plant species for which at least 200 individual visits were recorded. We found that for all of these species, the proportions of their general pollinator assemblages accounted for by particular functional groups showed spatial heterogeneity at the scale of tens of metres. This heterogeneity was connected either with no or only subtle changes of vegetation and flowering species composition. In five of these species, differences in conspecific plant density influenced the pollinator communities (with greater dominance of main pollinators at low-conspecific plant densities). The density of heterospecific plant individuals influenced the pollinator spectrum in one case. Our results indicate that the picture of plant-pollinator interactions provided by averaging data within large plots may be misleading and that within-site spatial heterogeneity should be accounted for in terms of sampling effort allocation and analysis. Moreover, spatially structured plant-pollinator interactions may have important ecological and evolutionary consequences, especially for plant population biology.