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.

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.

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.

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.