Mating structure and male production in Vespa analis and Vespa simillima (Hymenoptera: Vespidae)

We estimated queen mating frequency, genetic relatedness between workers and worker reproduction in the hornets Vespa analis and Vespa simillima using microsatellite DNA genotyping. The 20 V. analis colonies studied each contained a queen inseminated by a single male. Of the 15 V. simillima colonies studied, nine had a queen inseminated by a single male, four had a queen inseminated by two males, and two had a queen inseminated by three males. The estimated effective number of matings was 1.33 ± 0.74 (mean ± SD), with 75–85% of the offspring of the six multiply mated queens sired by single males. The values for genetic relatedness between the workers of V. analis and V. simillima were 0.739 ± 0.004 and 0.698 ± 0.013 (mean ± SD), respectively. We conclude that V. analis and V. simillima colonies are genetically monogynous and monandrous. When high relatedness between the workers occurs within colonies, kin selection theory predicts a potential conflict between queens and workers over male production. To determine whether males were derived from queens or workers, males from V. analis and V. simillima colonies were genotyped at four microsatellite loci and the level of ovary activation in workers was determined. None of the 787 V. analis workers and only 15 of 3520 V. simillima workers had developed ovaries. Furthermore, the genotyping identified no worker‐produced males in any colony. The presence of reproductive workers correlated positively with the number of workers within the colony. These results suggest that eusocial colonies with an annual life cycle tend to break down socially when they become large and are close to dying.     

The life cycle of Micromalthus debilisLeConte (1878) (Coleoptera: Archostemata: Micromalthidae): historical review and evolutionary perspective

Micromalthus debilis 33 ), has one of the most bizarre life cycles of any metazoan. Reproduction is typically by thelytokous, viviparous, larviform females, but there is also a rare arrhenotokous phase. The active first‐instar (triungulin) larva develops into a legless, feeding (cerambycoid) larva. This form either pupates, leading to a diploid adult female, or develops into any of three subsequent types of reproductive paedogenetic forms: (1) a thelytokous female that produces triungulins via viviparity; (2) an arrhenotokous female that produces a single egg that develops into the short‐legged (curculionoid) larva, eventually devouring its mother and becoming a haploid adult male; or (3) an amphitokous female that can follow either of the two above reproductive pathways. We speculate that Micromalthus is dependent on maternally transmitted bacteria for the ability to digest rotting wood, and that these bacteria are senescent in males, causing males to be obligately cannibalistic. Obligate male cannibalism, in turn, would have dramatically increased the cost of males, and have created a strong selective advantage for cyclic thelytoky and the other features of the Micromalthus life cycle that minimize the role of the male.     

Is the maximum reproductive rate of <Emphasis Type="Italic">Centris analis</Emphasis> (Hymenoptera,Apidae, Centridini) associated with floral resource availability?

Spatiotemporal variation in the availability of food resources may be a determining factor for reproductive success and maintenance of bees, but the extent of these variations is poorly understood. For management and conservation of bees, the first step is to know the behavior and the food resources used. Currently, urban areas are considered refuge zones for bees, and understanding the availability of floral resources and the influence on reproductive processes is very important for management of bees. We used the protocols applied in phenological studies with bees and plant species to evaluate both throughout the year in an urbanized area. At the same time, we used palynology protocols to analyze the pollen material collected from brood cells (food and feces) of immature Centris analis. These protocols allowed to evaluate the availability of floral resources in the studied area and the plant species effectively used by C. analis females to feed immature larvae during the reproductive period. The maximum reproductive period of C. analis was not associated with the highest floral resources availability. However, there was a strong selectivity of pollen in flowers of Malpighia emarginata (Malpighiaceae), which represented more than 59% of all the pollen grains provisioned throughout the year. This means that in the case of more specialized bees like C. analis, the availability of the preferred plants is more important than the overall floral resource availability in the area. Thus, to keep C. analis in the city, it is necessary to maintain or introduce Malpighiaceae species in the urban planning. On the other hand, at least 27% of the plant species found in the study area are pollinated by C. analis, emphasizing the importance of preserving this bee.     

Patterns of sexual size dimorphism in stingless bees: Testing Rensch’s rule and potential causes in highly eusocial bees (Hymenoptera: Apidae,Meliponini)

Eusocial insects offer a unique opportunity to analyze the evolution of body size differences between sexes in relation to social environment. The workers, being sterile females, are not subject to selection for reproductive function providing a natural control for parsing the effects of selection on reproductive function (i.e., sexual and fecundity selection) from other kinds of natural selection. Patterns of sexual size dimorphism (SSD) and testing of Rensch's rule controlling for phylogenetic effects were analyzed in the Meliponini or stingless bees. Theory predicts that queens may exhibit higher selection for fecundity in eusocial taxa, but contrary to this, we found mixed patterns of SSD in Meliponini. Non‐Melipona species generally have a female‐biased SSD, while all analyzed species of Melipona showed a male‐biased SSD, indicating that the direction and magnitude of the selective pressures do not operate in the same way for all members of this taxon. The phylogenetic regressions revealed that the rate of divergence has not differed between the two castes of females and the males, that is, stingless bees do not seem to follow Rensch's rule (a slope >1), adding this highly eusocial taxon to the various solitary insect taxa not conforming with it. Noteworthy, when Melipona was removed from the analysis, the phylogenetic regressions for the thorax width of males on queens had a slope significantly smaller than 1, suggesting that the evolutionary divergence has been larger in queens than males, and could be explained by stronger selection on female fecundity only in non‐Melipona species. Our results in the stingless bees question the classical explanation of female‐biased SSD via fecundity and provide a first evidence of a more complex determination of SSD in highly eusocial species. We suggest that in highly eusocial taxa, additional selection mechanisms, possibly related to individual and colonial interests, could influence the evolution of environmentally determined traits such as body size.     

Mating frequency and genetic relatedness of workers in the hornet Vespa analis (Hymenoptera: Vespidae)

Mating frequency of Vespa analis queens and the genetic relatedness of their workers was analyzed by DNA microsatellite genotyping. Of 20 colonies studied, 18 had a queen inseminated by a single male and two had queens each inseminated by two males. The estimated effective number of matings was 1.05 ± 0.037 (mean ± SE), with 75–85% of the offspring of the two multiply mated queens sired by a single male. The pedigree relatedness between nestmate workers averaged over the 20 colonies was estimated to be 0.74 ± 0.008, almost identical to the predicted value of 0.75 for colonies headed by a singly mated queen. Multiple matrilines; that is, the presence of workers not related to the current queens, were detected in six colonies, suggesting that queen replacement occurred via usurpation of the founding queens in these six colonies. These results demonstrate that the kin structure of V. analis is similar to that reported in other vespid species.     

Contest and scramble competitions in two bruchid species,Callosobruchus analis andC. phaseoli (Coleoptera: Bruchidae)

Larval competition between contest and scramble strategists was investigated using the two bruchid species, C. analis (contest species) and C. phaseoli (scramble species) with two different sized mung beans (large and small beans). In both sized beans, the adult emergences of each species dependen on total density of the initial larval densities of the two species and the ratio of the two densities. The emergence of one species was suppressed by the existence of the other species when the initial larval density per bean of the former species was less than that of the latter one. There were many cases in which both C. analis and C. phaseoli emerged from one bean in large beans, but such cases were quite rare in small beans. C. analis performed interference behavior only at late larval stages, whereas C. phaseoli was superior in exploitative competition all through their larval stages. These, combined with the niche segregation inside a bean, are throught to be the major factors of observed density- and frequency-dependent competition results. Based on the above experimental results, long-term competition results between the contest and scramble species were predicted.     

Visual Cues in Mate Recognition by Males of the Damselfly, Coenagrion puella (L.) (Odonata: Coenagrionidae)

Coenagrion puella males search actively for mates and are not aggressive to other males. To study the role of visual cues in male–female discrimination, four types of models were used: (1) bodies of intact insects, (2) models of painted males, (3) models of male–female chimerae, and (4) models of female body parts. Abdomen coloration pattern and presence of wings were the most important cues for sexual recognition by males. Step-by-step elimination of male coloration pattern leads to an increase in the tandem response rate. A female model painted as a male repelled males like the intact male model. The absence of either the head or the thorax slightly decreased the number of tandem responses, but models without both the head and the thorax were not recognized as a mate. Abdomen thickness larger than that of a normal female decreased the attractiveness of the model. Models of the gynochrome female were significantly more attractive than models of the androchrome one. Female models containing male parts were less attractive than models without any structure at this place. Using principal-components analysis, it is shown that models repelling males usually were those containing an intact male abdomen or a female abdomen painted with blue. The results indicate that C. puella males can distinguish males from females visually by morphological structures and coloration pattern.     

Centris pallida (Hymenoptera: Apidae) male body size decreases across five decades

Historical data suggest that many bee species have declined in body size. Larger-bodied bees with narrow phenological and dietary breadth are most prone to declines in body size over time. This may be especially true in solitary, desert-adapted species that are vulnerable to climate change such as Centris pallida (Hymenoptera: Apidae). In addition, body size changes in species with size-linked behaviours could threaten the prevalence of certain behavioural phenotypes long-term. C. pallida solitary bees are found in the Sonoran Desert. Males use alternative reproductive tactics (ARTs) and are dimorphic in both morphology and behaviour. C. pallida male body size has been studied since the 1970s in the same population. The authors collected body size data in 2022 and combined it with published records from 1974–2022. The authors find a persistent decline in the mean head width of patrolling males, and shifts towards smaller body sizes in the populations of males found foraging and hovering. Both morphs declined in average body size, and the proportion of large-morph males in the population decreased by 8%. Mating males did not decline in mean body size over the last five decades. The authors discuss hypotheses related to the decline in C. pallida male head width. Finally, the authors advocate for C. pallida as an excellent study system for understanding the stability of ARTs with size-linked behavioural phenotypes.     

Host-seeking behaviour of tracheal mites (Acari: Tarsonemidae) on honey bees (Hymenoptera: Apidae)

The behaviour of the endoparasitic tracheal mite, Acarapis woodi (Rennie) on honey bees (Apis mellifera L.) is a challenge to observe because of its small size. Through a microscope, we videotaped this mite's movement on young bees, dead bees and bees exposed to vegetable oil. Previous studies have shown that solid vegetable oil decreases mite infestations in a bee colony. We hypothesized that the oil alters mite behaviour to the detriment of the parasite, thus helping to safeguard the host. Habitat-seeking behaviour, identified as necessary for mites to locate a new host environment, was disrupted on both dead and oil-treated bees. Questing behaviour, which is associated with transfer between hosts, increased significantly on the dead and oily bees. The behaviours of mites were significantly different between all three treatments (x ²=494.96, p<0.001 on dead bees and x ²=851.11, p<0.001 on oily bees). Both questing and seeking behaviours were significantly different on each of the thoracic treatments (F _2,66=7.88, p<0.001 and F _2,66=21.28, p<0.001) and mite questing behaviour was not altered between males and females on live or oily bees (F _1,22=0.25, p<0.62), but habitat seeking was (F _1,22=7.42, p<0.012). The male questing and habitat-seeking behaviours were observed. We conclude that oil-treated bees gained protection from habitat-seeking mites because the normal behaviour of the mites seeking an oviposition site is interrupted.     

Sexual dimorphism documented in Reptalus iguchii (Matsumura) (Hemiptera: Fulgoromorpha: Cixiidae) with a description of males

Reptalus iguchii (Matsumura 1914) comb. nov., previously known from female specimens, is recognized for the first time in the Republic of Korea. This species is sexually dimorphic, based on observations of a pair in copula. Detailed morphological characters of both male and female were studied and are illustrated. Also, sequence data from ribosomal internal transcribed spacer 2 (ITS2) and cytochrome oxidase subunit I (COI) genes were used to distinguish males and females of the only other congener known from Korea, R. quadricinctus (Matsumura 1914). Artemisia and Prunus are recorded as potential hosts in the Republic of Korea for the first time.     

Male emergence timing and mating success in the funnel-web spider,Agelena limbata (Araneae: Agelenidae)

Field observations on the relationship between male mating success and emergence timing in the funnel-web spider,Agelena limbata, were conducted.Agelena limbata is an annual species and adult males appear slightly earlier than adult females in July. As males deposit a copulatory plug at the female epigynum after copulation, copulation with virgin females is important to males. The number of copulations in males with virgin females, which strongly correlates with the longevity of males and the number of females that males courted, did not correlate with the emergence timing of males. Early emerged males and females were significantly larger in size than later ones, but the correlation coefficient between the emerged date and the cephalothorax width was not strong. Males that emerged earlier did not have any advantage in copulating with larger and more fecund females. Furthermore, virgin females first copulated on average 7.9 days after their final molt and the mortality rate of adult males increased after the final molt. These factors may favor the smaller degree of protandry in male emergence timing inA. limbata.     

Foraging and courtship behaviour in males of the solitary bee Anthophora plumipes (Hymenoptera: Anthophoridae): thermal physiology and the roles of body size

Abstract.

• 1 The effects of climate and body size on male behaviour were examined in the solitary bee Anthophora plumipes (Hymenoptera: Anthophoridae), which shows resource-based polygyny at floral food sources in Britain in spring.
• 2 Larger males are able to fly at lower temperatures than smaller males, and can therefore court females under conditions in which smaller males cannot fly. This is predicted from patterns of endothermic ability at low temperatures already demonstrated within this species.
• 3 Video analysis of male competition for opportunities to initiate courtship with tethered females showed that larger males are also competitively superior, and can displace smaller males from favoured flight positions immediately behind females.
• 4 The mating system shown by male A plumipes is strongly dependent on male density. At low densities, males show exclusive territoriality at floral sources. As male density increases, this strategy is abandoned in favour of patrolling with considerable spatial overlap between males, and opportunistic Polygyny.
• 5 Despite high endothermic abilities, male behaviour is highly dependent on weather, and particularly ambient temperature. Males bask in the early morning and maintain high thoracic temperatures. Temperature data from freshly killed bees show that thoracic warming from solar sources effectively doubles the thermogenic power generated by the bee alone at low ambient temperatures.
• 6 Male strategies in A.plumipes are compared to female responses to climate. Having controlled for differences in body size there is no difference in endothermic abilities between the sexes. Males do not, however, fly under conditions in which females of the same size would remain active. These results are discussed in the light of differential dependence of reproductive success on flight activity for the two sexes.

     

Vibrating the food receivers: a direct way of signal transmission in stingless bees (Melipona seminigra)

An element common to the recruitment communication of eusocial bees (honey bees, stingless bees and bumble bees) are pulsed thorax vibrations generated by successful foragers within the nest. In stingless bees, foragers vibrate during the unloading of the collected food. In the present study on Melipona seminigra we demonstrate that during trophallactic contacts, the food receivers are directly vibrated by the foragers. As a consequence, both the temporal structure and the main frequency component of the forager’s vibrations are directly passed on to the receiver. The vibrations are attenuated by about 17 dB on their way from the forager’s thorax (velocity amplitude of the vibrations: ∼70 mm/s) to the receiver’s thorax (∼10 mm/s), the main amount of attenuation (about 12 dB) occurring during transmission from the head of the forager to that of the receiver. Vibrations conducted through the substrate between the forager and food receiver are comparatively small with velocity amplitudes of 0.3 mm/s. Possible ways of perception and the advantages of vibration transmission by direct contact within the recruitment context are discussed.     

Quality control of Cotesia flavipes (Cameron) (Hymenoptera: Braconidae) from different Brazilian bio-factories

The goal of this study was to evaluate the quality of Cotesia flavipes from different bio-factories as biological control agents. We evaluated biological characteristics of the parasitoids throughout their lifespan, and measured the body length and width, abdomen width, thorax width and width and length of the right forewing of female and male parasitoids. Our results showed that the number of males and pupal viability were similar among the bio-factories; the number of emerged females was greater in bio-factories I and II; the egg-pupa period and the pupal period were shorter in bio-factory IV; and a greater longevity was found in bio-factories II and III. Sex ratio (at approximately 60% females) was satisfactory (in terms of suitability for release) across all bio-factories. For morphometric measurements, the body, abdomen and wing widths were similar in males; however, thorax width was greater in the males from bio-factory I; bio-factory III produced females with the highest body length; bio-factory I produced females with the greatest abdomen width; bio-factories III and IV produced females with the greatest wing length. Among the bio-factories studied, bio-factory IV produced the best quality C. flavipes, with respect to the greatest number of parasitoids per pupal mass (a mean of 57% more parasitoids) in a satisfactory sex ratio, and with the shortest developmental time, which facilitates faster rearing in the laboratory. Studies such as this, which assess the quality of a mass-produced C. flavipes, are crucial for the continued use of this parasitoid in controlling Diatraea saccharalis in the field.     

Effects of flight behaviour on body temperature and kinematics during inter-male mate competition in the solitary desert bee Centris pallida

Abstract. Body temperatures and kinematics are measured for male Centris pallida bees engaged in a variety of flight behaviours (hovering, patrolling, pursuit) at a nest aggregation site in the Sonoran Desert. The aim of the study is to test for evidence of thermoregulatory variation in convective heat loss and metabolic heat production and to assess the mechanisms of acceleration and forward flight in field conditions. Patrolling males have slightly (1–3 °C) cooler body temperatures than hoverers, despite similar wingbeat frequencies and larger body masses, suggesting that convective heat loss is likely to be greater during patrolling flight than during hovering. Comparisons of thorax and head temperature as a function of air temperature (T_a) indicate that C. pallida males are thermoregulating the head by increasing heat transfer from the thorax to the head at cool T_a. During patrolling flight and hovering, wingbeat frequency significantly decreases as T_a increases, indicating that variation in metabolic heat production contributes to thermal stability during these behaviours, as has been previously demonstrated for this species during flight in a metabolic chamber. However, wingbeat frequency during brief (1–2 s) pursuits is significantly higher than during other flight behaviours and independent of T_a. Unlike most other hovering insects, C. pallida males hover with extremely inclined stroke plane angles and nearly horizontal body angles, suggesting that its ability to vary flight speed depends on changes in wingbeat frequency and other kinematic mechanisms that are not yet described.     

Solitary and social bees as pollinators of Wahlenbergia (Campanulaceae): single-visit effectiveness, overnight sheltering and responses to flower colour

Solitary bees often form specialised mutualisms with particular plant species, while honeybees are considered to be relatively opportunistic foragers. Thus, it may be expected that solitary bees are more effective pollinators than honeybees when foraging on the same floral resource. To test this, we studied two Wahlenbergia species (Campanulaceae) in South Africa that are visited by both social honeybees and solitary bees, and which are shown here to be genetically self-incompatible and thus reliant on pollinator visits for seed production. Contrary to expectation, the solitary bee Lipotriches sp. (Halictidae) and social bee Apis mellifera (Apidae), which were the two most frequent visitors to flowers of the study species, were equally effective pollinators in terms of the consequences of single visits for fruit and seed set. Both bee species preferentially visited female phase flowers, which contain more nectar than male phase flowers. Male solitary bees of several genera frequently shelter overnight in flowers of both Wahlenbergia species, but temporal exclusion experiments showed that this behaviour makes little contribution to either seed production or pollen dispersal (estimated using a dye particle analogue). Manipulation of flower colour using a sunscreen that removed UV reflectance strongly reduced visits by both bee groups, while neither group responded to Wahlenbergia floral odour cues in choice tests. This study indicates that while flowers of Wahlenbergia cuspidata and W. krebsii are pollinated exclusively by bees, they are not under strong selection to specialise for pollination by any particular group of bees.     

Comparisons between the biology of two co-occurring species of whiting (Sillaginidae) in a large marine embayment

Synopsis We compare the biology of the tropical species Sillago analis and the temperate species Sillago schomburgkii in Shark Bay, a large subtropical marine embayment on the west coast of Australia. This environment constitutes approximately the southernmost and northernmost limits of the distributions of these two species, respectively. The annuli visible in sectioned otoliths of S. analis and S. schomburgkii form annually. Their numbers were thus used to age the individuals of these two species, which are morphologically very similar and live in the same habitats. Although the growth rates of S. analis and S. schomburgkii are very similar until maturity is attained, they subsequently diverge, with S. schomburgkii investing relatively more energy into somatic growth. The maximum total lengths and ages of both the females (320 mm, 6 years) and males (283 mm, 8 years) of S. analis were not as great as those of the females (383 mm, 9 years) and males (302 mm, 9 years) of S. schomburgkii. In Shark Bay, S. schomburgkii spawns earlier and longer than S. analis, i.e. August–December vs. January–March, which would result in the juveniles of these two species recruiting into nursery areas at different times. In addition, S. schomburgkii spawns earlier and for longer in Shark Bay than in temperate marine waters 800 km further south, presumably reflecting the fact that, in that subtropical embayment, water temperatures over which this species typically spawns are attained earlier and last for longer. However, although environmental conditions in Shark Bay and those temperate marine waters differ markedly, the growth of the corresponding sexes of S. schomburgkii in these two water bodies is similar.     

Deceptive pollination in two subspecies of Disa spathulata (Orchidaceae) differing in morphology and floral fragrance

The non-rewarding flowers of Disa spathulata (L.f) Sw. have a highly elongated scent-producing lip that varies in size and form between the two subspecies of this orchid. We investigated whether this unusual morphology is associated with the evolution of a specialized scent-based pollination system. Field observations carried out in South Africa showed that flowers of D. spathulata are pollinated solely by bees belonging to the genus Tetraloniella (Anthophoridae). Flowers of D. spathulata subsp. spathulata were visited by Tetraloniella brevikeraia, while flowers of D. spathulata subsp. tripartita (Lindl.). H. P. Linder were visited by Tetraloniella junodi. The floral fragrance of D. spathulata is emitted by the lip blade and differs markedly between the two subspecies (only 24 [42.1] of the 57 compounds identified were shared). Fragrance of D. spathulata subsp spathulata is dominated by the fatty acid derivatives decyl actetate, octanol, and decanol, while that of D. spathulata subsp. tripartita is dominated by the terpenoids (E) - ocimene, caryophyllene, and (E) nerolidol. Both male and female Tetraloniella bees were strongly attracted to cut flowers of D. spathulata subsp spathulata, even at sites where the orchid does not naturally occur. Bees visit the orchids repeatedly despite the lack of rewards in their flowers. Sexual deception in D. spathulata can be excluded by the attraction of both male and female bees, and yet the manner in which bees are attracted seems too intense and species-specific to be characterized as food-deception. This implies the existence in D. spathulata of a system of scent-based deception of insects that has not been described previously in plants.     

Extreme thermotolerance and behavioral induction of 70-kDa heat shock proteins and their encoding genes in honey bees

Foraging honey bees frequently leave the hive to gather pollen and nectar for the colony. This period of their lives is marked by periodic extremes of body temperature, metabolic expenditure, and flight muscle activity. Following ecologically relevant episodes of hyperthermia between 33°C and 50°C, heat shock protein 70 (Hsp70) expression and hsp70/hsc70-4 activity in brains of nonflying laboratory-held bees increased by only two to three times baseline at temperatures 46–50°C. Induction was undetectable in thoracic–flight muscles. Yet, thorax hsp70 mRNA (but not hsc70-4 mRNA) levels were up to ten times higher in flight-capable hive bees and foraging bees compared to 1-day-old, flight-incapable bees, while brain hsp70/hsc70-4 mRNA levels were low and varied little among behavioral groups. These data suggest honey bee tissues, especially flight muscles, are extremely thermotolerant. Furthermore, Hsp70 expression in the thoraces of flight-capable bees is probably flight-induced by oxidative and mechanical damage to flight muscle proteins rather than temperature.