No full-text available
To read the full-text of this research,
you can request a copy directly from the authors.
Foraging and courtship behaviour in males of the solitary bee Anthophora plumipes (Hymenoptera: Anthophoridae): Thermal physiology and the roles of body size
Abstract
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.
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.
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.
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.
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.
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.
... Male-male competition often led to the evolution of antagonistic male behaviours such as fighting (McElligott et al. 1998;Brown et al. 2007), territorial defence (Jakobsson 1988;Frohschammer and Heinze 2009) or mate guarding (Endo and Doi 2002;Foitzik et al. 2002). Body size (Villabos and Shelly 1991;Haley et al. 1994;Stone et al. 1995;Bel-Venner and Venner 2006;Robbins et al. 2009) and early emergence/arrival on the mating site (Wiklund and Solbreck 1982;Carchini et al. 2005;Saino et al. 2012) was also found to be under positive sexual selection. In some species, both traits are traded-off (Zonneveld 1996), whereas in others, body size and emergence time are positively associated, pointing to a higher fitness of early and large males (Yoshimura et al. 2003). ...
... Larger males mated more often (but not generally longer) than smaller ones. A higher reproductive success of larger males has been reported in other Hymenopterans (Villabos and Shelly 1991;Stone et al. 1995;Abell et al. 1999) as in many other animal taxa. Depending on the mating system and ecology of a species, a higher reproductive success of larger males can have different causes: first, females preferentially choose larger males as mating partners because body size signals a high fitness (female choice; e.g., Robbins et al. 2009). ...
... Depending on the mating system and ecology of a species, a higher reproductive success of larger males can have different causes: first, females preferentially choose larger males as mating partners because body size signals a high fitness (female choice; e.g., Robbins et al. 2009). Second, larger males have a higher chance of encountering females because they are better dispersers (Stone et al. 1995;Abell et al. 1999). Third, large males perform better in fighting and in the defence of territories or mating partners (Haley et al. 1994;Bel-Venner and Venner 2006;Pitnick et al. 2009;Breuer et al. 2012). ...
Sexual selection has led to male morphologies and behaviours that either increase male attractiveness or their success in male–male competition. We investigated male traits under selection in the ant Hypoponera opacior, in which wingless males mate with pupal queens inside their natal colony and guard their partners for hours. The lack of female choice and fights among adult males makes this species an ideal study system to investigate sexual selection in the absence of these selective forces. We hypothesised that males, which emerge first and live longer, should have a higher mating success because of more mating opportunities, reduced competition and the ability to kill pupal competitors. We recorded the number and length of matings and tested whether these measures of male-mating success were associated with emergence order, lifespan and body size. Indeed, early emerged males mated more often and longer than their later-emerging rivals. Furthermore, longer-lived and larger males obtained more matings. Body size might be important because larger males either produce more sperm or perform better in mounting females. We found no evidence for a trade-off between body size and emergence time. Moreover, male removal manipulations revealed that males quickly adapt their guarding behaviour to changes in the competitive environment. Under reduced competition, males guarded their partners for shorter periods. In conclusion, these sib-mating ant males are under selection to develop fast, to live long, to be large and to be able to respond to the competitive situation in the nest.
... The males of this species, which live up to 6 weeks, are easily identifiable by the long hairs on the mid-tarsi, from which the name of the species comes (Falk 2019). The mating system involves two alternative behaviours: when the density of sexual competitors is low, males are territorial over flower patches, while as the density increases, they patrol overlapping areas and show opportunistic polygyny (Stone et al. 1995). Males have good visual acuity and flying skills as they chase females in flight. ...
... Their activity is also highly dependent on ambient temperature, with flight activity starting at a minimum ambient temperature of 10-14°C and a thorax temperature of around 24°C (Stone 1993). Interestingly, thermoregulatory and flight abilitiesand therefore mating success -are highly dependent on body size, with larger males able to fly at lower temperatures and take prime position immediately after females (Stone et al. 1995). Given their dependency on ambient temperature and size for reproductive success, males of A. plumipes are a good model to test possible functional trait shifts due to urbanisation. ...
Urbanisation, leading to the reduction and fragmentation of green areas and an increase in temperature (urban heat island
effect), is known to be a strong driver of intraspecific phenotypic variation in wild bees. However, the effects of urbanisation on
many functionally relevant morphological traits are still unstudied or debated. Here, males of the ground-nesting, solitary bee
Anthophora plumipes were sampled at nine sites in the metropolitan city of Milan (Italy). The aim was to test variation in body
size, head width, compound eye size, ommatidia density, antenna length, median ocellus size, galea length (proxy for tongue
length), wing area, wing loading, wing aspect ratio, and wing fluctuating asymmetry along an urbanisation gradient. We found
some of these traits to significantly shift across the gradient. Bees in hotter (urban) areas had longer galea, agreeing with
previous community-level studies showing more long-tongued wild bee species in urban areas. This may suggest that
urbanisation filters for longer mouthparts at both species and individual level. Ocelli were larger in males from more urbanised
sites, perhaps improving navigation in more fragmented habitats. Finally, bees in greener (less urban) areas had also lower wing
fluctuating asymmetry, suggesting that urbanisation may act as an early-life stressor in A. plumipes. None of the other analysed
morphological traits varied with urbanisation, which contrasts (especially for body size and wing size) with several previous
studies on other bee species in urban contexts. Such patterns highlight how difficult is to draw generalisable trends regarding
the effects of urbanisation on wild bees. Nonetheless, this study provides the first evidence of such effects on rarely studied
morphological traits (mouthparts, ocelli) in male bees.
... Possible perennial mix additions that are attractive to bees (assuming that appropriate soil and light conditions for their growth exist, and with pollen percentage protein by mass where available) include Anemone nemorosa [31], Corydalis spp. [32,125], Pulmonaria officinalis [126,127], Alkanna tinctoria and Symphytum spp. (17.5% protein; [54]) [125,126], Lamium album and Lamium purpureum [119] and Erysimum spp. ...
... [32,125], Pulmonaria officinalis [126,127], Alkanna tinctoria and Symphytum spp. (17.5% protein; [54]) [125,126], Lamium album and Lamium purpureum [119] and Erysimum spp. [119]. ...
Planted meadows are increasingly used to improve the biodiversity and aesthetic amenity value of urban areas. Although many 'pollinator-friendly' seed mixes are available, the floral resources these provide to flower-visiting insects, and how these change through time, are largely unknown. Such data are necessary to compare the resources provided by alternative meadow seed mixes to each other and to other flowering habitats. We used quantitative surveys of over 2 million flowers to estimate the nectar and pollen resources offered by two exemplar commercial seed mixes (one annual, one perennial) and associated weeds grown as 300m 2 meadows across four UK cities, sampled at six time points between May and September 2013. Nectar sugar and pollen rewards per flower varied widely across 65 species surveyed, with native British weed species (including dandelion, Taraxacum agg.) contributing the top five nectar producers and two of the top ten pollen producers. Seed mix species yielding the highest rewards per flower included Leontodon hispidus, Centaurea cyanus and C. nigra for nectar, and Papaver rhoeas, Eschscholzia californica and Malva moschata for pollen. Perennial meadows produced up to 20x more nectar and up to 6x more pollen than annual meadows, which in turn produced far more than amenity grassland controls. Perennial meadows produced resources earlier in the year than annual meadows, but both seed mixes delivered very low resource levels early in the year and these were provided almost entirely by native weeds. Pollen volume per flower is well predicted statistically by floral morphology, and nectar sugar mass and pollen volume per unit area are correlated PLOS ONE |
... Whatever the reasons for spatial distribution of males, our results clearly show that males did not establish territories and instead adopt a typical scramble competition strategy (Thornill & Alcock, 1983). Among the aculeate Hymenoptera, when receptive females are scarce and widely dispersed, males often engage in scramble competition for mates (Thornhill & Alcock, 1983;Stone et al., 1995;Alcock & Kemp, 2005;Paxton et al., 2005). On the other hand, when females emerge in clusters or when resources attracting females are aggregated, males usually adopt a female defence or resource defence polygyny strategy through territoriality (Thornhill & Alcock, 1983). ...
... Indeed, a lack of advantage (Larsson & Tengö, 1989;Seidelmann, 1999) or even a disadvantage (Stoks, 2000;Moya-Larano et al., 2002) for large males scrambling to find mates have been reported. Larger males of the bee Anthophora plumipes Pallas, however, tolerate lower temperatures better and can court females under conditions in which smaller males cannot fly, and they can also displace smaller males from favoured flight positions immediately behind females (Stone et al., 1995). We found an effect of body size on patrolling behaviour, which may suggest a possible advantage for large males of N. viduata. ...
The mating systems of mutillid wasps have rarely been studied. Here we present information on the mating system of Nemka viduata. At a site in southern Spain, many males of this species were seen flying over host (digger wasp) nest aggregations while searching for females. Male activity was greatest in the early morning and late afternoon, when females were more active searching for hosts, and on days when relatively large numbers of females were active. Males were not territorial but instead attempted to find emerging females before their competitors. As many as six males might arrive at a receptive female more or less simultaneously. Struggles to control access to females continued until one male copulated with the female on the ground or carried it off in flight to a location away from rival males. Male size seems to affect the patrolling behaviour (number of patrolled sites), but there is little evidence of an advantage for larger males, as expected in a scramble competition mating system. Scramble competition mating systems often evolve in species in which large numbers of males compete for scarce receptive females, a factor that makes male territorial defence of large areas highly costly.
... Although macroecological investigations encompass a wide variety of theoretical and empirical approaches (Brown, 1995(Brown, , 1999, often a distinction is drawn between univariate, bivariate and multivariate perspectives, and integration follows from joint consideration of the insights obtained from each (e.g. McGill, 2003;Hui & McGeoch, 2008; E.P. Storch et al., 2008). ...
... Acknowledging that there is substantial feedback between body size, physiological traits and life history variables (Blanckenhorn, 2000b Body size correlates with mortality from abiotic factors such as starvation, desiccation and low temperature (Lighton et al., 1994;Arnett & Gotelli, 2003;Lehmann et al., 2006;Colinet, Vernon & Hance, 2007), predation intensity (Nylin & Gotthard, 1998), predator guild composition , food particle size (Holter & Scholtz, 2005), fecundity (Honík, 1993;Taylor, Anderson & Peckarsky, 1998), mating and reproductive success (Stone, Loder & Blackburn, 1995;Taylor et al., 1998), activity/foraging time (Stone, 1994;Cerdá & Retana, 2000), the outcome of intraspecific competition (Heinrich & Bartholomew, 1979), flight ability (Dudley, 2000a), and various aspects of morphology (e.g. Feener et al., 1988;Green, 1999;Shingleton et al., 2008;Eberhard, 2009). ...
Body size is a key feature of organisms and varies continuously because of the effects of natural selection on the size-dependency of resource acquisition and mortality rates. This review provides a critical and synthetic overview of body size variation in insects from a predominantly macroecological (large-scale temporal and spatial) perspective. Because of the importance of understanding the proximate determinants of adult size, it commences with a brief summary of the physiological mechanisms underlying adult body size and its variation, based mostly on findings for the model species Drosophila melanogaster and Manduca sexta. Variation in nutrition and temperature have variable effects on critical weight, the interval to cessation of growth (or terminal growth period) and growth rates, so influencing final adult size. Ontogenetic and phylogenetic variation in size, compensatory growth, scaling at the intra- and interspecific levels, sexual size dimorphism, and body size optimisation are then reviewed in light of their influences on individual and species body size frequency distributions. Explicit attention is given to evolutionary trends, including gigantism, Cope's rule and the rates at which size change has taken place, and to temporal ecological trends such as variation in size with succession and size-selectivity during the invasion process. Large-scale spatial variation in size at the intraspecific, interspecific and assemblage levels is considered, with special attention being given to the mechanisms proposed to underlie clinal variation in adult body size. Finally, areas particularly in need of additional research are identified.
... One of the best known is body size, related to mating success and biological viability in other groups of bees (Villalobos and Shelly 1991;Stone et al. 1995;Alcock 1996). However, there is no evidence that body size is related to perfume traits neither larger males have higher tness in orchid bees (Eltz et al. 2015). ...
Male orchid bees (Euglossini) collect volatile and semi-volatile compounds from the environment for storage and accumulation in specialized hind legs. Later, these compounds form a perfume blend used during courtship to lure conspecific females for mating. It has been proposed that male orchid bees perfume plays an important role as a sexual signaling trait involved in pre-mating isolation of species, functioning as an indicator of male genetic quality. Eulaema nigrita Lepeletier is a common species in both forested and woody savannah physiognomies (Cerrado sensu stricto) of the Brazilian savanna biome. By identifying the chemical composition of male E. nigrita perfume, we tested for differences in the bouquet chemical profile in populations from remnants of seasonal semideciduous forest and woody savanna. In addition, we assessed the relation between perfume complexity and morphological traits associated with size and age of males. Our analysis showed a low effect of physiognomies on the differences in the perfume chemical profile of sampled males. Nevertheless, we observed significant differences in the chemical profile of individuals from two seasonal semideciduous forest remnants, which suggests an environmental effect in individual bouquet. Wing wear as a morphological measure of age was positively related to perfume complexity, in agreement with the premise that perfumes from older individuals are indicators of survival capacity in male orchid bees.
... Similarly, other traits related to " tness" can be used as indirect evidence that demonstrates the role of fragrances as an indicator of male quality in reproduction events. One of the best known is body size, related to mating success and biological viability in other groups of bees (Villalobos and Shelly 1991; Stone et al. 1995;Alcock 1996). ...
Male orchid bees (Euglossini) collect volatile and semi-volatile compounds from the environment for storage and accumulation in specialized hind legs. Later, these compounds form a perfume blend used during courtship to lure conspecific females for mating. It has been proposed that male orchid bee perfume plays an important role as a sexual signaling trait involved in pre-mating isolation of species, functioning as an indicator of male genetic quality. Eulaema nigrita Lepeletier is a common species in both forested and woody savannah physiognomies (Cerrado sensu stricto) of the Brazilian savanna biome. By identifying the chemical composition of male El. nigrita perfume, we tested for differences in the bouquet chemical profile in populations from remnants of seasonal semideciduous forest and woody savanna. In addition, we assessed the relation between perfume complexity and morphological traits associated with size and age of males. Our analysis showed a low effect of physiognomies on the differences in the perfume chemical profile of sampled males. Nevertheless, we observed significant differences in the chemical profile of individuals from two seasonal semideciduous forest remnants, which a priori suggests an environmental effect in individual bouquet. Wing wear as a morphological measure of age was positively related to perfume complexity, in agreement with the premise that fragrances from older individuals are indicators of survival capacity in male orchid bees.
... The composition of the local food plant spectrum and their typical growth habits creates very different selective environments for males that at one time or place may have the potential to monopolize resources when competitor density per patch is low (many small patches), but diminishingly so when patch size and rival density increases. The abandonment of an aggressive resource defense in favor of a scramble competition polygyny at high competitor densities has also been observed in the bee Anthophora plumipes (Stone et al. 1995) and seed bug Neacoryphus bicrucis (McLain 1992). In any case, the resource requirements of females to provision brood cells should provide opportunities for males of all sizes to search for mates. ...
The wool carder bee Anthidium manicatum is one textbook example of resource defense polygyny among solitary bees, known for intense male–male competition, forced copulations, and the extreme form of interspecific territoriality toward other flower visitors. This mating system depends on the spatial structure of the defended resource and requires several adaptations in males. The allocation of patches with host plants as well as male body size and phenology was analyzed over 3 years in the diverse habitat of a botanical garden. Anthidium manicatum males searched in groups up to 12 individuals a wide diversity of patches with various food plants of foraging females. Territories were established in small high-quality patches only. Males abandoned aggressive and territorial behavior in large patches. Available patches were occupied by males of the various body size fractions independently of each other according to patch profitability. The higher competitive weight of large males in small patches compared to spacious ones was balanced by an opposing correlation of patch profitability. Although the mating system in A. manicatum is clearly a resource defense polygyny, males were found to be plastic in their behavior, and territoriality was not consistently observed. Mate acquiring tactics, be they territory holder (bourgeois), sneaker, floater, or scrambler for mating, can be considered to be different behavioral phenotypes within one environmentally sensitive conditional strategy.
Significance statement
Territoriality is a rare and derived pattern in solitary bee mating behavior. In most cases of territoriality, males defend rendezvous places to meet freshly emerged, virgin females. While this type of mating behavior fits still into the framework of ancestral monandry of aculeate Hymenoptera, the continually polyandric resource defense polygyny found in the genus Anthidium is highly derived. Males occupy flower resources exploited for larval provisions and extort copulations from provisioning nesting females. Territoriality in Anthidium does not lead to a monopolization of females, the exclusion of many competitors from reproduction, and a reduction of sperm competition as is typical for resource-based mating systems. Contrary, Anthidium is a highly promiscuous species and both males and females are lifelong engaged in copulations with multiple mates. Also, the allocation of the resource fundamental to the defense polygyny was found to be more fairly balanced than expected. This study diversifies the mating system of anthidiine bees and demonstrates unusually high plasticity in the resource allocation of a territorial species.
... The size of adult insects may influence mating ability and is an essential parameter of reproductive success (Anderson, 1994;Blanckenhorn et al., 1998;Bonduriansky & Rowe, 2003;Fox & Czesak, 2006;Helinski & Harrington, 2011;Partridge et al., 1983;Stone et al., 1995;Teng & Zhang, 2009;Zamudio et al., 1995). In Tephritidae, the influence of size on mating success was also studied; within medfly, it was demonstrated that large males have higher mating success than small males (Blay & Yuval, 1999;Churchill-Stanland et al., 1986;Dukas, 2005;Kaspi et al., 2000;Orozco & Lopez, 1993;. ...
The Sterile Insect Technique (SIT) is a method of pest control used in the management of Ceratitis capitata (Medfly). This technique is based on the mating and transfer of ejaculate sperm to the wild female population from released sterile males. This transfer will induce sterility in the female through sperm dominant lethal mutations. Consequently, the factors influencing sperm transfer and sperm storage are issues that must be understood to enable successful SIT implementation. Age, size, nutritional status, copula duration, and irradiation are the most important factors that affect sperm transfer and storage. Inconsistent results from studies investigating the influence of these factors on sperm transfer/storage may be due to, the strain of fly studied and variability in the sexual experience of the flies used. In this review, we highlight the interaction between these factors, issues that still need to be addressed, and propose future directions for the improvement of medfly males’ sexual performance and the successful application of SIT against this pest.
... Why do some Acacia species release their pollen around dawn, and others in the afternoon? Activity of bees and other insects can be strongly affected by daily microclimatic fluctuations (Willmer 1983, Herrera 1990, Stone 1994, Stone et al. 1995, and this may place constraints on the pollinators available to acacias flowering at particular times of day. Is there a best time to dehisce? ...
... Inset: statistical results. 1980;Stone 1994;Stone et al. 1995;Kim 1997;Bosch & Vicens 2006;Seidelmann 2014). The vegetation structure of JK Campus-restored includes mass flowering plants, introduced as part of its restoration process, which may explain the larger proportion of females and the nests with larger internal diameters. ...
The reasons for the decline of bee diversity and abundance include the destruction and loss of natural habitats. Protected areas are created for biodiversity conservation, but these areas vary strongly in their level of vegetation disturbance. Using trap-nests, we assessed changes in solitary bee abundance, richness, and composition in areas ranging from naturally conserved to degraded. Solitary bees were sampled during an 18-month period in three areas of southeastern Brazil: a preserved area in Rio Preto State Park – PERP; a restored/altered area with exotic plants at the Federal University of the Jequitinhonha and Mucuri Valleys – JK Campus; and a degraded area in Biribiri State Park – PEBi. A total of seven species of bees built 115 nests. In the degraded area, only two nests were built. Abundance of built nests was higher in the preserved area (PERP), but diversity was higher in the restored area (JK Campus). Our results show that the solitary bee population responds positively to habitat complexity (local scale). The presence of a diverse solitary bee fauna in the restored area indicates that altered areas should also be protected as suitable areas for re-colonization of cavity-nesting bees.
... For example a study in the highly endemic cape floristic region of South Africa showed that climate change-induced impacts on species ranges varied from range expansion of 5-50% for two species of bees to substantial range contractions, between 32% and 99%, in another six species [48]. Seasonal shifts within [49] and across species [50,51] have also been detected in regions with distinct seasons and may simulate species turnover when local climatic conditions change. While the loss of specialist species due to environmental change may not have direct impacts on crop pollinator community, it entails lower rates of ecosystem processes, and some functions performed by specialists may not be carried out at all [43], potentially leading to greater biodiversity loss and ecosystem instability in the long run. ...
Our review looks at pollinator conservation and highlights the differences in approach between managing for pollination services and preserving pollinator diversity. We argue that ecosystem service management does not equal biodiversity conservation, and that maintaining species diversity is crucial in providing ecosystem resilience in the face of future environmental change. Management and policy measures therefore need to focus on species not just in human dominated landscapes but need to benefit wider diversity of species including those in specialised habitats. We argue that only by adopting a holistic ecosystem approach we can ensure the conservation and sustainable use of biodiversity and ecosystem services in the long-term.
... Note, however, that factors other than sexual selection for fighting ability can enable relatively large males to achieve a reproductive advantage. For example, large males of the apid Anthophora plumipes are more active under cool temperatures than smaller rivals (Stone, Loder, & Blackburn, 1995). Moreover, intense competition for mates does not always result in a reversed sexual size dimorphism in which some males are larger than the largest females. ...
Many attributes of male bees can be interpreted as the products of sexual selection, such as large male body size in species, in which male–male fighting is intense. Moreover, the diversity of locations chosen by searching males, the occurrence of alternative mating tactics, and the timing of mate searching also appear to have been shaped by sexual selection. Even errors in sexual discrimination made by males of many species may stem from a sexual selection in favor of individuals with a very low threshold for sexual activity. The female choice component of sexual selection has also had evolutionary consequences for males inasmuch as the mating success of different male tactics depends on whether females mate just once or with several different individuals. Females may select mating partners passively by permitting male competition to determine which individual inseminates them or actively by making the fertilization of their eggs dependent upon elements of male performance before, during, or after copulation.
... Given the lack of direct evidence for female choice, we employ an indirect approach. We test whether aspects of perfume phenotypes, i.e., the quantity and complexity of individual perfume loads, are related to certain traits that are related to mating success or viability fitness in other organisms (Archer et al. 2012;Bonduriansky et al. 2008;Friedl and Klump 2005;Moller and Alatalo 1999), including other groups of bees (Alcock 1996;Amin et al. 2012;Jaffe and Moritz 2010;Stone et al. 1995;Villalobos and Shelly 1991). These traits are male size (related to competitive strength), fluctuating asymmetry (related to developmental stability), and wing wear (related to age, i.e., survival ability). ...
Male neotropical orchid bees collect volatile substances from their environment and compose complex, species-specific odour blends in leg pouches. These perfumes are accumulated by the males over time and are exposed during pre-mating display. It has been hypothesized that perfumes are indicators of male genotypic quality and that females chose mates by the quality or intensity of their odour. Because direct experimental proof is lacking, we investigated whether the amount or complexity of male perfumes is related to male (1) size, (2) fluctuating (left–right) asymmetry, or (3) age, traits that are known to be sexually selected in other animals. We measured left and right forewings (cell node distances for 1 and 2, wing wear for 3) of Euglossa dilemma and Euglossa viridissima collected during dry and rainy seasons on the Yucatán peninsula, Mexico (total N = 768). Wing size was not related to the quantity or the complexity of perfume extracts in either species after excluding the effect of season, which positively affected both size and perfume load in E. dilemma. Wing asymmetry had also no effect, except in rainy season E. viridissima, where it was positively (not negatively) correlated with the quantity of perfume. Wing wear, an established age indicator of orchid bees, had the only consistent overall effect, being positively correlated with perfume amount and complexity in both species. This is in agreement with the idea that perfumes are an honest indicator of male survival capacity.
... Females also exhibited some variation in those features, but they were less conspicuous than in males. Intraspecific variation in body size as well as in color is not uncommon, and it is known to affect foraging as well as mating success in insects, including some anthophorine bees (e.g., Stone et al., 1995;Alcock et al., 2005;Chown and Gaston, 2010). In both bees and wasps, extensive facial yellow markings are more commonly found in males than in females. ...
We provide information on the nesting behavior, natural enemies, seasonality, and host plants of Anthophora (Dasymegilla) waltoni Cockerell at mid-elevations in southern China (Yunnan Province). Bees nested singly inside loose mortar joints of building walls. Nests were shallow and consisted of an unlined, unbranched tunnel ending in a single cell. Surveys conducted every two or three days from February to April, 2013, showed that the studied population is univoltine, protandrous, and active in early spring. Females were more commonly observed than males, and no bees were observed flying by the end of April. Both sexes of A. waltoni were often collected on flowers of Vicia cracca Linnaeus (Fabaceae).
... In most ectotherms, performance increases up to a thermal optimum and then falls away as body temperature approaches the critical maximum (Angilletta et al., 2002b). At lower temperatures, changes in locomotor performance can induce shifts in antipredator behaviours (Keogh and De Serto, 1994; Brodie and Russell, 1999; Whitaker and Shine, 1999; Cooper, 2000), foraging behaviour (Dıáz, 1994), general activity (Melville and Schulte, 2001), social behaviours (Mautz et al., 1992; Stone et al., 1995; Stutt and Willmer, 1998) and interspecific aggression (Magoulick and Wilzbach , 1998). Interestingly, several of these studies identified that locomotor performance is optimised within a range of temperatures rather than a single optimum (Mautz et al., 1992; Angilletta et al., 2002a). ...
1. I tested if individual variation in preferred body temperature (PBT) covaried with behaviours and male phenotypic traits (size and colouration) in Pseudemoia entrecasteauxii. 2. Individuals varied in their PBT and this variation was repeatable across days. PBT was not related to body size but males with orange ventral colour had higher PBT. 3. Males with orange ventral colour were more aggressive and dominated males with white venters. 4. Male behaviours were correlated and they covaried with PBT. Correlated behaviours such as these may be related to a shy–bold continuum.
... Under such circumstances, plants can either evolve tolerance of competition (e.g., through increased floral longevity:Levin 1978;Motten 1986;Rathcke 1988;Ashman and Schoen 1994;Ashman 2000), or partition the activity of shared pollinators on finer, daily, timescales (Levin and Anderson 1970;Ollerton and Lack 1992). Because pollinators commonly track pollen resource availability in daily time (Frankie et al. 1983;Stone 1994Stone , 1995Stone et al. 1995Stone et al. , 1996Stone et al. , 1998Stone et al. , 1999aHerrera 1997;Willmer and Stone 2004), and regularly remove pollen from their bodies (Gilbert 1981;Roubik 1989), the evolution of species-specific times of pollen release (dehiscence) has the potential to reduce competition for both pollinator visits and pollen purity (Armbruster and Herzig 1984;Armbruster 1985;Stone et al. 1996Stone et al. , 1998). The importance of daily structuring in competition for pollination is unclear, and most analyses to date lack the necessary resolution to examine daily patterns. ...
Competition for pollination is thought to be an important factor structuring flowering in many plant communities, particularly among plant taxa with morphologically similar and easily accessible flowers. We examined the potential for heterospecific pollen transfer (HPT) in a community of four Acacia species in a highly seasonal tropical habitat in Mexico. Partitioning of pollen flow among sympatric species appears to be achieved, in part, through segregation of flowering in seasonal time, and interspecific differences in pollinator guilds. However, two coflowering species (Acacia macracantha and Acacia angustissima) shared multiple flower visitors, raising the possibility of HPT. Each of these coflowering species showed high intraspecific daily synchrony in pollen release, but dehisce at different times of day. Pollinators rapidly harvested available pollen from one species before abandoning it to visit the flowers of the second later in the day. The activity of shared pollinators, predominantly bees, is thus structured throughout the day, and potential for HPT reduced. Suggestive evidence in favour of a resource partitioning explanation for this pattern is provided by the fact that A. macracantha showed significantly greater intraspecific synchrony when coflowering with a potential competitor (A. angustissima) than when flowering alone. We discuss our results in light of previous work on coflowering acacia assemblages in Tanzania and Australia.
... Understanding the causes of temporal structure requires understanding the relative contributions of sampling error and the top-down and bottom-up influences driving the timing of flower–visitor interactions . Revealing temporal structure imposed by topdown aspects of visitor biology requires finer resolution activity pattern and behavioral data than collected in this study (e.g., Stone et al. 1995 Stone et al. , 1998), but our data do reveal bottom-up constraints imposed by floral resources . Floral imposition of bottom-up control is sometimes very obvious: in our networks flowers of the morning glory Ipomoea kituiensis (Convolvulaceae) opened from 08:00 and were shriveled by noon, while flowers of Sida ovata and S. schimperiana (Malvaceae) opened only from 11:00 to 15:00. ...
Ecological interaction networks are a valuable approach to understanding plant-pollinator interactions at the community level. Highly structured daily activity patterns are a feature of the biology of many flower visitors, particularly provisioning female bees, which often visit different floral sources at different times. Such temporal structure implies that presence/absence and relative abundance of specific flower-visitor interactions (links) in interaction networks may be highly sensitive to the daily timing of data collection. Further, relative timing of interactions is central to their possible role in competition or facilitation of seed set among coflowering plants sharing pollinators. To date, however, no study has examined the network impacts of daily temporal variation in visitor activity at a community scale. Here we use temporally structured sampling to examine the consequences of daily activity patterns upon network properties using fully quantified flower-visitor interaction data for a Kenyan savanna habitat. Interactions were sampled at four sequential three-hour time intervals between 06:00 and 18:00, across multiple seasonal time points for two sampling sites. In all data sets the richness and relative abundance of links depended critically on when during the day visitation was observed. Permutation-based null modeling revealed significant temporal structure across daily time intervals at three of the four seasonal time points, driven primarily by patterns in bee activity. This sensitivity of network structure shows the need to consider daily time in network sampling design, both to maximize the probability of sampling links relevant to plant reproductive success and to facilitate appropriate interpretation of interspecific relationships. Our data also suggest that daily structuring at a community level could reduce indirect competitive interactions when coflowering plants share pollinators, as is commonly observed during flowering in highly seasonal habitats.
... Under such environmental conditions, Hylaeus often appear sluggish and will bask in the sun (E. Wilson, personal observation); this may be in part to utilize solar radiation to warm their flight muscles (Stone and Willmer 1989, Larsson 1991, Stone et al. 1995). Given the contrasting thermoregulatory abilities, it seems likely that yellowjackets encounter torpid Hylaeus on cool mornings when they are vulnerable to predation. ...
Variation in invasion success may result from the divergent evolutionary histories of introduced species compared to those of native taxa. The vulnerability of native biotas to ecological disruption may be especially great on oceanic islands invaded by continental species with unique ecological traits. In part because Hawaii lacks native eusocial insects, social invaders may threaten endemic taxa that are ecologically similar but solitary. Using a combination of field manipulations, molecular analyses, physiological data, and behavioral assays, we identify the mechanisms underlying the displacement of two genera of native solitary Hymenoptera in Hawaii by a social continental invader, the western yellowjacket (Vespula pensylvanica). Experimental removal of V. pensylvanica colonies resulted in increased densities of native Hymenoptera. Endemic Hylaeus bees directly suffer through predation by yellowjackets, and perhaps as a consequence, avoid floral resources occupied by V. pensylvanica. Native Nesodynerus wasps also avoid V. pensylvanica but are negatively affected by yellowjackets not through predation, but through exploitative competition for caterpillar prey. Displacement of native solitary Hymenoptera may be heightened by the ability of V. pensylvanica to prey upon and scavenge honey bees and to rob their honey stores, resources unavailable to endemic bees and wasps because of their specialized niches. Our study provides a unique example of an ecologically generalized social invader that restructures native assemblages of solitary Hymenoptera by interacting with endemic taxa on multiple trophic levels.
... The answer to the former question is that intraspecific diversity in male mating behav- iour has frequently been documented (e.g. Alcock et al., 1978;Stone et al., 1995;Willmer and Stone, 2005). Table II contains some spe- cies in two categories. ...
Considerable interspecific diversity exists among bees in the rendezvous sites where males search for females and in the behaviours employed by males in their efforts to secure matings. I present an evolutionary framework in which to interpret this variation, and highlight the importance for the framework of (i) the distribution of receptive (typically immediate post-emergence) females, which ordinarily translates into the distribution of nests, and (ii) the density of competing males. Other than the highly polyandrous honey bees (Apis), most female bees are thought to be monandrous, though genetic data with which to support this view are generally lacking. Given the opportunity, male bees are typically polygamous. I highlight intraspecific diversity in rendezvous site, male behaviour and mating system, which is in part predicted from the conceptual framework. Finally, I suggest that inbreeding may be far more widespread among bees than has hitherto been considered the case.
Male orchid bees collect volatile and semi-volatile compounds from the environment for storage and accumulation in specialized hind legs. Later, these compounds form a perfume blend used during courtship to lure conspecific females for mating. Male orchid bees perfume has been suggested to play an important role as a sexual signaling trait involved in pre-mating isolation of species, functioning as an indicator of male genetic quality. Eulaema nigrita Lepeletier (Apidae: Euglossini) is a common species in both forested and woody savanna (Cerrado stricto sensu) physiognomies of the Brazilian savanna biome. By identifying the chemical composition of male E. nigrita perfume, we tested for differences in the bouquet chemical profile in populations from remnants of seasonal semideciduous forest and woody savanna. In addition, we assessed the relation between perfume complexity and morphological traits associated with size and age of males. Our analysis showed a low effect of physiognomies on differences in the perfume chemical profile of sampled males. Nevertheless, we observed significant differences in the chemical profile of individuals from two seasonal semideciduous forest remnants, which suggests an environmental effect in individual bouquet. Wing wear measurements were positively related to perfume complexity, consistent with the premise that perfumes from older individuals are indicators of survival capacity in male orchid bees.
p>This thesis investigates size variation in the worker caste of social bumblebees ( Bombus spp.) by examining various morphological and behavioural characteristics in relation to body size. I confirm size variation in the worker castes of 23 Bombus species and that foragers are, on average, larger than in-nest bees and that workers of pocket-making species vary more than pollen-storers. It is believed that larger bees are suited to foraging tasks and small bees to in-nest tasks, large bees being able to collect more forage per trip. I confirm that larger workers gather nectar more efficiently. Workers show behavioural plasticity: smaller workers occasionally forage and larger bees often perform in-nest tasks and are superior brood carers. By manipulating the worker size distributions in artificial nests, I show that a workforce of large bees is superior to one of the small workers in terms of colony growth. Larger workers are supposedly better adapted to colder conditions than smaller workers. I show that bumblebees from cold regions have longer thoracic setae than those from hot regions and that, within species, foraging workers of races from cooler regions are often found to be larger than those from warmer regions. I also show that, in southern England, ambient temperature has no influence on nectar foraging efficiency or on the size of forager, but that pollen foraging trips are made in warmer conditions. Bumblebees obtain from flowers. I show that different plant species attract foragers of different mean size, that larger bees have longer tongues and handle deeper flowers more quickly and shallower flowers more slowly than smaller workers. That size variation enables a colony to efficiently harvest nectar and pollen from a variety of floral resources is the most encouraging adaptive explanation for size variation in bumblebee workers thus far.</p
Workers of many species of social Hymenoptera have functional ovaries and are capable of laying haploid, unfertilized eggs, at least in the absence of a queen. Except for honeybees, it remains largely unknown if worker‐produced males have the same quality as queen‐produced males and if workers benefit in direct fitness by producing their sons. Previous studies in the monogynous ant Temnothorax crassispinus revealed that a high proportion of males in natural and laboratory colonies are worker offspring. Here, we compare longevity, body size, sperm length, and sperm viability between queen‐ and worker‐produced males. We either split queenright colonies in queenright and queenless halves or removed the queen from a fraction of the queenright colonies and then examined the newly produced males. Male quality traits varied considerably among colonies but differed only slightly between queen‐ and worker‐produced males. Worker‐produced males outnumbered queen‐produced males and also had a longer life span, but under certain rearing conditions sperm from queen‐produced males had a higher viability.
RESUMEN Durante los años 1994, 1995 y 1997 se ha observado y verificado la existencia en Anthidium florentinum Fabricius (1775) de un sistema de apareamiento del tipo poligínico de defensa del recurso, sobre Rubus ulmifolius Schott, 1818 (Rosaceae). Esta especie muestra interesantes patrones en el comportamiento territorial derivado de este sistema de apareamiento. Se describen detalladamente estos patrones, en especial la relación entre el tamaño de los machos, el comportamiento que siguen y el éxito en el apareamiento, así como la defensa intraespecffica e interespecífica de] recurso, y el comportamiento reproductor. Por último, se revisan las similitudes y diferencias en las pautas de comportamien-to de esta y otras especies de Anthidium y se discuten algunos de los puntos más controvertidos derivados de la territorialidad, en relación a la visión general del comportamiento de otras abejas territoriales. Palabras clave: Anthidium, poliginia de defensa del recurso, comportamiento territorial.
ABSTRACT Territorial behaviour associate with the resource defense polygyny of Anthidium florentinum (Fabricius, 1775) (Hymenoptera, Megachilidae). The resource-based mating system of Anthidium florentinum (Fabricius, 1775) has been studied in Central Spain at patches of ftowermg Rubus ulmifolius Schott, 1818 (Rosaceae). This solitary bee displays a resource defense polygyny, like other congeneric species, with interesting behaviour patterns. In this paper, these patterns are showed and described, and specially, the relation among male body size, behaviour exhibited and reproductive success, as well as the aggressive and reproductive behaviour.
Fragmentation and loss of natural habitats are major threats to many bee species. Large, long-distance flying bees are predicted to be more efficient in utilizing resources and at the same time may function as important pollinators in a fragmented landscape. Using mark-recapture experiments, this study evaluates the movement of bees belonging to the "large, long-tongue" guild in a threatened, fragmented habitat. Bee movement between the sampling plots was limited, despite high recapture proportions within the plots. A maximum likelihood model has estimated a high degree (60 % of all marked bees) of site fidelity to the source plots and a mean traveling distance of 357 m for the bees that left the plots. Additional observations on the bees' foraging behavior suggest that some anthophorine bee species can be important pollinators in the studied habitat. We suggest that the bees' site fidelity and flower constancy are the main causes for their observed conservative movement pattern. solitary bees / habitat fragmentation / movement patterns / foraging behavior / site fidelity
Attempts by males of the solitary beeAnthophora plumipesPallas (Apoidea: Anthophoridae) to mate with foraging females at flowers of comfrey,Symphytum orientale, were studied. The mating system conformed to scramble competition polygyny. After an initial period of nectar foraging in the morning, males patrolled the comfrey and attempted to copulate with any females they encountered. The majority of females were unreceptive, and were able to reject the male after his initial pounce, which knocked the female to the ground. Females could experience rates of attempted copulation exceeding once every 3s, which significantly reduced their rate of visiting flowers and hence prolonged the period required to provision nest cells. Females responded to male harassment by evasive flight and physical repulsion and also by a more general change in foraging behaviour. Females abandoned flowers on the outer parts of plants frequented by males, and foraged from flowers inside the plant's growth where males did not often venture. Consideration of sugar rewards per flower and handling times suggested that male harassment halved the rate of reward for females from exposed outer flowers. When males were removed from the site, females abandoned the inner flowers and foraged from more profitable outer flowers. When males were released, the original pattern of behaviour was quickly re-established. During poor weather the ability of females to provision cells became marginal, and male harassment could thus have significant consequences for female fitness.
Under sexual conflict, males evolve traits to increase their mating and reproductive success that impose costs on females. Females evolve counteradaptations to resist males and reduce those costs. Sexual harassment is a form of sexual conflict in which males make repeated, costly attempts to mate. Costs to female foraging or predation risk have been measured in several species, but quantitative measurements of direct fitness costs are rare. In the alfalfa leafcutting bee, Megachile rotundata (Fabricius; Hymenoptera: Megachilidae), males harass females, and females resist all mating attempts. We placed bees in large, outdoor cages with various male-biased sex ratios. Harassment rate, nest progression, offspring production, temperature, and food availability were measured daily for 7 days. Harassment rates were highest at intermediate sex ratios. Harassment reduced the number of foraging trips and increased the duration of foraging trips made by females. Females produced offspring at a slower rate when subjected to higher rates of harassment. This shows a direct link from sex ratio to harassment to female fitness under natural conditions. We also discuss an alternative explanation that female resistance is a mechanism for mate choice for high-quality males, which would require that indirect benefits accrue through either daughters or grandsons, because all sons in haplodiploid species arise from unfertilized eggs.
This chapter discusses the behavioral, ecological, and physiological determinants of the activity patterns of bees. An activity pattern is the change in levels of a particular activity through time. Bees are an excellent taxon within which to investigate both the issues of constraint and currency. Their physiological constraints are better understood than those of almost any other invertebrate, allowing an unparalleled opportunity to integrate physiology into behavioral ecology. Many bees require elevated body temperatures (Tb) to fly, and hence thermal properties of their environment significantly constrain their activity. However, many species are heterothermic, being able to elevate their Tb endothermically when necessary, which gives some degree of escape from the usual thermal constraints on other entirely ectothermic insects. The chapter discusses the factors structuring activity patterns in a wide range of bees. Observed activity patterns are generated by interactions between properties of the bees themselves (intrinsic factors) and properties of their environment (extrinsic factors). Intrinsic factors include physiological differences between species and between the sexes. Extrinsic factors are the fluctuating properties of the environment, and are either abiotic or biotic.
Females of Dawson's burrowing bee (Amegilla dawsoni) are receptive to the males as they emerge but have become unreceptive by the time they begin to nest. In addition, there is a single emergence period per year lasting about a month. These factors are predicted to lead to protandry and males of Dawson's burrowing bee do tend to emerge earlier in the annual flight season than females. Moreover, even during a single day, emerging males tend to precede females. The degree of protandry, however, is size-dependent, with smaller males tending to precede larger ones, both over the course of the flight season and on any given day. Because small males are at a disadvantage in the fights that occur for females, the earlier emergence of minor males may be a sexually selected response that reduces the likelihood that they will be displaced from potential mates by larger rivals.
Most animals have to bridge some distances in space and time to provide all resources necessary for survival. Little is known about how the local and regional management of tropical agroforestry systems, differing in the availability of food resources and suitable nesting sites, determine foraging trip duration and density of bees and wasps (Hymenoptera Aculeata).
In this study, foraging trip duration and brood‐cell density (in trap nests exposed for a 15‐month period) were analysed for three species, which represent three guilds in 24 agroforestry systems in Central Sulawesi (Indonesia): the pollinator Heriades sp. aff. fulvescens (Apidae), the cacao caterpillar‐hunting predator Rhynchium haemorrhoidale umeroatrum (Eumenidae) and the spider‐hunting predator Auplopus levicarinatus (Pompilidae). The agroforestry systems were characterized by plant species richness, blossom cover of herbs, light intensity and distance from the nearest natural forest.
The correlation of foraging time to bee and wasp density showed the relative importance of food and nest‐site availability for the pollinator and the spider‐hunting predator, because both parameters are correlated with light intensity in the agroforestry systems. In contrast, foraging time and nest density of the eumenid predator were not correlated because of the distance between high quantities of food resources (in sites with dense cacao plants) and nesting sites (in adjacent natural forests).
The eumenid response to local and regional agroforestry management illustrates that species may survive only in landscapes that permit access to a multiple set of resources. Accordingly, habitat evaluations using only foraging time may lead to wrong conclusions, as key drivers of population dynamics may not be inside but outside the local systems, emphasizing the need for a landscape approach.
Males of Dawson’s burrowing bees (Amegilla dawsoni) search for virgin females at three locations: (1) open clay patches where females are emerging from underground brood cells,
(2) the vegetated peripheral zone adjacent to emergence areas (through which females pass after emerging), and (3) clusters
of flowering plants, which are often some distance from emergence areas. Males of Dawson’s burrowing bees exhibit a size dimorphism
with large major and small minors. Major males patrol only the open emergence sites, whereas minor males may be found in all
three locations. Although most females are mounted and presumably mated immediately upon emergence, some are not, and these
females make up a pool of potential mates for the small males patrolling the peripheral zone and flower patches. The density
of males at emergence sites and the probability of male-male aggression change over the course of a day and over the entire
flight season. When the level of competition is low, some minor males hunt for mates at emergence areas, where potential mates
are relatively numerous. But when the presence of many large rivals makes it unlikely that a small male can avoid being displaced
from emerging females, minors make the best of a bad job by shifting to areas where majors are absent.
The ecology of a species strongly influences the strategies with which males and females maximize their lifetime reproductive success. When males and females do not invest equally in offspring, the sex with the higher parental investment becomes a rare resource for the other. The spatial and temporal distribution of the limiting sex forms the basis of the mating system. In nest-constructing Aculeata such as the red mason bee, Osmia rufa, females perform intensive brood care, whereas males do not invest in their offspring but instead compete for access to mates. Receptive females of this species are widely distributed and do not assemble at certain places. Therefore, territorial behavior is not an advantageous mating tactic for males, which instead search for females within individual home ranges usually centered around food plants. The unmarked or defended home ranges of different males may completely overlap. Competitive searching leads to a random distribution of matings among males that is largely independent of body size. The mating system of O. rufa can be described as scramble competition polygyny.
The optimum maternal investment per offspring is determined by the relationship between the investment per offspring and offspring fitness. In the European beewolf, Philanthus triangulum, a solitary mass-provisioning sphecid wasp, offspring size correlates with the amount of provisions. We investigated whether the reproductive success of adult males depends on body size in a way that would influence the allocation of parental investment. Since the mating success of P. triangulum males cannot be determined by observation in the field, we assessed the influence of male size on characteristics of their territories, territorial behaviour and life history traits. Territory size was weakly correlated with male size, but a measure of territory quality (number of female nests in the vicinity) was independent of male size. Neither the duration of ownership nor the intensity of scent marking was correlated with male size. Territory owners were slightly smaller than nonterritorial males. The absolute amount of fat was positively correlated with size but, owing to allometric relationships, the energetic equivalent of the fat store appeared to be independent of size. Life span was not significantly influenced by body size under four different conditions (with and without food in the laboratory, in an outdoor flight cage and in the field). We discuss the discrepancy between these results and other studies that have mostly reported advantages to large males. We suggest that in noncontact male-male interactions, as seen in the European beewolf, body size might not be the key determinant for success in contests. We conclude that there is no evidence for a strong size dependence of male reproductive success. Thus the reproductive success of male progeny probably does not depend on parental investment in a way that would influence the investment allocation of females. Copyright 2000 The Association for the Study of Animal Behaviour.
Food limitation can reduce reproductive success directly, as well as indirectly, if foraging imposes a risk of predation or parasitism. The solitary bee Osmia pumila suffers brood parasitism by the cleptoparasitic wasp Sapyga centrata, which enters the host nest to oviposit while the female bee is away. I studied foraging and reproduction of O. pumila nesting within cages stocked with rich or sparse floral resources, and the presence or absence of S. centrata to test (1) the response of nesting female O. pumila to food shortages, (2) the response of nesting female O. pumila to the presence of parasites, and (3) whether brood produced under scarce resources are more likely to be parasitized by S. centrata. The rate of brood cell production was significantly lower in cages with sparse floral resources, although females in sparse cages did not produce significantly fewer brood cells overall. Sapyga centrata did not influence the rate of brood cell production, but females exposed to the cleptoparasites had marginally significantly lower reproductive output. Nests in parasite cages had significantly fewer brood cells than those in parasite free cages. The mean duration of foraging bouts made by female O. pumila in sparse cages was not significantly longer than that in rich cages. O. pumila spent less time in the nest between pollen and nectar foraging bouts in sparse cages with S. centrata than those in other cages suggesting that these individuals made more frequent food foraging trips. Despite the weak effects of parasites and bloom density on foraging behavior, O. pumila brood cells experienced a 5-fold higher probability of parasitism by S. centrata in cages with sparse bloom than in those with rich bloom [corrected]. These results support the hypothesis that indirect effects of food scarcity increase O. pumila susceptibility to brood parasitism, although the exact mechanism is not entirely clear yet.
Throughout the animal kingdom the innate nature of basic behaviour routines suggests that the underlying neuronal substrates necessary for their execution are genetically determined and developmentally programmed. Complex innate behaviours require proper timing and ordering of individual component behaviours. In Drosophila melanogaster, analyses of combinations of mutations of the fruitless (fru) gene have shown that male-specific isoforms (Fru(M)) of the Fru transcription factor are necessary for proper execution of all steps of the innate courtship ritual. Here, we eliminate Fru(M) expression in one group of about 60 neurons in the Drosophila central nervous system and observe severely contracted courtship behaviour, including rapid courtship initiation, absence of orienting and tapping, and the simultaneous occurrence of wing vibration, licking and attempted copulation. Our results identify a small group of median bundle neurons, that in wild-type Drosophila appropriately trigger the sequential execution of the component behaviours that constitute the Drosophila courtship ritual.
Groups of males of the nearctic Anthophora abrupta Say (Apoidea: Anthophorini) chew parsnip tissue, collect the odorous juice in unique adsorptive labral mustaches of fine, flattened hairs, and apply it to surrounding objects, apparently mixed with their mandibular gland secretion. Such perfumed areas outline an oval flight path. This resembles the fragrance-collecting and territorial behavior of male neotropical orchid bees (Apidae: Euglossinae).
Summary 1. We assess the importance of body mass and the minimum ambient temperature at which foraging occurs in determining the warm-up rates and thoracic temperatures in flight at an air temperature of 22 °C of 55 species of bee (Hymenoptera: Apoidea) from six families adapted to a variety of thermal environments. 2. To control for the effects of taxonomic differences in the relationships between these variables, we use multiple regression incorporated in the phylogen- etic regression method developed by Grafen (1989). 3. The prediction made by May (1976) that for very small heterotherms warm- up rate will correlate positively with body mass is confirmed when the effects of phylogeny and the thermal environment to which the bee is adapted have been controlled for. The relationship between warm-up rate and body mass within the Apoidea is thus not an extension to lower body masses of the relationship found for heterothermic vertebrates. 4. Having controlled for the effects of body mass in our analyses, we demonstrate that bees able to fly at lower ambient temperatures have higher thoracic temperatures and warm-up rates than bees adapted to wanner environ- ments. 5. There is some suggestion that kleptoparasitic bees, being freed from the need to forage in order to provision cells, have lower warm-up rates than provisioning species. 6. The significance of these relationships in the ecology of bees is discussed in relation to studies of body temperatures and warm-up rates in bees and other insects.
Summary 1. This study examines variation in thoracic temperatures, rates of pre-flight warm-up and heat loss in the solitary bee Anthophora plumipes(Hymenoptera; Anthophoridae). 2. Thoracic temperatures were measured both during free flight in the field and during tethered flight in the laboratory, over a range of ambient temperatures. These two techniques give independent measures of thermoregulat ory ability. In terms of the gradient of thoracic temperature on ambient temperature, thermoregulation by A. plumipes is more effective before flight than during flight. 3. Warm-up rates and body temperatures correlate positively with body mass, while mass-specific rates of heat loss correlate negatively with body mass. Larger bees are significantly more likely to achieve flight temperatures at low ambient temperatures. 4. Simultaneous measurement of thoracic and abdominal temperatures shows that A. plumipes is capable of regulating heat flow between thorax and abdomen. Accelerated thoracic cooling is only demonstrated at high ambient temperatures. 5. Anthophora plumipes is able to f ly at low ambient temperatures by tolerating thoracic temperatures as low as 25˚C, reducing the metabolic expense of endothermic activity. 6. Rates of heat generation and loss are used to calculate the thermal power generated by A. plumipes and the total energetic cost of warm-up under different thermal conditions. The power generated increases with thoracic temperature excess and ambient temperature. The total cost of warm-up correlates negatively with ambient temperature.
1. The foraging activities of the papilionid butterflies Ornithoptera priamus poseidon and Papilio ulysses, and the solitary bee Amegilla sapiens (Apoidea, Anthophoridae) on the shrub Stachytarpheta mutabilis were studied in highland Papua New Guinea. 2. The insects' activity patterns were analysed at three sites with differing diurnal microclimate variation. O. priamus and A. sapiens foraged in the morning (after a period of basking and wing-whirring) and late afternoon when temperatures were well below daily maxima, whereas P. ulysses showed foraging peaks during the hottest part of the day. 3. Site choice by all 3 species appeared to be determined primarily by temperature, but within the limits imposed by temperature, nectar supplies probably determined which site was visited. 4. P. ulysses showed interspersed foraging and courtship behaviour, and no behavioural switching was observed for this species. At high temperatures, both O. priamus and A. sapiens ceased foraging and showed territorial and courtship behaviour. This behavioural change allowed avoidance of heat stress, and occurred even when nectar supplies were maintained at high levels. 5. Thermal effects on behavioural switching in these insects are compared with related phenomena in other bees and butterflies.
Body temperatures during free flight in the field, warm-up rates during pre-flight warm-up, and temperatures during tethered flight are measured for four tropical solitary bee species at three sites of differing altitude in Papua New Guinea. All four species are capable of endothermic preflight warm-up; three species give slopes of thoracic temperature on ambient temperature of significantly less than 1, indicating regulation of thoracic temperature. In the kleptoparasitic Coelioxys spp. (Megachilidae) and Thyreus quadrimaculatus (Anthophoridae), warm-up rates and thoracic temperatures in flight are low by comparison with the two provisioning species Creightonella frontalis (Megachilidae) and Amegilla sapiens (Anthophoridae). In both C. frontalis and A. sapiens thoracic temperatures correlate positively and significantly with both ambient temperature and body mass. In A. sapiens, body mass increases with altitude; this can be interpreted as a response to lower ambient temperatures at higher altitude, an example of Bergmann's rule. In both A. sapiens and C. frontalis populations at higher altitude have higher thoracic temperatures independent of differences of body mass, suggestive of additional morphological or physiological adaptation to lower ambient temperatures. In A. sapiens there is no qualitative difference in body temperatures between males and females after controlling for body mass, while male C. frontalis have significantly lower thoracic temperatures than females of the species. This difference between A. sapiens and C. frontalis is discussed with reference to variation in mating systems found in the Apoidea.
Solitary bees (Centris pallida Fox; Hymenoptera: Anthophoridae) were observed during diurnal activity in a hot desert environment. Females foraged throughout the day in flowering paloverde trees; males divided their time between foraging and hovering in search of receptive females. Thoracic temperatures () of both sexes were high. Foraging males and females had averaging 44-46 and 43-45 C, respectively. The of hovering males averaged about 44 C in the early morning and 47 C from midmorning until activity ceased. Thoracic temperatures of both sexes changed by 3-4 C over an ambient temperature () range of 25-40 C. Abdominal temperature () was consistently about 4.5 C above , and there was no indication of augmented heat transfer between thorax and abdomen at high . Maximum tolerable was 50-51 C. Evidence for evaporative heat loss was equivocal for artificially heated bees, but free-flying animals showed little indication of use of evaporative cooling. Energy metabolism (measured as oxygen consumption) during flight (1. 1 W/g thorax) resembled that of other endothermic insects of similar mass; mass-specific thermal conductance (0.034 W/[g thorax - C]) was slightly larger than predicted from body mass. Centris pallida probably controls body temperature by reducing flight at high . Nevertheless, hovering males often attain within 2-3 C of lethality.
1. The insect wing appears to degenerate with use, so that wing wear increases with increased flight activity. Wing degeneration may affect an insect's mortality. 2. The rate of mortality is known to increase with age in worker bumble bees. This study examines whether wing wear can account for mortality in foraging bumble bees. 3. Workers in eight wild-foraging colonies of bumble bee (Bombus melanopygus Nylander) were divided into two treatment groups: clipped (the outer margin of each forewing trimmed, reducing wing surface area by an average of 18%) and unclipped controls. The mortality and behaviour of foraging individuals was assessed with colony watches and night censuses. 4. Treatment did not detectably affect the proportion of foraging bees, the lengths of foraging or within-nest bouts of foragers, or pollen load sizes, but wing-clipping did cause bees with the greatest amounts of initial wing wear to stop foraging. 5. Mortality was positively related to natural wing wear among unclipped bees. In addition, relative to control foragers, foragers with clipped wing margins experienced significantly reduced life expectancies. 6. These results support the hypothesis that wing wear is a proximate factor responsible for an increase in mortality in older workers. The wing-wear hypothesis therefore provides one functional reason for foragers to adopt a currency that maximizes the ratio of net benefit to cost (i.e. `efficiency').
Results add support to the argument that large males of C. pallida are at a great reproductive advantage over their smaller rivals in dense emergence sites. The success of large males persists from year to year and throughout any one flight season. The large male advantage can be quantified by contrasting the expected with the observed frequency of mating by males of different head- widths. A 5.7 mm head-width male is c7 times as likely to mate as a male of average size (5.1 mm). Females actively control the size of their progeny, producing added diversity in males because it is in their reproductive interest to do so. Small males faced with numerous larger opponents may switch reproduction tactics altogether to hover at waiting stations for passing females. -from Author
Bell (1986) predicted that plants should be able to grow a proportion of their flowers that do not produce any nectar, which thus escape the costs of nectar production, while gaining the benefits of insect pollination because most insect visitors either cannot discriminate, or would lose too much time attempting to do so. He worked out an ESS model predicting the proportion of 'cheating' flowers and discriminating insects: the proportion of cheaters should be D/H, where D is the discrimination time, and H the handling time of the insect visitors. We have tested Bell's hypothesis using the flowers of Cerinthe major L. (Boraginaceae) visited by Anthophora plumipes (Pallas) (Hymenoptera, Anthophoridae). From measurements of D and H, we predicted that 73% of flowers should be cheaters. Inspection of the ranked nectar production of individual flowers on individual days shows that between 75 and 85% are relatively low nectar producers. Reasons for this pattern are explored; in particular, the way in which nectar production varies with flower age may constitute a mechanism by which plants can play a mixed strategy.
Anthophora edwardsii Cresson, a vernal bee occurring throughout western North America, possesses several unusual biological characteristics for the genus. In an extensive nesting aggregation near Coalinga, California, females constructed their single-celled nests in compacted sandy soil on a flat hilltop. The nests were about 7.5 to 10 cm deep, lacked turrets, and were plugged with soil from the cell cap to the surface on completion. The bees do not spin cocoons. They pupate in the autumn and overwinter as adults in their natal cells. Females exhibit an apparent communal aggressive behavior towards intruders at the nest site. These bees show interpopulation variation in characteristics such as the number of cells per nest and whether they nest in flat ground or in vertical banks. Biotic enemies, other than mold, account for 18 to 37% mortality at three nest sites. A parasitic bee of the genus Melecta caused losses of 21 to 27% at two nest sites.
1. Energetic considerations of preflight warm-up in bees predict that bees should warm-up as fast as they can and that larger species should be able to warm-up more rapidly. Larger species should also be able to maintain higher thoracic temperatures during flight. These predictions are examined for 19 species in the solitary bee genus Anthopora (Hymenoptera; Apoidea; Anthophoridae). 2. The genus Anthophora has uniformly high warm-up rates and body temperatures in flight, and the highest warm-up rates measured in any heterotherm. 3. Under standard conditions, both warm-up rates and thoracic temperatures during flight increase with body mass in Anthophora, in keeping with the prediction that heterotherms should minimize the energetic cost of warm-up. 4. Having controlled for body mass effects, Anthophora species able to maintain flight activity at lower ambient temperatures have higher warm-up rates and body temperatures in flight, showing that even in the smallest endotherms there may still be room for selective modification within the limits imposed by body size. 5. Differences in warm-up rates between subgenera within Anthophora suggest that endothermic abilities have diverged in response to differences in the thermal environments which are the centres of diversity for the subgenera. 6. Similar importance of both body mass and the thermal environment in which the species is active is demonstrated over 40 bee species in six families. Because species over a wide taxonomic range do not constitute independent data points, phylogenetic effects in these analyses are controlled for using Grafen's phylogenetic regression. 7. The evolution of high levels of endothermy in Anthophora is discussed with reference to patterns of nectar secretion found in the arid environments which are current centres of diversity for the genus.
Attempts by males of the solitary bee Anthophora plumipes Pallas (Apoidea: Anthophoridae) to mate with foraging females at flowers of comfrey, Symphytum orientale, were studied. The mating system conformed to scramble competition polygyny. After an initial period of nectar foraging in the morning, males patrolled the comfrey and attempted to copulate with any females they encountered. The majority of females were unreceptive, and were able to reject the male after his initial pounce, which knocked the female to the ground. Females could experience rates of attempted copulation exceeding once every 3 s, which significantly reduced their rate of visiting flowers and hence prolonged the period required to provision nest cells. Females responded to male harassment by evasive flight and physical repulsion and also by a more general change in foraging behaviour. Females abandoned flowers on the outer parts of plants frequented by males, and foraged from flowers inside the plant's growth where males did not often venture. Consideration of sugar rewards per flower and handling times suggested that male harassment halved the rate of reward for females from exposed outer flowers. When males were removed from the site, females abandoned the inner flowers and foraged from more profitable outer flowers. When males were released, the original pattern of behaviour was quickly re-established. During poor weather the ability of females to provision cells became marginal, and male harassment could thus have significant consequences for female fitness.
A solitary anthophorid bee (Anthophora sp. nov.) in the Sinai desert showed sexually dimorphic diurnal activity patterns, whereby males defended territories containing clumps of the flower Alkanna orientalis between 0815 and 1430 hours while females fed exclusively on these same flowers only in the early morning up to 0900 hours and from about 1400 hours with virtually no male-female overlap on any one particular plant. Thus males defended the floral resources through long periods when conspecific females were absent. Female nests were usually within the male territories, however, and females copulated mainly with resident males. This system therefore has some similarities to resource defence polygyny, with territoriality functioning for mating advantage. The male strategy, however, was inappropriate merely to secure matings, as these occurred principally in the early morning and thereafter territories could have been abandoned for the day without loss of copulations; but by maintaining patrols throughout the day, a male would additionally ensure that there were better quality floral reserves in areas where his own females would feed to stock that day's nest-cells, and thereby he achieved matings with better-resourced females. The behaviour of the male Anthophora is therefore a form of territoriality that achieves potentially larger offspring, thus serving as pre-zygotic paternal investment.
Attempts by males of the solitary beeAnthophora plumipesPallas (Apoidea: Anthophoridae) to mate with foraging females at flowers of comfrey,Symphytum orientale, were studied. The mating system conformed to scramble competition polygyny. After an initial period of nectar foraging in the morning, males patrolled the comfrey and attempted to copulate with any females they encountered. The majority of females were unreceptive, and were able to reject the male after his initial pounce, which knocked the female to the ground. Females could experience rates of attempted copulation exceeding once every 3s, which significantly reduced their rate of visiting flowers and hence prolonged the period required to provision nest cells. Females responded to male harassment by evasive flight and physical repulsion and also by a more general change in foraging behaviour. Females abandoned flowers on the outer parts of plants frequented by males, and foraged from flowers inside the plant's growth where males did not often venture. Consideration of sugar rewards per flower and handling times suggested that male harassment halved the rate of reward for females from exposed outer flowers. When males were removed from the site, females abandoned the inner flowers and foraged from more profitable outer flowers. When males were released, the original pattern of behaviour was quickly re-established. During poor weather the ability of females to provision cells became marginal, and male harassment could thus have significant consequences for female fitness.
Patterns of activity at a large nesting aggregation and at foraging sites are described for females of the solitary bee Anthophora plumipes (Pallas). Changes in the quality and quantity of the resource collected by females provisioning cells are related to variation in female body mass and microclimate.
Activity at the nest site demonstrated relationships with aspects of the thermal environment experienced by A.plumipes. Measures of temperature showing significant relationships differed for females in different stages in the nesting cycle exhibiting characteristically different behaviour patterns.
Larger females emerged from nest tunnels earlier in the morning and provisioned cells at lower ambient temperatures than smaller individuals. Body size therefore predicts reproductive success at low ambient temperatures.
Pollen and nectar loads carried by females increased with ambient temperature. Because only one cell is completed per day and the size of offspring is determined by the quality and quantity of resource provided by the mother, the body size of individuals emerging in the following season will depend on interactions between climate and body size, in addition to any heritable component.
Variation in activity levels at foraging sites is attributed not only to thermal considerations, but also to variation in the quality of rewards available at different floral sources.
1. Cerceris arenaria (L.) females nest in sandy burrows, foraging for prey weevils to stock cells in which eggs are laid. Nests are easily distinguished and observed, and tunnel diameter correlates well with the occupying females' size (range 68–128 mg).
2. Foraging depends on ambient temperature and occurs only during periods of insolation. Large females can forage earlier in the day, and usually make a greater number of successful trips per day, than smaller ones.
3. Size differences of females affect heat exchange and resultant temperature excesses (Tex). At the highest ambient temperatures Tex for the largest wasps can exceed physiological tolerances, thus reducing foraging activity relative to the smaller females.
4. On very hot days wasps forage more frequently for nectar, probably to reduce their water deficits. Larval water balance is controlled less directly, by the architecture of burrows and the nature of cell walls and contents.
5. Temperature effects on foraging and nesting bring about a roughly two-fold increase in success as cell provisioners (and hence egg-layers) of larger over smaller females. Possible countering effects of very hot weather and of parasitism, either of which might favour small females in some conditions, are discussed.
1. In a 3-year study of the solitary bee Colletes cunicularius L. in Sweden, average body size and population density fluctuated greatly between years.
2. In this protandrous population, females mated just once and the sex ratio was slightly male biased. Males were smaller than females.
3. Size assortative mating (homogamy), associated with an increase in population density during the central days of female emergence and mating, was observed in two out of three years. Homogamy was also observed in pairs with remating males.
4. Most of the mating males had emerged the day they mated, but 42% were older. We found no support for a general large-male mating advantage.
5. Weight of emerging females and mating males were negatively correlated with ground temperature, indicating thermoregulatory influence on the process of sexual selection in this species.
Males of the wool-carder bee, Anthidium manicatum, patrol clumps of garden plants. Females of this species visit these plants for pollen, nectar, and pubescence; they also mate there. Females are polyandrous, with intervals between copulations as short as 35 s. Patrolling males defend their territories (0.1–1.3 m2) against other males and against other species of flower-visiting insects. Honey bees may be rendered unable to fly by the attacks of A. manicatum.Territory owners perform exploratory flights to other males' territories, changing territories often (median ownership 4–7 days; maximum 30 days) and flying up to 450 m to establish new territories. Territorial usurpations are nearly always by larger males.Female visitation rate is significantly correlated with number of flowers on a territory. The head size of territory-owner males shows significant correlation with territorial quality (measured by number of flowers, not area) and thus with number of female visits and copulatory opportunities. Some males fail to maintain territories and instead attempt to forage and copulate in other males' territories while the owners are otherwise occupied. Nonowner males are significantly smaller than owners, forage less often and from fewer flowers, and achieve significantly fewer copulations than owners. Females, however, do not reject smaller, nonowner males at a higher rate than they do larger, owner males; their choice for male size appears to be indirect, based instead on choice of food resource.The interval between a copulation and the male's next attempt with a different female is not shorter than that involving the same female. Males do not escort just-mated females about their teritories, as observed in Anthidium maculosum. Territorial behavior in this species most likely evolved through intrasexual competition for reproductive success which led to sexual dimorphism. The defense of a resourcebased territory is the mechanism used by a male to maximize his reproductive potential.
Comparative studies of the mating behaviour of coexisting but seasonally separated species may reveal microclimatic effects on insect mating systems. I studied two species of gregarious Hymenoptera, an early spring bee (Colletes cunicularius, Colletidae) and a digger wasp (Bembix rostrata, Sphecidae) emerging later in summer, during their mating periods in Sweden. Both species occupy sandy and sunny habitats, males are protandrous and polygynous, whereas females are monandrous. During mate searching males normally spent more than 50% of their active time in flight, patrolling the ground for emerging virgin females. In the bee, male mate searching was constrained by low ambient temperatures, in the wasp mainly by high temperatures. Measurements of thoracic flight temperatures indicated interspecific differences in thermoregulation between these two ectothermic species. While the bee raised its thoracic flight muscle temperature faster than through passive heating, the wasp was found to depress temperatures excess during flight at very high ambient temperatures, although it still had higher (2 ×) temperature excess values as compared to the bee. Large males were at an advantage during low ambient temperature, although for different reasons in the two species. Large bees were successful because they could generate enough heat to fly, whilst large wasps depressed their body temperature and avoided overheating only when it was not too hot. Some male wasps switched from patrolling to territoriality as a mate acquisition tactic during low male density and high ambient temperatures, probably to avoid overheating during flight.
Many elements of the behavioural repertoire of ectothermic animals depend on body temperature. Under differing thermal conditions, behaviours in insects, reptiles and other terrestrial ectotherms may therefore vary widely, and in any given thermal regime there may be simple physical and physiological character differences between individuals that lead to a predictable variation of behaviour. Where mating behaviour patterns are involved, digferential fitness may result. Recent studies show that the interactions of physiology with behavioural ecology should be a fruitful area for future research.
Preface Acknowledgments Part I. Introduction: 1. Approaches to tropical bee biology 2. Diversity of tropical bees Part II. Foraging and Pollination: 3. Resources gathered by bees 4. Mechanisms of resource collection 5. Foraging and flight activity 6. Pollination ecology Part III. Nesting and Reproduction Biology: 7. Bee nests 8. Nest predators, associates and defense 9. Mating and brood production Part IV. Community Ecology: 10. Seasonality, abundance and flower preference 11. Composition of bee assemblages 12. Roles of bees in communities References Index Appendices.
Thesis (M.S.)--Washington State University. Bibliography: leaves 27-28.
Thesis (Ph. D.)--Utah State University, Dept. of Zoology, 1963. Includes bibliographical references.
The view is examined that the adaptive value of conventional aspects of fighting behaviour is for assessment of relative RHP (resource holding power) of the combatants. Outcomes of aggressive disputes should be decided by each individual's fitness budget available for expenditure during a fight (determined by the fitness difference between adoption of alternative strategies, escalation or withdrawal without escalation) and on the rate of expenditure of the fitness budget if escalation occurs (determined by the RHPs of the combatants). Thus response thresholds for alternative strategies (“assessments”) will be determined by natural selection on a basis of which opponent is likely to expend its fitness budget first, should escalation occur. This “loser” should retreat (before escalation) and the winner should stay in possession of the resource. Many aggressive decisions depend on whether one is a resource holder, or an attacker. Assuming the RHP of the combatants to be equal, there are many instances of fitness pay-off imbalances between holder and attacker which should weight the dispute outcome in favour of one or other opponent by allowing it a greater expendable fitness budget. Usually the weighting favours the holder; the attacker therefore needs a correspondingly higher RHP before it may be expected to win. This is not invariably the case, and much observed data fits the predictions of this sort of model. If assessments are perfect and budget expenditure rates exactly predictable, then there would never seem to be any case for escalation. Escalation can be explained in terms of injury inflictions (expenditures) occurring as discrete events; i.e. as “bouts” won or lost during fighting. Assessment can give only a probabilistic prediction of the outcome of a bout. A simple model is developed to investigate escalation situations. Each combatant assesses relative RHP; this correlates with an absolute probability of winning the next bout (cabs). The stake played for is infliction of loss of RHP and is determined by the fitness budgets of the opponents. (Each individual plays for the withdrawal of its opponent.) This defines a critical probability of winning (ccrit) for each combatant, above which escalation is the favourable strategy (cabs > ccrit) and below which withdrawal is favourable (cabs < ccrit). Escalation should occur only where cabs-ccrit is positive for both combatants. This model gives predictions compatible with the observations, indicating that RHP loss alone can be adequate to explain withdrawal: escalation behaviour. Withdrawal tendency will be increased by low searching costs. Escalations should be restricted to closely matched RHP opponents if RHP disparity is the major imbalance. Outside the “escalation range” of a given individual, the higher RHP individual wins and the lower one loses (i.e. it should withdraw after conventional display). RHP disparity and holder: attacker imbalance should both interact to shape the observed pattern, though their relative importances will depend on species and situation. In some instances selection may favour immediate withdrawal from an occupied territory even without assessment of RHP.
Ausgewählte Untersuchungen zur Repro-duktionsbiologie der Solitärbienen-ArtenAnthophora acervorum(Linné 1758) undOsmia rufa(Linné 1758)
- U Thalmann
Ausgewahlte Untersuchungen zur Repro-duktionsbiologie der Solitarbienen-Arten (Linne, 1758) und (Linne, 1758)
- U Thalmann
Behavioural and physiological thermoregulation in solitary bees
- G N Stone
- Stone G.N.
- Stone G.N.
Notes on the life history, parasitcs and inquiline associates of A.abrupta, with some comparisons with the habits of certain other Anthophorinae
- Frison T.H.
The ethology of Andrena erythronii with comparative data on other species (Andrcnidae)
- Michcner C.D.
Biological observations on Anthophora urbana urbana Cresson
- Mayer D.F.
Nesting biology of Anthophora abrupta (Hymenoptera, Anthophoridae)
- Norden B.B.