Musculature, nervous system and glands of pregenital abdominal segments of the female of Nomia melanderi Ckll. (Hymenopters, Apoidea)

The gross anatomy of muscles, the topography of nerve tissues, and the histology of the pregenital abdominal glands of Nomia melanderi Ckll. are reported in detail. The movable and fixed points of muscle attachment were utilized in establishing a system of nomenclature for a typical abdominal segment. Names of nervules correspond to those of the tissues they innervate. The points of attachment of muscles of the fifth abdominal segment are essentially the same in both Nomia and Apis, except for the second tergo-sternal muscle which, in Nomia, has shifted its point of movable attachment to the membranous integument in front of the intersegmental membrane gland where it helps in relasing glandular secretion. The general plan of the nerves in the fifth abdominal segment in Nomia is more diffuse than in Apis, but there is no difficulty in establishing homology between the nervules of the two species. A pair of intersegmental stretch organs was found in abdominal segments 3–6. Glands of the sixth intersegmental membrane possess a reservoir with peripheral pouches both of which are absent in those of the fifth. Both types of glands have neither closing nor opening mechanisms, and neither is innervated. Release of glandular secretion is accomplished by the action of the tergo-sternal muscles.     

Fine structure of the intersegmental membrane glands of the sixth abdominal sternum of female Nomia melanderi (Hymenoptera,Apoidea)

The fine structure of the intersegmental glands of the sixth abdominal sternum in 1-week old females of Nomia melanderi is presented. The plasma membrane of the secretory cell is unfolded in many places and is covered by a basement membrane. The microvillous surface is invaginated to form a rather long sinuous cavity. The endoplasm is almost entirely filled by secretory granules. Many secretory granules are located close to the inner surface of the invaginated plasma membrane. The invagination contains a porous ductule, apparently of cuticulin origin, that is connected directly with the inner layer of the transport duct of the duct-forming cell. This type of arrangement allows the direct flow of the secretory substance to the outside in a continuous way. The cylindrical duct-forming cell, besides having typical cell organelles, contains a cuticular transport duct. This duct is composed of a thin cuticulin layer surrounded by a rather thick epicuticular one. The results suggest that the secretory cell has two secretory cycles. The first occurs while the gland is differentiating (at the pupal stage) and is involved in secretion of the cuticulin that forms the porous ductule. The second cycle, which starts by the beginning of nesting, is involved in the secretion of a substance that is carried to the outside via the transport duct of the duct-forming cell.     

The evolution of food-producing glands in eusocial bees (Apoidea,Hymenoptera)1

A food-producing role for cephalic exocrine glands has arisen independently in both taxa of highly eusocial bees, Apis and Meliponini. With several exceptions, there is little evidence that food is produced by glands of solitary bees or by most bees at lower levels of sociality. We suggest that this association with sociality is due to four adaptive features of these glands: (1) food from the glands allows feces from queens and larvae to have a small volume, (2) the queen's fecundity can be increased, (3) nutrient recovery via cannibalism can be facilitated, and (4) rearing of emergency replacement queens is accelerated. Acceleration of the rearing of other castes and of queens in the normal process of colony fission is not clearly an advantage ascribed to these glands. Trophic eggs produced by meliponine colony workers are analogous to the secretions from food-producing glands in Meliponini and Apis workers.     

Terga1 and inter–tergal metasomal glands of male braconine wasps (Insecta, Hymenoptera, Braconidae)

The distribution of cxocrine tergal and inter–tergal glands among males of more than sixty genera of Braconinac has been surveyed. The morphology of the glands and their reservoirs is described. Two gland types were found. Large, sac–like inter–tergal glands, usually most prominent between the 7th and 8th abdominal tergites, were found in all members of the Braconini, Adeshini, Aphrastobraconini, Glyptomorphini and the Atanycolus and Merinotus groups of genera. Inter–tergal glands were generally considerably smaller in members of the Bathyaulacini, Coeloidini. Euurobraconini and Iphiaulacini. Small pore–plate, vase–like or tubular tergal glands opening on the medio–lateral or antero–lateral selerotized parts of the 8th metasomal tergite were found in many genera, but were largely absent from members of the Braconini, Euurobraconini and Iphiaulacini ss. The phylogenetic implications of these findings are discussed.     

Mandibular glands of male European beewolves, Philanthus triangulum (Hymenoptera, Crabronidae)

Males of a solitary digger wasp, the European beewolf, Philanthus triangulum, possess large mandibular glands that have been reported to produce a scent marking pheromone. We analysed the morphology and ultrastructure of these glands using light microscopy as well as scanning and transmission electron microscopy. The paired glands are located laterally in the head and each side consists of a larger and a smaller part. Both parts possess a collecting duct each with distinct openings at the mandible base. However, the collecting duct of the larger part is additionally connected to the pharynx through a lateral extension. The collecting ducts are bordered by a monolayered epithelium lined with cuticle that exhibits conspicuous ramified protuberances. About 1400 acini consisting of class 3 gland cells surround the ducts and are connected to them through conducting canals. The main components in the cytoplasm of these gland cells are mitochondria, well-developed smooth endoplasmatic reticulum, and electron lucent vesicles suggesting a high secretory activity. The connection between the large gland parts and the pharynx suggests that the secretion of the mandibular glands might not only be delivered directly onto the mandibles but might also be transported to and stored in the postpharyngeal gland.     

Diversity and function of male antennal glands in Cynipoidea (Hymenoptera)

The functional anatomy of antennal glands located either on the 3rd or on the 3rd and 4th antennomeres in males of several species of cynipoids was investigated. SEM observations revealed variously modified antennomeres with elevated plates, tyloids and excavated areas. In all the cases, the antennomeres are equipped with cuticular pores, corresponding internally to cuticular ducts. TEM studies showed the presence of type III integumentary glands, as classified by Noirot & Quennedey. Each glandular unit is made up of an innermost secretory cell, producing the secretion, and an outermost canal cell, producing the evacuating duct. The secretion passes through the duct and reaches the cuticular pores, concentrated in a ventro-lateral portion of the antennomere called the 'release and spread structure'. Both in Cynipidae and in Eucoilinae (Figitidae), the courtship behaviour includes a pre-copulatory phase characterized by intense antennal stroking. Bioassays in the eucoilins Leptopilina boulardi and L. heterotoma showed that these glands are the production site of a contact sex recognition pheromone, necessary for the female to accept the male.     

A cladistic analysis of the nomadine bees (Hymenoptera: Apoidea)

Abstract. This study compares the results of Rozen's cladistic analysis of the larvae of fifteen genera of cleptoparasitic bees in the subfamily Nomadinae with an independent data set of adult characters for the same genera. Adult characters exhibited considerably higher levels of homoplasy and poorer resolution of cladistic relationships, with multiple equally parsimonious cladograms. However, comparison of a Nelson consensus tree based on adult characters with the cladogram based on larval characters reveals three components consistently supported in both analyses (the tribes Epeolini and Ammobatini, and Neopasites + Neolarra) , one component supported only by adult characters (Isepeolus + Protepeolus) , and one terminal component supported only by larval characters (Nomada + Ammobatini), as well as several more inclusive groupings based on larval characters that are difficult to compare with the adult consensus tree because it shows so much less resolution. When adult and larval characters are combined in a single data matrix, the resulting cladogram closely resembles the cladogram based on larval characters alone, although levels of homoplasy are considerably higher than in the larval analysis.
A preliminary analysis of adult characters for thirty-four genera in the Nomadinae also exhibited high levels of homoplasy and very large numbers of equally parsimonious cladograms. Nevertheless, certain consistent monophyletic groupings, most notably the Epeolini and Ammobatini, were also supported in this analysis. The one currently recognized tribe whose monophyly has received no support from any analysis is the Nomadini.
The relevance of these phylogenetic hypotheses to our understanding of host associations and variable features of egg morphology and oviposition behaviour in nomadine bees is briefly discussed.     

Quantitative Aspects of Antenna Grooming in Bees (Apoidea: Hymenoptera)

Antenna grooming in more than 100 species of bees (Apoidea), representing 34 genera of the 7 major bee families is recorded and quantitatively analysed. Most species of bees fall into one of two groups with respect to repetitive antenna cleaning: “Uniscrapers” predominantly clean their antennae with one stroke, “biscrapers” mostly with two subsequent strokes. Uniscrapers are more consequent in their behaviour than biscrapers. Most biscrapers occasionally clean their antenna with one or three strokes. Individual variation in the ratio of stroke repetition is considerably larger in bi- than in uniscrapers. In several species males and females differ with respect to their antenna cleaning behaviour, females tend to be more uniscraping. Most species of the families Colletidae, Halictidae, and Andrenidae, as well as the species of the genera Ceratina and Nomada (Anthophoridae) are biscrapers. Almost all species of Melittidae, Megachilidae, Apidae, and Anthophoridae (except Ceratina and Nomada) are uniscrapers. Bees with an antenna cleaner with ancestral (plesiomorphic) morphology are mostly, but not always, biscrapers, those with a derived antenna cleaner are always uniscrapers. Bees with a derived antenna cleaner perform on average less cleaning actions and strokes than those with an ancestral antenna cleaner. Uniscrapers with an ancestral strigilis do on average more cleaning actions per minute than biscrapers, thus they compensate partly for the fewer number of strokes. But nevertheless the uniscrapers do fewer strokes than the biscrapers (both with an ancestral strigilis). Females clean their antennae on average more often than males. It is interpreted that the behaviour of uniscraping and a derived morphology of the antenna cleaner result in greater efficiency than the status which is ancestral for Apoidea (biscraping and a plesiomorphic antenna cleaner).     

Musculature of the male genitalia in Rivellia (Diptera: Platystomatidae)

The musculature of male genitalia was studied hitherto only in two species of Tephritidae, one species of Platystomatidae, one species of Pallopteridae, and three species of Ulidiidae of the superfamily Tephritoidea. The split of the hypandrium from one structure into three (the hypandrium and two lateral sclerites) is traced. The hypandrial origin of the lateral sclerites of the hypandrial complex is shown based on the localization of muscle attachment sites. The subepandrial origin of the inner lobes of the surstyli is also confirmed.     

Leg tegumental glands and male rubbing behavior at leks inPolistes Dominulus (Hymenoptera: Vespidae)

In summer, males of Polistes dominulus defend adjacent territories at landmarks. Patrol flights are interrupted by stops at selected perches and conspicuous dragging movements of the last sternites and all the legs over the substrate, especially at a high intruder density. Leg rubbing is regularly associated with prolonged grooming sessions. Single secretory cells, resembling the sternal glands formerly described in this species (class 3 ducted glands), were found in all the legs (in the femur, the tibia, and especially the tarsus) and in both sexes. Leg tegumental gland cells, like sternal glands, are probably a source of pheromones. They have not previously been described in polistine wasps.     

Occurrence and morphometry of the hypopharyngeal glands inScaptotrigona postica Lat. (Hymenoptera, Apidae, Meliponinae)

As compared toApis mellifera where only workers have hypopharyngeal glands, inScaptotrigona postica, these glands occur in workers, queens and males. They are composed of two long axial ducts with many unicellular secretory alveoli interconnected by secretory canaliculi. The axial ducts are longer in males than in workers, but the alveolar areas of queens and males are generally smaller. In workers the alveoli have their greatest size in the nurses or middle-aged individuals while in queens and males they are larger in newly emerged individuals. The results indicate that the glands in workers may produce food for the brood as inA. mellifera, since they are well developed in the nurse workers. However, the function of the glands in queens and males remains to be clarified since these individuals have no part in brood care.     

Diversity in bee (Hymenoptera: Apoidea) and social wasp (Hymenoptera: Vespidae, Polistinae) community in "campos rupestres", Bahia, Brazil

Hymenoptera such as bees and social wasps are regular floral visitors in "campos rupestres" vegetation. A community of bees and social wasps was studied during floral visitation in an area of "campos rupestres", at Chapada Diamantina, BA, Brazil, from September 2001 to April 2002. The community was described in relation to diversity, evenness, and dominance rank, considering the individuals abundance (H' = 2.14/ J' = 0.55) and biomass (H' = 2.34/ J' = 0.60). Thirty nine bee (588 individuals/ 15.742 g) and 11 social wasp species (52 individuals/ 2.156 g) were collected, being the first report of social wasps for the Brazilian "campos rupestres". The main species regarding number of individuals were Trigona spinipes (Fabricius), Apis mellifera L., Frieseomelitta francoi (Moure), and Bombus brevivillus Franklin. About 48% of the species were represented by a single individual. There was an inversion in the dominance rank when the species biomass was considered. B. brevivillus, A. mellifera, T spinipes, and other species represented by 15 individuals or less, such as the social wasps Synoeca cyanea (Olivier), Polistes canadensis (L.) and Myschocyttarus drewseni (Saussure), and the bees Eufriesea nigrohirta (Friese), Xylocopa grisescens Lepeletier and Megachile (Pseudocentron) sp.l were the predominant species. The use of biomass in diversity analysis permitted to detect differences in the relative contribution of species in hierarchy dominance. The comparison between bee faunas from different areas indicates a large similarity of the sampled fauna in Palmeiras (Bahia State) with neighboring ecosystems, although with low values of similarity.     

Foraging patterns in three sympatric forest ant species, Prenolepis imparis, Paratrechina melanderi and Aphaenogaster rudis (Hymenoptera: Formicidae)

Abstract.

• 1 Ecological segregation was studied in Aphaenogaster rudis Emery, Paratrechina melanderi (Wheeler) and Prenolepis imparis (Say), the three commonest surface-foraging ants in a Maryland (U.S.A.) hardwood forest community.
• 2 P.imparis retreats from the surface during mid-summer, but shows more cool-season activity than do the other two species. P.melanderi is most active during the warmest months of the years. A.rudis is the most generalized of the three species in its seasonal activity pattern.
• 3 P.imparis is a highly aggressive species which forms large mobilizations at rich food sources. P.melanderi is a timid species which forms small mobilizations. A.rudis is intermediate in both respects.
• 4 Based on the results of baiting trials, P.imparis shows an apparent preference for protein-fat food, P.melanderi prefers carbohydrate food, and A.rudis exhibits no significant preference for either fclod type.
• 5 As a result of its extended seasonal activity, weak circadian periodicity, and lack of food specialization, A.rudis has the broadest annual niche of the three species studied.
• 6 The unusual seasonal activity pattern of P.imparis results in a relatively low degree of annual niche overlap between this species and either of the other two. Overlap between A.rudis and P.melanderi is substantially higher.
• 7 Experimental manipulations confirm that rates of resource removal by a behaviourally subordinate species (Amdis) are reduced in the presence of a dominant species (P.imparis).
• 8 Persistence of these three widespread, abundant, relatively generalized ant species in local sympatry may in part reflect their differential success in: (a) finding food, (b) defending food, and (c) handling food items of different sizes.