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.     

Temperature effects on the development and multiple generations in the alkali bee, Nomia melanderi cockerell

The range for development of post-diapausing prepupae of the alkali bee lies between 17 and 35°. Peak developmental temperature is approximately 29°. Although temperatures above 33° are outside of the favorable range of development and induce an obligatory aestivation, they are effective in causing a very rapid diapause breakage. Prepupae held at 35° for 110 days underwent no development, yet when removed to 29° they pupated in half of the time required by the controls at that temperature. Voltinism in the alkali bee is temperature dependent, and apparently the physiological system that triggers diapause is not activated unless the developing larvae or prepupae are subjected to temperatures below 27°. Males continue to emerge throughout the season, but a small percentage of the male population appears before any females to effect the phenomenon of proterandry.

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Zusammenfassung Der förderliche Bereich für die Postdiapause-Entwicklung der Präpuppen der Alkali-Biene liegt zwischen 17 und 35°. Die Optimaltemperatur beträgt annäherungsweise 29°. Obwohl Temperaturen über 33° außerhalb des förderlichen Entwicklungsbereichs liegen und eine obligatorische Ästivation induzieren, verursachen sie doch einen sehr raschen Bruch der Diapause. Präpuppen, die 110 Tage bei 35° gehalten wurden, entwickelten sich nicht weiter; nach 29° zurückgebracht, verpuppten sie sich jedoch in der Hälfte der Zeit, welche von den Kontrolltieren bei dieser Temperatur benötigt wirde. Mehrfach-Generationen der Alkali-Biene sind temperaturbedingt, und offensichtlich wird das physiologische System, welches Diapause auslöst, nur dann in Gang gesetzt, wenn die Larven oder Präpuppen Temperaturen unter 27° ausgesetzt werden. Männchen schlüpfen während der gesamten Brutsaison, doch erscheint ein kleiner Teil der männlichen Population vor den Weibchen und bewirkt so das Phänomen der Proterandrie.

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Technical Paper No. 1847, Oregon Agricultural Experiment Station, supported by National Science Foundation Grant No. G 15609.     

Phylogeny of the World Anthophorini (Hymenoptera: Apoidea: Apidae)

Abstract.  The phylogenetic relationships of members of the long-tongued bee tribe Anthophorini were investigated. Twenty-six ingroup taxa, representing all known genera and the most common subgenera of the Anthophorini, were included in a cladistic analysis of the tribe based on 51 morphological characters. The analysis yielded the generic relationships: (( Habrophorula , Elaphropoda ) ( Habropoda ( Deltoptila ( Pachymelus ( Amegilla , Anthophora ))))). The monophyly of the Anthophorini and each of its genera was confirmed. A second tribe, Habropodini, as postulated by some authors, is paraphyletic. On the basis of the results of the cladistic analysis and the biogeographical data of Anthophorini, an evolutionary scenario is postulated in which all genera of Anthophorini probably evolved in the Late Cretaceous, except Amegilla and Anthophora , which originated in the Oligocene. The northern part of south-east Asia (India to south-east China) is regarded as the most probable place of origin and radiation of the Anthophorini. The New World was probably colonized three times independently by Habropoda (upper Cretaceous to Tertiary), by an ancestral lineage of Deltoptila (upper Cretaceous to Tertiary) and by Anthophora (Tertiary to Quaternary).     

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.     

Phylogenetic relationships and host-plant evolution within the basal clade of Halictidae (Hymenoptera, Apoidea)

Bees are among the most important pollinators of angiosperm plants. Many bee species show narrow host‐plant preferences, reflected both in behavioral and morphological adaptations to particular attributes of host‐plant pollen or floral morphology. Whether bee host‐plant associations reflect co‐cladogenesis of bees and their host plants or host‐switches to unrelated host plants is not clear. Rophitinae is a basal subfamily of Halictidae in which most species show narrow host‐plant preferences (oligolecty). We reconstructed the phylogenetic relationships among the rophitine genera using a combination of adult morphology (24 characters) and DNA sequence data (EF‐1α, LW rhodopsin, wingless; 2700 bp total). The data set was analyzed by parsimony, maximum likelihood and Bayesian methods. All methods yielded highly congruent results. Using the phylogeny, we investigated the pattern of host‐plant association as well as the historical biogeography of Rophitinae. Our biogeographical analysis suggests a number of dispersal/vicariance events: (1) a basal split between North America and South America (most likely a dispersal from South America to North America), and (2) at least two subsequent interchanges between North America and Eurasia (presumably via the northern hemisphere land bridges). Our analysis of host‐plant associations indicates that Rophitinae specialized on a closely related group of angiosperm orders in the Euasterid I clade (mainly Gentianales, Lamiales and Solanales). However, there is little evidence of cocladogenesis between bees and plants and strong evidence of host switches to unrelated host plants. Based on our phylogenetic results we describe two new tribes of Rophitinae: Conanthalictini new tribe (including the genus Conanthalictus) and Xeralictini new tribe (including Xeralictus and Protodufourea). © The Willi Hennig Society 2007.     

Comparative analyses of lemon-smelling secretions from heads of Andrena F. (Hymenoptera,Apoidea) bees

• 1.1. Volatile cephalic secretions from male and female Andrena F. bees were analysed by capillary gas chromatography and mass spectrometry.
• 2.2. Eight species. A. clarkella (K.), A. fucata Smith, A. helvola (L.), A. praecox (Scop.), A. labiata F., A. fuscipes (K.), A. denticulata (K.) and A. nigriceps (K.), were studied, in A. nigriceps (K.) only females were available.
• 3.3. These species were found to contain some acyclic monoterpenes, especially geranial and neral, and/or straight-chain ketones with 8–14 carbon atoms.
• 4.4. The volatile compounds emanate from the mandibular glands.
• 5.5. Six of the species studied show large similarities in the composition of male and female cephalic secretions, whereas in one species. A. helvola (L.), characteristic differences were found between the sexes.
• 6.6. The possible role of the cephalic secretions in relation to territorial flight, population congregation, and sexual attraction is discussed.

     

New records for the bee fauna of Iran (Hymenoptera: Apoidea)

In a survery of the bee fauna of Horand forests in East Azerbaijan province of Iran during 2008 and 2009, 46 species belonging to the families Anthophoridae, Halictidae and Megachilidae were identified. Eight species are new to the fauna of Iran: Evylaeus albipes (Fabricius, 1781), E. corvinus (Morawitz, 1878), E. israelensis (Ebmer, 1974), Seladonia desertorum (Morawitz, 1876), Heriades crenulatus Nylander, 1856, Hoplitis anthocopoides (Schenck, 1853), H. manicata (Morice, 1901) and H. rufohirta (Latreille, 1811).     

Revision of the family Sphecidae (Hymenoptera: Apoidea) in South Korea

Korean species of the family Sphecidae are reviewed, and seventeen species in nine genera belonging to three subfamilies are confirmed. Among them, Ammophila campestris and Sceliphron caementarium are new to South Korea. Korean occurrence of Sceliphron madraspatanum is confirmed with an actual Korean specimen. This paper provides the first determination keys to higher taxa and species occurring in South Korea. Taxonomic information of each species including original publication of valid name and justifiable references for Korean occurrence is provided. Digital images and line drawings for selective characteristics are also available.