Parasitic flies (Conopidae, Diptera) may be important stress factors for the ergonomics of their bumblebee hosts

Bumblebees harbour a wide range of parasitic organisms that attack all stages of their life cycles (reviews in Postner, 1951; Pouvreau, 1973, 1974; Alford, 1975; Kistner, 1982). Among them, conopid flies (Conopidae, Diptera) are particularly interesting because they attack foraging bumblebees which are handling flowers, or even on the wing (Frison, 1926; Cumber, 1949; Postner, 1951; Howell, 1967; Askew, 1971). A single egg is attached to (Frison, 1926; Plath, 1934; Cumber, 1949) or inserted into (DeMeijre, 1904; Howell, 1967) the host's abdomen, where the larva hatches and feeds on haemolymph and internal organs. Within 6–10 days the larva passes through three recognizable stages (Pouvreau, 1974) before the fly pupates in situ within the abdomen. The host bee dies shortly before the parasite pupates (Postner, 1951; Smith, 1966) and the parasite overwinters in its puparium; the adult fly then emerges in early summer (Frison, 1926; Townsend, 1935; Cumber, 1949; Postner, 1951). Conopid flies as parasites of bumblebees are known from all major habitats where the hosts occur (e.g. Kröber, 1939; Smith, 1966). However, the effect of parasitism on distribution and abundance of bumblebees is not known. In this preliminary note we have estimated degrees of infestation and concomitant reduction of life span in affected workers. The results are compared with literature reports on infestation levels in Europe.     

The cephalic labial gland secretions of two socially parasitic bumblebees Bombus hyperboreus (Alpinobombus) and Bombus inexspectatus (Thoracobombus) question their inquiline strategy

Social parasitic Hymenopterans have evolved morphological, chemical, and behavioral adaptations to overcome the sophisticated recognition and defense systems of their social host to invade host nests and exploit their worker force. In bumblebees, social parasitism appeared in at least 3 subgenera independently: in the subgenus Psithyrus consisting entirely of parasitic species, in the subgenus Alpinobombus with Bombus hyperboreus, and in the subgenus Thoracobombus with B. inexspectatus. Cuckoo bumblebee males utilize species‐specific cephalic labial gland secretions for mating purposes that can impact their inquiline strategy. We performed cephalic labial gland secretions in B. hyperboreus, B. inexspectatus and their hosts. Males of both parasitic species exhibited high species specific levels of cephalic gland secretions, including different main compounds. Our results showed no chemical mimicry in the cephalic gland secretions between inquilines and their host and we did not identify the repellent compounds already known in other cuckoo bumblebees.     

Ontogeny of worker body size distribution in bumble bee (Bombus impatiens) colonies

1. Bumble bees exhibit worker size polymorphisms; highly related workers within a colony may vary up to 10‐fold in body mass. As size variation is an important life history feature in bumble bees, the distribution of body sizes within the colony and how it fluctuates over the colony cycle were analysed. 2. Ten commercially purchased colonies of Bombus impatiens (Cresson) were reared in ad libitum conditions. The size of all workers present and newly emerging workers (callows) was recorded each week. 3. The average size of bumble bee workers did not change with colony age, but variation in body size tended to decrease over time. The average size of callows did not change with population size, but did tend to decrease with colony age. In all measures, there was considerable variation among colonies. 4. Colonies of B. impatiens usually produced workers with normally distributed body sizes throughout the colony life cycle. Unlike most polymorphic ants, there was no increase in worker body size with colony age or colony size. This provides the first, quantitative data on the ontogeny of bumble bee worker size distribution. The potential adaptive significance of this size variation is discussed.     

Patterns of parasite infection in bumble bees (Bombus spp.) of Northern Virginia

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Larval development of Myzostoma cirriferum (Myzostomida)

The larval development of Myzostoma cirriferum is described by means of SEM, TEM, and cLSM. It is similar to that of other myzostomids and includes three stages: the protrochophore, the trochophore, and the metatrochophore. The protrochophore is a ball-shaped larva present in culture from 18-48 h after egg laying. It has no internal organs and its body is made of three cell types: covering cells and ciliated cells that are external and surrounded by a cuticle, and resting cells that fill the blastocoel. The trochophore is a pear-shaped larva that develops 20-72 h after egg laying; the body includes the same three cell types as the previous stage. The metatrochophore is a pear-shaped larva that develops between 40 h and 14 days and is characterized by the presence of two bundles of four chaetae. When fully developed, the metatrochophore has a digestive system (made of a pharynx, an esophagus, and a blind digestive pouch), two pairs of protonephridia, and a nervous system composed of a supraesophageal ganglion, circumesophageal connectives, and dorsal and ventral nerves. Metamorphosis generally occurs 7 days after egg laying. At that time, the metatrochophore loses its chaetae and becomes pleated ventrally. This ultrastructural analysis suggests that chaetae and the five ventral longitudinal nerve cords of M. cirriferum metatrochophores are homologous structures to those observed in some polychaete trochophores. Coupled with recent phylogenetic analyses, where the Myzostomida are placed outside the Annelida, homologies between myzostomid and polychaete larvae support the view that a trochophore appeared early during the spiralian evolution.     

Senescence of immune defence in Bombus workers

Abstract 1. Senescence in workers of social insects is a particularly intriguing life-history trait as the future fitness of workers relies primarily on age-dependent survival rate. The pattern of senescence of immune defence traits was investigated under laboratory conditions in workers of two bumble bees: Bombus terrestris and B. lucorum .
2. In both species, there was a significant decrease with age in the ability to encapsulate a foreign object (a global measure of the efficiency of immune systems). This pattern of senescence was observed in all colonies in B. terrestris (seven) and B. lucorum (eight) assayed, even though, for the latter, there was some heterogeneity among colonies.
3. In B. terrestris , two other measures of immune defence were taken: the relative percentage of fat body in the abdomen and the concentration of haemocytes (the immune defence cells). The quantity of fat body increased only slightly with age and there was no effect for the concentration of haemocytes. Interestingly, the concentration of haemocytes decreased strongly after an encapsulation response, regardless of the age of workers.
4. The importance of the senescence pattern observed for the immune defence traits is discussed in the context of the social biology of workers.     

Genetic differentiation of continental and island populations of Bombus terrestris (Hymenoptera: Apidae) in Europe

Ten microsatellite loci and a partial sequence of the COII mitochondrial gene were used to investigate genetic differentiation in B. terrestris, a bumble bee of interest for its high-value crop pollination. The analysis included eight populations from the European continent, five from Mediterranean islands (six subspecies altogether) and one from Tenerife (initially described as a colour form of B. terrestris but recently considered as a separate species, B. canariensis). Eight of the 10 microsatellite loci displayed high levels of polymorphism in most populations. In B. terrestris populations, the total number of alleles detected per polymorphic locus ranged from 3 to 16, with observed allelic diversity from 3.8 ± 0.5 to 6.5 ± 1.4 and average calculated heterozygosities from 0.41 ± 0.09 to 0.65 ± 0.07. B. canariensis showed a significantly lower average calculated heterozygosity (0.12 ± 0.08) and observed allelic diversity (1.5 ± 0.04) as compared to both continental and island populations of B. terrestris. No significant differentiation was found among populations of B. terrestris from the European continent. In contrast, island populations were all significantly and most of them strongly differentiated from continental populations. B. terrestris mitochondrial DNA is characterized by a low nucleotide diversity: 0.18%± 0.07%, 0.20%± 0.04% and 0.27%± 0.04% for the continental populations, the island populations and all populations together, respectively. The only haplotype found in the Tenerife population differs by a single nucleotide substitution from the most common continental haplotype of B. terrestris. This situation, identical to that of Tyrrhenian islands populations and quite different from that of B. lucorum (15 substitutions between terrestris and lucorum mtDNA) casts doubts on the species status of B. canariensis. The large genetic distance between the Tenerife and B. terrestris populations estimated from microsatellite data result, most probably, from a severe bottleneck in the Canary island population. Microsatellite and mitochondrial DNA data call for the protection of the island populations of B. terrestris against importation of bumble bees of foreign origin which are used as crop pollinators.     

Larval stop,delayed development and survival in overcrowded cultures of Drosophila melanogaster: Effect of urea and uric acid

Uric acid and urea added to non-crowded cultures of D. Melanogaster are able to reproduce the larval stop (cessation in development) detected in highly competitive situations. The quantitative analysis of media as well as of larvae and pupae reveals the presence of both compounds as natural waste products of nitrogen metabolism in Drosophila. The nature of their effect is discussed in terms of larval intoxication as a mechanism which may account for the effects usually observed in crowded cultures: development delay, lower survival and also larval stop (which can only be detected by interrupting the competitive process by an overfeeding technique).     

Impact of artificial photoperiodism on the colony development of the bumblebee Bombus terrestris

This study investigated the effect of the photoperiodic regimes 0 h light : 24 h dark (LD 0:24), LD 8:16, LD 16:8 and LD 24:0 at 28°C and 50% Relative Humidity (RH) on the colony development of hibernated (2.0, 2.5, 3.0, 3.5 and 4.0 months) bumblebee queens. The queens which had hibernated for 3.0 months and which were reared in a LD 8:16 photoperiod showed the highest rate of colony initiation (88.2%), colony foundation (67.6%) and progeny queen production (38.2%). The photoperiod LD 8:16 also produced the shortest period of colony initiation and colony foundation. The highest number of sexual males (171.2 ± 12.2) and queens (91.2 ± 9.9) were produced in the colonies when 3.5 and 3.0 month hibernated queens were kept in an LD 8:16 photoperiod. The results show that light regime and hibernation duration affect colony characteristics of Bombus terrestris.     

Effect of host plant on parasitism ofHelicoverpa armigera (Lep.: Noctuidae) byMicroplitis demolitor (Hym.: Braconidae)

Microplitis demolitor Wilkinson is an important larval parasitoid ofHelicoverpa armigera (Hübner) andH. punctigera (Wallengren) in Australia. The effect of host plant on parasitism of second instarH. armigera byM. demolitor was investigated in a glasshouse experiment. Parasitism was low (0%) on chickpea. Moderate to high levels of parasitism (22.4% to 75.4%) were recorded on sorghum, sunflower, maize, cotton and soybean. The results suggest that releases of larval parasitoids into chickpea are unlikely to enhance parasitismlevels during the first spring generation ofHelicoverpa spp.     

Queen-controlled sex ratios and worker reproduction in the bumble bee Bombus hypnorum,as revealed by microsatellites

Social insect colonies provide model systems for the examination of conflicts among parties with different genetic interests. As such, they have provided the best tests of inclusive fitness theory. However, much remains unknown about in which party's favour such conflicts are resolved, partly as a result of the only recent advent of the molecular tools needed to examine the outcome of these conflicts. Two key conflicts in social insect colonies are over control of the reproductive sex ratio and the production of male offspring. Most studies have examined only one of these conflicts but in reality they occur in tandem and may influence each other. Using microsatellite analyses, the outcome of conflict over sex ratios and male production was examined in the bumble bee, Bombus hypnorum. The genotypes were determined for mother queens, their mates and males for each of 10 colonies. In contrast to other reports of mating frequency in this species, all of the queens were singly mated. The population sex ratio was consistent with queen control, suggesting that queens are winning this conflict. In contrast, workers produced over 20% of all males in queen-right colonies, suggesting that they are more effective in competing over male-production. Combining these results with previous work, it is suggested that worker reproduction is a labile trait that may well impose only small costs on queen fitness.     

Larval development of the rhizocephalanSacculina polygenea (Crustacea: Cirripedia)

Larval development of the rhizocephalanSacculina polygenea (Crustacea: Cirripedia: Rhizocephala) parasitizing the coastal crabHemigrapsus sanguineus was studied in Vostok Bay, the Sea of Japan. At 22–23°C, the entire cycle of larval development takes 2.5 days and includes five naupliar stages and one cypris stage. Like other rhizocephalans, the larvae ofS. polygenea are lecithotrophic and only grow slightly in size in the course of development, and like all sacculinids, they have no flotation collar. The naupliar stages IV and V have a tubercle between the furcal rami; this tubercle is absent in the larvae of the genusPeltogasterella, but it has been described inS. carcini. The first seta of the antennule only disappears completely at the fourth stage, although it is markedly reduced at the third stage. No morphological differences, except differences in size, are found between male and female nauplii.     

Heritability of sperm length in the bumblebee Bombus terrestris

Sperm length is highly variable, both between and within species, but the evolutionary significance of this variation is poorly understood. Sexual selection on sperm length requires a significant additive genetic variance, but few studies have actually measured this. Here we present the first estimates of narrow sense heritability of sperm length in a social insect, the bumblebee Bombus terrestris. In spite of a balanced and straightforward rearing design of colonies, and the possibility to replicate measurements of sperm within single males nested within colonies, the analysis proved to be complex. Several appropriate statistical models were derived, each depending on different assumptions. The heritability estimates obtained ranged from h ² = 0.197 ± 0.091 to h ² = 0.429 ± 0.154. All our estimates were substantially lower than previous estimates of sperm length heritability in non-social insects and vertebrates.     

密林熊蜂雄性成蜂生殖系统发育动态

为了研究密林熊蜂Bombus patagiatus Nylander雄蜂出房后生殖系统发育变化过程, 本研究首次对密林熊蜂雄性成蜂（1-15日龄）生殖系统的精液量、 精子数、 性腺大小、 储精囊大小等进行了测定。结果显示： 雄性成蜂在1日龄时精液量最大, 达到5.95±0.39 μL; 镜检精子数在2日龄最多; 而精液中活动精子数出现波浪式的循环趋势, 9日龄达到最大值, 为25.93±1.06个; 精液中精子总数在9日龄达到最大值, 为160.67±17.11万个; 活动精子百分比以8、 9日龄最高, 分别达到86.38%±2.02%及86.45%±2.50%; 性腺长度在1日龄时最小, 为1 522.01±37.93 μm; 储精囊的中部宽度在7日龄最宽, 达86.38±2.96 μm, 但与9日龄相比无显著差异。由此可见, 8-9日龄雄性成蜂个体各指标较为稳定, 该日龄即为雄性成蜂性成熟且可以交配的时期。本研究结果为密林熊蜂人工繁育提供了理论依据。     

Duration of development of two species of carrion-breeding scuttle flies and forensic implications

A procedure for rearing carrion-breeding scuttle flies (Diptera: Phoridae) under naturally fluctuating temperatures is described. Data on the duration of development of the egg stage, egg plus feeding larval stages, postfeeding larvae and total development from egg to adult, at different temperature regimes (means of daily means) are tabulated for Megaselia giraudii (Egger) and M. rufipes (Meigen). The variation is greater than previously supposed and involves larvae derived from the same egg batch leaving the bait in successive waves. Despite this variation the tables allow the provision of percentage probabilities of estimations of the earliest oviposition dates for specimens collected from a forensic case.     

Larval development of the invasive charru mussel,Mytella strigata (Bivalvia: Mytilidae)

ABSTRACT

The South American charru mussel, Mytella strigata, was recently recorded in Singapore waters, possibly introduced into Southeast Asia through shipping. The mussels have rapidly spread across estuarine coastal mudflats. Adult mussels were collected, spawned in aquaria and larvae were successfully cultured to the juvenile stage in the laboratory. The larval morphology and development of M. strigata is described in this paper. D-shaped veligers were produced within 20 h of fertilization and were approximately 75 µm in shell length. These larvae were capable of settlement two weeks post fertilization. Given an adequate amount of food, they were able to grow up to 1 mm in shell length within 30 days. The larval shell of M. strigata possesses anterodorsal G2 hinge teeth as distinct wavy ledges, with a pitted resilial ridge clearly evident in the juvenile shell.     

A method for year-round rearing of cuckoo bumblebees (Hymenoptera: Apoidea: Bombus subgenus Psithyrus)

Summary. The study of the social interactions between host bumblebees and their inquiline species requires rearing them. Here we provide a simple method for rearing of cuckoo bumblebees (Bombus subgenus Psithyrus) in the laboratory. Two bumblebee cuckoo-host systems are used: Bombus (Psithyrus) vestalis hosted by B. (Bombus) terrestris and B. (Psithyrus) sylvestris hosted by B. (Pyrobombus) pratorum. First, the invasion of the Psithyrus female is performed in species-specific host colonies containing approximately 10 young workers. On average, a parasitized colony of B. terrestris produced 90 ± 9 young males and 21 ± 3 young females of B. vestalis, whereas a parasitized colony of B. pratorum produced 16 ± 2 young males and 5 ± 1 young females of B. sylvestris. One week after emergence, Psithyrus virgin females are placed in a flight cage exposed to natural light, with Psithyrus males of other colonies. After successful mating, the females are isolated and overwintered in a cold room at 4°C for 2–4 months. This method enables a year-round and mass breeding of Psithyrus species to facilitate studies of these rare species.     

Larval interspecific competition in two flea species parasitic on the same rodent host

Abstract.  1. The fleas Xenopsylla conformis and Xenopsylla ramesis exploit the same rodent host, Meriones crassus , and replace each other between two different habitats situated at the opposite sides of a steep precipitation gradient. It was hypothesised that the reason for this paratopic distribution is competition between larvae of the two species for food resources.
2. This hypothesis was tested by studying the performance of larvae of the two species in terms of their developmental success in mixed-species and single-species treatments under different air temperatures, relative humidities, substrate textures, and food abundance.
3. The number of individuals of X. conformis that survived until emergence depended significantly on the presence of competing species, being, in general, lower in mixed-species compared with single-species treatments. The decrease in developmental success of X. conformis in mixed-species treatments was found mainly during food shortage. In contrast, presence of the competitor did not affect the number of X. ramesis that survived until emergence. No effect of the presence of the competitor on duration of development or sex ratio was found in either species.
4. The results of this study, together with the results of our previous studies, provide an explanation for the paratopic distribution of X. conformis and X. ramesis that exploit the same host species.     

Larval development ofMacrophthalmus erato De Man, 1887 (Brachyura ocypodidae)

The zoeal and megalopal stage ofMacrophthalmus erato were obtained under laboratory conditions and are described and figured. Five zoeal and one megalopal stages were reared at 25; salinity and temperature of 29°C. Their features are compared with those of known species of the genus.