Social environment affects the life history tactic of a phoretic mite

Phoretic animals use their hosts for travelling to habitat patches suitable for reproduction. Some species, such as the mite Poecilochirus carabi, are phoretic as juveniles and cannot leave their habitat once they reach adulthood. Previous work has shown that mites exercise choice over the habitat in which they will mature and reproduce based on abiotic parameters, but it is hitherto unknown whether their social environment influences this choice. By manipulating the composition of their conspecific company we show that P. carabi perform the adult moult in the presence of prospective mating partners only. Furthermore, juvenile male mites do not moult in the presence of an adult competitor. Recently‐moulted males are severely disadvantaged in fighting, so such delayed moulting may allow juveniles to increase their chances of surviving and reproducing. Our results clearly demonstrate a strong influence of the social environment on a phoretic’s habitat choice and life history.     

Role of two phoretic mites in transmission of bluestain fungus, Ceratocystis minor

ABSTRACT.

• 1 Tarsonemus mites phoretic on southern pine beetles were shown to be hypervectors of bluestain fungus, Ceratocystis minor (Hedgcock 1906).
• 2 Beetles with phoretic mites had significantly more C.minor than did mite-free beetles.
• 3 Over 90% of tarsonemid mites collected from beetle-infested inner bark and 59% of mites phoretic on emerging beetles carried C.minor spores.
• 4 This association is an interesting example of a three-way mutualism among insects, mites and fungi.

     

Behind every great ant, there is a great gut

Ants are quite possibly the most successful insects on Earth, with an estimated 10 000 species worldwide, making up at least a third of the global insect biomass, and comprising several times the biomass of all land vertebrates combined. Ant species have diverse trophic habits, including herbivory, hunting/gathering, scavenging and predation and are distributed in diverse habitats, performing a variety of important ecosystem functions. Often they exert these functions while engaging in symbiotic associations with other insects, plants or microbes; however, remarkably little work has focused on the potential contribution of the ants’ gut symbionts. This issue of Molecular Ecology contains a study by Anderson et al. (2012) , who take a comparative approach to explore the link between trophic levels and ant microbiomes, specifically, to address three main questions: (i) Do closely related herbivorous ants share similar bacterial communities? (ii) Do species of predatory ants share similar bacterial communities? (iii) Do distantly related herbivorous ants tend to share similar bacterial communities? By doing so, the authors demonstrate that ants with similar trophic habits appear to have relatively conserved gut microbiomes, suggesting symbiont functions that directly relate to dietary preference of the ant host. These findings suggest an ecological role of gut symbionts in ants, for example, in metabolism and/or protection, and the comparative approach taken supports a model of co‐evolution between ant species and specific core symbiont microbiomes. This study, thereby, highlights the omnipresence and importance of gut symbioses—also in the Hymenoptera—and suggests that these hitherto overlooked microbes likely have contributed to the ecological success of the ants.     

Biology,demography and community interactions of Tarsonemus (Acarina: Tarsonemidae) mites phoretic on Dendroctonus frontalis (Coleoptera: Scolytidae)

1 Dendroctonus frontalis, the southern pine beetle, is associated with a diverse community of fungi and mites that are phoretic on the adult beetles. Tarsonemus ips, T. kranzti and T. fusarii (Acarina: Tarsonemidae) may interact within this community in ways that link the population dynamics of D. frontalis, the mites and three dominant species of fungi. We explored species associations by comparing the dietary suitability of different fungi for Tarsonemus spp. 2 All three mite species fed and reproduced at high rates when feeding on the bluestain fungus, Ophiostoma minus, which is an antagonist of D. frontalis larvae. 3 Mites also had positive population growth rates when feeding upon Ceratocystiopsis ranaculosus, one of the mycangial fungi, but could barely reproduce when feeding upon Entomocorticium sp. A, the mycangial fungus that is most suitable for D. frontalis. 4 During the time from colonization of a tree by D. frontalis adults until departure from the tree of their progeny (≈ 40 d at 30 °C), mite populations feeding upon O. minus can increase by factors of up to 209 (T. fusarii), 173 (T. ips) or 384 (T. krantzi). These high growth rates are allowed by rapid development (age of first reproduction = 8–9 d), high fecundity (≈ 1 egg/d) and high longevity (> 28 d). 5 Precocious mating increases the chance that females are mated prior to colonizing a new tree and arrhenotokous parthenogenesis permits reproduction by unmated females. 6 Tarsonemus mites may introduce negative feedback into D. frontalis population dynamics by generating indirect interactions between D. frontalis and O. minus.     

The natural history of a phoretic sphaerocerid Diptera fauna

ABSTRACT.

• 1 Three species of phoretic Sphaeroceridae kleptoparasitize the dung caches of scarab beetles in north Florida. Ceroptera sivinskii Marshall intercepts beetles under dung, then accompanies them into their burrows. Borborillus frigipennis (Spuler) rides on ball rolling and tunnelling scarabs. It perches on beetles both underground and in flight. B.singularis (Spuler) is similar in habits but is principally nocturnal.
• 2 Kleptoparasitism is probably a means of escaping environmental decay and competition in‘parent’faeces. Nonscarab competitors and predators are less abundant in beetle stored dung, and Borborillus spp. are among the few spaerocerids that flourish when large, rapidly developing calypterate flies are abundant.
• 3 Since oviposition in B.frigipennis occurs only underground, phoresy is an inexpensive and safe means to maintain contact with hosts. Males ride in order to find mates.

     

Sex ratio, sex-specific chick mortality and sexual size dimorphism in birds

It has been suggested that sexual size dimorphism (SSD) may influence sex ratios at different life stages. Higher energy requirements during growth associated with larger body size could lead to a greater mortality of the larger sex and ultimately to an overproduction of the smaller sex. To explore the associations between SSD and hatching and fledging sex ratio we performed a species-level analysis and a phylogenetically controlled analysis, based on 83 bird species. Overall, there was a significant inverse relationship between the degree of SSD and the proportion of males at hatching and fledging. Sex-specific mortality related to SSD showed a weak but persistent negative tendency, suggesting a mortality bias towards the larger sex. These results suggest that changes in relation to SSD may take place mainly at the conception stage, but could be adjusted during growth. However, conclusions should be treated cautiously as these relationships weaken when additional variables are considered.     

Sex allocation theory reveals a hidden cost of neonicotinoid exposure in a parasitoid wasp

Sex allocation theory has proved to be one the most successful theories in evolutionary ecology. However, its role in more applied aspects of ecology has been limited. Here we show how sex allocation theory helps uncover an otherwise hidden cost of neonicotinoid exposure in the parasitoid wasp Nasonia vitripennis. Female N. vitripennis allocate the sex of their offspring in line with Local Mate Competition (LMC) theory. Neonicotinoids are an economically important class of insecticides, but their deployment remains controversial, with evidence linking them to the decline of beneficial species. We demonstrate for the first time to our knowledge, that neonicotinoids disrupt the crucial reproductive behaviour of facultative sex allocation at sub-lethal, field-relevant doses in N. vitripennis. The quantitative predictions we can make from LMC theory show that females exposed to neonicotinoids are less able to allocate sex optimally and that this failure imposes a significant fitness cost. Our work highlights that understanding the ecological consequences of neonicotinoid deployment requires not just measures of mortality or even fecundity reduction among non-target species, but also measures that capture broader fitness costs, in this case offspring sex allocation. Our work also highlights new avenues for exploring how females obtain information when allocating sex under LMC.     

No seasonal sex-ratio shift despite sex-specific fitness returns of hatching date in a lizard with genotypic sex determination

Sex allocation theory predicts that mothers should adjust their sex-specific reproductive investment in relation to the predicted fitness returns from sons versus daughters. Sex allocation theory has proved to be successful in some invertebrate taxa but data on vertebrates often fail to show the predicted shift in sex ratio or sex-specific resource investment. This is likely to be partly explained by simplistic assumptions of vertebrate life-history and mechanistic constraints, but also because the fundamental assumption of sex-specific fitness return on investment is rarely supported by empirical data. In short-lived species, the time of hatching or parturition can have a strong impact on the age and size at maturity. Thus, if selection favors adult sexual-size dimorphism, females can maximize their fitness by adjusting offspring sex over the reproductive season. We show that in mallee dragons, Ctenophorus fordi, date of hatching is positively related to female reproductive output but has little, if any, effect on male reproductive success, suggesting selection for a seasonal shift in offspring sex ratio. We used a combination of field and laboratory data collected over two years to test if female dragons adjust their sex allocation over the season to ensure an adaptive match between time of hatching and offspring sex. Contrary to our predictions, we found no effect of laying date on sex ratio, nor did we find any evidence for within-female between-clutch sex-ratio adjustment. Furthermore, there was no differential resource investment into male and female offspring within or between clutches and sex ratios did not correlate with female condition or any partner traits. Consequently, despite evidence for selection for a seasonal sex-ratio shift, female mallee dragons do not seem to exercise any control over sex determination. The results are discussed in relation to potential constraints on sex-ratio adjustment, alternative selection pressures, and the evolution of temperature-dependent sex determination.     

Survival and colonization of nematodes in a composting process

Nematodes are omnipresent in composts and are active in virtually all stages of the composting process. Major shifts in species composition, life strategies, and feeding behavior occur during the composting process. Due to the heat peak, nematodes can be virtually absent, but several taxa appear immediately when the temperatures drop. These comprise both taxa present before the heat peak and new taxa. However, it is not known how nematodes populate the compost. In this study, we aimed to assess the survival and colonization capacity of nematodes in compost. Our results showed that composting processes inaccessible to insects or not in contact with soil did not significantly influence nematode succession during composting. However, differences between treatments were found for some specific taxa (i.e., for Acrostichus sp., Neodiplogasteridae sp., Nygolaimoides sp., and Rhabditidae sp. 1), illustrating the importance of insects for the dispersal of nematodes to compost. Experiments in the lab with the blue bottle fly as a possible carrier demonstrated actual transport of nematodes isolated from compost by the fly (i.e., Halicephalobus cfr. gingivalis, Diploscapter coronatus, Diplogasteritus sp., Acrostichus sp., and Mesorhabditis sp.). Juveniles and dauer stages of Aphelenchoides sp., Panagrolaimus sp., and rhabditids survived an experimentally induced temperature peak, while members of Tylenchidae did not. In conclusion, our results indicate that the rapidly changing nematode community in compost is the result of both differential survival and colonization capacities.     

Fitness costs of phoretic nematodes in the burying beetle,Nicrophorus vespilloides

Nicrophorusvespilloides is a social beetle that rears its offspring on decomposing carrion. Wild beetles are frequently associated with two types of macrobial symbionts, mites, and nematodes. Although these organisms are believed to be phoretic commensals that harmlessly use beetles as a means of transfer between carcasses, the role of these symbionts on N. vespilloides fitness is poorly understood. Here, we show that nematodes have significant negative effects on beetle fitness across a range of worm densities and also quantify the density‐dependent transmission of worms between mating individuals and from parents to offspring. Using field‐caught beetles, we provide the first report of a new nematode symbiont in N. vespilloides, most closely related to Rhabditoides regina, and show that worm densities are highly variable across individuals isolated from nature but do not differ between males and females. Next, by inoculating mating females with increasing densities of nematodes, we show that worm infections significantly reduce brood size, larval survival, and larval mass, and also eliminate the trade‐off between brood size and larval mass. Finally, we show that nematodes are efficiently transmitted between mating individuals and from mothers to larvae, directly and indirectly via the carcass, and that worms persist through pupation. These results show that the phoretic nematode R. regina can be highly parasitic to burying beetles but can nevertheless persist because of efficient mechanisms of intersexual and intergenerational transmission. Phoretic species are exceptionally common and may cause significant harm to their hosts, even though they rely on these larger species for transmission to new resources. However, this harm may be inevitable and unavoidable if transmission of phoretic symbionts requires nematode proliferation. It will be important to determine the generality of our results for other phoretic associates of animals. It will equally be important to assess the fitness effects of phoretic species under changing resource conditions and in the field where diverse interspecific interactions may exacerbate or reduce the negative effects of phoresy.     

Three new species of the genus Caesarodispus (Acari: Microdispidae) associated with ants (Hymenoptera: Formicidae), with a key to species

Three new species of the genus Caesarodispus (Acari: Heterostigmatina: Microdispidae) phoretic on ants are described from Iran: C. khaustovi Rahiminejad & Hajiqanbar sp. nov. , C. pheidolei Rahiminejad & Hajiqanbar sp. nov. and C. nodijensis Rahiminejad & Hajiqanbar sp. nov. All species were associated with alate ants of the subfamily Myrmicinae (Hymenoptera: Formicidae) from northern Iran. A key to all species of Caesarodispus is provided.     

Egg load is a cue for offspring sex ratio adjustment in a fig‐pollinating wasp with male‐eggs‐first sex allocation

Fig‐pollinating wasps (Agaonidae) only reproduce within fig tree inflorescences (figs). Agaonid offspring sex ratios are usually female‐biased and often concur with local mate competition theory (LMC). LMC predicts less female‐bias when several foundresses reproduce in a fig due to reduced relatedness among intra‐sexually competing male offspring. Clutch size, the offspring produced by each foundress, is a strong predictor of agaonid sex ratios and correlates negatively with foundress number. However, clutch size variation can result from several processes including egg load (eggs within a foundress), competition among foundresses and oviposition site limitation, each of which can be used as a sex allocation cue. We introduced into individual Ficus racemosa figs single Ceratosolen fusciceps foundresses and allowed each to oviposit from zero to five hours thus variably reducing their eggs‐loads and then introduced each wasp individually into a second fig. Offspring sex ratio (proportion males) in second figs correlated negatively with clutch size, with males produced even in very small clutches. Ceratosolen fusciceps lay mainly male eggs first and then female eggs. Our results demonstrate that foundresses do not generally lay or attempt to lay a ‘fixed’ number of males, but do ‘reset to zero’ their sex allocation strategy on entering a second fig. With decreasing clutch size, gall failure increased, probably due to reduced pollen. We conclude that C. fusciceps foundresses can use their own egg loads as a cue to facultatively adjust their offspring sex ratios and that foundresses may also produce more ‘insurance’ males when they can predict increasing rates of offspring mortality.     

The effect of season and sex on trophic levels of marine fishes

The diet of 76 species from Greek waters (north Aegean Sea) was examined for the determination of trophic level by species, season and sex. Trophic level estimates were derived from TrophLab software based on collected stomach content data. Such estimates are important for the implementation of marine management indicators, such as the marine trophic index. The mean ± s . e . difference between minimum and maximum trophic levels among seasons within species was 0·45 ± 0·04 and for 28·8% of the species this difference was larger than the corresponding s . e . of the trophic level of the pairs of seasons examined. Depending on which seasonal trophic level ( i.e. autumn, winter, spring or summer) was used for assigning a species into a functional trophic group 33·9 and 8·5% of the species changed one and more than one group, respectively. Finally, the mean ± s . e . absolute difference of male and female trophic level was 0·18 ± 0·02, and only for 6·6% of the species this difference was larger than the s . e . of trophic level for each sex separately. Depending on the trophic level used for assigning the species into a functional group ( i.e. male v. female), only 6·6 and 3·2% of the species changed one and more than one groups, respectively. It is therefore clear that all measures used in this study indicated that the effect of season on trophic level estimates was higher, compared to that of sex, the latter being of minor importance to feeding studies. Hence, future effort on stomach content analysis should focus on the examination of seasonal, small to moderate sample sizes.     

Selectable components of sex allocation in colonies of the honeybee (Apis mellifera L.)

Colonies of social insects that undergo fission as a componentof reproduction produce large excesses of males. Hypothesesto explain this phenomenon have assumed that the workers thatconstitute the entourage for the new queen (or queens) representinvestment in female reproductives. Selection for optimal colonysex allocation then leads to an increase in production of malesthat balances the investment in females based on their relativereproductive values. We show that the construction of comb dedicatedto the production of males (drone comb) versus workers (workercomb) is a component of sex investment under the control ofcolony workers. Relative comb construction was highly correlatedwith the relative investment in male and worker brood. Coloniesthat invested relatively more in their total numbers of malesinvested less in the dry weight of individual workers. Coloniesthat had more adult workers produced a greater number of malesand workers, but colony size did not affect the proportionalinvestment in drone comb or brood. Genetic variability was foundfor the number of adult workers in colonies, the amount of dronecomb produced, the amount of worker comb produced, and the dryweight of adult workers, suggesting that sex allocation is aselectable trait in honeybees.     

Split sex ratios in the social Hymenoptera: a meta-analysis

The study of sex allocation in social Hymenoptera (ants, bees,and wasps) provides an excellent opportunity for testing kin-selectiontheory and studying conflict resolution. A queen–workerconflict over sex allocation is expected because workers aremore related to sisters than to brothers, whereas queens areequally related to daughters and sons. If workers fully controlsex allocation, split sex ratio theory predicts that colonieswith relatively high or low relatedness asymmetry (the relatednessof workers to females divided by the relatedness of workersto males) should specialize in females or males, respectively.We performed a meta-analysis to assess the magnitude of adaptivesex allocation biasing by workers and degree of support forsplit sex ratio theory in the social Hymenoptera. Overall, variationin relatedness asymmetry (due to mate number or queen replacement)and variation in queen number (which also affects relatednessasymmetry in some conditions) explained 20.9% and 5% of thevariance in sex allocation among colonies, respectively. Theseresults show that workers often bias colony sex allocation intheir favor as predicted by split sex ratio theory, even iftheir control is incomplete and a large part of the variationamong colonies has other causes. The explanatory power of splitsex ratio theory was close to that of local mate competitionand local resource competition in the few species of socialHymenoptera where these factors apply. Hence, three of the mostsuccessful theories explaining quantitative variation in sexallocation are based on kin selection.     

Genetic components to caste allocation in a multiple-queen ant species

Reproductive division of labor and the coexistence of distinct castes are hallmarks of insect societies. In social insect species with multiple queens per colony, the fitness of nestmate queens directly depends on the process of caste allocation (i.e., the relative investment in queen, sterile worker and male production). The aim of this study is to investigate the genetic components to the process of caste allocation in a multiple-queen ant species. We conducted controlled crosses in the Argentine ant Linepithema humile and established single-queen colonies to identify maternal and paternal family effects on the relative production of new queens, workers, and males. There were significant effects of parental genetic backgrounds on various aspects of caste allocation: the paternal lineage affected the proportion of queens and workers produced whereas the proportions of queens and males, and females and males were influenced by the interaction between parental lineages. In addition to revealing nonadditive genetic effects on female caste determination in a multiple-queen ant species, this study reveals strong genetic compatibility effects between parental genomes on caste allocation components.     

SEX RATIO VARIATION IN A PARASITIC WASP I. REACTION NORMS

To understand genetic and phenotypic constraints on the sex ratio in a parasitic wasp that attacks fly pupae, I carried out a laboratory study of sex ratio variability in five strains of Muscidifurax raptor (Hymenoptera: Pteromalidae). I manipulated the environment through combinations of temperature and day length, and the numbers of females that attack a group of hosts. The change of phenotype in each strain over the range of environmental conditions describes the norm of each reaction for that strain, and measures how a strain responds to environmental variation to create phenotypic variability. Sex ratio in parasitic wasps is a complex trait that has several components—the numbers of eggs laid by an ovipositing wasp and the fraction of eggs that are fertilized (female). Further, sex ratio may be influenced by a female's reaction to other females exploiting the same hosts (superparasitism). I found no strain-environment interactions in either sex ratio or fecundity when I varied environmental conditions. Although strains differed in sex ratio and fecundity, all strains produced a more female-biased sex ratio and had higher fecundity when temperature and day length increased. Sex ratio and fecundity were phenotypically correlated, and strains with greater fecundity also produced a more female-biased sex ratio. All strains facultatively shifted sex ratio toward a higher fraction of males with increasing female density, despite apparent differences in superparasitism among strains. Males and females survived equally during development, so that mortality differences among strains and across environments could not account for sex ratio variability. This study indicates that sex ratio variability among strains is constrained by the correlation between sex ratio and fecundity, and that strains display similar facultative shifts in sex ratio as female density increases because sex ratio shifts are insensitive to differing levels of superparasitism.     

SEX RATIO AND AGE STRUCTURE OF GAMASID MITES FROM SMALL MAMMALS IN WESTERN YUNNAN,CHINA*

Abstract A total of 70 species of gamasid mites (7485 individuals) were collected from the body surface of 38 species of small mammals (3023 individuals) during an investigation of 13 counties of western Yunnan, China, from 1990 to 1996. Most species of gamasid mites have more females (80% 100%) than males (P <0.01). Exceptions are Haemogamasus gongshanensis where males outnumber females (P < 0.01) and Haemolaelaps glasgowi where there are similar numbers of males and females (P > 0.05). No larva was found in most of the mite species. The proportion of individuals at the immature stages (mainly protonymphs and deutonymphs) in most mite species is small (0–12.84%) compared to the adults (P < 0.01). Exceptions are Ornithonyssus bacoti and Haemolaelaps glasgowi where 62.84% and 49.48% respectively are nymphs. The biased sex ratios may reflect the probable different bloodsucking preferences and parthenogenesis of the gamasid mites. The absence of larvae and a low proportion of nymph stages in most mite species probably reflect that the larvae or nymphs of most mite species are non‐feeding or non‐blood sucking and are in their hosts' nests. Another possibility is that some species directly produce protonymphs or even deutonymphs without eggs and larvae which makes it difficult to find the larval stage.     

The effect of interspecific mating on sex ratios in the twospotted spider mite and the Banks grass mite (Acarina: Tetranychidae)

Mixed populations of the twospotted spider mite (TSM),Tetranychus urticae (Koch), and the Banks grass mite (BGM),Oligonychus pratensis (Banks), occur on corn and sorghum plants in late summer in the Great Plains. Interspecific matings between these arrhenotokous species occur readily in the laboratory but yield no female offspring. The effect of interspecific mating on female: male sex ratios was measured by examining the F₁ progeny of females that mated with both heterospecific and conspecific males in no-choice situations. TSM females that mated first with BGM males and then with TSM males produced a smaller percentage of female offspring than TSM females that mated only with TSM males (43.1±5.8 and 78.9±2.8% females, respectively). Similarly, BGM females mated with heterospecific males and then with conspecific males produced fewer female offspring than females mated only with BGM males (55.7±5.2 and 77.5±2.5%, respectively). Lower female: male sex ratios were produced also by BGM females that mated with TSM males after first mating with conspecifics (62.4±3.4%). In mixed populations containing males of both species, females also produced lower female: male sex ratios, but these ratios were not as low as expected based on mating propensities and progeny sex ratios observed in no-choice tests. These data suggest that interspecific mating may substantially reduce female fitness in both mite species by reducing the output of female offspring, but in mixed populations this effect is mitigated by unidentified behavioral mechanisms.