Do reproductive activities compromise immunological competence as measured by phenoloxidase activity? Field and experimental manipulation in females of two damselfly species

Recent studies of female insects indicate that reproductive activities, such as mating and oviposition, can impair immune ability. Using the two tropical damselfly species Argia anceps Garrison and Hetaerina americana (Fabricius), egg production and phenoloxidase (PO) activity, a key enzyme in insect immunity, are measured in mating, ovipositing and perching females in December and March. Perching females of both species have fewer eggs compared with mating and ovipositing females, which suggests that perching females are not engaged in reproduction. There is seasonal variation in egg number for the three categories in H. americana but not in A. anceps, which can be interpreted in terms of adaptive changes in egg production depending on female–male interactions in the former species but not in the latter species. There is no difference in PO activity among mating, ovipositing or perching females within either species, although measurements in December and March indicate distinct seasonal changes. Juvenile Hormone (JH) is known to reduce the effectiveness of the immune system by favouring the use of resources for reproduction. A possible role for JH is examined in H. americana, using the JH analogue methoprene to manipulate hormone activity, revealing that PO activity is reduced in methoprene‐treated H. americana females. Thus, although the results of the present study are indicative of possible hormone‐driven changes in PO, there is not necessarily a down‐regulation of immune function (as determined by PO activity) during mating or oviposition. The results complement some recent studies countering the idea that reproductive activities reduce the immune ability in insects.     

Molecular evidence for selection on female color polymorphism in the damselfly Ischnura graellsii

The significance of female color polymorphism in Odonata remains controversial despite many field studies. The importance of random factors (founder effects, genetic drift and migration) versus selective forces for the maintenance of this polymorphism is still discussed. In this study, we specifically test whether the female color polymorphism of Ischnura graellsii (Odonata, Coenagrionidae) is under selection in the wild. We compared the degree of genetic differentiation based on RAPD markers (assumed to be neutral) with the degree of differentiation based on color alleles. Weir and Cockerham's theta values showed a significant degree of population differentiation for both sets of loci (RAPD and color alleles) but the estimated degree of population differentiation (theta) was significantly greater for the set of RAPD loci. This result shows that some sort of selection contributes to the maintenance of similar color morph frequencies across the studied populations. Our results combined with those of previous field studies suggest that at least in some I. graellsii populations, density-dependent mechanisms might help to prevent the loss of this polymorphism but cannot explain the similarity in morph frequencies among populations.     

Odonates as indicators of the ecological integrity of the river corridor: Development and application of the Odonate River Index (ORI) in northern Italy

The assessment of the ecological conditions of rivers is crucial for their appropriate management and restoration. Bioindicators commonly used to evaluate the river status (i.e. diatoms, aquatic macrophytes, benthic macroinvertebrates and fish) detect alterations of water quality, but are not particularly sensitive to hydromorphological degradation, which is another relevant pressure in river systems. Furthermore, those bioindicators are usually applied only to flowing channels. We developed a new multimetric index, the Odonate River Index (ORI), to assess the conditions of the whole corridor in alluvial rivers. The ORI is a development of an evaluation system proposed in Austria, and based on the Odonate Habitat Index (OHI). Odonates were chosen as bioindicators for the ecological integrity of the river corridor, since this taxon provides information on the conditions of their aquatic breeding sites, as well as on the surrounding terrestrial areas, due to its amphibiotic life cycle. We used a case study of 18 reaches from six Italian Alpine rivers, characterized by different morphological conditions and level of human impact. Within each study reach, we selected four sites, both lotic and lentic sites. Dragonfly surveys consisted in field observation of adults, and collection of larvae and exuviae. To define the best sampling strategy, we compared the results of the ORI metrics obtained varying the input data by combining different sampling methods: the best compromise between effort and exhaustiveness was obtained coupling the observation of adults with the collection of exuviae. We found the ORI to be a robust and reliable tool to assess the status of the river corridor in a wide range of environmental conditions and river morphology, being particularly suitable to detect hydromorphological degradation and alterations of the structure of aquatic and riparian vegetation. We identified two limiting factors for the applicability of this index: low water temperatures of the main channel (i.e. mean annual value below 10 °C) and river reaches with no or scarce aquatic and riparian vegetation. In addition to the assessment of river conditions, the ORI could also be applied for monitoring the effects of river restoration actions.     

Stylus of the odonate endophytic ovipositor: a mechanosensory organ controlling egg positioning

Using light and scanning electron microscopy, a sensory field consisting of 15-20 campaniform sensillae is described on the base of the stylus of the endophytic ovipositor of Odonata. It is hypothesised that two symmetric styli equipped with this number of sensillae can function as a mechanosensory organ responsible for control of precise egg positioning in plant stems during oviposition. In laboratory experiments with females of damselflies Lestes sponsa and Lestes barbarus (Lestidae), it was demonstrated that the distance between laid eggs is not dependent on the presence of styli. Removal of styli from both sides did not influence a shift of oviposition to one side. Females with one removed stylus shifted the clutch line in the opposite direction toward the removed stylus. Additionally, removal of styli influenced positions of single eggs in egg sets, and disturbed the capacity for complex oviposition. Thus, both morphological and experimental data support the hypothesis that styli participate in the control of egg line and egg patterning in the clutch.     

Sexual comparisons in immune ability, survival and parasite intensity in two damselfly species

Recent evolutionary studies have suggested that females have a more robust immune system than males. Using two damselfly species (Hetaerina americana and Argia tezpi), we tested if females produced higher immune responses (as phenoloxidase and hydrolytic enzymes), had a higher survival (using a nylon implant inserted in the abdomen and measuring survival after 24h) and fewer parasites (gregarines and water mites) than males. We also tested whether immune differences should emerge in different body areas (thorax vs. abdomen) within each sex with the prediction that only females will differ with the abdomen having a higher immune response than their thorax since the former area, for ecological and physiological reasons, may be a target zone for increased immune investment. Animals were adults of approximately the same age. In both species, females were more immunocompetent than males, but only in H. americana females were immune responses greater in the abdomen than in the thorax. However, there were no differences in survival and parasite intensity or the probability of being parasitised between the sexes in either of the two species. Thus, this study lends partial support to the principle that females are better at defending than males despite the null difference in parasitism and survival.     

Survival rates in a natural population of the damselfly Ceriagrion tenellum: effects of sex and female phenotype

1. Ceriagrion tenellum females show genetic colour polymorphism. Androchrome (erythrogastrum) females are brightly (male‐like) coloured while gynochrome females (typica and melanogastrum) show cryptic colouration. 2. Several hypotheses have been proposed to explain the existence of more than one female morph in damselfly populations. The reproductive isolation and intraspecific mimicry hypotheses predict greater survival of gynochrome females, while the density dependent hypothesis predicts no differential survival between morphs. 3. Mature males had greater recapture probability than females while the survival probability was similar for both sexes. Survival and recapture rates were similar for androchrome and gynochrome females. 4. Gynochrome females showed greater mortality or migration rate than androchrome females during the pre‐reproductive period. This result is not predicted by the above hypotheses or by the null hypothesis that colour polymorphism is only maintained by random factors: founder effects, genetic drift, and migration.     

Fitness and body size in mature odonates

The relationship between body size and fitness components in odonates was examined using a meta‐analysis of 33 published studies. There was a positive and significant overall effect of body size on mating rate and lifetime mating success among males. There was also a weaker but still significant positive effect of body size on survivorship of males. The relationship between body size, mating rate, longevity, and lifetime mating success differed significantly between males of territorial and nonterritorial species. The effect of body size was significant for all fitness components in territorial species but significant only for longevity and lifetime mating success in nonterritorial species. Effect sizes appeared to be strongest on longevity in both sexes, and on male mating rate in territorial species. Other effect sizes, even when significant, were small. Despite a much smaller data set, female fitness also increased significantly with body size. Both clutch size and longevity showed a significant positive relationship with body size. These results suggest that there is a general fitness benefit to large size in odonates. Nevertheless, significant heterogeneity is apparent in this effect, which can be attributed to sex, mating system, and fitness component. Finally, these analyses point to inadequacies in the current data that need further study before the potentially rich patterns in size effects on fitness can be explored more thoroughly.     

Immune response to nylon filaments in two damselfly species that differ in their resistance to ectoparasitic mites

1. Insects commonly resist parasites using melanotic encapsulation. Many studies measuring immune response use the amount of melanin deposited on an artificial object that has been inserted into the animal as a proxy of the amount of resistance that the host is capable of mounting to natural parasites. 2. The relevance of this methodology to immune response in natural insect populations needs further study. Here, we examined two temperate damselfly species to elucidate the relationships among damselfly size, natural resistance to mites, and the immune response mounted by the same damselflies against nylon filaments. 3. The damselfly species that had high rates of melanotic encapsulation of mites in nature deposited more melanin on the nylon inserts than the species with low rates of natural resistance. 4. In females of this species, those that had resisted mites naturally melanised the nylon filament more aggressively than those that did not resist mites. 5. Our results show some support for the use of nylon filaments to assess natural patterns of immune response in these damselflies, but also suggest that caution should be used in interpreting the responses.     

Lentic odonates have larger and more northern ranges than lotic species

Aim We analysed latitudinal range, centres of distribution and northern range boundaries of dragonflies and damselflies occurring in Europe and North America with respect to larval habitat (standing water = lentic and running water = lotic). As lentic water bodies are thought to be less predictable in space and time than lotic habitats, species adapted to standing waters depend on effective dispersal ability for long‐term survival. If species occurring in lentic habitats have a higher propensity for dispersal, then larger range sizes in lentic than in lotic species, as well as an increase in the proportion of lentic species with latitude, would be expected. Location Europe, North America. Methods Distributional and habitat data were collected from published sources for all odonates of Europe and North America. Species were assigned to lentic and lotic habitats according to the habitat of the larvae. From distribution maps we estimated the latitudinal range, centre of distribution and northern range boundary of each species. Differences in these distribution variables between lentic and lotic species were evaluated using anova . We related the proportion of lentic species by latitudinal interval in Europe, and by political unit (state, province) in North America, to area, altitudinal range, longitude (only for North America) and latitude by means of generalized linear models. Results Lentic damselflies and dragonflies had larger latitudinal spans, and more northern distribution centres and range boundaries, than lotic species. The proportion of lentic species increased with latitude. These findings were consistent between continents. Main conclusions Our results support previous findings that distribution patterns of freshwater species depend on habitat preference. Evolution of dispersal propensity according to habitat characteristics is the most likely explanation. However, at present, alternative explanations, such as an increase in lentic habitats with latitude, cannot be ruled out.