Field estimates of reproductive success in a model insect: behavioural surrogates are poor predictors of fitness

Understanding, and therefore measuring, factors that determine fitness is a central problem in evolutionary biology. We studied a natural population of Coenagrion puella (Odonata: Zygoptera) over two entire breeding seasons, with over a thousand individuals uniquely marked and genotyped, and all mating events at the rendezvous site recorded. Using a parentage analysis, fitness of individuals in the first generation was quantified as the numbers of offspring that survived to maturity. Although mating behaviour can be predicted by environmental and demographical variables, the numbers of mature offspring produced (fitness) cannot, and crucially, are poorly correlated with behavioural observations of mating. While fitness of both sexes was positively related to mating behaviour and to female's ectoparasite burden, these behavioural observations explained little more variance in offspring production than environmental and demographical variables. Thus, we demonstrate that behavioural measures of reproductive success are not necessarily reliable estimates of fitness in natural populations.     

Habitat requirements and local persistence of three damselfly species (odonata: coenagrionidae)

Habitat requirements and population persistence were investigated in three damselfly species, all coastal plain pond specialists: Enallagma recurvatum, E. laterale, and E. pictum. Because of geographic restriction, two are of special concern to conservation, E. recurvatum and E. laterale. We surveyed more than 70 ponds on Cape Cod, Massachusetts, and collected adult presence-absence data during the summers of 1999 and 2000. We achieved a detection rate approaching 100% for each species by visiting each pond up to three times. We looked for relationships between the presence of each damselfly species and presence of specific aquatic vegetation, the presence of the other Enallagma species, and the number of ponds within various distances of the 72 surveyed ponds. Using stepwise logistic regression, we found the following significant associations: E. recurvatum with the rush Juncus militaris; E. laterale with water lilies (Nuphar variegatum and Brasenia schreberi) the damselfly E. pictum, and the number of ponds within 2 km; and E. pictum with the water lily Nymphaea odorata, the damselfly E. laterale, and the number of ponds within 1.5 km and 2.5 km. Presence-absence data were used to calculate turnover and local extinction rates for each species between the two years. E. recurvatum's turnover and local extinction rates (33.3% and 41% respectively) were much higher than either E. laterale (9.8%, 11.5%) or E. pictum (7.7%, 10%). These results suggest that E. recurvatum occurs in a metapopulation, and that patch colonization rates might be important to local population persistence.     

Developmental costs of rapid growth in a damselfly

Abstract.  1. Developmental costs of rapid growth in terms of increased fluctuating asymmetry are expected to contribute to the widespread occurrence of growth rates below the physiological maximum, but have rarely been demonstrated. Here, these costs are studied for the first time in an invertebrate, the damselfly Lestes viridis , using a rearing experiment where early- and late-hatched larvae of both sexes were reared at decreasing or permanent water levels.
2. Late-hatched animals were more asymmetrical than early-hatched animals except for males in the drying treatment. Also, females were more asymmetrical than males except in early-hatched animals in the drying treatment.
3. The data presented suggest that in females but not in males treatment groups with higher growth rates have more asymmetrical wings. However, at the individual level no relationship between growth rate and asymmetry was present.
4. Possible reasons why the suggested trade-off between growth and developmental instability was not present at the individual level, and at the group level only in females, are discussed.     

Reproductive behavior of females and its relation to the mating success of two male forms inMnais pruinosa (Zygoptera: Calopterygidae)

The behavior of females of the damselflyMnais pruinosa was observed in the breeding season. Males consisted of both territorial (esakii) and non-territorial (strigata) types in the study area. Females sometimes arrived at a stream for mating and oviposition. The staying time of females in the stream was 1–3 hours, while that of males was 5–8 hours. The proportion of the males that copulated with the females did not differ between the two male forms present in the stream. In the Calopterygidae, of which the females usually perform multiple copulation, oviposition without subsequent recopulation is considered to be advantageous for a mated male in order to avoid the risk of sperm displacement. From such a viewpoint, the time spent for oviposition was measured for females that arrived at the stream after copulation with different male forms. The ratio was 69.1: 11.5–30.9 betweenesakii andstrigata males. Assuming that the oviposition time is proportional to the number of fertilized eggs laid, this ratio would represent the relative reproductive success of the two male forms. In fact, the relative abundance of the two male forms was also biased in favor ofesakii males (61.7:38.3) in this population. The mechanism of coexistence of the two male forms is discussed in relation to their reproductive success.     

Intraguild predation, cannibalism, and microhabitat use in Calopteryx virgo and Somatochlora metallica larvae: a laboratory experiment

Intraguild predation (IGP) and cannibalism among co-occurring lotic odonate species was studied in Central Finland. A laboratory experiment was performed to assess the microhabitat use and cannibalism between intermediate and late instars of Calopteryx virgo larvae and predation by larger Somatochlora metallica larvae on the intermediate C. virgo instars. The experiment was run in small running-water aquaria where the larvae were able to divide their mutual habitat vertically by clinging onto artificial perches or crawling on the bottom. Life span of the small C. virgo larvae and attack rate on them were compared between the cannibalism and IGP treatments. The effect of predation on the activity, habitat use and spatial distribution of the small C. virgo larvae was examined. The IGP rate was 36%. The prey larvae spent the most of their time on the perches, whereas the S. metallica preferred the substrate. The large C.␣virgo larvae did not cannibalise smaller conspecifics. The presence of a predator (S. metallica) had no effect on the habitat use or activity of the prey (C. virgo) larvae. Habitat use differed more between those species than between conspecifics of different size classes of C. virgo. The spatial distribution between S.␣metallica and C. virgo showed a completely random pattern, whereas the two size classes of C. virgo aggregated in the vegetation. Absence of cannibalism and behavioural observations indicate that C. virgo may have a low tendency for intraspecific aggressions.     

A comprehensive test for negative frequency-dependent selection

Understanding the mechanisms that maintain genetic diversity within a population remains a primary challenge for evolutionary biology. Of the processes capable of maintaining variation, negative frequency-dependent selection (NFDS), under which rare phenotypes (or alleles) enjoy a high fitness advantage, is suggested to be the most powerful. However, few experimental studies have confirmed that this process operates in nature. Although a lot of suggestive evidence has separately been provided in various polymorphic systems, these are not enough to prove the existence of NFDS in each system. Here we present a general review of NFDS and point out some problems with previous works to develop reasonable alternative research strategies for testing NFDS. In the second half of this paper, we focused on NFDS in the common bluetail damselfly, Ischnura senegalensis, that shows female-limited genetic polymorphism. We show (1) the proximate causal mechanisms of the frequency-dependent process, (2) frequency-dependent inter-morph interaction, (3) rare morph advantage and (4) morph frequency oscillations in a natural population. These results provide unequivocal empirical support for NFDS in a natural system.     

Effects of eugregarine parasites on adult longevity in the polymorphic damselfly Mnais costalis Selys

Abstract.  1. The relationship between the abundance of midgut parasites (eugregarine trophozooites) and the survival of hosts (field-collected damselflies, Mnais costalis ) was investigated under laboratory conditions.
2. Males of M. costalis are morphologically and behaviourally polymorphic, typically existing as clear-winged non-territorial sneaks and orange-winged territorial fighters (the latter are larger in size). The survival cost of eugregarine infection for the two morphs was compared.
3. Orange-winged males showed shorter longevity compared to clear-winged males when they were fed at levels lower than the natural feeding rate, or when they were deprived of food. In contrast, morph longevity did not differ when they were fed until satiation every day.
4. Within each morph, the survival of damselflies was negatively correlated with the parasite abundance except when damselflies were fed until satiation.
5. The results suggest that the abundance of eugregarine parasites exerts a substantial cost, which is associated with the maintenance of alternative mating strategies, together with the higher developmental cost and energy expenditure of the fighter morph.     

The expression of a sexually selected trait correlates with different immune defense components and survival in males of the American rubyspot

Recent studies have suggested that courtship trait expression indicates immune strength. However, most studies have measured only one immune parameter, have not assessed individual differences in immune ability according to time and have not controlled for ecological differences among individuals after an immune challenge. In this work, we tested this hypothesis and controlled for these factors using males of the American rubyspot damselfly which bear a wing red spot whose size is evolutionarily maintained via male-male territorial competition. Our general hypothesis was that territorial, large-spotted males, had a better immune ability compared to nonterritorial, small-spotted males. We expected that the following variables were greater in territorial males compared to nonterritorial males: spot size, phenoloxidase (PO) and hydrolytic enzymatic (HE) activity in males challenged and nonchallenged with a nylon implant, PO and HE activity rate; PO activity after a Serratia marcescens challenge, and survival after a nylon challenge controlling for activity and feeding differences. We found that territorial males showed larger spot areas, greater PO and HE activity (independently of whether they were challenged or not), a higher rate of PO and HE activity (but only expressed at 8h), greater PO production after the bacterial challenge, and a higher survival after the challenge. These results corroborate that males with more pronounced sexual traits have a superior immune function.     

Adult Odonata conservatism as an indicator of freshwater wetland condition

There is a growing need to identify effective and efficient biological indicators for wetland assessment, and adult damselflies and dragonflies (Insecta: Odonata) possess several attributes that make them attractive for this application. We introduce a general indicator of freshwater wetland condition based on objectively estimated adult Odonata species conservatism, or sensitivity to human disturbances. We used an extensive opportunistic survey dataset from Rhode Island (USA) to empirically assign a coefficient of conservatism (CoC) to each of 135 Odonata species, based on their exclusivity to categories of degradation among 510 wetlands; the mean CoC of species observed in the adult stage was applied as an index of wetland integrity. An independent sample of 51 wetlands was also drawn from the opportunistic survey to test the performance of the index relative to human disturbance, as measured by multimetric rapid assessment and surrounding impervious surface area. The index was well predicted by both disturbance measures and showed no evidence of dependence on sampling effort, wetland size, or geomorphic class. Our findings suggest that conservatism of adult Odonata averaged across species may provide a robust indicator of freshwater wetland condition. And because adult Odonata are generally easy to identify, especially relative to larval Odonata, the index could be particularly useful for wetland assessment. Our straightforward empirical approach to CoC estimation could be applied to other existing spatially referenced Odonata datasets or to other species assemblages.     

Behavioral responses to fish kairomones and autotomy in a damselfly

The threat-sensitivity hypothesis predicts that prey species assess and adjust their behavior in accordance with the magnitude of the threat posed by a predator. A largely overlooked characteristic of a prey that will affect its sensitivity to predators is its history of autotomy. We studied threat-sensitive behavior to fish kairomones in larvae of Ischnura elegans damselflies, which had undergone autotomy, from a fishpond and from a fishless pond. In agreement with their higher perceived risk, larvae from the fishpond showed fewer rigid abdomen bends, foraged less and walked more slowly than larvae from the fishless pond. In line with their higher vulnerability to predators, larvae without lamellae spent less time foraging than larvae with lamellae. There was a decrease in swimming activity in the presence of fish kairomones except for larvae with lamellae from the fishless pond. This may reflect differences in vulnerability of larvae without lamellae between pond types. Such context-dependent responses in activity to kairomones should be kept in mind when evaluating the ability of a prey to recognize kairomones.