Negative body size-dependent resource allocation underlies conspicuous sexual ornaments in a territorial damselfly.

Sexual ornaments, signalling individual quality to choosy females or rival males, often show steeper body size scaling compared with non-sexually selected traits. Theory posits such steeper body size scaling is the result of differential resource allocation, reflecting trade-offs between different components of fitness. Yet, the process of resource allocation towards body size-dependent sexual ornaments has been rarely understood empirically. Using the Neotropical territorial damselfly Megaloprepus caerulatus, whose males and females carry wax-based, sex-specific white wing bands and white wing tips respectively, we investigated nutrition sensitivity and body size scaling of both traits by manipulating larval food availability and directly quantified both the fat allocated to wing ornaments and the fat reserve from which allocations are made. Both colour traits exhibited sensitivity to food availability during larval development and steeper body size scaling compared with control traits. Although the absolute amount of fat invested in developing the colour ornaments increased with body size, the proportion of total fat allocated to the ornaments decreased with body size, making exaggerated ornaments less affordable for smaller individuals. Our data demonstrate that measuring the proportion of resource pool from which an individual's ornaments are derived (i.e. its affordability) is essential for understanding the maintenance of honesty of sexual signals.

Tests of search image and learning in the wild: Insights from sexual conflict in damselflies.

Search image formation, a proximal mechanism to maintain genetic polymorphisms by negative frequency-dependent selection, has rarely been tested under natural conditions. Females of many nonterritorial damselflies resemble either conspecific males or background vegetation. Mate-searching males are assumed to form search images of the majority female type, sexually harassing it at rates higher than expected from its frequency, thus selectively favoring the less common morph. We tested this and how morph coloration and behavior influenced male perception and intersexual encounters by following marked Ischnura elegans and noting their reactions to conspecifics. Contrary to search image formation and associative learning hypotheses, although males encountered the minority, male-like morph more often, sexual harassment and clutch size were similar for both morphs. Prior mating attempts or copula with morphs did not affect a male's subsequent reaction to them; males rarely attempted matings with immature females or males. Females mated early in the day, reducing the opportunity for males to learn their identity beforehand. Once encountered, the male-like morph was more readily noticed by males than the alternative morph, which once noticed was more likely to receive mating attempts. Flexible behavior gave morphs considerable control over their apparency to males, influencing intersexual encounters. Results suggested a more subtle proximal mechanism than male learning maintains these color polymorphisms and call for inferences of learning to be validated by behavior of wild receivers and their signalers.

Mechanical and tactile incompatibilities cause reproductive isolation between two young damselfly species.

External male reproductive structures have received considerable attention as a cause of reproductive isolation (RI), because the morphology of these structures often evolves rapidly between populations. This rapid evolution presents the potential for mechanical incompatibilities with heterospecific female structures during mating and could thus prevent interbreeding between nascent species. Although such mechanical incompatibilities have received little empirical support as a common cause of RI, the potential for mismatch of reproductive structures to cause RI due to incompatible species-specific tactile cues has not been tested. We tested the importance of mechanical and tactile incompatibilities in RI between Enallagma anna and E. carunculatum, two damselfly species that diverged within the past ∼250,000 years and currently hybridize in a sympatric region. We quantified 19 prezygotic and postzygotic RI barriers using both naturally occurring and laboratory-reared damselflies. We found incomplete mechanical isolation between the two pure species and between hybrid males and pure species females. Interestingly, in mating pairs for which mechanical isolation was incomplete, females showed greater resistance and refusal to mate with hybrid or heterospecific males compared to conspecific males. This observation suggests that tactile incompatibilities involving male reproductive structures can influence female mating decisions and form a strong barrier to gene flow in early stages of speciation.

Predation by odonates depresses mosquito abundance in water-filled tree holes in Panama.

In the lowland moist forest of Barro Colorado Island (BCI), Panama, larvae of four common species of odonates, a mosquito, and a tadpole are the major predators in water-filled tree holes. Mosquito larvae are their most common prey. Holes colonized naturally by predators and prey had lower densities of mosquitoes if odonates were present than if they were absent. Using artificial tree holes placed in the field, we tested the effects of odonates on their mosquito prey while controlling for the quantity and species of predator, hole volume, and nutrient input. In large and small holes with low nutrient input, odonates depressed the number of mosquitoes present and the number that survived to pupation. Increasing nutrient input (and consequently, mosquito abundance) to abnormally high levels dampened the effect of predation when odonates were relatively small. However, the predators grew faster with higher nutrients, and large larvae in all three genera reduced the number of mosquitoes surviving to pupation, even though the abundance of mosquito larvae remained high. Size-selective predation by the odonates is a likely explanation for this result; large mosquito larvae were less abundant in the predator treatment than in the controls. Because species assemblages were similar between natural and artificial tree holes, our results suggest that odonates are keystone species in tree holes on BCI, where they are the most common large predators.

Population regulation of a tropical damselfly in the larval stage by food limitation, cannibalism, intraguild predation and habitat drying.

The relative importance of intraspecific, interspecific, and seasonal causes of larval mortality were investigated for aquatic larvae of the giant damselfly Megaloprepus coerulatus in Panama. These larvae live in water-filled holes in fallen and living trees, where they and three other common odonate species are the top predators. By mid wet season, M. coerulatus larvae were found in nearly half of all tree holes that harbored odonates. Although M. coerulatus were typically, but not always, eliminated from holes inhabited by larger hetero-specifics, M. coerulatus were more likely to encounter conspecifics than other odonate species. Hole with less than 11 of water rarely contained more than a single larva. In large holes where M. coerulatus was the only odonate species present, multiple larvae coexisted at a density of one larva per 1-21 of water. There the absence of 2-4 of the 5 larval size classes, despite a continuous input of eggs, suggested that cannibalism was a common cause of mortality. The size of the final instar, which determined adult body size, was correlated positively with tree hole volume for male, but not female, larvae. Experiments showed that when two larvae were placed together in 0.4-1 holes with abundant tadpole prey, the larger larva killed the smaller one. Often the larva that was killed was not eaten. Small larvae were more tolerant of each other than were pairs of medium or large larvae. Before killing occurred, the presence of larger larvae reduced the growth of smaller individuals, relative to controls. 'Obligate' killing was density-dependent. In 3.0-1 holes with ad libitum prey, conspecific killing occurred until the larval density stabilized at one larva per 1-1.5 I, similar to the density found in large holes under field conditions, For M. coerulatus, cannibalism functions to reduce the number of potential competitors for food in addition to providing nutrition. When interactions between paired larvae in small holes were experimentally prevented, competition for food reduced the growth of one or both larvae relative to controls. Holes that were watered during the dry season supported larval densities similar to those in the wet season. Thus, dry season mortality could not be attributed to a decrease in available prey. Rather, M. coerulatus larvae could not survive more than 1 month of complete drying. Because the dry season typically lasts more than 6 weeks, habitat drying is a secondary source of mortality, affecting second- or third-generation larvae that fail to emerge before tree holes dry out completely.

LIFETIME REPRODUCTIVE SUCCESS AND THE OPPORTUNITY FOR SELECTION IN A NONTERRITORIAL DAMSELFLY (ODONATA: COENAGRIONIDAE).

Major components of male and female lifetime reproductive success (LRS) were quantified for a damselfly that exhibits "scramble competition" for mates. The opportunity for selection on male reproduction was potentially 2.9 times that for females. Differential fertility/clutch and survivorship each accounted for about half of the total variation in female reproductive success. Variation in fertilization efficiency accounted for 7% of the total opportunity for selection on males. Although differences in survivorship and mating efficiency each contributed to about a third of the total opportunity for selection on male reproduction, both components appeared to be influenced by random factors. Survivorship was age-independent, and the mating distributions among males with equal mating opportunities were indistinguishable from those expected if matings were random with respect to male phenotype. Because the proportion of the standarized variance (I) in LRS that was attributed to sexual selection depended on the way the selective episodes were defined, the sample of individuals included in the partitioning analysis, and the degree of sexual selection on mated males that could be detected, my results caution against drawing conclusions about the dynamics of sexual selection on populations based on a superficial comparison of I values.