Physiological costs of compensatory growth in a damselfly

Little is known about physiological costs of rapid growth. We successfully generated compensatory growth to time stress and transient food stress in the damselfly Lestes viridis and studied the physiological correlates of the resulting reduced ability to cope with starvation. We found evidence for both mechanisms proposed to underlie the physiological trade-off: compensatory growth was associated with (1) a higher metabolic rate, as indicated by a higher oxygen consumption and a faster depletion of energy storage molecules (glycogen and triglycerides), and (2) a smaller investment in energy storage. The former may also explain why storage molecules after emergence were negatively affected by time stress and food stress, despite the successful compensation before emergence. These deferred physiological costs of rapid growth have the potential to couple larval stresses to adult fitness irrespective of age and size at emergence, and they may partly explain why many animals do not show their maximum achievable growth rate.     

A model of ecological and evolutionary consequences of color polymorphism

We summarize direct and indirect effects on fitness components of animal color pattern and present a synthesis of theories concerning the ecological and evolutionary dynamics of chromatic multiple niche polymorphisms. Previous endeavors have aimed primarily at identifying conditions that promote the evolution and maintenance of polymorphisms. We consider in a conceptual model also the reciprocal influence of color polymorphism on population processes and propose that polymorphism entails selective advantages that may promote the ecological success of polymorphic species. The model begins with an evolutionary branching event from mono- to polymorphic condition that, under the influence of correlational selection, is predicted to promote the evolution of physical integration of coloration and other traits (cf. multi-trait coevolution and complex phenotypes). We propose that the coexistence within a population of alternative ecomorphs with coadapted gene complexes promotes utilization of diverse environmental resources, population stability and persistence, colonization success, and range expansions, and enhances the evolutionary potential and speciation. Conversely, we predict polymorphic populations to be less vulnerable to environmental change and at lower risk of range contractions and extinctions compared with monomorphic populations. We offer brief suggestions as to how these falsifiable predictions may be tested.     

Body temperature and body size affect flight performance in a damselfly

Flight performance is undoubtedly an important factor for behavioral success in flying insects. Though it is well-known that the flight performance is influenced by body temperature and body size, the relative importance of these factors is not well-understood. We performed laboratory experiments using the male-polymorphic damselfly Mnais costalis with larger territorial males and smaller non-territorial males in a population. We analyzed the effects of body temperature and body size, measured as the thoracic temperature and left hind-wing length, respectively, on two indices of flight performance: maximum lifting force and size-corrected lifting force. The latter is an index of acceleration that is related to aerial agility. The results showed that higher body temperature produced both larger maximum lifting force and larger size-corrected lifting force. In contrast, while larger size produced a larger maximum lifting force, it produced a lower size-corrected lifting force. The results of field measurements showed that territorial males had variable thoracic temperatures depending on the insolation in their territories. In contrast, non-territorial males had less variable and generally higher thoracic temperatures than territorial males as they are mostly found in sunny spots. Until now, the influence of body temperature on behavioral performance has remained unclear although considerable studies have suggested such influence. We showed, here, for the first time, combined effects of body size and body temperature on flight performance. We also showed that body temperature was influenced by the mating strategies of a damselfly. These findings provide new insights into the cost and benefits of territorial behavior in ectothermic animals.     

Interspecific aggression in Leucorrhinia dragonflies: a frequency-dependent discrimination threshold hypothesis

Summary Male Leucorrhinia dragonflies defend territories from conspecific and heterospecific intruders. Defense against heterospecifics is surprising, as mating and oviposition are the only activities that occur on the territories, and heterospecific males are not expected to pose a reproductive threat. L. frigida and L. intacta males respond aggressively with equal frequency and intensity against intrusions by conspecifics and heterospecifics. In contrast, L. proxima males respond more aggressively against conspecifics. The apparent lack of species discrimination shown by L. frigida and L. intacta males may result because territorial males that assess intruders (as do L. proxima) suffer a tactical disadvantage from hesitating when an intruder flies in. This assessment process may lead to reduced fighting success by the territorial male. Thus there is a tradeoff between assessment cost incurred when the intruder is a conspecific and benefit gained from avoiding conflict when the intruder is a heterospecific. Given this assessment cost, males of species that only rarely encounter conspecific intruders are more likely to evolve species discrimination than males of species that commonly encounter conspecific intruders.     

Niche and fitness differences relate the maintenance of diversity to ecosystem function

The frequently observed positive correlation between species diversity and community biomass is thought to depend on both the degree of resource partitioning and on competitive dominance between consumers, two properties that are also central to theories of species coexistence. To make an explicit link between theory on the causes and consequences of biodiversity, we define in a precise way two kinds of differences among species: niche differences, which promote coexistence, and relative fitness differences, which promote competitive exclusion. In a classic model of exploitative competition, promoting coexistence by increasing niche differences typically, although not universally, increases the "relative yield total", a measure of diversity's effect on the biomass of competitors. In addition, however, we show that promoting coexistence by decreasing relative fitness differences also increases the relative yield total. Thus, two fundamentally different mechanisms of species coexistence both strengthen the influence of diversity on biomass yield. The model and our analysis also yield insight on the interpretation of experimental diversity manipulations. Specifically, the frequently reported "complementarity effect" appears to give a largely skewed estimate of resource partitioning. Likewise, the "selection effect" does not seem to isolate biomass changes attributable to species composition rather than species richness, as is commonly presumed. We conclude that past inferences about the cause of observed diversity-function relationships may be unreliable, and that new empirical estimates of niche and relative fitness differences are necessary to uncover the ecological mechanisms responsible for diversity-function relationships.     

Female oviposition-site preference and egg hatching success in the damselfly Calopteryx splendens xanthostoma

By removing males and controlling the rate of water flow past oviposition patches in an experimental area, we showed that female Calopteryx splendens xanthostoma preferred to oviposit in fast flowing water rather than slow flowing water. A series of manipulations revealed the fitness benefits to females for this preference: eggs placed in fast-flowing water developed significantly faster and showed significantly lower mortality than eggs placed in slow-flowing water. A major factor determining this difference was the deposition and growth of encrusting algae which prevented the successful hatching of eggs in slow-flowing water.     

Impacts of environmental heterogeneity on alternative mating tactics in the threadtail damselfly

Environmental heterogeneity, including variation in the physical environment, may be key to understanding the evolution and maintenance of alternative mating tactics, but its influence is rarely examined. Males of the threadtail damselfly Protoneura amatoria reversibly use two alternative mating tactics (perching vs. hovering) and have previously been found to modulate their use of these tactics in response to variation in both light conditions and the density of ovipositing females. Here, I show that mating success payoffs of the two tactics are differentially influenced by these factors. The payoff of the perching tactic was greater than that of the hovering tactic under low light conditions and at low densities of ovipositing females. The payoff of the hovering tactic was greater under high light conditions and higher densities of ovipositing females. The differential success of the two mating tactics in response to light conditions is discussed in light of flight dynamics, vision, and predation.     

Frequency-dependent male mate harassment and intra-specific variation in its avoidance by females of the damselfly Ischnura elegans

We focused on male harassment on different female color morphs of the damselfly Ischnura elegans and on variation in morph-specific mating avoidance tactics by females. In I. elegans, one of the female morphs is colored like the conspecific male (andromorphs) while the other morphs are not (gynomorphs). Our first goal was to quantify morph-specific male mating attempts, hence male harassment, in populations with manipulated population parameters (densities, sex ratios, and proportion of andromorphs). Second, we examined the female's perspective by looking for potential differences in morph-specific mating avoidance tactics and success of those tactics in a natural population. Differences in population conditions did influence the number of male mating attempts per morph. The less frequent female morph was always subject to fewer mating attempts, which contradicts earlier hypotheses on mimicry, but supports those that assume that males learn to recognize female morphs. Gynomorphs occupy less open habitat and often fly away when a male approaches, while andromorphs use more open habitat, do not fly large distances and directly face approaching males. Female morphs did not differ in the proportion of successful mating-avoidance attempts. Our results suggest that the maintenance of the color polymorphism is most probably the result of interactive selective forces depending on variation in all population conditions, instead of solely density- or frequency-dependent selection within populations.     

Underwater oviposition in a damselfly (Odonata: Coenagrionidae) favors male vigilance,and multiple mating by females

Summary Female Enallagma hageni oviposit underwater where they are inaccessible to males. I demonstrate that males guard submerged females rather than perch sites, and are behaviorally distinct from lone males at the water. In contrast to lone males, which always attempt to copulate with females presented to them, guarding males exhibit a conditional latency to remating which corresponds closely to the time required by females to oviposit a complete clutch of eggs. By ovipositing underwater, females decrease the risk that their eggs become exposed. Risks associated with submerged oviposition favor both mate guarding, and multiple, within-clutch matings by females. Both guarding mates and lone males rescue females that float on the water surface as the result of improper resurfacing. Such behavior reduces the mortality risk to females from 0.06 to 0.02 per oviposition bout. By remating between bouts, females benefit from the additional vigilance of lone males, who rescue floating females 1.4 times as often as original mates. A second consequence of multiple mating is an increase in the selective advantage of vigilance by mates. Because receptive females become scarce by early afternoon, whereas male density remains high, a male has little (3%) chance of encountering a second receptive female that day. However, he incurs a 42% risk of losing fertilizations if he abandons a mate. For male E. hageni mate guarding functions in the context of both natural and sexual selection. It insures that a mate lives to lay a complete egg clutch in addition to protecting a male's sperm investment.     

Lifetime mating success in a natural population of the damselfly,Enallagma hageni (Walsh) (Odonata: Coenagrionidae)

Summary Variance in lifetime mating success was measured for individuals of a population of Enallagma hageni, a non-territorial damselfly in northern Michigan. E. hageni is an explosive breeder with scramble competition for mates. Highly skewed operational sex ratios resulted in intense male-male competition which took the form of interference with tandem pairs. 41% of the males failed to mate in their lifetime as opposed to only 3.6% mating failure in females. The effect on mating success of size, age, longevity, and time spent at the breeding site were investigated. Intermediate sized males obtained the most matings, and male lifetime mating success was highly correlated with longevity.     

Differential use of stored sperm during oviposition in the damselfly Calopteryx splendens xanthostoma (Charpentier)

Female Calopteryx splendens xanthostoma (Charpentier) (Odonata), have two discrete sperm storage organs, the bursa copulatrix and the paired spermathecae. We used the Random Amplified Polymorphic DNA(RAPD) profiling technique to determine from which sperm storage organ spermatozoa were used to fertilise eggs during behaviourally distinct oviposition bouts. During oviposition bouts following remating sperm from the bursa copulatrix are largely used to fertilise eggs, but when females avoid remating before an oviposition bout the sperm used to fertilise eggs are derived mainly from the paired spermathecae. Our observations suggest that female C. s. xanthostoma can avoid male paternity assurance mechanisms by a combination of behavioural and anatomical adaptation: the reason(s) they do so is unclear. Received: 28 November 1995/Accepted after revision: 5 August 1996     

Population maintenance of the short-lived shrub Sambucus in a deciduous forest

This study quantitatively clarifies the life history of a shrub, Sambucus racemosa ssp. sieboldiana, in an old-growth forest, the Ogawa Forest Reserve, Japan, by a demographic approach using a projection matrix model that incorporates interactions between demographic parameters and canopy height dynamics. S. racemosa is a common deciduous shrub in central Japan and is known to grow predominantly at forest edges or roadsides. This indicates that it is a highly light-demanding species, and occurrence in gaps in old-growth stands suggests its "fugitive," gap-dependent life history in old-growth forests. We found that one distinctive feature of this species was that its seedlings can survive well in shaded conditions by alternating stems every year like perennial herb species. Matrix model analyses demonstrated that S. racemosa can continuously regenerate under the present disturbance regime of this forest and is highly adaptable to the structural dynamics of the old-growth forest. The maturity of S. racemosa shrubs depends on their size, and nearly all (>90%) of the mature (reproducing) individuals were found in gaps or near gaps. But wide seed dispersal by birds and the ability to form both seed banks and seedling banks, the latter of which has been regarded as a common characteristic of shade-tolerant climax species, probably increase the species' chances to encounter canopy gaps. Dynamic-canopied matrix models showed that the greatest elasticity is with shaded seedling survival. The frequent stem alternation of shaded seedlings often makes the growth rate negative, but the survival rate of seedlings in low light awaiting new gap creation is remarkably high (0.93 yr(-1)). The lower survival rate of the larger individuals and smaller minimum size to start reproduction than other canopy or subcanopy shade-tolerant species indicate that S. racemosa has the potential to reproduce before the closure of the encountered gaps and to complete its life history rapidly.     

Factors leading to the evolution and maintenance of a male ornament in territorial species

Male ornamentation is assumed to have evolved primarily from selection by female mate choice. Yet this is only one possible reason for ornament evolution. Ornaments might also be useful in aggressive competition by improving opponent assessment between males, or they might function to enhance signal detection by making males more conspicuous in the environment. We tested both these ideas in territorial Anolis lizards in which female choice is either absent or secondary to males competing for territories that overlap female home ranges. Male tail crests only evolved in species in which territory neighbors were distant, consistent with the signal detection hypothesis. Once the tail crest had evolved, however, it seems to have become a signal in itself, with variation in the frequency and size of tail crests within species correlating with variables predicted by the aggressive competition hypothesis. Our study presents an apparent example of a male ornament in which the selection pressure leading to variation among species in ornament expression is different from the selection pressure acting on variation within species. The Anolis tail crest is therefore likely to be an exaptation. We caution that conclusions made on the evolution of male ornaments are dependent on the phylogenetic perspective adopted by a study. Studies restricted to single species are useful for identifying selection pressures in contemporary settings (i.e., the current utility of traits), but may lead to erroneous conclusions on the factors that initially lead to the origin of traits.     

Tests for a biased sex ratio when the data are clustered

In behavioral ecology the overall sex ratio in a population of birds is often tested to see if it differs from a 50/50 ratio. In recent publications the binomial test or the ² test are carried out although the sexes of chicks within the same nest may not be independent. The lack of independence occurs since female birds can adjust the sex ratio in an adaptive way as demonstrated in recent studies. In order to take dependence into consideration the Wilcoxon signed rank test based on the within-brood differences between the proportions of sons and daughters was performed in a study investigating great tit hatchling sex ratios. We compare this test with a test based on an optimally weighted estimator recently proposed for medical studies with clustered binary data. According to our simulation results, this novel test is more powerful than the Wilcoxon signed rank test and should be used for the analysis of avian sex ratios. The methods are illustrated with real data from the great reed warbler.     

Estimating photophysiological condition of endosymbiont-bearing Baculogypsina sphaerulata based on the holobiont color represented in CIE L*a*b* color space

Symbiont-bearing large benthic foraminifers (LBFs) are widely distributed around coral reefs. If the physiological responses of LBFs to environmental changes can be recognized at an individual level, LBFs could serve as highly accurate bioindicators. In this study, chlorophyll a, respiration, and photosynthesis of Baculogypsina sphaerulata individuals were measured, and whether these physiological traits could be estimated based on the color of the holobiont (foraminifera and the diatom symbionts) was examined. Chlorophyll a content was estimated using a* and b* values of holobiont color represented in CIE L*a*b* color space. Photosynthetic performance decreased significantly with increasing whiteness (L*). These results indicated chlorophyll content as well as photosynthetic performance of Baculogypsina could be directly estimated using the holobiont color. The increased whiteness in color and decreased photosynthetic performance were mainly observed under low-light environment, possibly indicating symbiotic algae were shrunk into the central part of the host shell due to prolonged exposure to adverse conditions.     

Sperm competition in the damselfly Enallagma hageni Walsh (Odonata: Coenagrionidae): benefits of multiple mating to males and females

Summary Sperm competition was investigated in the non-territorial damselfly, Enallagma hageni. Using irradiated (sterile) male techniques, the last male to mate was found to fertilize up to 95% of the eggs of the first clutch laid after mating. Dissection of females collected before, during, and after copula showed that a male removes a maximum of 87% of the sperm by volume of a previous mate. These data verify an earlier estimate of lifetime reproductive success in this species which was based on mating success, and suggest that indirect dissection methods offer minimum estimates of sperm precedence. Male E. hageni have ample opportunity to benefit from sperm precedence, since at least 10% of the receptive females encountered had already mated once that day, but still contained complete or partial clutches of eggs. Female E. hageni benefit directly from high sperm precedence because it allows them to exchange matings for guarding service by males during oviposition bouts under water.     

Elytra color as a signal of chemical defense in the Asian ladybird beetle Harmonia axyridis

Carotenoid- and melanin-based colors are valuable indicators of quality in many vertebrates, but their signaling role in invertebrates remains relatively unexplored. The Asian ladybird beetle Harmonia axyridis is an ideal organism for studies of this nature because males and females exhibit highly variable red and black colors on their elytra and are chemically defended with an alkaloid (harmonine). We used digital photography to quantify elytra coloration and absorbance spectrophotometry and gas chromatography mass spectrometry analyses to quantify pigment and alkaloid concentrations, respectively, in wild-caught male and female H. axyridis. We predicted that extensive or intense coloration would be an aposematic signal of high-alkaloid stores. We found that carotenoid pigments largely controlled variation in red elytra coloration. There was no relationship between alkaloid content and either elytra redness or carotenoid pigment concentration in either sex. However, we found a positive correlation between the extent of elytra coloration and alkaloid content. Animals with proportionally more red (or less black) on the elytra were more alkaloid-rich; this relationship was particularly strong in females. We also found that females with lighter black spots had greater amounts of harmonine than those with darker spots. These results suggest that elytra color patterns have the potential to reveal information about chemical defensiveness to mates or predators. Prior studies in this species show that nonmelanic forms are typically less active and yet more sexually attractive than melanic forms, and both results may be explained by the fact that nonmelanics are better chemically defended.     

Equality of feeding roles and the maintenance of monogamy in tree swallows

Summary We investigated the division of labor in male and female tree swallows by measuring the rate of food delivery to nestlings at 36 nests. By observing natural nests and performing brood manipulations we found that males and females divided the feeding duties about equally and responded similarly to changes in brood size and age. Feeding rate was most highly correlated with brood mass. Manipulation and removal experiments showed that increased feeding rates could be elicited, but only for limited periods of time. Male and female tree swallows could only partially compensate in feeding nestlings when mates were removed. This, along with the higher mortality in enlarged broods and in those raised by single parents, indicates that both male and female are required to raise an entire brood to fledging. We argue that this requirement contributes to the absence of mate guarding and the maintenance of monogamy in the tree swallow.