A selective trade-off for territoriality and non-territoriality in the polymorphic damselfly Mnais costalis

Males of the damselfly Mnais costalis occur as territorial orange-winged 'fighter' males or non-territorial clear-winged 'sneaker' males. Their morph life histories differ considerably but the estimated lifetime reproductive success is the same for the two morphs. In this study we compared the developmental and reproductive costs associated with the two morphs. Orange-winged male and female reproductive costs resulted in a decline in adult fat reserves with increasing age. In contrast, the fat reserves of clear-winged males remained constant with adult age. Body size was positively correlated with mating success in orange-winged males, but had no influence on the mating success of clear-winged males. The orange-winged male flight muscle ratios (FMRs) were significantly higher than the clear-winged male and female FMRs. However, there was no difference in the size-corrected fat reserves of the two morphs; both had higher fat reserves than females. The gain in mass between eclosion and reproduction in orange-winged males and females was almost double the mass gained by clear-winged males, suggesting that clear-winged male development is less costly. An experiment in which pre-reproductive levels of nutrition were manipulated confirmed this.     

Male reproductive polymorphism and form-specific habitat utilization of the damselflyMnais pruinosa (Zygoptera: Calopterygidae)

Habitat utilization patterns were studied in damselfly males,Mnais pruinosa, which have two male forms with different reproductive behaviors. The ‘esakii’ (orange-winged males) were territorial around oviposition sites, while the ‘strigata’ (hyaline-winged males) were non-territorial, often sneaking into theesakii's territory or loitering on the foliage of vegetation along stream banks. The place in the stream in the study area where females frequently appeared was covered by reeds and had abundant oviposition sites. It was difficult for the territorialesakii to stay there because the crowded reeds prevented them from defending their territory and discovering the females. Furthermore, there was competition for the limited territorial space with another species,Mnais nawai. Strigata males concentrated in this place in direct proportion to the number of females. Females frequently mated withstrigata males and probably deposited eggs fertilized bystrigata sperm rather thanesakii sperm. The density ofstrigata was higher than that ofesakii in this study area. If the average reproductive success ofesakii andstrigata males is equal, this may indicate that the equilibrious point between the two male forms is biased towardstrigata.     

The genetic polymorphism linked to mate-securing strategies in the male damselfly<Emphasis Type="Italic"> Mnais costalis</Emphasis> Selys (Odonata: Calopterygidae)

Alternative male mate-securing strategies are widespread among animal taxa, but there are few well-documented examples of genetic polymorphisms for them. In the Japanese calopterygid damselfly Mnais costalis, males occur as either orange-winged territorial fighter males, or clear-winged non-territorial sneaker males. It has previously been suggested that this behavioral polymorphism is genetically controlled. However, there was no direct evidence for this. By rearing two groups of larvae from the same female but sired by different male morphs, I show that differential morph development is genetically controlled and consistent with a single-locus two-allele autosomal genetic polymorphism.     

Measuring male-female relationships during the mating season in wild Japanese macaques (Macaca fuscata yakui)

Heterosexual relationships during one mating season were examined in a wild troop of Japanese macaques (Macaca fuscata yakui) on Yakushima Island, Japan. Validation tests of putative mate choice behaviors demonstrated that female initiation and maintenance of proximity, female lookback at the male, and sexual presents to the male, were associated with increased mating. Male grooming the female was also associated with increased mating. Ten dyadic social behaviors were subject to principal components analysis to empirically define behavioral dimensions of male-female relationships. The analysis yielded four relationship dimensions: ‘Mutual Choice and Male Coercion,’ ‘Female Choice’ (two types), and ‘Mutual Choice’ Dyads tended to be characterized by more than one dimension. The results suggested that females sought matings with multiple males of various dominance ranks. Female relationships with high ranking males contained elements of male coercion and mate guarding, however, because these males attempted to inhibit females from mating with lower ranking males. The correlation between each relationship dimension and mating success depended, in part, on the dominance rank of males. Relationships involving high ranking males, which were most likely to contain elements of male coercion and mate guarding, were associated with mating success. Relationships involving low ranking males, which usually lacked such coercive elements. were less strongly correlated with mating success. These results, obtained from a wild troop, are compared to those previously obtained in captive and provisioned groups of Japanese macaques.     

Small plot mating disruption trials for tufted apple bud moth,Platynota idaeusalis, in Pennsylvania apple orchards

Mating disruption treatments for the tufted apple bud moth (TABM),Platynota idaeusalis (Walker), were tested in small plot trials in apple orchards in Pennsylvania. Treatments were evaluated by fruit injury and by capture of male TABM in traps baited with synthetic pheromone sources or virgin females. The TABM pheromone is a two component isomeric blend ofE-11-tetradecen-1-ol (E11-14:OH) andE-11-tetradecenyl acetate (E11-14:Ac). A 50∶50 ratio of these two components was used in standard monitoring septa and in mating disruption treatments released from either hollow fibers (‘fiber’) or PVC tubes (‘PVC’). Other pheromone blends tested included a 90∶10 ratio of E11-14: Ac and E11-14:OH (‘EAc’) and its reverse (‘EOH’), mixture ‘EAc’ with 30% of the Z-isomers (‘low AEc’), and a blend similar to the preceding with 2% Z9-12:Ac (‘generic’). These other blends were released from multi tube tape (‘tape’) or Shin-Etsu type rope (‘rope’) dispensers. Seasonal dispenser release rate in mg ha⁻¹ h⁻¹ was ca. 30 for the ‘rope’ dispensers, 14 for ‘PVC’ and 6 for ‘fiber’. ‘EAc-tape’ and ‘EOH-tape’ were equally effective in reducing catches of males in traps baited with synthetic lures and in traps baited with virgin females. Both treatments also reduced fruit injury. ‘EAc-rope’, ‘fiber’ and ‘PVC’ also were generally effective; whereas, the ‘low EAc’ and ‘generic’ treatments reduced trap capture less than 90% and did not reduce fruit injury. Dispenser density was positively correlated with reduction in trap capture for the ‘low EAc-rope’ and ‘genericrope’ treatments. Traps loaded with ‘fiber’ dispensers captured more male TABM than the other treatments in non-pheromone permeated environments. Trap capture of other tortricids was reduced in pheromone treatments. ‘EAc-rope’ and the ‘TABM’ treatments provided mean (s.e.) percent reduction in trap catch of 99.5 (0.4) and 42.9 (10.1), respectively, for the redbanded leafroller,Argyrotaenia velutinana (Walker), and 90.4 (6.8) and 90.4 (1.3), respectively, for the obliquebanded leafroller,Choristoneura rosaceana (Harris).     

Estimation of mortality by entomophages on exotic Harmonia axyridis versus native Adalia bipunctata in semi-field conditions in northern Italy

A semi-field experiment was carried out in two peach orchards in northern Italy to assess mortality due to predators and parasitoids on the exotic coccinellid Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in comparison with the native coccinellid Adalia bipunctata L. (Coleoptera: Coccinellidae). The experiments were conducted in cages to avoid the possible escape of the exotic ladybird (not yet established in Italy). Two kinds of cage experiments were included: ‘exclusion cages’ (access by walking predators impeded) and ‘free cages’ (walking predators free to enter). The cages, containing all the stages of the two ladybird species, were placed in two localities and left for 24 h. All ladybird stages used for the semi-field experiments came from a laboratory rearing. The eggs of H. axyridis experienced less mortality than those of A. bipunctata. The ant workers were the most frequent predators in ‘free cages’ but A. bipunctata cannibalism on eggs was also detected. Larvae of both coccinellid species were predated equally but larval predation of L1 and L2 was higher in comparison to predation of L3 and L4. Pupae and adults of both exotic and native ladybirds were never attacked by predators. Predation on younger larval stages was higher in the ‘free cages’ in comparison with ‘exclusion cages’. No ladybird parasitisation was observed. The ‘free cage’ technique seems to provide a standardised and realistic estimation of predation impact but more studies are needed to evaluate ladybird parasitisation in semi-field conditions.     

Reproductive parameters in relation to food supply in the whiskered tern ( Chlidonias hybrida )

It is generally accepted that breeding terns are sensitive to food supply and that their reproductive effort could be substantially affected by the availability and access to resources. In this study we examined reproductive parameters in the whiskered tern Chlidonias hybrida in relation to food supply during the courtship feeding period (food brought by males to females) over a 2-year period (2004–2005). We also studied whether the condition of the nestlings [body condition index (BCI)] was related to a proxy of the reproductive investment of the adults (the clutch size) during a season of food shortage. Behavioural observations showed a decrease in the intensity of male courtship feeding between years (2004 > 2005), and a strong shift in the relative abundance of the two prey groups (invertebrates/fish; invertebrate prey dropped from 88.0 to 49.3%) brought by males. This change in food delivery rates did not result in a delay in laying, but there was a significant difference in mean clutch size between years (2.71 ± 0.49 eggs in 2004 and 2.05 ± 0.78 eggs in 2005) without any within-year variation in relation to the laying dates. The egg size (volume and length) was related to the year (2004 < 2005), suggesting a trade-off in the quantity and the quality of eggs between the two seasons. We also found no evidence that the investment in a large clutch affected nestling BCI in the course of the food shortage season. Since many pairs (about 60%) interrupted breeding during the incubation stage, we assumed that parents that succeeded in rearing nestlings in these conditions were probably ‘high-quality’ individuals. Our results therefore showed that whiskered terns are sensitive to the varying food conditions they experienced throughout the courtship period. The diversity of prey types could be a key factor in the reproductive investment of this tern species.     

Male reproductive pattern in a polygynous ungulate with a slow life-history: the role of age, social status and alternative mating tactics

According to life-history theory age-dependent investments into reproduction are thought to co-vary with survival and growth of animals. In polygynous species, in which size is an important determinant of reproductive success, male reproduction via alternative mating tactics at young age are consequently expected to be the less frequent in species with higher survival. We tested this hypothesis in male Alpine ibex (Capra ibex), a highly sexually dimorphic mountain ungulate whose males have been reported to exhibit extremely high adult survival rates. Using data from two offspring cohorts in a population in the Swiss Alps, the effects of age, dominance and mating tactic on the likelihood of paternity were inferred within a Bayesian framework. In accordance with our hypothesis, reproductive success in male Alpine ibex was heavily biased towards older, dominant males that monopolized access to receptive females by adopting the ‘tending’ tactic, while success among young, subordinate males via the sneaking tactic ‘coursing’ was in general low and rare. In addition, we detected a high reproductive skew in male Alpine ibex, suggesting a large opportunity for selection. Compared with other ungulates with higher mortality rates, reproduction among young male Alpine ibex was much lower and more sporadic. Consistent with that, further examinations on the species level indicated that in polygynous ungulates the significance of early reproduction appears to decrease with increasing survival. Overall, this study supports the theory that survival prospects of males modulate the investments into reproduction via alternative mating tactics early in life. In the case of male Alpine ibex, the results indicate that their life-history strategy targets for long life, slow and prolonged growth and late reproduction.     

Environmental mediation of intraguild predation between the freshwater invader Gammarus pulex and the native G. duebeni celticus

Invasive species and environmental change often occur simultaneously across a habitat and therefore our understanding of their relative roles in the decline of native species is often poor. Here, the environmental mediation of a critical interspecific interaction, intraguild predation (IGP), was examined between invasive (Gammarus pulex) and native (G. d. celticus) freshwater amphipods. In the laboratory, IGP asymmetries (males preying on congeneric females) were examined in river water sourced from zones where: (1) the invader has completely displaced the native; (2) the two species currently co-exist, and (3) the native currently persists uninvaded. The invader was always a more effective IG predator, but this asymmetry was significantly weaker moving from ‘invader-only water’ through ‘co-existence water’ to ‘native-only water’. The constituent of the water that drives this mediation of IGP was not identified. However, balancing the rigour of laboratory experiments with field derived ‘environment’ has advanced understanding of known patterns in a native species decline, and its co-existence and persistence in the face of an invader.     

The influence of host fruit morphology on parasitization rates in the Caribbean fruit fly,Anastrepha suspensa

Among the host fruits of the Caribbean fruit fly there are a variety of sizes and shapes. These morphological differences may influence the vulnerability of the larvae to parasites. In the laboratory, Caribbean fruit fly larvae placed in the smaller of 2 different sizes of artificial ‘fruit’ (cloth spheres filled with a diet material) were parasitized at a higher rate by the braconid,Diachasmimorpha longicaudata (Ashmead) when spheres were presented separately. However, when parasites were simultaneously presented with 6 different sizes of ‘fruit’ there was no significant relationship between size and parasitization rate. This may be due to the parasites preference to search for larvae in larger ‘fruit’. In field collections of different species of host fruit, a significant inverse correlation exists between fruit radius and rate of parasitization. However, host fruit size accounts for only about 5% of the variance in yearly parasitization rates.      

Sperm Competition and Optimal Timing of Matings in Microcebus murinus

Female promiscuity is common in mammals and leads to sperm competition: the sperm of ≥2 males compete for ova. Scientists understand the possible role of optimal insemination periods for male reproductive success in many species as well as the impact of monopolization of receptive females. Information from experiments combined with detailed observations from the field that allow determining the relative impacts of the elements in the same species are rare. We studied sperm competition and the role of optimal insemination periods in gray mouse lemurs (Microcebus murinus), a small solitary nocturnal primate from Madagascar. We used controlled matings to identify the relative impact of both contest and scramble competition, which characterize their mating system, on paternity. Fifteen females mated with 3–6 males in quick succession. Our experiments revealed that the optimal insemination period is during early receptivity. Early but not first mating males are more likely to sire offspring. Comparison with our field data indicate that the timing of male monopolization efforts correspond with the optimal insemination period.     

Expression of a costly, plastic secondary sexual trait is correlated with age and condition in a damselfly with two male morphs

Males of the damselfly Mnais costalis Selys (Odonata: Calopterygidae) are morphologically and behaviourally polymorphic, typically existing as clear-winged non-territorial ‘sneaks’ and orange-winged territorial ‘fighters’. The amount of orange pigment in the wing, as measured with a chromameter, varied between individuals, and decreased as the reproductive season progressed. Young individuals maintained in the laboratory on high or low nutrient diets differed in the amount of pigment that developed in the wing. Males in the high nutrient group developed darker wings faster than those in the low nutrient group. Young adults of both sexes and morphs were fed ¹⁴C-radiolabelled tryptophan or tyrosine (precursors of the pigments ommochrome and melanin, respectively). Ommochrome was restricted to the pseudopterostigma of the males of both morphs and was not present in females. The presence of tyrosine in the wing cells of orange males, but not of clear males, indicated that the orange pigment is at least partly constituted from melanin. These data show that at least some pigment levels must be maintained continuously in the wings of orange males, and that maintenance is costly as it is compromised at low nutrient levels.     

Parasitism ofLeptinotarsa decemlineata [Coleoptera: Chrysomelidae] byEdovum puttleri [Hymenoptera: Eulophidae] in different cultivars of eggplant

Experiments were conducted to quantify parasitism of Colorado potato beetle,Leptinotarsa decemlineata (Say), by the egg parasitoid,Edovum puttleri Grissell, on 3 different cultivars of eggplant,Solanum melongena L. Levels of parasitism were higher (P<0.05) on ‘Black Pride’ than on other cultivars. The percentage of egg masses that were parasitized was 1.2-fold higher (P<0.05) on ‘Black Pride’ than on ‘Harris Special’ and ‘White’. The number of eggs per mass that were parasitized was 1.3- and 1.4- fold greater (P<0.05) on ‘Black Pride’ than on ‘Harris Special’ and ‘White’, respectively. The percentage of eggs that were parasitized per mass and percentage of emerged adult parasitoids did not differ (P>0.05) among cultivars; between 2.1- to 2.6- fold more females than males emerged from eggs on all cultivars during the growing season.Edovum puttleri suppressed the 2nd generation ofL. decemlineata on ‘Black Pride’ and ‘Harris Special’, but did not suppress populations on ‘White’.      

Aggressive and Docile Colony Defence Patterns in Apis mellifera. A Retreater–Releaser Concept

Colony defence in Apis mellifera involves a variety of traits ranging from ‘aggressive’ (e.g. entrance guarding, recruitment of flying guards) to ‘docile’ (e.g. retreating into the nest) expression. We tested 11 colonies of three subspecies (capensis, scutellata, carnica) regarding their defensiveness. Each colony was selected as reportedly ‘aggressive’, ‘intermediate’ or ‘docile’ and consisted of about 10,000 bees. We applied three stimulation regimes (mechanical disturbance, exposure to alarm pheromones, and the combination of both) and measured their behaviours by tracking the rates of outflying bees at the entrance sites of the test hives. We provided evidence that for mechanical disturbances the test colonies resolved into two response types, if the ‘immediate’ defence response, assessed in the first minute of stimulation, was taken as a function of foraging: ‘releaser’ colonies allocated flying guards, ‘retreater’ colonies reduced the outside-hive activities. This division was observed irrespective of the subspecies membership and maintained in even roughly changing environmental conditions. However, if pheromone and mechanical stimulation were combined, the variety of colony defensiveness restricted to two further types irrespective of the subspecies membership: six of nine colonies degraded their rate of flying defenders with increasing foraging level, three of the colonies extended their ‘aggressiveness’ by increasing the defender rate with the foraging level. Such ‘super-aggressive’ colonies obviously are able to allocate two separate recruitment pools for foragers and flying defenders.     

Coevolution of reproductive characteristics in three dioecious fig species and their pollinator wasps

This study investigates dioecious fig species using a pollinator introduction experiment. Our aims were to determine: (1) whether there was a significant difference in foundress distribution between sexes per fig species; (2) whether fig size and foundress number affect reproductive success of dioecious figs; and (3) who is the ‘controlling partner’ in the fig/pollinator mutualism. Three dioecious fig species: Ficus semicordata, Ficus hispida and Ficus tinctoria from Xishuangbanna, China, were selected for this experiment. We found that there was no significant difference of the foundress number in female and male figs of F. semicordata, F. hispida and F. tinctoria. Also, the foundress number did not depend on the fig diameter. The numbers and the proportions of fig seeds and female wasp offspring significantly increased with more foundresses; and fig seed number was significantly higher than female wasp offspring in F. semicordata and F. hispida, but not in F. tinctoria. Our results indicate that figs are generally the ‘controlling partner’ in fig-wasp mutualisms in species with large figs, but not with small figs. Compared with published studies of reproductive success in monoecious figs, the dioecious figs seem to be more efficient in producing both seeds and wasp offspring when there is a high number of foundress.     

Reproductive tactics and fertilization success of mature male Miyabe charr,Salvelinus malma miyabei

Synopsis Spawning behaviour, fertilization success and reproductive ecology of stream-resident and lake-run male Miyabe charr, Salvelinus malma miyabei, were studied under natural and artificial conditions. In Shikaribetsu Lake, Japan, the ratio of the two types of males is approximately 1:1, estimated by weir data. The majority of stream-resident males mature sexually at 2⁺ years of age, while the lake-run males begin to mature at 4⁺. In contrast to the large-bodied lake-run males, which paired with females, the smaller stream-resident males attempted to fertilize eggs by ‘streaking’ into the nest. We demonstrated through electrophoretic techniques that stream-resident males were able to fertilize eggs by streaking. Under artificial conditions, the estimated maximum proportion of eggs fertilized by a single stream-resident male was about 16.8. The evolutionary stability of the charr's dual mating system may be explained by Gross's theory for alternative life-history evolution, although further testing is necessary.     

What is the appropriate timescale for measuring costs of reproduction in a `capital breeder' such as the aspic viper?

Before we can quantify the degree to which reproductive activities constitute a cost (i.e., depress an organism's probable future reproductive output), we need to determine the timescale over which such costs are paid. This is straightforward for species that acquire and expend resources simultaneously (income breeders), but more problematical for organisms that gather resources over a long period and then expend them in a brief reproductive phase (capital breeders). Most snakes are capital breeders; for example, female aspic vipers (Vipera aspis) in central western France exhibit a 2- to 3-year reproductive cycle, with females amassing energy reserves for one or more years prior to the year in which they become pregnant. We use long-term mark-recapture data on free-living vipers to quantify the appropriate timescale for studies of reproductive costs. Annual survival rates of female vipers varied significantly during their cycle, such that estimates of survival costs based only on years when the females were ‘reproductive’ (i.e., produced offspring) substantially underestimated the true costs of reproduction. High mortality in the year after reproducing was apparently linked to reproductive output; low energy reserves (poor body condition) after parturition were associated with low survival rates in the following year. Thus, measures of cost need to consider the timescale over which resources are gathered as well as that over which they are expended in reproductive activities. Also, the timescale of measurement needs to continue long enough into the post-reproductive period to detect delayed effects of reproductive ‘decisions’. This revised version was published online in July 2006 with corrections to the Cover Date.     

Male barn swallows use different signalling rules to produce ornamental tail feathers

The evolution of secondary sexual characters is the subject of controversial debate between those defending their role as ‘viability indicators’ and those arguing that ornaments are purely ‘attractive traits’ selected by females. Recent theoretical studies suggest that these hypotheses are not mutually exclusive, as both viability and attractiveness can contribute to improve the reproductive success of progeny and could thus simultaneously underlie female choices. If that is the case, strategies of cheaper advertisement, allowing the expression of larger ornaments for the same cost, could proliferate even in species in which honest signalling of viability prevails. Under this scenario, different males could invest a different amount of resources per ornament unit of expression, thus using different signalling rules. We studied the relationship between tail feather length (a trait that is the subject of a female mate preference) and feather mass (a measure of investment in feather production) in a barn swallow Hirundo rustica population. Different males used different and consistent signalling rules when developing ornamental feathers. That is, to produce a feather of a given length, each male used a constant amount of resources across different years, but this amount varied between males. Although the amount of material invested in feathers (feather mass) is a condition-dependent trait, the organization of this material in ornamental feathers (i.e. the signalling rules) was not. Neither survival nor risk of feather breakage was related to the signalling rules. Thus, these results suggest that both ‘viability’ and ‘runaway’ mechanisms are independent determinants of the evolution of ornamental sexual feathers in the barn swallow. A preference for long tails will ensure that females either obtain a sire with high viability, or one transferring the capability to produce longer and more attractive tails at a lower cost of production to its offspring.     

Development of SCAR markers linked to self-incompatibility in <Emphasis Type="Italic">Brassica napus</Emphasis> L.

‘SI1300’ is a self-incompatible Brassica napus line generated by introgressing an S haplotype from B. rapa ‘Xishuibai’ into a rapeseed cultivar ‘Huayou No. 1’. Five S-locus specific primer pairs were employed to develop cleaved amplified polymorphic sequences (CAPS) markers linked the S haplotype of ‘SI1300’. Two segregating populations (F₂ and BC₁) from the cross between ‘SI1300’ and self-compatible European spring cultivar ‘Defender’, were generated to verify the molecular markers. CAPS analysis revealed no desirable polymorphism between self-incompatible and self-compatible plants. Twenty primer pairs were designed based on the homology-based candidate gene method, and six dominant sequence characterized amplified region (SCAR) markers linked with the S-locus were developed. Of the six markers, three were derived from the SRK and SP11 alleles of class II B. rapa S haplotypes and linked with S haplotype of ‘SI1300’. The other three markers were designed from the SLG-A10 and co-segregated with S haplotype of ‘Defender’. We successfully combined two pairs of them and characterized two multiplex PCR markers which could discriminate the homozygous and heterozygous genotypes. These markers were further validated in 24 F₃ and 22 BC₁F₂ lines of ‘SI1300 × Defender’ and another two segregating populations from the cross ‘SI1300 × Yu No. 9’. Nucleotide sequences of fragments linked with S-locus of ‘SI1300’ showed 99% identity to B. rapa class II S-60 haplotype, and fragments from ‘Defender’ were 97% and 94% identical to SLG and SRK of B. rapa class I S-47 haplotype, respectively. ‘SI1300’ was considered to carry two class II S haplotypes and the S haplotype on the A-genome derived from B. rapa ‘Xishuibai’ determines the SI phenotype, while ‘Defender’ carry a class I S haplotype derived from B. rapa and a class II S haplotype from B. oleracea. SCAR markers developed in this study will be helpful for improving SI lines and accelerating marker-assisted selection process in rapeseed SI hybrid breeding program.     

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.