Male and female condition influence mating performance and sexual receptivity in two tropical fruit flies (Diptera: Tephritidae) with contrasting life histories

Recent recognition of widespread polyandry in insects has generated considerable interest in understanding why females mate multiple times and in identifying factors that affect mating rate and inhibit female remating. However, little attention has been paid to understanding the question from both a female and male perspective, particularly with respect to factors that may simultaneously influence female remating rates. Here, we report on a study aimed at ascertaining the possible interactive effects that male and female size and diet, and female access to a host could have on mating latency, probability, and duration and female refractory period using two tropical fruit fly species with contrasting life histories. Of all factors tested, adult diet played the most significant role. Both Anastrepha ludens and Anastrepha obliqua males which had constant access to protein and sucrose mated more often, had shorter copulations and induced longer refractory periods in females than males fed a low quality diet (sucrose offered every third day). Female size and the interaction with male diet determined how quickly female A. ludens mated for the first time. Smaller females mated sooner with low quality fed males than with high quality fed males while there was no difference for large females, suggesting that male choice may be at play if high quality fed males discriminate against smaller females. Copulation duration also depended on both male and female nutritional condition, and the interaction between male diet and female size and diet. Large and high quality fed females had shorter copulations regardless of male condition. Importantly, for A. ludens, female refractory period depended on male size and the nutritional condition of both males and females, which could indicate that for this species, female receptivity does not depend only on the condition of the male ejaculate. For A. obliqua refractory period was associated with the interaction between male size and diet and male diet and host presence. We discuss our results in terms of male ability to inhibit female remating and the relative contribution of female condition to this behavior. We also address the importance of studying effects simultaneously on species with contrasting life histories.     

Life history and demography of the predatory mite,Amblyseius longispinosus Evans

The life history and demography of Amblyseius longispinosus Evans were studied under laboratory conditions 25–28°C and 65–85% RH using the red form of the twospotted spider mite, Tetranychus urticae Koch complex as prey. The entire development from egg to adult averaged 102.5 h for both sexes with a survival rate of 90.0%. Immature females consumed more prey eggs than males, averaging 7.9 and 6.4 eggs/day, respectively. The embryonic development time for male eggs was longer (average 45.2 h) than for female eggs (average 42.6 h). The average duration of succeeding stages did not differ between the sexes: larva, 15.7; protonymph, 21.1; and deutonymph, 23.0 h. Larvae were not observed to feed. Statistically significant differences between sexes were noted with respect to consumption of the deutonymphs (female, average 4.0±0.2; male, average 2.9±0.3 eggs/day) but not of the protonymphs. Mating occurred on the same day as adult emergence and was repeated several times during the reproductive life. Egg laying started on the second day after emergence. Parameters relating to oviposition were: fecundity, gross (50.7) and net (43.3) eggs/female; hatchability, 99.6% (gross) and 99.5% (net); the oviposition lasted at most 28 days and showed a peak in the first week. Net reproductive rate (R₀) was 36.7 female offspring/female/generation time (T) of 9.0 days. The sex ratio was biased toward the females (71.9%), while the intrinsic rate of increase (r_m) was 0.4 with a doubling time (DT) of 1.7 days. Males lived longer (36 days) than females (30 days) with a LT₅₀ for males of 26 days and for females of 22 days.     

Effects of host-plant quality on male two-spotted spider mite (Acari: Tetranychidae) mate location and guarding behavior

Effects of host-plant quality on two-spotted spider mite,Tetranychus urticae Koch, mate location and guarding behaviors were described using a no-choice bioassay. Males and quiescent deutonymphs were collected from lima bean leaves of one of two host qualities. High-chlorophyll (HC) leaves had been infested with spider mites for 6–10 days, while low-chlorophyll (LC) leaves had been infested for>21 days. Three parameters of maleT. urticae guarding behavior were quantified: approach arrestment, and arrestment duration. HC males approached quiescent deutonymphs more often than did LC males, even though host quality of females had no effect on male approach frequency. HC males were arrested more frequently by HC quiescent deutonymphs than were LC males, while LC males were arrested more often by LC females than were HC males. However, a different pattern was observed for arrestment duration. HC males were arrested for twice as long by LC quiescent deutonymphs than by HC females, while the LC-male arrestment durations elicited by HC and LC females did not differ. These results show that host-plant quality affectsT. urticae intersexual communication, in terms of both the female signal and the male response. Whether the differing male responses observed in this study indicate alternativeT. urticae mating strategies or are incidental by-products of host-induced physiological changes remains to be determined.     

Influence of host-plant density and male harassment on the distribution of female Euphydryas anicia (Nymphalidae)

Summary We studied behavioral mechanisms underlying the spatial distribution of Euphydryas anicia (Lepidoptera: Nymphalidae) females searching for larval host plants on a flat plain in Colorado. The rate of female movement, as represented by an empirically estimated diffusion coefficient, is affected by two factors. First, when searching in areas of high host-plant density, females tend to make sharper turns and fly for shorter distances between landings, compared to areas of low plant density. As a result, the rate of female displacement is lowered when they search in areas of high host-plant density, and thus females tend to aggregate in such areas. The second factor affecting female movement is the presence of males. Harassment by males induces females to increase the rate of their movement, and females are often chased out of host patches by males. Our results explain the observed spatial and temporal patterns of female distribution in relation to the distribution of host plants and males. In years when host plants are plentiful, harrassement by males in one host patch induces females to move to another. This pattern of female movement tends to disrupt the relationship between the spatial distributions of host plants and females, as well as females and males. In the year when hosts were concentrated in a single large patch, females could not easily avoid male harassment, and consequently we observed strong corelations of female-host distributions and male-female distributions.     

Male mate choice in the Andrew’s toad <Emphasis Type="Italic">Bufo andrewsi</Emphasis>: a preference for larger females

In amphibians, theory predicts that male mate choice with respect to female body size can be expected to occur when female fecundity is related to body size and when the time and energy invested into one mating are relatively large. Based on experimental observations, we tested whether male mate choice occurs in a population of the Andrew’s toad (Bufo andrewsi), a species in which both assumptions are likely to be met. When a male B. andrewsi was placed with a gravid female and a non-gravid similar-sized female, the male did not discriminate between them. When two gravid females with distinct size differences were provided to a male, the male preferred the larger one. In an experiment in which two different-sized gravid females were put in two separate transparent cylinders to exclude potential chemical cues, males spent more time in proximity to the larger gravid females and jumped more frequently towards the larger gravid females than the smaller ones. These findings suggest that male B. andrewsi recognizes female body size, exhibits mate choice, and prefers to mate with larger females that provide greater reproductive potential.     

Body size variation in <Emphasis Type="Italic">Gryon gallardoi</Emphasis> related to age and size of the host

This study aimed to evaluate the influence of Spartocera dentiventris (Berg) (Hemiptera: Coreidae) egg quality (regarding age, size and superparasitism) on male and female body size of Gryon gallardoi Brèthes (Hymenoptera: Scelionidae). It was also analysed host size influence on offspring sex choice and female oviposition order (within egg groups). Groups of 12 eggs, 2, 3, 4, 5, 6, 7, 8, and 12 days old were individually exposed to a female parasitoid for 2 h, and the order of each egg parasitisation was recorded. Size of adult parasitoids (head width and tibia length of the second leg pair), and parasitised and superparasitised host eggs were measured. Sexual dimorphism related to size (SDs), was estimated through a model II linear regression. Females have a larger head width but a smaller tibia length. Parasitoid adult size is significantly related to host age and size. Within an egg group, females did not exhibit preference for any egg size category either to start oviposition or to oviposit male or female eggs. The body size of both sexes diminished with host aging. There was a positive response in both sexes to host size increasing. Estimated SDs diminished with host aging and increased with host volume augmentation. The results suggest that adult body size of G. gallardoi, as well as size differences between males and females, are strongly associated to host quality.     

Interpatch movement of the red milkweed beetle,Tetraopes tetraophthalmus: individual responses to patch size and isolation

Individual movement patterns and the effects of host plant patch size and isolation on patch occupancy were examined for red milkweed beetles, Tetraopes tetraophthalmus, residing in a heterogeneous landscape. Male beetles were found to move both more often and farther between host plant patches than female beetles, and this difference affected the patterns of patch occupancy observed. Overall, unoccupied milkweed patches were smaller and more isolated than patches occupied by beetles. Patches uninhabited by females tended to be more isolated, but not necessarily smaller, than patches with female beetles, indicating that females may be affected more by patch isolation than patch size. Presence of male beetles on patches showed a stronger response to patch size than to patch isolation. Differences in movement between males and females illustrate the need for demographically based dispersal data. Comparisons of Tetraopes interpatch movement patterns between landscapes composed of patches of different size revealed that landscapes with overall smaller patches may have greater rates of interpatch movement.     

Male-biased sex ratios,female promiscuity,and copulatory mate guarding in an aggregating tropical bug,Dysdercus bimaculatus

The ecological and social bases of the mating system of the seed-feeding bug, Dysdercus bimaculatus(Hemiptera: Pyrrhocoridae), were studied in the lab and in aggregations at the host tree, Sterculia apetala(Malvales: Malvaceae), in Panama. On theoretical grounds, two factors are predicted to be of importance in determining the evolution of male mating tactics in Ms species: the operational sex ratio and the probability that undefended females will mate with other males, subjecting the gametes of deserters to sperm competition. Results of a study of a related species suggested that sperm displacement is probably substantial. Adult sex ratios at numerous sites were significantly male biased, and females whose mates were removed remated before oviposition (i. e., sperm utilization). These results predict that a mate defense tactic is likely to be superior to a nondefense tactic. The biological significance of the parameters is supported by observations that captive pairs often remained in copulafor several days, until just before oviposition. However, substantial variation in copulation duration was also observed, and possible causes of this variation are considered. Causes of male biased adult sex ratios were investigated by monitoring demographic changes within a single aggregation over 2 months. Both female juvenile and adult mortality rates were greater than male. In addition, dissections of reproductive adults showed that the flight muscles of females, but not males, had histolyzed, so that female reproduction is physiologically limited to a single site. Greater rates of immigration among both mature and young males suggests that an excess of males may also be found in the populations of bugs that subsequently colonize other host plants, so that female scarcity is typical of aggregations in all stages of development. The evolution of sex-limtied flight muscle histolysis may be explained by greater patchiness of females than males as mating resources, plus a lower energetic benefit/cost ratio of histolysis for males.     

Male Social Behavior and Dominance Hierarchy in the Sulawesi Crested Black Macaque (Macaca nigra)

In a 6-week study of the social behavior of wild Sulawesi crested black macaques (Macaca nigra), we found a linear and transitive dominance hierarchy among the six adult males in one social group. Dominance rank, as determined by the direction of supplantations, correlated strongly with percentage of time near more than four neighbors, frequency of grooming received from adult females, and percentage of time with an adult female as nearest neighbor. These results suggest that high-ranking males are socially attractive. Adult females sexually solicited high-ranking males more often than low-ranking males, but frequency of copulation was not correlated with dominance rank. Frequency and intensity of aggression between males are strongly correlated with rank distance, but aggression toward females was greatest for mid-ranking males. Males of all rank displayed significantly more aggression toward sexually receptive females than toward females in other estrous states. These data indicate that male Sulawesi crested black macaques display a social organization similar to that reported for multimale groups in other macaque species rather than the egalitarian social organization described for female Sulawesi macaques.     

Response of German cockroaches to aggregation pheromone emitted by adult females

The response of early, middle and late instar (mixed sexes), adult male, and gravid and non-gravid female German cockroaches to aggregation pheromone on filter papers conditioned by gravid and non-gravid females was studied. Cockroaches were attracted to and settled on each of the conditioned papers. Classes varied in the strength of their response to conditioned papers, relative response to gravid versus non-gravid female conditioned paper, and in clustering behavior. A significantly greater attraction of all classes except middle instars and non-gravid females to paper conditioned by non-gravid females suggests that non-gravid females emit more aggregation pheromone than gravid females. Possible explanations of differences in response with age/sex class, including an enhanced attraction of males from female sex phermone, are discussed.

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Résumé L'étude a porté sur les réponses de six groupes/âges/sexes de B. germanica à des papiers filtres imprégnés de phéromones émises par des femelles pleines ou non. Tous les groupes ont été attirés par les papiers traités, mais les attractions des stades larvaires moyens ou âgés ont été inférieures aux autres. 4 groupes sur 6 ont été plus attirés par le papier traité avec des femelles ne contenant pas d'oeufs mûrs que par les femelles pleines, ce qui suggère une plus forte émission par les femelles sans oeufs mûrs. Les stades larvaires moyens ont été les seuls à être plus attirés par le papier traité avec les femelles pleines. Les femelles ne contenant aucun oeuf ne manifestent aucune préférence. La tendance à former des amas inégaux était moins nette pour les stades larvaires moyens et âgés que pour les autres groupes. Les premiers stades ont montré une tendance très nette à se rassembler sur le même papier. Le rassemblement de mâles adultes sur du papier traité avec des femelles ne contenant pas d'oeufs peut avoir été partiellement une réponse à la phéromone sexuelle femelle qui avait imprégné le papier pendant le conditionnement.

     

Biological Attributes of the Nematode, Thripinema nicklewoodi, a Potential Biological Control Agent of Western Flower Thrips

Western flower thrips (WFT), Frankliniella occidentalis (Thysanoptera: Thripidae), is the most important insect pest for greenhouse flower crops, but chemical control is often difficult because of the thrips' location in flowers and buds. Thripinema nicklewoodi (Tylenchida: Allantonematidae) is an entomoparasitic nematode that attacks and sterilizes WFT, but its biology and impact on WFT are poorly understood. Methods to propagate and study T. nicklewoodi with a microscope slide arena for observation and a rolled bean leaf arena for rearing were developed. On average, 21.4 nematodes were excreted per day by a parasitized female WFT. The sex ratio of the excreted nematodes was 6.0/1.0 (female/male). After dissection of adult WFT, a maximum of 11 ovoid-shaped first-generation nematodes in a female thrips and 6 in a male thrips were found. There were more second-generation nematodes in the adult female WFT (192.6) than in the adult male WFT (93.7). When 50 healthy first instar WFT were exposed to 4 parasitized female WFT in a rolled bean leaf, a 75.3% mean parasitization rate in the adult stage of the thrips was obtained. In contrast to previous reports, male WFT can be parasitized as readily as females. Parasitism reduced the longevity of both adult female and adult male WFT, and the degree of reduction was higher in adult male WFT. T. nicklewoodi, when presented with various WFT life stages (first instar, second instar, prepupa, pupa, adult female, and adult male), achieved the highest attack rate in first and second instars and prepupa. The free-living nematodes excreted by the hosts actively migrated to a trichome on leaf disks in the observation arena and moved up this structure. Then, the nematode actively waved the anterior part of its body while attaching itself to the trichome with the posterior part of its body. After a nematode contacted the leg of a thrips, the nematode immediately moved up along the leg toward the abdomen of the host. Increased understanding of the biology of T. nicklewoodi is important to better assess its potential for biological control of WFT.     

Female Leadership During Migration and the Potential for Sex-Specific Benefits of Mass Spawning in the Brown Surgeonfish (Acanthurus nigrofuscus)

Spawning aggregations in coral-reef fishes have been hypothesized to confer any one of several mutually non-exclusive benefits, largely expected to serve the interests of both sexes simultaneously. Here we provide indication that in the brown surgeonfish, Acanthurus nigrofuscus, spawning aggregations may confer a sex-specific benefit. Following tagged individuals en-route to their daily spawning-aggregation site we found that while migrating groups (≤20 fish) consist of both males and females, females tend to occupy the lead position more often than expected by chance. In addition, we found evidence that female A. nigrofuscus divide their daily egg-stock among several spawning bouts within the aggregations. We propose that female leadership en-route to spawning aggregations, together with the potential benefits of multiple female mating, are consistent with a sex-specific benefit to spawning aggregations.     

Malarial parasitism and male competition for mates in the western fence lizard,Sceloporus occidentalis

Summary The effect of malarial parasitism on the ability of male western fence lizards, Sceloporus occidentalis, to compete for access to females was assessed experimentally. Pairs of male lizards, one infected with the malarial parasite, Plasmodium mexicanum, and the other not infected, were matched by size and color and placed in large seminatural outdoor enclosures along with an adult female lizard. Infected males displayed to females and to other males less often than did noninfected male lizards. Noninfected lizards were dominant in social interactions more often than malarious animals, based on duration and intensity of agonistic encounters toward the other male, and time spent with the female. Thus, malarial infection hinders the ability of male fence lizards to compete for mates.     

Distribution and fitness effects of the son-killer bacterium inNasonia

Summary Maternally inherited microorganisms that kill male (but not female) progeny are widespread in nature. Three hypotheses have been proposed for the evolution of male-killing microorganisms: inbreeding reduction, release of resources to remaining females and inoculum for horizontal transmission. The sonkiller bacterium,Arsenophonus nasoniae, is a maternally inherited bacterium that causes lethality of male embryos of infected females in the parasitoid wasp,Nasonia vitripennis. In this paper we describe the geographical distribution and frequency of the son-killer bacterium in North American populations ofN. vitripennis andNasonia longicornis. We tested the resource release hypothesis using the body size measurements of infected and uninfected females from natural populations. No evidence was found for a fitness increase of females infected with the bacterium compared to uninfected females. We propose a modification of the existing models, termed the incremental gain hypothesis. According to this model, the bacteria are maintained in host populations due to horizontal transmission and male killing provides an incremental gain in the fitness of infected females relative to females infected with non-male-killing bacteria.     

Strength in numbers: high parasite burdens increase transmission of a protozoan parasite of monarch butterflies (<Emphasis Type="Italic">Danaus plexippus</Emphasis>)

Parasites often produce large numbers of offspring within their hosts. High parasite burdens are thought to be important for parasite transmission, but can also lower host fitness. We studied the protozoan Ophryocystis elektroscirrha, a common parasite of monarch butterflies (Danaus plexippus), to quantify the benefits of high parasite burdens for parasite transmission. This parasite is transmitted vertically when females scatter spores onto eggs and host plant leaves during oviposition; spores can also be transmitted between mating adults. Monarch larvae were experimentally infected and emerging adult females were mated and monitored in individual outdoor field cages. We provided females with fresh host plant material daily and quantified their lifespan and lifetime fecundity. Parasite transmission was measured by counting the numbers of parasite spores transferred to eggs and host plant leaves. We also quantified spores transferred from infected females to their mating partners. Infected monarchs had shorter lifespans and lower lifetime fecundity than uninfected monarchs. Among infected females, those with higher parasite loads transmitted more parasite spores to their eggs and to host plant leaves. There was also a trend for females with greater parasite loads to transmit more spores to their mating partners. These results demonstrate that high parasite loads on infected butterflies confer a strong fitness advantage to the parasite by increasing between-host transmission.     

Male Sexual Signals and Female Choice in Drosophila grimshawi (Diptera: Drosophilidae)

Sexual selection and sexual signaling have been prominent topics in recent behavioral studies, but limited data have led to controversy regarding these topics. For example, the Hawaiian Drosophila are often cited as examples in which female choice has resulted in the evolution of elaborate male courtship signals, but relatively few data exist to test these claims adequately. We studied D. grimshawi, a lek-forming Hawaiian Drosophila, to determine whether there was evidence for female choice without male competition and to elucidate the possible cues females use to discriminate. Male mating success was found to be nonrandom and males that courted females intensely and deposited many pheromone-containing streaks on the substrate were the most successful. Hence, multiple cues seem to be involved in male mating success in this species. Some males performed only one display, however, and may represent an alternate male mating tactic. The protein content of the adult male diet significantly influenced the level of pheromone streak deposition, and thus, foraging environment may affect the outcome of sexual selection.     

The nutritional contribution of males affects the feeding behavior and spatial distribution of females in a bruchid beetle,<Emphasis Type="Italic"> Bruchidius dorsalis</Emphasis>

The relationship between the quality/quantity of male investment and the feeding behavior of females was investigated in a bruchid weevil, Bruchidius dorsalis (Fahraeus), whose males donate nutrition via seminal fluid to females. Experiments on the effect of feeding regimes of both sexes on the mating frequency of females showed that females mated at a higher frequency if given low-quality food or poor male investment. On the other hand, the experiment that examined the effect of male investment quality on female feeding behavior showed that females receiving the high-quality investment exhibited feeding behavior less often. These results suggest that male investment and feeding behavior play the same role for B. dorsalis females. These experiments also showed that there are sex-related asymmetries in mating and feeding behaviors: females mated more often but males fed more often. Moreover, a field census suggested that only males visited non-host flowers to feed on the pollen and nectar during the non-flowering period of the host plants; females always stayed on the host plants irrespective of the flowering phenology. These results suggest that in B. dorsalis courtship role reversal and sex-specific feeding modes are fairly fixed and obligatory, and that male investment, derived from sexual selection, could affect the feeding behavior and spatial distribution of both sexes, which may have far-reaching impact in various ecological contexts.     

Female Coalitions Against Male Aggression in Wild Chimpanzees of the Budongo Forest

In the wild, female chimpanzees (Pan troglodytes) are subject to male aggression that at times can be prolonged or particularly violent. There are no reports of cooperative retaliation to such aggression, a strategy observed in the congeneric Pan paniscus, from the wild despite >4 decades of detailed behavioral study across a number of populations and its occurrence among captive female chimpanzees. If the reports from captivity represent an inherent capacity, then the absence of similar reports from wild populations suggests that females may be able to form coalitions only under appropriate ecological and demographic conditions. During a study of male and female aggressive interactions among chimpanzees of the Budongo Forest, Uganda, wild adult female chimpanzees sometimes formed coalitions with one another to retaliate against male aggression. This may be possible because these females tend to be more gregarious than in other populations of East African chimpanzees, as other studies of the same population have suggested; the extent and variation of female chimpanzee social strategies may, therefore, need reconsideration. Further, my observations strengthen the argument that at least some of the differences between chimpanzees and bonobos may be more of degree than of kind.     

Colony dynamics in a primitively eusocial halictine beeLasioglossum (Dialictus) zephyrum (Hymenoptera: Halictidae)

Summary Proximate control of colony dynamics was studied in the primitively eusocial halictine beeLasioglossum (Dialictus) zephyrum using allozyme markers. The results indicate that workers produce on average 15% of the male brood (range=0–50%) in small laboratory colonies made up of unrelated, single-generation, uninseminated females. This proportion is not influenced by colony size, but is influenced by the relative size of the queen. Large queens are more successful in dominating their workers than are small queens, the queen being defined as the female that is the mother of most of the brood produced in the colony. Older and larger females tend to become queens. Thus, while small differences in age (up to 4 days) influence which female becomes a queen, her ability to control her workers is primarily influenced by her relative size. The proportion of reproduction that is co-opted by the queen is negatively correlated with colony reproductivity (the number of males/day/female). Colony reproductivity is also negatively correlated with the standard deviation in size among females.     

Thermal biology of reproduction in viviparous skinks,Eulamprus tympanum: why do gravid females bask more?

Summary In some reptiles, gravid females bask more, and/or maintain higher body temperatures than do males or non-gravid females. This phenomenon is usually explained in terms of the female or her offspring benefitting from accelerated embryogenesis and early birth, but the effect of increased basking on gestation period has not been studied. In a laboratory experiment, gestation periods of gravid female skinks (Eulamprus tympanum) decreased with the duration of access to radiant heat. Embryonic development was more rapid in the laboratory than in the field, and there were no apparent adverse effects of this accelerated gestation on females or offspring. Number and mass of offspring, survival rates of embryos, relative clutch mass and female mass before and after parturition were not influenced by the decrease in gestation period caused by increased basking. Females selected similar temperatures in the laboratory and field (32° C), despite the availability of higher temperatures in the laboratory. Thus, gestation in the laboratory was accelerated by spending longer periods at usual basking temperatures, rather than by selecting higher temperatures. In the field, mean and modal body temperatures of active animals were similar in gravid females, males and non-gravid females, but gravid females appear to bask more of the time, even in cloudy weather when other members of the population do not bask. Hence, an apparent similarity in thermal regimes of gravid and non-gravid animals may mask significant underlying differences in thermoregulatory strategies.