A field test of the Bruce effect in the monogamous prairie vole (Microtus ochrogaster)

The Bruce effect is a widely studied reproductive phenomenon in rodents in which exposure of pregnant females to unknown males causes termination of the current pregnancy. The Bruce effect has been reported from numerous studies in the laboratory, and one field study with the promiscuous gray-tailed vole, Microtus canicaudus, failed to support it. We conducted a field study with the monogamous prairie vole, M. ochrogaster, to determine if complete replacement of the male population every 10-14 days affected pregnancy and juvenile recruitment. The mean days to first parturition for control and treatment females were 36.8 and 44.4 days. Fifty-five percent of control females and 33% of treatment females conceived within the first 14 days of the study. All control females and 79% of treatment females successfully delivered at least one litter. These differences between treatment and control populations provide minimal support for the Bruce effect when compared with results from laboratory studies. Nulliparous females may have experienced some pregnancy disruption, but not parous females. Removal of mates, rather than exposure to strange males, may have contributed more to the lower reproductive success of treatment females than exposure to strange males. Treatment females, however, had lower juvenile recruitment than controls, which may have been due to infanticide from strange males. Our results are more similar to those of the field study of the gray-tailed vole than predicted, based on laboratory studies of prairie voles.     

Defensive volatile secretions of two diplopod species attract the carrion ball roller scarab <Emphasis Type="Italic">Canthon morsei</Emphasis> (Coleoptera: Scarabaeidae)

Summary. We report field and laboratory evidence indicating that the defensive compounds secreted by the millipedes Amplinus bitumidus (Diplopoda: Polydesmida) and Anadenobolus putealis (Diplopoda: Spirobolida) attract the carrion scarab Canthon morsei (Scarabaeidae: Scarabaeinae). In a Mexican tropical rain forest, C. morsei was mostly caught in pitfall traps baited with the carcasses of both millipede species. Only a few beetles were attracted to fish carrion. Experimental evidence indicates that C. morsei is attracted to freshly dead millipedes. Benzaldehyde and HCN were identified from Amplinus bitumidus; but 2-methyl-1,4-benzoquinone (toluquinone) and 2-methoxy- 3-methyl-1,4-benzoquinone from Anadenobolus putealis. Benzaldehyde attracted significantly more C. morsei individuals than fish meat, or millipedes of both species did after rinsing in hexane. This is the first report of Polydesmida attracting dung beetles.     

Interspecific attractiveness of structures built by courting male fiddler crabs: experimental evidence of a sensory trap

Male fiddler crabs Uca musica sometimes build sand hoods and male Uca beebei sometimes build mud pillars next to their burrows to which they attract females for mating. Mate-searching females preferentially approach these structures and subsequently mate with structure builders. Here we show that the preference for structures is not species-specific and argue that it may not have evolved for mate choice. When not near burrows, many species of fiddler crabs approach and temporarily hide near objects, suggesting that hoods and pillars may attract females because they elicit this general predator-avoidance behavior. To test this sensory trap hypothesis we individually released female U. musica, U. beebei and Uca stenodactylus, a non-builder, in the center of a circular array of empty burrows to which we added hoods and pillars and then moved a model predator toward the females. All species ran to structures to escape the predator and the two builders preferred hoods. Next, we put hood replicas on male U. beebei burrows and pillar replicas on male U. musica burrows. When courted, females of both species preferentially approached hoods as they did when chased with a predator. However, males of both species with hoods did not have higher mating rates than males with pillars perhaps because hoods block more of a male's visual field so he sees and courts fewer females. Sexual selection may often favor male signals that attract females because they facilitate general orientation or navigation mechanisms that reduce predation risk in many contexts, including during mate search.     

Size and age compositions and reproductive biology of the nervous shark Carcharhinus cautus in a large subtropical embayment, including an analysis of growth during pre- and postnatal life

The lengths-at-age of individuals of the nervous shark Carcharhinus cautus in Shark Bay, Western Australia, have been determined and used to explore the types of situation when it might be advisable to shift from employing a von Bertalanffy equation to a more complex equation for describing the growth of this species and of elasmobranchs in general. The reproductive biology of C. cautus was also examined in order to construct curves for describing growth throughout life from conception as well as from parturition. The presence, in November and early December, of fresh bite marks on the sides of mature females and of a very high proportion of spent individuals among mature males indicate that C. cautus copulates in late October/early November. Ovulation and conception occur in late November/early December and parturition takes place approximately 11 months later. Since mature non-pregnant females contain vitellogenic ova for 12-13 months, i.e. from November or December to the following December, and mature pregnant females contain embryos for 11 months, i.e. from December to October, C. cautus has a biennial reproductive cycle. By parturition, the females and males of C. cautus had reached ~28% and 32% of their lengths at their maximum observed ages, respectively. The maximum recorded total lengths and ages of females and males of C. cautus were 133 cm and 16 years and 111 cm and 12 years, respectively. Females and males reached maturity at ~101 and ~91 cm, respectively, and at least 50% of females and males had become mature by the end of their sixth and fourth years of life after parturition, respectively. The three-parameter, von Bertalanffy growth curves provided reasonably good fits to the lengths-at-age of females and males of C. cautus during just postnatal life and throughout the whole of pre- and postnatal life. While the four-parameter, Schnute growth curve significantly improved the fit to these data for both females and males from conception and for females from parturition, it was recognised that the likelihood ratio test is very sensitive when, as in these cases, there are a large number of data points. A number of interrelated factors were thus taken into account when discussing circumstances when it might be appropriate to switch from using a von Bertalanffy growth curve to the more complex Schnute growth curve.     

The energetic cost of protogynous versus protandrous sex change in the bi-directional sex-changing fish Gobiodon histrio

To investigate the relative cost of protogynous versus protandrous sex change we induced sex change in each direction in Gobiodon histrio (Gobiidae) and then compared growth, body condition and biochemical condition between sex-changed and non-sex-changed fish in treatment and control groups. Sex change in each direction was induced by establishing pairs of adult males and pairs of adult females on isolated coral colonies. Heterosexual pairs were used as controls. For both males and females, growth and body condition did not vary between fish that changed sex and those that did not. The relatively low cost of sex change, compared to the likely costs of searching for a mate of the correct sex, appears to explain the evolution of bi-directional sex change in coral-dwelling gobies. Lipid concentrations in the liver of males and females that changed sex were reduced by similar amounts (34% and 41%, respectively) compared to male and female controls that did not change sex. Therefore, changes in biochemical condition were approximately equal for fish that sex change in each direction and cannot explain the unequal frequency of sex change in each direction in natural populations of G. histrio.     

Abundance, growth and life cycle of the mesopelagic amphipod Primno abyssalis (Hyperiidea: Phrosinidae) in the Oyashio region, western subarctic Pacific

Abundance and life-cycle features of the mesopelagic hyperiid amphipod Primno abyssalis (formerly P. macropa) in the Oyashio region, western subarctic Pacific, were investigated using samples collected between July 1996 and July 1998. P. abyssalis was collected throughout the entire survey period, with abundance peaks occurring in spring to autumn. While all maturity stages of males and females were observed throughout the study period, the peak reproduction season was in summer. Instar analysis based on the segment number of the pleopod rami indicated that hatched juveniles molted 10 times before becoming adult males and 13 times before becoming adult females. Judging from the dry and ash-free dry weights of each instar, males and females continued to feed throughout the final instar stage. Based on cohort analysis of seasonal samples and laboratory observations on molting frequencies, growth in body length of P. abyssalis was linear with time, and estimated generation lengths were 2.3-3.8 years for females and 1.4-1.9 years for males. Brood size of females ranged from 66 to 337 and increased with increasing female body length. Lifetime fecundity, calculated as the sum of six successive broods, was 1,004. Compared with P. abyssalis in the southern Sea of Japan, those in the Oyashio region have a larger number of adult instars (six versus five for females, three 3 vs one for males), a lower growth rate (0.014 mm day^-1 vs 0.021 mm day^-1), and mature earlier (instar 13 vs instar 15 for females; instar 10 vs instar 11 for males). These characteristics are considered to be advantageous life-history traits to counteract higher niche competition within the mesopelagic community and higher predation pressure by mesopelagic fishes in the Oyashio region than in the Sea of Japan.     

Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences

Sexual selection and species recognition play important roles in mate choice; however, sexual selection preferences may overlap with traits found in heterospecifics, producing a conflict between sexual selection and species recognition. We examined female preferences in Xiphophorus pygmaeus for male traits that could provide both types of information to determine how females use multiple cues when preferences for these cues would conflict. We also examined X. pygmaeus behavior in the field to determine if females have the opportunity to choose mates. As no male-male competition was observed in the field, and females occasionally chased males from feeding areas, females apparently have the opportunity to exercise mate choice in their natural habitat. In the laboratory, female X. pygmaeus used body size as a sexual selection cue, preferring large heterospecifics (X. cortezi) to small conspecifics. Females also preferred barless X. cortezi over barred X. cortezi when males were size matched. Because X. pygmaeus males do not have bars, this preference suggests that X. pygmaeus females use vertical bars in species recognition, and that large body size and vertical bars are conflicting cues. However, X. pygmaeus females did not have a preference for males of either species when sexual selection and species recognition cues were presented concurrently. This result was surprising, because preferences for species recognition cues are often assumed to be stronger than sexual selection cues. We suggest that females may be using additional species-specific cues in mate choice to prevent hybridization.     

Competition,fluctuating asymmetry and sperm transfer in male gryllid crickets (<Emphasis Type="Italic">Gryllus bimaculatus</Emphasis> and <Emphasis Type="Italic">Gryllodes sigillatus</Emphasis>)

Sperm competition theory predicts that ejaculate expenditure by males should increase with the risk of competition from rivals. The tendency to modulate expenditure, however, may reflect various constraints acting on different individuals. We test this idea here by looking at the effects of competition and fluctuating asymmetry (FA) in two species of gryllid cricket. Male Gryllus bimaculatus and Gryllodes sigillatus modulated the sperm content of their spermatophores in relation to both apparent competition and FA in limb size. Males of both species increased the amount of sperm transferred when an apparent male competitor was present, sperm number increasing regardless of the species of apparent competitor in G. bimaculatus, but significantly more with a conspecific competitor in G. sigillatus. In addition, the relative size of the spermatophylax in G. sigillatus increased in the presence of apparent competition, while spermatophore transfer time decreased in G. bimaculatus. Sperm number showed opposite relationships with limb asymmetry in the two species, decreasing with increasing asymmetry in males in G. sigillatus, but increasing with asymmetry in both males and females in G. bimaculatus. G. bimaculatus, but not G. sigillatus, also transferred more sperm when paired with a larger female. Relationships between reproductive measures and limb asymmetry were significant only when competitors were present in G. bimaculatus, but were significant regardless of competition in G. sigillatus. The differences may reflect the additional burden of producing a spermatophylax in G. sigillatus.     

Population differences in male responses to chemical mating cues in the white-spotted longicorn beetle, Anoplophora malasiaca

Adult Anoplophora malasiaca beetles were collected from field populations inhabiting three different host plants: Citrus, Salix and Vaccinium. Male orientation responses towards odours of wounded branches of each host plant were examined. Males of all three populations were attracted most frequently to odours of their original hosts, indicating that male orientation behaviour was induced by the odours of the plants from which they were originally collected. We also observed the mating responses of the males to elytra extracts from young and mature females of the three populations and chemically analysed those extracts. Citrus males were highly responsive to female extracts, regardless of origin. Salix males were less reactive than Citrus males, but responded moderately to female extracts of all populations. Vaccinium males were never attracted to Citrus female extracts and responded more to extracts from mature Vaccinium and Salix females, indicating that males might distinguish female origin and maturity. All eight hydrocarbons, four ketones, and three lactones that were previously identified as female contact sex pheromones were found in the extracts of three populations and both ages. Higher amounts of lactones on mature females likely caused greater male responses. These results suggested that males would be more likely to approach a female feeding on the same host plant and subsequently recognise her sexual maturity based on the profile of contact sex pheromones on her elytra.     

Behavioural and electrophysiological responses of the olive fruit fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae), to male- and female-borne sex attractants

In the olive fruit fly, Bactrocera oleae, females attract males by producing 1,7-dioxaspiro[5.5]undecane (olean), the main component of the sex pheromone secreted by rectal glands. It has been recently demonstrated that males are able to produce (Z)-9-tricosene (muscalure) in rectal glands, a compound that selectively attracts females. In this study, a male grooming reaction that may transfer the male-borne compounds from rectal to urotergal glands was observed, suggesting that urotergal glands could be involved in B. oleae sexual communication. GC/MS, EAG, GC/EAD analyses and behavioural assays were carried out to compare the role of male rectal and urotergal glands during courtship. In both male glands, olean and muscalure amounts were age dependent. Extracts of rectal glands contained higher amounts of olean and/or muscalure than urotergal ones. Extracts of rectal and urotergal glands of males and females elicited EAG responses in both sexes. GC/EAD showed that female EAG response to male rectal extracts was mainly due to olean and muscalure. Synthetic compounds evoked EAG dose-dependent responses in both sexes, and the EAG response to muscalure was higher as compared to olean. Rectal and urotergal glands from old males were able to attract females, while urotergal glands from young males attracted only males. Overall, our results add knowledge to the mating system of B. oleae, giving first evidences on the electrophysiological activity of muscalure towards both sexes, as well as on the involvement of male urotergal glands in the chemical sexual communication of this pest.     

The assessment of female reproductive state during courtship and scramble competition in the fiddler crab, Uca paradussumieri

Male fiddler crabs, Uca paradussumieri, mate underground during a 4- to 7-day period each full and new moon. As soon as the tide recedes, males enter the burrows of females that will ovulate the following day ('pre-ovigerous' females). Males copulate with and guard these females until they ovulate. When interrupted by an intruding male, the first male to reach the female is usually able to defend her and successfully mate with her. In fiddler crabs, females mate multiply and there is last male sperm precedence. Before each semi-lunar mating period, male U. paradussumieri were more likely to court females with whom they would later mate than other nearby females with whom they did not mate. This suggests that males collect information on female reproductive state prior to the females becoming ovigerous. In this species, aggression was common between males that courted the same female. When previously courted females were approached by other males, the initial courter attempted to forcefully disrupt the courtship. This behavior may allow males the exclusive use of information on female reproductive condition. It also suggests a type of scramble competition between males over females. Furthermore, it indicates that males are able to locate receptive females prior to their becoming ovigerous. The shorter guarding period observed in this species, as compared with other fiddler crabs, is caused by females rejecting longer guarding periods. Male ability to assess female reproductive status may therefore be advantageous because it increases male mating success within a scramble type of competitive polygyny.     

The use of the tergal gland defensive secretion in male intrasexual aggression of the rove beetle,Aleochara curtula (Coleoptera: Staphylinidae), measured by closed-loop-stripping-analyses and tandem bioassay — mass fragmentography

Summary 2-methyl-1,4-benzoquinone as a target compound of the tergal gland secretion of the rove beetle,Aleochara curtula, was quantified from the reservoirs of individual beetles. Males store less secretion than females, and they evaporate more of the secretion as measured by adsorption from the air (closed-loop-stripping-analysis). The amount of emitted quinone is increased during aggressive interactions of rival males. The pulsed emission of the secretion during aggression is demonstrated by a novel technique combining observation of behavior with the on-line measurement of target fragment ions by mass-spectrometry (tandem bioassay — mass fragmentography). The emission of the secretion is used as a weapon in combats between males and may result in the repulsion of subordinate males from the mating site, but may also serve to mimic females chemically in order to avoid aggressive encounters.     

Influence of host plants on sexual communication in the herbivorous bug Lygocoris pabulinus

Summary. Host plant volatiles may be involved in the sexual communication of insects in several ways. In the pheromone-producing sex, these volatiles may affect pheromone production or release and, in the receptive sex, plant volatiles may have a synergistic effect on the attraction to sex pheromone. We conducted three types of experiments to determine if and how plant volatiles are involved in the sexual communication of Lygocoris pabulinus (L.) (Heteroptera: Miridae), the females of which attract males. In a one-choice cylinder bioassay, females were offered to males on two different plants, belonging to different plant families, i.e. potato leaves and goosefoot leaves, to determine if specific plant volatiles were involved in the attraction of males towards females. Females on potato leaves were as attractive as females on goosefoot leaves, but significantly more attractive than females without plant material. The latter result suggests an interaction between females and potato leaves. However, in two-choice flying and walking bioassays, using delta traps in a wind tunnel and a vertical Y-track olfactometer, males were attracted to females irrespective of the presence of potato leaves. This difference in result is probably due to the fact that in the latter assays females were confined with pollen as an alternative food source, while females in the one-choice assay had access to water only, so that they may have suffered from malnutrition. Males in the one-choice assay were also attracted to potato leaves from which females had been removed, indicating that attractive components from females are deposited and adsorbed to the substrate. Plants are probably only indirectly involved in sexual communication, their surface merely functioning as a substrate from which pheromone is released. Males may subsequently be attracted to such plants or substrates. Clean plant material was not attractive to L. pabulinus males, hence plant volatiles alone do not seem to be used by these males as possible mate location cues.     

Reproductive biology of a small isopod symbiont living on a large isopod host: from the maternal marsupium to the protective grip of guarding males

Mating systems of many symbiotic crustaceans are characterised by a high degree of mate guarding. A peculiar case of mate guarding has been reported for small symbiotic janirid isopods where males mate with immature females. Field samples of individual hosts and laboratory experiments were conducted to reveal the mating behaviour of the symbiont in a natural environment, that is, on their hosts. Along the coast of the Magellan Strait, Chile, the janirid isopod Iais pubescens was frequently found on the shore-living isopod Exosphaeroma gigas. Symbiont prevalence (percent hosts occupied) was high at eight of the nine sampling sites. Mean symbiont intensity was very low at one site (<<1 individual host^-1), intermediate at two sites (1-10 individuals host^-1) and high at the other sites (10-40 individuals host^-1). The mean sex ratio (males:females) was male biased at most sampling sites (n=7). Females of I. pubescens reached substantially larger sizes (1.5-3.0 mm body length, BL) than males (1.1-1.9 mm BL). The majority of males were carrying small juveniles (66.15%), and males with juveniles were significantly larger than males without juveniles - this suggests that males prefer virgin juveniles to adult females and that they compete for small juveniles. In laboratory observations, males were seen to manipulate the marsupium of adult females that were about to release small juveniles. Males obtained virgin juveniles in this manner. Juveniles were carried for ~7 days, and they moulted shortly before being fertilised and released by males. The high proportion of juveniles carried by males in the field (68.2%) supports previous observations that males initially are not able to distinguish male and female juveniles. It is suggested that the mating system of symbiotic janirid isopods with long-term sperm storage and continuous receptivity in females and male mating with virgin females has evolved in response to highly unpredictable encounter probabilities between the sexes. Mate guarding and manipulation of small virgin juveniles may be favoured on the highly mobile hosts of symbiotic janirid isopods. Furthermore, adult females may gain by leaving their emerging offspring in the protective grip of guarding males, thereby reinforcing the maintenance of this peculiar mating system.     

The effects of two acanthocephalan parasites, Pomphorhynchus laevis and Polymorphus minutus, on pairing success in male Gammarus pulex (Crustacea: Amphipoda)

Parasites can decrease male mating success in host species in various ways, in particular by affecting male competitive ability for access to females. However, male-male competition can take different forms (i.e. interference vs exploitation competition) and which type of competition is most affected by parasites is not always clear. We investigated the influence of two acanthocephalan parasites Pomphorhynchus laevis and Polymorphus minutus on the pairing success of their intermediate host, Gammarus pulex, using field-based studies and complementary laboratory-based studies. We first studied male pairing success in the field using four large samples of paired and unpaired individuals collected at different dates. In three of the samples, the effects of size and parasite infection were significant, whereas for one sample only male size had a significant effect. There was no difference in size distributions between infected and uninfected gammarids. Large males were paired more often than smaller males, and uninfected males were paired more frequently than infected males, the pairing success of P. minutus-infected males being more severely affected than that of P. laevis-infected males. We then experimentally tested the ability to enter into precopula with a receptive female in the presence or absence of competitors. In competitive situations, the pairing success of P. laevis- and P. minutus-infected males was significantly lower than that of uninfected males, with pairing success being more affected in P. laevis-infected than in P. minutus-infected males. In the absence of competition, males infected with P. laevis were significantly less likely to enter into precopula compared with uninfected males and P. minutus-infected males, whereas there was no difference between uninfected and P. minutus-infected males in their inclination to pair with a receptive female. However, for both parasites, latency time to pair formation was significantly shorter for uninfected than for infected males. In a third experiment, we tested for a potential effect of vertical segregation on the pairing success of infected and uninfected males, but found no evidence for it. We conclude that infected males may be less competitive than uninfected males in competition by exploitation between males for females.     

The economics of harem maintenance in the sac-winged bat, Saccopteryx bilineata (Emballonuridae)

Saccopteryx bilineata has a polygynous mating system in which males defend females in a harem territory. Harem defense and courtship include energetically costly flight maneuvers and hovering displays. We tested if (1) harem males have a greater field metabolic rate than non-harem males or females and if (2) the field metabolic rate of harem males is correlated with the number of females in a harem territory. We measured the energy budget in 32 S. bilineata with the doubly labeled water method and compared these estimates with behavioral observations in the daytime roost. Among adult bats, field metabolic rate varied with body mass by an exponent of approximately two. We found no significant difference in field metabolic rate or mass-specific field metabolic rate between harem and non-harem males. The mass-specific field metabolic rate of harem-males increased with harem size. The latter finding supports the hypothesis that the energy costs of courtship display and territorial defense influence the energy budget of harem males. Overall, field metabolic rates of S. bilineata were lower than those of similarly sized bats of the temperate zone and only 2.3 times above the basal metabolic rate recorded for this species. We suggest that male S. bilineata did not take advantage of their metabolic capacity because a prudent allocation of energy to activities of harem maintenance is an adaptive strategy for males in this mating system.     

Maturation and diel reproductive periodicity of round scad (Carangidae: Decapterus punctatus)

The diel reproductive periodicity of Carangidae is poorly known but appears to be highly variable between species. Some species spawn during the day, others are believed to spawn at night, and it is demonstrated here that round scad, Decapterus punctatus, spawn at dusk. We collected D. punctatus in the eastern Gulf of Mexico during three April cruises (1995, 1996, and 1997). Based on histological criteria, size at 50% maturity was 113 mm fork length (FL) for males and 128 mm FL for females. The gonad-somatic index (GSI) of mature males was significantly different between hours and appeared to show diel periodicity. Diel periodicity was also observed in changes in female GSIs, whole oocyte diameters, and ovarian histology. The average GSI of mature females fluctuated two-fold between day and night, and the size distribution of whole oocytes in some fish was bimodal (at 0.3-0.4 and 0.7-0.8 mm diameter) at dusk rather than unimodal during most of the diel cycle. Histological preparations revealed that these rapid changes in ovarian GSIs and oocyte size distributions were the result of final oocyte maturation. Germinal vesicle migration was observed from 0900 to 1400 hours eastern standard time (EST), germinal vesicle breakdown was evident as early as 1100 hours EST, and ovulation occurred as early as 1800 hours EST. Spawning frequency (approximately every 5 days) was similar whether calculated from the proportion of females with hydrated oocytes during the afternoon or from the proportion of females with postovulatory follicles during the morning. Batch fecundity correlated with fish size and ranged from 5,500 to 34,700 hydrated eggs per individual. These findings do not support published hypotheses that young-of-the-year D. punctatus reproduce before their first winter or that D. punctatus reproductive output is bimodal within a year.     

A histological study of reproduction in the serpulids<Emphasis Type="Italic"> Pomatoceros triqueter</Emphasis> and<Emphasis Type="Italic"> Pomatoceros lamarckii</Emphasis> (Annelida: Polychaeta)

The reproductive cycles of the serpulid polychaetes Pomatoceros lamarckii (Quatrefages, 1865, as Vermilia) and P. triqueter (Linnaeus, 1767, as Serpula triquetra) were studied by histological examination during 1999 and 2000 at Bantry Bay, south-west Ireland. Gametogenesis, sex ratios and hermaphroditism were also investigated. The male/female ratio in P. lamarckii was approximately equal, but in P. triqueter it deviated significantly in favour of females. There was no significant variation in the sex ratios throughout the year in either species. Results confirm that the species are protandric hermaphrodites. In P. lamarckii, but not in P. triqueter, sexual dimorphism was exhibited in body size, the females being larger than the males of the same species. In both species, the proportion of females increased with increasing body size. Simultaneous hermaphroditism was recorded in P. lamarckii, but not in P. triqueter. There was no clearly defined annual gametogenic cycle in either species. Both species appear to have an extended reproductive season, with numerous small-scale peaks in reproductive maturity that can vary annually. Spawning was broadly synchronous between sexes. Visual observation alone (i.e. without histology) was insufficient to accurately assess reproductive condition in individuals not in, or close to, a ripe state.     

Chemical composition and pheromonal function of the defensive secretions in the subtribe Stizopina (Coleptera, Tenebrionidae, Opatrini)

The chemical composition of the defensive secretions of 52 species from 15 genera of the tenebrionid subtribe Stizopina was analyzed. The secretions of all species contained 1,4-benzoquinones, 1-alkenes, and monoterpene hydrocarbons, only one species was lacking the latter. Methyl- and ethyl-1,4-benzoquinone were ubiquitous, mostly accompanied by smaller amounts of 1,4-benzoquinone as well as isopropyl- and propyl-1,4-benzoquinone. 1-Alkenes were dominated by 1-undecene with varying admixtures of other 1-alkenes. The monoterpene hydrocarbons always consisted of a mixture of α-pinene, camphene, β-pinene and limonene, but also p-cymene, α-terpinene or α-phellandrene were found in some species. Furthermore, some species synthesized additional compounds such as phenols, ketones, 2,5-dihydroxy-6-methylbenzoate, 2-hydroxy-4-methoxyacetophenone and naphthoquinones. Bioassays showed that the defensive secretion co-functioned as an aggregation pheromone in the subtribe Stizopina. All nine tested species from six genera were attracted to defensive secretion of Stizopina species, but they did not distinguish between defensive secretions of different Stizopina species. This lack of discrimination might be the initial step for the formation of interspecific aggregations and the evolution of cleptoparasitism within the Stizopina.     

Effects of size and nuptial gifts on butterfly reproduction: can females compensate for a smaller size through male-derived nutrients?

In butterflies and other insects, fecundity generally increases with female adult weight. Hence, most butterflies are essentially "capital breeders", because nutrients acquired during the larval stage are stored and subsequently used for egg production during the adult stage. However, in some species, males transfer a large nutritious ejaculate to the female at mating. These females can partly be characterized as "income breeders", and female mass can potentially be decoupled from fecundity to some extent. In the gift-giving green-veined white butterfly Pieris napi, it has been shown that female fecundity and longevity increase with number of matings and also that females mature at smaller size under poor food conditions compared to males. So it has been suggested that females can compensate for their smaller size through nuptial feeding. Here we test this hypothesis in P. napi by assessing female fecundity and longevity in relation to female mass and polyandry. The results showed no support for the hypothesis. Smaller females were not capable of increasing their mating rate to compensate for a low weight at eclosion. Instead, larger females remated sooner. Also, smaller females suffered from both a reduced daily and total fecundity compared to larger females and this decrease in fecundity was independent of female mating status, i.e. females allowed to mate only once and multiply mated females suffered to the same extent from their smaller size.