Males, but not females, mate with multiple partners: a laboratory study of a primitively eusocial wasp Ropalidia marginata

The intense interest in social Hymenoptera, on account of their elaborate sociality and the paradox of altruism, has often suffered from considerable gender imbalance. This is partly due to the fact that worker behaviour and altruism are restricted to the females and partly because males often live off the nest. Yet, understanding the males, especially in the context of mating biology is essential even for understanding the evolution of sociality. Mating patterns have a direct bearing on the levels of intra-colony genetic relatedness, which in turn, along with the associated costs and benefits of worker behaviour, are central to our understanding of the evolution of sociality. Although mating takes place away from the nest in natural colonies of the primitively eusocial wasp Ropalidia marginata, mating can be observed in the laboratory if a male and a female are placed in a transparent, aerated plastic container, and both wasps are in the range of 5-20 days of age. Here, we use this setup and show that males, but not females, mate serially with multiple partners. The multiple mating behaviour of the males is not surprising because in nature males have to mate with a number of females, only a few of whom will go on to lay eggs. The reluctance of R. marginata females to mate with multiple partners is consistent with the expectation of monogamy in primitively eusocial species with totipotent females, although the apparent discrepancy with a previous work with allozyme markers in natural colonies suggesting that females may sometimes mate with two or three different males remains to be resolved.

Females of a tree cricket prefer larger males but not the lower frequency male calls that indicate large body size

In the tree cricket Oecanthus henryi, females are attracted by male calls and can choose between males. To make a case for female choice based on male calls, it is necessary to examine male call variation in the field and identify repeatable call features that are reliable indicators of male size or symmetry. Female preference for these reliable call features and the underlying assumption behind this choice, female preference for larger males, also need to be examined. We found that females did prefer larger males during mating, as revealed by the longer mating durations and longer spermatophore retention times. We then examined the correlation between acoustic and morphological features and the repeatability of male calls in the field across two temporal scales, within and across nights. We found that carrier frequency was a reliable indicator of male size, with larger males calling at lower frequencies at a given temperature. Simultaneous playback of male calls differing in frequency, spanning the entire range of natural variation at a given temperature, revealed a lack of female preference for low carrier frequencies. The contrasting results between the phonotaxis and mating experiments may be because females are incapable of discriminating small differences in frequency or because the change in call carrier frequency with temperature renders this cue unreliable in tree crickets. (C) 2012 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Virgin wasps develop ovaries on par with mated females, but lay fewer eggs

In the primitively eusocial wasp Ropalidia marginata, mating is not necessary for a female wasp to develop her ovaries, lay eggs, and even to become the sole egg layer of her colony despite the presence of other mated nestmates. Here, we show that virgin wasps do not differ from their mated counterparts in the extent and rapidity of their ovarian development, in the proportion of individuals that build a nest and laid eggs, and in the time taken to do so. However, a significantly larger proportion of virgin females showed resorbing oocytes, and laid fewer eggs as compared to mated individuals. Thus, virgin females have the ability to develop ovaries and lay eggs but also to refrain from necessarily laying all mature eggs produced, before mating opportunities arise. This dual ability would be adaptive in haplodiploid, tropical species with perennial nesting cycles and frequent opportunities for workers to become replacement queens or solitary nest foundresses throughout the year.

Rhythmic egg-laying behaviour in virgin females of fruit flies Drosophila melanogaster

Fruit fly Drosophila melanogaster females display rhythmic egg-laying under 12: 12 h light/dark (LD) cycles which persists with near 24 h periodicity under constant darkness (DD). We have shown previously that persistence of this rhythm does not require the neurons expressing pigment dispersing factor (PDF), thought to be the canonical circadian pacemakers, and proposed that it could be controlled by peripheral clocks or regulated/triggered by the act of mating. We assayed egg-laying behaviour of wild-type Canton S (CS) females under LD, DD and constant light (LL) conditions in three different physiological states; as virgins, as females allowed to mate with males for 1 day and as females allowed to mate for the entire duration of the assay. Here, we report the presence of a circadian rhythm in egg-laying in virgin D. melanogaster females. We also found that egg-laying behaviour of 70 and 90% females from all the three male presence/absence protocols follows circadian rhythmicity under DD and LL, with periods ranging between 18 and 30 h. The egg-laying rhythm of all virgin females synchronized to LD cycles with a peak occurring soon after lights-off. The rhythm in virgins was remarkably robust with maximum number of eggs deposited immediately after lights-off in contrast to mated females which show higher egg-laying during the day. These results suggest that the egg-laying rhythm of D. melanogaster is endogenously driven and is neither regulated nor triggered by the act of mating; instead, the presence of males results in reduction in entrainment to LD cycles.

Finding hidden females in a crowd: Mate recognition in fig wasps

Multi-species mating aggregations are crowded environments within which mate recognition must occur. Mating aggregations of fig wasps can consist of thousands of individuals of many species that attain sexual maturity simultaneously and mate in the same microenvironment, i.e, in syntopy, within the close confines of an enclosed globular inflorescence called a syconium - a system that has many signalling constraints such as darkness and crowding. All wasps develop within individual galled flowers. Since mating mostly occurs when females are still confined within their galls,, male wasps have the additional burden of detecting conspecific females that are ``hidden'' behind barriers consisting of gall walls. In Ficus racemosa, we investigated signals used by pollinating fig wasp males to differentiate conspecific females from females of other syntopic fig wasp species. Male Ceratosolen fusciceps could detect conspecific females using cues from galls containing females, empty galls, as well as cues from gall volatiles and gall surface hydrocarbons. In many figs, syconia are pollinated by single foundress wasps, leading to high levels of wasp inbreeding due to sibmating. In F. racemosa, as most syconia contain many foundresses, we expected male pollinators to prefer non-sib females to female siblings to reduce inbreeding. We used galls containing females from non-natal figs as a proxy for non-sibs and those from natal figs as a proxy for sibling females. We found that males preferred galls of female pollinators from natal figs. However, males were undecided when given a choice between galls containing non-pollinator females from natal syconia and pollinator females from non-natal syconia, suggesting olfactory imprinting by the natal syconial environment. (C) 2013 Elsevier Masson SAS. All rights reserved.

Ovarian development in a primitively eusocial wasp: Social interactions affect behaviorally dominant and subordinate wasps in opposite directions relative to solitary females

In many primitively eusocial wasp species new nests are founded either by a single female or by a small group of females. In the single foundress nests, the lone female develops her ovaries, lays eggs as well as tends her brood. In multiple foundress nests social interactions, especially dominance-subordinate interactions, result in only one `dominant' female developing her ovaries and laying eggs. Ovaries of the remaining `subordinate' cofoundresses remain suppressed and these individuals function as workers and tend the dominant's brood. Using the tropical, primitively eusocial polistine wasp Ropalidia marginata and by comparing wasps held in isolation and those kept as pairs in the laboratory, we demonstrate that social interactions affect ovarian development of dominant and subordinate wasps among the pairs in opposite directions, suppressing the ovaries of the subordinate member of the pair below that of solitary wasps and boosting the ovaries of dominant member of the pair above that of solitary females. In addition to being of physiological interest, such mirror image effects of aggression on the ovaries of the aggressors and their victims, suggest yet another mechanism by which subordinates can enhance their indirect fitness and facilitate the evolution of worker behavior by kin selection. (C) 2014 Elsevier B.V. All rights reserved.

The interaction of N-trifluoroacetylgalactosamine and its derivatives with winged bean (Psophocarpus tetragonolobus) basic agglutinin reveals differential mechanism of their recognition: a fluorine-19 nuclear magnetic resonance study

Here, we show the binding results of a leguminosae lectin, winged bean basic agglutinin (WBA I) to N-trifluoroacetylgalactosamine (NTFAGalN), methyl-alpha-N-trifluoroacetylgalactosamine (Me alpha NTFAGalN) and methyl-beta-tifluoroacetylgalactosamine (Me beta NTFAGalN) using (19) F NMR spectroscopy. No chemical shift difference between the free and bound states for NTFAGalN and Me beta NTFAGalN, and 0.01-ppm chemical shift change for Me alpha NTFAGalN, demonstrate that the Me alpha NTFAGalN has a sufficiently long residence time on the protein binding site as compared to Me beta NTFAGalN and the free anomers of NTFAGalN. The sugar anomers were found in slow exchange with the binding site of agglutinin. Consequently, we obtained their binding parameters to the protein using line shape analyses. Aforementioned analyses of the activation parameters for the interactions of these saccharides indicate that the binding of alpha and beta anomers of NTFAGalN and Me alpha NTFAGalN is controlled enthalpically, while that of Me beta NTFAGalN is controlled entropically. This asserts the sterically constrained nature of the interaction of the Me beta NTFAGalN with WBA I. These studies thus highlight a significant role of the conformation of the monosaccharide ligands for their recognition by WBA I.

Silent katydid females are at higher risk of bat predation than acoustically signalling katydid males

Males that produce conspicuous mate attraction signals are often at high risk of predation from eavesdropping predators. Females of such species typically search for signalling males and their higher motility may also place them at risk. The relative predation risk faced by males and females in the context of mate-finding using long-distance signals has rarely been investigated. In this study, we show, using a combination of diet analysis and behavioural experiments, that katydid females, who do not produce acoustic signals, are at higher risk of predation from a major bat predator, Megaderma spasma, than calling males. Female katydids were represented in much higher numbers than males in the culled remains beneath roosts of M. spasma. Playback experiments using katydid calls revealed that male calls were approached in only about one-third of the trials overall, whereas tethered, flying katydids were always approached and attacked. Our results question the idea that necessary costs of mate-finding, including risk of predation, are higher in signalling males than in searching females.

Males and females of the social wasp Ropalidia marginata do not differ in their cuticular hydrocarbon profiles and do not seem to use any long-distance volatile mate attraction cues

Sex pheromones are vital in communication between individuals belonging to opposite sexes and form an integral part of the reproductive biology of various species. Among insects, sexual dimorphism in CHCs has been reported from diverse taxa spanning seven different orders, and thereby CHCs have been implicated as sex pheromones. Because males and females of the primitively eusocial wasp Ropalidia marginata touch each other with their antennae during mating, before engaging in sperm transfer, a sex pheromone that is perceived via contact chemosensation through the antennae can possibly exist in this species. Since CHCs have been implied as sex pheromones in various insects (including hymenopterans), and since sexual dimorphism of CHCs should be an obligatory prerequisite for them to act as sex pheromones, we investigated whether males and females of R. marginata differ in their CHC profiles. We found only nonvolatile CHCs, and our results show absence of sexual dimorphism in CHCs, suggesting that CHCs do not function as sex pheromone in this species. A behavioral assay failed to show presence of mate attraction at a distance, thereby showing the absence of volatile long-distance mate attraction cues (that may originate from sources other than and in addition to CHCs).

On the methods of determining size/weight characteristics of food organisms of fish. (Determination of the length of Copepoda in larval stages by egg-laying females of the free living freshwater Copepoda). [Translation from: Voprosy Ikhtiologii 14, 182-194, 1960. ]

As is known, copepods play an important role in the nutrition of fish. Therefore with a view to facilitating research on the study of the quantitative side of feeding, there have recently appeared a considerable number of papers devoted to the development of methods for determining the wet. weight of these crustaceans. For the further facilitating of research in the nutrition of fish it would be of great interest to clarify the problem, is there not some kind of rule in the growth of the crustaceans during metamorphosis, and if there is such a rule is it not possible, to determine the length of the larvae at each stage, not by measuring them, but by using the formulae derived on the basis of these rules. This article examines the growth curves of different species of freshwater Copepoda, obtained on the basis of experimental observations in cultures or by way of measurement of mass material at all stages of development in samples from water-bodies. The authors study in particular the ratio of the mean diameter of the eggs to the mean length of the egg-bearing females.

Changes in the organism of females of some Cladocera in transition to gamogenesis. [Translation from: Zoologicheskie Zhurnal 49 1573-1575, 1970. ]

In species of Cladocera not forming ephippia, the latent eggs have a sheath formed by glands of the reproductive canals. Representatives of the families Daphniidae and Moinidae Goulden, 1968, in connection with the formation of their complex-structured ephippia, lost these glands. It was investigated whether there are such glands in species the latent eggs of which are enclosed in primitive ephippia. For this, with the help of histological methods, 55 females of Acroperus elongatus (Sars) (Chydoridea) and 88 females of Lathonura rectirostris (O. F. Muller), (Macrothricidae) were collected near Leningrad, were studied.

Action of ecdysterone on the moulting of amphipod females: Gammarus pulex (L.) and G.fossarum Koch. Early results. [Translation from: Crustaceana 28, 86-88, 1975.]

In studying sexual attraction in gammarids of the group pulex, it has seemed necessary to dissociate the processes of moulting and ovogenesis in order to recognize their respective effects on this phenomenon. For this purpose a synthetic hormone, ecdysterone, was utilized. In the first instance the author followed the action of the hormone on isolated females in vitellogenesis. It was proved that the behaviour of Gammarus pulex and Gammarus fossarum vis-a-vis the ecdysterone used proves to be very close to that of isopods that was observed in Orchestia gammarellus in earlier research. Although they were in vitellogenesis, the females saw their intermoult cycle shortened.

Identification of Sex and Female’s Reproductive Stage in Commercial Fish Species through the Quantification of Ribosomal Transcripts in Gonads

The estimation of maturity and sex of fish stocks in European waters is a requirement of the EU Data Collection Framework as part of the policy to improve fisheries management. On the other hand, research on fish biology is increasingly focused in molecular approaches, researchers needing correct identification of fish sex and reproductive stage without necessarily having in house the histological know-how necessary for the task. Taking advantage of the differential gene transcription occurring during fish sex differentiation and gametogenesis, the utility of 5S ribosomal RNA (5S rRNA) and General transcription factor IIIA (gtf3a) in the molecular identification of sex and gametogenic stage was tested in different economically-relevant fish species from the Bay of Biscay. Gonads of 9 fish species (, Atlantic, Atlantic-chub and horse mackerel, blue whiting, bogue, European anchovy, hake and pilchard and megrim), collected from local commercial fishing vessels were histologically sexed and 5S and 18S rRNA concentrations were quantified by capillary electrophoresis to calculate a 5S/18S rRNA index. Degenerate primers permitted cloning and sequencing of gtf3a fragments in 7 of the studied species. 5S rRNA and gtf3a transcript levels, together with 5S/18S rRNA index, distinguished clearly ovaries from testis in all of the studied species. The values were always higher in females than in males. 5S/18S rRNA index values in females were always highest when fish were captured in early phases of ovary development whilst, in later vitellogenic stages, the values decreased significantly. In megrim and European anchovy, where gonads in different oogenesis stages were obtained, the 5S/18S rRNA index identified clearly gametogenic stage. This approach, to the sexing and the quantitative non-subjective identification of the maturity stage of female fish, could have multiple applications in the study of fish stock dynamics, fish reproduction and fecundity and fish biology in general.