The form of sexual selection arising from male-male competition depends on the presence of females in the social environment

Sexual selection arises from social interactions, and if social environments vary so too should sexual selection. For example, male-male competition often occurs either in the presence or in the absence of females, and such changes in the social environment could affect the form and strength of sexual selection. Here we examine how the presence of a female influences selection arising from male-male competition in a leaf-footed cactus bug, Narnia femorata, which has a resource defence mating system. Males compete for territories on cacti because females lay eggs on the cactus plants. Females are not always present when this competition first occurs; however, the presence or absence of the female matters. We found that both the form and strength of selection on male traits, those traits that influenced success in intrasexual competition, depended on the social context. When a female was not present, male size and the area of the sexually dimorphic hind legs was only marginally important to winning a contest. However, males with larger overall size and leg area were more likely to win in the presence of a female. There was also positive quadratic selection on these traits when a female was present with both the largest and the smallest males winning. The implication is unexpected alternative strategies when females are present. Our results support the notion that sexual selection should be studied under all relevant social contexts.     

Primary homology assessment of structures in the female atrial system among species of the Polycystididae (Rhabditophora, Eukalyptorhynchia)

Abstract. The female atrial system of members of the Polycystididae has been studied at the light microscopical level and compared among the constituent taxa. Based on the criteria of position and conjunction, hypotheses of homology are put forward and compared with the assessments of homology found in earlier literature. Contradictory terminology is synonymized and adapted to fit new findings that recognize homology among structures in different taxa. Based on differences in position and structure, 2 types of female duct are recognized: female duct type I and female duct type II. The term “female bursa” is restricted to a sperm resorbing organ at the proximal end of the female duct type I and/or at the end of a common oviduct. Some species have bundles of glands at the place where the oviduct(s) enter the female duct type I, which are considered homologous among these species. Different types of seminal receptacles are recognized. The term “insemination duct” is restricted to the ducts which in some species, in addition to the oviducts, connect the ovaries with the female duct type I. A single muscular duct, the common oviduct, connects the joined oviducts with the female duct type I in species of Duplacrorhynchus and is considered homologous with the similar duct present in some other species. A uterus is always present, entering the common genital atrium through its frontal wall, or entering a female duct type II. This morphological review of the female atrial system reveals a mosaic distribution of characters if applied to previous phylogenies that are based primarily on the male reproductive system.     

Female sterility in hybrids between Anopheles gambiae and A. arabiensis, and the causes of Haldane's rule

Although F1 female hybrids between Anopheles gambiae and A. arabiensis are fully fertile, sterility is present in backcross females. Here we report the results of a study into the genetic basis of backcross female sterility. Using 23 markers, we performed quantitative trait loci (QTL) mapping analyses to identify chromosomal regions involved in hybrid female sterility. We found that female sterility in backcrosses in both directions is primarily caused by interspecific interactions between a heterozygous X chromosome and recessive autosomal factors. In addition, our data provide support for two theories implicated in Haldane's rule in a single taxon. A comparison with data from a previous study shows that male hybrid sterility QTL are present in higher numbers than female hybrid sterility QTL. Furthermore, autosomal female sterility factors tend to be recessive, supporting the dominance theory for female sterility. Finally, our data indicate a very large effect of the X chromosome from both species on hybrid female sterility, despite the fact that the X chromosome represents less than 9% of the genome. However, this could be the result of a lack of introgression of the X chromosome between A. gambiae and A. arabiensis, rather than a faster evolution of sterility factors on the X chromosome.     

Strategic Investment in Sperm Removal Behaviour in a Bushcricket

Multiple mating by females is widespread and generates sperm competition among the ejaculates of rival males over fertilization. One way in which males can avoid or reduce sperm competition is by displacing or removing previous males’ sperm from female sperm stores. An apparent example of this occurs in the bushcricket Metaplastes ornatus. Males perform a specialised sperm removal behaviour (SRB), using their highly-derived subgenital plate, with which they remove sperm from the female’s spermatheca during the early phases of mating before transferring a spermatophore of their own. Here we investigated whether males strategically invest in SRB according to the amount of previously stored sperm present in females. Each male was tested twice, once with a female containing sperm (‘filled’ condition) and once with a female from whom most previously deposited sperm had recently been removed by another male (‘emptied’ condition). For comparison, a separate group of males was paired with virgin females. Males did not discriminate between non-virgin females in the ‘emptied’ or ‘filled’ conditions in terms of their investment in SRB, suggesting they may not able to perceive the amount of sperm present in the female’s spermatheca. By contrast, male investment in SRB was significantly reduced in pairings with virgin females, indicating that males are sensitive to some aspect of a female’s mating status. Our results thus suggest that males modulate SRB in response to female-mediated cues, possibly chemical cues left by previous males, which would not be present on virgin but would be on non-virgin females.     

Hyperparasitism in Amblyomma rotundatum (Acari: Ixodidae)

In the present study, we report a case of hyperparasitism in Amblyomma rotundatum. During examination of live ticks immediately after collecting them from Boa constrictor snakes held in a reptile facility in Mossoró, RN, northeastern Brazil, 1 unengorged tick female was seen attached to the venter of a partially engorged female. The hypostome and chelicerae of the unengorged female had penetrated the integument of the partially engorged female to the level of the basis capitulli and the palps were splayed outward. To our knowledge, we present the second report of hyperparasitism for the genus Amblyomma.     

Identification of volatile compounds from cheek glands of lesser bandicoot rats and assessment of behavioural response for identified compounds

Check gland secretions were collected from sexually mature male and female rats to identify the volatile compounds using gas chromatography linked mass spectroscopy with a modified head space technique. Twenty one volatile fractions including alkanes, aliphatic acids, esters and alcohols were found. Three compounds viz. 3-octen-1-ol (E) (I), cyclopentane undecanoic acid (II) and 2,4,6,8 tetramethyl-l-undecane (III) were present in very high concentrations in both male and female rats. In addition, the estrus female contained another compound, L-alanine, 1,1-dimethyl ethyl ester. On the basis of odour preference tests the compounds I, II and III were found to attract both male and female of the homotypic species. In contrast, the fourth compound present in the female attracts the opposite sex. The results suggest that the volatile fractions have unique function and the level of attraction varies from compound to compound.     

Sexually dimorphic and isomorphic glomeruli in the antennal lobes of the sphinx moth Manduca sexta

Antennal lobes of adult male and female Manduca sexta were compared in order to investigate the nature and extent of sexual dimorphism of the primary olfactory center of this lepidopteran species. Complete identification of the glomeruli led to the conclusion that all female glomeruli have homologous male counterparts. Thus, there is no sex-specific glomerulus present in one sex and absent in the other. Sexual dimorphism (i.e. glomeruli present but morphologically different in males and females), however, does occur in the three glomeruli composing the male macroglomerular complex. The female homologs of this complex consist of two previously identified 'large female glomeruli' and one newly identified normal-sized glomerulus. The lateral and medial large female glomeruli are interpreted to be homologous to the first two macroglomerular-complex glomeruli-the cumulus and toroid 1. The third male component, the toroid 2, was tentatively identified with a normal-sized spheroidal glomerulus of the female, called here the 'small female glomerulus'. The 60 'ordinary' glomeruli that make up the rest of the glomerular neuropil were found to be homologous in males and females, with the exception of two anomalous (or uncertain) glomeruli. Some variations in relative position and size observed among those glomeruli suggest a diffuse, quantitative kind of sexual dimorphism.     

Resource attractiveness of the male beaugregory damselfish and his decision to court or defend

We examined the decisions territorial male beaugregory damselfish (Stegastes leucostictus) make when presented with mating and aggressive situations simultaneously. Specifically, we testedhow males responded to simultaneous and consecutive presentationsof conspecific males and females in bottles. We hypothesizedthat (1) territorial males would show lower levels of courtshiptoward females when a competing male is also present comparedto when only the female is present and (2) territorial maleswould show higher levels of aggression toward intruding maleswhen a female is also present on their territories than whenonly the intruder is present. We predicted that males withhigh-quality breeding sites would court females and attackcompetitors more vigorously than males with low-quality breedingsites. Males on low-quality breeding sites exhibited fewerconspicuous courtship displays toward bottled females and alower bite rate toward bottled males than did the males onhigh-quality breeding sites. We concluded that poor breeding-sitequality was responsible for the decline in the most conspicuous aspects of both courtship and aggression. When presented simultaneouslywith a bottled male and a bottled female, defenders of bothbreeding-site types drastically reduced the number of courtshipdisplays and the amount of time they spent near the female.Thus, all aspects of courtship were reduced in the presenceof a male intruder. Time spent near the bottled male also decreased in these simultaneous presentations; however, this decreasewas less dramatic than the reduction in time spent near bottledfemales. Habitat quality did not affect these trade-off decisions,but the bite rate exhibited toward the bottled male increasedin the simultaneous presentations compared to the single presentations.Due to this increase in a territorial male's defensive behaviorwhen a female is present, we conclude that the presence of afemale temporarily increased the value of a territory. We suggestthat the possible loss of a territory to an intruding maletakes priority over the potential benefits of successfullycourting a single female and that it is a male's future reproduction,as well as his current reproduction, by which he assesses hisbenefits.     

Artificial vaginocervical stimulation induces a conditioned place preference in female rats

Female rats express a conditioned place preference (CPP) for a context paired with mating. During a mating encounter, the female rat is exposed to several different types of stimuli, including, but not limited to, vaginocervical stimulation and social contact. The present experiment tested the hypothesis that two components of the mating interaction, vaginocervical stimulation or social contact, each induce a CPP in female rats. During conditioning rats received nonpaced mating, artificial vaginocervical stimulation, social interaction or a control treatment. Rats expressed a CPP for the context paired with nonpaced mating or artificial vaginocervical stimulation whereas social interaction and the control treatment did not induce a CPP. The present findings highlight the important role that vaginocervical stimulation plays in the reinforcing effects of mating in female rats.     

An olfactory basis for maternal discrimination of sex of offspring in rats (rattus norvegicus)

Lactating Long-Evans rats (Rattus norvegicus) were presented with three-day-old male and female pups that were either untreated, coated with collodion in the anogenital region or on the neck, or perfumed; or with female pups odorized with either male or female pup urine. As measured by related t-tests of maternal anogenital licking, male and female pups were discriminated in the untreated condition, but not when masking stimuli were present. Female pups treated with male urine elicited more licking than female pups treated with female urine. Overall, maternal licking was decreased by olfactory masking agents and increased by pup urine. It was concluded that olfactory stimuli from pups both stimulate maternal licking and serve as a basis for discriminating male and female offspring.     

Influence of body size on fecundity and sperm management in the parasitoid wasp Anisopteromalus calandrae

A large body size is considered to be advantageous to the reproductive success of females as a result of several factors, such as the allocation of more resources to reproduction and the efficient management of sperm transferred by males. In the present study, the effects of female body size, female mating status and additional food availability on fecundity and the offspring sex ratio are investigated in the parasitoid wasp Anisopteromalus calandrae Howard (Hymenoptera: Pteromalidae). Because of haplodiploid sex determination, females must fertilize eggs to produce female offspring but not to produce male offspring. As predicted, female fecundity and the number of female offspring are positively correlated with body size. However, although the volume of the spermatheca increases with female body size, the amount of sperm stored in the spermatheca is relatively constant, irrespective of body size. Consequently, larger females produce a greater proportion of male offspring, especially at the end of the oviposition sequence, suggesting that larger females that possess more resources for reproduction and produce a larger number of offspring are more likely to suffer sperm depletion. The results of the present study also show that mated females have an increased fecundity compared with virgin females, although the opportunity to feed on honey along with host feeding has no impact upon fecundity or the sex ratio.     

Effects of gonadotropin-releasing hormones, corticotropin-releasing hormone, and vasopressin on female sexual behavior

The effects of intracerebroventricular (icv) infusion of four neuropeptides on female sexual behavior were examined in the female musk shrew (Suncus murinus). In the first experiment, (icv) infusion of 100 ng of the mammalian form of gonadotropin-releasing hormone (mGnRH) facilitated rapid display of receptivity. Gonadotropin-releasing hormone-infused females had shorter latencies to rump present and tail wag, compared with controls. In a second experiment, icv administration of the other form of GnRH present in musk shrew brain, the chicken GnRH-II form, produced no changes in female behavior relative to the control condition. In Experiment 3, icv delivery of corticotropin-releasing hormone (CRH) facilitated female sexual behavior, relative to vasopressin and controls. The females treated with CRH had shorter latencies to display rump present, tail wag, and for the receipt of the first missed intromission compared with females in the other treatment groups. Vasopressin increased female scent marking relative to that of CRH-treated females. These data indicate that neurohormones of the hypothalamic-pituitary-gonadal and the hypothalamic-pituitary-adrenal axes can facilitate reproductive behavior in S. murinus.     

Mating‐induced ovulation in loggerhead sea turtles,Caretta caretta

Mating‐induced ovulation is common in mammals, but has been rarely described in other taxa. Observations of several mature female loggerhead sea turtles, Caretta caretta, held in captivity seemed to indicate that ovulation did not occur in the absence of a male. This study was designed to determine whether this was an effect of captivity or an effect of the absence of a male. Two mature female loggerheads were followed over six annual reproductive cycles. Ultrasound exams were performed approximately every 2 weeks to follow the development of follicles in the ovaries. During the first two seasons, no male was present, in the next two seasons, a mature male was present without mating, and in the final two seasons a mature male was present, mating with one or both females. When no male was present or when present without mating, ovarian follicles developed to full size, but ovulation did not occur and the follicles gradually began to decrease in diameter and undergo changes evident on ultrasound. In the fifth season, only one of the females mated, dropping two eggs after 7 days, and continued to oviposit throughout the following months (total 275 eggs). The unmated female did not ovulate, showing the same pattern as earlier seasons. In the final season, both females mated and ovulated, dropping eggs for the next four months (total 539 eggs). The following year, no males were present and neither female ovulated. This study provides clear evidence that ovulation in loggerhead sea turtles is induced by mating. Zoo Biol 27:213–225, 2008. © 2008 Wiley‐Liss, Inc.     

The development of two flanking SCAR markers linked to a sex determination locus in Salix viminalis L

Most studies of sex determination systems in plants involve dioecious annuals that have known sex chromosomes. Despite the absence of such structures in the majority of dioecious plants, gender seems to be under relatively strict genetic control in some species. Genetic markers linked to a female sex-determination locus in Salix viminalis L. have been discovered through bulked segregant analysis of three full-sib families using approximately 1,000 arbitrary primers. Two RAPD markers that were present in the common female parent as well as in predominantly female progeny of these families were subsequently sequenced and converted to sequence characterized amplified region (SCAR) markers. The two SCAR markers are correlated with gender in the three full-sib families and are present in 96.4% of the female progeny and 2.2% of the males, providing evidence of linkage to a putative female-specific locus associated with gender determination in S. viminalis. Estimates of recombination suggest that the two markers are flanking a putative sex determination locus, SDL-II, in certain families of S. viminalis.     

Pregnancy in a patient with adrenocortical carcinoma during treatment with Mitotane - a case report

We present the case of a female patient with virilising adrenocortical carcinoma treated surgically who conceived during adjuvant treatment with mitotane. We discuss the frequently erroneous routine treatment with oral hormonal contraception without thorough differential diagnosis in female patients with oligo-/amenorrhea and subsequent delay in the proper diagnosis of adrenocortical carcinoma.     

Effects of spatial proximity and alliances on dominance relations among female ring-tailed lemurs (Lemur catta) at Berenty Reserve, Madagascar

In the present study, we describe a change in the dominance rank of the top-ranking female in a wild troop of ring-tailed lemurs (Lemur catta) at Berenty Reserve, Madagascar. After the top-ranking female fell to the bottom-ranking position, she was able to outrank a low-ranking female with the support of her adult daughter or an unrelated high-ranking female. These results indicate that, as in cercopithecine monkeys such as macaques and baboons, close proximity and alliances influence dominance relations among adult females in a wild troop of ring-tailed lemurs.     

Are you Paying Attention? Female Wolf Spiders Increase Dragline Silk Advertisements When Males do not Court

Females of many spider species invest in chemical advertisements to attract males, yet variation in investment relative to the presence or quality of males remains poorly understood. Males of the wolf spider Pardosa milvina court females longer and more intensively when in contact with female silk and also court more intensively when encountering silk from virgin rather than mated females; therefore, females may use silk as a medium to advertise their receptivity to mate. We estimated female investment in advertisements by measuring variation in the quantity and type of silk deposited by females in the presence or absence of males and among males that varied in their courtship intensity. We measured dragline silk, cord silk, and attachment disk deposition from females on gridded sheets of paper in response to four stimuli over a 30‐min period (n = 39/treatment): (1) an intensively (high) courting male with access to female silk, (2) a weakly courting or non‐courting male (low) without access to female silk, (3) no male present, but female silk present (silk control), and (4) no stimulus present (control). Females produced significantly more dragline silk and significantly less cord silk in the presence of low‐courting males compared to any other treatment, but we found no difference in attachment disk deposition across treatments. Our results suggest that females invest more heavily in dragline deposition when encountering low‐courting males. Additional studies are necessary to determine the relative role of different silk types in male–female sexual communication.     

Sugars in the gut of the sandfly Phlebotomus orientalis from Dinder National Park, Eastern Sudan

The sandfly Phlebotomus orientalis Parrot (Diptera: Psychodidae) is the vector of visceral leishmaniasis in eastern and Upper Nile regions of Sudan, where vector infection rates of over 7% have been reported. Sugars are known to be important for development of the parasite and for increasing the survival and oviposition rates of several species of sandflies. In the present study we have analysed the sugars present in the guts of individuals and groups of male and female P. orientalis and compared these with sugars from several potential local plant sources: Acacia seyal, Balanites aegyptiaca and Combretum kordofanum. The distribution of these trees in Sudan is closely correlated with that of P. orientalis. Only 20% of individually analysed female sandflies had significant amounts of sugars present suggesting that P. orientalis either digest their sugar meal quickly or do not require regular sugar meals. Interestingly, the sugars present in the males were significantly different to those found in the females, indicating that they had fed on different sugar sources. There was evidence that fruit sugars from Balanites aegyptiaca, Combretum kordofanum and aphid or coccid honeydew are utilized by male and female P. orientalis. There was evidence to indicate that female P. orientalis feeds directly on honeydew. There was no evidence to indicate that direct feeding on leaves is a typical source for the sugar meal. There was no melizitose and only a very small amount of turanose present in the male, suggesting that honeydew was not an important sugar source for males.     

Morphometric comparison of the midgut epithelial cells in male and female Aedes aegypti L. (Insecta, Diptera).

Midgut epithelial cells of male and female Aedes aegypti, 3 days after emergence, were compared morphometrically. The results of the present investigation concerning the female, are in good agreement with those of a previous study (Hecker et al., 1974), demonstrating that morphometric investigation of midgut epithelia in A. aegypti can successfully be reproduced, and that the mosquito strain used did not show quantitative morphological changes due to laboratory rearing. In males, the cells of the anterior (A) and posterior part (P, 'stomach') of the midgut differ in their quantitative composition. Higher values are found for the microvilli and for the basal labyrinth in the A-part. On the other hand a higher volume density of the mitochondria is present in the P-part. No significant differences are found in the A-part between males and females. Significant differences, however, are present in the P-part. Distinctly more rer in the female stomach can be correlated with the synthesis of enzymes for blood digestion, which are absent in the male. In addition, the more complex functions of the female P-part are also reflected by higher values for other organelles and membrane systems (e.g. mitochondria, basal labyrinth).     

Who's Watching Me: Female Siamese Fighting Fish Alter Their Interactions in Response to an Audience

It is well established that interactions between conspecifics are often influenced by the presence of passive bystanders. Individuals have been found to alter their behavior in a variety of contexts, from foraging to aggression, based on the presence, sex, or identity of an audience. This audience effect may influence not only the nature of a signaling event but also the evolution of signal structure as signals may have to convey information across a distance. Additionally, audience individuals may use information obtained by watching in later encounters with these individuals, which may act as a selection pressure on communication. Communication networks in Siamese fighting fish, Betta splendens, are particularly well studied, with audience effects influencing both male–male interactions and male–female interactions. However, the effects of an audience on female–female interactions have not been examined in this species or any other. This study examined the interactions of pairs of females in three different audience conditions (male, female, and no audience). The results suggest that female–female interactions are affected by the presence of an audience as interactant‐directed gill flaring, the most commonly performed behavior, was performed more with an audience present. Additionally, the sex of the audience seemed to be influential, reflected by a difference in the frequency of interactant‐directed behaviors when a female vs. a male audience was present. This study is one of the first to demonstrate that females modify their behavior as a result of being watched and stresses the importance of examining audience effects in a variety of social contexts.