Differential allocation in a gift-giving spider: males adjust their reproductive investment in response to female condition.

BACKGROUND: When males are selective, they can either reject low-quality females or adjust their reproductive investment in response to traits that indicate female quality (e.g., body size or condition). According to the differential allocation hypothesis, males increase their reproductive investment when paired with high-quality females (positive differential allocation) or increase their reproductive investment when paired with low-quality females (negative differential allocation). This hypothesis has been proposed for monogamous species with biparental care, and most empirical studies focus on birds. Here we used the polygamous spider Paratrechalea ornata, in which males offer prey wrapped in silk as nuptial gifts, to test whether males adjust their reproductive investment in gift size, pre-copulatory and copulatory courtship, and sperm transfer in response to female body condition. RESULTS: Males exposed to females in good body condition added more flies to the gift, stimulated these females longer with abdominal touches during pre-copulatory courtship, and had longer pedipalp insertions than males exposed to females in poor body condition. Female condition affected neither silk investment in nuptial gift wrapping nor the quantity of sperm transferred by males. Finally, females in good body condition oviposited faster after copulation and laid more eggs than females in poor body condition. CONCLUSIONS: We provide experimental evidence that males of a gift-giving spider exhibit positive differential allocation in three key aspects of their reproductive investment: the size of the nutritious gift, duration of pre-copulatory courtship, and duration of pedipalp insertions, which is regarded as a form of copulatory courtship in spiders. This positive differential allocation is likely associated with the benefits of copulating with females in good body condition. These females are more fecund and oviposit faster after copulation than females in poor body condition, which under natural field conditions probably reduces the risk of multiple matings and thus the level of sperm competition faced by the males. As a final remark, our findings indicate that the hypothesis of differential allocation also applies to species with a scramble competition mating system, in which males heavily invest in nuptial gift construction, but not in parental care.

Dinner and a show: The role of male copulatory courtship song and female blood-feeding in the reproductive success of Lutzomyia longipalpis from Lapinha, Brazil.

Lutzomyia longipalpis is the main vector of visceral Leishmaniasis in the Americas and is composed of a species complex. Males of this sand-fly produce acoustic signals during copulation and different patterns are observed among Brazilian populations. Such acoustic signals are commonly involved in species recognition. However, since the song is only produced during copulation it is not clear how it affects mating success or contributes to sexual isolation. Another aspect that may affect reproductive success is the presence of food. Since hematophagy is such an important aspect of L. longipalpis biology, we wanted to test if blood-feeding can influence the reproductive behaviour of this insect. We performed crossing experiments removing males' wings (silencing them) and playing back either the homo-specific or the hetero-specific song to either unfed or blood-fed females. Our results showed that both songs and blood-feeding affect insemination success, but not the frequency of copulation. In trials where females were not blood-fed song clearly affected insemination; males with wings, and males with homo-specific song playback had a higher insemination success than wingless males (no song) and trials with hetero-specific song. Blood-feeding females prior to the trials increased insemination in all groups including the control group which suggests that mating happens simultaneously with, or immediately after, the blood meal. Blood-fed females also seemed to discriminate less against the wrong song or the lack of song (wingless) one day after feeding, however trials with the correct song still had higher insemination rates. Altogether, our results show that both the male copulatory courtship songs and female blood-feeding are important for reproductive success and as such are important components of the sexual behaviour of L. longipalpis.

Sexual selection on bushcricket genitalia operates in a mosaic pattern.

In most species with internal fertilization, male genitalia evolve faster than other morphological structures. This holds true for genital titillators, which are used exclusively during mating in several bushcricket subfamilies. Several theories have been proposed for the sexual selection forces driving the evolution of internal genitalia, especially sperm competition, sexually antagonistic coevolution (SAC), and cryptic female choice (CFC). However, it is unclear whether the evolution of genitalia can be described with a single hypothesis or a combination of them. The study of species-specific genitalia action could contribute to the controversial debate about the underlying selective evolutionary forces. We studied female mating behaviors in response to experimentally modified titillators in a phylogenetically nested set of four bushcricket species: Roeseliana roeselii, Pholidoptera littoralis littoralis, Tettigonia viridissima (of the subfamily Tettigoniinae), and Letana inflata (Phaneropterinae). Bushcricket titillators have several potential functions; they stimulate females and suppress female resistance, ensure proper ampulla or spermatophore attachment, and facilitate male fixation. In R. roeselii, titillators stimulate females to accept copulations, supporting sexual selection by CFC. Conversely, titillator modification had no observable effect on the female's behavior in T. viridissima. The titillators of Ph. l. littoralis mechanically support the mating position and the spermatophore transfer, pointing to sexual selection by SAC. Mixed support was found in L. inflata, where manipulation resulted in increased female resistance (evidence for CFC) and mating failures by reduced spermatophore transfer success (evidence for SAC). Sexual selection is highly species-specific with a mosaic support for either cryptic female choice or sexually antagonistic coevolution or a combination of both in the four species.

Demonstrating sexual selection by cryptic female choice on male genitalia: What is enough?

Rapid divergence in external genital structures occurs in nearly all animal groups that practice internal insemination; explaining this pattern is a major challenge in evolutionary biology. The hypothesis that species-specific differences in male genitalia evolved under sexual selection as courtship devices to influence cryptic female choice (CFC) has been slow to be accepted. Doubts may stem from its radical departure from previous ideas, observational difficulties because crucial events occur hidden within the female's body, and alternative hypotheses involving biologically important phenomena such as speciation, sperm competition, and male-female conflicts of interest. We assess the current status of the CFC hypothesis by reviewing data from two groups in which crucial predictions have been especially well-tested, Glossina tsetse flies and Roeseliana (formerly Metrioptera) roeselii bushcrickets. Eighteen CFC predictions have been confirmed in Glossina and 19 in Roeseliana. We found data justifying rejection of alternative hypotheses, but none that contradicted CFC predictions. The number and extent of tests confirming predictions of the CFC hypothesis in these species is greater than that for other generally accepted hypotheses regarding the functions of nongenital structures. By this criterion, it is reasonable to conclude that some genital structures in both groups likely involved sexual selection by CFC.

Genitalic stridulation during copulation in a species of crane fly, Tipula (Bellardina) sp. (Diptera: Tipulidae) / Estridulación genital durante la cópula en una especie de mosca grulla, Tipula (Bellardina) sp. (Dípteros: Tipulidae)

The male genitalia of many animal groups have elaborate and species-specific forms. One hypothesis  to explain why this is so is that male genitalia function as stimulatory devices that are under sexual selection by cryptic female choice. This report is based on a videotaped observation of a single male of an unidentified species of Tipula (Bellarina) from San José Province, Costa Rica. A male crane fly stridulated (produced vibrations) during copulation using sustained and stereotyped movements of file and scraper structures on his genitalia. Males of related species have similar file and scraper structures on their genitalia, suggesting that they probably also stridulate during copulation. Rev. Biol. Trop. 57 (Suppl. 1): 251-256. Epub 2009 November 30.

Un macho de Tipula (Bellardina) sp. estriduló (produjo vibraciones) por medio de movimientos estereotipados de estructuras de sus órganos genitales durante varios minutos de la cópula. Los machos de algunas especies relacionadas poseen estructuras similares, lo cual sugiere que también estridulan durante la cópula.

Why do male Xylographus contractus Mellié (Coleoptera: Ciidae) present abdominal fovea? Evidence of sexual pheromone secretion / Por que machos de Xylographus contractus Mellié (Coleoptera: Ciidae) apresentam fóvea abdominal? Evidências de secreção de feromônio sexual

Most male ciids present a fovea in the first urosternite. We tested two alternative hypotheses for the function of male abdominal fovea in Xylographus contractus Mellié: (i) the fovea secretes sexual pheromone, and (ii) the fovea secretes aggregation pheromone. For this, 59 specimens were submitted, separately, to two stimuli (fovea extracts and control) in an y-shaped glass olfactometer. For the analyses, we considered the 47 individuals that moved toward one of the longer olfactometer branches. The probability of moving toward the fovea extract was greater for females than for males (chi2 = 3.94, P = 0.047). The proportion of males that moved toward any of the olfactometer branches was the same (chi2 = 0.29, P = 0.59). We concluded that the male's abdominal fovea secretes a sexual pheromone and that there is no aggregation pheromone being secreted by this structure. We suggested that this sexual pheromone is used for short-range communication, acting in pre-copulatory courtship.

A maioria dos machos de ciídeos possuem uma fóvea no primeiro urosternito. Foram testadas duas hipóteses alternativas para explicar a função da fóvea abdominal em machos de Xylographus contractus Mellié: (i) a fóvea secreta feromônio sexual, e (ii) a fóvea secreta feromônio de agregação. Para isso, avaliou-se a resposta de 59 indivíduos, separadamente, a dois estímulos olfativos (extrato da fóvea e controle) em olfatômetro tipo Y. Nas análises, consideraram-se as respostas dos 47 indivíduos que se dirigiram a um dos braços do olfatômetro. A probabilidade de se deslocar em direção ao extrato de fóvea foi maior para fêmeas do que para machos (chi2 = 3,94, P = 0,047). A proporção de machos que se dirigiram para cada um dos braços do olfatômetro foi a mesma (chi2 = 0,29, P = 0,59). A fóvea abdominal dos machos secreta feromônio sexual, não sendo secretado nenhum feromônio de agregação por essa mesma estrutura. Sugere-se que o feromônio sexual seja usado em comunicação a curta distância, mediando mecanismos pré-copulatórios.

COPULATORY BEHAVIOR, GENITAL MORPHOLOGY, AND MALE FERTILIZATION SUCCESS IN WATER STRIDERS.

Recent theoretical and empirical interest in postmating processes have generated a need for increasing our understanding of the sources of variance in fertilization success among males. Of particular importance is whether such postmating sexual selection merely reinforces the effects of premating sexual selection or whether other types of male traits are involved. In the current study, we document large intraspecific variation in last male sperm precedence in the water strider Gerris lateralis. Male relative paternity success was repeatable across replicate females, showing that males differ consistently in their ability to achieve fertilizations. By analyzing shape variation in male genital morphology, we were able to demonstrate that the shape of male intromittent genitalia was related to relative paternity success. This is the first direct experimental support for the suggestion that male genitalia evolve by postmating sexual selection. A detailed analysis revealed that different components of male genitalia had different effects, some affecting male ability to achieve sperm precedence and others affecting male ability to avoid sperm precedence by subsequent males. Further, the effects of the shape of the male genitalia on paternity success was in part dependent on female morphology, suggesting that selection on male genitalia will depend on the frequency distribution of female phenotypes. We failed to find any effects of other morphological traits, such as male body size or the degree of asymmetry in leg length, on fertilization success. Although males differed consistently in their copulatory behavior, copulation duration was the only behavioral trait that had any significant effect on paternity.