Eviction-driven infanticide and sexually selected adoption and infanticide in a neotropical parrot.

Infanticide and adoption have been attributed to sexual selection, where an individual later reproduces with the parent whose offspring it killed or adopted. While sexually selected infanticide is well known, evidence for sexually selected adoption is anecdotal. We report on both behaviors at 346 nests over 27 y in green-rumped parrotlets (Forpus passerinus) in Venezuela. Parrotlets are monogamous with long-term pair bonds, exhibit a strongly male-biased adult sex ratio, and nest in cavities that are in short supply, creating intense competition for nest sites and mates. Infanticide attacks occurred at 256 nests in two distinct contexts: 1) Attacks were primarily committed by nonbreeding pairs (69%) attempting to evict parents from the cavity. Infanticide attacks per nest were positively correlated with population size and evicting pairs never adopted abandoned offspring. Competition for limited nest sites was a primary cause of eviction-driven infanticide, and 2) attacks occurred less frequently at nests where one mate died (31%), was perpetrated primarily by stepparents of both sexes, and was independent of population size. Thus, within a single species and mating system, infanticide occurred in multiple contexts due to multiple drivers. Nevertheless, 48% of stepparents of both sexes adopted offspring, and another 23% of stepfathers exhibited both infanticide and long-term care. Stepfathers were often young males who subsequently nested with widows, reaching earlier ages of first breeding than competitors and demonstrating sexually selected adoption. Adoption and infanticide conferred similar fitness benefits to stepfathers and appeared to be equivalent strategies driven by limited breeding opportunities, male-biased sex ratios, and long-term monogamy.

Neural mechanisms involved in female mate choice in invertebrates.

Mate choice is a critical decision with direct implications for fitness. Although it has been recognized for over 150 years, our understanding of its underlying mechanisms is still limited. Most studies on mate choice focus on the evolutionary causes of behavior, with less attention given to the physiological and molecular mechanisms involved. This is especially true for invertebrates, where research on mate choice has largely focused on male behavior. This review summarizes the current state of knowledge on the neural, molecular and neurohormonal mechanisms of female choice in invertebrates, including behaviors before, during, and after copulation. We identify areas of research that have not been extensively explored in invertebrates, suggesting potential directions for future investigation. We hope that this review will stimulate further research in this area.

My path to primatology: some stories from the field.

In this paper, I summarize the major facets of my 50-year career as a primatologist. I briefly describe the aspects of my upbringing and early education that led me to the study of primate behavior, first in captive settings and, later, in the wild. My research on the Arashiyama West Japanese macaques and my interactions with Japanese primatologists was a formative stage in my career, and I present the background of this international project and how it led to my growing focus on female life history studies. After a couple of failed attempts to establish a long-term study of primates in their native habitats, I began the Santa Rosa Primate Project in Costa Rica in 1983, which focuses mainly on white-faced capuchins, and to some extent on howlers and spider monkeys. The Santa Rosa project has expanded over the past four decades and continues to this day, with the participation of a large team of colleagues, local field assistants and students. I present some of the major findings of our Santa Rosa monkey research in the areas of female reproduction, sexual conflict and conservation of primates in a regenerating tropical dry forest. I also briefly describe how and why I came to develop a sideline of research on gender and science.

Queens remate despite traumatic mating in stingless bees.

Males can control female reproduction using genital plugs to impede access by rivals. In social bees, ants, and wasps, plugging may involve traumatic mating, with females being harmed. In stingless bees, chances are that plugs may promote ovarian activan, and are thought to ensure single mating-a general tendency among the social Hymenoptera. However, understanding on relationships between mating plugs, traumatic mating, and mating systems in stingless bees remains limited. To address this, we (1) compared mated queens of 7 Neotropical species to understand the patterns of copulatory marks in females and (2) compared pre- and post-mating genitalia of males and females in Melipona fasciculata to depict plug functional morphology. Data revealed an unprecedented consequence of mating in stingless bees: the characteristic marks left by mating plugs on female abdomens and the inferences that can be made from them. To our surprise, in 1 species M. fasciculata we found that queens retain the plug long after mating, and may carry it for the rest of their lives. All the other 6 species retained the plug for only a short period. Remated queens were only found in M. seminigra, whose multiple copulatory marks match previous findings of polyandry in this species. Our study shows that queens can remate, and suggests that male genital morphology may determine in part the time persistence of plugs. We conclude that traumatic mating plugs do not fully prevent remating in stingless bees and that mating systems are not uniform in this group. Nonetheless, exceptional cases of facultative polyandry in social insects-for example, when mating plugs fail-may confirm a general tendency for single mating in close link with efficient mating plugs.

Ontogenetic colour change of a sexual ornament in males of a damselfly: female mimicry, crypsis or both?

Female mimicry by males is a widespread phenomenon in several taxa and may be involved in aggression avoidance or facilitated access to resources. In early developmental stages, female mimicry may be a mechanism involved in signalling sexual immaturity or, when coupled with strategies related to visual camouflage, may be involved in the avoidance of male-male agonistic interactions. Here, we addressed whether the delayed colour maturation of a sexual ornament in males of Mnesarete pudica damselflies might be a case of crypsis, female mimicry or both. We analysed how conspecifics and predators perceive the pigmented wings of juvenile males by contrasting the wing spectra against a savannah background and the wings of both juvenile and sexually mature males and females. Our results based on the modelled visual system of conspecifics and predators suggest that the colour maturation of juvenile males may function as both crypsis and female mimicry. We discuss whether these results related to age- and sexual-dichromatism might be a mechanism to avoid unwanted intraspecific interactions or to avoid territorial and aggressive males. We conclude that the female mimicry and crypsis in juvenile males of M. pudica are mechanisms involved in avoidance of predators and unwanted intraspecific interactions, and the signalling of sexual maturity.

Morphology of parenchymally implanted foreign bodies indicates copulatory wounding in a planarian.

Traumatic mating, that is, copulation that involves wounding the partner's body, is a widespread phenomenon that is particularly prevalent in hermaphroditic animals. Traumatic mating is generally a collateral side effect of diverse strategies (from physical anchorage to injection of substances to manipulate the partner), but the trauma could also be adaptive by itself if it delays remating by the injured partner. In the Tricladida (the clade of planarian flatworms), reciprocal sperm transfer is often assumed to occur by means of a 'regular' nontraumatic copulation, that is, insertion of the penis through the partner's gonopore and deposition of the ejaculate into its genital atrium, with subsequent sperm migration to the oviducts. However, while studying the anatomy of Brazilian land planarians for taxonomic purposes, we found foreign bodies, reminiscent of spermatophores, implanted within the parenchyma of Choeradoplana albonigra (Riester, 1938). Herein, we describe and illustrate several lines of morphological evidence indicating that these foreign bodies likely represent a novel case of intragenital copulatory wounding (e.g., structural and histochemical similarity to land planarians spermatophores; implantation at the level of the gonopore; vestiges of rupture of the genital atrium's wall), corroborating that traumatic mating is pervasive and underreported in Metazoa. We also propose two different hypotheses to explain such copulatory wounding, viz., that it concerns (1) a regular mating strategy or (2) an accidental effect of the copulation. In any event, this land planarian may prove useful as a novel, noninsect terrestrial model organism to investigate the evolution of traumatic mating.

How Efficient Is Fertilization by Traumatic Insemination in Orius insidiosus (Hemiptera: Anthocoridae)?

Traumatic insemination (TI) can be injurious to females, and females have evolved various paragenital structures to mitigate these impacts. We examined the mating behavior of Orius insidiosus (Say) and the consequences of single and double matings for female fitness. A total of 100 virgin females (4-6-d old) were directly observed while they mated with virgin males. Some of these females were mated a second time with a different, nonvirgin male 3-5 d later, after they oviposited in sunflower stems. Females were held in isolation, fed eggs of Ephestia kuehniella Zeller, and reproductive success was tracked for 30 d. Six females died during their first copulation (6%), and another within 48 h, without laying eggs. Four percent of the females died during their second copulations. Copulations lasting less than 90 s usually did not result in successful fertilization, and duration of copula was positively correlated with egg fertility in singly-mated females. Duration of copula was more than halved in second matings, twice as variable, and negatively correlated with 30 d fecundity. Thirty-seven percent of singly-mated females and 31% of twice-mated females were infertile, with fewer than half of all females producing 88% of all eggs. We conclude that O. insidiosus females are likely monandrous in the wild, and that TI in this species is inefficient, contributing to high variation in female fitness. Thus, mating involves a significant mortality risk for females, despite their possession of complex paragenital structures that ostensibly mitigate copulatory injury.

The role of extragroup encounters in a Neotropical, cooperative breeding primate, the common marmoset: a field playback experiment.

In cooperatively breeding species, encounters with intruders may serve multiple functions ranging from reaffirming group territory ranges to facilitating assessments for additional breeding opportunities. While these distinctive events offer the opportunity to investigate the delicate balance of these social dimensions within animal societies, their unpredictable occurrence makes witnessing and controlling these events in the wild particularly challenging. Here we used a field playback approach to simulate conspecific territorial incursions in cooperatively breeding common marmosets (Callithrix jacchus) to distinguish between the three following non-mutually exclusive functions of intergroup encounters in this species of New World primate: territorial defense, mate defense, and assessment of breeding opportunities. For these experiments, we systematically broadcast species-typical long-distance contact calls - phees - commonly used in intergroup interactions from the core and periphery of the groups' territories using either male or female vocalizations. Consistent with a territorial defense hypothesis, a group's reaction was independent of the simulated intruder's sex and the response strength was greater when the playback stimulus was broadcast from the core areas of groups' territories relative to stimulus broadcast from periphery areas. However, sex differences in some facets of their responses suggest that this is not the only potential function for these encounters. Mated males and females started to move first in response to simulated intruders of the opposite sex, suggesting that these events offered opportunities to assess extra-pair breeding opportunities, while the occurrence of females' piloerection towards simulated female intruders is suggestive of mate-guarding. These data provide unique experimental evidence for the theory that excursions by conspecific intruders may serve multiple functions in a cooperatively breeding vertebrate and are reflective of the known complexities of common marmoset sociobiology.

Males choose to keep their heads: Preference for lower risk females in a praying mantid.

Male reproductive success is obviously mate limited, which implies that males should rarely be choosy. One extreme case of a reproductive (or mating) cost is sexual cannibalism. Recent research has proposed that male mantids (Parastagmatoptera tessellata) are choosy and not complicit in cannibalism and that they modify behavior towards females based on the risk imposed by them. Since female cannibalism depends on females' energetic state (i.e. hunger) we investigated whether male mantids are capable of using environmental cues that provide information regarding the energetic state of females to make their mate choices. Under laboratory conditions, males were confronted individually with three options: a female eating a prey, a female without a prey, and a male eating a prey (as a control for the presence of prey). Each subject comprising a choice was harnessed and placed in the corners of a triangular experimental arena at an equidistant distance from the focal male. The prey was a middle size cricket that subjects ate in approximately twenty minutes. The behavior of focal males was recorded for six hours. Females were under the same deprivation regime and, in line with previous studies, consuming one cricket did not significantly increase females' abdomen girth. Male mantids significantly preferred females that were eating a prey. In all cases choices were made after the females consumed the whole prey. This suggests that males did not use the prey as a direct way to avoid being cannibalized by keeping the female busy. The preference for females that had recently fed may have evolved because of the potential reduction in sexual cannibalism.

Origins of female genital diversity: Predation risk and lock-and-key explain rapid divergence during an adaptive radiation.

The study of male genital diversity has long overshadowed evolutionary inquiry of female genitalia, despite its nontrivial diversity. Here, we identify four nonmutually exclusive mechanisms that could lead to genital divergence in females, and potentially generate patterns of correlated male-female genital evolution: (1) ecological variation alters the context of sexual selection ("ecology hypothesis"), (2) sexually antagonistic selection ("sexual-conflict hypothesis"), (3) female preferences for male genitalia mediated by female genital traits ("female-choice hypothesis"), and (4) selection against inter-population mating ("lock-and-key hypothesis"). We performed an empirical investigation of all four hypotheses using the model system of Bahamas mosquitofish inhabiting blue holes that vary in predation risk. We found unequivocal support for the ecology hypothesis, with females exhibiting a smaller genital opening in blue holes containing piscivorous fish. This is consistent with stronger postmating female choice/conflict when predators are present, but greater premating female choice in their absence. Our results additionally supported the lock-and-key hypothesis, uncovering a pattern of reproductive character displacement for genital shape. We found no support for the sexual conflict or female choice hypotheses. Our results demonstrate a strong role for ecology in generating female genital diversity, and suggest that lock-and-key may provide a viable cause of female genital diversification.

Sexual coevolution of spermatophore envelopes and female genital traits in butterflies: Evidence of male coercion?

Signa are sclerotized structures located on the inner wall of the corpus bursa of female Lepidoptera whose main function is tearing open spermatophores. The sexually antagonistic coevolution (SAC) hypothesis proposes that the thickness of spermatophore envelopes has driven the evolution of the females signa; this idea is based in the fact that in many lepidopterans female sexual receptivity is at least partially controlled by the volume of ejaculate remaining in the corpus bursa. According to the SAC hypothesis, males evolved thick spermatophore envelopes to delay the post-mating recovery of female sexual receptivity thus reducing sperm competition; in response, females evolved signa for breaking spermatophore envelopes faster, gaining access to the resources contained in them and reducing their intermating intervals; the evolution of signa, in turn, favored the evolution of even thicker spermatophore envelopes, and so on. We tested two predictions of the SAC hypothesis with comparative data on the thickness of spermatophore envelopes of eleven species of Heliconiinae butterflies. The first prediction is that the spermatophore envelopes of polyandrous species with signa will be thicker than those of monandrous species without signa. In agreement with this prediction, we found that the spermatophore envelopes of a polyandrous Heliconius species with signa are thicker than those of two monandrous Heliconius species without signa. The second prediction is that in some species with signa males could enforce monandry in females by evolving "very thick" spermatophore envelopes, in these species we predict that their spermatophore envelopes will be thicker than those of their closer polyandrous relatives with signa. In agreement with this prediction, we found that in two out of three comparisons, spermatophore envelopes of monandrous species with signa have thicker spermatophore envelopes than their closer polyandrous relatives with signa. Thus, our results support the idea that selective pressures arising from sexually antagonistic interactions have been important in the evolution of spermatophore envelopes, female signa and female mating patterns.