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Male choice, willingness to mate and body size in seaweed flies (Diptera: Coelopidae).

The mating system of seaweed flies is characterized by scramble competition and a premating struggle during which females vigorously attempt to remove mounted males and prevent copulation. Here we investigate factors affecting the willingness of males to mate in five species of Coelopa : C. frigida, C. nebularum, C. ursina, C. pilipes and C. vanduzeei. The data from 2000 individual observations of pairs of flies revealed no association between female size and the willingness of males to mate, suggesting that males do not exhibit mate choice for larger, more fecund females. Male willingness to mount was associated with male size in C. frigida, C. nebularum and C. ursina, although not in C. pilipes and C. vanduzeei. There is considerable intraspecific variation in male size in the former three species, including the presence of two almost discrete size classes within C. nebularum. We suggest that males of different sizes in these species may be adopting alternative reproductive strategies. Large males may benefit from increased longevity and their ability to withstand female rejection responses, whereas small males are more active and thus likely to encounter and mount more females. In addition small males develop faster and thus gain earlier access to females. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Male mating patterns in wild multimale mountain gorilla groups.

Although mountain gorillas, Gorilla gorilla beringei, are classified as having a one-male mating system, approximately 40% of the social units are multimale groups. I observed two multimale groups of mountain gorillas at the Karisoke Research Center, Rwanda, Africa, for 17 months to determine male mating patterns and male-male mating harassment in relation to both male dominance rank and female reproductive status. Dominant males mated significantly more than did individual subordinate males, and dominant males mated more with cycling adult and pregnant females. The dominant males participated in 47 and 83% of observed matings in the two groups. Subordinate males were more likely than dominant males to mate with subadult females. Eleven of 14 females were observed to mate with more than one male, and multiple males mated with three of the five females observed at the probable time of conception. Mating harassment was initiated and received by both dominant and subordinate males. Mating harassment occurred infrequently (during 30 and 22% of matings in each group), usually consisted of mild aggression, and usually terminated copulations by subordinate males, but not those by dominant males. These results suggest that multimale mountain gorilla groups can be favourable environments for subordinate males to obtain mating opportunities. Dominant males may be unable or unwilling to prevent subordinate males from mating. Based on behavioural observations, mountain gorillas can have a multimale mating system but further research on the role of females in male mating success and paternity determination is needed to understand fully this species' mating system. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Mate, neighbour and stranger songs: a female song sparrow perspective.

We investigated discrimination by female song sparrows Melospiza melodia, between different categories of male song using the copulation solicitation display as a preference assay. Females responded most strongly to songs recorded from their mates, less strongly to songs of neighbouring males and least strongly to songs of stranger males. Among the stranger songs, however, females preferred songs that were most similar structurally to song types in their mates' repertoires (matching songs). These results are interpreted as evidence that females can recognize individual males based on the songs in their repertoires. Moreover, the observed female preferences for nonmatching neighbour and matching stranger song over nonmatching stranger song, suggest that any male with songs structurally similar to mate songs or even to nonmate but local neighbourhood songs, will be at an advantage in sexual interactions with local females. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Floater males gain reproductive success through extrapair fertilizations in the stitchbird.

We used minisatellite DNA profiling to assign parentage to stitchbird, Notiomystis cincta, chicks from a breeding population on Tiritiri Matangi Island off the coast of the North Island of New Zealand. The small population size allowed samples to be collected from all potential parents and nearly (33/34 nestlings) complete assignment of paternity. Analysis revealed that 35% of nestlings (12/34) were the result of extrapair copulation and that extrapair young were present in 80% of nests (8/10). About half of the extrapair nestlings were the offspring of unpaired males. This is substantially higher than predicted from the literature, which suggests that extrapair paternity is typically gained by paired males. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Proximate factors associated with high levels of extraconsort fertilization in polygynous grey seals.

Behavioural estimates of male mating success in polygynous grey seals, Halichoerus grypus, may be misleading as females are known to be promiscuous. At Sable Island, Nova Scotia, we collected behavioural observations and skin samples for paternity analysis from 56 females and their attending males. Twenty-four of these females were found in the following year and their offspring were sampled. Using seven hypervariable microsatellite loci, we excluded the consort male as the father in 43% of the cases. The probability of exclusion of these seven loci was 98.2%. Contrary to expectations, inland females had higher rates of extraconsort fertilizations (ECFs) (70%) than beach females (23%). Younger females (<9 years) had slightly more ECFs than older females, but this was not significant. The duration of male consortship did not differ between females with ECFs and females fertilized by their consort male. Two explanations may account for the inland females having more ECFs: a higher ratio of females to tenured males inland may provide a greater opportunity for nonconsort males to obtain copulations; and inland females travel greater distances to depart for the ocean and may attract more males. These results are more consistent with the hypothesis that ECFs are a by-product for females of male strategies to maximize reproductive success than with hypotheses concerned with either material or genetic benefits gained by females. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Central males instead of multiple pairs in redfronted lemurs, Eulemur fulvus rufus (Primates, Lemuridae)?

In group-living Malagasy primates (Lemuriformes), certain demographic and morphological traits deviate both from theoretical expectations derived from sexual selection theory and from patterns in better-known anthropoid primates. Lemurs lack sexual dimorphism in body and canine size and live in relatively small multimale-multifemale groups with, on average, even adult sex ratios, despite a polygynous mating system. In addition, the majority of gregarious lemurs are cathemeral, that is, they are regularly active both day and night. The social system of cathemeral lemurs has been considered an adaptation to their activity pattern. If so, groups should consist of multiple pairs that aggregate during diurnal activity and range separately at night in response to predation and infanticide risk, respectively. Moreover, mating privileges should exist between pair partners. We tested this hypothesis with data collected on two groups of wild redfronted lemurs in western Madagascar during 1023 focal animal hours spanning 4 months including the mating season. In addition, data on spatial relations were collected on 12 nights. We found no differences in group cohesion between day and night and no behavioural evidence for multiple male-female pairs. Instead, one male in each group monopolized social interactions with all females. However, females copulated with all resident males, although most often with the central male. Thus, basic predictions of the cathemerality hypothesis are unsupported. Multiple matings and indications of oestrous synchrony can be viewed as female counterstrategies against infanticide, whereas reproductive strategies of redfronted lemur males remain obscure. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Male infanticide in captive plains zebra, Equus burchelli.

On the assumption that infanticide exists in plains zebra, as reported for horses, Equus caballus, we tested the following hypothesis. Introducing a new zebra male into a herd of breeding females should increase foal mortality in comparison with herds in which the sire of the foals is still present. The younger the foal, the more likely infanticide should be. We collected data from five herds in two zoological gardens in the Czech Republic. We found nine records of infanticide in plains zebra and three cases of abortions that were probably induced by forced copulation. We analysed additional indirect data to investigate the possibility of introduced males causing other abortions. Abortions were three times more likely in herds with introduced males than with only fathers present. Postnatal mortality of the foals was four times greater with introduced males than with fathers. No indication of a sex preference was observed for infanticide by a new male for either abortions or postpartum deaths. When we combined all records involving introduced males, the probability of foal death was greatest when the new male joined the herd just after conception and decreased with increasing time between conception and date of the new male introduction (the chance of a foal surviving was less than 5% just after conception and more than 50% at the time of delivery). Mortality of foals did not depend on whether the new male was introduced before or after the foal was born. Survival increased to more than 60% after the foal reached 1 month of age. Our results suggest that captive plains zebra show the highest occurrence of infanticide reported among ungulates. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Lack of pubertal influences on female dispersal in muriqui monkeys, Brachyteles arachnoides.

The hormonal mediation of dispersal in female mammals is poorly understood, in part because of the difficulties of detecting the onset of ovarian cycling and puberty in dispersing individuals. We used noninvasive methods of faecal steroid assays to determine the timing of dispersal relative to puberty and ovarian cycling in wild female muriqui monkeys, a species in which males are philopatric and nearly all females transfer from their natal groups. Natal females had a mean+/-SE age of 73.4+/-7.2 months (N=18) at the time of their transfers. Intergroup transfers occurred when one or more sexually active adult females were present, but did not show any seasonal patterns. Faecal progesterone and oestradiol profiles from nine natal females prior to transfer and four non-natal females that transferred into our study group demonstrate unequivocally that dispersal occurs prior to puberty in this species. All females showed baseline oestradiol levels and low progesterone levels compared with cycling adult females. Immigrants were first observed to copulate at 11.2+/-2.2 months of age (N=4), prior to the onset of normal ovarian cycles, and gave birth to their first offspring at 33.8+/-7.3 months (N=4) after transferring. Mean cortisol levels did not differ between natal emigrants or recent immigrants, and were within the range of those of adult males during the nonbreeding season in 10 of the 11 prepubertal females sampled. These results indicate that female dispersal is not triggered by activational hormones associated with puberty or escape from reproductive suppression in this species. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Extraterritorial forays and male parental care in hooded warblers.

Extrapair paternity is common among many songbird species yet few studies have quantified male extraterritorial foray (ETF) effort and examined potential trade-offs. One potentially important constraint for males is the need to provide parental care. Current models of male extrapair mating tactics propose that males reduce extraterritorial foray effort later in the breeding season because they face a trade-off between feeding nestlings versus pursuing extrapair matings. However, detailed field studies examining the trade-off between paternal care and male extraterritorial forays are lacking. We used radiotelemetry to quantify male extraterritorial foray effort in hooded warblers, Wilsonia citrina, to test the widely held predictions that: (1) males make significantly fewer and shorter forays during the nestling stage relative to other stages (i.e. fertile and incubating stages); and (2) male extraterritorial foray effort is negatively correlated with parental effort. Males made 0.87+/-0.09 forays/h and spent on average 12.2% of their time foraying off territory. Results were equivocal; some data suggested male foray effort decreased in relation to parental care, while other data suggested otherwise. Pairwise tests controlling for (1) extrapair mating opportunity among males and (2) male, territory and social mate quality revealed a possible trade-off between the mean duration and percentage of time in extraterritorial foray versus providing parental care. Conversely, results also revealed (1) no difference in foray rate, foray duration or percentage of time spent off territory over the various stages of the breeding season, (2) no relationship between male foray effort and male feeding rate, and (3) no difference in foray rate in pairwise comparisons, controlling for variability in extrapair mating opportunity and male quality. Overall, the trade-off between providing male parental care and pursuing alternative mating tactics may not be as strong for male hooded warblers as once hypothesized because males dedicated relatively little time to seeking extrapair copulations off territory. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Behavioural isolation between two closely related Hawaiian Drosophila species: the role of courtship.

Behavioural isolation plays a critical role in several recent models of speciation. A detailed understanding of the process of speciation requires analysis of taxonomic groups that have not completed reproductive isolation. We studied D. silvestris and D. heteroneura because they are still in the process of divergence: behavioural isolation between them is incomplete, and neither postzygotic nor ecological isolation has been detected. Behavioural isolation is due to the failure of courtships between male D. silvestris and female D. heteroneura: there is no postzygotic isolation from either parental species. The F1 hybrids are as successful in courtship with parental individuals as same-species pairs, which suggests that the hybrids resemble male D. heteroneura or female D. silvestris in some behaviour patterns that are crucial to mating success. We searched for this crucial resemblance by examining courtship between F1 hybrids and the parental adults. We found that successful F1 males are somewhat more similar than unsuccessful F1 males to D. heteroneura males, but nevertheless they were intermediate between males of the two species. We also found that in both species the presence of female wings is necessary for courtship to proceed to copulation. These results reinforce an earlier report that behavioural isolation between these species is largely attributable to the decision as to whether to court at all, rather than to the details of courtship. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Reproductive biology of captive male cottontop tamarin monkeys as a function of social environment.

The cottontop tamarin, Saguinus oedipus oedipus, is a cooperatively breeding monkey in which mature male and female offspring serve as helpers to assist in rearing younger siblings. Generally, only one female per social group reproduces; breeding restriction is mediated in postpubertal female offspring through low and acyclic levels of reproductive hormones. We investigated (1) reproductive activity of postpubertal male offspring, and (2) whether aggression towards male offspring and a cortisol-mediated stress response might restrict breeding of male offspring in the natal group. We examined sexual behaviour, olfactory communication and urinary hormone levels (testosterone, dihydrotestosterone, luteinizing hormone, cortisol) of the subject males while we manipulated their social environment from housing in natal groups to pairing with a novel female, and after the production of their own offspring. Mounting and erection rates of the male subjects were as high in the natal group as when paired with a novel female. However, most mounts in the natal group were directed towards other males, and complete copulation sequences did not occur with natal-group females. Social environment had no significant effect on olfactory investigation of breeding females. Although hormone levels increased significantly after the subjects were removed from the natal group, the elevation was transient; the hormone levels of subjects in their natal groups did not differ from the levels shown by the same males when successfully producing their own offspring. Male offspring received more contact aggression in the natal group than when paired with the novel female. However, most of the aggression was received from siblings rather than the breeding pair, and levels of cortisol did not correspond with levels of aggression. Thus, at both a behavioural and endocrine level, mature male offspring in captive natal groups were potentially fertile, but sexual activity with natal-group females appeared to be behaviourally restricted and directed instead towards group males. In wild cottontop tamarin groups, this reproductive potential may allow male helpers flexibility to respond to breeding opportunities. Copyright 2001 The Association for the Study of Animal Behaviour.

Journal Article↗

Sp4, a member of the Sp1-family of zinc finger transcription factors, is required for normal murine growth, viability, and male fertility.

We report the cloning, characterization, and targeting of an Sp1-related zinc finger transcription factor gene from the distal arm of mouse chromosome 12. This gene, previously identified in rats and humans and designated sp4, is homologous to the Drosophila buttonhead (btd) gene, which is expressed in the head region of developing flies. Similarly, in situ hybridizations show that sp4 is highly expressed in mouse embryos in the developing central nervous system (CNS). Expression of sp4 is seen as early as Day 9 of development, where transcripts are abundant in the posterior neuropore. Expression in later embryos is detected throughout the CNS as well as in other structures, including the nasal mucosa, the vomeronasal organ, the epithelium of the lung and intestinal tract, the testes, and the developing teeth. Northern blot analysis showed sp4 expression in the adult brain and other tissues. Gene targeting by homologous recombination was used to determine the role of sp4 during mouse development. Two-thirds of homozygous mutants die within the first few days after birth and those that survive are smaller than their wild-type littermates. While fertility of the female mutants appears normal, homozygous mutant males do not breed, despite having histologically intact testes containing mature sperm. sp4/sp4 mutant males fail to copulate, indicating that this gene is required for normal male reproductive behavior.

Amino Acid Sequence↗

Analyses of reproductive interactions that occur after heterospecific matings within the genus Caenorhabditis.

Formation of zygotes in internally fertilizing organisms requires a number of successful interactions between oocytes and sperm within a receptive female reproductive tract. These interactions are usually assumed to be species-specific. For most species, it is either not possible to inseminate females with sperm from a different species or not possible to observe the consequences of such an insemination because the female is opaque. Nematodes of the genus Caenorhabditis are optically transparent and prior work indicates copulation between individuals of two different species is possible. We have used a series of vital stains and other cytological methods to analyze sperm after cross-species mating. We present here a series of analyses of the postcopulatory, prefertilization interactions among three Caenorhabditis species and find that reproductive biology is conserved, to varying degrees, among all three species. This approach allows investigation into which in vivo interactions between sperm and both oocytes and the somatic gonad have been maintained during the reproductive isolation that accompanies speciation.

Animals↗

Developmental toxicity of inhaled trans-1,2-dichloroethylene in the rat.

The developmental toxicity of trans-1,2-dichloroethylene (t-DCE), a component of certain Freon cleaning agents, was examined in pregnant rats. t-DCE was administered by inhalation 6 hr daily on Days 7-16 of gestation (the day copulation was confirmed was termed Day 1 of gestation) at exposure levels of 0, 2000, 6000, or 12,000 ppm. The offspring were then examined on Day 22 of gestation. Overt maternal toxicity was expressed as a significant reduction in weight gain at 12,000 ppm and in feed consumption at 6000 and 12,000 ppm. During the exposure period, lacrimation and stained periocular hair, and signs of ocular irritation, were observed in all groups. In addition, increased incidences of alopecia, lethargy, and salivation were observed in the high-dose dams. Significant increases in the mean number of resorptions per litter were seen in the litters of dams exposed to 6000 and 12,000 ppm of t-DCE; however, these values are within the range of historical controls and not considered to be treatment related. The mean combined and female fetal weights were significantly reduced in the litters of dams exposed to the highest concentration (12,000 ppm) of t-DCE. Marginal effects on feed consumption, unaccompanied by other changes and reflective of the pattern seen at higher doses, were seen at 2000 ppm. Thus, marginal maternal toxicity was seen at 2000 ppm and exposures to 6000 ppm t-DCE or higher caused frank maternal toxicity while the fetus was affected only at 12,000 ppm. Therefore, t-DCE is not considered to be uniquely toxic to the rat conceptus.

Abnormalities, Drug-Induced↗

Developmental toxicity of inhaled N-methylformamide in the rat.

The developmental toxicity of N-methylformamide (MMF), an industrial chemical intermediate used in the production of agrichemicals, was examined in pregnant rats. MMF was administered by nose-only inhalation, 6 hr daily on Days 7-16 of gestation (the day copulation was confirmed was termed Day 1 of gestation, Day 1G) at exposure concentrations of 0, 15, 50, or 150 ppm. Dams were regularly monitored throughout gestation for body weight gain, feed consumption, and clinical signs. Cesarean sections were performed on Day 22G and the offspring were examined. Maternal toxicity was evident in dams exposed to 50 or 150 ppm; one dam exposed to 150 ppm died on Day 14G (considered to be treatment-related) and dams in the 50 and 150 ppm groups exhibited concentration-related clinical findings. Clinical signs of wheezing and rattling were observed both during and after the exposure period. The 150 ppm group also showed significant decreases in weight gain and feed consumption. A significant increase in the mean number of resorptions per litter at the 150 ppm level indicated an embryolethal effect. Developmental toxicity was apparent by a significant decrease in mean fetal body weight and increases in fetal malformations (subcutaneous cysts on the head, microphthalmia, anophthalmia, fused ribs and/or vertebra, and distended brain ventricles) and variations (misaligned and fused sternebrae) due to retarded development at 150 ppm. Significant fetal body weight decreases were also present at 50 ppm. Thus, in this study, the no-observable-adverse-effect level for both dam and fetus was 15 ppm MMF, indicating that for the parameters included in this study, the conceptus is not uniquely sensitive to MMF.

Administration, Inhalation↗

The sexually dimorphic medial preoptic nucleus of quail: a key brain area mediating steroid action on male sexual behavior.

About 10 years ago, a sexually differentiated nucleus was identified in the preoptic area (POA) of the Japanese quail in the course of studies analyzing the dimorphic mechanisms involved in the activation of sexual behavior. In this species, males exposed to testosterone copulate while females never show this masculine behavior. The present paper reviews anatomical, neurochemical, and functional data that have been collected since that time about the quail dimorphic nucleus. The medial preoptic nucleus (POM) is significantly larger in adult male than in adult female quail. Its volume is also steroid-sensitive in adulthood: it decreases when circulating levels of testosterone are low (castration, exposure to short-days) and it increases when testosterone levels are high (treatment with testosterone, exposure to long-days). The POM is a necessary and sufficient site of steroid action for the activation of male copulatory behavior. The volumetric difference of the POM results from a difference in the adult hormonal milieu of males and females (activational effect) and is not affected by embryonic treatments that permanently modify sexual behavior (no organizational effects on POM). In contrast, the size of neurons in the dorsolateral part of POM appears to be irreversibly affected by embryonic steroids and this feature is therefore a better correlate of the behavioral sex difference. The POM is characterized by the presence of a wide variety of neurotransmitters, neuropeptides, and receptors. It can, in addition, be specifically distinguished from the surrounding POA by the presence of aromatase-immunoreactive cells, by a high density of alpha 2-adrenergic receptors, and by a dense vasotocinergic innervation. Some of these neurochemical markers of the dimorphic nucleus are themselves modulated by steroids. In particular, the aromatase-immunoreactive cells of the lateral POM appear to be a key target for steroids in the activation of male copulatory behavior. The POM is bidirectionally connected to many brain areas. It receives inputs from a variety of sensory areas and from a number of regulatory areas (e.g., catecholaminergic cell groups). This nucleus also sends outputs to "neurovegetative" centers and to brain regions directly connected to the motor pathways. These connections fully support the role of the POM as an integrative center for the control of male sexual behavior. The available data indicate that there is a high degree of steroid-induced neuronal plasticity in the POM, including changes in neuronal function, in protein synthesis, and in specific inputs. These phenomena can easily be studied in the POM because they are of a large magnitude, they are localized in a specific brain site, and they develop rapidly after exposure to steroids. They are also directly related to a clear functional output, the activation of male sexual behavior. The quail POM therefore constitutes an exceptional model for the analysis of steroid-induced brain plasticity in a functionally relevant context.

Androgens↗

Plasma LH and steroid hormones in King penguin (Aptenodytes patagonicus) during the onset of the breeding cycle.

Temporal changes in plasma levels of luteinizing hormone (LH), testosterone, estradiol, and progesterone were measured throughout molt and the onset of reproduction in free-living male and female King penguins (Aptenodytes patagonicus) at Crozet Island (46 degrees S, 51 degrees E). In both sexes, LH concentrations began to rise as soon as molting ended, before the departure for refeeding at sea, suggesting that the beginning of reproductive activity is not associated with refeeding and/or courtship behavior. LH remained at high values throughout courtship and decreased to lower levels during incubation. Gonadotrophin secretion covaried with sustained plasma testosterone levels in males and increased estradiol and progesterone levels (peaking at copulation) in females.

Animals↗

Reproductive endocrinology and weight change in relation to reproductive success in the magellanic penguin (Spheniscus magellanicus).

The Magellanic penguin is a colonial monogamous species that lays only a single clutch of two eggs per year. However, failed breeders remain at the colony and engage in nest building, fights, and copulations without relaying. The seasonal changes in reproductive hormones and body weight through the nesting cycle were studied, with respect to the reproductive success or failure of individuals. Body weight changed dramatically in both sexes through the season, in response to fasting during incubation, and high body weight in males at the onset of incubation was a strong predictor of eventual reproductive success. Circulating steroid hormones had a biphasic seasonal pattern, with elevated levels during the sexual phase of breeding (prior to egg laying), declining to low, stable levels during the parental phase after eggs were laid. Luteinizing hormone levels were elevated in females, but not in males, prior to egg laying. Both sexes responded to reproductive failure by increasing the secretion of testosterone, and females also increased the secretion of estradiol, a response that would be expected of a species that can renest following failure. However, renesting is extremely rare, and this hormonal response to failure may instead serve to promote maintenance of pair bonds and also territory ownership across years.

Animals↗