Egg coloration is correlated with female condition in eastern bluebirds (Sialia sialis)

Egg coloration has been hypothesized to reflect female condition. Because of the proposed physiological costs associated with deposition of biliverdin pigments and because of their conspicuousness, eggs with blue-green coloration may reliably convey information about female or brood quality. We tested the hypothesis that expression of blue-green coloration of eastern bluebird (Sialia sialis) eggs positively correlates to female condition. First, we documented the incidence of egg color polymorphism within the population. We observed that 98% of females laid blue-green eggs while less than 2% laid white eggs and less than 1% laid pink eggs. In a subset of clutches, we used full spectrum reflectance spectrometry (300–700 nm) to compare eggshell coloration to measures of female condition. We found that the color of eggs within clutches was more similar than the color of eggs from different clutches, and that the blue-green eggs have spectral peaks that are consistent with the characteristic absorbance spectra of biliverdin pigmentation. Females in better body condition and older females laid more colorful eggs. Moreover, individual females laid more colorful eggs later in the laying sequence. Overall, these data indicate that egg coloration covaries with female condition, suggesting that egg coloration could function as a reliable signal of female quality or that egg coloration may allow females to recognize eggs laid by conspecific brood parasites.     

Multiple maternity and paternity in single broods of apparently monogamous eastern bluebirds (Sialia sialis)

Summary In an effort to assess genealogical relationships between breeding adults and their putative offspring, we collected and analyzed blood from 257 wild, uniquely color-banded eastern bluebirds (Sialia sialis). Electrophoretic analysis of the protein products of two loci, esterase-2 (EST-2) and nucleoside phosphorylase (NSP), revealed that at least 5% of adult males and 15% of adult females were caring for at least one offspring not their own. Multiple parentage was evident in 9% of all broods sampled. Twenty-five percent of complete families sampled evidenced multiple parentage. Biparental care of nestlings by adult eastern bluebirds is not necessarily a reflection of the genetic kinship between putative parents and their offspring. The consort pattern among eastern bluebirds does not completely correlate with the pattern of gametic contributions, leading to a reconsideration of their mating system as possibly polygamous. This is among the first unambiguous observations of multiple patenity in a wild bird population, and of multiple maternity in a wild passerine.     

Cave molly females (Poecilia mexicana, Poeciliidae, Teleostei) like well-fed males

We examined the preference of Atlantic molly females (Poecilia mexicana) to associate with a well-fed or a starved male in simultaneous choice tests. Females from three different populations were tested in three treatments: (1) the females could choose on the basis of multiple cues from the males (visual plus non-visual); (2) only non-visual cues could be perceived in darkness, (3) only visual cues were presented. The three tested populations differ clearly in their ecology: one population occurs in a typical river habitat, the second one in a milky sulfur creek outside a cave, and the third population occurs in a cave habitat (cave molly). In the river-dwelling population, females never showed a preference. In the population from the sulfur creek, females preferred to associate with the well-nourished male when visual cues from the males were available. Only cave molly females exhibited a strong preference for well-nourished males in all treatments. A morphological comparison demonstrated that wild-caught males from river habitats are typically in a good nutritional state. In the sulfur creek, males showed signs of starvation. Cave molly males were in an even worse nutritional state. In the cave population, saturated males probably indicate high fitness, thereby driving the evolution of the preference for good male nutritional state.Communicated by K. Lindström     

The effect of rearing environment on blue structural coloration of eastern bluebirds (<Emphasis Type="Italic">Sialia sialis</Emphasis>)

We used a brood-size manipulation to test the effect of rearing environment on structural coloration of feathers grown by eastern bluebird (Sialia sialis) nestlings. Ultraviolet (UV)-blue structural coloration has been shown to be sexually selected in this species. Our experimental design took advantage of the growth of UV-blue wing feathers in nestlings that are retained as part of the first nuptial plumage. We cross-fostered nestlings to create enlarged and reduced broods with the purpose of manipulating parental feeding rates and measured the effect on nestling growth and plumage coloration. Brood size influenced feeding rates to offspring, but the effect varied with season. In general, male nestlings reared in reduced broods were fed more often, weighed more, and displayed brighter structural plumage compared to nestlings reared in enlarged broods. Female nestlings appeared to experience less adverse affects of brood enlargement, and we did not detect an effect of brood-size manipulation on the plumage coloration of female nestlings. Measures of plumage coloration in both males and females, however, were correlated to hatching date and nestling mass during feather development. These data provide empirical evidence that environmental quality can influence the development of the blue structural coloration of feathers and that males may be more sensitive to environmental fluctuations than females.     

Inter-sexual conflicts over copulations in the European starling: evidence for the female mate-guarding hypothesis

Recent studies have suggested that conflicts of interests between the sexes may lead to variation in copulation behavior among pairs. We examined differences in the rate and timing of copulation solicitations and copulations among females of different mating status in the facultatively polygynous starling in an attempt to explain why females copulate repeatedly with their mate. All within-pair copulations were female-solicited indicating that females control copulations in the starling. Before egg laying, females solicited copulations at a high rate (usually more than 2 per hour). In contrast to most other species studied so far, females continued to solicit copulations throughout the egg laying period, and also solicited after egg laying (the latest solicitation occurred when the nestlings were 4 days old). Primary females solicited more copulations than monogamous females both during and after the fertile period. Many copulation solicitations of primary females occurred at the nestbox where their male was singing to obtain an extra female. Both primary and secondary females solicited more copulations after the egg-laying period than monogamous females. A large proportion of female copulation solicitations was refused by the male partner: female solicitation resulted in more male refusals in primary females than in monogamous females. Primary females were more likely to be chased aggressively when they solicited a copulation than monogamous females; most aggressive chases occurred when primary females flew towards their male to solicit copulation when he was singing at another nestbox. Overall, our results demonstrate that there is a conflict over copulation between males and females in polygynous pairs. The conflict presumably relates to the cost of sharing male parental investment: females use copulation solicitation behavior to interrupt their singing males apparently in an attempt to prevent them from becoming polygynous. We present the first empirical evidence that female songbirds use copulation solicitation behavior as a form of mate guarding, often in a non-reproductive context. We did not find a positive relationship between copulation rate during the fertile period and the amount of male parental care as is predicted by the paternity confidence hypothesis.     

Sneak copulations in long-tailed macaques (Macaca fascicularis): no evidence for tactical deception

Sexual competition is highly prevalent within multi-male multi-female primate groups and may lead to copulations in absence of potentially interfering bystanders. Such avoidance of bystanders may result from tactical deception or from simpler mechanisms such as taking advantage of encountered situations without bystanders, operant conditioning or a peripheral positioning of non-alpha males. We investigated which individuals are avoided as bystanders, how individuals react to the presence of bystanders and whether copulation partners separate themselves from the group in a tactical way. Our observations of a group of 15 female and seven male long-tailed macaques housed in three interconnected, but visually separated compartments revealed that both males and females can interrupt sexual behaviour and that bystanders of both sexes were avoided during copulations (n?=?256). The strength of the effect of bystanders tended to decrease with the dominance rank of male bystanders, but did not depend on the dominance rank of female bystanders. The audience effects of non-alpha individuals did not depend on the strong audience effect of the alpha male in combination with proximity with the alpha male. The effects that we found for separate bystanders suggest that sexual competition concerns rank dependent male–male competition and rank independent female–female competition. Additionally, both male and female copulation partners paid attention to the presence of bystanders and conducted fewer copulation solicitations in their presence. The timing of a male and female’s separation from the group suggests that exploitation of the peripheral position of non-alpha males, and not tactical deception, may cause these audience effects.     

Male philopatry,extra-pack copulations and inbreeding avoidance in Ethiopian wolves (Canis simensis)

Monogamous pairings have been regarded as the fundamental social unit in all canid species, including those living in packs. In Ethiopian wolves, however, habitat saturation limits dispersal, which raises the question of whether they avoid inbreeding and, if so, by what mechanism. In two study areas Ethiopian wolf packs had stable memberships. Each pack comprised two to eight adult males, one to three adult females, including a clear-cut dominant individual of each sex, together with one to six yearlings and up to six pups (n = 9 packs). Males remained in their natal packs, apparently throughout their lives. Some females also failed to disperse while others dispersed in their second or third year and became floaters. Dominant females monopolized breeding, and were succeeded either by their most dominant daughters (three cases) or by floaters (two cases). In the former case there is potential for incest; however, 70% of 30 copulations observed were between the dominant female of one pack and a male from an adjoining pack. In Ethiopian wolves, under conditions where dispersal is constrained and the potential for inbreeding is high, extra-pack matings (and associated multiple paternity) result in outbreeding. We raise the possibility that extra-pair copulations may be widespread in canid societies and that the monogamy supposedly fundamental to the family may be more sociological than genetic. Received: 15 October 1994/Accepted after revision: 9 December 1995     

Alternative mating tactics in males of Polistes dominulus (Hymenoptera: Vespidae)

Summary In summer, males of Polistes dominulus form large aggregations at sunny landmarks. We identified two size-correlated behavioural categories: residents (R) and transient (T). R males, which constitute 20%–25% of the total population, are larger than T males, territorial, aggressive, and more site-faithful, while T males range more widely, are non-aggressive, and show little site tenacity. Field and laboratory data suggest that R males have an advantage in mating, particularly if they engage in frequent flights while on their territories. These alternative mating tactics within the same population are combined with behavioural flexibility in some individuals, which switch from one option to the other.     

Underwater oviposition in a damselfly (Odonata: Coenagrionidae) favors male vigilance,and multiple mating by females

Summary Female Enallagma hageni oviposit underwater where they are inaccessible to males. I demonstrate that males guard submerged females rather than perch sites, and are behaviorally distinct from lone males at the water. In contrast to lone males, which always attempt to copulate with females presented to them, guarding males exhibit a conditional latency to remating which corresponds closely to the time required by females to oviposit a complete clutch of eggs. By ovipositing underwater, females decrease the risk that their eggs become exposed. Risks associated with submerged oviposition favor both mate guarding, and multiple, within-clutch matings by females. Both guarding mates and lone males rescue females that float on the water surface as the result of improper resurfacing. Such behavior reduces the mortality risk to females from 0.06 to 0.02 per oviposition bout. By remating between bouts, females benefit from the additional vigilance of lone males, who rescue floating females 1.4 times as often as original mates. A second consequence of multiple mating is an increase in the selective advantage of vigilance by mates. Because receptive females become scarce by early afternoon, whereas male density remains high, a male has little (3%) chance of encountering a second receptive female that day. However, he incurs a 42% risk of losing fertilizations if he abandons a mate. For male E. hageni mate guarding functions in the context of both natural and sexual selection. It insures that a mate lives to lay a complete egg clutch in addition to protecting a male's sperm investment.     

Fifteen-spined stickleback (Spinachia spinachia) females prefer males with more secretional threads in their nests: an honest-condition display by males

In the fifteen-spined stickleback (Spinachia spinachia), the male alone builds a nest and provides care for the eggs until they hatch. The nest is made of filamentous algae held together with shiny secretional threads of a glycoprotein, here called tangspiggin. Tangspiggin is produced by transformed kidney cells. I investigated how tangspiggin production was related to food intake by giving males low or high food rations. Males in the high-food group produced significantly more tangspiggin than those in the low-food group. Since low food rations did not lead to a significant drop in resting metabolic rate (oxygen consumption), tangspiggin production appears to be one of the first energy-demanding processes to be dropped during food deprivation. Among males given equal food rations, females preferred to mate with those with elevated levels of tangspiggin in their nests. Tangspiggin threads may signal safety for the offspring by keeping the eggs within the nest and protecting them from egg predators. Moreover, the threads become less shiny with age, due to particle deposition, and older nests that may house older (more valuable) eggs are thereby well camouflaged. Tangspiggin provides an honest signal of male condition, acting as an extra-bodily ornament, signaling direct benefits to females or possibly also indirect benefits of "good genes" from a male who is a successful forager.     

Fragrant legs in Paysandisia archon males (Lepidoptera, Castniidae)

Paysandisia archon was accidentally introduced into Europe where it damages endemic and ornamental palms, also threatening the date palm from North Africa. Little was known about sex pheromones in the Castniidae day-flying moth before a recent paper that concluded on the absence of female sex pheromone in P. archon. A putative identification of a short-range male pheromone, present on fore- and hindwings, was reported. In this paper, we describe the original structure of the male androconia located on the tarsi of the mid-legs. The extracts of mid-legs were analysed by GC/MS and the chemical structure of the male androconia component was identified as E2,Z13-18:OH. After extraction in solvent, biological activity of the extracts was assessed by EAG. The chemistry and the morphology of the androconia reinforce the current classification of the Castniidae in the Cossoid/Sesioid assemblage and provide new information on the chemical ecology in day-flying Lepidoptera and suit the recent paper describing the courtship behaviour     

Female yellowhammers (Emberiza citrinella) prefer yellower males: a laboratory experiment

The importance of male plumage coloration as a signal of male dominance and a cue for female choice in the monogamous yellowhammer, Emberiza citrinella, was tested in two sets of experiments in an indoor aviary. Dominance was tested by introducing two individuals, with no previous experience of each other, in an aviary with food and water. Aggression occurred more often between two males than between a male and a female and more frequently between pairs of males including at least one old male than between two young males. Dominance was not related to male colour in trials between males of the same age class, but it was in trials between an old and a young male, often differing in colour. Thus, age may be a more important determinant of dominance than colour. Female preference for more colourful males was tested by allowing hormone-induced females to choose between a more and a less colourful dummy male. Females spent more time in front of more colourful than drabber males and also more often perched beside colourful males than duller individuals. Although male colour cannot be ruled out as a dominance signal, the results suggest that male colour is primarily used as a signal in mate choice. Female choice may hence be responsible for maintenance of bright plumage in the male yellowhammer.     

The morphology and behavior of dimorphic males in Perdita portalis (Hymenoptera : Andrenidae)

Summary In Perdita portalis, a ground nesting, communal bee, males are clearly dimorphic. The two male morphs are easily distinguished based on head size and shape into (1) a flight-capable, small-headed (SH) morph that resembles the males of other closely related species and (2) a flightless, large-headed (LH) morph that possesses numerous derived traits, such as reduced compound eyes, enlarged facial foveae and fully atrophied indirect flight muscles. The SH morph occurs exclusively on flowers while the LH morph is found only in nests with females. While on flowers, SH males are aggressive, fighting with conspecific males and heterospecific male and female bees, and they mate frequently with foraging females. Using artificial observation nests placed in the field, I observed the behavior of females and LH males within their subterranean nests. LH males are aggressive fighters; males attacked each other with mandibles agape, and male-male fights always ended in the death of one male. LH males are highly attentive to the reproductive behavior of females; they spend increasing amounts of time near open cells during cell provisioning, and mating only takes place immediately prior to oviposition when females are forming the accumulated pollen and nectar into a ball. Based on larvae reared to adulthood in the laboratory, the two male morphs occur in equal proportions. The behavior of males in closely related species, especially P. texana, and the origin and maintenance of male dimorphism are discussed.     

Mustached males in a tropical poeciliid fish: emerging female preference selects for a novel male trait

One possible mechanism for the (co-)evolution of seemingly novel male traits and female preferences for them is that males exploit pre-existing female biases, and livebearing fishes (Poeciliidae) have been at the forefront of this research for almost two decades. Here, using 13 poeciliid species from four different genera, we tested whether mustache-like rostral filaments found in males of some Mexican molly (Poecilia sphenops) populations could have evolved due to exploitation of a pre-existing female bias. While Mexican mollies were the only species with a significant female association preference for mustached males, we also did not find any species exhibiting significant aversion for mustached males; rather, variance in female preference scores was large throughout. For example, more than 25% of females spent twice as much time with the mustached male compared to the non-mustached male in most species, but even 31% of Mexican molly females spent more time near the non-mustached male. Hence, a comparison of the strength of preference was inconclusive. We discuss the possibility that the female preference of P. sphenops for mustached males could be due to a female pre-existing bias (sensu lato), even if population means were not significant for species other than P. sphenops. This highlights the importance of distinguishing between population means and individual preferences when interpreting mate choice, and thus, adds depth to the concept of mating preferences as a motor for evolutionary change.     

UV wavelengths make female three-spined sticklebacks (<Emphasis Type=Italic>Gasterosteus aculeatus</Emphasis>) more attractive for males

Numerous fishes possess UV vision and show UV patterns, which often play a role in social communication, especially during courtship. UV light is scattered strongly in water and thus might be used for intra-specific close-range communication without being detected by predators. In addition to the red-belly colouration and blue-coloured iris, male three-spined sticklebacks possess UV reflectance on their skin, and females prefer males presented with UV light rather than without. To investigate whether UV also influences male visual preference for females in this species, we used a dichotomous mate choice design in which one female could be viewed by a male in two visual conditions. Courting male sticklebacks preferred females that were presented in full-spectrum conditions including UV to the same females presented in conditions lacking this light component. Furthermore, control trials with neutral-density filters indicated that male preference in the UV treatment was not affected by a difference in achromatic brightness between the UV+ and UV− stimuli. Reflectance measurements of gravid females suggest an enhanced UV contrast between the dark bar pattern, which is characteristic of gravid females, and silvery body parts.     

Reproduction and condition of female Octopus mimus (Mollusca: Cephalopoda)

The relationship between reproduction and condition was studied in a 15 mo sample of 919 maturing, mature and post-spawning female Octopus mimus (388 to 3714 g) caught in Iquique (North Chile). O. mimus is a semelparous species, with reproduction taking place all year round. Investment in reproductive tissues was, on average, 9.9% of mature female body weight, independent of season. However, somatic growth during maturation was dependent on season and varied between 26 and 63% of the whole-body growth in weight. The condition of females did not vary markedly until spawning, although seasonal variations were apparent, winter being the most unfavourable. Condition deteriorated dramatically after spawning, during parental care of the eggs. During this period, somatic-tissue depletion, mainly from the muscles, was >25% of the total body weight of mature females. The fecundity of O. mimus was probably limited by the costs associated with parental care of the eggs.     

Digit ratio (2D:4D) and dominance rank in female rhesus macaques (Macaca mulatta)

Female rhesus macaques exhibit matrilineal dominance structures, and high dominance rank confers fitness benefits across a lifetime and across generations. Rank effects are “inherited” through social processes that are well understood; however, biological mechanisms that might impact these processes are not well known. Recently, it has been shown that prenatal androgens appear to be implicated in supporting dominance rank hierarchies in some mammals. In humans, interindividual differences in the second (index) to fourth (ring) digit ratio (2D:4D) have been linked indirectly to variation in prenatal androgens, with low 2D:4D in both sexes associated with higher inferred prenatal androgen effects. 2D:4D has also been related to dominant social behavior and has been shown to co-vary with social systems across nonhuman primate species. Here, we investigate how 2D:4D co-varies with socially inherited dominance rank in female rhesus macaques. Low 2D:4D was associated with higher-ranking females, while higher 2D:4D was associated with lower-ranking females. Similar relationships were also shown between ranked families within matrilines. This is the first study to show such a relationship between 2D:4D and dominance rank in a nonhuman primate and suggests that prenatal androgen effects could be involved in the maintenance of dominance rank in female cercopithecine primates.