Copulation duration,but not paternity share,potentially mediates inbreeding avoidance in Drosophila montana

Studying the incidence of inbreeding avoidance is important for understanding the evolution of mating systems, especially in the context of mate choice for genetic compatibility. We investigated whether inbreeding avoidance mechanisms have evolved in the malt fly, Drosophila montana, by measuring mating latency (a measure of male attractiveness), copulation duration, days to remating, offspring production, and the proportion of offspring sired by the first (P₁) and second (P₂) male to mate in full-sibling and unrelated pairs. SNP markers were used for paternity analysis and for calculating pairwise relatedness values (genotype sharing) between mating pairs. We found 18 % inbreeding depression in egg-to-adult viability, suggesting that mating with close relatives is costly. Copulation duration was shorter between previously mated females and their brothers than with unrelated males. Based on an earlier study, shorter copulation is likely to decrease the number of inbred progeny by decreasing female remating time. However, shorter copulations did not lead to lower paternity (P₂) of full-sibling males. Progeny production of double-mated females was lower when the second male was a full-sibling as compared to an unrelated male, but we could not distinguish between inbreeding depression and lower female reproductive effort after mating with a relative. Relatedness estimates based on 34 SNPs did not detect any quantitative effect of relatedness variation on copulation duration and progeny production. We suggest that inbreeding depression has been strong enough to select for inbreeding avoidance mechanisms in our Finnish D. montana population.     

Disturbance influences oyster community richness and evenness, but not diversity

Foundation species in space-limited systems can increase diversity by creating habitat, but they may also reduce diversity by excluding primary space competitors. These contrasting forces of increasing associate diversity and suppressing competitor diversity have rarely been examined experimentally with respect to disturbance. In a benthic marine community in central California, where native oysters are a foundation species, we tested how disturbance influenced overall species richness, evenness, and diversity. Surprisingly, overall diversity did not peak across a disturbance gradient because, as disturbance decreased, decreases in overall species evenness opposed increases in overall species richness. Decreasing disturbance intensity (high oyster abundance) led to increasing species richness of sessile and mobile species combined. This increase was due to the facilitation of secondary sessile and mobile species in the presence of oysters. In contrast, decreasing disturbance intensity and high oyster abundance decreased the evenness of sessile and mobile species. Three factors likely contributed to this decreased evenness: oysters reduced abundances of primary sessile species due to space competition; oysters supported more rare mobile species; and oysters disproportionately increased the relative abundance of a few common mobile species. Our results highlight the need for further studies on how disturbance can differentially affect the evenness and richness of different functional groups, and ultimately how these differences affect the relationship between overall diversity and ecosystem function.     

Extra-pair paternity in eastern bluebirds: effects of manipulated density and natural patterns of breeding synchrony

The causes of variation in rates of extra-pair paternity among avian populations remain unclear, but could include environmental factors such as breeding density and synchrony. By experimentally manipulating nest site availability, we tested the effects of breeding density on the frequency of extra-pair paternity in eastern bluebirds (Sialia sialis). We also examined the role of breeding synchrony on extra-pair paternity using natural timing of nests. Microsatellite analysis revealed 34 of 305 nestlings (11.2%) were the result of extra-pair fertilizations; and 21 of 79 broods (26.6%) had at least one extra-pair nestling. Several measures of breeding density had independent effects on extra-pair paternity. First, experimental plot type affected extra-pair paternity, with 28 of 34 (82.4%) extra-pair young from nests in high density areas, and only six (17.6%) from nests in low density areas. Independently of plot type, the number of breeding neighbors within a 320-m radius was a significant predictor of the likelihood of extra-pair paternity at the nest. Extra-pair paternity was associated with temporal factors such as absolute timing of breeding and natural levels of local breeding synchrony, but only in bivariate comparisons. We found a positive interaction between density of neighbors within a 320-m radius and local breeding synchrony; this term reduced the main effects of synchrony and number of neighbors, but not experimental treatment. Our results demonstrate the importance of utilizing multiple aspects of proximity in breeding density analyses and testing for interactions between ecological factors that can influence the behavioral events leading to extra-pair fertilizations.     

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.     

Extrapair copulations in western bluebirds (Sialia mexicana): female receptivity favors older males

. Extrapair paternity has been the focus of numerous studies of socially monogamous birds, but the dynamics of extrapair male-female interactions have been explored in relatively few. In western bluebirds (Sialia mexicana), a species with 20% extrapair paternity, two to four observers monitored extrapair copulations (EPCs) occurring on the female's home territory while her social mate was detained in a cage near the nest. Detention elevated the frequency and visibility of EPCs, so we could score receptivity of females to identifiable extrapair males without the interference of intermale aggression. Most intruding extrapair males were neighboring breeders visiting outside their own mate's fertile period. Local male relatives of the detained male attempted nonincestuous EPCs, but neighboring sons did not attempt to copulate with their mothers and, in some cases, chased away intruders. Female receptivity was not influenced by whether the detained mate was visually present or hidden behind cloth and did not increase with the persistence or relative size of the extrapair male. Female receptivity increased with the age difference between the extrapair male and the female's social mate, suggesting that females prefer older males as extrapair partners. This study suggests that costs of resistance are relatively unimportant in determining female receptivity in western bluebirds, but identifies preferential receptivity to older males as a potential factor in the outcome of inter- and intrasexual conflict over paternity.     

Trichinella spiralis infection in voles alters female odor preference but not partner preference

Females may choose mates based on secondary sex traits that reflect disease resistance. Accordingly, females should be able to distinguish between unparasitized and parasitized males, and should prefer to mate with unparasitized individuals. Mate and odor preferences for uninfected males or males infected with the nematode, Trichinella spiralis, were examined among prairie voles (Microtus ochrogaster) and meadow voles (M. pennsylvanicus). In a 15-min odor preference test, only female meadow voles distinguished between bedding from parasitized and unparasitized conspecific males, and preferred to spend time with bedding from unparasitized males. Although T. spiralis infection influenced odor preference in female meadow voles, there was no effect of infection status on mate preference among either species. Testosterone and corticosterone concentrations were not different between parasitized and unparasitized males. However, among prairie voles, males that spent an increased amount of time with females during the mate preference test had elevated testosterone concentrations. Taken together, these data suggest that (1) female meadow voles can discriminate between unparasitized and parasitized males, (2) the effects of infection on steroid hormone concentrations may be masked by the effects of social interactions, and (3) parasites may represent a selective constraint on partner preference in voles; however, the life cycle of parasites may influence female preference and should be considered in studies of female preference. Received: 23 April 1998 / Accepted after revision: 25 October 1998     

Growth rate and hatching date in ostrich chicks reflect humoral but not cell-mediated immune function

A tradeoff between immune response and life history traits, in particular growth rate, has been documented in various bird species. Ostriches are fast-growing birds and a typical feature of cohorts is that offspring often differ greatly in size. We investigated the relationship between hatching date and growth rate of chicks and both cell-mediated (measured using a phytohaemagglutinin (PHA) injection) and humoral immune responses in ostrich chicks maintained on a research farm. Chicks with higher growth rates had intermediate responses to both diphtheria and tetanus toxoids. By contrast, no relation between growth rates and responses to PHA injection were found. We conclude that chick growth rate variation may be explained beyond a certain threshold by a tradeoff between the humoral response and growth. Both responses to PHA injection and humoral responses in chicks were found to decrease with chick hatching date. Within the context of ostrich farming, these results could partially explain size variations observed in cohorts of chicks, as well as high mortality rates during their first 3 months of age.     

Males are selective too: mating, but not courtship, with sequential females influences choosiness in male field crickets (Gryllus bimaculatus)

Mate choice experiments have generally focused on female choice; few have considered that males can also be selective. We examined courtship in male field crickets sequentially introduced to four females of differing size. Large (L) and small (S) females were introduced in order of either LSLS or SLSL. We demonstrate that naive males invest equally (courtship effort) in the first female they meet, regardless of her size, but show greater courtship effort when they subsequently encounter large compared with small females. Moreover, we demonstrate that males show this discriminatory behavior when they are permitted to mate with females (i.e. use a spermatophore) but are less choosy when the female is removed before spermatophore transfer.     

Replacement males in the western bluebird: opportunity for paternity, chick-feeding rules, and fitness consequences of male parental care

In western bluebirds (Sialia mexicana), most pairs remain together for life and share equally in post-hatching parental care. We removed resident males of socially monogamous pairs during laying and after clutch completion to examine chick-feeding rules used by replacement males and current and future fitness consequences of paternal care. Replacement males were not infanticidal and 7 (47%) fed nestlings. Feeding replacement males and the females they joined fed at rates similar to controls. Females without a feeding replacement male compensated by feeding more themselves so that overall feeding rates were not compromised, but they reduced their brooding time. Unlike assisted females, unassisted females exhibited reduced nesting success and their 14-day-old chicks weighed less than controls. Field metabolic rates of unassisted females were 17% higher than those of control females, but the difference was not statistically significant. Older females were better able to raise young without the male's help than were yearling females. Female condition was not affected by male provisioning and unassisted females were as likely to survive to the next breeding season as assisted females. We found no future benefits of provisioning by replacement males; those that fed were no more likely to breed with the female on her subsequent attempt than were males that did not feed, and subsequent clutch sizes were not reduced for females rearing young without the male's help. These experiments suggest that male parental care increases nesting success in western bluebirds and that replacement males use an all-or-none rule to determine whether or not to feed chicks: if they are present during the fertile period they feed at typical rates; if they are not, they usually do not feed at all. Because chick-feeding by males is tied to opportunity for paternity, influences success in the current nest, and does not affect the male's future breeding success, it appears to be parental rather than mating effort. Received: 8 May 1998 / Accepted after revision: 23 July 1998     

Predator type, not body condition, influences positioning within whirligig groups

The anti-predator responses of an animal may depend on the species of predator attacking. For example, ground-dwelling monkeys respond with different warning calls in response to attacks from hawks from above vs attacks from leopards at the side. In grouping animals, it has usually been assumed that the best anti-predator behavior against most predators is to move toward the center of a group. However, little work has been done to study positioning of individuals in response to different predators and whether it might be influenced by gender or body condition. An optimality model is developed that gives predictions for how the needs of predator avoidance, drafting, and foraging could lead to one optimal location within the group. We then conducted a laboratory experiment with individually marked whirligig beetles (Coleoptera: Gyrinidae). We exposed them to an aerial (bird) or aquatic (fish) attack and determined changes in their distance to the front of groups at different speeds of water. We found that female whirligigs occupied the front of a group in response to a fish attack in fast water, but the back after an aerial bird attack, while the reverse was true for males. In slow water, different results were found; in response to a bird attack, females were more likely to be found in the front compared with males. Body condition was not correlated to the front/back position of either males or females before or after attacks in our study.     

Female control of offspring paternity in a western population of red-winged blackbirds (Agelaius phoeniceus)

Extra-pair copulations, which occur when individuals that have formed social relationships to breed copulate outside their pairbond, now are recognized as an important component of reproductive success in many species. In situations where both males and females benefit from extra-pair copulations without incurring much risk, an inevitable conflict arises between pairbonded mates. In this study I investigated the conflict of interest between male and female reproductive strategies in a western population of red-winged blackbirds (Agelaius phoeniceus). Female red wings in this population initiate extra-pair copulations, which resulted in a 35% rate of extra-pair fertilization. Females initiated the majority (78%) of extra-pair copulations away from their nesting territory where pairbonded individuals typically copulate, and females that engaged in extra-pair copulations spent a significantly greater amount of time off the marsh during peak fertilization compared to females that did not. In addition, females that nested in areas with a large number of potential extra-pair partners produced significantly more extra-pair fertilized young compared to females that nested on marshes with few male neighbors. Males’ strategies to protect paternity were limited primarily to patrolling territory boundaries and to opportunistically preventing extra-pair copulations off the marsh when they were visible. In this population females appear to use behavioral means to control nestling paternity, which in turn directly affected their mate’s reproductive success, and males were restricted to using strategies that were largely ineffective at preventing the threat of extra-pair paternity. Received: 23 December 1994/Accepted after revision: 17 December 1995     

The high frequency of extra-pair paternity in tree swallows is not an artifact of nestboxes

A common criticism of nestbox studies is one of creating artificial nesting conditions and breeding behavior different from what would be seen under natural conditions. We assessed the frequency of extra-pair paternity (percentage of broods with at least one extra-pair young) in 25 families of tree swallows (Tachycineta bicolor) nesting in natural cavities and compared it to that in a nestbox population. We found that 84% of females nesting in natural cavities obtained fertilizations from extra-pair males. These extra-pair males fathered 69% of all nestlings. Studies of tree swallows breeding in nestboxes have shown that 50–87% of broods contained extra-pair young, with extra-pair males fathering 38–53% of all the young. In broods with extra-pair paternity, natural cavities contained a significantly greater proportion of extra-pair young than did nestboxes. Despite differences in nesting habitat and female age structure, the frequency of extra-pair paternity did not differ significantly between the natural-cavity and nestbox populations. Therefore, the presence of extra-pair paternity in tree swallows is not an artifact of nestboxes or of artificial nesting conditions. Received: 2 May 1995/Accepted after revision: 14 January 1996     

Increase in Hair Lead, but not Blood Lead Content of Occupationally-Exposed Workers

This study attempted to investigate whether increased hair lead concentrations due to environmental exposure are accompanied by parallel increases in blood lead concentrations. A sample set consisting of both occupationally-exposed individuals and persons from the general population was investigated. The lead content of scalp and pubic hair from the general population (n=189), traffic police personnel (n=27) and battery workers (n=22) were analysed. The hair samples were taken from the nape of the neck by cutting 23 cm closest to the scalp and cleaned using a washing procedure developed in our laboratory. The effectiveness of the washing procedure was partly determined by examining the scanning electron micrographs of the hair samples. Venous samples of blood were taken from each volunteer's arm.The traffic police and battery plant operators sampled had significantly higher average scalp hair and pubic hair lead levels than the non occupationally-exposed general population. No significant differences were found between the blood lead values of these workers and the general population. Compared to the general population both occupationally-exposed groups had significantly more symptoms of lead toxicity.Lead particles were still observed on the hair shafts after the washing procedure. The amount of such lead contamination was difficult to quantify, the problem being further exacerbated by the difference in hair texture of the persons sampled. This exogenous contamination therefore detracts from the usefulness of hair as an indicator of lead exposure.     

Declining extra-pair paternity with laying order associated with initial incubation behavior,but independent of final clutch size in the blue tit

Although functional explanations for female engagement in extra-pair copulation have been studied extensively in birds, little is known about how extra-pair paternity is linked to other fundamental aspects of avian reproduction. However, recent studies indicate that the occurrence of extra-pair offspring may generally decline with laying order, possibly because stimulation by eggs induces incubation, which may suppress female motivation to acquire extra-pair paternity. Here we tested whether experimental inhibition of incubation during the laying phase, induced by the temporary removal of eggs, resulted in increased extra-pair paternity, in concert with a later cessation of laying, in blue tits (Cyanistes caeruleus). As expected, experimental females showed a more gradual increase in nocturnal incubation duration over the laying phase and produced larger clutches than controls. Moreover, incubation duration on the night after the first egg was laid predicted how extra-pair paternity declined with laying order, with less incubation being associated with more extra-pair offspring among the earliest eggs in the clutch. However, incubation duration on this first night was unrelated to our experimental treatment and independent of final clutch size. Consequently, the observed decline in extra-pair paternity with laying order was unaffected by our manipulation and larger clutches included proportionally fewer extra-pair offspring. We suggest that female physiological state prior to laying, associated with incubation at the onset of laying, determines motivation to acquire extra-pair paternity independent of final clutch size. This decline in proportion of extra-pair offspring with clutch size may be a general pattern within bird species.     

The natives are restless, but not often and mostly when disturbed

The argument that the threat posed by introduced species is overblown is often buttressed by the observation that native species sometimes also become invasive. An examination of the literature on plant invasions in the United States shows that six times more nonnative species have been termed invasive than native species, and that a member of the naturalized nonnative pool is 40 times more likely than a native species to be perceived as invasive. In the great majority of instances in which a native plant species is seen as invasive, the invasion is associated with an anthropogenic disturbance, especially changed fire or hydrological regime, livestock grazing, and changes wrought by an introduced species. These results suggest that natives are significantly less likely than nonnatives to be problematic for local ecosystems.     

Ammonium, but not nitrate, stimulates an increase in glutamine concentration in the haemolymph of Tridacna gigas

The concentration of glutamine in Tridacna gigas haemolymph increased >35-fold following exposure to sea water supplemented with ammonium (20 μM), but no increase was observed with nitrate (20 μM). Lack of a diel cycle, no decrease in haemolymph glucose levels, the expression patterns of glutamine synthetase in zooxanthellae and host, and the lack of glutamine release in response to nitrate supplementation all support the proposition that the increase in haemolymph glutamine is a product of the host and not the zooxanthellae. Unlike ammonium, nitrate accumulates rapidly in the haemolymph. It has no effect on the concentration of glutamine in the haemolymph, but there is an increase in arginine, histidine and lysine in the haemolymph, suggesting the release of these essential amino acids from zooxanthellae. Glutamine synthetase (GS) activity decreased markedly in the gill and less so in the mantle over a period of 6 d exposure to elevated ammonium (20 μM). In contrast, GS activity in zooxan- thellae doubled. The response of zooxanthellae in situ was confirmed by incubating freshly isolated zooxanthellae for 4 d in ammonium, which resulted in a ten-fold increase in GS activity. Comparison of the in situ response of zooxanthellae with that obtained in vitro indicates that the symbionts are likely to be exposed to ammonium concentrations lower than that found in the haemolymph. Received: 14 November 1997 / Accepted: 28 April 1998