Sex ratio varies with egg investment in the red-necked phalarope (Phalaropus lobatus)

Fisher’s sex ratio theory predicts that on average parents should allocate resources equally to the production of males and females. However, when the cost/benefit ratio for producing males versus females differs, the theory predicts that parents may bias production, typically through underproduction of the sex with greater variation in fitness. We tested theoretical predictions in the red-necked phalarope, a polyandrous shorebird with sex-role reversal. Since females are larger and therefore potentially more expensive to produce and may have greater variation in reproductive success, we predicted from Fisher’s hypothesis a male bias in population embryonic sex ratio, and from sex allocation theory, female biases in the clutches of females allocating more resources to reproduction. We measured eggs and chicks and sexed 535 offspring from 163 clutches laid over 6 years at two sites in Alaska. The embryonic sex ratio of 51.1 M:48.9 F did not vary from parity. Clutch sex ratio (% male) was positively correlated with clutch mean egg size, opposite to our prediction. Within clutches, however, egg size did not differ by sex. Male phalarope fitness may be more variable than previously thought, and/or differential investment in eggs may affect the within-sex fitness of males more than females. Eggs producing males were less dense than those producing females, possibly indicating they contained more yolk relative to albumen. Albumen contributes to chick structural size, while yolk supports survivorship after hatch. Sex-specific chick growth strategies may affect egg size and allocation patterns by female phalaropes and other birds.     

Females prefer larger leks: field experiments with ruffs (Philomachus pugnax)

Summary Preference by females for choosing mates at male aggregations has been hypothesized as the primary selective pressure favoring the formation of leks, but alternative hypotheses account for lek formation without invoking female preference. Observational studies to determine whether male mating success increases with lek size, as predicted under the female preference hypothesis, have produced inconsistent results, possibly due to covariation of lek size with other variables or to male-male or intersexual conflict over lek size. We tested whether females prefer larger leks in a field experiment with ruffs (Philomachus pugnax), a lekking sandpiper, in which male group size, composition, and location were controlled. Wild females chose the larger of two adjacent groups often enough such that males in larger groups had significantly higher per capita rates of female visitation (Table 3). Such behavior would probably lead to higher per male mating rates at larger leks, which is generally considered a necessary condition for female choice to select for lek display (Fig. 2). Lek size in nature will reflect both female preference for larger leks and competition among males, which may favor smaller lek size. All else being equal, however, female ruffs preferred to visit larger groups strongly enough to maintain lekking by males.     

Mate guarding,copulation strategies and paternity in the sex-role reversed,socially polyandrous red-necked phalarope<Emphasis Type="Italic"> Phalaropus lobatus</Emphasis>

In a recent review, Westneat and Stewart (2003) compiled evidence that extra-pair paternity results from a three-player interaction in which sexual conflict is a potent force. Sequentially polyandrous species of birds appear to fit this idea well. Earlier breeding males may attempt to use sperm storage by females to obtain paternity in their mates subsequent clutches. Later-breeding males may consequently attempt to avoid sperm competition by preferring to pair with previously unmated females. Females may bias events one way or the other. We examined the applicability of these hypotheses by studying mating behavior and paternity in red-necked phalaropes (Phalaropus lobatus), a sex-role reversed, socially polyandrous shorebird. Male red-necked phalaropes guarded mates more strongly than other shorebirds. Males increased within-pair copulation attempts during their mates fertile period, and maintained or further increased attempts towards the end of laying, suggesting an attempt to fertilize the females next clutch; these attempts were usually thwarted by the female. Paired males sought extra-pair copulations with females about to re-enter the breeding pool. Multilocus DNA fingerprinting showed that 6% of clutches (4/63) each contained one chick sired by a male other than the incubator, producing a population rate of these events of 1.7% (n=226 chicks). Male mates had full paternity in all first clutches (n=25) and 15 of 16 monogamous replacement clutches. In contrast, 3 of 6 clutches of second males contained extra-pair young likely fathered by the females previous mate. Previously mated female phalaropes may employ counter-strategies that prevent later mating males from discriminating against them. The stability of this polyandrous system, in which males provide all parental care, ultimately may depend on females providing males with eggs containing primarily genes of the incubating male, and not a previous mate.Communicated by M. Webster     

Factors regulating annual mating success and reproductive success in spotted sandpipers (Actitis macularia)

Summary We investigated factors affecting annual mating success (MS) and reproductive success (RS) of spotted sandpipers (Actitis macularia) from 13 years of a 17-year study at Little Pelican Island, Leech Lake, Minnesota. Analyses were restricted to ages 1–3. Mean annual female MS varied from 1.3 to 2.7 mates, and the MS pattern was indistinguishable from random. However, female MS increased with age and was affected by arrival date, territory size, and beach size. Female RS also increased with age, and number of mates and year effects were the most significant explanatory variables in each age. Older female RS was increased by priority on a territory and presence of a previous mate. Territory size and beach size varied with population density and did not predictably affect RS. The strong year affect on RS was associated with annual variation in sex ratio and predation. Males produce only one successful clutch per year, so MS greater than one is a result of nest loss and does not increase RS. Neither male MS nor RS changed with age. Male reproductive failure rate varied by year. Given that a male produced young, the degree of RS was affected by year, arrival date, priority on a territory, territory size, and beach size. In years with early-season predation, late arrivals had higher RS; territory and beach size effects varied by year. Neither the presence, nor degree, of female care was associated with male RS. Male RS was more subject to annual environmental variability than was female RS, probably because of relatively low annual potential RS among males.Offprint requests to: L.W. Oring at the current address     

Conditional lekking in ruff (Philomachus pugnax)

Summary In our study of ruff, a lekking shorebird, we found that the lek ratio — the proportion of the total population of males occurring on leks was low, averaging 12% over the breeding season (Fig. 1D). Off-lek males spent approximately the same proportion of time as lek males in displaying to females (Fig. 4). We defined three spacing tacties that male ruffs use to position themselves to court females: Following — directly pursuing females, and two types of lekking behavior, Intercepting — waiting for females at resource-rich sites, and True Lekking — waiting for females at places without any resources other than the males themselves. Males switched among these tactics, causing the lek ratio to vary over the season (Fig. 1 D). Lek ratio increased when the number of females present in the study area plummeted at the end of May, and was positively correlated throughout the season with the copulation rate of the preceding day, suggesting that males were tracking the behavior, as well as the number, of females available (Table 1, Fig. 1). Early in the season, males off leks spent most of their time feeding (Fig. 4), and lek ratio was positively correlated with temperature (Table 1), suggesting that some males may have been unable to lek during cold weather. Males on leks mated at significantly higher rates than Followers (Table 4). On average, males at our interception lek were less site faithful and less peristent than males at true leks, and the interception lek itself disappeared after females stopped coming to use its adjacent resource (Table 2, Fig. 5). The most successful individuals in our population were the True Lekking males, rather than the Interceptors.In addition to the conditinal lekking tactics described here, ruff display a dimorphism in behaviorat leks. Independent males defend small courts on leks, while Satellite males share courts and mutually display with independents Both independents and satellites may use all three conditional tactics. We propose that satellites evolved as specialized. Followers, adept at tracking the movements of females among leks.     

Interannual differences in the relative timing of southward migration of male and female western sandpipers (Calidris mauri)

In the shorebird subfamily Calidridinae, one of the parents shortens parental care and initiates southward migration before the other. We estimated the difference in passage date between male and female western sandpipers (Calidris mauri) at their first major stopover on the southward migration from breeding areas in Alaska, in 18 years between 1978 and 2000. Overall, adult females preceded adult males by 1.22 days. A novel finding was that among juveniles, which migrate approximately a month later than adults, females preceded males by similar magnitude (1.14 days). There was wide variation among years, however, and males actually preceded females in years with late hatch. We relate these findings to hypotheses for female-first southward migration in sandpipers.     

Providing parental care entails variable mating opportunity costs for male Temminck’s stints

Males of many species theoretically face a fitness tradeoff between mating and parental effort, but quantification of this is rare. We estimated the magnitude of the mating opportunity cost paid by incubating male Temminck’s stints (Calidris temminckii), taking advantage of uniparental care provided by both sexes in this species. “Incubating males” provide all care for an early clutch, limiting subsequent mating possibilities. “Non-incubating” males include males that failed to obtain, lost to predation, or actively avoided incubating clutches. These males were free to pursue extrapair copulations and to mate with females laying later clutches, which females usually incubate themselves. Male incubation classes did not differ in measures of quality, and many individuals changed classes between years, suggesting the use of conditional reproductive tactics. However, specialist non-incubators may also exist. Using microsatellites to assign parentage, we compare males’ total fertilizations and the subset “free of care” fertilizations between incubation classes. Incubators were more likely to gain at least one fertilization per season and averaged one more per season than non-incubators. However, successful non-incubators were more likely to gain “free of care” fertilizations, averaging two more than successful incubators. The relative success of male incubation classes also changed with local sex ratios. With higher female proportions, non-incubators gained disproportionately more offspring, suggesting that the use of tactics should be partly determined by the availability of potentially incubating females. Overall, we estimate the opportunity cost of incubating to be 13–25 % of the potential annual reproductive output.     

Mortality-minimizing sandpipers vary stopover behavior dependent on age and geographic proximity to migrating predators

Ecological theory for long-distance avian migration considers time-, energy-, and mortality-minimizing tactics, but predictions about the latter have proven elusive. Migrants must make behavioral decisions that can favor either migratory speed or safety from predators, but often not both. We compare the behavior of adult and juvenile western sandpipers Calidris mauri during the course of their temporally segregated passages at a major stopover site. Here, the passage and winter arrival of an important predator, the peregrine falcon Falco peregrinus begins near the end of the adult sandpiper passage (July) and increases rapidly through the juvenile passage (August). The mortality-minimizing hypothesis predicts that as the falcon front is distant but approaching, sandpipers should initially increase the fuel-loading rate (lowered vigilance and predator apprehension) to increase migration speed and so maintain their head start. As the falcon front gains proximity to and passes over the stopover site, sandpipers should become increasingly cautious. Our measurements show that adults decreased vigilance during the period prior to falcon arrival, and had lower vigilance overall than juveniles. Juveniles were more apprehensive, flying further and longer in response to disturbance by a falcon silhouette. This trend was reversed in response to a human approach. Both groups were more vigilant and more apprehensive in a study year with earlier falcon arrival. These results suggest that late (juvenile) and early (adult) migrants minimize mortality on migration in different ways, adults by increased migratory speed at the expense of caution on stopover sites, and juveniles by increased caution at the expense of speed.