Differential deposition of antimicrobial proteins in blue tit (Cyanistes caeruleus) clutches by laying order and male attractiveness

Female birds can influence offspring fitness by varying the relative quantities of egg components they deposit within and between clutches. Antimicrobial proteins (lysozyme, ovotransferrin, and avidin) are significant components of the avian albumen and likely aid in defense of embryos from microbial infection. Within clutches, females may enhance antimicrobial defense of early-laid eggs to protect them from the high risk of infection incurred before the onset of incubation. Among entire clutches, females may invest more resources in young sired by more attractive males because they have higher reproductive value. We tested these hypotheses by quantifying antimicrobial protein distribution within and among clutches in blue tit eggs. Contrary to our hypothesis, clutches showed no differential deposition of lysozyme or avidin within clutches, but eggs laid in the middle of the sequence had higher concentrations of ovotransferrin than eggs in the beginning and end. Consistent with our second hypothesis, we found that females produced eggs with higher concentrations of lysozyme (although not ovotransferrin or avidin) when mated to more attractive (more UV-reflective) males. Furthermore, females mated to polygynous males deposited less lysozyme than those mated to monogamous males. These data suggest that allocation of lysozyme at the clutch level may be a maternal effect mediated by male qualities.     

Brood size manipulation affects frequency of second clutches in the blue tit

The reproductive trade-off hypothesis predicts that the investment made in current reproduction determines the breeders’ future fitness as a consequence of intra-or inter-generational reproductive costs. Long-lived species are expected to favour their own reproductive value at the expense of their offspring, hence incurring in inter-generational costs, whereas short-lived species are expected to invest in the current breeding attempt even at the expense of their own survival, thus incurring in intra-generational costs. We tested whether intensity of current reproductive effort has intra-or inter-generational costs in a short-lived bird, the blue tit Parus caeruleus, with a brood size manipulation experiment. We expected more intra-generational (parental reproduction and/or survival) than inter-generational (offspring quality and survival) reproductive costs. We found that parental effort, measured as the hourly rate of parental visits to nests, increased gradually with experimental manipulation. Brood size manipulation resulted in a gradual increase in the number of fledglings per nest from reduced to increased treatments. We found an effect of the manipulation on the probability of making a second clutch, with adults rearing enlarged broods being less likely to undertake such a second reproduction during the season compared to those rearing control or decreased broods. We found no evidence of other reproductive costs; neither as adult weight after manipulation, apparent parental local survival, apparent offspring local survival or local recruitment. Although the results seem to support the a priori expectations, alternative explanations are discussed.Communicated by M. Soler     

Extra-pair paternity,sperm competition and the evolution of testis size in birds

Sperm competition should select for increased sperm production if the probability of fertilization by a specific male is proportional to the relative number of sperm inseminated. A review of the literature generally supports the predicted positive association between sperm production or allocation and various measures of the presumed intensity of sperm competition. However, it is not clear how increased sperm competition is related to extra-pair paternity, and it remains unknown whether certainty of paternity should be associated with relative testis size. Based on a large sample of bird species with information on extra-pair paternity gathered from the literature, we demonstrate that testis mass is related positively to the level of extra-pair paternity, after controlling for body size and phylogeny. Although large testes may be necessary to avoid sperm depletion in species in which males frequently engage in multi-pair copulations, we argue that selection has favoured increased testis mass in situations of more intense sperm competition in order to retaliate against copulations by rival males. The fact the males are not always successful in retaliating against rival ejaculates further suggests that females may largely control the allocation of paternity in birds and that increased sperm production by males may simply be a male strategy to make the best of a bad situation.     

Search costs influence the spatial distribution,but not the level,of extra-pair mating in tree swallows

Costs of searching for a mate are an important component of models of sexual selection, yet they have rarely been examined in wild populations of vertebrates. In this paper, we report an experiment in which we handicapped female tree swallows by clipping some flight feathers. This manipulation increased the costs of flight and searching for extra-pair mates. Despite these costs, handicapped females had the same level of extra-pair mating (percentage of extra-pair young, percentage of broods with extra-pair young, and the number of extra-pair sires per brood) as control females. However, handicapped females were more likely to have young sired by extra-pair males that lived closer to her nest than control females. This change in the distribution of extra-pair mating was most likely due to female choice rather than male coercion, and it suggests that extra-pair mating has significant benefits to females. One important implication of our study is that ecological and social factors that influence search costs could affect the spatial distribution of extra-pair sires and, consequently, the intensity of sexual selection. These effects may have been overlooked in previous studies that did not identify extra-pair sires.     

Paternal care and egg survival both increase with clutch size in the fathead minnow,Pimephales promelas

Summary In some species of fishes with paternal care, females prefer to spawn with males who are already defeding eggs; moreover, in many species, paternal care increases with the number of eggs that a male is defending. If egg survival depends on the level of paternal care, and is largely independent of egg number, then egg survival should increase with clutch size. This result would provide a potential adaptive mechanism for female preference for males with eggs. I examined the effects of clutch size on paternal care and egg survival in the fathead minnow, Pimephales promelas, and found that both increased with male clutch size.     

Maternal effects on begging behaviour: an experimental demonstration of the effects of laying sequence, hatch order, nestling sex and brood size

Differential resource allocation by females across the laying sequence has been hypothesised as a mechanism through which females could either compensate nestlings that hatch last in asynchronous broods or promote brood reduction. In this study we artificially incubated eggs and cross-fostered offspring to manipulate nestlings’ position in the hatching order, to identify whether the competitive ability of nestlings is dependent on position in the laying sequence. In both control and experimentally reversed broods, first hatched chicks had a higher survival than last hatched siblings. Yet, nestlings that hatched from eggs laid in the second half of a clutch begged with a greater intensity than nestlings hatched from eggs laid in the first half of a clutch. In natural broods, the greater begging competitiveness of nestlings from later-laid eggs led to a moderation of sibling competition and these nestlings achieved the same body size and weight as nestlings from eggs laid in the first half of the clutch. The lack of a substantial difference in the size and condition of surviving nestlings in respect to laying order suggests that differential resource allocation across the egg-laying sequence partially compensates for hatching last in asynchronous broods and reduces the negative effects of the nestling size hierarchy. The effect of laying order, brood size and experimental treatment also differed for male and female nestlings. Our study highlights the need to be aware of the complex and subtle effects of nestling sex and laying sequence when investigating genetic and environmental influences on individual fitness.     

Egg size and laying order in relation to offspring sex in the extreme sexually size dimorphic brown songlark,<Emphasis Type="Italic"> Cinclorhamphus cruralis</Emphasis>

In some bird species, mothers can advantage the offspring of one sex either by elevating them in the laying order to promote earlier hatching or by allocating greater resources to eggs of the preferred sex. In size dimorphic species, the predictions as to which sex should benefit most from such pre-laying adjustments are ambiguous. The smaller sex would benefit from an initial size advantage to help compensate for the faster growth rate of the larger sex. However, an early advantage to offspring of the larger sex might have a greater effect on their lifetime reproductive success than an equivalent advantage to offspring of the smaller sex. We investigated these hypotheses in the polygynous brown songlark, Cinclorhamphus cruralis, which is one of the most sexually size dimorphic birds known. We conducted within-clutch comparisons and found that females hatched from larger eggs and were initially heavier (but not structurally larger) than their brothers. This may afford females an early competitive advantage, as egg volume remained correlated with chick mass until at least 5 days of age. Similarly, we found that hatch order was still positively associated with nestling mass and size when the brood was 10 days of age, but there was no clear relationship between offspring sex and hatching order. During this study, food was plentiful and there were few obvious cases of nestling starvation. When food is limited, we suggest that the greater nutrient reserves of female hatchlings could not only help compensate for their slower growth, but could also give them a survival advantage over their brothers early in the nestling period. Consequently, egg size dimorphism may be an adaptation that facilitates an early shift in brood sex-ratio towards cheaper daughters in conditions of low food availability.     

Patterns,correlates, and paternity consequences of extraterritorial foray behavior in the field sparrow (Spizella pusilla): an automated telemetry approach

Behavioral Ecology and Sociobiology - ᅟMales of fiddler crabs (genus: Uca) construct courtship structures using mud or sand to attract mate-searching females for underground mating. A sensory...     

Roles of male residence and relative size in the social behavior of Iberian rock lizards, Lacerta monticola

Many studies on contest competition used residency asymmetry as a discrete variable. However, the probability of winning an interaction may change as a continuous function of the value of the location where the encounter occurs. We performed a field study to examine the importance of location within a home range and relative body size to the outcomes of agonistic interactions between male lizards, Lacerta monticola. The distances to activity centers (the most used locations based on a density function of sightings) and relative size play important roles in agonistic interactions and had interacting effects in natural conditions. On the other hand, previous studies with lizards suggested that inferior competitors are able to avoid agonistic interactions in the field. Thus, we staged encounters in the laboratory to examine the behavioral responses of smaller individuals. The responses of each focal smaller male were measured in its own home cage (resident), in the cage of a larger male (intruder) and in a cage in which no male was previously present (control). The predominant behavioral tactics of smaller males were avoidance when they are the intruders and displaying when they are the residents. Submissive displays by smaller males may help reduce the costs of agonistic encounters.     

Heterotrophic activity associated with particulate size fractions in a Spartina alterniflora salt-marsh estuary,Sapelo Island,Georgia, USA,and the continental shelf waters

An investigation was undertaken to determine the distribution and activity of the heterotrophic microplankton associated with particles from a Georgia salt marshestuarine ecosystem to the western edge of the Sargasso Sea. Heterotrophic activity was determined by the uptake of ¹⁴C-glucose. More than 80% of the activity was associated with detritus greater than 3 m in creek, river and coastal (within 4 km of shore) waters. In the Gulf Stream, approximately 80% of the heterotrophic activity was in the fraction less than 3 m. In the estuary, total heterotrophic activity fluctuated with the tides; the greatest activity occurred near low ebb tide at all locations. The lowest activity was measured at slack low and high tides. In creek water most of the heterotrophic activity was associated with particles between 14 and 180 m, whereas in coastal waters (less than 4 km from shore) most of the activity was in the fraction greater than 180 m.Contribution No. 331 from the University of Georgia Marine Institute, Sapelo Island, Georgia 31327, USA.     

Sex allocation in a monomorphic seabird with a single-egg clutch: test of the environment, mate quality, and female condition hypotheses

Sex allocation theory posits that mothers should preferentially invest in sons when environmental conditions are favorable for breeding, their mates are of high quality, or they are in good body condition. We tested these three hypotheses in rhinoceros auklets (Cerorhinca monocerata), monomorphic seabirds that lay a single-egg clutch, in 2 years that differed in environmental conditions for breeding. Results supported the environment and mate quality hypotheses, but these effects were interactive: offspring sex was independent of paternal traits in the poor year for breeding, while females mated to larger and more ornamented males reared more sons in the better year. Conversely, offspring sex was unrelated to female condition, as indexed by hatching date. We propose that good rearing conditions enable females to rear sons possessing the desirable phenotypic attributes of their mates. Results also supported two critical assumptions of sex allocation theory: (1) dimorphism in offspring condition at independence: daughters fledged with higher baseline levels of corticosterone than sons and (2) differential costs of rearing sons versus daughters: mothers rearing sons when environmental conditions were poor completed parental care in poorer condition than mothers rearing daughters in the same year and mothers rearing either sex when conditions were better. These novel results may help to explain the disparate results of previous studies of avian sex allocation.     

Chinstrap penguins alter foraging and diving behavior in response to the size of their principle prey, Antarctic krill

Penguins may exhibit plasticity in their diving and foraging behaviors in response to changes in prey availability. Chinstrap penguins are dependent predators of Antarctic krill in the Scotia Sea region, but krill populations have fluctuated in recent years. We examined the diet of chinstrap penguins at Livingston Island, South Shetland Islands, in relation to their diving and foraging behavior using time-depth recorders over six breeding seasons: 2002–2007. When krill were smaller, more chinstrap penguins consumed fish. In these years, chinstrap penguins often exhibited a shift to deep dives after sundown, and then resumed a shallower pattern at sunrise. These night dives were unexpectedly deep (up to 110 m) and mean night dive depths sometimes exceeded those from the daytime. The average size of krill in each year was negatively correlated to mean night dive depths and the proportion of foraging trips taken overnight. Based on these patterns, we suggest that when krill were small, penguins increasingly targeted myctophid fish. The average krill size was negatively correlated to the time chinstrap penguins spent foraging which suggests that foraging on smaller krill and fish incurred a cost: more time was spent at sea foraging.     

Extra-pair paternity and the evolution of testis size in a behaviorally monogamous tropical mammal,the large treeshrew (<Emphasis Type="Italic">Tupaia tana</Emphasis>)

Monogamy is rare in mammals (<5% spp.) but occurs in greater frequency among primates (15%) and their close relatives, the treeshrews (100%; Order: Scandentia). Two genetic studies of parentage in monogamous primates revealed high rates of extra-pair paternity (EPP), but to date parentage has not been studied in a treeshrew species. I analyzed the genetic parentage of 22 offspring from two populations of large treeshrews in Sabah, Malaysia (NE Borneo), using seven autosomal microsatellite loci and one mitochondrial DNA marker. Half of these offspring were sired by males that were not the presumed partner of the mother (50% EPP), and three litters exhibited evidence of multiple paternity. However, comparative analysis indicated that the high rate of EPP in Tupaia tana is not associated with intense sperm competition. Relative testis size of treeshrews was similar to testis size in 22 primate species with uni-male mating systems but smaller than 44 primates with multi-male mating systems. After factoring out the effects of body size and phylogeny, I also found that the evolution of multi-male mating systems was significantly associated with the evolution of larger testis size. Male–female pairs of T. tana occupy joint territories but forage and sleep alone (“dispersed pair-living”), and I argue that this form of behavioral monogamy renders mate guarding ineffective. The adaptive advantages of behavioral monogamy likely differ from the advantages driving EPP in large treeshrews. However, small testis size suggests that behavioral monogamy is not masking a dispersed multi-male mating system in this species.     

Diel variation in the abundance of epifauna associated with seagrasses of the Indian River,Florida, USA

Day-night comparisons of the motile epifauna in the canopy of seagrass beds in the Indian River lagoon, Florida revealed that abundances were not stable over the diel period. Collections were made between February and June, 1982. Higher densities of epifauna occurred on seagrasses at night. For both of the numerically-dominant gastropod and crustacean groups, four of the five top ranked species followed this diel pattern. Crustaceans showed particularly large changes in abundance, averaging a twofold increase in nocturnal samples. Faunal similarity between day and night collections remained high, however, indicating that community structure was preserved during diel abundance changes. Equating epifauna collected from seagrass blades with those associated with the seagrass habitat in general therefore may be erroneous, particularly if collections are made during daylight. Changes in prey availability due to the diel pattern are likely to extend to important community trophic functions.     

Carotenoids in the egg yolks of collared flycatchers (<Emphasis Type="Italic">Ficedula albicollis</Emphasis>) in relation to parental quality,environmental factors and laying order

Birds may influence the fitness of their offspring by transmission of different amounts of carotenoids to their eggs. Carotenoids play crucial roles in antioxidant protection and immune defence mechanisms, but they may be available to females in limiting amounts. Therefore, their allocation to the eggs may be influenced by the female’s condition, age and environmental circumstances. Furthermore, the quality of the male parent, which affects the reproductive value of the offspring, may also influence this investment. In this correlational study, we investigated proximate and ultimate factors that may lead to variation in yolk lutein, zeaxanthin and β-carotene concentrations among and within clutches of a wild passerine, the collared flycatcher (Ficedula albicollis). We found that carotenoid concentration was positively associated with caterpillar supply at the time of egg formation, which suggests a proximate constraint of carotenoid availability on yolk composition. Neither female condition, body size, age, nor male plumage ornamentation, age and body size correlated with carotenoid deposition. Yolk β-carotene concentration was found to be positively linked to yolk testosterone concentration. We suggest that females allocated more β-carotene to their eggs to mitigate the potentially detrimental effects of elevated steroid concentration. We found that concentration of β-carotene increased with laying order. The possible function of this pattern may be to enhance the resistance to oxidative stress and pathogens of the disadvantaged last-hatching nestling, suggesting that collared flycatchers pursue a compensatory, “brood survival” strategy.     

High degree of multiple paternity in the viviparous Shiner Perch, <Emphasis Type="Italic">Cymatogaster aggregata</Emphasis>, a fish with long-term female sperm storage

The Shiner Perch (Cymatogaster aggregata Gibbons) exhibits a viviparous reproductive mode and long-term female sperm storage, two biological features that may predispose this fish species for both intense sperm competition and frequent multiple paternity within broods. To test these hypotheses, we used polymorphic microsatellite markers to identify sires and quantify paternal contributions to the progeny arrays of 27 pregnant females from a natural population of C. aggregata. The number of sires per brood ranged from one to eight (mean 4.6), typically with skewed distributions of fertilization success by the fathers but no correlation between sire number and brood size. The extraordinarily high incidences of multiple paternity in this species probably are due in part to high rates of mate encounter, but selection pressures related to the avoidance of maternal–fetal incompatibility may further have promoted the evolution of polyandrous mating behaviors in this female-pregnant species. Our genetic data are consistent with the hypothesis that viviparity, long-term sperm storage, and extreme polyandry are interrelated reproductive phenomena that should promote the evolution of post-copulatory sperm competition and/or cryptic female choice in these fishes.     

Effect of pair-bonding with adults on the ontogenetic manifestation of aggressive behavior in a primate,Saguinus fuscicollis

Summary Injurious aggression, scent marking and some visual displays shown in situations analogous to territorial defense were studied in young saddleback tamarins who had lived with nonrelated adult and juvenile social partners from 6 months of age under the experimental conditions outlined below. Twenty male and 20 female tamarins were removed from their natal families when 6 months old, and were used to form pairs and trios of the following composition: (1) one juvenile female and one juvenile male (10 pairs); (2) one juvenile female and one adult male (4 pairs); (3) one juvenile female, one juvenile male, and one adult male (6 trios); (4) one adult female and one juvenile male (4 pairs). These groups cohabited permanently and when the subjects were between 10 and 12 months old, their social interactions with adult conspecific strangers were studied, using series of 10-min social encounter tests. During these tests injurious aggression (attacks, fights, chases), a number of visual hostile displays (threat face, ruffle, arch) and scent marking were recorded.Young males and females living with adults of the opposite sex showed more injurious aggression against strangers and scent marked more frequently than young subjects living with partners of their own age. Males living with adult females also showed more facial threat than males living with young females (Figs. 1 and 2). Young males living in trios with adult males and young females scent marked less frequently than males living only with a young female. Aggression, scent marking and facial threat shown by subjects living with adult sex partners were at the level of socially and sexually experienced, pair-bonded adults (Fig. 3). No differences in ruffling and arching scores were found. The injurious aggression scores of the subjects were correlated with those of their mates. These results show that tamarins can fulfill the behavioral roles of pair bonded adults and also breed (Epple and Katz 1980) at an unexpectedly early age, when they live under the influence of an adult sex partner. Young animals cohabiting with an adult of the same sex, on the other hand, show a trend toward inhibition of adult-like behaviors.     

Biokinetics of plutonium,americium and californium in the marine isopod Cirolana borealis,with observations on its feeding and molting behavior

In a series of laboratory experiments carried out during 1982, the biokinetics of ²³⁷Pu (IV), ²³⁷Pu (V), ²⁴¹Am (III), and ²⁵²Cf (III) were studied in the benthic marine isopod Cirolana borealis Lilljeb. Following a 3 wk exposure to labelled seawater, isopods reached non-equilibrium concentration factors of 52±15, 54±12, 176±14, and 185±53 for Pu (IV), Pu (V), Am and Cf, respectively. Uptake of these radionuclides occurred to a great extent by surface adsorption, with the major fraction of the accumulated radionuclides being associated with exoskeleton and lesser percentages located in gut, digestive gland, muscle and haemolymph. The pattern of radiotracer depuration was in all cases biphasic, with the long-lived retention compartment retaining the largest fraction of whole-body radioactivity. Biological half-lives for radionuclide turnover in the long-lived compartment were 60±3 d, 87±5 d, 261±12 d and 288±60 d for Pu (IV), Pu (V), Am and Cf, respectively. Examination of radionuclide distribution among tissues following longterm depuration showed that the radionuclides were eliminated from internal tissues more rapidly than from the exoskeleton. Uptake and depuration studies indicated that the biokinetics of americium and californium were similar to each other, but that they differed significantly from those of oxidized and reduced plutonium. In general, these experimental results support previous reports that americium and californium are more bioavailable than plutonium in the marine waters. The fraction americium and californium assimilated into tissue from food was very low and did not exceed 5%. Unassimilated labelled food was often retained in the gut for more than 2 mo and released only infrequently in a loosely bound condition. In the event of contamination, this particular feeding-digestion strategy in highly mobile, scavenging isopods could act as a potential biological mechanism for the widespread dispersion of radioactive wastes.     

Simultaneous behavior of skipjack (Katsuwonus pelamis), bigeye (Thunnus obsesus), and yellowfin (T. albacares) tunas, within large multi-species aggregations associated with drifting fish aggregating devices (FADs) in the equatorial eastern Pacific Ocean

Ten separate experiments monitoring the simultaneous behaviors of 26 skipjack (Katsuwonus pelamis), 26 bigeye (Thunnus obesus), and 33 yellowfin (T. albacares) tunas within large multi-species aggregations associated with drifting fish aggregating devices (FADs) were investigated using ultrasonic telemetry in the equatorial eastern Pacific Ocean. Experiments were conducted during a research cruise aboard a chartered purse seine vessel. Purse seine sets were made on the tuna aggregations associated with FADs at the termination of six of the ten experiments. Seventeen of the 44 tagged tunas were not recaptured indicating the transient nature of the associative behavior of tunas with FADs. Although there was considerable overlap in the depths of the three species, by day and night, there were some species-specific differences and diel differences within species. While we documented spatial and temporal differences in the schooling behavior of the three tuna species, the differences do not appear sufficient such that modifications in purse seine fishing practices could effectively avoid the capture of small bigeye and yellowfin tunas, while optimizing the capture of skipjack tuna in purse seine sets on FADs.