Experimentally manipulated brood sex ratios: growth and survival in the black-headed gull (Larus ridibundus), a sexually dimorphic species

In sexually size dimorphic species, individuals of the larger sex often suffer from enhanced mortality during the nestling period. This has been attributed to higher nutritional requirements of the larger sex, which may render this sex more vulnerable to adverse food conditions. However, sex-biased mortality might not exclusively depend on the differences in food demand but also on other phenotypic differences, e.g., in competitiveness. Interference competition between the sexes and position in the laying sequence in particular may be essential components contributing to biased mortality.By creating synchronously-hatched unisex broods in the sexually size dimorphic black-headed gull, we specifically tested the effect of sex-specific food demand by excluding interference competition between the sexes as well as hatching asynchrony. To test the effect of egg quality, which varies with the position in the laying sequence, we composed each nest of chicks from eggs of all different positions in the laying sequence.All-male nests showed significantly enhanced mortality compared to all-female nests from the beginning of the development of the sexual size dimorphism onwards. This underlines the role of a higher food demand in biased mortality of the larger sex.In males but not females, asymptotic body mass and skeletal size were negatively associated with position in the laying sequence, while survival was not affected by position. As a consequence, sexual size dimorphism at the end of the nestling period was less pronounced compared to the natural situation. These data show that, although male growth is more sensitive to a decrease in egg quality, the higher mortality of last hatched chicks in natural nests is mainly due to hatching asynchrony and egg size but not egg content.     

Unequal sex ratios and their consequences in Herring Gulls (Larus argentatus)

Summary We observed the sex ratios and age composition of courting, nesting, and nonbreeding birds (in clubs) in four breeding colonies of Herring Gulls in Maine and New York. Breeding pairs, with eggs or chicks, often contained an immature male, but rarely contained an immature female. Similarly, courting pairs contained immature males, but rarely immature females. A higher proportion of courting pairs contained immatures compared with pairs with eggs or chicks. In all four colonies there was an excess of females in the clubs. A simulated colony which assumed unequal survival for males and females had an age ratio for unpaired birds that was similar to the observed values. We conclude that in Herring Gulls, mate competition exists and stems in part from an unequal tertiary sex ratio.     

Parental condition, brood sex ratio and differential young survival: an experimental study in gulls (Larus fuscus)

Empirical evidence is growing that the offspring sex ratio in birds can be biased in relation to the body condition of parents during breeding. The sex ratio bias may come about because (1) the actual production of the two sexes may be skewed and/or (2) there may be a sex bias in early nestling mortality contingent on parental condition. By manipulating parental condition and giving them a control brood to rear, thereby eliminating effects operating via the eggs, we examined the extent to which parental condition influences the post-hatching survival of male and female lesser black-backed gulls, Larus fuscus. We found that the pre-fledging survival of male chicks was strongly reduced in all-male broods reared by parents in poor condition. Pre-fledging survival of female chicks was, however, unaffected by parental condition or brood sex composition. Thus, independently of any production biases, sex differences in nestling mortality alone can bias the offspring sex ratio at fledging in relation to the prevailing rearing conditions. In other studies on gulls we have, however, also shown that females in poor condition at laying preferentially produce female eggs. Clearly a bias in fledging sex ratio can occur within the same species due to a combination of differential production and differential post-laying mortality; the latter can involve a differential effect of poor egg quality on male and female offspring, differential effects of brood sex composition on their survival and a difference in the capacity of parents to rear males and females. All of these processes need to be taken into account in attempting to understand offspring sex ratios. Received: 15 February 2000 / Revised: 7 August 2000 / Accepted: 26 August 2000     

Estimate of removal rate ofNereis virens (Polychaeta: Nereidae) from an intertidal mudflat by gulls (Larus spp.)

The feeding behavior of herring gulls (Larus argentatus), ringed-billed gulls (L. delawarensis) and great blackbacked gulls (L. marinus) on an intertidal mudflat in Maine, USA, was investigated. Remains of fish, mussels, crabs, insects, and the polychaeteNereis virens were recovered from gull feces. Forty-three percent of the fecal samples containedN. virens jaws, setae, or both. A comparison of jaws from fecal material and from worms collected from the natural community demonstrates that gulls preferentially preyed upon larger worms. Feeding was largely confined to 3 h around low tide, and birds fed mostly in the low intertidal and below mean low water where the largest worms were located. Individual birds remained on the flat for a mean of 28.0 min and consumed a mean of 19.2 worms per visit. It was calculated that gulls remove a mean of 808 largeN. virens from the flat per tide from June to October, representing an estimated 0.04% of the standing crop of largeN. virens.     

Inter-sexual differences in T-cell-mediated immunity of black-headed gull chicks (<Emphasis Type="Italic">Larus ridibundus</Emphasis>) depend on the hatching order

Hatching asynchrony in avian species leads to age and size differences between nestlings within a brood, handicapping last-hatched chicks in the sibling rivalry. Starvation due to this competitive disadvantage has been regarded as the primary cause of an increase in mortality with hatching order. However, for gulls it has also been suggested that disease is the cause of mortality for last-hatched chicks, possibly through reduced immunocompetence and thereby an enhanced susceptibility to infection. In addition, the male-biased mortality reported for several gull species may be related to a higher vulnerability to diseases in males compared to females. To determine the potential influence of the immune system on these mortality patterns, we investigated the T-cell-mediated immunity (CMI) of black-headed gull chicks in relation to hatching order and sex. We found a significant decrease in the CMI with hatching order. This result may be causally related to systematic changes in maternal yolk steroids and carotenoids within the laying sequence. For second-laid eggs, male CMI was significantly lower than female CMI. This is possibly linked to higher plasma levels of testosterone in male embryos which might have an immunosuppressive effect. If so, this effect is masked in eggs of either high (first egg) or low (last egg) quality. Chicks with low CMI showed enhanced mortality rates. Thus the differences in immune response are likely to contribute to the observed mortality patterns. However, hatching order significantly affected mortality independently of CMI, suggesting that competitive disadvantage due to hatching asynchrony is also important.Communicated by M. Webster     

Egg coloration in ring-billed gulls (<Emphasis Type="Italic">Larus delawarensis</Emphasis>): a test of the sexual signaling hypothesis

Although many avian eggs appear to be cryptically colored, many species also lay vibrant blue green eggs. This seemingly conspicuous coloration has puzzled biologists since Wallace, as natural selection should favor reduced egg visibility to minimize predation pressure. The sexual signaling hypothesis posits that blue green egg coloration serves as a signal of female quality and that males exert post-mating sexual selection on this trait by investing more in the nests of females laying more intensely blue green eggs. This hypothesis has received mixed support to date, and most previous studies have been conducted in cavity-nesting species where male evaluation of his partner’s egg coloration, relative to that of other females, may be somewhat limited. In this study, we test the sexual signaling hypothesis in colonially nesting ring-billed gulls (Larus delawarensis) where males have ample opportunity to assess their mate’s egg coloration relative to that of other females. We used correlational data and an experimental manipulation to test four assumptions and predictions of the sexual signaling hypothesis: (1) blue green pigmentation should be limiting to females; (2) extent of blue green egg coloration should relate to female quality; (3) extent of blue green egg coloration should relate to offspring quality; and (4) males should provide more care to clutches with higher blue green chroma. Our data provide little support for these predictions of the sexual signaling hypothesis in ring-billed gulls. In light of this and other empirical data, we encourage future studies to consider additional hypotheses for the evolution of blue green egg coloration.     

Sex-specific costs of hatching last: an experimental study on herring gulls (<Emphasis Type="Italic">Larus argentatus</Emphasis>)

An organism’s pattern of development can have important long-term fitness effects. In species where the sexes differ in size or other phenotypic traits, they may also have different optimal developmental rates. This influences both parental sex allocation strategies and susceptibility of the sexes to early developmental conditions. However, sex differences in developmental rate and vulnerability to environment during the embryonic period are not well understood. In birds, sibling competition and hatching asynchrony may select for accelerated embryonic development of the last offspring in order to reduce their competitive disadvantage after hatching. They may advance their hatching in response to vocal stimuli by the older siblings. It is, however, unclear whether this flexibility in developmental rates is sex specific. In this study, we experimentally manipulated between-embryo contact and tested whether this affected the pre-natal developmental rate and post-hatching performance of male and female offspring from last-laid eggs in the herring gull. Post-hatching performance was measured both in competitive and non-competitive situations. Among young incubated in isolation, males hatched faster than females, but both sexes fledged in similar, relatively good condition. Among young incubated with normal between-embryo contact, hatching time did not differ between sexes, but males fledged in poorer condition than females, regardless of whether they were reared singly or in a brood. These results suggest that male and female offspring differ in their ability to mitigate the costs of hatching asynchrony.     

Intra-specific interactions influence egg composition in the lesser black-backed gull (Larus fuscus)

Egg composition, which is under maternal control, can have a profound effect on offspring fitness. The presence of maternal testosterone and carotenoids in avian egg yolk, for example, is thought to enhance the development and competitive ability of the offspring and protect the hatching and growing chick against oxidative stress. Egg quality often differs between females and such variation can be due to differences in maternal social environment, e.g. breeding density. However, this is confounded by the possibility that the quality of individuals breeding in high- or low-density areas may vary. We tested if maternal social environment influences egg composition in a colonial seabird, the lesser black-backed gull (Larus fuscus). To control for confounding effects of female quality, we experimentally manipulated maternal social environment during egg formation. We increased the frequency of intra-specific interactions (i.e. aggressive encounters with conspecifics other than nest mates) in which the females were involved, by placing an elevated platform in their territory. Females that took part in more intra-specific interactions produced a heavier last egg, but the yolk testosterone concentration in eggs laid by control and experimental females did not differ. Differences in yolk testosterone concentration in relation to embryo sex were found neither in the control nor in the experimental group. In contrast, within the control group, eggs with a male embryo contained more carotenoids than eggs with a female embryo. Moreover, experimental females that had been involved in more intra-specific interactions produced female eggs with higher carotenoid levels compared to female eggs of control birds. An experimental increase in carotenoid levels was not observed in eggs containing a male embryo. Our results suggest that intra-specific interactions experienced by female birds during egg formation can influence conditions for embryonic development.Communicated by J. Graves     

Regional differences in behaviour in bank voles (Clethrionomys glareolus): dyadic encounters

North Scandinavian bank voles (Clethrionomys glareolus) differ from south Scandinavian conspecifics both in population dynamics and in activity, social behaviour, density-related sensitivity in sexual maturation and in reproduction. Some ecologists think that cyclically fluctuating rodents are particularly aggressive, at least in certain cycle phases. This hypothesis was tested in dyadic encounters between animals from southern and northern Sweden, respectively. Univariate analyses showed significant differences between regions in four separate behavioural traits, but not in behaviour related to aggression. Laboratory breeding for several generations did not affect the level of aggressivity. Multivariate analyses revealed two main components of activity and sociability, both with regional variation. Activity components (also including “freezing” behaviour) were chiefly related to age while sociability showed mainly regional variation. Differences observed may be due to the geographic location or earlier cyclicity, as the cyclic pattern was weak at the time when the test animals were sampled. An adaptation to high-density situations appears likely as the sympatric field vole (Microtus agrestis) shows similar regional differences although it lives in a different habitat. Cyclic animals appear to disperse at increasing densities while non-cyclic animals show clumped distributions. Received: 19 February 1996/Accepted after revision: 17 August 1996     

Lifetime reproductive success in the spotted sandpiper (Actitis macularia) : sex differences and variance components

Summary Lifetime reproductive success (LRS) of male (N=103) and female (N= 66) spotted sandpipers (Actitis macularia) was studied for 13 years of a 17-year study at Little Pelican Island, Leech Lake, Minnesota. There was no sex difference in longevity, but females had significantly more mates, eggs, chicks, fledged young, and young returning in subsequent years than did males. Variance in LRS was partitioned into five life-history components: longevity (L), mates per year (M), eggs per mate (E), proportion eggs hatched (H), and proportion of chicks fledged (F). For both sexes, F accounted for the greatest proportion of variance in LRS (males, 43%; females, 47%), followed by L (males, 26%; females, 43%) and H (males, 21%; females, 28%). Positive covariance between H and F was consistent with predator-caused clutch and brood loss. Contrary to our expectations, males had a higher coefficient of variation in reproductive success than did females. This was because males were relatively more likely than females to produce no young.Offprint requests to: L.W. Oring at the current address     

Sexual behavior among parasitic Megarhyssa wasps (Hymenoptera: Ichneumonidae)

Summary Male wasps of three sympatric species of Nearctic Megarhyssa (Hymenoptera: Ichneumonidae) typically form mixed-species (61.7%) aggregations composed of up to 28 males over sites of female emergence. Observations of 107 aggregations over two seasons in New York state revealed two male mating strategies: postemergence copulation on the tree surface and preemergence insemination accomplished by male abdominal insertion into the female emergence hole. Insertions by one or more males (as many as ten consecutively but not more than two simultaneously) occurred at 88% of the aggregations, with each inserting for an average of 24.8 min. Insemination as a result of preemergence insertion had a success rate of 83% while postemergence copulation attempts were successful 30% of the time (average duration 61.9 s). Individual male reproductive success was very low and not correlated with size, arrival order at emergence sites, or tenacity at the site. Males frequently aggregated at sites of male emergences as well as those of nonconspecifics, though conspecifics to the emerger ultimately played the major role at aggregations. Male-male aggression in aggregations was not apparent, though jostling occurred during the insertion period and during postemergence copulation attempts.     

Chemical defense and evolutionary trends in biosynthetic capacity among dorid nudibranchs (Mollusca: Gastropoda: Opisthobranchia)

Summary. An evolutionary scenario incorporating recent advances in phylogenetic research begins with an opisthobranch-pulmonate common ancestor that was herbivorous and had some diet-derived chemical defense. The Nudibranchia and their closest relatives, the Notaspidea, form a lineage the ancestors of which had switched to feeding upon sponges and deriving protection from metabolites contained in them. Subsequently there have been repeated shifts in food and defensive metabolites, and trends are evident in the ability to detoxify, sequester and utilize metabolites from food, as well as to synthesize defensive compounds de novo. The Notaspidea display a minor adaptive radiation that foreshadows a more extensive one in the various lineages of nudibranchs. This review emphasizes changes that have occurred within the Holohepatica, or dorid nudibranchs (order Doridacea). Their sister-group, the Cladohepatica, consists of three other orders, Dendronotacea, Arminacea, and Aeolidiacea, in which there has been a shift from sponges to Cnidaria as food. The Dendronotacea often feed upon Octocorallia, which combine spicules, chemical defense, and stinging capsules and thereby suggest a transition from feeding on sponges. A previous diet of Octocorallia is suggested by the defensive use of prostaglandins in the dendronotacean Tethys fimbria, which eats crustaceans. A shift to bryozoans in some Arminacea is accompanied by use of different metabolites. Dorid nudibranchs evidently began as sponge-feeders, but some lineages have shifted to a variety of other food organisms, and others have specialized in the kind of sponges they feed on and how they do it. There have been shifts to bryozoans (Ectoprocta) and ascidians (Chordata: Urochordata) that track metabolites rather than the taxonomy of the food. There is a crude correlation between the genealogy and the defensive metabolites of the sponge-feeding dorids. De novo synthesis is well documented in this order and the metabolites are appropriately positioned so as to have an adaptive effect. The hypothesis that the capacity for de novo synthesis was acquired by gene transfer across lineages is rejected, partly on the basis of different chirality of metabolites in the nudibranchs and their food organisms. Instead it is proposed that there has been a preadaptive phase followed by evolution in a retrosynthetic mode, with selection favoring enzymes that enhance the yield of end products that are already present in the food. Received 5 February 1999; accepted 26 July 1999     

Sediment processing in Capitella spp. (Polychaeta: Capitellidae): strain-specific differences and effects of the organic toxicant fluoranthene

The effects of the widespread polycyclic aromatic hydrocarbon (PAH) fluoranthene on sediment-processing rates in the infaunal polychaete Capitella spp. were investigated by comparing five populations of this deposit-feeding species complex: Capitella sp. I from New York, USA; Capitella sp. M from Milos, Greece; Capitella sp. S from Sylt, Germany; and two unidentified Capitella populations from salmon farm sediments –Capitella population K from Kilmelford, Scotland and Capitella population C from Cranford, Ireland. Replicate worms from each strain were exposed to 0, 10, and 95 μg (g dry wt sediment)⁻¹ fluoranthene (=μg/g fluoranthene) for a period of 16 days. Initial and final wet and dry weights (mg) of worms and worm-specific growth rates (WSGRs) were calculated. Sediment processing was measured as the sum of the total dry weight of pellets produced during the experiment, and we estimated size-specific processing rates (SSPRs) as a measure of sediment processed per mg worm dry weight per day. The five populations of Capitella spp. differed significantly in body size, WSGR, and sediment-processing rates. Capitella sp. I grew faster than all of the other populations. Capitella population C and Capitella population K from fish farm sediments, with the largest body lengths (up to 52.0 ± 27.2 mm), had the highest processing rates, whereas the small Capitella sp. S (up to 17.1 ± 5.6 mm) had the lowest. There were also significant differences in SSPR among populations with Capitella sp. I having a higher SSPR (about 12 × body wt/day) than Capitella population C (about 5 × body wt/day) and Capitella population K (3 × body wt/day). The fluoranthene concentrations used in the present study, while representing moderately to highly contaminated conditions, had only marginal effects on sediment-processing and growth rates of all of the Capitella populations examined. Processing of contaminated sediment by Capitella spp. may be important in the remediation of PAH-contaminated sediment. Received: 16 January 2000 / Accepted: 28 August 2000     

Population differences in gonad and pyloric caeca cycles of the New Zealand seastarSclerasterias mollis (Echinodermata: Asteroidea)

Inshore and offshore populations ofSclerasterias mollis (Hutton, 1872) were sampled on the outer continental shelf off the Otago Peninsula, New Zealand, and their nutritional and reproductive cycles are described from 1985–1986. Histological changes in the gonads are generally typical of other asteroids. The gametogenic cycle takes 12 mo. The gonad and pyloric caeca indices in both females and males had an inverse relationship.S. mollis accumulates nutrients in the pyloric caeca during summer and early autumn. The gonads develop in the autumn and winter. Offshore seastars were much larger and had significantly higher gonad and pyloric caeca indices than inshore individuals. These differences in body sizes and organ indices arise from differences in food availability and/or population density.     

A test of neuroecological predictions using paperwasp caste differences in brain structure (Hymenoptera: Vespidae)

Adaptive brain architecture hypotheses predict brain region investment matches the cognitive and sensory demands an individual confronts. Social hymenopteran queen and worker castes differ categorically in behavior and physiology leading to divergent sensory experiences. Queens in mature colonies are largely nest-bound while workers depart nests to forage. We predicted social paperwasp castes would differ in tissue allocation among brain regions. We expected workers to invest relatively more than queens in neural tissues that process visual input. As predicted, we found workers invested more in visual relative to antennal processing than queens both in peripheral sensory lobes and in central processing brain regions (mushroom bodies). Although we did not measure individual brain development changes, our comparative data provide a preliminary test of mechanisms of caste differences. Paperwasp species differ in the degree of caste differentiation (monomorphic versus polymorphic castes) and in colony structure (independent- versus swarm-founding); these differences could correspond to the magnitude of caste brain divergence. If caste differences resulted from divergent developmental programs (experience-expectant brain growth), we predicted species with morphologically distinct queens, and/or swarm-founders, would show greater caste divergence of brain architecture. Alternatively, if adult experience affected brain plasticity (experience-dependent brain growth), we predicted independent-founding species would show greater caste divergence of brain architecture. Caste polymorphism was not related to the magnitude of queen-worker brain differences, and independent-founder caste brain differences were greater than swarm-founder caste differences. Greater caste separation in independent-founder brain structure suggests a role for adult experience in the development of caste-specific brain anatomy.     

Spermiogenesis and sperm structure in the shrimp Parapenaeus longirostris (Crustacea: Dendrobranchiata): comparative aspects among decapods

Early spermatids of the dendrobranchiate shrimp Parapenaeus longirostris (Lucas, 1846) have a spherical nucleus with large patches of heterochromatin, surrounded by a cytoplasmic mass that contains the conspicuous proacrosomal vesicle. The highly polarized mid spermatid mainly consists of the nuclear region, displaying a discontinuous nuclear envelope, and a large proacrosomal vesicle located at the opposite side of the cell. The most recent spermiogenic transformations primarily concern elongation of the proacrosomal vesicle to form a tapering spike. This results in the typically tack-shaped sperm of natantian decapods. The initial steps of spermiogenesis in the two studied dendrobranchiates prove to be parallel to reptant spermiogenesis in some respects, namely rupture of the nuclear envelope, chromatin decondensation and differentiation of electron-dense regions within the proacrosomal vesicle content. Specifically, whereas the anteriormost condensation gives rise to the operculum in brachyurans, in dendrobranchiates it becomes the apical portion of the spike. Despite an unquestionable morphological similarity between the sperm of carideans and dendrobranchiates, spermiogenesis in both groups displays meaningful differences. Spermatids of caridean shrimps lack a distinct proacrosomal vesicle. In the course of spermiogenesis, the spike arises from aggregated cytosolic materials; hence it is not membrane-bound. Unlike in other decapods, caridean sperm do not undergo a conventional acrosome reaction, since exocytotic events are not involved in this process. The above arguments suggest that, in the Decapoda, separation into three sperm classes is more suitable than the two traditionally accepted classes. The dendrobranchiate and reptant sperm types share a number of spermiogenic and functional features, while the caridean sperm type appears to represent an independent evolutive line with regard to sperm development and function.     

Seasonal modulation of flight speed among nocturnal passerine migrants: differences between short- and long-distance migrants

Migrating birds are expected to fly at higher airspeeds when minimizing time rather than energy costs of their migratory journeys. Spring migration has often been suggested to be more time selected than autumn migration, because of the advantage of early arrival at breeding sites. We have earlier demonstrated that nocturnal passerine migrants fly at higher airspeeds during spring compared to autumn, supporting time-selected spring migration. In this study, we test the hypothesis that seasonal airspeeds are modulated differently between short- and long-distance migrants, because of a stronger element of time selection for autumn migration over long distances. In support of this hypothesis, we demonstrate that the seasonal difference in airspeed is significantly larger (spring airspeed exceeding autumn airspeed by a factor of 1.16 after correcting for the influence of altitude, wind and climb/descent on airspeed) among short-distance compared to long-distance (factor 1.12) migrants. This result is based on a large sample of tracking radar data from 3 years at Falsterbo, South Sweden. Short-distance migrants also tend to fly with more favourable winds during autumn, indicating relaxed time constraints (being able to afford to wait for favourable winds) compared to long-distance migrants. These results indicate surprisingly fine-tuned seasonal modulation of airspeed and responses to wind, associated with behavioural strategies adapted to different levels of time selection pressures during spring and autumn migration.     

Territoriality and monogamy among agile gibbons (Hylobates agilis)

Summary I simulated territorial intrusions by mated pairs, solitary males and solitary females through song playback to investigate the behaviors that agile gibbons (Hylobates agilis) use to maintain territoriality and monogamy. Mated male and female gibbons approached and duetted in response to a simulated territorial intrusion by a neighboring group. Strange solitary females elicited similar approach and singing responses from mated pairs. In contrast, the simulated presence of a solitary male consistently evoked approaches by mated males only. Females and males did not contribute equally to pair movements or to answering songs; males led the majority of approaches, while females initiated all duets. I hypothesize that carrying small, vulnerable infants may inhibit females from leading approaches. These results are consistent with previous studies which suggest that female territoriality prevents male gibbons from becoming polygynous.