Experimentally induced polygyny in a monogamous bird species: prothonotary warblers and the polygyny threshold

Summary Hypotheses regarding evolution of polygyny were tested using prothonotary warblers (Protonotaria citrea), through experimental manipulations of habitat quality. Two experiments were performed. Experiment 1 involved supplying nest boxes in different breeding habitats: flooded riparian areas (high food availability) and adjacent dry bottomland (low food availability). Nine bigamous matings were induced in this experiment, all occurring in flooded habitats even though monogamous mating opportunities existed in dry areas. Costs of polygyny for secondary females in flooded habitat were similar to costs incurred by monogamous females in dry habitat. Thus, a polygyny threshold apparently existed, possibly based on differences in food abundance in the two habitats. However, male quality covaried with habitat quality, as males in flooded areas were older and larger than males in dry habitat. Secondary females may have chosen polygyny in the best breeding situation, a combination of male and territory quality. In Experiment 2 nest box density was varied within flooded habitat to allow males to monopolize different numbers of nest-sites. Monogamous females settled earlier on territories with a large number of nest-sites, and males that defended many nest-sites were more likely to become polygynous. Male physical characteristics were not related to occurrence of polygyny in flooded habitat. Limitation of suitable nest-holes ultimately constrains occurrence of polygyny in prothonotary warblers.     

Sperm allocation in relation to male traits, female size, and copulation behaviour in freshwater crayfish species

Sperm competition is a well-recognised agent in the evolution of sperm and ejaculate structure, as well as variation in female quality. Models of the evolution of ejaculate expenditure predict that male body condition, female fecundity and the risk and intensity of sperm competition may be the ultimate factors shaping optimal ejaculate size. We investigated sperm allocation in Austropotamobius italicus, a freshwater crayfish exhibiting a coercive mating system and external fertilisation, in relation to male and female traits and copulation behaviour under laboratory conditions. We found that mating males were sensitive to female size and produced larger ejaculates when mating with larger females, which were more fecund in terms of number of eggs produced. We found no evidence for female egg production being sperm-limited, as the number of eggs was not dependent on male sperm expenditure. Copulation duration and number of ejaculations reliably predicted the amount of sperm transferred, and both these behavioural measures positively covaried with female body size. These results indicate that male freshwater crayfish can modulate their sperm expenditure in accordance with cues that indicate female fecundity. In addition, a novel finding that emerged from this study is the decrease in sperm expenditure with male body size, which may either suggest that large, old male crayfish are better able than small males to economise sperm at a given mating to perform multiple matings during a reproductive season, or that they experience senescence of their reproductive performance.     

The importance of biparental care in a precocial,monogamous bird,the bar-headed goose (Anser indicus)

Summary This study presents the first direct evidence of benefit derived from the male's parental effort during rearing of young in a monogamous, precocial bird species. We compared mothers accompanied by a mate (paired mothers) with single mothers, some of which were artificial widows, in a semicaptive flock of bar-headed geese (Anser indicus). Paired mothers were less often displaced by other birds and suffered fewer interruptions of brooding. Their goslings spent more time grazing and survived with a higher probability. Although one beneficial result of the male's parental effort may be the continued availability and survival of his mate, a more direct benefit arises from higher offspring survival between hatching and fledging.     

Incubation start and egg size in relation to body reserves in the common eider

Avian incubation is often initiated before all eggs are laid. In altricial birds this has been proposed to facilitate brood reduction through asynchronous hatching. However, in precocial birds eggs normally hatch synchronously even if incubation has started before all eggs are laid. Patterns of incubation start may be the adaptive trait selected for both in altricial and precocial species. Several hypotheses have been proposed to explain the timing of incubation start in birds. Decreasing egg-size after incubation start may be adaptively related to an early incubation start, either to ensure synchronous hatching or to decrease fitness cost of late hatched eggs. We have measured individual body condition, egg size and start of incubation in common eider Somateria mollissima, a precocial sea-duck which does not feed during the incubation period. Females in poor body condition start to incubate earlier in the laying sequence than those in good body condition. Furthermore females in poor body condition lay smaller final eggs than females in good body condition. The laying of smaller eggs late in the sequence is therefore probably related to energetic or nutritional state. We propose that females in poor body condition start to incubate early to shorten the nest period in order to reduce their mass loss, but at the cost of reduced size and growth of the ducklings from the eggs laid after incubation start. Females in good body condition on the other hand postpone incubation start at the cost of a longer incubation period and a higher mass loss to the benefit of synchronized hatching and a higher survival of ducklings.     

Macrophyte species composition and structure along littoral region in relation to limnological variables of a tropical wetland ecosystem

The composition and structure of macrophyte communities of the littoral region of Bhindawas wetland (India) was investigated. Two hypotheses were tested: (1) if limnological variables such as water chemistry and soil parameters are important determinants of species composition and abundance and (2) what is the variance in species richness and composition within the wetland. Twenty-eight macrophyte species belonging to 17 different families were identified with relatively low species diversity and richness (H′?=?2.09; Margalef index?=?1.12). Eichhornia crassipes, attained the highest relative frequency (22.69%), followed by Salvinia natans, Paspalum paspaloides, Phragmites karka, Cynodon dactylon and Cyperus alopecuroides. The first and second axis of CCA ordination explained 38.4% and 28.5% of the total variance of species–limnological variables relationship, respectively. Water-related variables (dissolved oxygen, pH and nitrate concentration) and soil nutrient factors (soil carbon and phosphorus levels) were significantly correlated to macrophyte distribution. Sampling sites formed two distinct groups based on cluster analysis, indicating varying ecological conditions. Generally, composition and abundance of macrophytes evince highly productive nature of the lake, while dominance of invasive species like E. crassipes and S. natans indicates high influence of increased nutrient levels and anthropogenic activities in the catchment on lake macrophyte composition.     

Fecundity and survival in relation to resistance to oxidative stress in a free-living bird

Major life history traits, such as fecundity and survival, have been consistently demonstrated to covary positively in nature, some individuals having more resources than others to allocate to all aspects of their life history. Yet, little is known about which resources (or state variables) may account for such covariation. Reactive oxygen species (ROS) are natural by-products of metabolism and, when ROS production exceeds antioxidant defenses, organisms are exposed to oxidative stress that can have deleterious effects on their fecundity and survival. Using a wild, long-lived bird, the Alpine Swift (Apus melba), we examined whether individual red cell resistance to oxidative stress covaried with fecundity and survival. We found that males that survived to the next breeding season tended to be more resistant to oxidative stress, and females with higher resistance to oxidative stress laid larger clutches. Furthermore, the eggs of females with low resistance to oxidative stress were less likely to hatch than those of females with high resistance to oxidative stress. By swapping entire clutches at clutch completion, we then demonstrated that hatching failure was related to the production of low-quality eggs by females with low resistance to oxidative stress, rather than to inadequate parental care during incubation. Although male and female resistance to oxidative stress covaried with age, the relationships among oxidative stress, survival, and fecundity occurred independently of chronological age. Overall, our study suggests that oxidative stress may play a significant role in shaping fecundity and survival in the wild. It further suggests that the nature of the covariation between resistance to oxidative stress and life history traits is sex specific, high resistance to oxidative stress covarying primarily with fecundity in females and with survival in males.     

Extra-pair paternity and the opportunity for sexual selection in a socially monogamous bird (Dendroica petechia)

We used multi-locus DNA fingerprinting to characterise the genetic mating system of the socially monogamous yellow warbler (Dendroica petechia). Over 2 years there were no instances of brood parasitism, but 59% of families (n = 90) contained extrapair sired young and 37% of offspring (n = 355) were of extra-pair paternity. Most hypotheses for extra-pair mating in monogamous species assume a paternity benefit to extra-pair sires, and focus on the benefit(s) to females. However, the assumption of male benefit has been little tested. Among yellow warblers, known extra-pair sires were just as likely to be cuckolded as any male in the population, and there was at least one reciprocal exchange of extra-pair paternity. Nevertheless, among known extra-pair sires, the paternity gains from extra-pair paternity were, on average, greater than the losses in their own families. These results show there is a paternity benefit to certain males. However, the benefit is not absolute but relative and therefore more difficult to measure. The results also suggest that patterns of extra-pair fertilisation are not determined by female choice alone. Most confirmed extra-pair mates were territorial neighbours, but some resided as far as three territories apart, and greater spatial separation was implied in other cases. Thus, the opportunity for extra-pair mating is great. We estimate that as a result of extra-pair fertilisations, variance in male mating success is increased somewhere between 3-fold and 15-fold over that which would result from within-pair reproduction alone. These findings affirm the potential importance of extra-pair reproduction for sexual selection in monogamous species and they support earlier suggestions that extra-territorial forays by male yellow warblers are for the purpose of extra-pair mating.     

Ingestion rate,assimilation efficiency and energy balance in Mytilus chilensis in relation to body size and different algal concentrations

Basing on a quantification of filtration, ingestion, assimilation, biodeposition, excretion and respiration rate, energy budgets were established in Mytilus chilensis Hupé in relation to body size and three different food concentrations of the unicellular green alga Dunaliella marina. The present quantifications revealed that in M. chilensis the ingestion rate only increases slightly with an increase in food concentration which, however, is counterbalanced by a significant decrease in assimilation efficiency in such a way that assimilation rate finally is nearly constant and independent of the food concentrations tested. The quantifications of these results are given by the a-values of the general allometric growth equation P=aW^b relating the energy disposable for growth and reproduction (P; cal d^-1 to body size (W; dry-tissue wt, g). The best energy budget was obtained at the lowest food concentration tested (0.8 mg algal dry wt l^-1; at 12°C and 30 S) with an a-value of 58.8, while the energy budget at the highest food concentration (2.14 mg l^-1) was only slightly lower with an a-value of 49.8; the b-values were 0.49 and 0.51, respectively. The net growth efficiencies (K₂) decreased with increasing body size (from 20 mg to 3 000 mg drytissue wt) from 76.7 to 47.9% at the lowest food level and from 72.6 to 44.0% at the highest food level tested. These relatively high net growth efficiencies seem to reflect optimal experimental conditions. Furthermore, by a comparison of estimated growth (calculated on the basis of the best energy budget) with growth actually quantified in culture raft mussels in the south of Chile during the highest production period of the year, it is obvious that the energy budgets established really reflect the conditions experienced by the mussels in their natural environment.This research was supported by grants S-80-3 and C-80-1 of the D.I.D.-UACH, by CONICYT, Found. Volkswagenwerk, Found. Fritz-Thyssen, by the GTZ, DFG and by the DAAD     

Plasticity in size and age at maturity in a monogamous fish: effect of host coral size and frequency dependence

The number and maximum body size of the gobioid fish, Paragobiodon echinocephalus, increase with the size of its obligate host coral, Stylophora. Only the largest two individuals breed monogamously in each coral head, and the reproductive success of each spawning is positively correlated with body size. In this study, the plasticity in size and age at maturity in P. echinocephalus was examined. We analyzed life history data from gobies 15–20 mm TL (total length) at their initial marking. Gobies found in larger corals were of lower rank in size order and began to breed later at a larger size, usually upon moving to other corals. The size at maturity ranged widely from 17.2 to 36.0 mm TL. After maturation, growth rates decreased. Mortality, however, was not affected by the timing of maturation. The host coral size did not affect the growth and mortality of marked fish, but the mortality rate of juveniles prior to marking appeared to be higher in smaller corals. The estimated lifetime reproductive success did not differ between the gobies inhabiting corals of different size. Thus the plasticity in size and age at maturity in this species may be maintained by frequency-dependent selection in choosing a host coral size that affects an individual’s social status. Received: 5 April 1995/Accepted after revision: 18 February 1996     

Neophobia and social tolerance are related to breeding group size in a semi-colonial bird

In semi-colonial species, some individuals choose to breed in isolation while others aggregate in breeding colonies. The origin and the maintenance of this pattern have been questioned, and inherited phenotype dependency of group breeding benefits has been invoked as one of the possible mechanisms for the evolution of semi-coloniality. Using field observations and behavioural tests in the semi-colonial barn swallow (Hirundo rustica), we tested the hypothesis that breeding group size is related to personality. We measured neophobia (the fear and avoidance of new things) and social tolerance of adults and showed that these two independent traits of personality are strongly related to breeding group size. The biggest colonies hosted birds with higher neophobia, and larger groups also hosted females with higher social tolerance. This parallel between group size and group composition in terms of individual personality offers a better understanding of the observed diversity in breeding group size in this species. Further studies are, however, needed to better understand the origin of the link between individual personality and group breeding strategies.     

Pre-reproductive survival in a tropical bird and its implications for avian life histories

The factors that affect survival until reproduction are essential to understanding the organization of life histories within and among species. Theory predicts, for example, that survival until reproduction influences the optimum level of reproductive investment by parents, which might partly explain prolonged parental care in species with high first-year survival. Tests and refinements of life-history theory have been hampered, however, by a lack of field-based estimates of pre-reproductive survival, especially for tropical species, which have been the subject of many comparative analyses. Tropical species are predicted to have higher first-year survival and delayed reproduction compared to Northern Hemisphere species. We estimated survival until reproduction, age at first reproduction, and sources of variation in juvenile survival in a Neotropical passerine, the Western Slaty-Antshrike (Thamnophilus atrinucha), in central Panama. We observed that fledged antshrikes had 76% survival through the dependent period and 48% survival to the age of 1 year; survival rate was lowest during the first week after leaving the nest. Timing of fledging within the breeding season, fledgling mass, and age at dispersal influenced survival, while sex of offspring and year did not. Individuals did not breed until two years of age, and post-fledging pre-reproductive survival was 41% of annual adult survival. High survival until reproduction in antshrikes balanced their low annual productivity, resulting in a stable population. Survival during the post-fledging period of dependence and the first year of independence in the Western Slaty-Antshrike exceeded estimates for Northern Hemisphere species. This difference appears to be associated with the extended post-fledging parental care, delayed dispersal, low costs of dispersal, and the less seasonal environment of antshrikes.     

Anaerobic heat production of bivalves (Polymesoda caroliniana andModiolus demissus) in relation to temperature,body size,and duration of anoxia

Anaerobic heat-production rates of two co-occurring species of estuarine bivalves (a clam and a mussel) were measured with double-twin heat-flow calorimeters, one at 20°C, the other at 30°C. There is no significant difference between the two species in metabolic rates. There is evidence of initial aerobic metabolism in some individuals, as shown by high initial rates exponentially decreasing with time, while others had fluctuating but stable average metabolic activity from the beginning. During aerobic as well as anaerobic metabolism, the bivalves showed rhythmic periods of activity and quiescence. The two species differed in their rhythmic pattern of active and resting metabolism. In the case ofPolymesoda caroliniana, periods of resting metabolism tend to be longer and periods of active metabolism shorter at 30°C than at 20°C. There is a similarity between thermograms ofModiolus demissus at 20° and 30°C. Following acute temperature changes from 5° to 20° and 30°C, the bivalves showed stable metabolic rates in a matter of hours. The stabilized average rates [pooled averages for both species of 1.34×10^-4 (standard error of the mean=0.17×10^-4) W g^-1 dry weight of tissue at 20°C and 2.10×10^-4 (SE=0.20×10^-4) W g^-1 at 30°C] signify a temperature coefficient (Q₁₀) of 1.56 between 20° and 30°C, or partial temperature acclimation. Subtracting heat production as a result of physical activity, i.e., considering only resting metabolism, the corresponding means and standard errors of the means are 1.24×10^-4 and 0.14×10^-4 W g^-1 at 20°C and 1.91×10^-4 and 0.077×10^-4 W g^-1 at 30°C. Anaerobic heat production rate at 20°C is proportional to body size (r=0.84, 9 degrees of freedom, DF). ForM. demissus, measured anaerobic heat production is on the order of 7.5% of the level of aerobic respiration reported in the literature.     

Group size and ectoparasitism affect daily survival probability in a colonial bird

Little is known in general about how group size or ectoparasitism affect survival in colonial animals. We estimated daily within-season survival probabilities for nesting adult and recently fledged juvenile cliff swallows (Petrochelidon pyrrhonota) at 239 colonies from 1983 to 2003 in southwestern Nebraska, USA. Some colonies had been fumigated to remove ectoparasites. We conducted mark-recapture at each colony site to estimate daily survival. There were no systematic differences between males and females in daily survival. Adults and juveniles occupying parasite-free colonies had, on average, 4.4% and 62.2% greater daily survival, respectively, than their counterparts in naturally infested colonies. Daily survival of all birds increased with colony size for both parasite-free colonies and those under natural conditions, although the effect was stronger for adults at fumigated sites and for juveniles. Average daily survival probability for adults tended to increase during warmer and drier summers. Although daily survival varied at some sites over the course of the nesting cycle, there were no strongly consistent within-year temporal effects on survival. Even small differences in daily survival probability can translate into large effects on mean lifespan. The deleterious effects of ectoparasites on daily survival within the season represent a previously unknown cost of ectoparasitism. The increase in within-season survival with colony size reflects the net effects of many costs and benefits associated with colony size. Ectoparasitism is probably the most important cost that tends to partly balance the positive effects of large colonies. The greater survival of cliff swallows in the larger colonies is a previously unknown advantage of colonial nesting.Communicated by P. Heeb     

Oxygen uptake,ammonia excretion and phosphate excretion by krill and other Antarctic zooplankton in relation to their body size and chemical composition

Oxygen uptake, ammonia excretion and phosphate excretion were measured in 14 Antarctic zooplankton species, including various size classes of krill (Euphausia superba), during a cruise to the Antarctic Ocean adjacent to Wilkes Land in the summer of 1980. Elemental composition (C, N and P) was also determined on the specimens used in these metabolic rate measurements. The values obtained for C, N and P were 4.7 to 47.5%, 1.2 to 12.5% and 0.09 to 1.23% of dry weight, respectively. Regression analyses of metabolic rates on different measures of body weight (fresh, dry, C, N and P) were made on krill, salps and other zooplankton as arbitrarily defined groups and also on the combined groups to determine the best measure of body weight for intra- and interspecific comparison of metabolic rates. The correlations were highly significant in all regressions, although no common measure of body weight provided the best correlation for the three groups of animals. Except for the regression of ammonia excretion on C and N weight, all other regressions of metabolic rates and body weights were significantly different within these three groups. In the combined group, oxygen uptake and ammonia excretion were better correlated to C and N weights than to dry and P weights. For phosphate excretion in the combined groups, dry weight gave the best correlation. Despite these results, the choice of a particular measure of body weight was shown to be important in a comparison of the rates between krill and salps because of their widely different chemical compositions. Our results of rate measurements are compared with those of previous workers for some Antarctic zooplankton, particularly krill. Some of the previous data are in good agreement with ours, while others are not. Possible contributing factors are considered in the latter case. The ratios between the rates (O:N, N:P and O:P) fell within the general ranges reported for zooplankton from different seas. The O:N ratio was consistently low (7.0 to 19.8, by atoms) in all species, suggesting the importance of protein in their metabolic substrates. Protein-oriented metabolism was also supported by the results of C and N analyses which indicated no large deposition of lipid in these animals. From the results of metabolic rate measurements and elemental analyses, daily losses in bodily C, N and P for Antarctic zooplankton in summer were estimated as 0.4 to 2.8%, 0.6 to 2.5% and 1.3 to 19.4%, respectively. These values are approximately one order of magnitude lower than those reported for subtropical and tropical zooplankton.