Plasma levels of androgens and gonadal development of breeders and helpers in the bell miner (Manorina melanophrys)

Plasma concentration of androgens and gonad development were studied in wild cooperatively breeding bell miners (Manorina melanophrys) of different age, sex and social status. Plasma levels of androgens increased with age of birds. For sexually mature (SM) bell miners, androgen levels were higher in breeders than in nonbreeding (NB) helpers (P < 0.005), and they were marginally higher in females than in males (P < 0.10). Thus female breeders showed the highest plasma levels of androgens. This pattern may be explained by an asymmetry between the sexes in the levels of competition needed to reach and defend breeding status; the degree of intra-sexual competition seems to be higher among female bell miners than among males. In general, male breeders had larger gonads than male SMNB helpers; male breeder gonads produced spermatozoa whereas most helpers' gonads did not. However, old (i.e. 34 months) SMNB male helpers had large gonads producing spermatozoa in spite of having low plasma levels of androgens. We suggest that young male bell miners may be under a voluntary mechanism of reproductive suppression perhaps favoured by inbreeding avoidance; although some aggressive interactions observed between very young helpers (about 2 months) and adult members of the breeding unit suggests that a certain degree of imposition also occurs at young ages. Older male helpers may avoid aggression with the male breeder and imposed reproductive suppression, hence the low circulating levels of androgens, but they have the potential to sneak extra-pair copulations. Gonad size decreased during the non-breeding season in both sexes.     

Co-operative breeding by the Australian Bell Miner Manorina melanophrys Latham: A test of kin selection theory

Summary A population of Bell Miners Manorina melanophrys cosisted of two social units (clans), each consisting of one or more monogamous paris and numerous helpers. Individuals of a breeding pair occupied one home range, while non-breeding individuals occupied partially overlapping home ranges. Attendants (i.e. parents and helpers) showed clear preferences as to whom they aided (fed) and how much aid they provided. A significant correlation was found between the amount of aid given by an attendant and the genetic relationship between the attendant and the recipient. This provides strong quantitative evidence for the theory of kin selection. Only 14% of eggs laid produced juveniles which lived longer than 3 months. It is suggested that the extremely high juvenile mortality favours individuals which help relatives, even while such individuals have dependent offspring of their own.     

Sociality and cooperative breeding of red-cockaded woodpeckers,Picoides borealis

Summary Thirty groups of red-cockaded woodpeckers (Picoides borealis) were studied from 1976–1982 to (1) determine the demographic structure of groups, (2) identify the role helpers play in reproductive activities, and (3) investigate the selective pressures promoting sociality and helping behavior. Groups had only 1 mated pair and 0–2 helpers. Approximately half of all groups had helpers and a given group had helpers some years but not others. Helpers, with rare exception, were males 1 or 2 years old and progeny of 1 or both members of the breeding pair. As a result of higher nestling survival, groups with helpers fledged significantly more young per year than unassisted pairs ( , SD=0.97,n=43 vs. , SD=1.01,n=50). Nesting success was also associated with size and quality of nesting period home range, but evidence suggested that the increased number of young fledged resulted directly from the action of helpers. There was a significant positive correlation between reproductive success and experience of breeding females among unassisted pairs but a significant negative correlation among pairs with helpers. In groups with experienced females, helpers were assisting both their mothers and fathers and, therefore, were related to the offspring on the average by 0.50. In groups with inexperienced females, helpers were assisting their fathers and unrelated females and were related to the offspring by 0.25. The red-cockaded woodpecker's unique habit of excavating nest and roost cavities in living pines and the extended period of time required for excavation may be an important ecological constraint that promotes the retention of helpers. Because helpers are related to the offspring they help rear, kin selection and gains in indirect fitness may provide a partial explanation of why red-cockaded woodpecker helpers help. However, the negative correlation between the efficacy of helping behavior and the helpers' relatedness to the offspring they help rear implies that helpers are least effective in producing offspring which would represent greatest gains to indirect fitness. This raises questions about the relative importance of kin selection and indirect fitness in the evolution of helping behavior among red-cockaded woodpeckers.     

Cooperatively breeding carrion crows adjust offspring sex ratio according to group composition

In cooperatively breeding species, parents should bias offspring sex ratio towards the philopatric sex to obtain new helpers (helper repayment hypothesis). However, philopatric offspring might increase within-group competition for resources (local resource competition hypothesis), diluting the benefits of helper acquisition. Furthermore, benefits of offspring sex bias on parents’ fitness may depend on different costs of production and/or different breeding opportunities of sons and daughters. Because of these counteracting factors, strategies of offspring sex allocation in cooperative species are often difficult to investigate. In carrion crows Corvus corone corone in northern Spain, sons are more philopatric and more helpful at the nest than daughters, which disperse earlier and have higher chances to find a breeding vacancy. Consistent with the helper repayment hypothesis, we found that crows fledged more sons in groups short of subordinate males than in groups with sufficient helper contingent, where daughters were preferred. Crow females also proved able to bias primary sex ratio, allocating offspring sex along the hatching sequence in a way that provided the highest fledging probability to sons in the first breeding attempt and to daughters in the following ones. The higher cost of producing male offspring may explain this pattern, with breeding females shifting to the cheapest sex (female) as a response to the costs generated by previous reproductive attempts. Our results suggest complex adjustments of offspring sex ratio that allowed crows to maximize the value of daughters and sons.     

Sex ratio,monogamy and breeding success in the Midas cichlid (Cichlasoma citrinellum)

Summary The Midas cichlid (Cichlasoma citrinellum) is a large, aggressive monogamous fish from the Great Lakes of Nicaragua. It lives in an ecosystem where breeding sites are at a premium and where breeding success can be low. I tested the strength of the pair bond by manipulating the sex ratios in captive groups, using ratios of 1:1, 2:1, 4:1, 1:2 and 1:4 (:). No polygamy was seen at any of the ratios. Departure from a ratio of 1:1 decreased breeding success for those pairs that formed. Both sexes appeared to enforce the pair bond but did so in different ways: females attacked males and females equally but males attacked other males significantly more than they did females. These results, coupled with field observations, indicate the Midas cichlid is obligated to monogamy by the demands of competition for breeding sites, the need for two parents to protect the fry and by energetic constraints.     

Intra-group relatedness affects parental and helper investment rules in offspring care

In any system where multiple individuals jointly contribute to rearing offspring, conflict is expected to arise over the relative contributions of each carer. Existing theoretical work on the conflict over care has: (a) rarely considered the influence of tactical investment during offspring production on later contributions to offspring rearing; (b) concentrated mainly on biparental care, rather than cooperatively caring groups comprising both parents and helpers; and (c) typically ignored relatedness between carers as a potential influence on investment behavior. We use a game-theoretical approach to explore the effects of female production tactics and differing group relatedness structures on the expected rearing investment contributed by breeding females, breeding males, and helpers in cooperative groups. Our results suggest that the breeding female should pay higher costs overall when helpful helpers are present, as she produces additional offspring to take advantage of the available care. We find that helpers related to offspring through the breeding female rather than the breeding male should contribute less to care, and decrease their contribution as group size increases, because the female refrains from producing additional offspring to exploit them. Finally, within-group variation in helper relatedness also affects individual helper investment rules by inflating the differences between the contributions to care of dissimilar helpers. Our findings underline the importance of considering maternal investment decisions during offspring production to understand investment across the entire breeding attempt, and provide empirically testable predictions concerning the interplay between maternal, paternal and helper investment and how these are modified by different relatedness structures.     

Offspring sex ratios reflect lack of repayment by auxiliary males in a cooperatively breeding passerine

The repayment hypothesis posits that primary sex ratios in cooperative species should be biased towards the helping sex because these offspring “repay” a portion of their cost through helping behavior and therefore are less expensive to produce. However, many cooperatively breeding birds and mammals do not show the predicted bias in the primary sex ratio. Recent theoretical work has suggested that the repayment hypothesis should only hold when females gain a large fitness advantage from the presence of auxiliary adults in the group. When auxiliaries provide little or no fitness advantage, competition between relatives should lead to sex ratios biased towards the dispersing (non-helping) sex. We examined the benefits auxiliaries provide to females and corresponding offspring sex ratios in the red-backed fairy-wren (Malurus melanocephalus), a cooperatively breeding Australian bird with male auxiliary helpers. We found that auxiliaries provide little or no benefit to female reproductive success or survival. As predicted, the population primary sex ratio was biased towards daughters, the dispersing sex, and females with auxiliaries produced female-biased broods whereas females without auxiliaries produced unbiased broods. Moreover, offspring sex ratios were more strongly biased toward females in years when auxiliaries were more common in the population. These results suggest that offspring sex ratios are associated with competition among the non-dispersing sex in this species, and also that females may use cues to assess local breeding opportunities for their offspring.     

Siblicide,parent-offspring conflict,and unequal parental investment by egrets and herons

Summary O'Connor (1978) generated behavioral predictions from Trivers's (1974) theory of parent-offspring conflict (POC) in relation to avian brood reduction, especially for species using escalated sibling aggression. In general, when daily mortality rates fall between certain values (the mortality difference thresholds for parents and surviving offspring: O'Connor 1978), POC is expected. At such times parents and the early-hatching senior offspring are expected to disagree on the necessity, or at least timing, of the victim chick's death. Because that death is promoted directly by the senior chicks and because parents have realistic means of nullifying those efforts, behavioral conflict is expected. I examined the activities of all family members in one siblicidal (great egrets, Casmerodius albus) and one nonsiblicidal brood-reducing species (great blue herons, Ardea herodias) of ardeidae in Texas nesting colonies. Daily mortality rates over two breeding seasons approximated the predicted POC zone for four-chick (B/4) heron broods and three-chick (B/3) egret broods. As predicted, egret sibling aggression was significantly lower in two-chick broods. Herons seldom fought, regardless of brood size. Special attention was paid to the egret B/3 data for behavioral manifestations of POC. Parents did not interfere overtly with sibling fights (99.2% of 2,829), but parents seemed to have a subtle mollifying effect on the aggression (fewer sever fights occurred when parents were present). Parents showed no favoritism in food distribution to dying victim chicks. The mixed results are consistent with two rather different POC interpretations: (1) that significant conflict exists and the offspring are winning or (2) that there is no significant evolutionary conflict. In the latter case, parents are regarded as creating asymmetries among offspring (e.g., via hatch asynchrony), then leaving competition among siblings to run its course without further intervention (laissezfaire policy). This view explicitly recognizes that siblicide and other investment-skewing selfish chick behaviors may serve parental interests well-i.e., that equal allocation of resources is not likely to be the parental optimum. Using the relative reproductive values of each nestling as an index of the parent's unequal parental investment optimum produces a closer fit to the observed food distribution skew, but does not resolve the question of whether conflict remains.     

Cooperatively breeding superb fairy-wrens show no facultative manipulation of offspring sex ratio despite plausible benefits

We report a long-term study of offspring sex ratios in the cooperatively breeding superb fairy-wren Malurus cyaneus. Detailed study of this species had revealed a suite of potentially strong selection pressures on the sex ratio. First, females gain substantial fitness benefits from the presence of helpers; so females without male helpers would benefit from any strategy that increased the probability of recruiting help, such as overproduction of sons (local resource enhancement hypothesis), but large numbers of helper males compete among themselves, favouring the production of daughters (local resource competition). Second, daughters fledged early in the season have far greater chances of recruitment to the breeding population than late-fledged daughters, so mothers would benefit from production of daughters early in the breeding season (early bird hypothesis). Third, extra-group mate choice imposes strong sexual selection on males, suggesting that females mating with attractive sires could benefit from investing in sons (sexual selection hypothesis). However, the predictions from these and other sex ratio hypotheses were rejected. The only convincing evidence for manipulation of the sex ratio was a slight bias towards sons (11 sons to 10 daughters) that occurred regardless of context. This result does not support current theory.     

A Preliminary Study on Soil–Gas Radon Geochemistry According to Different Bedrock Geology in Korea

Three study areas of Kwanak campus (Seoul National University), Boeun (Choongbuk) and Gapyung (Kyonggi) were selected and classified according to their bedrock types in order to investigate soil–gas radon concentrations. The mean values of radon concentrations decreased in the order of Gapyung (40BqL^–1) > Kwanak campus (30BqL^–1) > Boeun (22BqL^–1), and decreased in the order of granite gneiss > banded gneiss > granite > black slate–shale > mica schist > shale–limestone > phyllite schist according to bed rock types. Variation in radon emanation with water content in soils and with soil grain size was assessed by the modified Morse (1976) 3min method. Soil–gas radon concentrations increased with increasing water content in the range of less than about 6–16wt.%, but decreased above 6–16wt.%. Radon concentrations also increased with decreasing soil grain size. Radioactivity analysis of radionuclides of ²³⁸U series in some soil samples indicated their possible radioactive disequilibrium between ²²⁶Ra and ²³⁸U due to the differing geochemical behaviour of intermediate radionuclides. Thus, a radioactive isotope geochemical approach should be necessary for soil–gas radon study.     

Flexible helper structure as an ecological adaptation in the pied kingfisher (Ceryle rudis rudis L.)

Summary The pied kingfisher has two types of helpers at the nest: primary ones, helping their own parents, and secondary ones, helping birds other than parents. Primary helpers are always accepted by breeding pairs, secondary helpers only in poor environmental conditions where the time and energy budget of parents is not sufficient for rearing the offspring alone. Under these conditions, helpers increase the breeding success of pairs (Tables 2 and 3) by providing additional food for the young (Table 4). Thus the flexible helper structure can be seen as an ecological adaptation. It is argued that-originating from a skewed sex ratio and breeding in colonies-this adaptation evolved through the combined effects of individual and kin selection.     

Clutch loss affects the operational sex ratio in the strawberry poison frog Dendrobates pumilio

Both modelling and field data from three breeding seasons show that an environmental factor, clutch loss (CL), affects the operational sex ratio (OSR) and therefore male mating frequency in strawberry poison frogs. Clutch loss affects the length of reproductive cycles of both sexes: with increasing clutch losses, males spend proportionately more time than females in parental investment activities. Because of this, males spend relatively less time in the mating pool, i.e. exhibit proportionately more time-out than females in comparison to a situation with low or no clutch loss. Hence, clutch loss leads to a less male-biased OSR, coupled with a decrease in the opportunity for sexual selection. Furthermore, this study resolves an apparent paradox, the negative correlation between mating frequency and reproductive success (=number of produced tadpoles) of individual males in one breeding season. Clutch loss decouples the correlation between mating frequency and reproductive success because females re-enter the mating pool when they lose their offspring. However, clutch loss diminishes the reproductive output. Similar consequences of clutch loss on the OSR may be true for many species where both sexes reproduce frequently in one breeding season.Communicated by J. Christensen-Dalsgaard     

Parentage analysis of multi-male social groups of tasmanian native hens (Tribonyx mortierii): genetic evidence for monogamy and polyandry

Accurate estimates of male reproductive success are essential to understanding the evolution of polyandrous mating systems. Here, we use multilocus DNA fingerpinting to assess parentage in an island population of Tasmanian native hens (Tribonyx mortierii), which often live in multi-male and/or multi-female social groups. This isolated population presented special challenges to this technique because it was artificially founded from a small number of individuals in the recent past. DNA profiles from four multilocus minisatellite probes were analyzed for adults and offspring from six social groups using two methods: (1) significance of band-sharing coefficients and (2) distribution among a group's offspring of fragments unique to certain adults. Traditional band-sharing analyses did not provide sufficient resolution to establish parentage in this population due to the high level of band-sharing between adults within groups. In contrast, the distribution of unique fragments suggests that in most cases, all offspring within a group have the same male and female genetic parents, so that monogamy may be the predominant genetic mating system of this species. This forces a rexamination of the evolutionary basis of polyandry in these birds. It also demonstrates some of the difficulties in using these highly polymorphic genetic markers for parentage analyses when putative parents are closely related.     

The “red tides” of Malta

A 5 year survey of Marsamxetto Harbour, Malta, was carried out to investigate some factors contributing to red tides and to identify the causative organisms. Red tides occurred usually in May but, less predictably, also during June – August. Causative organisms included the diatom Chaetoceros sp., and the dinoflagellate genera Prorocentrum, Peridinium, Ptychodiscus, Gonyaulax, Ceratium and Cochlodinium. Early summer red tides occurred at a time of increasing insolation, and water temperature, prevailing onshore winds and salinities in the range 34.4 to 36.6 A consistent feature was a quick rise in sea temperature of more than 2 C^o during the week preceding red water. Red tides later in the year coincided with higher temperatures and salinities and calm weather.     

Carotenoids in the tiger prawn Penaeus esculentus during ovarian maturation

Female Penaeus esculentus Haswell were collected by 15 to 20 min duration trawls during 1990. Carotenoids were analysed in the digestive gland, abdominal muscle, the remainder of the body (hereafter called integument) and ovary of prawns in Stage 2 through Stage 4 (fully mature) of maturation. The only oxycarotenoids (xanthophylls) identified were astaxanthins or astaxanthin esters; occasionally low levels of -carotene were detected in the digestive gland. The concentrations of astaxanthin monoesters (AM) and diesters (AD) were highest, with only minor amounts of free astaxanthins (Ast), except in the maturing ovaries, where free astaxanthins predominated (up to 80% of the total carotenoid). Of the total carotenoid, 82 to 94% was in the integument, but at maturity the digestive gland contained 10.7±3.4% and the ovary 5.6±0.9% of the total carotenoid. Only the ovary increased in mass during maturation, reaching up to 5.2% of total prawn mass. During this period, digestive gland concentrations of AM, AD and Ast all increased (tota 20 to 120 g g^-1); levels in the muscle and integument varied little throughout maturation (total 0.4 and 100 g g^-1, respectively); ovary AM levels remained low throughout (1.5 to 1.2 g g^-1), AD increased from only 2 to 5 g g^-1, but Ast increased from 2 to 34 g g^-1. Apart from the ovary, AM concentrations were the most variable. In common with other decapod Crustacea, the maturing ovary of P. esculentus contained high levels of carotenoids, indicating that these may have an important role in early development. The natural diet of P. esculentus includes a variety of carotenoids, but except for a little -carotene, the digestive gland, where absorption occurs, contained astaxanthins, with only an occasional trace of -carotene. This suggests that the conversion of dietary carotenoids to astaxanthin occurs soon after ingestion.     

Reproductive competition in queenless honey bee colonies (Apis mellifera L.)

Previously we reported that there are subfamily differences in drone production in queenless honey bee colonies, but these biases are not always explained by subfamily differences in oviposition behavior. Here we determine whether these puzzling results are best explained by either inadequate sampling of the laying worker population or reproductive conflict among workers resulting in differential treatment of eggs and larvae. Using colonies composed of workers from electrophoretically distinct subfamilies, we collected samples of adult bees engaged in the following behavior: true egg laying, false egg laying, indeterminate egg laying, egg cannibalism, or nursing (contact with larvae). We also collected samples of drone brood at four different ages: 0 to 2.5-h-old eggs, 0 to 24-h-old eggs, 3 to 8-day-old larvae, and 9 to 14-day-old larvae and pupae. Allozyme analyses revealed significant subfamily differences in the likelihood of exhibiting egg laying, egg cannibalism, and nursing behavior, as well as significant subfamily differences in drone production. There were no subfamily differences among the different types of laying workers collected from each colony, suggesting that discrepancies between subfamily biases in egg-laying behavior and drone production are not due to inadequate sampling of the laying worker population. Subfamily biases in drone brood production within a colony changed significantly with brood age. Laying workers had significantly more developed ovaries than either egg cannibals or nurses, establishing a physiological correlate for the observed behavioral genetic differences. These results suggest there is reproductive conflict among subfamilies and individuals within queenless colonies of honey bees. The implications of these results for the evolution of reproductive conflict, in both queenright and queenless contexts, are discussed.     

Ecological constraints on the behaviour of mother long-tailed macaques (Macaca fascicularis)

Summary The ecological consequence of the effects of different social conditions on the behaviour of females with infants were examined in two different-sized groups of long-tailed macaques (Macaca fascicularis) that inhabited adjacent home ranges in Gunung Leuser National Park in Indonesia. This study aims to test predictions of the within-group competition/predation avoidance hypothesis of social organization. Females, particularly those carrying an infant, are considered to experience the most direct effects of environmental constraints on fitness prospects. Data on maintenance behaviour, height in canopy and spatial position were collected for five mothers in a small group and eight mothers in a large group. Comparisons were made on the basis of group size and dominance rank. Mothers from the large group spent less time feeding on clumped fruits and more time foraging on dispersed food items. Lower-ranking mothers, undergoing the highest rate of within-group competition, foraged most. For macaques in the study area, predation risk is considered to decrease with height in the canopy. Mothers in the small group, assumed to be more susceptible to predation, remained substantially higher in the canopy than mothers in the large group. In both groups, mothers adjusted their height to their spatial position in the group. Lower-ranking mothers were more often found without neighbours, presumably to avoid feeding competition. As a result they stay higher in the canopy than higher-ranking mothers. The results clearly demonstrate that individual females in the same reproductive state and living in the same area, but in different social environments, adapted their behaviour according to the hypothesis. Correspondence to: D.R. Vos     

Experimental evidence for helping and hindering by previous offspring in the cooperative-breeding Seychelles warbler Acrocephalus sechellensis

Prebreeding Seychelles warblers (Acrocephalus sechellensis) frequently act as helpers on their natal territory, aiding in territory defence, predator mobbing, nestbuilding, incubation (only females) and feeding dependent young of their parents. In some cases helpers could attain breeding status (e.g. by joint-nesting) in their natal group and become co-breeders. Comparisons of group size and reproductive success on a given quality territory suggest that the presence of alloparents (helpers and cobreeders) significantly affects the reproductive success of their parents. The influence of alloparents on reproductive success was examined by removing alloparents from breeding units and comparing the success of natural-sized and artificially reduced groups. Removal experiments, controlled for territory quality, group size and breeder age, showed that the presence of one alloparent significantly improved the reproductive success of its parents. Analysis strongly suggests that this was entirely due to helping behaviour (i.e. providing care to offspring of their parents), thereby improving the helper's inclusive fitness benefits from staying at home. However, these experiments showed also that the presence of two or more alloparents in medium-quality territories significantly decreased reproductive success, compared with groups with one alloparent. Several lines of evidence suggest that this may have been due to the joint-nesting and reproductive competition that could occur in breeding groups, or simply to resource depression when a large number of previous offspring remained on their natal territory.     

Cadmium in Halobates micans from the central and south Atlantic Ocean

The concentrations of cadmium in 428 samples of the sea-skater Halobates micans from 57 locations in the central and south Atlantic Ocean ranged from 1.7 to 122 g Cd g^-1 dry wt. The concentrations in females, males and nymphs were not significantly different. The data suggest that the cadmium concentrations in these insects are somehow related to surface currents. The highest values (50 to 70 g g^-1, maximum value 122 g g^-1) were recorded from the South Equatorial Current region; intermediate values (10 to 20 g g^-1) from the Equatorial Counter Current and the North Equatorial Current; and relatively low concentrations (>10 g g^-1) from the Canary Current in the north and the Benguela Current in the south. A remarkable feature of these data lies in the fact that they reveal areas of high Cd content which correspond closely with those indicated by data obtained by Bull et al. (1977) from a different set of samples, analyzed by different methods and obtained in different years.     

Genetic evidence for monogamy in the cooperatively breeding red-cockaded woodpecker

We examined the genetic relationship among putative parents, offspring, and helpers in 224 red-cock-aded woodpeckers (Picoides borealis) from the Sandhills of North Carolina. Comparison of DNA similarity with a pedigree constructed from 3,823 individually-marked birds observed from 1979 to 1992 provided verification of observed relatedness in the sampled population (R ² = 0.87, df = 14, P < 0.001). In this population, breeding pairs may or may not have helpers, most of which are males that remain on their natal territory. Our sample included helpers assisting their parents, helpers assisting a related male and an unrelated female, and helpers assisting an unrelated pair. Band by band comparison and examination of similarity among DNA profiles indicated that all offspring from non-helped nests were sired by their putative parents (n = 28 families). Similarly, all but one offspring in helped nests were also sired by their putative parents (n = 16 families). In the exceptional case, the offspring evidently was sired by a male external to the group. Analysis of similarity values supported the conclusion that matings by helpers or extra-group males are rare or non-existent. Our results indicate that in this species advantages gained by individuals remaining on their natal territories as helpers do not generally include siring offspring. Correspondence to: S. Haig