The costs of male parental care and its evolution in a neotropical frog

Summary Parental care is practiced exclusively by males of the Puerto Rican frog, Eleutherodactylus coqui. Males brood clutches of direct-developing eggs in non-aquatic nest sites and defend eggs against cannibalistic nest intruders. Here, I report on energetic and mating costs incurred by males that provide parental care, and suggest how these proximate costs affect male fitness and the evolution of male parental care in this species. Energetic costs are small for brooding males in comparison to non-brooding, calling males. Brooding males had a higher frequency of empty stomachs and lost small, but significant, fractions of their initial body mass during parental care. Abdominal fat bodies of brooding males during the middle third of parental care were significantly smaller than those of calling males; those of males brooding eggs in earlier or later stages were not different. The mating cost of parental care is greater. Most brooding males cease calling during parental care. However, gravid females are available (i.e., known to mate) on most nights during the principal breeding season; hence non-calling males miss potential opportunities to mate. A mating cost was estimated by calculating nightly mating probabilities for calling males in a plot where nightly calling male densities and daily oviposition schedules were known. On average, a male exhibiting normal calling behavior would be expected to obtain a new mate once every 35.7 days. Hence a brooding male that ceased calling for a 20-day parental care period would miss, on average, 0.56 additional mates. Males that were more successful than average in attracting mates could miss up to 1.63 matings. A marginal value model (Fig. 1) is used to analyze the net effect on male fitness of parental care benefits and costs in E. coqui (Fig. 3). The model indicates that males garner the highest reproductive success by providing care from oviposition through hatching. There is no stage during the pre-hatching period at which a desertion strategy would yield higher reproductive success. In fact, the model suggests that males should provide full parental care even in the face of much higher mating costs than currently obtain in the system.     

Consequences of paternal care on pectoral fin allometry in a desert-dwelling fish

Positive static allometry is a scaling relationship where the relative size of traits covaries with adult body size. Traditionally, positive allometry is thought to result from either altered physiological requirements at larger body size or from strongly condition-dependent allocation under sexual selection. Yet, there are no theoretical reasons why positive allometry cannot evolve in fitness-related traits that are solely under the influence of natural selection. We investigated scaling and sexual dimorphism of a naturally selected trait, pectoral fin size, in comparison to a trait important in male–male combat, head width in natural populations of a fish, the desert goby Chlamydogobius eremius. Male desert gobies provide uniparental care and use their pectoral fins to fan the brood (often under hypoxic conditions); hence, larger fins are expected to be more efficient. Male pectoral fins do not appear to fulfil a signalling function in this species. We found that, for both pectoral fin size and head width, males exhibited positive allometric slopes and greater relative trait size (allometric elevation) than females. However, for head width, females also showed positive allometry, albeit to a lesser degree than males. Because fin locomotory function typically does not result in positive allometry, our findings indicate that other naturally selected uses, such as paternal care, can exaggerate trait scaling relationships.     

Mating effort or paternal investment? Incorporation rate and cost of male donations in the wartbiter

Summary Provision of nuptial gifts and incorporation of male-derived substances occurs in several insects. There has been some discussion as to whether these gifts should be regarded as a form of paternal investment or part of mating effort. One assumption of the paternal investment hypothesis is that the donating male is expected to father all, or the majority of, the offspring. The rate of incorporation into developing eggs is therefore assumed to be rapid enough to assure the donating male of paternity of the offspring in which he is investing. Spermatophore production in a paternal investment system is also expected to be costly, since males are providing females with highly nutritious gifts during copulation, thereby increasing female fecundity. Therefore, under conditions of food limitation males are expected to alter either mating frequency or spermatophore size. Experiments on the wartbiter (Decticus verrucivorus), using radioactive isotopes, show that the incorporation rate of male-derived substances is so slow that females become sexually receptive before any material received from the last male have been incorporated into eggs laid by the female. Since sperm mixing occurs in this species and since the label is present in eggs laid up to more than 30 days after mating, males might potentially benefit another male's offspring. The nuptial gift (the spermatophylax) was found, regardless of mating frequency and diet, to have a low protein content. These findings coincide with previous results showing that consumption of the gift had no effect on female fecundity. Instead, wartbiter males keep mating frequency high, are capable of mating every day regardless of food availability, invest the same proportion of the body weight in spermatophore production regardless of size, and produce a spermatophylax that correlates with ampulla size. These results indicate that the nuptial gift mainly functions to increase fertilization success in the wartbiter.     

Allocation of reproductive effort in Mus domesticus: responses of offspring sex ratio and quality to social density and food availability

Summary Female mammals in good condition can maximize their inclusive fitness by investing more in male offspring than in female offspring during periods of poor environmental quality. To test this hypothesis, we measured the effects of undernutrition and crowding before and during gestation on the sex ratio and weight of offspring at parturition and at weaning in Mus domesticus. Sex ratio was not significantly affected by density. Dams altered the sex ratio of their offspring in response to food availability, but only if variance in competitive success within the experimental subpopulation was evident. Thus ad lib fed females produced litters with an unbiased sex ratio, competitively successful females under moderate food availability produced a male-biased sex ratio, and severely food deprived females produced litters with a female-biased sex ratio. In groups that experienced competition for food, successful dams favoured male offspring during lactation. These results are consistent with the predictions of Trivers and Willard (1973). Analysis of within-cell variance and covariance suggests that the interaction of social structure and food availability provides specific cues for the dams' tactical reproductive choices.     

The relationship between ant-tending and maternal care in the treehopper Publilia modesta

We examined the relationship between maternal care and ant-tending in the treehopper species, Publilia modesta. Ant-tending increased the number of nymphs surviving to adulthood by approximately fivefold. Maternal care increased the number of nymphs surviving to adulthood by approximately 50% in the presence of ants. Maternal care had either no effect or the opposite effect in the absence of ants; when ants were not present survivorship was 30% lower in the presence of mothers. Maternal care increased the average number of ants tending nymph aggregations approximately threefold early in the summer; consequently, we suggest that the primary benefit of maternal care is to attract ants while nymphs are small and few in number. Because there was no benefit of maternal care in the absence of ants, we conclude that ant-tending provides an ecological context that favors maternal care in this treehopper species.     

Baboons adjust secondary sex ratio in response to predictors of sex-specific offspring survival

Summary Most theories of sex ratio adjustment assume that parents will adjust the sex ratio of births (secondary sex ratio) in a manner that maximizes offspring reproductive success (as long as this does not jeopardize parental reproductive success). Survival to maturity is typically the largest component of variance in offspring reproductive success. This should make environmental predictors of sex-specific offspring survival strong predictors of secondary sex ratio adjustment. We tested this survivorship maximization hypothesis for secondary sex ratio adjustment using data from a 17-year demographic study of 315 yellow baboon infants at Mikumi National Park, Tanzania. Sex differences were found in the degree to which several social and ecological conditions affected infant survival to 1 year. Female, but not male, infant survival was inversely correlated with birth order and the proportion of infant females in their birth cohort. Male, but not female, infant survival was inversely correlated with the amount of rainfall early in the year (January) and the proportion of infant males in their birth cohort; male survival also was positively correlated with maternal dominance rank. The consistency and timing of these effects across years suggested that such information was, in fact, available to females around the time of conception. Most importantly, social and ecological conditions that predicted improved survivorship of a given sex also were positively correlated with production of that sex. Early births were female-biased; and low January rainfall was correlated with a male-biased sex ratio, becoming increasingly female-biased as January rainfall increased. However, no sex ratio effects were correlated with maternal rank. Data supported the hypothesis that females adjusted secondary sex ratio in a manner that maximized sex-specific infant survival. This hypothesis also offered a plausible explanation for some of the contradictory data that have arisen from studies of maternal rank effects on sex ratio both within and between species.Correspondence to: S. K. Wasser     

Parentage and paternal care: consequences of intersexual selection in Savannah sparrows?

Empirical relationships between parentage and male parental care are commonly interpreted in the context of life-history models that consider increased offspring survivorship as the only benefit of paternal effort. However, indirect benefits associated with male care can also influence a male's response to cuckoldry: if females allocate paternity according to their prior experience with male parental care, it may pay for males to provision extra-pair young in early broods. Here, I assess the relationship between first-brood parentage and paternal care in a population of Savannah sparrows (Passerculussandwichensis) where a male's fertilization success in the second brood appears to be influenced by his prior parental performance. Based on the multi-locus DNA fingerprinting of 17 first broods, male feeding effort was influenced by parentage (percent of brood resulting from within-pair fertilizations) but not by brood size, male mating status (monogamous versus polygynous), timing of breeding (hatching date), structural size (wing length) or condition (mass). Males provided more care to broods that contained few within-pair young. This result supports the idea that males provision young to increase their future mating success, but alternative hypotheses involving male quality and timing of breeding cannot be excluded. Received: 13 August 1996 / Accepted after revision: 22 February 1997     

Litter reductions reveal a trade-off between offspring size and number in brown bears

Experimental manipulations have revealed positive effects of litter reduction on offspring mass in small mammals, but little is known about this trade-off in large mammals. We examined the determinants of natural litter size variation and quantified the effects of litter size, maternal characteristics, and litter composition on yearling mass using 24?years of data on marked brown bears (Ursus arctos) in Sweden. Infanticide by adult males is a major cause of cub-of-the-year mortality, leading to litter size reductions. Litter size (n?=?265) at den emergence ranged from one to four cubs (average, 2.7) and increased with maternal age. Litter size, however, appeared independent of maternal size, population density, interlitter interval, study area, or previous litter sex ratio. Yearling body mass increased with maternal body size but was independent of litter sex ratio. Litter size and yearling mass were negatively correlated, mostly because singletons were about 30?% heavier than yearlings from litters of two to four cubs. In reduced litters, survivors were on average 8?% heavier as yearlings than individuals from intact litters, suggesting that sibling competition reduces growth. Trade-offs between litter size and yearling mass in bears appear similar in magnitude to those found in small mammals.     

Parents adjust care in response to weather conditions and egg dehydration in a Neotropical glassfrog

Parental hydration of terrestrially developing eggs has evolved repeatedly among frogs and is thought to buffer embryos from environmental variation. While many anurans offer relevant opportunities to study parental care, research on how parents respond to environmental variation and offspring conditions are lacking. In this study, we investigated the interrelationships of weather, embryo hydration demands, and parental provisioning in a wild population of the glassfrog Hyalinobatrachium (‘Centrolenella’) fleischmanni in Oaxaca, Mexico. We determined whether males modify parental behavior in response to changes in weather conditions that effect embryo dehydration, how variation in both weather and parental hydration affect egg water balance and embryonic mortality, and whether parental provisioning is related to the hydration levels of egg clutches. We found that male H. fleischmanni compensate for environmental variation in offspring conditions by adjusting the frequency of parental care in response to both weather and egg dehydration. Using a male removal experiment, we examined the function of paternal care and made comparisons with previous research, finding that both the adaptive value of parental care and flexibility in parental behavior are impacted by spatial and temporal conditions. We present observations that indicate a direct conflict between providing parental care and multiple matings. In summary, this research demonstrates that the variable frequency of paternal care in H. fleischmanni is a response to the fluctuating nature of the climate and resulting hydration requirements of embryos in combination with the allocation of effort to parental care versus mating activity.     

Use of mercury in agriculture and its relationship to environmental pollution

Linuron samples at 75mg/L concentration were photolyzed with ultraviolet lamps or with natural sunlight. Ten photoproducts were identified and yields were determined for the more significant photoproducts resulting from either sunlight or ultraviolet lamp photolysis.     

Experimental evidence of a testosterone-induced shift from paternal to mating behaviour in a facultatively polygynous songbird

Previous studies have suggested that testosterone (T) profiles of male birds reflect a trade-off between mate attraction behaviours (requiring high T levels) and parental care activities (requiring low T levels). In this study, we experimentally elevated T levels of monogamous males in the facultatively polygynous European starling (Sturnus vulgaris), and compared mate attraction and paternal behaviour of T-treated males with those of controls (C-males). T-males significantly reduced their participation in incubation and fed nestlings significantly less often than C-males. Females paired to T-treated males did not compensate for their mate’s lower paternal effort. The observed reduction in a male’s investment in incubating the eggs was accompanied by an increased investment in typical female-attracting behaviours: T-males spent a significantly higher proportion of their time singing to attract additional females. They also occupied more additional nestboxes than C-males, although the differences just failed to be significant, and carried significantly more green nesting materials into an additional nestbox (a behaviour previously shown to serve a courtship function). T-males also behaved significantly more aggressively than C-males. During the nestling period, the frequency of mate-attracting behaviours by T-treated and control males no longer differed significantly. Despite the reduced paternal effort by T-males and the lack of compensation behaviour by females, hatching and breeding success did not differ significantly between T- and C-pairs. Received: 7 February 2000 / Revised: 10 August 2000 / Accepted: 3 September 2000     

Predator foraging behavior in response to perception and learning by its prey: interactions between orb-spinning spiders and stingless bees

Although rewarded bees learn and remember colors and patterns, they have difficulty in learning to avoid negative stimuli such as decorated spider webs spun by Argiope argentata. A. argentata decorates its web with silk patterns that vary unpredictably (Fig. 1) and thus foraging insects that return to sites where spiders are found encounter new visual cues daily. Stingless bees can learn to avoid spider webs but avoidance-learning is slowed or inhibited by daily variation in web decorations (Figs. 3,4; Tables 1,2). In addition, even if bees learn to avoid decorated webs found in one location, they are unable to generalize learned-avoidance responses to similarly decorated webs found at other sites. A. argentata seems to have evolved a foraging behavior that is tied to the ways insects perceive and process information about their environment. Because of the evolutionary importance of bee-flower interdependence, the predatory behavior of web-decorating spiders may be difficult for natural selection to act against.     

Reproductive synchrony in a habitat-forming kelp and its relationship with environmental conditions

Key biological processes such as the timing and synchrony of reproduction, are sensitive to fluctuations in the environment, as well as alterations in the physiology and behaviour of individuals and the ecology of populations. The main aim of this study was to identify patterns in reproductive timing and relationships with prevailing environmental conditions, for the major habitat-forming alga in temperate Australia, Ecklonia radiata (C. Agardh) J. Agardh. We observed strong synchronous patterns of zoospore production across spatial scales from hundreds of metres to tens of kilometres, with populations of E. radiata reproducing from mid-summer to the end of autumn. High zoospore densities were positively related to sea temperature and E. radiata released zoospores in temperatures well above previously documented thermal tolerance limits for other Laminariales. Reproductive timing was also negatively related to seasonal increase in wave exposure. We conclude that E. radiata undergoes synchronous reproduction which can be linked to its annual growth cycle and natural environmental fluctuations, to promote growth and survival of recruits.     

The air speed of migrating birds and its relationship to the wind

Summary This paper deals with the flight speeds of migrating birds observed by radar over a Swiss Alpine pass. The distributions of air speeds for different classes of birds and the influence of the wind on air speeds were investigated. Our findings differ greatly from observations of bird migration over the North Atlantic Ocean and the North American continent. Our data reveal: (1) that the air speeds of the bulk of migrating birds were in the range of 8–18 m/s and that the amount of slow flyers (air speed below 5 m/s) was less than 5%; (2) that there was an obvious influence of the wind on air speeds and, moreover, the data showed a distinctly variable compensatory behavior among different bird classes; (3) that the wind component in the direction of the bird's heading was a better predictor for air speed than the wind direction. Although we do not speculate about possibilities and mechanisms of wind detection, a simple argument shows how birds could estimate wind directions if they do use the moving surface as a reference system.     

Growth of the rock lobster Jasus lalandii and its relationship to benthos

Growth increments of the male rock lobster Jasus lalandii vary systematically with location within a small (38 km²) fishing ground off the South African west coast. High growth rates were recorded from an area where the availability of benthos suitable as rock-lobster food was also high. Increments could also be shown to vary significantly in different years, but over the size range examined, growth did not appear to be a function of size. Benthic biomass was found to be inversely related to depth. However, comparisons between biomass values from the same depths in two different areas of the fishing ground suggested that biomass was higher in areas which were in proximity to shallow water, where extensive kelp beds were found. Mussels (Aulacomya magellanica) formed an important constituent of the rock lobster's diet; these were prevalent at depths between 20 and 40 m, and especially in areas where rock-lobster growth rates were high.     

平原河网地区大型底栖动物群落结构及其与环境因子的关系

大型底栖无脊椎动物是河流生态系统的重要组成生物,具有活动范围相对固定、生命周期较长、生活习性相对稳定等特点,可有效指示河流水质和水生态系统健康状况。有关大型底栖生物群落结构特征及其与水环境之间的关系研究一直是河流生态系统的研究热点。底栖动物具有较强的地域性,其群落结构特征受当地的自然环境特征和社会经济发展程度的双重因素影响。为研究典型平原河网地区的底栖动物群落结构及对水环境状况的响应特征,在上海市选取了83个河道断面进行大型底栖动物采样分析,共获取底栖动物20个分类单位（种）,其中软体动物10种,环节动物6种,节肢动物4种。上海河道底栖动物群落结构总体呈现结构单一、物种丰富度低、高度耐污性等显著特征,环节动物的栖息密度和出现频率均最高。城市化水平的空间差异性直接影响底栖动物空间分布的异质性,由市区至近郊、远郊,物种数从11种逐渐上升至15种,而栖息密度则由8 776.3个/m2下降至690.3个/m2。底栖动物群落结构的空间分布特征与水质理化指标的空间分布特征具有良好的对应性,水质空间变化导致环节动物和软体动物、节肢动物的空间分布存在一定交错性,耐污性较强的环节动物主要分布在有机污染重、溶解氧低的市区河道断面,而喜清洁环境的软体动物和节肢动物主要分布在水质较好的郊区断面。寡毛纲物种的栖息密度与溶解氧质量浓度呈显著负相关性。     

Reproductive status of Octopus pallidus, and its relationship to age and size

Age-specific information on individual octopus reproductive development and investment from wild populations has until recently been unobtainable. Using daily-formed increments within stylets (internal shells) the individual ages of 503 wild Octopus pallidus were determined. In addition, detailed reproductive information was collected for each of the aged octopus, along with reproductive data for an additional 925 octopus. All of the octopus were collected from Bass Strait waters in south-eastern Australia from November 2004 to November 2006. This information was used to investigate seasonal trends in reproductive scheduling and investment, fecundity and egg size. Maturation in O. pallidus primarily depends on size with little relationship to age and is highly variable between genders, with females >350 days still maturing in comparison to all males >142 days being mature. Size at 50% maturity for females was approximately 473 g, which is considerably larger than male 100% maturity at <250 g. This indicates that for females at least, maturity does not necessarily come with age. Seasonal scheduling in reproductive investment between genders revealed an optimal spawning period between late summer and early autumn. These results reinforce the view that individual growth and maturity is highly variable in cephalopods.