Opposite shifts in size at metamorphosis in response to larval and metamorph predators

Predation risk can cause organisms to alter the timing of life history switch points. Theory suggests that increased risk in an early life stage should select for switching earlier and smaller, while increased risk in the subsequent stage should select for switching later and larger. This framework has frequently been applied to metamorphosis in amphibians, with mixed results. Few studies examining the effect of larval predation risk on metamorphosis have observed the predicted pattern, and no studies, to our knowledge, have examined the effect of increased risk during and after metamorphosis on the timing of this switch point. Here we examine the effect of larval and post-metamorphic predation risk on metamorphosis in the red-eyed treefrog, Agalychnis callidryas. We raised tadpoles in the presence or absence of cues from caged water bugs fed larvae and cues from spiders fed emerging metamorphs. Water bugs are effective larval predators, while spiders are poor larval predators but prey on metamorphs. Furthermore, since spiders forage on the water surface it is possible that tadpoles could assess future risk from this predator. Predators induced opposite shifts in life history. Tadpoles emerged smaller and less developed in response to water bugs, but later and larger in response to spiders. Interestingly, predator effects on larval duration were not independent; tadpoles delayed emerging in response to spiders, but only in the absence of water bugs.     

Food allocation rules vary with age and experience in a cooperatively breeding parrot

Although it is known that parents can differ in their optimal resource allocation to offspring in size-structured broods, the mechanisms determining differences in allocation rules of carers are not yet clarified. In cooperatively breeding species, breeders and non-reproductive helpers often differ in their fitness payoffs of providing care and in their breeding experience. Cooperative breeders thus provide an appropriate system to examine two hypotheses originally proposed to explain differences in food allocation among parents: (i) food allocation between carers differs because of the distinct cost-benefit ratio of selective feeding (i.e. breeders and helpers are expected to differ in food allocation) and (ii) carers differ in their ability to feed selectively (i.e. differences in food allocation are expected between experienced adults and inexperienced yearlings). We compared feeding rates with which breeders, old helpers and yearling helpers provisioned nestlings of different hatching rank. The influence of experience upon food allocation was further assessed by comparing food allocation of yearlings early and late during nesting. We show that allocation rules differ between age classes because breeders and old helpers fed the youngest chicks most, whereas yearlings showed the opposite pattern. The role of experience was supported by the fact that yearlings adjusted food allocation to that observed in experienced adults during the breeding season. We thus suggest that food allocation in El Oro parakeets depends either on differential skills of adults to transfer food to the youngest chick or on their ability to recognize nestling needs.     

Trill performance components vary with age, season, and motivation in the banded wren

Acoustic displays with difficult-to-execute sounds are often subject to strong sexual selection because performance levels are related to the sender’s condition or genetic quality. Performance may also vary with age, breeding stage, and motivation related to social context. We focused on within-male variation in four components of trill performance in banded wren (Thryophilus pleurostictus) songs: note consistency, frequency bandwidth, note rate, and vocal deviation. The latter is a composite measure reflecting deviation from the performance limit on simultaneously maximizing both frequency bandwidth and note rate. We compared the changes in these song parameters at three time scales: over the course of years, across the breeding season, and at different times of the day with contrasting agonistic contexts. Vocal deviation decreased and note consistency increased between years, suggesting that experience may improve individual proficiency at singing trills. Consistency also increased across the season, confirming that practice is important for this parameter. Although there was no significant seasonal change in vocal deviation, one of its components, note rate, increased during the season. Neither vocal deviation nor consistency varied with agonistic context. However, note rate increased during playback experiments simulating territorial intrusions compared to dawn chorus singing. The magnitude of a male’s increase in note rate was positively correlated with his aggressive behavior during the playback experiment. Thus consistency, bandwidth, and vocal deviation indicate age, whereas trill rate flexibly indicates the singer’s aggressive motivation. We also found evidence of a within-male trade-off between vocal deviation and consistency.     

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     

Rapid changes in growth,condition, size and age of small pelagic fish in the Mediterranean

Since 2007, the ecosystem of the Gulf of Lions has shifted to a different regime, characterised by a low anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) biomass and a remarkably high sprat (Sprattus sprattus) biomass. Surprisingly, the abundance and recruitment of anchovy and sardine remained high. To understand which processes (bottom-up or top-down control, etc.) could have caused this shift, we studied the changes in body condition, growth and size and age of anchovy, sardine and sprat over 1984–1985 and 1992–2012, using data from scientific surveys. The annual age structure of anchovy and sardine was estimated using Bayesian mixture models based on size frequency data with priors on the age–length relationship derived from independent otolith readings. The results indicated periods during which anchovy and sardine were in an average (1992–2004), good (2005–2007) or poor (2008–2012) overall state of condition. For sardine, the shift towards smaller fish observed during these past 4 years was explained by a combination of slower growth and the disappearance of older individuals (ages 2+). Despite the increase in biomass of sprat since 2008, indications were found that sprat was also smaller than in the past. As growth and condition decreased and overexploitation has not been documented or suspected for those three species in this area, we propose that the current decline in sardine and anchovy biomass could be due to qualitative and/or quantitative modifications in the planktonic production (i.e. a bottom-up control) or mass mortalities of adults due to an epidemic disease.     

The relationship between dewlap size and performance changes with age and sex in a Green Anole (Anolis carolinensis) lizard population

Ornaments are believed to signal an individual's ability to reproduce successfully and/or survive. Since an individual's fitness is often influenced by multiple traits (e.g. number of copulations, ability to acquire nest sites or to escape predators), which are difficult to quantify simultaneously, we examine performance traits (bite force, jumping performance) believed to be relevant to an individual's fitness. Specifically, we ask whether variation in dewlap size is related to variation in body size, bite force and jumping ability in the lizard Anolis carolinensis. Our results show that dewlap size is correlated with jumping capacity across all individuals, whereas the relationships between dewlap size, body size and bite force differ depending on sex/age class. We argue that selection against relatively large dewlaps at the transition between small mature and large mature males might be responsible for the lack of a relationship within large males. The absence or presence of a correlation between dewlap size and bite force, on the other hand, might be explained by differences in behaviour, such as territory establishment, anti-predator tactics, and/or mate choice. Our work thus suggests that selective forces influencing the evolution of ornaments may operate differently on different sexes and life-history stages.     

Litter sex ratios in the golden hamster vary with time of mating and litter size and are not binomially distributed

Summary Pregnancy rates, litter sizes, and litter sex ratios vary strongly with the time in the estrous cycle at which female golden hamsters (Mesocricetus auratus) are mated. Early matings tend to produce relatively high pregnancy rates, large litters, and female-biased sex ratios, while late matings tend to produce low pregnancy rates, small litters, and male-biased sex ratios. Time of mating and litter size are therefore correlated, but each appears to have an independent effect on litter sex ratio: time of mating and sex ratio are positively correlated, holding litter size constant, while litter size and sex ratio are negatively correlated, holding time of mating constant. At each litter size greater than two, the variance of litter sex ratios is less than the binomial variance expected on the hypotheses of independent sampling with a constant probability of producing a male. The main features of the distribution of litter sex ratios can be generated from a causal model in which different probabilities of producing a male apply to early and late conceptions within each litter. The relationship between litter size and mean litter sex ratio is potentially consistent with several different models for the evolution of adaptive sex-ratio variation. Offrint request to: U.W. Huck     

Spermatophore characteristics in bushcrickets vary with parasitism and remating interval

Male bushcrickets provide females with a nuptial gift, a spermatophore, which is transferred to females at mating. The spermatophore consists of a gelatinous mass, the spermatophylax, and the sperm-containing ampulla. Male spermatophore size is positively correlated with insemination rate and female refractory period and therefore with male reproductive success. In this study, we examined spermatophylax weight, ampulla weight and sperm number in males of Poecilimon mariannae parasitized by the acoustically orienting fly Therobia leonidei. We show that in parasitized males, spermatophylax weight decreases with the level of parasitism. In line with the hypothesis that parasitism is a cost to reproduction, we found that spermatophylax weight was reduced at remating. In contrast, the replenishment of the spermatophylax in unparasitized males was complete after 2 days and was increased no further after 3 days. Both sperm number and ampulla weight showed an increase over time since last mating and sperm production was estimated at a constant rate of 500,000 per day in all individuals, regardless of parasitism. The allocation of investment in components of the spermatophore varies greatly with parasitism and remating. Both factors had rather independent effects on spermatophore constitution, revealing functional constraints acting on spermatophore characteristics in bushcrickets, which are important for understanding the selection pressures working on its components. Received: 13 September 1999 / Received in revised form: 4 February 2000 / Accepted: 13 March 2000     

Preference for male traits in female wolf spiders varies with the choice of available males,female age and reproductive state

Individual variation in female preference for male traits may influence mate choice, especially if benefits and costs of choosiness vary with the range of available males or reproductive timing. We examined variation in female preference for male leg tuft size in Schizocosa ocreata (Hentz) wolf spiders with video playback. Dichotomous (simultaneous) choice test experiments included all possible combinations of four stimuli (i.e., modified versions of the same video male stimulus): average tuft size (control), reduced (−25%), enlarged (+25%), and no tufts (removed). Females exhibited a directional preference for larger tuft size independent of the nature of the choice (except for reduced tufts vs no tufts where no difference was seen). Female preference in the short term (over a period of 4 days) was also highly repeatable for control vs reduced tufts, but not for control vs enlarged tufts. Responses of females in ‘no-choice’ presentations of a single (control) male stimulus varied with age post-maturity; females were less receptive in weeks 1 and 2, highly receptive at week 3, and less thereafter. Mated females were least receptive and most aggressive towards a male stimulus. Females offered choices repeatedly at different ages post-maturity consistently preferred the control male vs reduced tufts over all 3 weeks but varied in their preference for enlarged tufts vs control male. In the first 2 weeks, females preferred the enlarged tuft male stimulus, but showed no preference by the third week. Females tested in week 4 showed no preference in either choice. Results suggest that the potential interaction between female preference for male traits and female reproductive timing may be a critical consideration in mate choice.     

Soil C and N changes with afforestation of grasslands across gradients of precipitation and plantation age

Afforestation, the conversion of unforested lands to forests, is a tool for sequestering anthropogenic carbon dioxide into plant biomass. However, in addition to altering biomass, afforestation can have substantial effects on soil organic carbon (SOC) pools, some of which have much longer turnover times than plant biomass. An increasing body of evidence suggests that the effect of afforestation on SOC may depend on mean annual precipitation (MAP). The goal of this study was to test how labile and bulk pools of SOC and total soil nitrogen (TN) change with afforestation across a rainfall gradient of 600-1500 mm in the Rio de la Plata grasslands of Argentina and Uruguay. The sites were all former grasslands planted with Eucalyptus spp. Overall, we found that afforestation increased (up to 1012 kg C x ha(-1) x yr(-1)) or decreased (as much as 1294 kg C x ha(-1) x yr(-1)) SOC pools in this region and that these changes were significantly related to MAP. Drier sites gained, and wetter sites lost, SOC and TN (r2 = 0.59, P = 0.003; and r2 = 0.57, P = 0.004, respectively). Labile C and N in microbial biomass and extractable soil pools followed similar patterns to bulk SOC and TN. Interestingly, drier sites gained more SOC and TN as plantations aged, while losses reversed as plantations aged in wet sites, suggesting that plantation age in addition to precipitation is a critical driver of changes in soil organic matter with afforestation. This new evidence implies that longer intervals between harvests for plantations could improve SOC storage, ameliorating the negative trends found in humid sites. Our results suggest that the value of afforestation as a carbon sequestration tool should be considered in the context of precipitation and age of the forest stand.     

煤矸石风化物不同粒级中重金属镉含量及其形态变化

对抚顺西露天矿煤矸石山表层煤矸石风化母岩进行多点取样,风化物筛分成>2 mm、2～0.25 mm和<0.25 mm等3个粒级,分析测定了矸石山表层母岩及其风化物不同粒级中Cd含量及其形态变化特征.结果表明:煤矸石母岩及其风化物中cd含量超过土壤环境质量2级标准,页岩、泥岩及其风化物pH值7.2～8.6,但燃后泥岩pH值4.9.碳质页岩2～0.25 mm粒级风化物中Cd损失最多,为73.6%;<0.25 mm细粒级Cd损失较少,为17.0%,这与细颗粒的表面吸附作用有关;泥岩及其各粒级风化物Cd元素不易损失,其Cd含量变化也相对较小;但燃后泥岩风化物中的Cd较容易损失.煤矸石风化物各样品类型中的Cd均以残渣态为主,可还原态和弱酸可提取态所占比例次之,可氧化态所占比例最小.弱酸可提取态Cd在细颗粒风化物中所占比例较高.     

Seasonal changes in the diet and feeding behaviour of a top predator indicate a flexible response to deteriorating oceanographic conditions

Shifts in the diet of top predators can be linked to changes in environmental conditions. In this study, we tested relationships between environmental variation and seasonal changes in diet of a top predator, the grey-headed albatross Thalassarche chrysostoma, breeding at Bird Island, South Georgia in an austral summer of 1999/2000. Oceanographic conditions in that year around South Georgia were abnormal (i.e. anomalously high sea surface temperature to a relative 19-year long-term mean). The diet of grey-headed albatrosses showed high seasonal variation, shifting from cephalopods (42.9 % by mass) in late February to Antarctic krill Euphausia superba (58.3 %) in late April, and grey-headed albatrosses breeding performance was low (16.8 %). This study shows these albatrosses did not manage to find sufficient alternative prey and highlight the risk to top predators if there is an increase in the frequency or severity of food shortages in Antarctic waters.     

Threshold to maturity in a long-lived reptile: interactions of age, size, and growth

Thresholds to sexual maturity—either age or size—are critical life history parameters. Usually investigated in short-lived organisms, these thresholds and interactions among age, size, and growth are poorly known for long-lived species. A 34-year study of captive green turtles (Chelonia mydas) that followed individuals from hatching to beyond maturity provided an opportunity to evaluate these parameters in a long-lived species with late maturity. Age and size at maturity are best predicted by linear growth rate and mass growth rate, respectively. At maturity, resource allocation shifts from growth to reproductive output, regardless of nutrient availability or size at maturity. Although captive turtles reach maturity at younger ages than wild turtles, the extensive variation in captive turtles under similar conditions provides important insights into the variation that would exist in wild populations experiencing stochastic conditions. Variation in age/size at maturity should be incorporated into population models for conservation and management planning.     

Seasonal cycles in whole-body proximate composition and energy content of forage fish vary with water depth

Quantifying the nutritional quality of forage fish is integral for understanding upper trophic levels as forage fish are the dominant prey for top predator fish, marine mammals, and sea birds. Many existing reports documenting body composition of forage species are not comparable due to confounding effects. This study systematically assessed the variability in proximate composition and energy content of 16 forage species in southeastern Alaska (57.2626 N/133.7394 W) between 2001 and 2004. Variation in energy and lipid contents was related to habitat, epipelagic planktivores varying most, mesopelagics intermediate, and demersal species relatively invariable. Season was the greatest source of variation as a result of short growing seasons at high latitude and energy allocation strategies for reproduction and growth. Among species that varied seasonally, energy and lipid increased over summer and declined during winter. Annual differences in body composition occurred during periods of peak energy content. Sampling recommendations and guidance for bioenergetics models are provided.     

Experimental and natural changes in the peacock's (Pavo cristatus) train can affect mating success

Petrie et al. (1991) demonstrated a correlation between the degree of elaboration of peacocks' trains and their mating success, and also showed that this correlation occurred because females preferred to mate with the male that had the most elaborate train of those sampled on the lek. Although these data suggest that female choice is responsible for non-random mating in this species, they do not conclusively show that train morphology is the cue that females respond to, because they do not rule out the possible influence of another unidentified variable which is correlated with train elaboration. This paper presents an experimental test of the importance of the peacock's train in determining male mating success. If the number or arrangement of eye-spots in the peacock's train influences mating success, then changing the number of eye-spots should change mating success. This prediction was tested in an experiment where the trains of male peafowl (Pavo cristatus) were manipulated by removing a number of eye-spots between mating seasons. Peacocks with eye-spots removed showed a significant decline in mating success between seasons compared with a control group. This result, together with the observational data, supports the hypothesis that the peacock's train has evolved, at least in part, as a result of female choice.     

Waste management in the leaf-cutting ant Acromyrmex echinatior: the role of worker size, age and plasticity

Division of labour is the hallmark of the success of many social animals. It may be especially important with regard to waste management because waste often contains pathogens or hazardous toxins and worker specialisation can reduce the number of group members exposed to it. Here we examine waste management in a fungus-farming, leaf-cutting ant, Acromyrmex echinatior, in which waste management is necessary to protect their vulnerable fungal crop. By marking ants with task-specific paint colours, we found clear division of labour between workers that engage in waste management and those that forage, at least during the fine timescale of the 3-day marking period. This division of labour was influenced by both age and size, with waste management workers tending to be smaller and younger than foragers. The role of preventing contaminated ants from entering the colony was fulfilled mainly by medium-sized workers. When the level of waste was experimentally increased, most of the ants that responded to remove the waste were workers previously engaged in tasks inside the nest rather than external waste workers or foragers. These responding workers tended to be young and medium-sized. Surprisingly, the responding ants were subsequently able to revert back to working within the fungus garden, but the probability of them doing so depended on their age and the length of time they were exposed to waste. The results demonstrate the importance of division of labour with regard to waste management in A. echinatior and show that this is adaptable to changing needs.     

Correlating gene expression with larval competence,and the effect of age and parentage on metamorphosis in the tropical abalone<Emphasis Type="Italic"> Haliotis asinina</Emphasis>

The non-geniculate crustose coralline alga (CCA) Mastophora pacifica can induce the metamorphosis of competent Haliotis asinina (Vetigastropoda) larvae. The ability to respond to this natural cue varies considerably with larval age, with a higher proportion of older larvae (e.g. 90 h) able to metamorphose in response to M. pacifica than younger larvae (e.g. 66 h). Here we document the variation in time to acquisition of competence within a larval age class. For example, after 18 h of exposure to M. pacifica, approximately 15 and 36% of 84 and 90-h-old H. asinina larvae had initiated metamorphosis, respectively. This age-dependent response to M. pacifica is also observed when different aged larvae are exposed to CCA for varying periods. A higher proportion of older larvae require shorter periods of exposure to CCA than younger larvae in order to initiate metamorphosis. In this experiment, as in the previous, a small proportion of young larvae were able to respond to brief periods of CCA exposure, suggesting that they had developed the same state of competency as the majority of their older counterparts. Comparisons of the proportions of larvae undergoing metamorphosis between families reveals that parentage also has a significant (P<0.05) affect on whether an individual will initiate metamorphosis at a given age. These familial differences are more pronounced when younger, largely pre-competent larvae (i.e. 66 h old) are exposed to M. pacifica, with proportions of larvae undergoing metamorphosis differing by as much as 10 fold between families. As these data suggest that variation in the rate of development of the competent state has a genetic basis, and as a first step towards identifying the molecular basis to this variation, we have identified numerous genes that are differentially expressed later in larval development using a differential display approach. Spatial expression analysis of these genes suggests that they may be directly involved in the acquisition of competence, or may play a functional role in the postlarva following metamorphosis.Communicated by M.S. Johnson, Crawley     

Preparation and activation: determinants of age at reproductive maturity in male baboons

Age at maturity is a particularly important life history trait, but maturational data are rare for males in natural populations of mammals. Here we provide information on three maturational milestones and their social and demographic correlates among 43 wild male baboons, Papio cynocephalus, in a natural population in Amboseli National Park, Kenya. We examined (1) age at testicular enlargement, which signals puberty and the onset of subadulthood, (2) age at attainment of adult dominance rank, which we consider to be the beginning of adulthood, and (3) age at first sexual consortship, which is the best measure available for age at first reproduction in male baboons. Testicular enlargement (median age = 5.69 years) occurred earlier among sons of high ranking mothers, and was not influenced by rainfall or seasonality. Attainment of adult dominance rank (median age = 7.41 years) was also accelerated among sons of high-ranking mothers, and among males whose mothers had died while the males were juveniles. First sexual consortship (median age = 7.92 years) was not influenced directly by maternal characteristics, but attainment of adult dominance rank always preceded first consortship. The lag time between attainment of adult rank and first consortship (median = 2.5 months; range = 5–526 days), was predicted by the number of sexually cycling females in the group when the male attained rank, and by how high ranking the male became in his first months as an adult. We suggest that the age at which a male baboon is ready to begin reproducing is influenced by a relatively stable maternal characteristic that exerts its influence early in development, but the timing with which this potential is realized depends on activation by more proximate, often stochastic triggers such as female availability. This two-level organization of influences is likely to contribute to the variance both in age at first reproduction and in lifetime fitness. Differences in the relative magnitude of the two levels will lead to both intra- and interspecific variability in the opportunity for maternal selection and sexual selection.     

Carer provisioning rules in an obligate cooperative breeder: prey type, size and delivery rate

Providing food to developing offspring is beneficial for offspring but costly for carers. Understanding patterns of provisioning thus yields important insights into how selection shapes (allo-) parental care strategies. Broadly, offspring development will be influenced by three components of provisioning (prey type, size and delivery rate). However, all three variables are rarely considered simultaneously, leading to suggestions that the results of many studies are misleading. Additionally, few studies have examined the provisioning strategies of breeders and non-breeding helpers in obligate cooperative breeders, wherein reproduction without help is typically unsuccessful. We investigated these components of provisioning in obligately cooperative chestnut-crowned babblers (Pomatostomus ruficeps). Prey type was associated with size, and delivery rate was the best predictor of the overall amount of food provided by carers. As broods aged, breeders and helpers similarly modified the relative proportion of different prey provided and increased both prey size and delivery rate. Breeding females contributed less prey than male breeders and adult helpers, and were the only carers to load-lighten by reducing their provisioning rates in the presence of additional carers. While our results suggest that breeders and helpers follow broadly comparable provisioning rules, they are also consistent with the idea that, in obligately cooperative species, breeding females benefit more from conserving resources for future reproduction than do helpers which have a low probability of breeding independently.     

Between-area differences in age and length at first maturity of the orange roughy Hoplostethus atlanticus

Mean age and length at onset of maturity were estimated for orange roughy (Hoplostethus atlanticus) populations off New Zealand, Namibia, and Great Britain. Significant between-area differences were apparent in both these parameters. Implications of the between-area differences on stock structure of orange roughy in New Zealand waters are discussed. A proportional relationship between age at onset of maturity and modal size of fish in the mature population is demonstrated. This is consistent with later-maturing fish experiencing a longer period at a pre-maturity growth rate that is relatively faster than the rate of somatic growth after maturity. Received: 22 December 1997 / Accepted: 5 May 1998