The interaction between reproductive cost and individual quality is mediated by oceanic conditions in a long-lived bird

Environmental variability, costs of reproduction, and heterogeneity in individual quality are three important sources of the temporal and interindividual variations in vital rates of wild populations. Based on an 18-year monitoring of an endangered, recently described, long-lived seabird, Monteiro's Storm-Petrel (Oceanodroma monteiroi), we designed multistate survival models to separate the effects of the reproductive cost (breeders vs. nonbreeders) and individual quality (successful vs. unsuccessful breeders) in relation to temporally variable demographic and oceanographic properties. The analysis revealed a gradient of individual quality from nonbreeders, to unsuccessful breeders, to successful breeders. The survival rates of unsuccessful breeders (0.90 +/- 0.023, mean +/- SE) tended to decrease in years of high average breeding success and were more sensitive to oceanographic variation than those of both (high-quality) successful breeders (0.97 +/- 0.015) and (low-quality) nonbreeders (0.83 +/- 0.028). Overall, our results indicate that reproductive costs act on individuals of intermediate quality and are mediated by environmental harshness.     

Age- and state-dependent reproductive effort in male mountain goats, Oreamnos americanus

In mammalian polygynous mating systems, male reproductive effort consists mainly of male–male competition and courting of females, which entail substantial somatic costs. Males are thus expected to adjust their reproductive effort according to their age and condition. In this study, we examined how activity budgets of male mountain goats (Oreamnos americanus), a polygynous ungulate, varied with age in a marked population over two periods: (1) summers 1995–2006 and (2) ruts 2004–2006. We then assessed if the proportions of time spent in male–male competition and courtship behaviors were influenced by age-specific body mass and social rank during the rut. Males spent most of their time foraging and resting during summer, and rested more and foraged less with increasing age. During the rut, pronounced shifts in activity budgets occurred as juveniles (1–2 years) increased time spent foraging, whereas adults (≥3 years) increased standing and time spent in social interactions at the expense of foraging. At old age, reproductive effort either stabilized or decreased slightly, providing weak support for the ‘mating strategy–effort’ hypothesis, predicting that courtship behaviors should peak in prime-aged males. Age-specific body mass did not affect time spent in male–male competition, but was positively related with time spent in courtship behaviors, providing support for the ‘individual quality’ hypothesis, predicting that males with more resources at the start of the rut should spend more time in mating-related activities. Age-specific social rank did not affect reproductive effort. Surviving to prime age while increasing mass each year should thus allow male ungulates to gain greater ability to court estrus females.     

Arrival timing and seasonal reproductive performance in a long-distance migratory landbird

The date when a landbird migrant arrives on its breeding grounds may have reproductive consequences. Generally, early arriving individuals begin breeding earlier and consequently experience greater seasonal reproductive performance. Here, we describe relationships between arrival timing and seasonal reproductive performance in the American redstart (Setophaga ruticilla), a long-distance passerine migrant, arriving at northerly breeding grounds in Michigans eastern Upper Peninsula. Evidence suggests that both males and females benefited from early arrival at the breeding grounds. Early males appeared to settle on higher quality territories and hatched nestlings sooner than later arrivals. Early females began their clutches early, produced heavier nestlings and possibly laid more eggs than later arrivals. Larger clutches and heavier offspring increase the likelihood of offspring recruiting into the breeding population. The findings of this study point to fitness consequences arising from when a bird arrives at its breeding grounds. These results also have implications for understanding how events occurring during spring migration influence reproductive performance as migratory delays likely influence arrival timing.Communicated by: W. Wiltschko     

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.     

Early nutritional conditions, growth trajectories and mate choice: does compensatory growth lead to a reduction in adult sexual attractiveness?

It is well documented that poor early nutrition can have profound negative effects on adult life-history traits. However, it has also been demonstrated that organisms can undergo compensatory resource allocation strategies (such as an accelerated growth rate) if food availability improves, so as to mitigate the effects of the poor early conditions. Previous research has indicated that elevated growth rates can incur costs in the longer term, such as an increased rate of senescence and shorter lifespan. We tested whether a phase of compensatory growth after a period of reduced food availability earlier in life affected the sexual attractiveness of adult male green swordtails Xiphophorus helleri, a species in which it has previously been documented that females prefer larger-bodied and longer-tailed males. The experiment compared the attractiveness of size-matched brothers that had experienced contrasting growth trajectories as juveniles; the experiments were initially conducted in the middle of a male’s sexually reproductive life and were then repeated towards the very end of life. At both ages, males that had undergone compensatory growth were equally as attractive as their brothers that had grown normally. These results suggest that the growth compensation benefits males through an increase in their attractiveness over that which they would have had if they had remained on their original growth trajectory. The lack of change in relative attractiveness with age indicates that the compensation does not cause greater deterioration in secondary sexual characters at older ages than in continuously well-fed males.     

Elaborately ornamented males avoid costly parental care in the house finch (<Emphasis Type="Italic">Carpodacus mexicanus</Emphasis>): a proximate perspective

Selection should favor flexibility in reproductive tactics when the combination of sexual traits and reproductive behaviors that achieve the highest fitness differs between males within a population. Understanding the functional significance of variation in male reproductive tactics can provide insight into their evolution. Male house finches (Carpodacus mexicanus) in a Montana population display continuous variation in parental tactics: males with more elaborated (redder) plumage color provide little or no parental care compared to less elaborated (dull) males. Here, we first determined whether elevation of prolactin (a pituitary hormone) was related to variation in male parental tactics and, second, we used the relationship between prolactin levels and parental behavior to investigate why redder males avoid a high investment in parental care. We found that prolactin elevation was closely associated with paternal care. In addition, males with redder plumage color had low prolactin levels, whereas dull males, which provision twice as frequently, had high levels of prolactin. We also found that male condition was unrelated to plumage color but negatively related to prolactin levels. These results suggest that the low provisioning of redder males was not due to physiological constraints, but instead reflected a tactic to avoid the costs associated with parental care. The condition benefits accrued by redder males may explain their higher post-breeding survival compared to dull males. Moreover, dull males were previously shown to have higher pairing success than redder males, suggesting that the relationship between male plumage color and parental care may reflect individually optimized parental tactics.Communicated by H. Kokko     

Material, energy and environmental performance of technological and social systems under a Life Cycle Assessment perspective

Selected energy and material resource conversion systems are compared in this paper under an extended LCA point of view. A multi-method multi-scale assessment procedure is applied in order to generate consistent performance indicators based on the same set of input data, to ascertain the existence of constraints or crucial steps characterized by low conversion efficiency and to provide the basis for improvement patterns. Optimizing the performance of a given process requires that many different aspects are taken into account. Some of them, mostly of technical nature, relate to the local scale at which the process occurs. Other technological, economic and environmental aspects are likely to affect the dynamics of the larger space and time scales in which the process is embedded. These spatial and time scale effects require that a careful evaluation of the relation between the process and its surroundings is performed, so that hidden consequences and possible sources of inefficiency and impact are clearly identified. In this paper we analyse and compare selected electricity conversion systems, alternative fuels and biofuels, waste management strategies and finally the time evolution of an urban system, in order to show the importance of a multiple perspective point of view for the proper evaluation of a system's environmental and resource use performance.     

Worldviews,institutions and sustainability: An introduction to a co-evolutionary perspective

This work analyzes hierarchical and causal relationship between worldview, institutions, technology and economic behaviour under conditions of informational uncertainty.

In our pursuit of sustainable living, institutions function both as constraints and as primary instruments of change of socio-economic behaviour. The process of radical institutional change is contingent upon prior change in the dominant worldview, while its dynamics resembles that of a paradigm change in political and scientific arenas. Therefore, the timely and adaptive outcome of institutional change with respect to global environmental problems remains intrinsically uncertain.     

Alternative male mating tactics in a cichlid, Pelvicachromis pulcher: a comparison of reproductive effort and success

Pelvicachromis pulcher is a small African cichlid which breeds in holes. Males may either reproduce monogamously (pair males), polygynously (harem males), or be tolerated as helpers in a harem territory (satellite males). These helpers share in defence of the territory against conspecifics, heterospecific competitors and predators. There are two male colour morphs that are fixed for life and are apparently genetically determined. These differ in their potential mating strategy. Red morph males may become harem owners, while yellow morph males may become satellite males, and males of both morphs may alternatively pair up monogamously. We compared the reproductive effort and success of these three male reproductive strategies. Effort was measured as attack rates, time expenditure and the risk of being injured or killed when attacking competitors or predators of three sympatric fish species. Reproductive success was measured by observing how many eggs were fertilized by each male when this was possible, and by using genetic markers. The number of fry surviving to independence of parental care was used as a criterion of success. The reproductive success of harem males was 3.3 times higher than that of pair males and 7 times higher than that of the average satellite male. Dominant satellite males, however, were as successful as monogamous pair males, using the measure of fertilized eggs. To our knowledge, this has not been found previously in any fish species. Both harem and pair males had lower parental defence costs per sired offspring, however, than males using the alternative satellite tactic. Defence effort was significantly related to the risk of injury. Received: 17 January 1996 / Accepted after revision: 9 June 1997     

Racketed tail of the male and female turquoise-browed motmot: male but not female tail length correlates with pairing success,performance, and reproductive success

Both males and females of many avian species maintain elaborate plumage traits, and elaborate monomorphic plumage may convey adaptive benefits to one or both sexes as inter- or intraspecific signals. Both sexes of the turquoise-browed motmot (Eumomota superciliosa) are elaborately plumed with long racket-tipped tail. I investigated whether the racketed tail functions as a sexually selected signal in one or both sexes by testing the predictions that males and/or females with the largest tails have: (1) greater pairing success, (2) greater reproductive performance (clutch-initiation date, clutch size, and hatching success), and (3) greater reproductive success. Yearling males with longer denuded rachises (wires) on the central tail feathers had greater pairing success. In addition, adult males with longer wires paired with females who laid larger clutches, had greater hatching success independent of clutch size, and fledged more young. There was no relationship between female tail plumage and pairing success, reproductive performance, or fledgling success. These results are consistent with the hypothesis that male tail plumage functions as a mate choice or status signal, but that the tail of the female does not function in a sexually selected context. I discuss alternative hypotheses for the evolutionary maintenance of the elaborate female tail plumage.     

Philopatry, male presence and grooming reciprocation among female primates: a comparative perspective

Competition for food and a safe location in a group are considered to be the main determinants of variation in social relationships among female primates. The effect of the presence of males is usually overlooked, however. Here we put forward two hypotheses connecting the (relative) number of males in a group, a statistic measuring the strength of positive relationships among females (the degree of reciprocity of grooming) and female residence. Under the first hypothesis, we assumed that philopatric females suffer especially from competition for males. Because females of these species are restrained to their native group and thus do not move to groups where better conditions (i.e. more males) prevail, a higher socionomic sex ratio would imply reduced competition and this should be reflected in better relationships among females (i.e. in a higher degree of reciprocation). The second hypothesis is based on the contention of Wrangham (1987) that groups with more males are better able to defend large food sources: in these groups female relationships would suffer less from within-group competition for food. We therefore also expected a positive correlation between the absolute number of males and grooming reciprocation. To test the hypotheses, matrices of grooming interactions were collected by going through the primatological literature. For ten female-resident and four female-transfer primate species, the degree of grooming reciprocation was quantified using a specially adapted matrix statistic. The results favoured the first (competition for males) hypothesis. The degree of grooming reciprocation among females was positively correlated with socionomic sex ratio, but neither with group size nor with the absolute number of males. Correlations between the degree of reciprocity and sex ratio were confirmed at three levels: within the same group at different points in time, between groups of the same species, and between species. In addition, the degree of reciprocation increased with sex ratio more strongly in typical single/variable male than in multi-male species. This we interpreted as a direct consequence of the lower social availability of males for females in multi-male groups than single-male groups. In turn, this effect may be due to males competing for females in multi-male groups. Received: 28 May 1997 / Accepted after revision: 9 November 1997     

The phalloid organ, orgasm and sperm competition in a polygynandrous bird: the red-billed buffalo weaver (Bubalornis niger)

The buffalo weavers, Bubalornis spp., are unique amongst birds in possessing a phalloid organ, a phallus-like structure anterior to the cloaca. We studied the red-billed buffalo weaver Bubalornis niger, to determine whether the phalloid organ has evolved in response to sperm competition. The phalloid organ was significantly longer in males that were resident at nests than in non-resident males, and among resident males was significantly longer in those males with a harem than in those without. Red-billed buffalo weavers bred colonially and had either a cooperatively polygynandrous (usually two unrelated males and several females) or a polygynous (one male and several females) mating system. Cooperative polygynandry provided females with the opportunity to copulate with more than one male and paternity analyses using DNA fingerprinting revealed that 63% of 16 multiple-offspring broods, comprising 43 offspring, had multiple sires, which included both nest-owning males and extra-group males. Sperm competition was therefore intense. Observations and experiments with buffalo weavers in captivity revealed that the phalloid organ was not intromittent during copulation, but functioned as a stimulatory organ which necessitated protracted copulation in order to induce male 'orgasm' and ejaculation, a feature apparently unique to this species.     

A dusk chorus effect in a nocturnal bird: support for mate and rival assessment functions

The dawn chorus is a striking feature of spring mornings and a characteristic behaviour of many bird species, particularly the passerines. Dawn singing has been considered a reliable signal of male quality for mate and rival assessment. Singing is presumed to be relatively costly at dawn both because air temperatures are relatively low and because birds have not fed overnight. Models of optimal daily routine predict the existence of a “dusk chorus” in nocturnal birds, although this prediction has received little empirical attention. Nocturnal birds at dusk may be energy-limited because of a lack of daytime feeding, and singing at dusk may thus ensure signal reliability. Here, we used an observational and experimental approach to study vocal behaviour at dusk and dawn in a nocturnal raptor, the little owl Athene noctua. We assess whether male little owls adjust their vocal behaviour according to feeding stage (i.e. period of the night), ambient air temperature and territorial context (i.e. spontaneous calling behaviour vs elicited calling by intrusion). Across different temperatures, we find that both spontaneous vocal activity and inter-individual variability in call duration increased at dusk, clearly indicating a dusk chorus phenomenon. Results from playback presentations suggest that food, rather than air temperature, is likely to be more constraining at dusk. We discuss how comparing dusk and dawn choruses in nocturnal and diurnal species can provide insights into both mechanistic and functional aspects of signalling behaviour.     

Variation in homeward orientation performance in the sleepy lizard (Tiliqua rugosa): effects of sex and reproductive period

Sleepy lizards are monogamous skinks which show high pair fidelity. This study reveals inter- and intrasexual differences in homeward orientation performance in this lizard. Male and female lizards were displaced during three phases of the spring activity period, the pre-pairing, pairing/mating, and post-pairing periods. All groups (with the exception of post-pairing males) were significantly oriented homewards, but males were significantly better oriented towards home than females during the pairing period. Furthermore, males were significantly better homeward oriented during the pre-pairing and pairing periods than in the post-pairing period. Similar results were observed for rate of movement away from the release site. In sleepy lizards, sex-based differences in homing behaviour are unlikely to be attributable to differences in the area of familiarity, or availability of orientation mechanisms. However differences in homing motivation may explain these differences. Males may miss mating if absent from the home range during the pre-pairing and pairing periods, while females may still be able to obtain a mating even when absent. Females however may be more motivated than males to return to the familiar home range during the post-pairing period to ensure efficient feeding during internal embryo development. Received: 16 February 1998 / Accepted after revision: 28 March 1998     

Comparing discriminant analysis, neural networks and logistic regression for predicting species distributions: a case study with a Himalayan river bird

We assessed the occurrence of a common river bird, the Plumbeous Redstart Rhyacornis fuliginosus, along 180 independent streams in the Indian and Nepali Himalaya. We then compared the performance of multiple discrimant analysis (MDA), logistic regression (LR) and artificial neural networks (ANN) in predicting this species’ presence or absence from 32 variables describing stream altitude, slope, habitat structure, chemistry and invertebrate abundance. Using the entire data (=training set) and a threshold for accepting presence in ANN and LR set to P≥0.5, ANN correctly classified marginally more cases (88%) than either LR (83%) or MDA (84%). Model performance was assessed from two methods of data partitioning. In a ‘leave-one-out’ approach, LR correctly predicted more cases (82%) than MDA (73%) or ANN (69%). However, in a holdout procedure, all the methods performed similarly (73–75%). All methods predicted true absence (i.e. specificity in holdout: 81–85%) better than true presence (i.e. sensitivity: 57–60%). These effects reflect species’ prevalence (=frequency of occurrence), but are seldom considered in distribution modelling. Despite occurring at only 36% of the sites, Plumbeous Redstarts are one of the most common Himalayan river birds, and problems will be greater with less common species. Both LR and ANN require an arbitrary threshold probability (often P=0.5) at which to accept species presence from model prediction. Simulations involving varied prevalence revealed that LR was particularly sensitive to threshold effects. ROC plots (received operating characteristic) were therefore used to compare model performance on test data at a range of thresholds; LR always outperformed ANN. This case study supports the need to test species’ distribution models with independent data, and to use a range of criteria in assessing model performance. ANN do not yet have major advantages over conventional multivariate methods for assessing bird distributions. LR and MDA were both more efficient in the use of computer time than ANN, and also more straightforward in providing testable hypotheses about environmental effects on occurrence. However, LR was apparently subject to chance significant effects from explanatory variables, emphasising the well-known risks of models based purely on correlative data.     

Different tactics,one goal: initial reproductive investments of males and females in a small Arctic seabird

Despite a great number of studies on extra-pair paternity in birds, the actual roles of males and females in extra-pair contacts is poorly understood, as detailed behavioural studies comparing the reproductive performance of the two sexes prior to egg laying are relatively scarce. Here, we investigated mating behaviour (copulations and aggressive interactions), time budget and body condition (size-adjusted body mass and baseline corticosterone level) in the little auk (Alle alle), a monogamous and highly colonial, Arctic seabird. We performed the study in a large breeding colony of the little auk in Hornsund (Spitsbergen). We found that the males frequently attempted extra-pair copulations (EPCs), although these contacts were almost always unsuccessful, mostly because of the females’ rejection behaviour. These results clearly indicate that genetic monogamy is maintained through female control. Nevertheless, males tried to protect their paternity by staying in close proximity to their females and aggressively intervening when their mates became involved in EPCs. Compared to females, males also spent more time in the colony guarding nest sites. Despite the apparent sex differences in the time budget and frequency of aggressive interactions, body condition was similar in the two sexes, indicating comparable parental investments during the mating period.     

Alternative reproductive tactics in male common shrews: relationships between mate-searching behaviour,sperm production,and reproductive success as revealed by DNA fingerprinting

The common shrew (Sorex araneus) is a solitary small mammal with a promiscuous mating system. Previous studies of this species suggest that females typically mate multiply, and that males may adopt alternative mate-searching tactics. We studied two generations of common shrews in a population near Oxford, England. Males were found to adopt two different mate-searching tactics. Those classed as type A occupied relatively small exclusive ranges during March, and made repeated long-distance movements to visit female ranges around the time of first oestrus in April. Males classed as type B established large overlapping ranges in areas of relatively high female density during March, and maintained these ranges throughout April. Type B males were larger than type A males at an early stage of sexual maturation, but there was no difference in the adult body size of the two types of male. Type A males had significantly higher epididymal sperm counts than type B males. Paternity analyses of litters born during the first year of the study reveal that the mean number of offspring fathered by type B males was greater than the mean number fathered by type A males. It is concluded that different mate-searching tactics may be conditional upon the timing of sexual maturation. Differences in sperm production are discussed in relation to sperm competition theory.     

Timing of copulation in the molting and reproductive cycles in a grapsid crab,Gaetice depressus (Crustacea: Brachyura)

From the seasonal changes in the amount of semen in the sperm ducts of males and in the fullness of spermatheca of females, the copulation season of Gaetice depressus in central Japan is considered to approximately coincide with the oviposition season, from March to October. The sperm ducts of all the large males examined contained semen, the amount of which did not vary obviously with stages of the molt cycle or with seasons. The genital apertures of females were normally covered with calcified immobile opercula in this species. The opercula decalcified after molting and again shortly before oviposition, and retained mobility for 1 to 3 d. Shortly after molting, soft-shelled females with mobile opercula did not have well-developed ovaries nor did they copulate with males even if they had a chance. In contrast, all of the hard-shelled females with mobile opercula had well-developed ovaries, and laid eggs within 2 d after the decalcification of opercula regardless of the occurrence of copulation. Laboratory experiments showed that the females normally copulated with males before oviposition. Crabs examined in this study were collected from Shirahama, Wakayama Prefecture, Japan between March 1988 and March 1989.     

The status of heavy metal levels in a Ramsar site,Kuala Gula bird sanctuary: the impact of the anthropogenic inputs

Distributions of Cd, Cu, Pb, and Zn in the surface sediments from two sites of the Kuala Gula Bird Sanctuary, Malaysia were monitored for a period of 6 months from October 2006 to March 2007. In December 2006, the concentration of Zn in one location was significantly (p??1) than the other metals at both sites, but in the oxidizable organic fraction it was highest at both sites during October; with mean concentrations of 18?mg?kg^?1 at both locations. In the acid-reducible fraction, high concentrations of Pb (2.3?mg?kg^?1) were detected at station 2 in February 2007, being highest among all four metals at both stations. The acid-reducible fraction found in Pb ranged from 0.10% to 3.1% in both stations. Percentages ranging from 51% to 96% were observed for all four metals in the resistant fraction throughout the sampling period. These results indicate low contributions from anthropogenic sources. The findings constitute a baseline data archive for future reference.