Life-history States Research Articles

Integrated Population Modeling Provides the First Empirical Estimates of Vital Rates and Abundance for Polar Bears in the Chukchi Sea

Large carnivores are imperiled globally, and characteristics making them vulnerable to extinction (e.g., low densities and expansive ranges) also make it difficult to estimate demographic parameters needed for management. Here we develop an integrated population model to analyze capture-recapture, radiotelemetry, and count data for the Chukchi Sea subpopulation of polar bears (Ursus maritimus), 2008–2016. Our model addressed several challenges in capture-recapture studies for polar bears by including a multievent structure reflecting location and life history states, while accommodating state uncertainty. Female breeding probability was 0.83 (95% credible interval [CRI] = 0.71–0.90), with litter sizes of 2.18 (95% CRI = 1.71–2.82) for age-zero and 1.61 (95% CRI = 1.46–1.80) for age-one cubs. Total adult survival was 0.90 (95% CRI = 0.86–0.92) for females and 0.89 (95% CRI = 0.83–0.93) for males. Spring on-ice densities west of Alaska were 0.0030 bears/km2 (95% CRI = 0.0016–0.0060), similar to 1980s-era density estimates although methodological differences complicate comparison. Abundance of the Chukchi Sea subpopulation, derived by extrapolating density from the study area using a spatially-explicit habitat metric, was 2,937 bears (95% CRI = 1,552–5,944). Our findings are consistent with other lines of evidence suggesting the Chukchi Sea subpopulation has been productive in recent years, although it is uncertain how long this will continue given sea-ice loss due to climate change.

Hypothalamic and liver transcriptome from two crucial life-history stages in a migratory songbird.

What is the central question of this study? What are the molecular underpinnings of the seasonal adaptation in a latitudinal migratory songbird? What is the main finding and its importance? We found changes in mRNA levels after a photoperiod-induced alteration of seasonal state in a captive long-distance latitudinal avian migrant. The hypothalamus and liver transcriptomes revealed genes involved in the regulatory and functional pathways between non-migratory and migratory states. Our results provide insights into mechanisms underlying homeostasis during seasonal changes that are conserved across most species, including humans. Very little is understood about genetic mechanisms underlying the onset of spring migration in latitudinal avian migrants. To gain insight into the genetic architecture of the hypothalamus and liver tissues of a long-distance migrant, we examined and compared the transcriptome profile of captive night-migratory black-headed buntings (Emberiza melanocephala) between photoperiod-induced winter non-migratory (WnM) and spring migratory (SM) life-history states under short and long days, respectively. High-throughput 454 pyrosequenced transcripts were mapped initially with reference to the genome of two phylogenetically close species, Taeniopygia guttata and Ficedula albicollis. The F. albicollis genome gave higher annotation results and was used for further analysis. A total of 216 (78 in hypothalamus; 138 in liver) genes were found to be expressed differentially between the WnM and SM life-history states. These genes were enriched for physiological pathways that might be involved in the regulation of seasonal migrations in birds. For example, genes for the ATP binding pathway in the hypothalamus were expressed at a significantly higher level in SM than in the WnM life-history state. Likewise, upregulated genes associated with the myelin sheath and focal adhesion were enriched in the hypothalamus, and those with cell-to-cell junction, intracellular protein transport, calcium ion transport and small GTPase-mediated signal transduction were enriched in the liver. Many of these genes are a part of physiological pathways potentially involved in the regulation of seasonal migration in birds. These results show molecular changes at the regulatory and metabolic levels associated with seasonal transitions in a long-distance migrant and provide the basis for future studies aimed at unravelling the genetic control of migration in birds.

Photic and non-photic regulation of growth, development, and metamorphosis in giant tree frog (Rhacophorus maximus) tadpoles

Amphibians are seasonal breeders living in seasonal environments having time constraints to achieve their annual life history states. Their survival is strongly dependent on successful timing of certain critical life history decisions. Larval amphibian development, growth, and metamorphosis are strongly time-limited due to variations in the seasonal factors. Photoperiod influences wide range of physiological processes in seasonal breeder. Being ectothermic, temperature may also play critical role in development of amphibian larvae. Food also acts as an ultimate factor in many organisms. Most studies of rearing conditions on growth, development, and metamorphosis of amphibian’s tadpoles have been focused on laboratory species; genus Xenopus. Less is known about other seasonally breeding amphibians. In present study, we addressed the effects of photic (light quality) and non-photic cues (temperature and food quality) on growth, development and metamorphosis of giant tree frog (Rhacophorus maximus). Three experiments have been performed. In experiment one effect of light quality (light wavelength), in experiment two effect of food quality and in experiment three effect of temperature on growth, development, and metamorphosis of giant tree frog tadpoles were studied. Our results suggest that light and food quality and high temperature promotes faster growth, development, and metamorphosis in R. maximus tadpoles.

Adrenal responses of large whales: Integrating fecal aldosterone as a complementary biomarker to glucocorticoids

Until now, physiological stress assessment of large whales has predominantly focused on adrenal glucocorticoid (GC) measures. Elevated GC concentrations in feces (fGC) are known to reflect stressful disturbances, such as fishing gear entanglement and human-generated underwater noise, in North Atlantic right whales (Eubalaena glacialis). However, there can be considerable variation in GC production as a function of sex and life history stage, which may confound the interpretation of fGC levels. Additionally, GC antibodies used in immunoassays can cross-react with other fecal metabolites (i.e., non-target steroids), potentially influencing fGC data. Here, aldosterone concentrations (fALD; aldosterone and related metabolites) were measured in fecal samples from right whales (total n=315 samples), including samples from identified individuals of known life history (n=82 individual whales), to evaluate its utility as a complementary biomarker to fGC for identifying adrenal activation. Concentrations of fALD were positively correlated with fGCs in right whales (r=0.59, P<0.001), suggesting concurrent secretion of these hormones by the adrenal gland. However, fALD levels were less influenced by concentrations of reproductive steroids in feces, minimizing the potential confounder of assay cross-reactivity in samples with highly skewed hormone ratios. Across different life history states for right whales, fALD concentrations showed similar patterns to those reported for fGC, with higher levels in pregnant females (35.9±7.6ng/g) followed by reproductively mature males (9.5±0.9ng/g) (P<0.05), providing further evidence of elevated adrenal activation in these groups of whales. The addition of fALD measurement as a biomarker of adrenal activation may help distinguish between intrinsic and external causes of stress hormone elevations in large whales, as well as other free-living wildlife species, providing a more comprehensive approach for associating adrenal activation with specific natural and anthropogenic stressors.

When do acoustic cues matter? Perceived competition and reproductive plasticity over lifespan in a bushcricket

Individuals often modify their behaviour in response to environmental cues and their own condition. Here we asked whether males modify ejaculates based on information from the sociosexual environment and their physiological age, and how those two factors may interact. We used two populations of the chorusing bushcricket, Ephippiger diurnus, to test whether males strategically adjust large, costly spermatophores they transfer to females during mating based on experience of rivals' calls, and whether males change their investment strategies with age. Males broadcast highly consistent, rhythmically repeated calls of syllables in daily choruses. Populations differ in average syllable number, an important trait under selection via female preference, and females from our two populations prefer calls with more syllables than the population mean. We reared males in one of five acoustic environments that varied in call syllable number. We then mated males twice, as young and old adults, measuring spermatophore size each time. We found that acoustic experience, age and their interaction all significantly influenced male investment, resulting in reaction norms with different slopes. Young males differentially invested in spermatophores in response to acoustic experience, whereas old males generally invested in larger spermatophores across environments. We then tested for a broad pattern of age-related investment with eight different field-collected populations, finding the majority of old males significantly increased spermatophore investment. Our findings demonstrate that both environmental context and an individual's life history state influence plasticity in reproductive investment, and such adjustments may optimize their reproductive success.

Genetic Drift, Not Life History or RNAi, Determine Long-Term Evolution of Transposable Elements.

Transposable elements (TEs) are a major source of genome variation across the branches of life. Although TEs may play an adaptive role in their host’s genome, they are more often deleterious, and purifying selection is an important factor controlling their genomic loads. In contrast, life history, mating system, GC content, and RNAi pathways have been suggested to account for the disparity of TE loads in different species. Previous studies of fungal, plant, and animal genomes have reported conflicting results regarding the direction in which these genomic features drive TE evolution. Many of these studies have had limited power, however, because they studied taxonomically narrow systems, comparing only a limited number of phylogenetically independent contrasts, and did not address long-term effects on TE evolution. Here, we test the long-term determinants of TE evolution by comparing 42 nematode genomes spanning over 500 million years of diversification. This analysis includes numerous transitions between life history states, and RNAi pathways, and evaluates if these forces are sufficiently persistent to affect the long-term evolution of TE loads in eukaryotic genomes. Although we demonstrate statistical power to detect selection, we find no evidence that variation in these factors influence genomic TE loads across extended periods of time. In contrast, the effects of genetic drift appear to persist and control TE variation among species. We suggest that variation in the tested factors are largely inconsequential to the large differences in TE content observed between genomes, and only by these large-scale comparisons can we distinguish long-term and persistent effects from transient or random changes.

Seasonal neuronal plasticity in song-control and auditory forebrain areas in subtropical nonmigratory and palearctic-indian migratory male songbirds.

This study examines whether differences in annual life-history states (LHSs) among the inhabitants of two latitudes would have an impact on the neuronal plasticity of the song-control system in songbirds. At the times of equinoxes and solstices during the year (n = 4 per year) corresponding to different LHSs, we measured the volumetric changes and expression of doublecortin (DCX; an endogenous marker of the neuronal recruitment) in the song-control nuclei and higher order auditory forebrain regions of the subtropical resident Indian weaverbirds (Ploceus philippinus) and Palearctic-Indian migratory redheaded buntings (Emberiza bruniceps). Area X in basal ganglia, lateral magnocellular nucleus of the anterior nidopallium (LMAN), HVC (proper name), and robust nucleus of the arcopallium (RA) were enlarged during the breeding LHS. Both round and fusiform DCX-immunoreactive (DCX-ir) cells were found in area X and HVC but not in LMAN or RA, with a significant seasonal difference. Also, as shown by increase in volume and by dense, round DCX-ir cells, the neuronal incorporation was increased in HVC alone during the breeding LHS. This suggests differences in the response of song-control nuclei to photoperiod-induced changes in LHSs. Furthermore, DCX immunoreactivity indicated participation of the cortical caudomedial nidopallium and caudomedial mesopallium in the song-control system, albeit with differences between the weaverbirds and the buntings. Overall, these results show seasonal neuronal plasticity in the song-control system closely associated with annual reproductive LHS in both of the songbirds. Differences between species probably account for the differences in the photoperiod-response system between the relative refractory weaverbirds and absolute refractory redheaded buntings. J. Comp. Neurol. 524:2914-2929, 2016. © 2016 Wiley Periodicals, Inc.

Diurnal variation in corticosterone release among wild tropical forest birds.

BackgroundGlucocorticoids are adrenal steroid hormones essential to homeostatic maintenance. Their daily variation at low concentrations regulates physiology and behavior to sustain proper immunological and metabolic function. Glucocorticoids rise well above these baseline levels during stress to elicit emergency-state responses that increase short-term survival. Despite this essence in managing life processes under both regular and adverse conditions, relationships of glucocorticoid release to environmental and intrinsic factors that vary at daily and seasonal scales are rarely studied in the wild.MethodsThis study on 41 passerine species of the Ecuadorian Chocó applied a standardized capture-and-restraint protocol to examine diurnal variation in baseline and stress-related release of corticosterone, the primary avian glucocorticoid. Tests for relationships to relative body mass, hemoglobin concentration, molt status and date complemented this evaluation of the time of day effect on corticosterone secretion in free-living tropical rainforest birds. Analyses were also partitioned by sex as well as performed separately on two common species, the wedge-billed woodcreeper and olive-striped flycatcher.ResultsInterspecific analyses indicated maximum baseline corticosterone levels at the onset of the active phase and reductions thereafter. Stress-related levels did not correspond to time of day but accompanied baseline reductions during molt and elevations in birds sampled later during the September - November study period. Baseline corticosterone related negatively to hemoglobin in the wedge-billed woodcreeper and stress-related levels increased with body mass in the olive-striped flycatcher. There were no substantial sex-related differences.ConclusionsThe results of this study suggest a diurnal rhythmicity in baseline corticosterone release so robust as to emerge in pooled analyses across a highly variable dataset. While this detection in nature is singular, correspondent patterns have been demonstrated outside of the tropics in captive model species. Congruity in daily rhythms and links to physiological and life-history state across disparate taxa and environments may promote the yet unresolved utility of corticosterone release as a global metric for population health. However, certain results of this study also deviate from laboratory and field research at higher latitudes, cautioning generalization. Environmental distinctions such as high productivity and tempered seasonality may precipitate unique life-history strategies and underlying hormonal mechanisms in tropical rainforest birds.Electronic supplementary materialThe online version of this article (doi:10.1186/s12983-016-0151-3) contains supplementary material, which is available to authorized users.

Biomass partitioning in grassland plants along independent gradients in temperature and precipitation

How plants allocate biomass to different parts strongly affects vegetation dynamics and ecosystem processes and services such as productivity and carbon storage. We tested the hypothesis that plant size explains the majority of variation in the size of plant parts (as predicted by allometric partitioning theory, APT) and that additional variation is explained by optimal responses for a given individual reproductive state and environment (as predicted by optimal partitioning theory, OPT) for alpine-lowland species pairs from three genera of grassland plants (Veronica, Viola and Carex) sampled along orthogonal gradients in temperature and precipitation. We found general patterns of allometric scaling (allometric exponents) of roots, stems, leaves and flowers, more or less as predicted by APT, and these patterns remained fairly constant across temperature and precipitation gradients. In contrast, basic allocation (allometric coefficients) was clearly related to climate, such as less allocation to leaves but more to roots, stems and flowers with increasing temperatures, in accordance with OPT. Furthermore, our results show that basic allocation is related to habitat affinity (alpine, lowland) and individual life-history states (reproductive or not) and that there is greater variability in small plants, which suggests that biomass partitioning theory should consider both the life-history and ecology of small plants to accurately predict climate-related grassland plant allocation and its implications.

Annual life-history dependent seasonal differences in neural activity of the olfactory system between non-migratory and migratory songbirds

Present study investigated seasonal plasticity in neural activity of the olfactory system, and assessed whether this was influenced by differences in seasonal life-history states (LHSs) between the non-migratory and migratory birds. Brains of non-migratory Indian weaver birds and migratory redheaded buntings were processed for ZENK immunohistochemistry, a marker of neuronal activation, at the times of equinoxes (March, September) and solstices (June, December), which correspond with the periods of different seasonal LHSs during the year. Immunoreactivity was quantified in brain regions comprising the olfactory system viz. olfactory bulb (OB), anterior olfactory nucleus (AON), prepiriform cortex (CPP), lateral olfactory tract (LOT) and olfactory cortex (piriform cortex, CPI; lateral olfactory cortex, LOC). In weaver birds, ZENK-like immunoreactive (ZENK-lir) cells were significantly higher in all the brain areas during post-breeding season (September) than during the other seasons; OBs had higher neuronal activity in the breeding season (June), however. A similar neural activity pattern but at enhanced levels was found in migratory buntings almost all the year. These results for the first time show LHS-associated differences in the seasonal plasticity of a sensory system between the non-migratory and migratory songbirds.

Circadian timing in central and peripheral tissues in a migratory songbird: dependence on annual life-history states.

Predictable seasonal change in photoperiod triggers a sequential change in the daily activity-rest pattern, adaptive for migration in several bird species. The night-migratory black-headed bunting (Emberiza melanocephala) is day active under short photoperiods (8 h light:16 h dark, short day sensitive). Under long photoperiods (16 h light:8 h dark), the buntings are initially day active (long day premigratory) but subsequently become intensely night active (long day migratory) and after few weeks again return to a day active pattern (long day refractory). However, it is unclear how the daily expression of circadian genes changes during photoperiod-induced seasonal life-history states (LHSs). We measured period 2 (Per2), cryptochrome 1 (Cry1), brain and muscle arnt-like protein 1 (Bmal1), and circadian locomotor output cycles kaput (Clock) mRNA expressions in various neural and peripheral tissues of buntings in different LHSs and discovered differences of ∼2 to 6 h in the phase and 2- to 4-fold in amplitude of circadian oscillations of Per2, Cry1, and Bmal1 between photoperiod-induced LHSs. Phase relationship in mRNA oscillations was altered between oscillator components in the circadian pacemaker system (retina, pineal, hypothalamus) as well as in the peripheral (liver, muscle) tissues. These results show for the first time altered waveforms of clock gene expressions in all tissues in parallel with behavioral shifts and suggest the involvement of circadian system in photoperiod induction of seasonal LHSs in a migratory species.

Hypothalamic gene switches control transitions between seasonal life history states in a night-migratory photoperiodic songbird

This study investigated photoperiodic plasticity in hypothalamic expression of genes implicated in the photoperiodic light perception (rhodopsin, melanopsin, neuropsin and peropsin), transduction (pax6, bmal1, clock, per2 and casr), induction (eya3, tshβ, dio2 and dio3, gnrh and gnih) and metabolism (NPY, sirtuin1, foxO1, hmgcr, citrate synthase and dehydrogenases) in photosensitive and photorefractory redheaded buntings. There was a significant increase in eya3, tsh β, dio2, pax6 and rhodopsin and decrease in dio3 mRNA expression at hour 15 and/or 19 on the day photosensitive buntings were subjected to a 13- or 16 h, but not to 8- and 11 h light exposure. Downstream reproductive and metabolic gene expression was not altered, except for an increase in those genes coding for succinate and malate dehydrogenase enzymes involved in lipogenesis. Photorefractory buntings had high dio3 mRNA expression which significantly declined after 1 short day exposure, suggesting possible involvement of dio3 in the maintenance of photorefractoriness. Positive correlation of rhodopsin on eya 3 and tshβ indicates its role in photoperiodic timing, perhaps involving the peropsin and pax6 genes. These results suggest that rapid switching of hypothalamic gene expression underlies photoperiod-induced seasonal plasticity and regulates transitions from photosensitive to photostimulated and from photorefractory to photosensitive states in migratory songbirds.

Annual life history-dependent gene expression in the hypothalamus and liver of a migratory songbird: insights into the molecular regulation of seasonal metabolism.

Birds seasonally switch from one life history state (LHS) to another to maximize their fitness. Accordingly, they exhibit distinct differences in their physiological and behavioral phenotypes between seasons. Possible molecular mechanisms underlying changes through the seasons have scarcely been examined in migratory birds. The present study measured key genes suggested to be involved in the metabolic regulation of 4 photoperiodically induced seasonal LHSs in a long-distance migratory songbird, the blackheaded bunting (Emberiza melanocephala). Buntings were held under short days (8 h light:16 h darkness, 8L:16D), during which they maintained the winter nonmigratory phenotype. Then they were exposed for several weeks to long days (13L:11D). Differences in the activity-rest pattern, body fattening and weight gain, testis size, organ (heart, intestine) weights, and blood glucose and triglyceride levels confirmed that buntings sequentially exhibited spring migration-linked premigratory, migratory, and postmigratory LHSs under long days. The mRNA levels of circadian genes involved in metabolism (Bmal1, Clock, Npas2, Rorα, and Rev-erbα) and of genes that encode for proteins/enzymes involved in the regulation of glucose (Sirt1, FoxO1, Glut1, and Pygl) and lipids (Hmg-CoA; Pparα, Pparγ; Fasn and Acaca) showed LHS-dependent changes in their light-dark expression patterns in the hypothalamus and liver. These initial results on genetic regulation of metabolism in a migratory species extend the idea that the transitions between LHSs in a seasonal species are accomplished by changes at multiple regulatory levels. Thus, these findings promise new insights into the mechanism(s) of adaptation to seasons in higher vertebrates.

Life at a different pace: annual itineraries are conserved in seasonal songbirds.

The duration of life history state (LHS) reflects the adaptive strategy a species has evolved to cope with a changing environment. Inhabitants at different latitudes may thus have significant differences in the rates of metabolic and physiological processes underlying LHSs. Birds, in order to maximize their fitness in the environment in which they live, seasonally switch from one LHS to another during the year. The present study investigated whether an annual itinerary of a species would determine its rate of reaction to inductive long days. We compared the photoinduced cycles of changes in body mass and testes, as indices of migratory and reproductive LHSs, between two long day breeding species, the migratory redheaded bunting and non-migratory Indian weaverbird. Changes in body mass and testis size were measured in photosensitive buntings and weaverbirds (n = 7 each) on short days (LD 8:16) subjected first to 0.5 h weekly light increments until the light period was 13 h per day, and then maintained on LD 13:11 for another 32 weeks. A similar observation was recorded on a group of buntings (n = 14) and weaverbirds (n = 9) maintained on increasing natural day lengths (NDL; Lucknow, 26°55' N, 80°59' E) for 47 weeks. As predicted, the rates of induction of seasonal cycles under an identical inductive photoperiod were significantly faster in temperate buntings with five annual LHSs than in the subtropical weaverbirds with three annual LHSs. This suggests that annual itineraries of songbirds with which they may have evolved with at their breeding latitudes, determine their response to the external photoperiodic environment.

Flight Performance and Feather Quality: Paying the Price of Overlapping Moult and Breeding in a Tropical Highland Bird

A temporal separation of energetically costly life history events like reproduction and maintenance of the integumentary system is thought to be promoted by selection to avoid trade-offs and maximize fitness. It has therefore remained somewhat of a paradox that certain vertebrate species can undergo both events simultaneously. Identifying potential costs of overlapping two demanding life history stages will further our understanding of the selection pressures that shape the temporal regulation of life history events in vertebrates. We studied free-living tropical Slaty brush-finches (Atlapetes schistaceus), in which individuals spontaneously overlap reproduction and moult or undergo both events in separation. To assess possible costs of such an overlap we quantified feather quality and flight performance of individuals in different states. We determined individual’s life history state by measuring gonad size and scoring moult stage, and collected a newly grown 7th primary wing feather for later analysis of feather quality. Finally, we quantified flight performance for each individual in the wild. Overlapping individuals produced lighter and shorter wing feathers than individuals just moulting, with females decreasing feather quality more strongly during the overlap than males. Moreover, overlapping individuals had a reduced flight speed during escape flights, while their foraging flight speed was unaffected. Despite overlappers being larger and having a smaller wing area, their lower body mass resulted in a similar wing load as in breeders or moulters. Individuals measured repeatedly in different states also showed significant decreases in feather quality and escape flight speed during the overlap. Reduced escape flight speed may represent a major consequence of the overlap by increasing predation risk. Our data document costs to undergoing two life history stages simultaneously, which likely arise from energetic trade-offs. Impairments in individual quality and performance may represent important factors that select for temporal separation of life history stages in other species.

Ascaroside signaling in C. elegans

Over the past 10 years, the relevance of small-molecule signaling for many aspects of C. elegans development and behavior has become apparent. One prominent group of small-molecule signals are the ascarosides, which control dauer entry and exit as well as a variety of sex-specific and social behaviors, including male attraction, hermaphrodite repulsion, olfactory plasticity, and aggregation. This wide range of biological functions is facilitated by a great diversity of ascaroside chemical structures. These are based on the sugar ascarylose, which is linked to fatty acid-like side chains of varying lengths and often decorated further with building blocks derived from amino acids, folate, and other primary metabolites. Different ascarosides or combinations of ascarosides mediate different phenotypes, and even small differences in chemical structures are often associated with strongly altered activity profiles. Additional complexity arises from concentration-dependent effects and synergism between different ascarosides. The ascarosides are sensed by several types of chemosensory head neurons, including the ASK, ASI, and ADL neurons as well as the male-specific CEM neurons. Ascaroside perception is mediated by diverse families of G-protein coupled membrane receptors that act upstream of conserved signal transduction pathways, including insulin/IGF-1 signaling and transforming growth factor beta (TGF-β) signaling. Biosynthesis of the ascarosides appears to integrate input from several primary metabolic pathways, including peroxisomal β-oxidation of long-chain fatty acids and amino acid catabolism. Life stage, sex, as well as food availability and other environmental factors affect ascaroside biosynthesis, suggesting that ascaroside signaling communicates detailed information about life history and metabolic state.

The influence of winter severity, predation and senescence on moose habitat use

Habitat use is widely known to be influenced by abiotic and biotic factors, such as climate, population density, foraging opportunity and predation risk. The influence of the life-history state of an individual organism on habitat use is less well understood, especially for terrestrial mammals. There is good reason to expect that life-history state would affect habitat use. For example, organisms exhibiting poor condition associated with senescence have an increased vulnerability to predation and that vulnerability is known to alter habitat use strategies. We assessed the influence of life-history stage on habitat use for 732 moose (Alces alces) killed by wolves (Canis lupus) over a 50-year period in Isle Royale National Park, an island ecosystem in Lake Superior, USA. We developed regression models to assess how location of death was associated with a moose's life-history stage (prime-aged or senescent), presence or absence of senescent-associated pathology (osteoarthritis and jaw necrosis), and annual variation in winter severity, moose density and ratio of moose to wolves, which is an index of predation risk. Compared to senescent moose, prime-aged moose tend to make greater use of habitat farther from the shoreline of Isle Royale. That result is ecologically relevant because shoreline habitat on Isle Royale tends to provide better foraging opportunities for moose but is also associated with increased predation risk. During severe winters prime-aged moose tend to make greater use of habitat that is closer to shore in relation to senescent-aged moose. Furthermore, moose of both age classes were more likely to die in riskier, shoreline habitat during years when predation risk was lower in the preceding year. Our results highlight a complicated connection between life history, age-structured population dynamics and habitat-related behaviour. Our analysis also illustrates why intraspecific competition should not be the presumed mechanism underlying density-dependent habitat use, if predation risk is related to density, as it is expected to be in many systems.

From interference to predation: type and effects of direct interspecific interactions of small mammals

Indirect exploitative competition, direct interference and predation are important interactions affecting species coexistence. These interaction types may overlap and vary with the season and life-history state of individuals. We studied effects of competition and potential nest predation by common shrews (Sorex araneus) on lactating bank voles (Myodes glareolus) in two seasons. The species coexist and may interact aggressively. Additionally, shrews can prey on nestling voles. We studied bank vole mothers’ spatial and temporal adaptations to shrew presence during summer and autumn. Further, we focused on fitness costs, e.g. decreased offspring survival, which bank voles may experience in the presence of shrews. In summer, interference with shrews decreased the voles’ home ranges and they spent more time outside the nest, but there were no effects on offspring survival. In autumn, we found decreased offspring survival in enclosures with shrews, potentially due to nest predation by shrews or by increased competition between species. Our results indicate a shift between interaction types depending on seasonal constraints. In summer, voles and shrews seem to interact mainly by interference, whereas resource competition and/or nest predation by shrews gain importance in autumn. Different food availability, changing environmental conditions and the energetic constraints in voles and shrews later in the year may be the reasons for the varying combinations of interaction types and their increasing effects on the inclusive fitness of bank voles. Our study provides evidence for the need of studies combining life history with behavioural measurements and seasonal constraints.

The role of androgen receptors in regulating territorial aggression in male song sparrows

This paper examines the role that androgen receptors (ARs) play in modulating aggressive behavior in male song sparrows, Melospiza melodia morphna. Song sparrows are seasonally breeding, territorial birds that maintain year-round territories with male–female pair bonds formed during the spring breeding season. Plasma testosterone levels peak as territories are established and mates acquired. In late summer, testosterone levels fall and remain basal during the non-breeding season. We examined the role of ARs in regulating territorial aggression in captive song sparrows under short- and long-day conditions as well as just prior to, and at the start of the breading season in freely living birds using the nonsteroidal antiandrogen flutamide to block AR function. Birds were implanted with either empty or drug filled silastic implants for 18 to 42 days and then challenged with a novel male decoy to assess the individual birds level of male–male aggression. Freely living birds remained on their home territory and underwent a simulated territorial intrusion, whereas laboratory-held birds were assessed using a laboratory simulated territorial intrusion and remained in their home cage. Experimental treatment of male song sparrows decreased aggressive behavior during the pre-breeding life history substage (March–April) in freely living birds as well as in laboratory-held birds under long-day (16L:8D) conditions. During the early breeding substage (April–May) there was no measurable effect of flutamide treatment on aggressive behavior, nor was there a difference in behavior in the (8L:16D) laboratory birds. This demonstrates that ARs are an important component of the neuroendocrine control of aggressive behavior. Given that flutamide only affected aggression during the pre-breeding substage and in LD birds, the results suggest that AR dependent control of aggressive behavior changes as song sparrow life history states change.

Sampling design considerations for demographic studies: a case of colonial seabirds

For the purposes of making many informed conservation decisions, the main goal for data collection is to assess population status and allow prediction of the consequences of candidate management actions. Reducing the bias and variance of estimates of population parameters reduces uncertainty in population status and projections, thereby reducing the overall uncertainty under which a population manager must make a decision. In capture-recapture studies, imperfect detection of individuals, unobservable life-history states, local movement outside study areas, and tag loss can cause bias or precision problems with estimates of population parameters. Furthermore, excessive disturbance to individuals during capture-recapture sampling may be of concern because disturbance may have demographic consequences. We address these problems using as an example a monitoring program for Black-footed Albatross (Phoebastria nigripes) and Laysan Albatross (Phoebastria immutabilis) nesting populations in the northwestern Hawaiian Islands. To mitigate these estimation problems, we describe a synergistic combination of sampling design and modeling approaches. Solutions include multiple capture periods per season and multistate, robust design statistical models, dead recoveries and incidental observations, telemetry and data loggers, buffer areas around study plots to neutralize the effect of local movements outside study plots, and double banding and statistical models that account for band loss. We also present a variation on the robust capture-recapture design and a corresponding statistical model that minimizes disturbance to individuals. For the albatross case study, this less invasive robust design was more time efficient and, when used in combination with a traditional robust design, reduced the standard error of detection probability by 14% with only two hours of additional effort in the field. These field techniques and associated modeling approaches are applicable to studies of most taxa being marked and in some cases have individually been applied to studies of birds, fish, herpetofauna, and mammals.