Spatial and temporal factors associated with nest survival of Gray Flycatchers in managed ponderosa pine forests

Gray Flycatchers (Empidonax wrightii) breed in a variety of habitats in the arid and semi‐arid regions of the western United States, but little is known about their breeding biology, especially in the northern portion of their range where they nest in ponderosa pine (Pinus ponderosa) forests. From May to July 2014 and 2015, we conducted surveys for singing male Gray Flycatchers along the eastern slope of the Cascade Range in Washington, U.S.A, monitored flycatcher nests, and quantified nest‐site vegetation. We used a logistic‐exposure model fit within a Bayesian framework to model the daily survival probability of flycatcher nests. During the 2 yr of our study, we monitored 141 nests, with 93% in ponderosa pines. Mean clutch size was 3.6 eggs and the mean number of young fledged per nest was 3.2. Predation accounted for 90% of failed nests. We found a positive association between daily nest survival and both nest height and distance of nest substrates from the nearest tree. Flycatchers that locate their nests higher above the ground and further from adjacent trees may be choosing the safest alternative because higher nests may be less exposed to terrestrial predators and nests in trees that are farther from other trees may be less exposed to arboreal predators such as jays (Corvidae) that may forage in patches with connected canopies. Nests in trees farther from other trees may also allow earlier detection of approaching predators and thus aid in nest defense.     

Variation in maternal effects and embryonic development rates among passerine species

Embryonic development rates are reflected by the length of incubation period in birds, and these vary substantially among species within and among geographical regions. The incubation periods are consistently shorter in North America (Arizona study site) than in tropical (Venezuela) and subtropical (Argentina) South America based on the study of 83 passerine species in 17 clades. Parents, mothers in particular, may influence incubation periods and resulting offspring quality through proximate pathways, while variation in maternal strategies among species can result from selection by adult and offspring mortality. Parents of long-lived species, as is common in the tropics and subtropics, may be under selection to minimize costs to themselves during incubation. Indeed, time spent incubating is often lower in the tropical and subtropical species than the related north temperate species, causing cooler average egg temperatures in the southern regions. Decreased egg temperatures result in longer incubation periods and reflect a cost imposed on offspring by parents because energy cost to the embryo and risk of offspring predation are both increased. Mothers may adjust egg size and constituents as a means to partially offset such costs. For example, reduced androgen concentrations in egg yolks may slow development rates, but may enhance offspring quality through physiological trade-offs that may be particularly beneficial in longer-lived species, as in the tropics and subtropics. We provide initial data to show that yolks of tropical birds contain substantially lower concentrations of growth-promoting androgens than north temperate relatives. Thus, maternal (and parental) effects on embryonic development rates may include contrasting and complementary proximate influences on offspring quality and deserve further field study among species.     

Mutual ornamentation and the parental behaviour of male and female Collared Flycatchers Ficedula albicollis during incubation

One of the benefits of mate choice based on sexually selected traits is the greater investment of more ornamented individuals in parental care. The choosy individual can also adjust its parental investment to the sexual signals of its partner. Incubation is an important stage of avian reproduction, but the relationship between behaviour during incubation and mutual ornamentation is unclear. Studying a population of Collared Flycatchers Ficedula albicollis, we monitored the behaviour of both sexes during incubation in relation to their own and their partner's plumage traits, including plumage‐level reflectance attributes and white patch sizes. There was a marginally positive relationship between male feeding rate during incubation and female incubation rate. Female but not male behavioural traits were associated with the laying date of the first egg and clutch size. The behaviour of the two sexes jointly determined the relative hatching speed of clutches and the hatching success of eggs. Females with larger white wing patches spent less time incubating eggs and left the nestbox more frequently. Males with larger white wing patches fed females less frequently, whereas males with brighter white plumage areas visited the nestbox more regularly without feeding. Females tended to leave the nest less often when mated to males with larger wing patches, and females spent less time incubating when males had more UV chromatic plumage. The behaviour of both partners during incubation therefore predicted hatching patterns and was correlated with their own and sometimes with their partner's plumage ornamentation. These results call for further studies of mutual ornamentation and reproductive effort during incubation.     

Growth rate variation among passerine species in tropical and temperate sites: an antagonistic interaction between parental food provisioning and nest predation risk

Causes of interspecific variation in growth rates within and among geographic regions remain poorly understood. Passerine birds represent an intriguing case because differing theories yield the possibility of an antagonistic interaction between nest predation risk and food delivery rates on evolution of growth rates. We test this possibility among 64 Passerine species studied on three continents, including tropical and north and south temperate latitudes. Growth rates increased strongly with nestling predation rates within, but not between, sites. The importance of nest predation was further emphasized by revealing hidden allometric scaling effects. Nestling predation risk also was associated with reduced total feeding rates and per-nestling feeding rates within each site. Consequently, faster growth rates were associated with decreased per-nestling food delivery rates across species, both within and among regions. These relationships suggest that Passerines can evolve growth strategies in response to predation risk whereby food resources are not the primary limit on growth rate differences among species. In contrast, reaction norms of growth rate relative to brood size suggest that food may limit growth rates within species in temperate, but not tropical, regions. Results here provide new insight into evolution of growth strategies relative to predation risk and food within and among species.     

Are species differences in maternal effects arising from maternal care adaptive?

Parental care benefits offspring through maternal effects influencing their development, growth and survival. However, although parental care in general is likely the result of adaptive evolution, it does not follow that specific differences in the maternal effects that arise from care are also adaptive. Here, we used an interspecific cross‐fostering design in the burying beetle species Nicrophorus orbicollis and N. vespilloides, both of which have elaborate parental care involving direct feeding of regurgitated food to offspring, to test whether maternal effects are optimized within a species and therefore adaptive. Using a full‐factorial design, we first demonstrated that N. orbicollis care for offspring longer regardless of recipient species. We then examined offspring development and mass in offspring reared by hetero‐ or conspecific parents. As expected, there were species‐specific direct effects independent of the maternal effects, as N. orbicollis larvae were larger and took longer to develop than N. vespilloides regardless of caregiver. We also found significant differences in maternal effects: N. vespilloides maternal care caused more rapid development of offspring of either species. Contrary to expectations if maternal effects were species‐specific, there were no significant interactions between caretaker and recipient species for either development time or mass, suggesting that these maternal effects are general rather than optimized within species. We suggest that rather than coadaptation between parents and offspring performance, the species differences in maternal effects may be correlated with direct effects, and that their evolution is driven by selection on those direct effects.     

Sex,growth rate,rank order after brood reduction,and hatching date affect first‐year survival of long‐lived Herring Gulls

Among most species of birds, survival from hatching throughout the first year of life is generally lower than subsequent survival rates. Survival of young birds during their first year may depend on a combination of selection, learning, unpredictable resources, and environmental events (i.e., post‐fledging factors). However, knowledge about post‐fledging development in long‐lived species is usually limited due to a lengthy immature stage when individuals are generally unobservable. Therefore, pre‐fledging characteristics are often used to predict the survival of young birds. We assessed effects of nestling growth rates, hatching date, hatching asynchrony, brood size and rank order after brood reduction, and sex on first‐year survival of 137 fledglings using a mark‐resighting analysis. We found that the survival probability (Φ_1yr = 0.39) of first‐year Herring Gulls (Larus argentatus) in our study colony located at the outer port of Zeebrugge (Belgium) was lower than that of older individuals (Φ_>1yr = 0.75). All 10 models best supported by our data included nestling growth rate, suggesting that variability in first‐year survival may be linked primarily to individual variation in growth. First‐year survival was negatively correlated with hatching date and rank order after brood reduction. Hence, carry‐over effects of breeding season events such as timing of breeding, early development, and social status had an influence on survival of Herring Gulls after fledging. Furthermore, we found sex‐biased mortality in first‐year Herring Gulls, with females (Φ_1yr = 0.45) surviving better than males (Φ_1yr = 0.38). Although adult survival is generally regarded as the key parameter driving population trajectories in long‐lived species, juvenile survival has recently been acknowledged as an important source of variability in population growth rates. Thus, increasing our knowledge of factors affecting age‐specific survival rates is necessary to improve our understanding of population dynamics and ultimately life‐history variation.     

Distinct preoptic‐BST nuclei dissociate paternal and infanticidal behavior in mice

Paternal behavior is not innate but arises through social experience. After mating and becoming fathers, male mice change their behavior toward pups from infanticide to paternal care. However, the precise brain areas and circuit mechanisms connecting these social behaviors are largely unknown. Here we demonstrated that the c‐Fos expression pattern in the four nuclei of the preoptic‐bed nuclei of stria terminalis (BST) region could robustly discriminate five kinds of previous social behavior of male mice (parenting, infanticide, mating, inter‐male aggression, solitary control). Specifically, neuronal activation in the central part of the medial preoptic area (cMPOA) and rhomboid nucleus of the BST (BSTrh) retroactively detected paternal and infanticidal motivation with more than 95% accuracy. Moreover, cMPOA lesions switched behavior in fathers from paternal to infanticidal, while BSTrh lesions inhibited infanticide in virgin males. The projections from cMPOA to BSTrh were largely GABAergic. Optogenetic or pharmacogenetic activation of cMPOA attenuated infanticide in virgin males. Taken together, this study identifies the preoptic‐BST nuclei underlying social motivations in male mice and reveals unexpected complexity in the circuit connecting these nuclei.     

Brood discrimination, nest mate discrimination, and determinants of social behavior in facultatively quasisocial beetles (Nicrophorus spp.)

In this study we investigated ecological determinants of socialityin burying beetles (Nicrophorus spp.), potential conflicts ofinterest among reproductive females, and the effects of nestingfailure and costs of fighting on cooperation. Burying beetlesare known to form monogamous pairs when exploiting small vertebratecarcasses. More complex social behavior in this group is poorlyunderstood. We conducted experiments in which one or two females(N. defodiens, N. orbicollis) were provided small or large carcasseson which to breed. On large but not on small carcasses, twofemales often formed cooperative breeding associations (jointlyprepared a carcass and fed young). In N. defodiens, but notN. orbicollis, two females produced a larger brood than singlefemales on large carcasses. In both species, the reproductiveoutput per female was less for two than for one female. Thepresence of a second female did not decrease the preparationtime of a carcass (discovery of resource to egg hatch). Conflictwas evident between females. Trials employing females of similarsize were more likely to result in injury than trials usingfemales of dissimilar size (N. tomentosus, N. defodiens, N.orbicollis). In N. tomentosus, those associations that persistedthe longest resulted in the fewest injuries. After care of youngwas initiated, conflict among familiar nest mates was not observed.There was no evidence that breeding females could discriminatebetween brood; use of a genetic marker (N. orbicollis) demonstratedthat females fed related and unrelated young alike. Femalesof similar size (high potential cost of fighting for the dominantindividual) were not more likely to form cooperative breedingassociations than females of dissimilar size (low cost of fightingfor dominant). Females of a species subject to a high rate ofnest failure (N. defodiens) were more likely to cooperate thanfemales of a species with a low rate of nest failure (N. orbicollis).It is argued that limited reproductive opportunities, difficultyin controlling rivals' access to a large carcass, and the superabundantlarval food supply represented by a large carcass, but not kinselection, have contributed to the evolution of cooperativebehavior in this group. In addition, we hypothesize that beetlesmight initially tolerate consexual rivals on large carcasseswhen there is a high likelihood of nesting failure, therebyavoiding potentially costly conflicts.     

Home‐ranges,population densities,vocal behavior,and post‐fledging movements of Cipo Canasteros (Asthenes luizae,Furnariidae), a rock‐specialist endemic of the highlands of eastern Brazil

Cipo Canasteros (Asthenes luizae, Furnariidae) have a fragmented and limited range restricted to the campos rupestres (rupestrian grasslands) habitat in the Brazilian highlands of the Espinhaço Range, and little is known about their behavior, ecology, and population biology. From March 2009 to November 2010, we monitored birds (24 banded and 22 radio‐tracked) at two study sites at Serra do Cipó in the state of Minas Gerais, Brazil, to estimate their home‐range sizes and population density, and describe their habitat use, natal dispersal behavior, and vocal behavior. We found an average density of 8.7 paired adults/km² in our study areas or 22.9 paired adults/km² when considering only used habitats. The sex ratio was male‐biased (males/total adults = 0.68), adults exhibited high site fidelity, home‐ranges averaged 4.0 ha (fixed kernel 95%) or 3.5 ha (95% minimum convex polygon) in size, and both sexes defended territories year‐round. We recorded four main types of songs, including two uttered more often during the breeding season. We monitored the natal dispersal of two males and one female who moved maximum distances of 1238 m, 780 m, and 1056 m, respectively, from natal areas. Our results confirm that Cipo Canasteros are restricted to the rocky‐outcrop habitat of the campos rupestres. In part due to their habitat specialization, Cipo Canasteros are considered Near Threatened, but other factors contributing to their demographic fragility include the small number, and probably low survival, of females and low reproductive success due to predation and brood parasitism by cowbirds. Given these threats, along with their specialized habitat and restricted range, the future conservation of Cipo Canasteros will likely depend on the extent to which their campos rupestres habitat can be conserved.     

Lifetime survival rates and senescence in northern elephant seals

The aim of this study was to extend 40 yr of prior demographic work on northern elephant seals (Mirounga angustirostris) at Año Nuevo, California, by including the oldest animals. We used a Bayesian mark‐recapture analysis to estimate lifelong survival and lifespan of a cohort of 372 weaned pups branded in 1985–1987 and resighted until 2008. Annual survival probability of females averaged 86.3%/yr at ages 5–16, then declined until age 21, the age of the oldest female. Male survival was lower, averaging 67.7%/yr from age 1 to age 15, the age of the oldest male. Northern elephant seal females in the expanding population at Año Nuevo live longer than southern elephant seal females (M. leonina) at colonies whose populations are declining. This comparison suggests that high survival of females is a key factor in population growth.     

Regional and temporal variation in diet and provisioning rates suggest weather limits prey availability for an endangered raptor

Understanding variation in food requirements of wild animals is of central importance in population ecology and conservation, as it helps to identify where and when food may be limiting. Studies on diet variation or prey provisioning rates may give useful insights when direct information on prey availability is lacking. We assess spatial and temporal variation in the diet of an endangered predator, the Black Harrier Circus maurus. This raptor is endemic to southern Africa and specializes on small mammals but also feeds on birds and reptiles as alternative prey. Using data on 1679 prey identified in 953 pellets collected in inland and coastal regions from 2006 to 2015, we show that diet composition changed little throughout the breeding season in the coastal region, whereas there was a marked seasonal decline in the occurrence of small mammal prey in the inland region, with a concomitant increase in alternative prey. The proportion of small mammals in the diet declined with increasing maximum temperature, the latter being highest at the inland region late in the breeding season. Using camera recordings at nests in 2014, we further analysed daily patterns of prey provisioning to nestlings. A marked reduction in small mammal provisioning rates occurred during the middle of the day in the hotter inland region but not in the cooler coastal region. Reduced availability of the primary prey, small mammals, in hotter conditions, through a reduction in activity or overall abundance, could explain these patterns. Finally, we show a positive relationship between winter rainfall and interannual differences in the proportion of small mammals in the diet of Black Harriers breeding in the coastal region, suggesting relationships between diet and prey abundance that are mediated through rainfall. We discuss the need to consider spatial variation in food availability in conservation strategies.     

Factors that influence vital rates of Seaside and Saltmarsh sparrows in coastal New Jersey,USA

As saltmarsh habitat continues to disappear, understanding the factors that influence the population dynamics of saltmarsh breeding birds is an important step in the conservation of these declining species. Using 5 yrs (2011–2015) of demographic data, we evaluated and compared apparent adult survival and nest survival of Seaside (Ammodramus maritimus) and Saltmarsh (A. caudacutus) sparrows at the Edwin B. Forsythe National Wildlife Refuge in New Jersey, USA. We determined the effect of site management history (unditched vs. ditched marshes) on adult and nest survival to aid in prioritizing future management or restoration actions. Apparent adult survival (61.6%, 95% CI: 52.5–70.0%) of Seaside Sparrows averaged > 1.5 times greater than that of Saltmarsh Sparrows (39.9%, 95% CI: 34.0–46.2%). Nest survival and predation and flooding rates did not differ between species, and predation was the primary cause of nest failure for both species. Apparent adult survival and nest survival did not differ between unditched and ditched marshes for either species, indicating that marsh ditching history may not affect the quality of breeding habitat for these species. Because predation was the primary cause of nest failure for both species in New Jersey, we suggest that future studies should focus on identifying predator communities in salt marshes and the potential for implementing predator‐control programs to limit population declines.     

The effect of gut microbiota elimination in Drosophila melanogaster: A how‐to guide for host–microbiota studies

In recent years, there has been a surge in interest in the effects of the microbiota on the host. Increasingly, we are coming to understand the importance of the gut microbiota in modulating host physiology, ecology, behavior, and evolution. One method utilized to evaluate the effect of the microbiota is to suppress or eliminate it, and compare the effect on the host with that of untreated individuals. In this study, we evaluate some of these commonly used methods in the model organism, Drosophila melanogaster. We test the efficacy of a low‐dose streptomycin diet, egg dechorionation, and an axenic or sterile diet, in the removal of gut bacteria within this species in a fully factorial design. We further determine potential side effects of these methods on host physiology by performing a series of standard physiological assays. Our results showed that individuals from all treatments took significantly longer to develop, and weighed less, compared to normal flies. Males and females that had undergone egg dechorionation weighed significantly less than streptomycin reared individuals. Similarly, axenic female flies, but not males, were much less active when analyzed in a locomotion assay. All methods decreased the egg to adult survival, with egg dechorionation inducing significantly higher mortality. We conclude that low‐dose streptomycin added to the dietary media is more effective at removing the gut bacteria than egg dechorionation and has somewhat less detrimental effects to host physiology. More importantly, this method is the most practical and reliable for use in behavioral research. Our study raises the important issue that the efficacy of and impacts on the host of these methods require investigation in a case‐by‐case manner, rather than assuming homogeneity across species and laboratories.     

Evidence for rapid evolution of phenology in an invasive grass

Evolutionary dynamics of integrative traits such as phenology are predicted to be critically important to range expansion and invasion success, yet there are few empirical examples of such phenomena. In this study, we used multiple common gardens to examine the evolutionary significance of latitudinal variation in phenology of a widespread invasive species, the Asian short‐day flowering annual grass Microstegium vimineum. In environmentally controlled growth chambers, we grew plants from seeds collected from multiple latitudes across the species' invasive range. Flowering time and biomass were both strongly correlated with the latitude of population origin such that populations collected from more northern latitudes flowered significantly earlier and at lower biomass than populations from southern locations. We suggest that this pattern may be the result of rapid adaptive evolution of phenology over a period of less than one hundred years and that such changes have likely promoted the northward range expansion of this species. We note that possible barriers to gene flow, including bottlenecks and inbreeding, have apparently not forestalled evolutionary processes for this plant. Furthermore, we hypothesize that evolution of phenology may be a widespread and potentially essential process during range expansion for many invasive plant species.     

An evaluation of behavior inferences from Bayesian state‐space models: A case study with the Pacific walrus

State‐space models offer researchers an objective approach to modeling complex animal location data sets, and state‐space model behavior classifications are often assumed to have a link to animal behavior. In this study, we evaluated the behavioral classification accuracy of a Bayesian state‐space model in Pacific walruses using Argos satellite tags with sensors to detect animal behavior in real time. We fit a two‐state discrete‐time continuous‐space Bayesian state‐space model to data from 306 Pacific walruses tagged in the Chukchi Sea. We matched predicted locations and behaviors from the state‐space model (resident, transient behavior) to true animal behavior (foraging, swimming, hauled out) and evaluated classification accuracy with kappa statistics (κ) and root mean square error (RMSE). In addition, we compared biased random bridge utilization distributions generated with resident behavior locations to true foraging behavior locations to evaluate differences in space use patterns. Results indicated that the two‐state model fairly classified true animal behavior (0.06 ≤ κ ≤ 0.26, 0.49 ≤ RMSE ≤ 0.59). Kernel overlap metrics indicated utilization distributions generated with resident behavior locations were generally smaller than utilization distributions generated with true foraging behavior locations. Consequently, we encourage researchers to carefully examine parameters and priors associated with behaviors in state‐space models, and reconcile these parameters with the study species and its expected behaviors.     

Late‐life and intergenerational effects of larval exposure to microbial competitors in the burying beetle Nicrophorus vespilloides

Intergenerational effects can have either adaptive or nonadaptive impacts on offspring performance. Such effects are likely to be of ecological and evolutionary importance in animals with extended parental care, such as birds, mammals and some insects. Here, we studied the effects of exposure to microbial competition during early development on subsequent reproductive success in the burying beetle Nicrophorus vespilloides, an insect with elaborate parental care. We found that exposure to high levels of microbial competition both during a female's larval development and during her subsequent reproduction resulted in females rearing smaller broods than those exposed to lower levels of microbial competition. To determine whether these differences arose before or after offspring hatching, a cross‐fostering experiment was conducted. Our results demonstrate that the impact of larval competition with microbes for resources extends into adult life and can negatively affect subsequent generations via impacts on the quality of parental care provided after hatching. However, we also find evidence for some positive effects of previous microbial exposure on prehatch investment, suggesting that the long‐term results of competition with microbes may include altering the balance of parental investment between prehatch and post‐hatch care.     

Variation at phenological candidate genes correlates with timing of dispersal and plumage morph in a sedentary bird of prey

Polymorphic genes involved in the conserved molecular signalling of circadian and circannual clocks may play important roles in governing the timing of breeding and dispersal and thereby affect fitness in vertebrates. However, relatively few studies have explored associations between phenological candidate genes and behaviour, and these are somewhat biased towards particular taxonomic groups such as passerine birds and salmonid fish. Consequently, we assayed microsatellite polymorphisms within the exonic and 3′ untranslated regions of the regulatory genes CLOCK, NPAS2, ADCYAP1 and CREB1 in the common buzzard (Buteo buteo), a polymorphic raptor species with three plumage morphs that differ in key life history traits including lifetime reproductive success. In contrast to studies of passerines, CLOCK poly‐glutamine (poly‐Q) was found to be monomorphic in 976 common buzzard nestlings as well as in three other Buteo species. Moreover, none of the candidate genes were significantly associated with fledging dates, although intermediately melanized females were found to lay earlier on average than light or dark morph individuals, and their offspring carried longer ADCYAP1 alleles. In contrast, all three candidate genes explained significant variation in one or more measures of juvenile buzzard dispersal (resighting probability, timing of dispersal and distance dispersed). Our findings contribute towards a broader body of work on the adaptive significance of CLOCK polymorphism, while also building upon previous studies that have documented links between ADCYAP1 variability and the timing of migration.