Geology topics

Katie M. Dugger

Publications and source records attributed to Katie M. Dugger.

At least 37 records · Page 2Linked to original sources

USGS2023

Estimating northern spotted owl (Strix occidentalis caurina) pair detection probabilities based on call-back surveys associated with long-term mark-recapture studies, 1993–2018

The northern spotted owl ( Strix occidentalis caurina ; hereinafter NSO) was listed as “threatened” under the Endangered Species Act in 1990 and population declines have continued since that listing. Given the species’ protected status, any proposed activities on Federal lands that might impact NSO require consultation with U.S. Fish and Wildlife Service and part of that consultation often includes surveys to determine presence and occupancy status of the species in the proposed activity area. The objective of this report is to present study-area specific estimates of the probability of detection for NSO pairs from twelve 2-week seasonal survey periods using data from a recent range-wide meta-analysis. These estimates were a by-product of pair occupancy modeling but might provide insight into potential changes in the effect of the invasive barred owl on NSO detection rates. We used two-species multi-season occupancy models to estimate the probability of detection for NSOs on each of 11 study areas for each 2-week survey period and relative to the range-wide effect of barred owl presence or absence. Detection probabilities within the season generally increased from the earliest surveys in March through mid-season, decreasing again in the late season on five study areas. For three other study areas, detection rates were highest during the earliest survey periods in late March or early April. Estimates of cumulative seasonal detection of NSO (across a maximum of six within-season surveys) were less than 0.90 when barred owls (BO) were present on all but one study area, regardless of when surveys were conducted within a season. However, despite low detection rates, the probability that a territory was occupied when an NSO pair was not detected over six within-season surveys was also very low. When BO are not present on a territory, a six-survey protocol had a high probability of detecting an NSO pair at least once during the season on all study areas, except for the very lowest per-survey estimates. Conducting most surveys earlier in the season, when the probability of detecting pairs is highest (through May on most areas) could improve seasonal detection rates. However, alternative methods of population monitoring—such as the use of passive acoustic recorders—may be needed to continue monitoring NSO for research and management.

California, Oregon, Washington↗

USGS2023

High-resolution recording of foraging behaviour over multiple annual cycles shows decline in old Adélie penguins’ performance

Age-related variation in foraging performance can result from both within-individual change and selection processes. These mechanisms can only be disentangled by using logistically challenging long-term, longitudinal studies. Coupling a long-term demographic data set with high-temporal-resolution tracking of 18 Adélie penguins ( Pygoscelis adeliae , age 4–15 yrs old) over three consecutive annual cycles, we examined how foraging behaviour changed within individuals of different age classes. Evidence indicated within-individual improvement in young and middle-age classes, but a significant decrease in foraging dive frequency within old individuals, associated with a decrease in the dive descent rate. Decreases in foraging performance occurred at a later age (from 12–15 yrs old to 15–18 yrs old) than the onset of senescence predicted for this species (9–11 yrs old). Foraging dive frequency was most affected by the interaction between breeding status and annual life-cycle periods, with frequency being highest during returning migration and breeding season and was highest overall for successful breeders during the chick-rearing period. Females performed more foraging dives per hour than males. This longitudinal, full annual cycle study allowed us to shed light on the changes in foraging performance occurring among individuals of different age classes and highlighted the complex interactions among drivers of individual foraging behaviour.

Proceedings of the Royal Society B↗

USGS2023

Understanding the spatiotemporal distribution of snow refugia in the rain-snow transition zone of north-central Idaho

Knowledge of snow cover distribution and disappearance dates over a wide range of scales is imperative for understanding hydrological dynamics and for habitat management of wildlife species that rely on snow cover. Identification of snow refugia, or places with relatively late snow disappearance dates (SDDs) compared to surrounding areas, is especially important as climate change alters snow cover timing and duration. The purpose of this study was to increase understanding of snow refugia in complex terrain spanning the rain-snow transition zone at fine spatial and temporal scales. To accomplish this objective, we used remote cameras to provide relatively high temporal and spatial resolution measurements on snowpack conditions. We built linear models to relate SDDs at the monitoring sites to topoclimatic and canopy cover metrics. One model to quantify SDDs included elevation, aspect, and an interaction between canopy cover and cold-air pooling potential. High-elevation, north-facing sites in cold-air pools (CAPs) had the latest SDDs, but isolated lower-elevation points also exhibited relatively late potential SDDs. Importantly, canopy cover had a much stronger effect on SDDs in CAPs than in non-CAPs, indicating that best practices in forest management for snow refugia could vary across microtopography. A second model that included in situ hydroclimate observations (December – February (DJF) temperature and March 1 snow depth) indicated that March 1 snow depth had little impact on SDD at the coldest winter temperatures, and that DJF temperatures had a stronger effect on SDD at lower snow depths, implying that the relative importance of snowfall and temperature could vary across hydroclimatic contexts in their impact on snow refugia. This new understanding of factors influencing snow refugia can guide forest management actions to increase snow retention and inform management of snow-dependent wildlife species in complex terrain.

Environmental Research Letters↗

USGS2023

Faster growth and larger size at crèche onset are associated with higher offspring survival in Adélie Penguins

We conducted the first assessment of Adélie Penguin ( Pygoscelis adeliae ) chick survival that accounts for imperfect resighting. We found that when chicks are larger in size when they enter the crèche stage (the period when both parents forage at the same time and chicks are left relatively unprotected), they have a higher probability of survival to fledging. We investigated the relationships between growth, crèche timing, and chick survival during one typical year and one year of reduced food availability. Chicks that hatched earlier in the season entered the crèche stage older, and chicks that both grew faster and crèched older entered the crèche at a larger size. These relationships were stronger in the year of reduced food availability. Thus, parents increased their chicks’ chance of fledging if they provided sufficient food for faster growth rates and/or extended the length of the brood-guarding period. Early nest initiation (i.e., early hatching) provided parents with the opportunity to extend the guard period and increase chick survival. However, to extend the guard stage successfully, they must provide larger meals and maintain higher chick growth rates, even if just one parent at a time is foraging, which previous work has shown is not possible for all individuals. We show that the factors governing tradeoffs in chick-rearing behavior of Adélie Penguin parents may vary in accord with environmental conditions, a result from which we can better understand species’ adaptations to environmental changes.

Ornithology↗

USGS2023

A mixture of Nalbuphine, Azaperone, and Medetomidine for Immobilizing Ringtail (Bassariscus astutus)

We evaluated a combination of nalbuphine HCl (40 mg/mL), azaperone tartrate (10 mg/mL), and medetomidine HCl (10 mg/mL), a combination known as NAM or NalMed-A, in 23 ringtails ( Bassariscus astutus ) during 29 handling events for a radio-collaring study in southern Oregon, US, from August 2020 to March 2022. The combination was delivered to ringtails by hand injection at 0.075 mL NAM per estimated 1 kg body mass. The mean (± standard deviation, SD) dosage calculated post hoc was 3.366 (±0.724) mg/kg nalbuphine, 0.841 (±0.181) mg/kg medetomidine, and 0.841 (±0.181) mg/kg azaperone. All captured ringtails were effectively immobilized with a mean (SD) induction time of 13.24 (±3.57) min. The medetomidine and nalbuphine components were antagonized with a combination of atipamezole and naltrexone HCl with a mean (SD) recovery time of 2.48 (±1.94) min. This combination appeared to be safe and effective for immobilizing ringtails with a low volume dose, smooth antagonism, and rapid recovery. In addition, NAM does not contain any drugs that are US Drug Enforcement scheduled, which makes it useful for immobilization procedures by wildlife professionals in the US.

Journal of Wildlife Diseases↗

USGS2022

Range-wide sources of variation in reproductive rates of northern spotted owls

We conducted a range-wide investigation of the dynamics of site level reproductive rate of northern spotted owls using survey data from 11 study areas across the sub-species geographic range collected during 1993–2018. Our analytical approach accounted for imperfect detection of owl pairs and misclassification of successful reproduction (i.e., at least one young fledged) and contributed further insights into northern spotted owl population ecology and dynamics. Both nondetection and state misclassification were important, especially because factors affecting these sources of error also affected focal ecological parameters. Annual probabilities of site occupancy were greatest at sites with successful reproduction in the previous year and lowest for sites not occupied by a pair in the previous year. Site-specific occupancy transition probabilities declined over time and were negatively affected by barred owl presence. Overall, the site-specific probability of successful reproduction showed substantial year-to-year fluctuations and was similar for occupied sites that did and did not experience successful reproduction the previous year. Site-specific probabilities for successful reproduction were very small for sites that were unoccupied the previous year. Barred owl presence negatively affected the probability of successful reproduction by northern spotted owls in Washington and California, as predicted, but the effect in Oregon was mixed. The proportions of sites occupied by northern spotted owl pairs showed steep, near-monotonic declines over the study period, with all study areas showing the lowest observed levels of occupancy to date. If trends continue it is likely that northern spotted owls will become extirpated throughout large portions of their range in the coming decades.

Ecological Applications↗

USGS2022

Temporal mismatch in space use by a sagebrush obligate species after large-scale wildfire

The increase in size and frequency of wildfires in sagebrush steppe ecosystems has significant impacts on sagebrush obligate species. We modeled seasonal habitat use by female greater sage-grouse ( Centrocercus urophasianus ) in the Trout Creek Mountains of Oregon and Nevada, USA, to identify landscape characteristics that influenced sage-grouse habitat selection and to create predictive surfaces of seasonal use 1 and 7 years postfire. We developed three resource selection function models using GPS location data from 2013 to 2019 for three biologically distinct seasons (breeding, n = 149 individuals: 8 March–12 June; summer, n = 140 individuals: 13 June–20 October; and winter, n = 94 individuals: 21 October–7 March). For all seasons, by the fourth or fifth year postfire, sage-grouse selected for unburned patches more than all other burn severity patches and the use of unburned areas in comparison with burned areas increased through time. During the breeding season, sage-grouse selected for low-sagebrush ( Artemisia arbuscula )-dominated ecosystems and areas with low biomass (normalized difference vegetation index). During summer, sage-grouse selected for areas with higher annual and perennial grasses and forb cover, and areas that had higher biomass. During winter, sage-grouse selected for areas of intact sagebrush on less rugged terrain. For the winter and breeding season, there was a positive linear relationship between annual grasses and forb cover through time. Seven years postfire (2019), the area predicted to have a high probability of use in each seasonal range decreased (breeding: 16.4%; summer: 12.2%; and winter: 4.2%), while the area predicted to have low or low-medium probability of use increased (breeding: 14.5%; summer: 22.5%; and winter: 22.8%) when compared to the first year following the wildfire (2013). Our results demonstrated a 4- to 5-year time lag before female sage-grouse adapted to a disturbed landscape began avoiding burned areas more than intact, unburned habitats. This mismatch in ecological response may imply declines in habitat availability for sage-grouse and may destabilize population vital rates. Spatially explicit models can aid in identifying priority areas for restoration efforts and conservation actions to mitigate the impacts of future disturbances.

Nevada, Oregon↗

USGS2021

Acute and lagged fitness consequences for a sagebrush obligate in a post mega-wildfire landscape

Species responses to disturbance influence their extinction risks. Greater sage- grouse ( Centrocercus urophasianus ) are bioindicators of sagebrush ecosystem health and the loss of sagebrush ( Artemisia spp.) due to wildfire, can cause long- term declines in sage- grouse populations and other sagebrush obligate species. We examined the de - mographic response of a greater sage- grouse population following a mega- wildfire using stochastic age- structured female- based matrix models over 6 years (2013– 2018). Notably, chick survival (range = 0.18– 0.38) and female survival (yearling range: 0.20– 0.68; adult range: 0.27– 0.75) were low compared to values reported for greater sage- grouse in other parts of their distribution. Greater sage- grouse displayed vari - ation in demographic tactics after the fire; however, adult female survival explained most of the variation in λ during each year, which reflected a declining population in 3 of 6 years with more uncertainty observed in 2015 when populations may have been increasing, and 2017 and 2018, when populations may have been declining. The continued annual population decline observed since 2016 suggested there were ad - ditional strong environmental impacts that may have been compounded by the fire effects, prolonging recovery of greater sage- grouse. Our results support others that reported negative effects to greater sage- grouse demographics from broad- scale fire and provide a baseline for understanding how this species responds to loss of sage - brush cover based on their life history strategy.

Nevada, Oregon↗

USGS2021

Feather corticosterone reveals developmental challenges in a long-term study of juvenile northern spotted owls

Corticosterone is a steroid hormone integral to a variety of physiological pathways and is strongly associated with the vertebrate stress response. In avian species, circulating corticosterone is sequestered into developing feathers and is used as an indicator of energy allocation during feather growth and widely applied in conservation physiology. The northern spotted owl Strix occidentalis caurina is a federally threatened old-growth forest obligate of conservation concern endemic to the Pacific Northwest of the United States and Canada. The effects of landscape characteristics and individual variation on early development in spotted owls remain unstudied despite long recognition of this knowledge gap and its potential importance to species conservation. We quantified corticosterone concentrations in 4,720 feathers from 1,056 juvenile spotted owls across seven study areas between 2001 and 2017. We used an information-theoretic approach to examine the environmental and individual factors related to feather corticosterone in juvenile spotted owls as an indicator of challenges during early development. Feather corticosterone was positively related to temperature and precipitation, and negatively related to juvenile mass at banding. We found strong support for an interaction between mass and precipitation, with greater amounts of precipitation being associated with higher levels of feather corticosterone in lighter juveniles. The temperature and precipitation metric with the strongest relationship with feather corticosterone occurred during the fledging period, suggesting that this period presents an energetic challenge for juvenile spotted owls. Greater juvenile mass decreased the effect of precipitation, suggesting that greater mass was important for juveniles to maintain homeostasis during fledgling. Feather corticosterone in juvenile spotted owls provided insights to the challenges faced during early development, adding to our understanding of spotted owl life history and potential for population recovery.

Oregon, Washington↗

USGS2021

Differential landscape use by forest owls two years after a mixed-severity wildfire

Owls are important avian predators in forested systems, but little is known about landscape use by most forest-adapted owl species in environments impacted by mixed-severity wildfire. To better understand species-specific patterns of post-wildfire landscape use within an owl guild, we used passive acoustic monitoring using autonomous recording units. The technology is effective for multi-species surveys, especially if some species are rare, nocturnal, or difficult to detect by traditional means. In 2017, we surveyed the interior and adjacent unburned areas of a 10,700-ha mixed-severity wildfire that burned in 2015 in southwest Oregon. We used occupancy modeling to identify patterns of landscape use by five species of forest owls: barred owls ( Strix varia ), great horned owls ( Bubo virginianus ), western screech-owls ( Megascops kennicottii ), northern pygmy-owls ( Glaucidium gnoma ), and northern saw-whet owls ( Aegolius acadicus ). Our results showed a positive relationship between increasing fire severity and probability of use by western screech-owls and a similar but somewhat weaker relationship for northern pygmy-owls. Barred owls were rarely detected in severely burned areas and their use decreased with increased fire severity. We observed generally low landscape use for great horned owls, which decreased with increased fire severity and at higher elevations. Thus, four out of the five species appeared to use recently burned forests at different levels, with only northern saw-whet owls showing near-complete avoidance of the burned area. These findings increase our understanding of the basic ecology of each species and highlight the varied use of burned areas within this community. These previously undocumented patterns of landscape use in burned landscapes should provide insights to managers and policymakers in the Pacific Northwest as climate shifts, and fires may increase in size, frequency, and severity.

Oregon↗

USGS2021

Fire refugia in old-growth forests: Predicting habitat persistence to support land management in an era of rapid global change

Recent stand-replacing wildfires in late-successional and old-growth (LSOG) forests have increased land manager interest in fire refugia, which could provide vital habitat for threatened and endangered species during a time of rapid change. The overall goal of this project was to model, map, and share information essential for the conservation of LSOG forest ecosystems in the U.S. Pacific Northwest, within a diverse co-production team of state and federal land managers. We developed statistical models of contemporary (2002-2017) fire refugia, non-stand-replacing fire (NSR), and high-severity fire based on topography, fuels, fire weather, fire behavior and climate. Independent models were built for two ecoregions (Figure 1), one encompassing the Douglas-fir/western hemlock forests of the northwestern portion of our study area and the other encompassing dry-mixed conifer forests of the eastern Cascades and Klamath-Siskiyou region. We used these models to produce probability surface maps for fire refugia, NSR, and high-severity fire under low, moderate, and extreme fire weather and fire growth scenarios. These maps and associated products provide timely information about the likely persistence, change, and loss of LSOG forests under current and future climate conditions.

California, Oregon, Washington↗

USGS2021

Conspecific and congeneric interactions shape increasing rates of breeding dispersal of northern spotted owls

Breeding dispersal, the movement from one breeding territory to another, is rare for philopatric species that evolved within relatively stable environments, such as the old-growth coniferous forests of the Pacific Northwest. Although dispersal is not inherently maladaptive, the consequences of increased dispersal on population dynamics in populations whose historical dispersal rates are low could be significant, particularly for a declining species. We examined rates and possible causes of breeding dispersal based on a sample of 4,118 northern spotted owls ( Strix occidentalis caurina ) monitored in seven study areas over 28 yr, 1990–2017, in Oregon and Washington, USA. Using a multistate mark–resight analysis, we investigated the potential impacts of an emergent congeneric competitor (barred owl Strix varia ) and forest alteration (extrinsic factors), and social and individual conditions (intrinsic factors) on 408 successive and 1,372 nonsuccessive dispersal events between years. The annual probability of breeding dispersal increased for individual owls that had also dispersed in the previous year and decreased for owls on territories with historically high levels of reproduction. Intrinsic factors including pair status, prior reproductive success, and experience at a site, were also associated with breeding dispersal movements. The percent of monitored owls dispersing each year increased from ˜7% early in the study to ˜25% at the end of the study, which coincided with a rapid increase in numbers of invasive and competitively dominant barred owls. We suggest that the results presented here can inform spotted owl conservation efforts as we identify factors contributing to changing rates of demographic parameters including site fidelity and breeding dispersal. Our study further shows that increasing rates of breeding dispersal associated with population declines contribute to population instability and vulnerability of northern spotted owls to extinction, and the prognosis is unlikely to change unless active management interventions are undertaken.

California, Oregon, Washington↗

USGS2021

The influence of subcolony-scale nesting habitat on the reproductive success of Adélie penguins

Group-size variation is common in colonially breeding species, including seabirds, whose breeding colonies can vary in size by several orders of magnitude. Seabirds are some of the most threatened marine taxa and understanding the drivers of colony size variation is more important than ever. Reproductive success is an important demographic parameter that can impact colony size, and it varies in association with a number of factors, including nesting habitat quality. Within colonies, seabirds often aggregate into distinct groups or subcolonies that may vary in quality. We used data from two colonies of Adélie penguins 73 km apart on Ross Island, Antarctica, one large and one small to investigate (1) How subcolony habitat characteristics influence reproductive success and (2) How these relationships differ at a small (Cape Royds) and large (Cape Crozier) colony with different terrain characteristics. Subcolonies were characterized using terrain attributes (elevation, slope aspect, slope steepness, wind shelter, flow accumulation), as well group characteristics (area/size, perimeter-to-area ratio, and proximity to nest predators). Reproductive success was higher and less variable at the larger colony while subcolony characteristics explained more of the variance in reproductive success at the small colony. The most important variable influencing subcolony quality at both colonies was perimeter-to-area ratio, likely reflecting the importance of nest predation by south polar skuas along subcolony edges. The small colony contained a higher proportion of edge nests thus higher potential impact from skua nest predation. Stochastic environmental events may facilitate smaller colonies becoming “trapped” by nest predation: a rapid decline in the number of breeding individuals may increase the proportion of edge nests, leading to higher relative nest predation and hindering population recovery. Several terrain covariates were retained in the final models but which variables, the shapes of the relationships, and importance varied between colonies.

Scientific Reports↗

USGS2021

Invader removal triggers competitive release in a threatened avian predator

Invasive species can cause extinctions of native species and widespread biodiversity loss. Invader removal is a common management response, but the use of long-term field experiments to characterize effectiveness of removals in benefitting impacted native species is rare. We used a large-scale removal experiment to investigate the demographic response of a threatened native species, the northern spotted owl, to removal of an invasive competitor species, the barred owl. Removal of barred owls had a strong, positive effect on survival of spotted owls, which arrested long-term population declines of spotted owls. The results demonstrate that the long-term persistence of spotted owls will depend heavily on reducing the negative impacts of barred owls while simultaneously addressing other threats, such as habitat loss.

California, Oregon, Washington↗

USGS2021

Maintenance of nest quality in Adélie penguins Pygoscelis adeliae: An additional benefit to life in the center

In colonial seabirds, differences in the nesting or fledging success have been associated with differences in nest position within the breeding aggregation (subcolony): less successful nests are located on the periphery, with more successful nests closer to the center. For Pygoscelid penguins, central nests tend to be larger, with nest size being an indicator of individual quality because stones must be gathered singly, so more stones reflect more individual effort. Competition for nest materials, including the collection of materials from another’s nest, has also frequently been described in penguins and other colonial seabirds . We used the data collected during the incubation stage from a total of 20 subcolonies at two separate breeding colonies of Adélie penguins ( Pygoscelis adeliae) on Ross Island (Antarctica) to test the influence of nest position on breeding success. We also investigated how competition for nest stones could occur at different intensities depending on size of the subcolony, nest position, and quality within a subcolony. We found that peripheral nests experienced lower breeding success and higher number of individuals attempting to remove stones with higher removal success rates than from nests toward the center. The higher costs associated with maintaining and defending nests that incur higher removal pressure could be an additional factor involved in the lower breeding success of peripheral nests.

Polar Biology↗

USGS2021

Nest microclimates of Greater Sage-Grouse in a post-megafire landscape: does selection equate to success?

Temperature at fine spatial scales is an important driver of nest site selection for many avian species during the breeding season and can influence nest success. Sagebrush ( Artemisia spp. ) communities have areas with high levels of vegetation heterogeneity and high thermal variation; however, fire removes vegetation that provides protection from predators and extreme environmental conditions. To examine the influence of microclimates on Greater Sage-Grouse ( Centrocercus urophasianus ) nest site selection and nest success in a fire affected landscape, we measured black bulb temperature (Tbb) and vegetation attributes (e.g. visual obstruction) at 3 spatial scales (i.e. nest bowl, microsite, and landscape) in unburned and burned areas. Nest bowls exhibited greater buffering of Tbb than both nearby microsites and the broader landscape. Notably, nest bowls were warmer in cold temperatures, and cooler in hot temperatures, than nearby microsites and the broader landscape, regardless of burn stage. Nest survival was higher for nests in unburned areas compared to nests in burned areas (unburned NS = 0.43, 95% CI: 0.33 to 0.54; burned NS = 0.24, 95% CI: 0.10 to 0.46). Amount of bare ground was negatively associated with nest survival, but effects diminished as the amount of bare ground reached very low levels. Shrub height and visual obstruction were positively associated with nest survival during the entire study period whereas, minimum Tbb had a weaker effect. Our findings demonstrate that thermoregulatory selection by Greater Sage-Grouse at nest sites had marginal effects on their nest survival. However, given that increases in vegetation structure (e.g. shrub height) provides thermal refuge and increases nest survival, vegetation remnants or regeneration in a post-fire landscape could be critical to Greater Sage-Grouse nesting ecology.

Oregon, Nevada↗

USGS2021

Range-wide declines of northern spotted owl populations in the Pacific Northwest: A meta-analysis

The northern spotted owl ( Strix occidentalis caurina ) inhabits older coniferous forests in the Pacific Northwest and has been at the center of forest management issues in this region. The immediate threats to this federally listed species include habitat loss and competition with barred owls ( Strix varia ), which invaded from eastern North America. We conducted a prospective meta-analysis to assess population trends and factors affecting those trends in northern spotted owls using 26 years of survey and capture-recapture data from 11 study areas across the owls' geographic range to analyze demographic traits, rates of population change, and occupancy parameters for spotted owl territories. We found that northern spotted owl populations experienced significant declines of 6–9% annually on 6 study areas and 2–5% annually on 5 other study areas. Annual declines translated to ≤35% of the populations remaining on 7 study areas since 1995. Barred owl presence on spotted owl territories was the primary factor negatively affecting apparent survival, recruitment, and ultimately, rates of population change. Analysis of spotted and barred owl detections in an occupancy framework corroborated the capture-recapture analyses with barred owl presence increasing territorial extinction and decreasing territorial colonization of spotted owls. While landscape habitat components reduced the effect of barred owls on these rates of decline, they did not reverse the negative trend. Our analyses indicated that northern spotted owl populations potentially face extirpation if the negative effects of barred owls are not ameliorated while maintaining northern spotted owl habitat across their range.

California, Oregon, Washington↗

USGS2021

Age-related reproductive performance of the Adélie penguin, a long-lived seabird exhibiting similar outcomes regardless of individual life-history strategy

Age-related variation in reproductive performance in long-lived iteroparous vertebrate species is common, with performance being influenced by within-individual processes, such as improvement and senescence, in combination with among-individual processes, such as selective appearance and disappearance. Few studies of age-related reproductive performance have compared the role of these drivers within a metapopulation, subject to varying degrees of resource competition. We accounted for within- and among-individual changes among known-aged Adélie penguins Pygoscelis adeliae during 17 years (1997-2013), at three clustered colonies of disparate size, to understand patterns in age-related reproductive success during early and late adulthood. Age at first reproduction (AFR) was lowest, and number of breeding attempts highest, at the largest colony. Regardless of AFR, success improved with early post-recruitment experience. For both oldest and youngest recruitment groups, peak performance occurred at the end of their reproductive life span indicating a possible cost of reproduction. Intermediate recruitment groups reached peak performance in their mid-reproductive life span and with intermediate breeding experience, before decreasing. Breeding success was lowest for the initial breeding attempt regardless of AFR, but we observed subsequent variation relative to recruitment age. Gaining experience by delaying recruitment positively influenced reproductive performance early in the reproductive life span and was most evident for the youngest breeders. Oldest recruits had the highest initial and peak breeding success. Differences in AFR resulted in trade-offs in reproductive life span or timing of senescence but not in the overall number of breeding attempts. Patterns differed as a function of colony size, and thus competition for resources. Early life improvement in performance at the larger colonies was primarily due to within-individual factors and at the largest colony, AFR. Regardless of colony size late-life performance was positively related to the age at last reproduction, indicating selective disappearance of lower performing individuals. These results highlight that different life-history strategies were equally successful, indicating that individuals can overcome potential trade-offs associated with early- and late-life performance. These results have important implications for understanding the evolution of life-history strategies responsible for driving population change.

Beaufort Island ,Ross Island↗