Variation In First-year Survival Research Articles

Partitioning variance in population growth for models with environmental and demographic stochasticity.

How demographic factors lead to variation or change in growth rates can be investigated using life table response experiments (LTRE) based on structured population models. Traditionally, LTREs focused on decomposing the asymptotic growth rate, but more recently decompositions of annual 'realized' growth rates using 'transient' LTREs have gained in popularity. Transient LTREs have been used particularly to understand how variation in vital rates translate into variation in growth for populations under long-term study. For these, complete population models may be constructed to investigate how temporal variation in environmental drivers affect vital rates. Such investigations have usually come down to estimating covariate coefficients for the effects of environmental variables on vital rates, but formal ways of assessing how they lead to variation in growth rates have been lacking. We extend transient LTREs to further partition the contributions from vital rates into contributions from temporally varying factors that affect them. The decomposition allows one to compare the resultant effect on the growth rate of different environmental factors, as well as density dependence, which may each act via multiple vital rates. We also show how realized growth rates can be decomposed into separate components from environmental and demographic stochasticity. The latter is typically omitted in LTRE analyses. We illustrate these extensions with an integrated population model (IPM) for data from a 26 years study on northern wheatears (Oenanthe oenanthe), a migratory passerine bird breeding in an agricultural landscape. For this population, consisting of around 50-120 breeding pairs per year, we partition variation in realized growth rates into environmental contributions from temperature, rainfall, population density and unexplained random variation via multiple vital rates, and from demographic stochasticity. The case study suggests that variation in first year survival via the unexplained random component, and adult survival via temperature are two main factors behind environmental variation in growth rates. More than half of the variation in growth rates is suggested to come from demographic stochasticity, demonstrating the importance of this factor for populations of moderate size.

Demographic responses to climate change in a threatened Arctic species.

The Arctic is undergoing rapid and accelerating change in response to global warming, altering biodiversity patterns, and ecosystem function across the region. For Arctic endemic species, our understanding of the consequences of such change remains limited. Spectacled eiders (Somateria fischeri), a large Arctic sea duck, use remote regions in the Bering Sea, Arctic Russia, and Alaska throughout the annual cycle making it difficult to conduct comprehensive surveys or demographic studies. Listed as Threatened under the U.S. Endangered Species Act, understanding the species response to climate change is critical for effective conservation policy and planning. Here, we developed an integrated population model to describe spectacled eider population dynamics using capture–mark–recapture, breeding population survey, nest survey, and environmental data collected between 1992 and 2014. Our intent was to estimate abundance, population growth, and demographic rates, and quantify how changes in the environment influenced population dynamics. Abundance of spectacled eiders breeding in western Alaska has increased since listing in 1993 and responded more strongly to annual variation in first‐year survival than adult survival or productivity. We found both adult survival and nest success were highest in years following intermediate sea ice conditions during the wintering period, and both demographic rates declined when sea ice conditions were above or below average. In recent years, sea ice extent has reached new record lows and has remained below average throughout the winter for multiple years in a row. Sea ice persistence is expected to further decline in the Bering Sea. Our results indicate spectacled eiders may be vulnerable to climate change and the increasingly variable sea ice conditions throughout their wintering range with potentially deleterious effects on population dynamics. Importantly, we identified that different demographic rates responded similarly to changes in sea ice conditions, emphasizing the need for integrated analyses to understand population dynamics.

Early-life behaviour predicts first-year survival in a long-distance avian migrant.

Early-life conditions have critical, long-lasting effects on the fate of individuals, yet early-life activity has rarely been linked to subsequent survival of animals in the wild. Using high-resolution GPS and body-acceleration data of 93 juvenile white storks (Ciconia ciconia), we examined the links between behaviour during both pre-fledging and post-fledging (fledging-to-migration) periods and subsequent first-year survival. Juvenile daily activity (based on overall dynamic body acceleration) showed repeatable between-individual variation, the juveniles' pre- and post-fledging activity levels were correlated and both were positively associated with subsequent survival. Daily activity increased gradually throughout the post-fledging period, and the relationship between post-fledging activity and survival was stronger in individuals who increased their daily activity level faster (an interaction effect). We suggest that high activity profiles signified individuals with increased pre-migratory experience, higher individual quality and perhaps more proactive personality, which could underlie their superior survival rates. The duration of individuals' fledging-to-migration periods had a hump-shaped relationship with survival: higher survival was associated with intermediate rather than short or long durations. Short durations reflect lower pre-migratory experience, whereas very long ones were associated with slower increases in daily activity level which possibly reflects slow behavioural development. In accordance with previous studies, heavier nestlings and those that hatched and migrated earlier had increased survival. Using extensive tracking data, our study exposed new links between early-life attributes and survival, suggesting that early activity profiles in migrating birds can explain variation in first-year survival.

Diagnosing the timing of demographic bottlenecks: sub-adult survival in red-billed choughs

1. Determining the precise timing and location of major demographic bottlenecks, such as periods of low survival, is key to identifying ecological causes of variation in population growth rate. Such understanding is key to designing efficient and effective mitigation. 2. In a protected population of red-billed chough Pyrrhocorax pyrrhocorax on Islay, Scotland, variation in population growth rate largely reflects among-year variation in first-year survival. First-year survival was unprecedentedly low during 2007–2010, threatening population viability. 3. We used colour-ring resightings to estimate monthly survival probabilities (Φm) throughout the first year from fledging for eight chough cohorts (totalling 519 individuals) representing the full observed range of variation in first-year survival. We thereby identify the time and location of recent low survival. 4. On average across all cohorts, Φm varied among months, being low during the first month after ringing (May–June, accounting for c. 24% of all first-year mortality) and high during the last 4 months of the first year (January–May, accounting for c. 6% of all first-year mortality). Most mortality (c. 70%) occurred after fledglings dispersed from natal territories. 5. The 2007–2009 cohorts experienced low Φm during July–December. This represents an additional low survival period compared to previous cohorts rather than decreased Φm across all months or further decreases through periods when Φm was low across all cohorts. 6. Synthesis and applications. These data have general relevance in showing that dramatically low annual survival, which needs to spark rapid management action, can reflect different and unanticipated periods of low survival rather than exaggeration of typical variation. With specific regard to conserving Islay’s chough population, our data show that sub-adult survival has recently been low during July–December, probably reflecting conditions on key grassland foraging areas. Managers aiming to increase population viability should increase invertebrate diversity, abundance and availability at these times and locations, thereby increasing foraging options available to choughs.

Evidence of density- and size-dependent mortality in hatchery-reared juvenile white sturgeon (Acipenser transmontanus) in the Kootenai River

We evaluated effects of stocking level and size-at-release on survival rates of hatchery-reared juvenile white sturgeon ( Acipenser transmontanus ) in the Kootenai River using Cormack–Jolly–Seber and related models implemented in Program MARK. A total of 119 768 marked and unmarked hatchery juveniles were released from 1992 to 2006, of which 2938 passive integrated transponder (PIT)-tagged fish were subsequently recaptured. Annual survival rates of marked groups ranged from 0.01 to 0.84 (mean = 0.45) during the first year at large, from 0.48 to 1.0 (mean = 0.84) in the second year, and averaged 1.0 during all subsequent years. First year survival rates declined substantially in recent years, particularly for small fish (<25 cm fork length). Approximately 59% of the variation in first year survival was explained by a negative relationship with estimated juvenile abundance (linear regression, P < 0.01). Length-at-release of individuals explained a substantial proportion of the within-year variation in survival during the first year at large. Our results provide strong evidence of density- and size-dependent mortality in hatchery-reared juvenile white sturgeon in the Kootenai River. Management actions that prioritize the release of fewer, larger-sized fish will likely improve first year survival rates and subsequent recruitment to the spawning-age population.

The demographic impact of extreme events: stochastic weather drives survival and population dynamics in a long‐lived seabird

1. Most scenarios for future climate change predict increased variability and thus increased frequency of extreme weather events. To predict impacts of climate change on wild populations, we need to understand whether this translates into increased variability in demographic parameters, which would lead to reduced population growth rates even without a change in mean parameter values. This requires robust estimates of temporal process variance, for example in survival, and identification of weather covariates linked to interannual variability. 2. The European shag Phalacrocorax aristotelis (L.) shows unusually large variability in population size, and large-scale mortality events have been linked to winter gales. We estimated first-year, second-year and adult survival based on 43 years of ringing and dead recovery data from the Isle of May, Scotland, using recent methods to quantify temporal process variance and identify aspects of winter weather linked to survival. 3. Survival was highly variable for all age groups, and for second-year and adult birds process variance declined strongly when the most extreme year was excluded. Survival in these age groups was low in winters with strong onshore winds and high rainfall. Variation in first-year survival was not related to winter weather, and process variance, although high, was less affected by extreme years. A stochastic population model showed that increasing process variance in survival would lead to reduced population growth rate and increasing probability of extinction. 4. As in other cormorants, shag plumage is only partially waterproof, presumably an adaptation to highly efficient underwater foraging. We speculate that this adaptation may make individuals vulnerable to rough winter weather, leading to boom-and-bust dynamics, where rapid population growth under favourable conditions allows recovery from periodic large-scale weather-related mortality. 5. Given that extreme weather events are predicted to become more frequent, species such as shags that are vulnerable to such events are likely to exhibit stronger reductions in population growth than would be expected from changes in mean climate. Vulnerability to extreme events thus needs to be accounted for when predicting the ecological impacts of climate change.

Density effects at multiple scales in an experimental plant population

1 In sessile organisms, each individual in a population experiences a different local conspecific density, and this density itself will vary depending on the scale at which it is measured (converging on a 'mean field' population density only at coarse scales). 2 The effects of density on any ecological process (e.g. competition, herbivory, mortality) may therefore be scale-specific, and different processes may respond to density at different spatial scales. 3 A field experiment was set up in which focal Silene latifolia plants were subjected to a range of densities at each of four spatial scales (squares with sides of 16 cm, 80 cm, 4 m and 20 m); data were collected on growth, herbivory, flowering and survival. The Akaike information criterion was used to determine which scale displayed the strongest relationship between density and each response variable. 4 The variables showed peak responses to density at different scales and with various relationships (positive, negative and non-monotonic). Some (e.g. herbivory by mammals) showed contrasting responses at different scales. 5 Growth rate showed surprisingly little response to conspecific density, presumably owing to balancing effects of intra-specific and inter-specific competition from the surrounding vegetation. However, 25% of variation in first-year survival was explained by a model including a positive effect of density at the 80-cm scale. 6 Herbivory was mainly due to slugs and showed negative responses to density at the 4-m and 20-m scales, in accordance with hypotheses about resource limitation, mobility and spatial scaling. Mammal herbivory responded with contrasting patterns at the 80-cm and 20-m scales. Congruence between the responses of herbivory and survival suggest that herbivory was an important cause of mortality. 7 Plant population patterns may thus reflect density effects interacting at different characteristic scales. Generalizations about density-dependence in spatially structured populations may be misleading if they fail to recognize that density is scale-dependent.

Diagnosing a decline in return rate of 1-year-old cormorants: mortality, emigration or delayed return?

1. In long-lived birds with delayed recruitment, variation in prebreeding population parameters is difficult to study, although potentially important. Cormorants (Phalacrocorax carbo sinensis) often return to their natal colony when 1 year old, enabling direct study of first-year survival and emigration. Density-dependent regulation mechanisms may be strong at this stage of the life cycle. 2. In the Danish cormorant colony Vorsø, 11 000 chicks have been colour-ringed between 1977 and 1997. Returning birds have been resighted in the colony and dead recoveries have been collected. Since 1990, the proportion of a cohort observed in the colony as 1-year-olds (return rate) has declined from 0·40 to 0·10. Possible explanations include increased mortality, increased emigration and a later age of first return to the colony. Breeding success has also declined strongly as a consequence of low food availability. 3. We used capture-recapture analysis of recovery and resighting data to investigate variation in first-year survival, emigration and resighting probabilities. Survival fluctuated widely (0·42-0·75, mean 0·58); emigration increased in the 1990s from 0·05 to 0·15; resighting probability declined from 0·75 to 0·20 after 1990. 4. First-year survival was particularly low in 1993 and 1996. The causes of the year-to-year variation were not clear. Survival may have been affected by food availability during the post-fledging period. 5. The declining return rate was caused mainly by decreasing resighting probability of 1-year-olds, although increasing emigration also contributed. The biological mechanism was that increasing numbers of cormorants did not return to the natal colony as 1-year-olds. We suggest that this was a consequence of low physiological condition among newly fledged young, caused by low food availability in the 1990s. 6. We conclude that declining food availability has had several consequences for colony dynamics. First-year emigration and the age of first return to the colony have increased and, in the worst years, survival has decreased. If the decline in food availability was due to cormorant predation, this would constitute an example of density-dependent regulation of immature colony attendance.

Survival rates of Redshank Tringa totanus wintering on the Moray Firth

Long-term mark–recapture data were used to estimate the annual survival rates of Redshank wintering on the Moray Firth in Scotland. Survival modelling required the exclusion of all birds caught during the main passage months (August, March and April), and a highly variable annual catching effort limited the precision of annual survival estimates. Survival rates of juvenile Redshank (between the first and second winters of life) varied markedly from year to year and averaged 43% (se 3.6%). Adult survival rates were less variable between years and were age-dependent, with 67% (se 5.0%) surviving and returning between the second and third winters of life, compared to 74% (se 1.4%) for older birds. Year-to-year variation in adult survival was weakly (and negatively) related to the number of snow days in winter. Year-to-year variation in first-year survival was non-linearly related to winter rainfall, with low survival during dry (and cold) winters, higher survival during winters with average rainfall and lower survival during wet winters. Having accounted for these weather relationships there was no evidence that survival was related to the size of the local wintering Redshank population. Organized annual ringing programmes of wintering waders on British estuaries have the potential to monitor long-term changes in survival rates and productivity. Although constant effort sampling may be difficult to achieve for wintering waders, the utility of mark–recapture data collected on estuaries is likely to depend heavily on careful study design.

Bet-hedging theory and eigenelasticity: a comparison of the life histories of loggerhead sea turtles (Caretta caretta) and snapping turtles (Chelydra serpentina)

The life table for snapping turtles (Chelydra serpentina) generated from a 16-year study in Algonquin Park, Ontario, suggests that the population is declining. We use a stage-based matrix model based on this life table to simulate population management options. In addition we analyze the demographic sensitivities of the Algonquin Park population life table and a life table recently published for a population of snapping turtles at the E.S. George Reserve in Michigan. The results are compared with a similar study of loggerhead sea turtles (Caretta caretta). We use these inter- and intra-specific comparisons to test bet-hedging life-history theory. Bet-hedging theory predicts that the long lives and low annual reproductive effort of turtles reduces the effect of low, stochastic juvenile survival on an individual's reproductive success. We test this prediction using proportional sensitivity of the intrinsic rate of increase to variation in life-table parameters (eigenelasticity) to compare the two populations of snapping turtles with each other and with loggerhead sea turtles. Annual adult survival is shown to be the variable most predictive of sensitivity to variation in first-year survival.

Root growth potential, seedling morphology and bud dormancy correlate with survival of loblolly pine seedlings planted in December in Alabama.

First-year survival of December-planted loblolly pine seedlings sampled from 20 nurseries ranged from 36 to 86%. Various characteristics of the seedlings including root growth potential (RGP), shoot/root ratio, root weight and stem length were correlated with survival. A regression model using the number of new roots >/= 0.5 cm and the shoot/root ratio accounted for 80% of the variation in first-year survival. Alone, the number of new roots >/= 0.5 cm accounted for 76% of the variation in survival. Foliar nutrients were not correlated with survival. Seedlings with a high proportion of quiescent buds at planting had a higher RGP and better survival than seedlings with dormant (endodormant) buds. The data suggest that seedlings with dormant buds should not be planted on sites where rapid new root growth is needed for survival.