A review of the population dynamics of mule deer and black‐tailed deer Odocoileus hemionus in North America

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Density‐independent predation affects migrants and residents equally in a declining partially migratory elk population

Migration is expected to benefit individuals through exposure to higher quality forage and reducing predation rates more than non‐migratory conspecifics. Previous studies of partially migratory ungulates (with migrant and resident individuals) have focused on bottom–up factors regulating resident and migrant segments, yet differential predation between strategies could also be a density‐dependent regulatory mechanism. Our study tested for density‐dependence in mortality, as well as mechanisms of ­bottom–up or top–down regulation in the resident and migrant portions of the partially migratory Ya Ha Tinda elk population. We tested for density dependence in adult female and juvenile survival rates, and then discriminated between predator‐ and food‐regulation hypotheses by testing for density‐dependence amongst mortality causes for adult female elk. Notably, the population declined almost 70% from near previously published estimates of carrying capacity over 10 years, providing ideal conditions to test for density dependence. In contrast to predictions, we found only weak support for density dependence in adult survival and juvenile survival. We also found few differences between migrant and resident elk in adult or juvenile survival, though juvenile survival differences were biologically significant. Predation by humans and grizzly bears was density dependent, but similar between migratory strategies. Predation by wolves was the leading known cause of mortality, yet remained constant with declining elk density equally for both migrant and resident elk, indicating wolf predation was density‐independent. Instead of being strongly regulated by food or predation, we found adult female survival was driven by density‐independent predation and climatic factors. The few differences between migratory strategies suggest equivalent fitness payoffs for migrants and residents. This population is being limited by density‐independent predation leading to declines of both migratory strategies. Our results challenge classical predator–prey theory, and call for better integration between predator–prey and migration theory.     

Home range dynamics of the yellow‐footed rock‐wallaby (Petrogale xanthopus celeris) in central‐western Queensland

Analyses of the interspecific differences in macropod home range size suggest that habitat productivity exerts a greater influence on range size than does body mass. This relationship is also apparent within the rock‐wallaby genus. Lim reported that yellow‐footed rock‐wallabies (Petrogale xanthopus xanthopus) inhabiting the semi‐arid Flinders Ranges (South Australia) had a mean home range of 170 ha. While consistent with the hypothesis that species inhabiting less productive habitats will require larger ranges to fulfil their energetic requirements, the ranges reported by Lim were considerably larger than those observed for heavier sympatric macropods. The aim of the current study was to document the home range dynamics of P. x. celeris in central‐western Queensland and undertake a comparison with those reported for their southern counterparts. Wallaby movements were monitored at Idalia National Park, between winter 1992 and winter 1994. Male foraging ranges (95% fixed kernel; 15.4 ha, SD = ±7.8 ha) were found to be significantly larger than those of female wallabies (11.3 ha, SD = ±4.9 ha). Because of varying distances to the wallabies' favoured foraging ground (i.e. an adjacent herb field), the direction in which the wallabies moved to forage also significantly affected range size. Mean home range size was estimated to be 23.5 ha (SD = ±15.2 ha; 95% fixed kernel) and 67.5 ha (SD = ±22.4 ha; 100% minimum convex polygon). The discrepancy between these two estimates resulted from the exclusion of locations, from the 95% kernel estimates, when the wallabies moved to a water source 1.5 km distant from the colony site. The observed foraging and home ranges approximated those that could be expected for a macropod inhabiting the semi‐arid zone (i.e. 2.4 times larger‐than‐predicted from body mass alone). Possible reasons for the disparity between the current study and that of Lim are examined.     

Rain,rats and pythons: Climate‐driven population dynamics of predators and prey in tropical Australia

Abstract: All natural populations fluctuate in abundance and age structure through time; understanding why they do so is a critical step towards their effective management and conservation. However, the long‐term data sets needed for such an understanding are rarely available, especially for tropical organisms. A 17‐year capture‐mark–recapture study yielded detailed information on the demography of water pythons (Liasis fuscus) and their main prey, the dusky rat (Rattus colletti), on the Adelaide River flood plain in tropical Australia. The link between annual rainfall patterns and rat demography was highly non‐linear. Rat numbers were low during years with low and high rainfall at the end of the wet season (April). Numbers of both predators and prey fluctuated considerably among years. Annual fluctuations in rat numbers generated a corresponding variation in rates of female python reproduction, python body condition and survival. Although variation in recruitment, survival and prey abundance all had a significant impact on annual fluctuations in python numbers, our analyses suggest that recruitment constituted the main determinant in driving the population dynamics of these large tropical predators. In combination with our other studies on this system, the data show that population dynamics of the water python population is ultimately driven by annual variation in rainfall, mediated via shifts in prey availability. The water pythons and the dusky rats of the Adelaide River flood plain thus demonstrate an unusually clear and direct link between an abiotic factor (rainfall) and predator–prey population dynamics.     

Density‐dependent survival and recruitment in a long‐distance Palaearctic migrant,the Sand Martin Riparia riparia

Long‐term studies can provide powerful insights into the relative importance of different demographic and environmental factors determining avian population dynamics. Here we use 23 years of capture–mark–recapture data (1981–2003) to estimate recruitment and survival rates for a Sand Martin Riparia riparia population in Cheshire, NW England. Inter‐annual variation in recruitment and adult survival was positively related to rainfall in the sub‐Saharan wintering grounds, but unrelated to weather conditions on the breeding grounds. After allowing for the effects of African rainfall, both demographic rates were negatively density‐dependent: adult survival was related to the size of the western European Sand Martin population (probably reflecting competition for resources in the shared wintering grounds) while recruitment was related to the size of the local study population in Cheshire (potentially reflecting competition for nesting sites or food). Local population size was more sensitive to variation in adult survival than to variation in recruitment, and an increase in population size after 1995 was driven mainly by the impact of more favourable conditions in the African wintering grounds on survival rates of adults. Overwinter survival in this long‐distance Palaearctic migrant is determined partly by the amount of suitable wetland foraging habitat in the sub‐Saharan wintering grounds (which is limited by the extent of summer rainfall) and partly by the number of birds exploiting that habitat.     

Population demography of frillneck lizards (Chlamydosaurus kingii,Gray 1825) in the wet‐dry tropics of Australia

We explore the effects of biotic and abiotic factors on the population demography of frillneck lizards (Chlamydosaurus kingii) in the Australian wet‐dry tropics. Annual growth rates of males were significantly higher across all body sizes compared to females, resulting in a significant larger maximum body size in males. Both male and female lizards were highly philopatric and 81% of the among‐year recapture distances were less than 200 m. Juvenile and adult frillnecks were subjected to low but highly variable annual survival rates. Both proportion of juveniles and relative proportion of reproductive females showed extensive among‐year variations. No relationship was, however, observed between proportion of gravid females and that of juveniles captured during the subsequent year. High rainfall in January was negatively correlated with recruitment most likely caused by increased egg/embryo mortality due to flooding of nest sites. We therefore suggest that the lack of association between female reproduction and juvenile recruitment was due to the effects of stochastic variation in January rainfall. Lizard numbers increased during the first five years of the study followed by a decline during the subsequent four years. Our analyses show that annual variation in survival constituted the main determinant in driving the annual change in frillneck numbers. Surprisingly, no relationship was observed between fillneck population dynamics and annual variation in juvenile recruitment. We suggest that the 7‐years over which these analyses were conducted were insufficient to detect any significant effects of recruitment on lizard numbers, demonstrating the need for long‐term studies to accurately document vertebrate population demographic processes in areas experiencing stochastic variations rainfall such as the Australian wet‐dry tropics.     

Life history and population dynamics of a tree species in tropical semi‐arid climate: A case study with Cordia oncocalyx

In semi‐arid climates, plant population dynamics are strongly influenced by the amount and temporal distribution of rainfall. We monitored a population of the tree species Cordia oncocalyx (Boraginaceae) for 24 months in the dry thorny woodland of semi‐arid northeastern Brazil, to investigate which life‐history traits allow this tree to be locally dominant. We used horizontal life tables and a Lefkovitch matrix and tested for relationships among demographic parameters of seedling, infant, juvenile, immature, virginile and reproductive ontogenetic stages with rainfall and canopy openness. Germination and recruitment occurred in the rainy months, and dry‐season mortality occurred only in seedlings (76% and 100%, first and second years, respectively) and infants (3% and 6%). Juveniles showed greater height growth under more open canopies (Spearman correlation coefficient = 0.24), suggesting that light availability influences growth. The population growth rate was λ = 1.0336, and the highest sensitivity occurred in the infant‐juvenile transition. Our results show light as a restrictive growth factor for plants in the juvenile stage and confirm the strong influence of rainfall on the dynamics of trees in a seasonally dry environment. The formation of a persistent seed bank with germination concentrated at the rainfall onset but spreading over the rainy season are strategies that hedge bets before establishment. The formation of a bank of infants, which can resume growth as soon as there is water, hedges bets after establishment. We attribute the positive population growth rate of Cordia oncocalyx to survival strategies allowing bet‐hedging both before and after establishment.     

The role of El Niño–Southern Oscillation in the dynamics of a savanna large herbivore population

Our understanding of large‐scale climatic phenomena and dynamics of large herbivore populations comes principally from research in northern regions with temperate, seasonal climate and animal communities with relatively low species diversity. To assess the generality of that perspective, we investigated effects of El Niño–Southern Oscillation (ENSO) on population dynamics of African buffalo Syncerus caffer inhabiting a semi‐arid savanna with variable rainfall. We used linear and nonlinear‐threshold models to investigate relationships between population parameters and explanatory variables affecting forage conditions (seasonal rainfall, Southern Oscillation Index [SOI]). El Niño‐related droughts in 1982–1983 and 1991–1992 were associated with strongly negative population change, a pattern expected to coincide with a decrease in normally high and constant adult survival. Consistent with that nonlinear pattern, we detected threshold relationships between wet‐season rainfall and population change. Juvenile recruitment was described best by linear relationships with dry‐season. Because ENSO operates primarily through wet‐season rainfall, whereas population dynamics were also related to dry‐season rainfall, SOI did not have the predictive ability of individual weather components.     

The founding of a southern elephant seal colony

The only large mainland colony of southern elephant seals (Mirounga leonina) is on Península Valdés, at 42°S, in Argentine Patagonia. Censuses of pups have been carried out regularly there since 1970, and the population grew five‐fold by 2010. Here we use Bayesian modeling tools to make rigorous estimates of the rate of population growth, r, and to estimate survival and recruitment parameters that could account for the growth, incorporating observation error across different census methods. In the 1970s, r= 8%/yr, but has slowed to <1%/yr over the past decade. Using explicit demographic models, we established that the high growth of the 1970s was consistent with adult and juvenile survival at the upper end of published values (0.87/yr adult female survival; 0.40 juvenile survivorship to age four); the decline in the rate of population growth from 1970 to 2010 can be described by density‐dependent reductions in adult and juvenile survival that fall well within published variation. Extrapolating empirical models of population growth rate backwards illustrates that the population could have been an established colony, with 100 pups born per year, between 1915 and 1945, consistent with qualitative observations prior to 1950. We conclude that the Valdés colony was founded by a few immigrants early in the 20th century and has been growing mostly by internal recruitment, with unknown density‐dependent processes causing a reduction in growth and stabilization at 15,000–16,000 pups born.     

Large‐scale predator control increases population viability of a rare New Zealand riverine duck

The introduction of mammalian predators to oceanic islands has led to dramatic declines in the abundance of many native species. Conservation management of these species often relies on low‐cost predator control techniques that can be implemented over large scales. Assessing the effectiveness of such management techniques is difficult, but using population viability analyses (PVA), which identify the population growth rate (λ) and extinction risk of threatened species, may offer a solution. PVA provide the opportunity to compare the relative effectiveness of various management options and can identify knowledge gaps to prioritize research efforts. We used PVA to assess the population viability of whio (Hymenolaimus malacorhynchos), a rare riverine duck endemic to New Zealand. Current populations are threatened by introduced mammalian predators and are rapidly declining in both distribution and abundance. Whio conservation management is dominated by large‐scale, low‐intensity predator control, targeting introduced stoats (Mustela erminea). There is evidence that such control increases whio productivity but it is unknown if this increase is sufficient for long‐term population persistence. We undertook a stochastic PVA to assess the viability of whio populations under different management scenarios using data obtained from a 6‐year study of whio demographic responses to predator control. Populations with no predator control and low productivity will rapidly decline to extinction. Increasing productivity through predator control increased population viability but populations still showed a declining trajectory. A perturbation analysis showed that the growth rate of whio populations was largely driven by adult survival. Therefore, future research should target obtaining more robust estimates of adult survival, particularly how it is affected by predator control. Overall, our analysis indicated that large‐scale predator control increases the short‐term viability of whio populations but is insufficient for long‐term population persistence.     

Avian cholera,post‐hatching survival and selection on hatch characteristics in a long‐lived bird,the common eider Somateria mollisima

Infectious diseases can have dramatic impacts on animal population dynamics, but how they influence vital rates remains understudied. We took advantage of the appearance of an avian cholera epizootic in an arctic colony of common eiders Somateria mollissima to study variation in juvenile survival and selection on hatch characteristics in relation to this highly infectious disease. Avian cholera is one of the most important infectious diseases affecting wild birds and is thought to primarily affect adult survival. Here, we show that avian cholera was associated with a 90% decline in duckling survival, leading to almost zero recruitment. Before the cholera outbreak, there was significant stabilizing selection on hatching date and significant positive directional selection on hatching mass. During cholera outbreaks, selection on hatch characteristics was no longer significant. These results were based on a low sample of surviving ducklings in cholera years, but suggested that date and mass at hatching did no longer affect duckling survival in the presence of cholera. These effects of avian cholera on post‐hatching survival were likely not only the consequence of the disease per se, but also a consequence of an increase in predation rates that followed the emergence of avian cholera. Our results emphasize the dramatic direct and indirect impacts that infectious disease can have on vital rates, and thus population dynamics.     

A wide window of migration phenology captures inter‐annual variability of favourable conditions: Results of a whole‐lake experiment with juvenile Pacific salmon

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Radio‐transmitters have no impact on survival of pre‐fledged American Woodcocks

American Woodcocks (Scolopax minor) are a high priority species of conservation need across most of their breeding range due to long‐term population declines. Survival of juveniles may be key to understanding these population declines, but there have been few direct estimates of juvenile woodcock survival rates, and no recent assessment of the possible effect of radio‐tagging on juvenile survival. In 2011 and 2012, we radio‐tagged 73 juvenile American Woodcocks in west‐central Minnesota and compared survival rates of radio‐tagged (N = 58) and non‐radio‐tagged (N = 82) juveniles during the period from hatching to fledging. We compared survival rates of juveniles with known fates and used logistic‐exposure models to assess the potential impact of radio‐transmitters on survival. We evaluated variables related to juvenile survival including age, hatch date, maximum and minimum temperature, precipitation, and year to assess the possible effects of radio‐transmitters. The best‐supported model of survival rate of juvenile American Woodcocks included the interaction of age and year and a negative effect of precipitation (β = ?0.76, 85% CI: ?1.08 to ?0.43), but did not include a negative effect of transmitters. Our results suggest that radio‐transmitters did not impact survival of juvenile American Woodcocks and that transmitters are a reliable tool for studying survival of juvenile American Woodcocks, and perhaps other precocial shorebirds.     

Demographic dynamics of the afro-tropical pig-nosed frog, <Emphasis Type="Italic">Hemisus marmoratus</Emphasis>: effects of climate and predation on survival and recruitment

We studied the population ecology of the West African pig-nosed frog, Hemisus marmoratus, to understand the relative contributions of adult survival and recruitment to population growth rate in savannah frogs using mark-recapture modelling. We marked a total of 821 adult frogs over 6 years and recaptured 74 at least once between years. Between-year adult survival was sex-specific and varied between 0.06 and 0.53 for males and 0.07–0.41 for females. Adult survival was significantly associated with annual rainfall and is cause for concern if rainfall declines further in the study region as predicted by changes in the global climate. There was a significant interaction between rainfall and sex with dry weather having a stronger negative effect on males than females. Pig-nosed frogs experienced boom and bust years with a single decline more dramatic than increases. Recruitment (in situ and immigration; 0.67–0.88) was substantially more important than adult survival (0.12–0.33) in determining realised population growth. In situ recruitment was highly variable between years with 1–36% of eggs and tadpoles released by females into the pond surviving to metamorphosis. Years of low tadpole survival were associated with high numbers of predatory tortoises. Thus, like other pond-breeding anurans, pig-nosed frogs showed highly variable juvenile recruitment, low adult survival and density-independent effects on population growth by predators and weather.     

The influence of pre‐settlement and early post‐settlement processes on the adult distribution and relative dominance of two invasive mussel species

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A field trial to test effects of watering,seed addition and disturbance on perennial species recruitment in Belah woodland

Failure of perennial species to regenerate is a significant threat to semi‐arid woodlands across south‐eastern Australia. High grazing pressure eliminates the recruitment of many perennial species in semi‐arid woodlands, but little is known about requirements for regeneration under low grazing pressure. We tested the effects of addition of water (irrigation to match the largest rainfall events of the last century), seed, soil disturbance and fire within a grazing exclosure in Belah (Casuarina pauper) woodland in the Murray‐Sunset National Park, Victoria. Recruitment was observed in 13 perennial species and was dominated by chenopods. Addition of water, seed and soil disturbance increased abundance of juvenile perennial species above the low‐level background recruitment that occurred in the prevailing drought conditions. This supports the view that continuous recruitment occurs for many semi‐arid perennials. Low seed availability and an inability to maintain soil moisture conditions matching that of regeneration events are likely factors in the lack of recruitment for tree species and limited response of shrubs in this experiment.     

Can multitrophic interactions and ocean warming influence large‐scale kelp recovery?

Ongoing changes along the northeastern Atlantic coastline provide an opportunity to explore the influence of climate change and multitrophic interactions on the recovery of kelp. Here, vast areas of sea urchin‐dominated barren grounds have shifted back to kelp forests, in parallel with changes in sea temperature and predator abundances. We have compiled data from studies covering more than 1,500‐km coastline in northern Norway. The dataset has been used to identify regional patterns in kelp recovery and sea urchin recruitment, and to relate these to abiotic and biotic factors, including structurally complex substrates functioning as refuge for sea urchins. The study area covers a latitudinal gradient of temperature and different levels of predator pressure from the edible crab (Cancer pagurus) and the red king crab (Paralithodes camtschaticus). The population development of these two sea urchin predators and a possible predator on crabs, the coastal cod (Gadus morhua), were analyzed. In the southernmost and warmest region, kelp forests recovery and sea urchin recruitment are mainly low, although sea urchins might also be locally abundant. Further north, sea urchin barrens still dominate, and juvenile sea urchin densities are high. In the northernmost and cold region, kelp forests are recovering, despite high recruitment and densities of sea urchins. Here, sea urchins were found only in refuge habitats, whereas kelp recovery occurred mainly on open bedrock. The ocean warming, the increase in the abundance of edible crab in the south, and the increase in invasive red king crab in the north may explain the observed changes in kelp recovery and sea urchin distribution. The expansion of both crab species coincided with a population decline in the top‐predator coastal cod. The role of key species (sea urchins, kelp, cod, and crabs) and processes involved in structuring the community are hypothesized in a conceptual model, and the knowledge behind the suggested links and interactions is explored.     

Long‐term changes in semi‐arid vegetation: Invasion of an exotic perennial grass has larger effects than rainfall variability

Abstract. Questions: This paper examines the long‐term change in the herbaceous layer of semi‐arid vegetation since grazing ceased. We asked whether (1) there were differences in the temporal trends of abundance among growth forms of plants; (2) season of rainfall affected the growth form response; (3) the presence of an invasive species influenced the abundance and species richness of native plants relative to non‐invaded plots, and (4) abundance of native plants and/or species richness was related to the time it took for an invasive species to invade a plot. Location: Alice Springs, Central Australia. Methods: Long‐term changes in the semi‐arid vegetation of Central Australia were measured over 28 years (1976–2004) to partition the effects of rainfall and an invasive perennial grass. The relative abundance (biomass) of all species was assessed 25 times in each of 24 plots (8 m × 1 m) across two sites that traversed floodplains and adjacent foot slopes. Photo‐points, starting in 1972, were also used to provide a broader overview of a landscape that had been intensively grazed by cattle and rabbits prior to the 1970s. Species’abundance data were amalgamated into growth forms to examine their relationship with environmental variation in space and time. Environmental variables included season and amount of rainfall, fire history, soil variability and the colonization of the plots by the exotic perennial grass Cenchrus ciliaris (Buffel grass). Results: Constrained ordination showed that season of rainfall and landscape variables relating to soil depth strongly influenced vegetation composition when Cenchrus was used as a covariate. When Cenchrus was included in constrained ordination, it was strongly related to the decline of all native growth forms over time. Univariate comparisons of non‐invaded vs impacted plots over time revealed unequivocal evidence that Cenchrus had caused the decline of all native growth form groups and species richness. They also revealed a contrasting response of native plants to season of rainfall, with a strong response of native grasses to summer rainfall and forbs to winter rainfall. In the presence of Cenchrus these responses were strongly attenuated. Discussion: Pronounced changes in the composition of vegetation were interpreted as a response to removal of grazing pressure, fluctuations in rainfall and, most importantly, invasion of an exotic grass. Declines in herbaceous species abundance and richness in the presence of Cenchrus appear to be directly related to competition for resources. Indirect effects may also be causing the declines of some woody species from changed fire regimes as a result of increased fuel loads. We predict that Cenchrus will begin to alter landscape level processes as a result of the direct and indirect effects of Cenchrus on the demography of native plants when there is a switch from resource limited (rainfall) establishment of native plants to seed limited recruitment.     

Variable rainfall has a greater effect than fire on the demography of the dominant tree in a semi‐arid Eucalyptus savanna

Rainfall, fire and competition are emphasized as determinants of the density and basal area of woody vegetation in savanna. The semi‐arid savannas of Australia have substantial multi‐year rainfall deficits and insufficient grass fuel to carry annual fire in contrast to the mesic savannas in more northern regions. This study investigates the influence of rainfall deficit and excess, fire and woody competition on the population dynamics of a dominant tree in a semi‐arid savanna. All individuals of Eucalyptus melanophloia were mapped and monitored in three, 1‐ha plots over an 8.5 year period encompassing wet and dry periods. The plots were unburnt, burnt once and burnt twice. A competition index incorporating the size and distance of neighbours to target individuals was determined. Supplementary studies examined seedling recruitment and the transition of juvenile trees into the sapling layer. Mortality of burnt seedlings was related to lignotuber area but the majority of seedlings are fire resistant within 12 months of germination. Most of the juveniles (≤1 cm dbh) of E. melanophloia either died in the dry period or persisted as juveniles throughout 8.5 years of monitoring. Mortality of juveniles was positively related to woody competition and was higher in the dry period than the wet period. The transition of juveniles to a larger size class occurred at extremely low rates, and a subsidiary study along a clearing boundary suggests release from woody competition allows transition into the sapling layer. From three fires the highest proportion of saplings (1–10 cm dbh) reduced to juveniles was only 5.6% suggesting rates of ‘top‐kill’ of E. melanophloia as a result of fire are relatively low. Girth growth was enhanced in wet years, particularly for larger trees (>10 cm dbh), but all trees regardless of size or woody competition levels are vulnerable to drought‐induced mortality. Overall the results suggest that variations in rainfall, especially drought‐induced mortality, have a much stronger influence on the tree demographics of E. melanophloia in a semi‐arid savanna of north‐eastern Australia than fire.     

Spatial patterns in age- and colony-specific survival in a long-lived seabird across 14 contrasting colonies

Demographic rates such as recruitment and survival probability can vary considerably among populations of the same species due to variation in underlying environmental processes. If environmental processes are spatially correlated, nearby populations are expected to have more similar demographic rates than those further apart. Breeding populations and foraging ranges are spatially segregated in colonial seabirds, making them ideal for studying spatial patterns in demographic rates and their effects on local population dynamics. Here we explored variation in age-dependent survival probabilities across 14 colonies of Herring Gulls Larus argentatus breeding along the Dutch North Sea coast. We used long-term mark–recapture data of marked fledglings to estimate survival, and estimated spatial autocorrelation of survival probabilities. We assessed whether survival until recruitment age or until 10 years old (close to their expected lifespan) explained variation in population trajectories of each colony. Juvenile and adult survival showed a strong, but different, north-to-south gradient in survival probability, with lower juvenile but higher adult survival in northern colonies than southern colonies, whereas the spatial pattern of immature survival was less distinct. Neither recruitment nor the proportion of 10-year-old adults alive predicted whether a colony collapsed, declined, remained stable or increased. The distinct spatial pattern in survival suggests variation in regional food availability, which do not seem to drive local population dynamics. The absence of a link between survival and colony trajectories implies that connectivity between populations plays an important role affecting population dynamics.