Factors driving territory size and breeding success in a threatened migratory songbird, the Canada Warbler

Successful conservation of migratory birds demands we understand how habitat factors on the breeding grounds influences breeding success. Multiple factors are known to directly influence breeding success in territorial songbirds. For example, greater food availability and fewer predators can have direct effects on breeding success. However, many of these same habitat factors can also result in higher conspecific density that may ultimately reduce breeding success through density dependence. In this case, there is a negative indirect effect of habitat on breeding success through its effects on conspecific density and territory size. Therefore, a key uncertainty facing land managers is whether important habitat attributes directly influence breeding success or indirectly influence breeding success through territory size. We used radio-telemetry, point-counts, vegetation sampling, predator observations, and insect sampling over two years to provide data on habitat selection of a steeply declining songbird species, the Canada Warbler (Cardellina canadensis). These data were then applied in a hierarchical path modeling framework and an AIC model selection approach to determine the habitat attributes that best predict breeding success. Canada Warblers had smaller territories in areas with high shrub cover, in the presence of red squirrels (Tamiasciurus hudsonicus), at shoreline sites relative to forest-interior sites and as conspecific density increased. Breeding success was lower for birds with smaller territories, which suggests competition for limited food resources, but there was no direct evidence that food availability influenced territory size or breeding success. The negative relationship between shrub cover and territory size in our study may arise because these specific habitat conditions are spatially heterogeneous, whereby individuals pack into patches of preferred breeding habitat scattered throughout the landscape, resulting in reduced territory size and an associated reduction in resource availability per territory. Our results therefore highlight the importance of considering direct and indirect effects for Canada warblers; efforts to increase the amount of breeding habitat may ultimately result in lower breeding success if habitat availability is limited and negative density dependent effects occur.

Breeding success and traits of equal and unequal pairs of little penguins (Eudyptula minor) on Phillip Island during 2001-2008

According to life-history theory, individuals optimize their decisions in order to maximize their fitness. This raises a conflict between parents, which need to cooperate to ensure the propagation of their genes but at the same time need to minimize the associated costs. Trading-off between benefits and costs of a reproduction is one of the major forces driving demographic trends and has shaped several different parental care strategies. Using little penguins (Eudyptula minor) as a model, we investigated whether individuals of a pair provide equal parental effort when raising offspring and whether their behavior was consistent over 8 years of contrasting resource availability. Using an automated identification system, we found that 72% of little penguin pairs exhibited unforced (i.e., that did not result from desertion of 1 parent) unequal partnership through the postguard stage. This proportion was lower in favorable years. Although being an equal pair appeared to be a better strategy, it was nonetheless the least often observed. Individuals that contributed less than their partner were not less experienced (measured by age), and gender did not explain differences between partners. Furthermore, birds that contributed little or that contributed a lot tended to be consistent in their level of contribution across years. We suggest that unequal effort during breeding may reflect differences in individual quality, and we encourage future studies on parental care to consider this consistent low and high contributor behavior when investigating differences in pair investment into its offspring. Key words: attendance patterns, individual quality, meal size, parental care, reproductive costs, seabirds.

Increased reproductive success in older female red kangaroos and the impact of harvesting

Wildlife harvesting has a long history in Australia, including obvious examples of overexploitation. Not surprisingly, there is scepticism that commercial harvesting can be undertaken sustainably. Kangaroo harvesting has been challenged regularly at Administrative Appeals Tribunals and elsewhere over the past three decades. Initially, the concern from conservation groups was sustainability of the harvest. This has been addressed through regular, direct monitoring that now spans > 30 years and a conservative harvest regime with a low risk of overharvest in the face of uncertainty. Opposition to the harvest now continues from animal rights groups whose concerns have shifted from overall harvest sustainability to side effects such as animal welfare, and changes to community structure, genetic composition and population age structure. Many of these concerns are speculative and difficult to address, requiring expensive data. One concern is that older females are the more successful breeders and teach their daughters optimal habitat and diet selection. The lack of older animals in a harvested population may reduce the fitness of the remaining individuals; implying population viability would also be compromised. This argument can be countered by the persistence of populations under harvesting without any obvious impairment to reproduction. Nevertheless, an interesting question is how age influences reproductive output. In this study, data collected from a number of red kangaroo populations across eastern Australia indicate that the breeding success of older females is up to 7-20% higher than that of younger females. This effect is smaller than that of body condition and the environment, which can increase breeding success by up to 30% and 60% respectively. Average age of mature females in a population may be reduced from 9 to 6 years old, resulting in a potential reduction in breeding success of 3-4%. This appears to be offset in harvested populations by improved condition of females from a reduction in kangaroo density. There is an important recommendation for management. The best insurance policy against overharvest and unwanted side effects is not research, which could be never-ending. Rather, it is a harvest strategy that includes safeguards against uncertainty such as harvest reserves, conservative quotas and regular monitoring. Research is still important in fine tuning that strategy and is most usefully incorporated as adaptive management where it can address the key questions on how populations respond to harvesting.

Stable breeding despite variable feeding in two sympatric auk species

Capsule Despite substantial inter-annual and inter-specific variance in the composition of chick diet, the breeding success of Guillemots (Common Murres) Uria aalge and Razorbills Alca torda remained constant from 2008 to 2010.
Aims To examine inter-specific and inter-annual differences in breeding success, chick provisioning behaviour and predation between two sympatric auk species.
Methods Focal observations of breeding auks at Rathlin Island, Northern Ireland, during 2008, 2009 and 2010 recorded reproductive success, reasons for breeding failure, prey composition and quality and chick provisioning rates.
Results Breeding success of both species was stable over the three years, despite significant variance in the composition and quality of the diet provided to chicks. Razorbills experienced greater rates of failure than Guillemots owing to chick loss and had lower overall breeding success.
Conclusion Guillemot and Razorbill breeding success was independent of the composition and quality of prey items delivered to chicks. Inter-specific differences in reproductive success may have been attributed to greater rates of predation at Razorbill rather than Guillemot nests.

Foraging trip time-activity budgets and reproductive success in the black-legged kittiwake

Seabirds are central place foragers during the breeding season and, as marine food resources are often patchily distributed, flexibility in foraging behaviour may be important in maintaining prey delivery rates to chicks. We developed a methodological approach using a combination of GPS data loggers and temperature-depth recorders that allowed us to describe the behaviour of surface-feeding seabirds. Specifically, we tested whether differences in foraging behaviour of black-legged kittiwakes Rissa tridactyla could be linked with reproductive success by comparing 2 consecutive years at 2 sites. At Rathlin Island (Northern Ireland) during 2010, foraging differed markedly from that during 2009 and from that at Lambay Island (Republic of Ireland) during both years. Birds exhibited foraging trips of greater duration, travelled a greater total distance, spent more time in transit and spent longer recuperating on the surface of the water. This notable shift was associated with a decline in breeding success, with greater loss of eggs to predation and lower prey delivery rates, resulting in the starvation of 15 % of chicks. We suggest that food resources were reduced or geographically less accessible during 2010, with suitable foraging areas located further from the colony. Birds did not invest greater amounts of time attempting to catch prey. Thus, our results indicate that kittiwakes at Rathlin modulated their foraging behaviour not by increasing foraging effort through feeding more intensively within prey patches but by extending their range to increase the probability of encountering more profitable prey patches. © Inter-Research 2012.

Breeding parameters of great cormorants (Phalacrocorax carbo carbo) at mixed species colonies on Prince Edward Island, Canada

Breeding parameters of Great Cormorants (PkaZac/iOCOfiCLX CCUibo dCUtbo) and Double-crested Cormorants (P. CLU/uXuA CMJhLtllb) were examined at two mixed species colonies at Cape Tryon and Durell Point, Prince Edward Island from 1976 to 1978. Differential access to nests at the two colony sites resulted in more complete demographic data for P. CCUibo than for P. CLUJiituA. In 1911j P. CCtfibo was present at both colonies by 21 March, whereas P. auAAJtuA did not return until 1 April and 16 April at Cape Tryon and Durell Point, respectively. Differences in the arrival chronology by individuals of each species and differences in the time of nest site occupation according to age, are suggested as factors influencing the nest site distribution of P. CXUtbo and P. aiVtituA at Cape Tryon. Forty-eight P. dOJtbo chicks banded at the Durell Point colony between 19 74 and 19 76 returned there to nest as two- to four-year olds in 19 77 and 19 78. Unmarked individuals with clutch-starts in April were likely greater than four years old as all marked two to four-year olds (with one possible exception) in 19 77 and 1978 had clutch-starts in May and June. Seasonal variation in the breeding success of P. dOJibo individuals was examined at Durell Point in 1977. Mean clutch-size, hatching success and fledging success exhibited a seasonal decline. Four- and 5-egg clutches represented the majority (75%) of all P. CCUibo clutches at Durell Point in 1977 and had the highest reproductive success (0.48 and 0.43 chicks fledged per egg laid respectively). Smaller clutches produced small broods with significantly higher chick mortality while larger clutches suffered high egg loss prior to clutch completion.

Breeding Biology of Grassland Birds in Western New York: Conservation and Management Implications

Declining grassland breeding bird populations have led to increased efforts to assess habitat quality, typically by estimating density or relative abundance. Because some grassland habitats may function as ecological traps, a more appropriate metric for determining quality may be breeding success. Between 1994 and 2003 we gathered data on the nest fates of Eastern Meadowlarks (Sturnella magna), Bobolinks (Dolichonyx oryzivorous), and Savannah Sparrows (Passerculus sandwichensis) in a series of fallow fields and pastures/hayfields in western New York State. We calculated daily survival probabilities using the Mayfield method, and used the logistic-exposure method to model effects of predictor variables on nest success. Nest survival probabilities were 0.464 for Eastern Meadowlarks (n = 26), 0.483 for Bobolinks (n = 91), and 0.585 for Savannah Sparrows (n = 152). Fledge dates for first clutches ranged between 14 June and 23 July. Only one obligate grassland bird nest was parasitized by Brown-headed Cowbirds (Molothrus ater), for an overall brood parasitism rate of 0.004. Logistic-exposure models indicated that daily nest survival probabilities were higher in pastures/hayfields than in fallow fields. Our results, and those from other studies in the Northeast, suggest that properly managed cool season grassland habitats in the region may not act as ecological traps, and that obligate grassland birds in the region may have greater nest survival probabilities, and lower rates of Brown-headed Cowbird parasitism, than in many parts of the Midwest.

Was the removal of rabbits and house mice from Selvagem Grande beneficial to the breeding of Cory's shearwaters Calonectris diomedea borealis?

The breeding success of Cory's shearwaters Calonectris diomedea borealis at its important Atlantic colony on Selvagern Grande has been monitored periodically at two study plots since 1982. A successful eradication programme was implemented to remove two alien invasive mammals, rabbits Oryctolagus cuniculus and house mice Mus musculus, from the island during 2002. The availability of long-term breeding data for Cory's shearwaters on Selvagern Grande provided a unique opportunity to study the effects of the removal of rabbits and mice on seabird breeding. Annual observation of approximately 400 Cory's nests showed that significantly more birds fledged from both study sites in the five breeding seasons after the eradication than in the 13 seasons prior to it for which reliable breeding data were available. The numbers of young birds present at the time of fledging were an average of 47 and 23% greater than pre-eradication numbers at the two study sites. The eradication of rabbits and mice was simultaneous and, therefore, it was impossible to attribute the increased breeding success of Cory's shearwaters to the removal of one or other species. However, both are known to have adverse impacts on the breeding of nesting seabirds. These observations provide important justification for the implementation of further programmes for the removal of alien invasive mammals from oceanic islands.

Song development in birds: the role of early experience and its potential effect on rehabilitation success

Environmental conditions during the early life stages of birds can have significant effects on the quality of sexual signals in adulthood, especially song, and these ultimately have consequences for breeding success and fitness. This has wide-ranging implications for the rehabilitation protocols undertaken in wildlife hospitals which aim to return captive-reared animals to their natural habitat. Here we review the current literature on bird song development and learning in order to determine the potential impact that the rearing of juvenile songbirds in captivity can have on rehabilitation success. We quantify the effects of reduced learning on song structure and relate this to the possible effects on an individual's ability to defend a territory or attract a mate. We show the importance of providing a conspecific auditory model for birds to learn from in the early stages post-fledging, either via live- or tape-tutoring and provide suggestions for tutoring regimes. We also highlight the historical focus on learning in a few model species that has left an information gap in our knowledge for most species reared at wildlife hospitals.

Nest spacing and breeding performance in Short-toed Eagle Circaetus gallicus in northeast Greece

Aims: To describe the phenology and breeding success of one of the densest populations of Short-toed Eagle in Europe. Methods All nests in the Dadia-Lefkimi-Soufli forest in northeast Greece were located and visited regularly throughout the 1996-98 breeding seasons. Data on every stage of the breeding cycle were collected and related to among-year variation in the weather conditions during March to June. Results: A total of 58 pairs were located during the three-year study spread across 22 territories (the same territories are usually occupied each year). The nests were evenly spaced (mean of 2.7 km between nests). Adults arrived between mid-March and mid-April. Only one egg per nest was laid. Nestlings fledged on average after 68.9 days. Eagles departed between 8 September and 2 October. Conclusions: Arrival date determines laying date. The population size appears to be stable but the species has a relatively low reproductive rate and takes three to four years to mature, consequently it may be susceptible to stochastic or human-mediated factors.

Nest location, clutch size and nest success in the Scarlet Ibis Eudocimus ruber

Breeding success and nest-site characteristics were studied during the 1996-1997 breeding season in a colony of Scarlet Ibises Eudocimus ruber in south-eastern Brazil to test the hypothesis that nest-site characteristics and clutch size affect nest success. Two nesting pulses produced young, the earlier being more successful. Predation accounted for most failures during the first pulse, wind destruction during the second. A third pulse with few nests produced no young. Adult Ibises abandoned nests when they lost sight of other incubating birds. Logistic regression analysis indicated that nest success during the first pulse was positively related to clutch size, number of nests in the nest tree and in the nearest tree, and negatively to the distance to the nearest neighbour. During the second pulse there were significant negative associations between success, nest height and distance to the fourth nearest nest, and a positive association between success and nest cover. The results agree with the 'selfish herd' hypothesis, indicating that nest aggregation may increase breeding success, but the nest-site characteristics affecting success can differ over the course of one breeding season.

Influence of habitat on nest location and reproductive output of Montagu’s Harriers breeding in natural vegetation

We examined distribution and breeding success of semi-colonial Montagu’s Harriers (Circus pygargus) in relation to habitat in Castellón province (eastern Spain). Breeding areas used by harriers at a 1-km2 scale were characterised by having intermediate percentages of scrub cover, their nesting habitat, and also had intermediate coverage of herbaceous crops and non-irrigated orchards. Out of all habitat variables considered, only the percentage of herbaceous crops within 500 m from individual nests had a positive and significant effect on breeding output of the species, suggesting that this habitat may be efficiently used by harriers to forage. Breeding output was also related to laying date and number of breeding neighbours within 500 m around nests, with pairs laying later and having a higher number of breeding neighbours showing lower fledged brood sizes. Number of neighbours (but not laying date) was positively related to scrub cover within 500 m and to cover of herbaceous crops within 2,000 m. Conservation actions for Montagu’s Harrier in the study area should be aimed at preserving areas of scrub with nearby presence of herbaceous crops or natural grasslands. However, habitat improvement for semi-colonial species such as Montagu’s Harrier may not result in a change of species distribution area, and good habitat areas may remain unoccupied, as social factors like presence of conspecifics play an important role in breeding area selection for these species.

(Table 1) Characteristics of breeding male Adélie penguins (Pygoscelis adeliae) treated with varying degrees of corticosterone, Adélie land, Antarctica

Breeding individuals enter an emergency life-history stage when their body reserves reach a minimum threshold. Consequently, they redirect current activity toward survival, leading to egg abandonment in birds. Corticosterone (CORT) is known to promote this stage. How and to what extent CORT triggers egg abandonment when breeding is associated with prolonged fasting, however, requires further investigation. We manipulated free-living male Adelie penguins with CORT-pellets before their laying period. We then examined their behavioral response with respect to nest abandonment in parallel with their prolactin levels (regulating parental care), and the subsequent effects of treatment on breeding success in relieved birds. Exogenous CORT triggered nest abandonment in 60% of the treated penguins -14 days after treatment and induced a concomitant decline in prolactin levels. Interestingly, prolactin levels in treated penguins that did not abandon their nest were higher at the point of implantation and also after being relieved by females, when compared with abandoning penguins. Among successful birds, the treatment did not affect the number of chicks, nor the brood mass. Our results show the involvement of CORT in the decision-making process regarding egg abandonment in Adelie penguins when incubation is associated with a natural long fast. However, we suggest that CORT alone is not sufficient to trigger nest abandonment but that 1) prolactin levels need to reach a low threshold value, and 2) a rise in proteolysis (i.e. utilization of protein as main energy substrate) seems also to be required.

Modelling the effect of plant water use traits on yield and stay-green expression in sorghum

Post-rainy sorghum (Sorghum bicolor (L.) Moench) production underpins the livelihood of millions in the semiarid tropics, where the crop is affected by drought. Drought scenarios have been classified and quantified using crop simulation. In this report, variation in traits that hypothetically contribute to drought adaptation (plant growth dynamics, canopy and root water conducting capacity, drought stress responses) were virtually introgressed into the most common post-rainy sorghum genotype, and the influence of these traits on plant growth, development, and grain and stover yield were simulated across different scenarios. Limited transpiration rates under high vapour pressure deficit had the highest positive effect on production, especially combined with enhanced water extraction capacity at the root level. Variability in leaf development (smaller canopy size, later plant vigour or increased leaf appearance rate) also increased grain yield under severe drought, although it caused a stover yield trade-off under milder stress. Although the leaf development response to soil drying varied, this trait had only a modest benefit on crop production across all stress scenarios. Closer dissection of the model outputs showed that under water limitation, grain yield was largely determined by the amount of water availability after anthesis, and this relationship became closer with stress severity. All traits investigated increased water availability after anthesis and caused a delay in leaf senescence and led to a ‘stay-green’ phenotype. In conclusion, we showed that breeding success remained highly probabilistic; maximum resilience and economic benefits depended on drought frequency. Maximum potential could be explored by specific combinations of traits.

Climate forcing on avian life history

In the 21st century, human-induced global climate change has been highlighted as one of the most serious threats to ecosystems worldwide. According to global climate scenarios, the mean temperature in Finland is expected to increase by 1.8 4.0°C by the end of the century. The regional and seasonal change in temperature has predicted to be spatially and temporally asymmetric, where the High-Arctic and Antarctic areas and winter and spring seasons have been projected to face the highest temperature increase. To understand how species respond to the ongoing climate change, we need to study how climate affects species in different phases of their life cycle. The impact of climate on breeding and migration of eight large-sized bird species was studied in this thesis, taking food availability into account. The findings show that climatic variables have considerable impact on the life-history traits of large-sized birds in northern Europe. The magnitude of climatic effects on migration and breeding was comparable with that of food supply, conventionally regarded as the main factor affecting these life-history traits. Based on the results of this thesis and the current climate scenarios, the following not mutually exclusive responses are possible in the near future. Firstly, asymmetric climate change may result in a mistiming of breeding because mild winters and early spring may lead to earlier breeding, whereas offspring are hatching into colder conditions which elevate mortality. Secondly, climate induced responses can differ between species with different breeding tactics (income vs. capital breeding), so that especially capital breeders can gain advantage on global warming as they can sustain higher energy resources. Thirdly, increasing precipitation has the potential to reduce the breeding success of many species by exposing nestlings to more severe post-hatching conditions and hampering the hunting conditions of parents. Fourthly, decreasing ice cover and earlier ice-break in the Baltic Sea will allow earlier spring migration in waterfowl. In eiders, this can potentially lead to more productive breeding. Fifthly, warming temperatures can favour parents preparing for breeding and increase nestling survival. Lastly, the climate-induced phenological changes in life history events will likely continue. Furthermore, interactions between climate and food resources can be complex and interact with each other. Eiders provide an illustrative example of this complexity, being caught in the crossfire between more benign ice conditions and lower salinity negatively affecting their prime food resource. The general conclusion is that climate is controlling not only the phenology of the species but also their reproductive output, thus affecting the entire population dynamics.