Presence of fish affects lake use and breeding success in ducks

Several previous studies indicate that presence of fish has negative effects on waterbirds breeding on lakes, owing either to competition for common invertebrate prey or fish predation on ducklings/chicks. However, others have reported results to the contrary and it remains unresolved what factors trigger, inhibit, and modulate fish–waterbird interactions. The present study was designed to test the effect of fish presence per se, with a minimum of variation in possibly confounding environmental variables. Thus, after stratifying for area, depth, altitude, pH, and total phosphorus we compared 13 lakes with and 12 without fish (mainly pike Esox lucius and perch Perca fluviatilis) with respect to (i) general species richness of waterbirds, (ii) species-specific utilization and breeding success of two dabbling ducks (mallard Anas platyrhynchos and teal Anas crecca) and a diving duck (goldeneye Bucephala clangula). General species richness of waterbirds was higher on fishless lakes. Overall use (bird days) and brood number of teal and goldeneye were higher on fishless lakes. The latter also had more benthic and free-swimming prey invertebrates compared to lakes with fish. Mallard use, mallard brood number, and abundance of emerging insects did not differ between lake groups. Generalized linear models including fish presence as factor and considering seven environmental variables as covariates, confirmed that all waterbird variables except mallard days and broods were negatively correlated to fish presence. There was also a residual positive relationship of lake area on general species richness, teal days, and teal broods. Our data demonstrate a stronger effect of fish presence on diving ducks and small surface feeding ducks than on large surface-feeding ducks. We argue that observed patterns were caused by fish predation on ducks rather than by fish–duck competition for common prey.     

Habitat associations of ducks during different phases of the breeding season

Habitat use of mallard Anas platvrhynchos , teal A crecca. wigeon A penelope and goldeneye Bucephala clangula was studied m southern Finland during 1988–1991 Emphasis included evaluation of the importance of food and vegetation structure and revealing of the patterns of habitat use throughout the breeding season Average lake scores of pairs, broods, and juveniles of all species were on the luxunant end of a principal component analysis axis describing habitat structure A more detailed examination, however, revealed clear differences in habitat distributions both between and within species Generally, habitat distributions seemed to shift toward the luxuriant from pairs to broods However, goldeneye juveniles used both the most luxuriant and the poorest habitats more than expected When all phases of the breeding cycle are considered, habitat luxuriance seemed to be more important for dabbling ducks and nektonic invertebrates more important for the goldeneye Emerging insects were most important to the teal     

Pike predation affects breeding success and habitat selection of ducks

1. Fish and ducks often belong to the same local food web, and several studies indicate that there is a general negative effect of fish on breeding ducks. This pattern has so far been addressed mainly within the framework of competition for common invertebrate prey, while predation by large fish as a force behind settlement and abundance patterns in ducks remains largely unknown. This is the first study to address the effect of fish predation on breeding ducks, isolated from that of competition, and the first experiment to explore the ability of ducks to identify and avoid lakes with high risk of fish predation. 2. We used a before–after control–impact design and 11 naturally fishless lakes. Waterfowl on the lakes were surveyed during the breeding season of 2005. Large adult pike (Esox lucius) were added to two lakes in early spring 2008, and waterfowl surveys were repeated on all 11 lakes. 3. Pike introduction did not affect the number of pairs on lakes during the nesting season in any of three focal duck species (mallard Anas platyrhynchos, teal Anas crecca, and goldeneye Bucephala clangula). During the brood‐rearing season, however, there was a decrease in duck days in teal and goldeneye in lakes with pike, with similar trends observed in mallard. The number of goldeneye ducklings was also significantly lower in lakes with pike. We were unable to determine whether the response was attributable to direct pike predation or to broods leaving experimental lakes, but in either case, our study demonstrates high fitness costs for ducks breeding on lakes with pike. 4. The apparent inability of nesting ducks to detect pike and the clear fitness implications may influence the annual recruitment of ducks on a larger scale as pike are both common and widespread. Vegetation complexity and food abundance are likely to be of overriding importance when breeding ducks are choosing a nesting site. As pike have a strong influence on breeding birds, relying on vegetation and cues of food abundance, while ignoring indicators of predation risk from fish, could lead to lakes with pike acting as an ecological trap.     

The beaver as an ecosystem engineer facilitates teal breeding

Positive interactions are important in influencing communities. Facilitation is one such influence in which a species makes the environment more suitable for others species. Ecological engineers are powerful facilitators as they physically modify habitats in a way that alters resource availability. We studied the effect of beaver flooding on breeding success of a dabbling duck, the teal. Our long-term (1988-2001) study was conducted in a boreal area where dabbling ducks face resource limitation. We focused on two aspects of facilitation: resource enhancement and habitat amelioration. We found that pond use by teal broods systematically increased upon beaver flooding. Beaver ponds harboured more resource, i.e. aquatic invertebrates than undisturbed waters, and invertebrate abundance was reflected as enhanced teal brood density. Beaver inundated shores were shallower than those of other waters, making their habitat structure more favourable to ducklings. Consequently, teal brood mortality was lower in beaver ponds than in waters unaffected by beaver. In boreal areas beaver facilitate teal breeding by enhancing both resource abundance and habitat structure.     

Density-dependent reproductive output in relation to a drastically varying food supply: getting the density measure right

When a limiting resource (e.g. food) varies drastically between years, and population density is measured in the conventional way as individuals per area, demographic processes such as productivity and survival may erroneously be considered density-independent. We tested the hypothesis that if the variation in a limiting resource is not taken into account in the density measure, this may lead to erroneous conclusions about the density-dependence of demographic variables. We studied the food-related variation in productivity of bramblings Fringilla montifringilla , an insectivorous passerine bird, using 19 years of standardised insect censusing, bird censusing and mist-netting of birds in subalpine birch forest in Swedish Lapland. The yearly variation in our measure of brambling per capita productivity (numbers of juveniles per adult trapped) was explained to 30–40% by the larvae abundance of the moth Epirrita autumnata . Taking larvae density into account, no other environmental variable (inferred predation pressure, breeding phenology, and summer temperature) was significantly related to variation in reproductive output. There was no effect of brambling population density on per capita productivity, that is, when density was measured the conventional way, productivity seemed density-independent. However, per capita productivity was significantly and negatively correlated to the food-related population density (population density divided by larval density), supporting the hypothesis that not including a limiting resource into the density measure may indeed lead to erroneous conclusions about the density-dependence of demographic variables.     

Density-dependent decline of breeding success in an introduced, increasing mute swan Cygus olor population

An increasing population of the mute swan Cygnus olor , was studied from its very establishment in 1976 until 1998. As the number of pairs increased, there was a decline in all production parameters, including the average number of clutches per pair, the average number of broods per clutch, and the average number of fledged young per brood. In a multiple regression analysis covering the whole breeding season, the number of pairs and the average number of clutches per pair explained 71% of the variation in the average number of fledglings per pair. During the brood season, the average number of fledglings per brood was an additionally important parameter in explaining fledgling production. Habitat quality seemed to affect breeding: in places occupied earlier more cygnets fledged from a clutch than in habitats inhabited later. The decreased production of young very likely reflects density-dependent effects on reproduction. This density dependence seems to operate on the breeding grounds, since winter harshness did not affect breeding success. Density-dependent processes started acting already when pairs were beginning to nest, and continued during the brood period. Density dependence has apparently not been detected at the pair stage in earlier studies of bird populations.     

Seasonal patterns in reproductive success of temperate‐breeding birds: Experimental tests of the date and quality hypotheses

For organisms in seasonal environments, individuals that breed earlier in the season regularly attain higher fitness than their late‐breeding counterparts. Two primary hypotheses have been proposed to explain these patterns: The quality hypothesis contends that early breeders are of better phenotypic quality or breed on higher quality territories, whereas the date hypothesis predicts that seasonally declining reproductive success is a response to a seasonal deterioration in environmental quality. In birds, food availability is thought to drive deteriorating environmental conditions, but few experimental studies have demonstrated its importance while also controlling for parental quality. We tested predictions of the date hypothesis in tree swallows (Tachycineta bicolor) over two breeding seasons and in two locations within their breeding range in Canada. Nests were paired by clutch initiation date to control for parental quality, and we delayed the hatching date of one nest within each pair. Subsequently, brood sizes were manipulated to mimic changes in per capita food abundance, and we examined the effects of manipulations, as well as indices of environmental and parental quality, on nestling quality, fledging success, and return rates. Reduced reproductive success of late‐breeding individuals was causally related to a seasonal decline in environmental quality. Declining insect biomass and enlarged brood sizes resulted in nestlings that were lighter, in poorer body condition, structurally smaller, had shorter and slower growing flight feathers and were less likely to survive to fledge. Our results provide evidence for the importance of food resources in mediating seasonal declines in offspring quality and survival.     

Regulation of numbers in the Great tit (Aves: Passeriformes)

The census data of the Great tit collected by Perrins (1965) and others in Marley Wood near Oxford are analysed for density-dependence. Clutch size and hatching success are density-dependent and sufficiently so to regulate the population at the observed level (assuming that there is in addition a fairly large density-independent mortality). There may also be some weak density-dependent mortality outside the breeding season. The density-dependent variations in clutch size are probably in the main due to shortage of available food and density-dependent hatching failure is caused by predation. Territorial behaviour has been shown experimentally to determine breeding density, and may produce a density-dependent effect outside the breeding season. These three factors are responsible for regulation of the Great tit population in Marley Wood.     

Effects of food abundance, density and climate change on reproduction in the sparrowhawk Accipiter nisus

The reproductive success of predators depends on abiotic environmental conditions, food abundance and population density, and food abundance, density and their interactions may respond to changes in climatic conditions. Timing of reproduction by five of the eight numerically most common prey of the sparrowhawk Accipiter nisus advanced significantly since 1971, during a period of temperature increase. There was no evidence that mean laying date or any other reproductive parameter of sparrowhawks changed consistently during the study period 1977–1997. Laying date advanced and percentage of unsuccessful female sparrowhawks decreased with beech mast in the current year, an index of food abundance for avian prey. Mean laying date of sparrowhawks was advanced in warmer springs, and although mean clutch size was not larger in warm than in cold springs, mean brood size of successful pairs and breeding success increased in such springs, showing that sparrowhawks enjoyed a fitness gain when reproducing early. The timing of sparrowhawk reproduction with respect to the peak in abundance of fledgling prey increased, from a good match between mean timing of fledging by prey and maximum demand for food by the predator in 1977, to reproduction occurring later than the peak in fledging prey availability in 1997. The size of the breeding population of sparrowhawks was not predicted by mean spring temperature, the size of the breeding population the previous year or beech mast crop. The size of the post-breeding population was predicted by size of the breeding and post-breeding population the previous year and by the proportion of unsuccessful females the current year. These findings imply that sparrowhawks did not respond to change in climate, although climate changed the timing of reproduction by the main prey species.     

Of mice and mallards: positive indirect effects of coexisting prey on waterfowl nest success

Coexisting prey species interact indirectly via their shared predators when one prey type influences predation rates of the second prey type. In a temperate system where the predominant shared predator is a generalist, I studied the indirect effects of rodent populations on waterfowl nest success, both within the nesting season among sites and among years. Among six to ten upland fields (14 to 27 ha), mallard ( Anas platyrhynchos ) nest success was positively correlated with rodent abundance in all three years of the study. After removing year effects, mallard nest success remained positively correlated with the relative abundance of rodents. Of the rodent species present, California voles ( Microtus californicus ) were the most important coexisting prey type influencing nest success. Among years, mallard nest success was positively correlated with vole abundance; the asymptotic relationship suggests a threshold response to vole abundance, beyond which predators become satiated and additional voles do little to affect nest success. I tested and rejected three alternative explanations for the observed positive correlation between mallard nest success and rodent abundance that do not involve an indirect effect of coexisting prey populations. The influences of dense nesting cover, nesting density, and predator activity did not explain the observed patterns of nest success. These results suggest that rodent populations buffer predation on waterfowl nests, both within and among years, via the behavioral responses of shared predators to coexisting prey.     

Assessment of territory quality and its effects on breeding success in a migrant passerine, the Wheatear Oenanthe oenanthe

This paper examines how a returning migrant assesses the quality of an area as a breeding territory before the period of peak food demand and how effective the assessment is in terms of breeding success. Male Wheatears Oenanthe oenanthe return from Africa to choose territories in the Breckland of eastern England about March, females arriving shortly after males. The food supply was predictable: prey densities during the breeding period (egg-laying to chick independence) were strongly correlated with prey densities at the same sites during the period of arrival and territory establishment. Prey densities were also related to vegetation structure, averaging highest on short turf. Male arrival date and territory size were not significantly related to prey density but were strongly related to vegetation structure, implying that birds used vegetation as an indirect clue to prey availability. Neither territory size nor nest spacing appeared to affect nest losses caused by predators. The major variations in number of young fledged (other than predation) were caused by the number of nestlings hatched and presence of a second brood. Both early arrival and an early first brood improved first-brood success and were necessary for a second brood. Not all birds which arrived early bred early enough for a second brood. First-brood hatching date was strongly negatively correlated with pre-breeding prey availability but not significantly related to vegetation structure. Thus by using vegetation as a clue to habitat quality, some pairs suffered reduced breeding success. This result implies that birds may not be able adequately to assess prey density directly at the time of territory establishment. The critical period for food availability may not be the period of peak demand (nestling period), when food is relatively abundant, but is probably the pre-breeding period when females must accumulate reserves for eggs and when the food supply is poor. Food supply during this period may determine the timing of breeding and the ability to rear a second brood, and may thus have a greater effect on breeding success.     

Population trends in bird species at Dungeness, Kent

Most wintering wildfowl species increased in numbers on the Dungeness Reserve during the period 1969-87, particularly teal, gadwall and wigeon. Wigeon, teal and gadwall showed the fastest rates of increase. An exponential model provided a better fit to the data for smew and gadwall than a linear one. Populations were more or less as variable in number in the early years of management (1969-77) as in subsequent years (1978-86).
Of wintering and passage wader species, turnstone, redshank, black-tailed godwit and common sandpiper increased. Snipe showed a decline in abundance in spring. Numbers of black terns, the only non-wading migrant analysed, remained variable over the period.
Breeding tufted duck and great-crested grebe increased in numbers during the period 1969-86 whereas mallard declined. Black-headed gull numbers increased, as did exponentially-breeding common and sandwich terns. Of four passerine species analysed, only breeding reed warblers increased during the period 1968-86, whereas numbers of breeding sedge warbler and whitethroat remained stable. Wheatear numbers declined in early years but have increased since 1983.
Increases in species richness were documented in wintering and passage, and breeding species.     

Population variability among three small mammal species in the semiarid Neotropics: the role of density-dependent and density-independent factors

We addressed the role of density-dependent (direct and delayed) and density-independent (precipitation) factors in shaping the dynamics of fluctuating populations of three small mammal species. Using a stepwise regression procedure, we tested the effects of nonlagged population density (log₁₀ N_t-1), lagged population density (log₁₀ N_t-2), and annual precipitation on the per capita rate of population change of Phyllotis darwini, Akodon olivaceus , and Thylamys elegans in two habitat types of a semiarid region of Chile. The most irruptive species ( P. darwini ) showed direct and delayed density-dependent effects in equatorial subpopulation, and only direct density-dependence in polar subpopulation. The per capita rates of population change of A. olivaceus showed direct density-dependent and precipitation effects in both habitats types, while T. elegans showed direct density-dependence and precipitation effects in the equatorial subpopulation but only a marginal effect of direct density-dependence in the polar subpopulation. The presence of delayed density-dependent strongly suggests the importance of biological interactions in shaping the dramatic irruptions exhibited by P. darwini.     

Hatching in dabbling ducks and emergence in chironomids: a case of predator–prey synchrony?

It has been hypothesized that dabbling ducks (Anas spp.) time breeding to coincide with annual regional peaks in emerging dipterans, especially Chironomidae, which are important prey for newly hatched ducklings. However, this hypothesis has never been evaluated in a replicated lake-level study, including year effects in emergence patterns. We collected duck and invertebrate data from 12 lakes during the nesting seasons 1989–1994 in a watershed in southern Finland. The oligotrophic study lakes are typical of the boreal Holarctic, as are the three focal duck species: mallard Anas platyrhynchos L., widgeon Anas penelope L and teal Anas crecca L. Hatching of ducklings showed a clear peak in relation to ambient phenology (annual ice-out date of lakes), whereas chironomid emergence was more erratic and showed no clear peak at the lake level, although total watershed-level emergence was somewhat higher before and long after the duck hatching peak. Thus, we find no evidence that ducklings hatch in synchrony with abundance peaks of emerging chironomids. There was large within-year temporal variation in chironomid emergence among lakes, but this was not correlated with ambient temperature. The rank of individual lakes with respect to the abundance of emerging chironomids was consistent among as well as within years, a predictability that ought to make adaptive lake choice by ducks possible. On the lake level, there was a positive correlation between the total amount of emerging chironomids and brood use. We argue that emergence patterns of chironomids on typical boreal lakes are neither compressed nor predictable enough to be a major selective force on the timing of egg-laying and hatching in dabbling ducks. Despite spatial (among-lake) patterns of abundance of emerging chironomids being predictable within and among years, the observed pattern of brood use suggests that other factors, e.g. habitat structure, also affect lake choice.     

Factors affecting species number and density of dabbling duck guilds in North Europe

We addressed how species number and pair density in guilds of co-existing species is related to habitat structure, and to the abundance and diversity of food resources using the assemblage of seven species of dabbling ducks (genus Anas ) breeding in 60 lakes distributed over six regions in temperate north Europe
Partial correlation and multiple regression revealed that species richness was best predicted by habitat structural diversity as indexed by a principal component analysis based on 18 vegetation and lake characteristics, and by the abundance of aquatic and emergent prey We found no effect of lake size or prey size diversity on species richness Pair density was correlated with the percentage of shoreline with horsetails ( Equisetum ), by habitat structural diversity and by the abundance of emergent invertebrate prey Neither prey size diversity nor abundance of aquatic prey correlated with pair density Species richness and pair density in North European duck guilds vary both with habitat structure and prey availability     

Breeding in the stable boreal landscape: lake habitat variability drives brood production in the teal (Anas crecca)

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Marine copepod diversity patterns and the metabolic theory of ecology

The ongoing climate change has improved our understanding of how climate affects the reproduction of animals. However, the interaction between food availability and climate on breeding has rarely been examined. While it has been shown that breeding of boreal birds of prey is first and foremost determined by prey abundance, little information exists on how climatic conditions influence this relationship. We studied the joint effects of main prey abundance and ambient weather on timing of breeding and reproductive success of two smaller (pygmy owl Glaucidium passerinum and Tengmalm’s owl Aegolius funereus) and two larger (tawny owl Strix aluco and Ural owl Strix uralensis) avian predator species using long-term nation-wide datasets during 1973–2004. We found no temporal trend either in vole abundance or in hatching date and brood size of any studied owl species. In the larger species, increasing late winter or early spring temperature advanced breeding at least as much as did high autumn abundance of prey (voles). Furthermore, increasing snow depth delayed breeding of the largest species (Ural owl), presumably by reducing the availability of voles. Brood size was strongly determined by spring vole abundance in all four owl species. These results show that climate directly affects the breeding performance of vole-eating boreal avian predators much more than previously thought. According to earlier studies, small-sized species should advance their breeding more than larger species in response to increasing temperature. However, we found an opposite pattern, with larger species being more sensitive to temperature. We argue that this pattern is caused by a difference in the breeding tactics of larger mostly capital breeding and smaller mostly income breeding owl species.     

Quantifying density dependence in a bird population using human disturbance

Although density dependence has long been recognised as vital to population regulation, there have been relatively few studies demonstrating it spatially in wildlife populations, often due to the confounding effects of variation in habitat quality. We report on a study of woodlarks Lullula arborea, a species of European conservation concern, breeding on lowland heath in Dorset, England. We take the novel approach of utilising the birds’ response to human disturbance, which resulted in much of the variation in density but had no direct impact on demographic rates. Within years, in sites with greater density there were smaller mean chick masses, lower post-fledging survival, and higher rates of nestling mortality attributed to starvation. The effects on clutch size and fledging success were confounded by the area of grassland within a site. There was no effect on brood size. Density dependence also operated within sites between years: as density increased there were reductions in mean chick mass and post-fledging survival, while nestling mortality attributed to starvation increased. Density-dependent effects on clutch size were only weakly regulatory, whereas density-dependent starvation and post-fledging mortality rates contributed strongly to differences in overall breeding output. Heavier chicks (when 7 days old) were significantly more likely to fledge and less likely to starve. Broods with heavier chicks were more likely to supply recruits to the breeding population. Nestling mass was not a factor in survival in the immediate post-fledging period, suggesting that density-dependent processes act independently on this stage. We conclude that the number of birds per hectare of suitable habitat is a valid means of expressing density, and that habitat acts as a surrogate for food abundance through which density dependence operates on the woodlark population.     

Changes in the wildfowl populations wintering on the Severn Estuary

Monthly winter counts of seven major wildfowl species show that, since 1960, Bewick's swan, shelduck, wigeon, teal and pintail have significantly increased in both the Severn and Britain as a whole; European White-fronted geese have shown no trend on the Severn but have significantly declined in Britain; mallard show no trend on the Severn but have increased in Britain. All except shelduck show a correlation between the trends in Britain and the Severn Estuary, while all but mallard and pintail rank as internationally important on the Severn. In recent years the Severn has assumed a considerable importance for pochard and tufted ducks. The status of many species in the Severn Estuary is dependent on the management of key areas.     

Experimental evidence for density-dependent survival in mallard (Anas platyrhynchos) ducklings

It is unresolved to what extent waterfowl populations are regulated by density-dependent processes. By doing a 2-year crossover perturbation experiment on ten oligotrophic boreal lakes we addressed the hypothesis that breeding output is density dependent. Wing-clipped mallard (Anas platyrhynchos) hens were introduced with their own brood and then monitored for 24 days. Predicted responses were that per capita duckling and hen survival would be lower in high-density than in low-density treatments. Survival was evaluated by model fitting in program MARK. Density, year, and lake were used as main effects, while day after introduction, a weather harshness index, and presence of hens were covariates. Daily survival in ducklings was lower in the high-density treatment, but this effect was year dependent. The highest-ranking model for duckling survival also included a positive effect of duckling age and presence of hens, and a negative effect of harsh weather. Density did not affect female survival although there was a prominent year effect. The highest-ranking model for female survival also included negative effects of day after introduction and harsh weather. This is the first study to report density-dependent survival in experimentally introduced ducklings in a natural setting. Implications for population dynamics and management of harvested populations are far-reaching if such regulation occurs in some years, but not in others.