Habitat selection rules in breeding mallards (Anas platyrhynchos): a test of two competing hypotheses

Ideal preemption and conspecific attraction are alternative hypotheses of the habitat selection rules used by individuals. According to the former an occupied site is assumed to be preempted and therefore not available for later arriving individuals, whereas according to the latter individuals are assumed to be attracted by conspecifics to occupied sites, rather than avoiding them. We studied these competing hypotheses in breeding mallards (Anas platyrhynchos) by a cross-over experiment in 2 years, introducing birds onto lakes before migratory wild mallards arrived. If mallards use the ideal preemptive rule, breeding density of wild mallards in experimental lakes should be lower and they should be occupied less frequently than control lakes, but if mallards use the conspecific attraction rule the reverse should be true. Our results allowed us to reject the ideal preemptive rule whereas the conspecific attraction rule was to some extent supported. We discuss these findings in relation to population limitation. The results suggest that the local breeding population studied is not limited by spacing behaviour related to habitat selection. Received: 14 July 1997 / Accepted: 17 November 1997     

Interspecific interactions and co-existence in dabbling ducks: observations and an experiment

We studied the possible role of resource limitation and interspecific competition in assemblages of dabbling ducks on breeding lakes in Finland and Sweden with observational and experimental data. After initial vegetation mapping and yearly censuses of ducks in 1985–1990, we collected observational data in 1991–1994 from 28 lakes with natural populations of mallard Anas platyrhynchos and teal A. crecca. Mallard and teal co-occur over vast areas in the Holarctic and they are the only breeding dabbling ducks on many oligotrophic lakes. Both species are migratory in our study regions, teal arriving later in spring than mallards. Log-linear model analysis of observational presence/absence data revealed a positive, not a negative, association between the species. This association was independent of habitat diversity as well as of lake size. Mallard-teal interaction was also studied in a cross-over introduction experiment in 32 other lakes in two years. Wing-clipped mallards were introduced to breeding lakes before the arrival of teal to induce resource limitation and interspecific competition, hypothesized to reduce lake use by teal. The density of mallard pairs on experimental lakes was 2.9–8.0 times higher than on controls, but there was no negative response by teal to the treatment. This is the first combined observational-experimental demonstration of lack of interspecific competition in waterfowl. Our results indicate that heterospecific attraction may affect species co-existence in dabbling ducks. Received: 28 October 1996 / Accepted: 13 January 1997     

Within-season sequential density dependence regulates breeding success in mallards Anas platyrhynchos

Density dependence in vital rates is a key issue in population ecology but remains largely unexplored experimentally. We studied breeding success, lake use, and prey availability in wild mallards Anas platyrhynchos on small nemoral lakes in a replicated, two-year cross-over experiment in which pair density was increased. The number of wild mallards that settled on lakes prior to introductions of extra pairs did not differ between control and introduction years. Introductions led to a lake-level reduction in the number of broods observed. However, the number of stage 2+ (almost fledged) ducklings did not differ between treatments, nor did lake utilization by nonbreeding adults, broods and ducklings. Prey resource availability differed greatly among lakes, but it did not correlate with breeding success. Partialling out the possible effect of food competition from wild adult nonbreeding mallards did not change this conclusion. Our study demonstrates sequential density dependence in breeding success; introductions caused a decrease in brood number, but despite fewer broods a similar number of nearly fledged ducklings were produced. We suggest that predation and/or lake change of broods soon after hatching created these patterns. We conclude that using a single and late measure of breeding success such as fledged birds can mask regulatory processes. Implications of density dependence and its relation to individual reproductive success are understood better if breeding success is decomposed into nest success, duckling survival and fledgling survival.     

Breeding success of sympatric dabbling ducks in relation to population density and food resources

Breeding success in sympatric mallard Anas platyrhynchos , teal A. crecca and wigeon A. penelope in a boreal watershed in Finland was studied for 12 years. Benthic and surface-emerging prey animals were trapped to obtain annual indices of food abundance. Mallard and teal were equally abundant over the years, being roughly twice as numerous as wigeon.
Pair density, brood:pair ratio and duckling:pair ratio were used to test the hypothesis that per capita breeding success decreases in a density-dependent fashion as either pair density or the number of nesting pairs per available food unit increases. In mallard we found no density-dependent patterns at all. In teal per capita brood production decreased as prey animals became relatively scarcer, but this interpretation may not be robust. In wigeon, however, there were two independent significant patterns of direct density-dependence in a temporal succession, i.e. between pair density and per capita brood production in the early part of the breeding season, and then between per capita abundance of surface-emerging insect prey and the number of ducklings per pair. Despite wide dietary overlap and frequent co-occurrence on single lakes among species in the guild, we found no evidence for interspecific density-dependent effects. We hypothesize that there is no or infrequent food limitation for breeding dabblers in this system, and that behavior may be the process behind the pattern of density-dependence in wigeon.     

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.     

Dynamics of habitat distribution in breeding mallards: assessing the applicability of current habitat selection models

The applicability of ideal free, ideal despotic and ideal preemptive habitat selection models to explain dynamics of habitat distribution of breeding mallards ( Anas platyrhynchos ) was explored. Data from 35 lakes studied between 1985 and 2000 were used to examine overall habitat distribution of breeding pairs, breeding success in different habitats, within-year order of habitat occupation, and density dependence of habitat distribution and breeding success. Two habitat types, rich and poor, were defined based on the structure and luxuriance of shore vegetation; each lake belonged to one or the other of the habitat types. Breeding pairs used the rich habitat more than expected, breeding density also being higher there than in the poor habitat. Both average brood density and breeding success were higher in the rich habitat than in the poor. Breeding success was not density dependent, neither when analysed separately for the habitat types, nor in the study area in general. Within season, arriving mallard pairs did not occupy rich lakes earlier than poor lakes. An isodar analysis based on between-year variation of the breeding density in rich and poor habitats revealed that habitat distribution of breeding pairs was not density dependent. By contrast, density in the rich habitat increased and proportional use of the poor habitat decreased with increasing overall population density, i.e. the rich habitat got increasingly crowded. None of the habitat selection models considered was applicable to explain the dynamics of habitat distribution of breeding mallards.     

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.     

Wetland Selection by Mallard Broods in Canada's Prairie-Parklands

ABSTRACT Although brood survival has a pronounced effect on population growth in mallards (Anas platyrhynchos), knowledge of brood ecology is more limited than for other vital rates. During 1993–1997 we collected wetland selection data from 210 radiomarked mallard broods on 15 study areas located throughout the Canadian Prairie-Parklands. We used information-theoretic approaches to select the best-approximating model of habitat selection in relation to wetland characteristics. Wetland permanence, cover type, width of flooded emergent vegetation, and interactions between these variables and date, moisture level, and dominant species of emergent vegetation were all important predictors of wetland selection. Mallard broods selected deeper wetlands, especially later in the brood-rearing season. Mallard broods also selected wetlands with large central expanses of open water and wide peripheral zones of flooded emergent cover. These habitat characteristics can most easily be met in landscapes that already contain an abundance and diversity of natural wetland habitats. Where such wetlands are unavailable, restoration or management of deeper wetlands may be necessary to meet the habitat requirements of mallard ducklings.     

Prior knowledge about spatial pattern affects patch assessment rather than movement between patches in tactile-feeding mallard

1. Heterogeneity in food abundance allows a forager to concentrate foraging effort in patches that are rich in food. This might be problematic when food is cryptic, as the content of patches is unknown prior to foraging. In such case knowledge about the spatial pattern in the distribution of food might be beneficial as this enables a forager to estimate the content of surrounding patches. A forager can benefit from this pre-harvest information about the food distribution by regulating time in patches and/or movement between patches. 2. We conducted an experiment with mallard Anas platyrhynchos foraging in environments with random, regular, and clumped spatial configurations of full and empty patches. An assessment model was used to predict the time in patches for different spatial distributions, in which a mallard is predicted to remain in a patch until its potential intake rate drops to the average intake rate that can be achieved in the environment. A movement model was used to predict lengths of interpatch movements for different spatial distributions, in which a mallard is predicted to travel to the patch where it expects the highest intake rate. 3. Consistent with predictions, in the clumped distribution mallard spent less time in an empty patch when the previously visited neighbouring patch had been empty than when it had been full. This effect was not observed for the random distribution. This shows that mallard use pre-harvest information on spatial pattern to improve patch assessment. Patch assessment could not be evaluated for the regular distribution. 4. Movements that started in an empty patch were longer than movements that started in a full patch. Contrary to model predictions this effect was observed for all spatial distributions, rather than for the clumped distribution only. In this experiment mallard did not regulate movement in relation to pattern. 5. An explanation for the result that pre-harvest information on spatial pattern affected patch assessment rather than movement is that mallard move to the nearest patch where the expected intake rate is higher than the critical value, rather than to the patch where the highest intake rate is expected.     

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.     

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.     

Social Indices of Breeding Productivity in Parkland Mallards

Abstract Social indices were developed to assess breeding productivity of waterfowl based on weekly roadside surveys of social groupings (i.e., pairs, lone M, flocked M). We calculated social indices for mallard (Anas platyrhynchos) populations breeding on 16 study sites in the Canadian parklands from 1993 to 1998 using 7 previously developed indices. We also calculated duckling:pair ratios from our roadside counts, and we obtained independent measures of nesting effort, nesting success, female success, and fledging rate for these same 16 sites from a concurrent telemetry study. Social indices were correlated (r² = 0.28-0.67) with telemetry-based measures of breeding productivity in 5 of 7 cases, with the strongest relationships deriving from indices that emphasized renesting effort. The 2 ineffective social indices (r² ≤ 0.13) both measured early onset of nesting activity. Duckling:pair ratios could be calculated more easily from the same survey data and also were correlated (r² = 0.26-0.48) with measures of breeding productivity. Because surveys measuring late-nesting effort also can enumerate early hatched ducklings, we recommend that waterfowl researchers use duckling:pair ratios rather than social indices because ducking:pair ratios are more easily interpretable. Development of sightability-adjustment factors for pair and duckling surveys could further enhance the utility of duckling:pair ratios as indices of breeding productivity in mallards.     

Looking like mother makes mallard ducklings dominant over their siblings

Colour variation in time and space among animals may affect social relationships such as pairing and dominance interactions. For instance, some birds are naturally sensitive to leg colour, with some colours being more visible or attractive than others. The colour of the leg-rings used to mark birds may thus be related to behavioural and reproductive variables. Most studies have investigated this effect for adults during reproduction, but leg-ring colour may also affect the behaviour of young birds. We tested the potential effect of leg-ring colours on the within-brood dominance hierarchy of mallard (Anas platyrhynchos) ducklings while each brood formed a stable and exclusive family unit with its mother. Ducklings did not acquire a within-brood dominance rank according to the colour of their own ring. This result suggests that mallards may not have a sensory bias for a given colouration. However, ducklings wearing a ring of the same colour as one of the two rings of their mother were dominant over their siblings. We discuss the potential behavioural and methodological implications of this result.     

Land Cover Switching in Autumn by Female Mallards in Ohio

Autumn waterfowl habitat management often focuses on providing high energy food resources to attract and concentrate waterfowl for harvest. Similarly, many waterfowl conservation plans assume food resources are the primary, controllable limiting factor influencing waterfowl distribution during migration; however, hunting-related disturbance also influences waterfowl distribution in autumn. We investigated factors influencing mallard (Anas platyrhynchos) movements in an intensively hunted and food-rich landscape in Ohio, USA, during autumn of 2015 and 2016. We used locations from female mallards equipped with global positioning system (GPS) back-pack-transmitters to determine the probability of mallards switching land cover types based on seasonal and daily patterns of hunting disturbance and to determine the distances mallards moved between cover types that offer refuge and those that offer food resources as evidence for or against food resource depletion during the hunting season. Mallards switched cover types to exploit food-rich but intensively hunted locations nocturnally and cover type switching during times subject to disturbance increased significantly from the early segment to the late segment of hunting season. Distances mallards moved between refuge cover types and food-rich cover types did not change over the duration of the study. Hunting disturbance is a key variable influencing autumn movements and distribution of mallards, and mallards in a food-rich and intensively hunted landscape likely employ nocturnal foraging as a strategy to survive autumn migration. Nocturnal foraging behavior has consequences for waterfowl managers tasked with providing quality waterfowl hunting opportunities because ducks that forage only at night are largely unavailable to hunters. © 2020 The Wildlife Society.     

Clutch size and body size at first reproduction in Daphnia pulicaria at different levels of food and predation

Field data from seven alpine lakes in Serra da Estrela. Portugal.show that reproduction in Daphnia may be as efficiently controlledby fish predation and copepod predation on eggs in brood cavitiesas it is by food limitation. Body length and clutch size estimatesin Daphnia pulicaria revealed high inter- and intra-populationvariability in maturation size (body size at first reproduction).and in number of eggs per clutch. Daphnia at first maturationin lakes stocked with rainbow trout were half the size of thosefound in fishless lakes (body length of 0.86–0.95 and1.55–1.81 mm. respectively). The mean number of eggs perclutch was reduced to a similar degree by food limitation, predationby fish and copepod predation on eggs in brood cavities, butthe underlying mechanisms of this reduction were different.Food limitation caused smaller clutch sizes in all individuals,so variation remained the same. Fish predation caused the selectiveremoval of individuals with maximum clutches, so variation decreased.Copepod predation caused removal of eggs from brood cavitiesof randomly infested females, so that variation increased, particularlyat a high food level when non-infested females carried largeclutches of eggs.     

Within-brood social status and consequences for winter hierarchies amongst Mallard Anas platyrhynchos ducklings

Juveniles of many birds establish dominance hierarchies within family social units, only to leave and compete to acquire dominance status in new social groups. Little is known about the role of sex, body mass, size or experience during the duckling period on subsequent dominance rank and adult social relationships. We used captive Mallard Anas platyrhynchos ducklings to test for the role of individual characteristics and growth parameters in establishing within-brood hierarchies, the maintenance of within-brood hierarchies in the subsequent wintering group and differences in social ranks between broods. Strong stable linear hierarchies were present within each brood and, later, within each phase of the winter. There was a reorganisation of the hierarchical order between the duckling period and early winter, but only few modifications afterwards during the winter. None of the tested “hatching”, “duckling” and “adult” traits explained either the within-brood or the winter hierarchies, but winter rank was related to brood of origin with ducklings from the same brood having similar social ranks. These differences between broods were maintained through the whole winter in most cases, though one brood drastically progressed in the hierarchy during late-winter. These results suggest that the factors affecting the establishment of social relationships within broods differ from those in winter groups, and that brood-related mechanisms influence social relationships during winter. We discuss our results in the light of direct and indirect maternal influence.     

Helper effects in the azure‐winged magpie Cyanopica cyana in relation to highly‐clumped nesting pattern and high frequency of conspecific nest‐raiding

In avian cooperative breeding systems, many benefits obtained by social pairs from the presence of helpers have been uncovered. However, until now, the factors that determine the type of assistance helpers provide and the responses of social pairs have not been well illustrated. We examined the contribution of helpers to cooperative groups and the relevant responses of dominant pairs in the azure‐winged magpie Cyanapica cyana which breeds on the Tibetan Plateau. We used the capture–mark–recapture method to identify helpers. Results showed that helpers were mostly the yearling sons of dominant pairs. They mainly contributed to the cooperative group in three ways, courtship‐feeding the incubation female, provisioning the brood, and defending the nest. For responses of dominant pairs, we unexpectedly found that clutch size was not influenced by the presence of helpers at the nest. However, cooperative groups had higher brood feeding rates than biparental nests and their feeding pattern also differed to that of the latter. Consequently, nestlings in cooperative groups had larger fledging body mass than that in biparental nests. By examining reasons for nest failure, we revealed that conspecific nest‐raiding contributed to more nest failure than any other natural predators. Because of the contribution of helpers in defending against both predators and conspecific nest‐raiders, cooperative groups had higher survival rate than biparental nests. Thus, our findings suggest that in a highly‐clumped nesting pattern, factors concerning the risk of nest predation, rather than that influencing food supply, play an important role in determining helper effects and responses of aided dominant pairs.     

Long-Term Foraging Expeditions (Molongo) among the Baka Hunter-Gatherers in the Northwestern Congo Basin, with Special Reference to the “Wild Yam Question”

While “Pygmy” hunter-gatherers have generally been assumed to be the original inhabitants of the central African rainforest, recent studies dispute the viability of subsistence in tropical rainforests through foraging alone. There are, however, few studies that are based on sound data of actual long-term foraging. This paper examines the viability of “pure” foraging in tropical rainforests, based on data from participant observation of a long-term foraging expedition among the Baka in the northwestern Congo Basin. The group was observed to subsist solely on wild food resources, particularly wild yams, for two-and-half months. The study was carried out during the dry season when food resources are generally thought to be scarce in tropical rainforests.     

An Attraction Pheromone from Heads of Worker Vespula germanica Wasps

Vespula germanica is a social wasp that efficiently exploits food resources. Odor cues, derived from substances located in wasp heads, have been proved to be central in conspecific attraction in this species. However, it remains unknown whether this attraction is related to foraging or defense responses. In this study we analyze conspecific attraction under two different contexts: at the nest entrance (defense) and under foraging conditions. We also test wasp response with two dosages of head extract and crosschecked the attractiveness of extracts obtained from different populations. We found no evidence of alarm response to head extracts either at the nest entrance or under foraging conditions. Moreover, no differences in attractiveness were found to both doses tested. Head extracts attract similarly in the same or a different population suggesting that conspecific attraction is not restricted to colony nestmates.