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     

Experimental duck plague in blue-winged teal and Canada geese

Ten adult blue-winged teal (Anas discors) and six Canada goose (Branta canadensis) goslings were inoculated with liver tissue from a natural case of duck plague in a wild mallard (Anas platyrhynchos). Four additional teal were placed in contact with the inoculated ducks. Inoculated teal died 63.5-68 hr after inoculation; two of the contact teal died 161-162 hr after exposure. Three of the goslings died 119-133 hr after inoculation, the others were killed when moribund 90-133 hr postinfection. The clinical course of disease was extremely rapid in both species. Signs were limited to sudden onset of profound weakness, ataxia, tremors and terminal convulsions. The only consistent gross lesion in the teal was a small dark spleen; half the teal also had inconspicuous foci of epithelial necrosis in the distal esophagus and in the cloaca. Goslings had more severe lesions, with focal hepatic necrosis visible in all, and mucosal necrosis over the intestinal lymphoid tissue, and intestinal hemorrhage in three birds each. Microscopic lesions in both species were similar to those reported in mallards. Duck plague might be overlooked at necropsy of blue-winged teal, because of the paucity of gross lesions.     

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.     

Differences in endozoochorous dispersal between aquatic plant species, with reference to plant population persistence in rivers

Summary 1. In river ecosystems, populations are continuously subjected to unidirectional downstream currents resulting in a downstream movement of populations. To ensure long-term population persistence in rivers, organisms must have a mechanism for upstream dispersal, which allows them to re-colonise upstream areas.
2. In this study we assessed differences in the potential for endozoochorous seed dispersal of Sparganium emersum and Sagittaria sagittifolia , two aquatic plant species with different seed morphologies, by mallard ( Anas platyrhynchos ) and teal ( Anas crecca ), two duck species with different body weights.
3. We found no significant differences in seed retrieval (the proportion of ingested seeds retrieved after gut passage) and seed retention time (time between seed ingestion and retrieval), between mallard and teal, despite the difference in body weights. We did find a significantly higher germination (%) over retention time of S. emersum seeds retrieved from teal compared with mallard, most likely related to a more efficient removal of the seed coat during passage through the gut of teal.
4. There were large differences between S. emersum versus S. sagittifolia in: (i) seed retrieval (22.65 ± 20.8% versus 1.60 ± 2.4%, respectively); (ii) seed retention time in duck gut, with a maximum of 60 h versus 12 h; (iii) the effect of gut passage on seed germination, with an increase of approximately 35% versus a decrease of 25%; and (iv) the effect of gut passage on seed germination rate, with an acceleration of 10 days versus a delay of 3 days on average. The results show that S. emersum has a higher potential for endozoochorous dispersal by ducks and postdispersal establishment than S. sagittifolia .
5. We propose that, in rivers, bird-mediated seed dispersal may promote re-colonisation of upstream areas, enabling long-term plant population persistence.     

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.     

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     

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

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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.     

Avian influenza viruses and paramyxoviruses in wintering and resident ducks in Texas

Cloacal swabs were collected from teal (Anas crecca, Anas cyanoptera, Anas discors), mottled duck (Anas fulvigula) and northern pintail (Anas acuta) in Brazoria County, Texas, USA, during February 2001, mottled ducks during August 2001, and blue-winged teal (A. discors) during February 2002. Prevalence of avian influenza virus (AIV) infections during each sampling period were 11, 0, and 15%, respectively. The hemagglutinin (H) subtypes H2 and H7 were detected in both years, while the H8 subtype was detected in 2001 and the H1 subtype was detected in 2002. Avian paramyxovirus type 1 (APMV-1) was isolated from 13% of mottled ducks sampled in August 2001 and 30.7% of teal in February 2002. The season of isolation of both viruses and the majority of the AIV subtypes detected in this study are not typical based on previous reports of these viruses from North American ducks.     

Variation in home-range size and movements of wintering dabbling ducks

Despite a long historical record of radio-tracking analyses, basic home-range information is still lacking for most common waterfowl species, especially during the winter. We investigated how dabbling duck home ranges and daily foraging movements are influenced by extrinsic (site, temperature, date) and intrinsic factors (species, sex, age). We radio-tagged and monitored 125 individuals of three duck species (mallard Anas platyrhynchos, Eurasian teal A. crecca crecca and northern pintail A. acuta) in three French wetlands over four winters. Home-range sizes for a given species varied greatly among our study sites. Moreover, species differed according to home-range structure and distance traveled to reach their foraging grounds (teal had a more patchy home range and traveled farther distances than mallards). Foraging distances increased with temperature and time (over the winter season), but this effect differed among species, suggesting that they behave differently in response to food depletion and/or cold weather. The commuting behavior (i.e., the decision to leave the roost at night for foraging) differed among species and season. Teals were more risk-prone because they were more likely to leave the roost at night. In our study, ducks foraged at distances of 1–2 km from roosts, whereas distances of 2–48 km have been recorded in North America. We suggest that food supply, hunting pressure or population density may account for these inter-continental differences.     

Fish–duck interactions in boreal lakes in Finland as reflected by abundance correlations

We studied the hypothesis that fish play an important role in lake use by ducks (pairs and broods) in boreal lakes. The study was based on densities of different duck and fish species in 28 boreal lakes in southern Finland. We focused on the three most common duck species (mallard Anas platyrhynchos, green-winged teal A. crecca and common goldeneye Bucephala clangula) and on the three most common fish species (perch Perca fluviatilis, roach Rutilus rutilus and pike Esox lucius) in the region. We considered both competitive and predatory interactions between ducks and fish, the perch and roach being potential competitors with ducks and the pike a potential predator of ducks. We found a negative association between green-winged teal brood density and total fish density, the other duck species having only a weak association with total fish density. When the three fish species were considered separately, a negative association, suggesting food competition, was found between perch, green-winged teal and goldeneye, whereas the role of roach as a food competitor seemed to be of minor importance. We did not find any clear signs of predatory effects of pike on ducks. Our results suggest that food competition is a more important factor than pike predation in affecting lake use by ducks in oligotrophic boreal environments in southern Finland.     

Gastrointestinal helminths of the Mexican duck, Anas platyrhynchos diazi Ridgway, from north central Mexico and southwestern United States

Twenty-five species of helminths, recovered from the gastrointestinal tracts of 129 Mexican ducks from Mexico and the United States, were all new host records. The species included: Echinoparyphium recurvatum, Echinostoma revolutum, Hypoderaeum conoideum, Notocotylus attenuatus, Prosthogonimus cuneatus, Zygocotyle lunata, Anomotaenia ciliata, Cloacotaenia megalops, Diorchis bulbodes, Diorchis sp., Drepanidotaenia lanceolata, Echinocotyle rosseteri, Fimbriaria fasciolaris, Fimbriarioides sp., Hymenolepis sp. 1, Hymenolepis sp. 2, Sobolevicanthus gracilis. Corynosoma constrictum, Polymorphus minutus, Amidostomum acutum, Echinuria sp., Epomidiostomum crami, Hystrichis varispinosus, Rusguniella arctica, and Tetrameres sp. Fimbriarioides sp. occurred predominantly in ducks from south-central Chihuahua, Mexico. The distributions for the other 24 species of the helminths were not significantly different among the four collecting areas. The helminth fauna for the 32 complete specimens and 97 intestinal tracts was distributed relatively evenly among the hosts with a calculated mean evenness of 0.77 +/- 0.15 and 0.89 +/- 0.03 respectively. The parasite fauna was more similar to those of the black duck Anas rubripes Brewster of eastern North America (53%), the mallard, Anas platyrhynchos (L.) (49%), and the mottled duck, Anas fulvigula Ridgway, from Florida (45%), than to the green-winged teal, Anas crecca (L.) (36%), the gadwall, Anas strepera (L.) (30%), and the American wigeon, Anas americana Gmelin (26%), collected in the Southwest.     

Recolonization of breeding waterfowl communities by the whooper swan: vacant niches available

Breeding population of the whooper swan Cygnus cygnus has increased dramatically in Finland during the last 50 yr, from ca 15 to 1500 pairs. We studied if the recent recolonization by the whooper swan into waterfowl communities in Finland has had negative effects on Anas species belonging to the same foraging guild. Using data of foraging behaviour and ecomorphology of the whooper swan and Anas species, we studied the position of the former in the dabbling guild. We used long-term duck census data from different parts of Finland to study changes in breeding numbers of Anas species in lakes that have been recently colonized by the whooper swan and in control lakes that have not been colonized. Overall, foraging and ecomorphological similarity between the whooper swan and the other species decreased in the following order: pintail A. acuta (most similar), mallard A. platyrhynchos , shoveler A. clypeata , garganey A. querquedula. wigeon A. penelope. teal A. crecca (least similar). We did not find evidence of adverse impact by the whooper swan colonization on population densities of the other species. Neither did we find any indication that species relatively more similar with the whooper swan in terms of foraging ecology would show relatively more negative response to the whooper swan colonization. Our results suggest that whooper swans have occupied vacant niche space when colonizing boreal waterfowl communities.     

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.     

Disturbance, Herbivory, and Propagule Dispersal Control Dominance of an Invasive Grass

Despite the dramatic changes invasive plants cause to ecosystems and communities, factors that control dominance of invasive species after establishment in a community are poorly understood. Most active management relies on catastrophic disturbances of invasive-dominated communities to increase richness and diversity of plant communities. This study examines the importance of propagule dispersal and deer herbivory on continued dominance of Phalaris arundinacea after a non-catastrophic, short-term disturbance to monotypic stands of this invasive grass. The disturbance caused no change in P. arundinacea cover among treatments during any year of the study and, thus, simulates disturbance intensity more likely to be encountered in unmanaged settings. Despite the small disturbance, the combinations of disturbance + seeding and disturbance + seeding + deer exclusion caused greater species richness than controls even three years after disturbance. Increased invasion of P. arundinacea stands caused few effects on the dominant, as P. arundinacea biomass was unaffected after the first year. Selective herbivory by deer of species other than P. arundinacea increased the effects of disturbance and seeding, and aided in continued dominance of the grass. The tolerance of P. arundinacea for direct anthropogenic effects, including poor water quality and hydroperiod fluctuations, and indirect effects, such as increased herbivory by historically high deer populations, indicates the complexity of determining persistence of invasive species.     

Spatio-temporal dynamics in waterbirds during the non-breeding season: Effects of local movements,migration and weather are monthly,not yearly

Predicting population sizes and dynamics of mobile organisms is important for trend estimation, but this is difficult when the origin of individuals cannot be determined, i.e. residents and migrants are indistinguishable in the field.Here we examine fluctuations in populations during the non-breeding season (autumn to spring) of seven waterbird species, which co-occur on 122 neighbouring water bodies in south-west Germany. We asked whether site-level abundance patterns can be predicted by weather variables, and whether these effects vary over months and years. We used a spatially explicit hidden Markov model to estimate the effects of local and regional movement on population fluctuations.Although the species varied in their dynamics, with different amounts of movement estimated between sites, several patterns were common across species: density effects were highly month-specific, but with little variation in the strength of effects over years. The abundance of most species was positively related to temperature, especially in winter.The common teal Anas crecca and common pochard Aythya ferina were the most site-fidelic species once they were present in the study area, but also exhibited the strongest regional migration. The mallard Anas platyrhynchos, tufted duck Aythya fuligula and great crested grebe Podiceps cristatus each behaved more like a single population, as individuals more frequently moved between sites and abundance fluctuations at sites were not explained by migratory movements alone.Our study shows that the strength of population parameters and environmental forces can be decomposed into monthly and yearly effects. Estimating the unknown origin and movement of individuals may show that commensurate populations of mobile species may have different underlying dynamics, while responding similarly to environmental factors.     

绿翅鸭、琵嘴鸭、斑嘴鸭越冬期的生存能

在野鸭越冬期逐旬测定了绿翅鸭、琵嘴鸭、斑嘴鸭笼养下的生存能。结果表明,三种野鸭越冬期每天摄食量分别为绿翅鸭32.4克,琵嘴鸭54.9克、斑嘴鸭80.2克。食物利用率依次为75.6％;73.7％;73.8％。生存代谢能依次为102.7千卡/只·天;173.6千卡/只·天;248.2千卡/只·天。总能量的摄入有三个高峰,分别出现在秋季迁来后、冬季寒潮后和春季迁飞前。 三种野鸭越冬期体重呈现出二低二高的变化形式。1983年与1963年野外野鸭体重相比较,绿翅鸭平均降低了20％,斑嘴鸭降低了12％,表明太湖湖泊生态系统对野鸭的栖息越冬有恶化的趋势。 本文分三个季节拟合了体重与生存能的关系式,比较了不同季节生存能与静止代谢能的差异,表明野鸭是通过提高体廓的隔热能力,增加体内能量的储存以及减少活动和生产的能量消耗来渡过寒冷又缺少食物的冬季的。     

Sitting ducklings: Timing of hatch,nest departure,and predation risk for dabbling duck broods

For ground‐nesting waterfowl, the timing of egg hatch and duckling departure from the nest may be influenced by the risk of predation at the nest and en route to wetlands and constrained by the time required for ducklings to imprint on the hen and be physically able to leave the nest. We determined the timing of hatch, nest departure, and predation on dabbling duck broods using small video cameras placed at the nests of mallard (Anas platyrhynchos; n = 26), gadwall (Mareca strepera; n = 24), and cinnamon teal (Anas cyanoptera; n = 5). Mallard eggs began to hatch throughout the day and night, whereas gadwall eggs generally started to hatch during daylight hours (mean 7.5 hr after dawn). Among all species, duckling departure from the nest occurred during daylight (98%), and 53% of hens typically left the nest with their broods 1–4 hr after dawn. For mallard and gadwall, we identified three strategies for the timing of nest departure: (a) 9% of broods left the nest the same day that eggs began to hatch (6–12 hr later), (b) 81% of broods left the nest the day after eggs began to hatch, and (c) 10% of broods waited 2 days to depart the nest after eggs began to hatch, leaving the nest just after the second dawn (27–42 hr later). Overall, eggs were depredated at 10% of nests with cameras in the 2 days prior to hatch and ducklings were depredated at 15% of nests with cameras before leaving the nest. Our results suggest that broods prefer to depart the nest early in the morning, which may best balance developmental constraints with predation risk both at the nest and en route to wetlands.     

Evidence of the trade-off between starvation and predation risks in ducks

The theory of trade-off between starvation and predation risks predicts a decrease in body mass in order to improve flight performance when facing high predation risk. To date, this trade-off has mainly been validated in passerines, birds that store limited body reserves for short-term use. In the largest avian species in which the trade-off has been investigated (the mallard, Anas platyrhynchos), the slope of the relationship between mass and flight performance was steeper in proportion to lean body mass than in passerines. In order to verify whether the same case can be applied to other birds with large body reserves, we analyzed the response to this trade-off in two other duck species, the common teal (Anas crecca) and the tufted duck (Aythya fuligula). Predation risk was simulated by disturbing birds. Ducks within disturbed groups were compared to non-disturbed control birds. In disturbed groups, both species showed a much greater decrease in food intake and body mass during the period of simulated high risk than those observed in the control group. This loss of body mass allows reaching a more favourable wing loading and increases power for flight, hence enhancing flight performances and reducing predation risk. Moreover, body mass loss and power margin gain in both species were higher than in passerines, as observed in mallards. Our results suggest that the starvation-predation risk trade-off is one of the major life history traits underlying body mass adjustments, and these findings can be generalized to all birds facing predation. Additionally, the response magnitude seems to be influenced by the strategy of body reserve management.