Robust estimation of survival and contribution of captive‐bred Mallards Anas platyrhynchos to a wild population in a large‐scale release programme

The survival of captive‐bred individuals from release into the wild to their first breeding season is crucial to assess the success of reintroduction or translocation programmes, and to assess their potential impact of wild populations. However, assessing the survival of captive‐bred individuals following their release is often complicated by immediate dispersal once in the wild. Here, we apply Lindberg's robust design model, a method that incorporates emigration from the study site, to obtain true estimates of survival of captive‐bred Mallards Anas platyrhynchos, a common duck species released on a large scale in Europe since the 1970s. Overall survival rate from release in July until the onset of the next breeding season in April was low (0.18 ± 0.07 se) and equivalent to half the first‐year survival of local wild Mallards. Higher overall detectability and temporary emigration during the hunting period revealed movements in response to hunting pressure. Such low survival of released Mallards during their first year may help prevent large‐scale genetic mixing with the wild population. Nevertheless, by combining our results with regional waterfowl counts, we estimated that a minimum of 34% of the Mallards in the region were of captive origin at the onset of the breeding season. Although most released birds quickly die, restocking for hunting may be of sufficient magnitude to affect the wild population through genetic homogenization or loss of local adaptation. Robust design protocols allow for the estimation of true survival estimates by controlling for permanent and temporary emigration and may require only a moderate increase in fieldwork effort.     

Minimal evidence of interspecific hybridisation between the Yellow-billed Duck and introduced Mallard in central and northwestern South Africa

Hybridisation is the interbreeding of genetically distinct groups that can lead to introgression – an exchange of genetic material between species. Hybridisation is of conservation concern when an alien invasive species is involved, as it can lead to a loss of local genetic adaptations and genetic diversity. Hybridisation is a significant threat for many dabbling ducks where interbreeding with the closely related invasive Mallard Anas platyrhynchos is extremely common. Phenotypic evidence suggests that Mallard populations in South Africa hybridise with the indigenous Yellow-billed Duck Anas undulata. The aim of this study was to determine the incidence of hybridisation between Yellow-billed Ducks, occurring in central and northwestern South Africa, and introduced Mallards. Genetic variation between Mallards, Yellow-billed Ducks, and their inferred hybrids was assessed using mitochondrial and microsatellite DNA markers. All samples inferred to be hybrids based on the phenotype were found to have Yellowbilled Duck mitochondrial DNA and showed minimal evidence of admixture across the microsatellite markers. Thus, these results do not support the notion that hybridisation between Mallards and Yellow-billed Ducks is prevalent in central and northwestern South Africa. However, hybridisation could be occurring where Mallards are found in higher abundance, such as in the Western Cape Province. Therefore, continued monitoring of this potential hybridisation should be performed frequently and throughout South Africa.     

Environmental Change and Extended Phenotypes: Does Eutrophication Influence Nest Building in Sticklebacks?

Many species use extended phenotypes, such as purpose‐built nests, to increase their reproductive success. These traits have to be adjusted to local environmental conditions to maximize fitness. An important question is whether species are able to adjust their extended phenotypes to human‐induced rapid environmental changes. We investigated whether populations of threespine stickleback, Gasterosteus aculeatus, exposed to different degrees of human‐induced eutrophication during the last decades, have differentiated phenotypically in their nest‐building behaviour. Stickleback males build nests that they use both in mate attraction and for offspring protection and whose characteristics vary with environmental conditions. We allowed males from parallel pairs of mildly and severely eutrophied habitats to build nests under standardized conditions in the laboratory. We recorded the time it took the males to build a nest, the size and neatness of the completed nest and the use of nest ornaments. We found eutrophication at the site of capture to influence nest‐building time – males from eutrophied habitats built faster. However, eutrophication did not alter nest structure or the use of nest ornaments. This is probably because the nests are concealed in vegetation or under stones and females cannot evaluate them before they have followed the male to the nest, and predators cannot detect them before close to the nest. Thus, reduced long‐range visibility does not influence the use of nests as mate choice cues or for offspring protection. This contrasts with a recorded effect of eutrophication on courtship behaviour, whose efficiency depends on long‐range visibility. This suggests that traits are adjusted to eutrophication depending on the influence of eutrophication on the function of the traits.     

Low survival after release into the wild: assessing “the burden of captivity” on Mallard physiology and behaviour

Captive-reared animals used in reinforcement programs are generally less likely to survive than wild conspecifics. Digestion efficiency and naive behaviour are two likely reasons for this pattern. The Mallard is a species with high adaptability to its environment and in which massive reinforcement programs are carried out. We studied physiological and behavioural factors potentially affecting body condition and survival of captive-reared Mallards after being released. Digestive system morphology and an index of body condition were compared among three groups: captive-reared birds remaining in a farm (control), captive-reared birds released into the wild as juveniles (released) and wild-born birds (wild). We also compared behaviour and diet of released vs. wild Mallards. Finally, we conducted a 1-year survival analysis of captive-reared birds after release in a hunting-free area. Gizzard weight was lower in control Mallards, but the size of other organs did not differ between controls and wild birds. The difference in gizzard weight between released and wild birds disappeared after some time in the wild. Diet analyses suggest that released Mallards show a greater preference than wild for anthropogenic food (waste grain, bait). Despite similar time-budgets, released Mallards never attained the body condition of wild birds. As a consequence, survival probability in released Mallards was low, especially when food provisioning was stopped and during harsh winter periods. We argue that the low survival of released Mallards likely has a physiological rather than a behavioural (foraging) origin. In any case, extremely few released birds live long enough to potentially enter the breeding population, even without hunting. In the context of massive releases presently carried out for hunting purposes, our study indicates a low likelihood for genetic introgression by captive-reared birds into the wild population.     

Factors affecting reproductive performance in captive Mallard ducks

Propagation of wild-strain Mallards (Anas platyrhynchos) in captivity is hindered by low egg fertility. Therefore, we studied the effects of captivity, age, mate choice and isolation on reproductive parameters of Mallards. Captive drakes had smaller immature testes than free-flying Mallard drakes. Captive yearling ducks weighed less than adult ducks at the beginning of the breeding season, but no differences were found between their initial clutch size, egg volume or number of clutches laid. Yearling pairs had lower egg fertility (7%) than adult pairs (80%). Egg fertility was higher (51 vs 21%) in self-chosen pairs than in randomly assigned pairs. Isolation of ducks, however did not influence egg production.     

Competitive foraging in mallards: “Ideal free’ ducks

Mallards (Anas platyrhynchos L.) distribute themselves between two patches of food in a close approximation to the distribution predicted by the ideal free model. However an important assumption of this model is violated since the despotic behaviour of some individuals results in different birds receiving unequal payoffs. The distribution of the birds between the food patches is influenced by the distribution of these despots. Evidence is presented to suggest that the ducks initially use the frequency of supply of food items at a patch to assess its profitability, but they can, over a longer time scale, use other cues.     

Viral replication, persistence in water and genetic characterization of two influenza A viruses isolated from surface lake water

Water-borne transmission has been suggested as an important transmission mechanism for Influenza A (IA) viruses in wild duck populations; however, relatively few studies have attempted to detect IA viruses from aquatic habitats. Water-isolated viruses have rarely been genetically characterized and evaluation for persistence in water and infectivity in natural hosts has never been documented. In this study, we focused on two IA viruses (H3N8 and H4N6 subtypes) isolated from surface lake water in Minnesota, USA. We investigated the relative prevalence of the two virus subtypes in wild duck populations at the sampling site and their genetic relatedness to IA viruses isolated in wild waterbirds in North America. Viral persistence under different laboratory conditions (temperature and pH) and replication in experimentally infected Mallards (Anas platyrhynchos) were also characterized. Both viruses were the most prevalent subtype one year following their isolation in lake water. The viruses persisted in water for an extended time period at constant temperature (several weeks) but infectivity rapidly reduced under multiple freeze-thaw cycles. Furthermore, the two isolates efficiently replicated in Mallards. The complete genome characterization supported that these isolates originated from genetic reassortments with other IA viruses circulating in wild duck populations during the year of sampling. Based on phylogenetic analyses, we couldn't identify genetically similar viruses in duck populations in the years following their isolation from lake water. Our study supports the role for water-borne transmission for IA viruses but also highlights that additional field and experimental studies are required to support inter-annual persistence in aquatic habitats.     

Direct and indirect effects of winter harshness on the survival of Mallards Anas platyrhynchos in northwest Europe

To understand population dynamics it is necessary to understand vital rates, which may be affected by a wide range of factors including environmental variables such as weather. Weather conditions can affect birds’ vital rates directly through increased mortality due to impaired conditions, or indirectly via changes in body condition and/or behaviour. Most understanding of direct and indirect effects of weather comes from studies of breeding birds, whereas the situation in non‐breeding periods is less clear. Here, we analysed annual survival of non‐breeding Mallard Anas platyrhynchos, the most hunted waterfowl species in Europe, and assessed whether survival is related directly to winter harshness and/or indirectly via changes in winter recovery distributions. Recovery data on Mallards, initially marked in southeast Sweden, were analysed with an information‐theoretic approach using program mark . Over 10 000 Mallards were marked in two time periods, 1964–1982 and 2002–2008, of which 13.3 and 4.7%, respectively, were later recovered. Mallards had lower annual survival in the early trapping period (0.58–0.63) than in the later period (0.69–0.71), with no clear effects of sex, age or year. Within each study period, winter harshness did not directly correlate with survival. However, milder winters may have contributed indirectly to higher survival in the second period, as winter harshness data were correlated with the distances to recovery positions for females, and also because winter recovery areas have shifted northeast during the past decades, possibly indicating a shortened migratory distance. Migration is costly, and there is therefore a likely linkage between migration behaviour and survival of dabbling ducks, in which direct as well as indirect effects of winter harshness may play a role. Other factors, such as hunting pressure, are also likely to have changed in the past decades, and may also have contributed to improved survival of Mallards in northwest Europe.     

Sex differences in body composition of wintering Mallards (Anas platyrhynchos): possible implications for survival and reproductive performance

Capsule Wintering female Mallards have a higher fat load than males which may compensate for their lower body size and ultimately improve their fitness.

Aims To investigate whether sex-related differences in biological processes in winter are associated with differences in body nutrient storage and flight capabilities and, if so, the possible implications for survival and reproductive performance.

Methods We analysed the body composition and the flight parameters of 35 male and 35 female Mallards wintering in western Europe on the Rhine river.

Results Males were 15% heavier and 18% larger than females, but the latter carried 27% more fat per unit body mass. Relative to body lipid mass, body protein mass was 11% lower for females. Wing loading was 7% lower in females and power loading was similar in both sexes.

Conclusions High lipid mass gives females survival capabilities to food shortage similar to males. This helps sustain them through their prebasic moult and breeding success. Heavier muscles in males compensate for their higher wing loading and thus improve pairing success. Wintering and reproductive strategies in Mallard seem to be closely linked through body reserves.     

Nesting ecology of a naturalized population of Mallards Anas platyrhynchos in New Zealand

Investigating the reproductive ecology of naturalized species provides insights into the role of the source population's characteristics vs. post‐release adaptation that influence the success of introduction programmes. Introduced and naturalized Mallards Anas platyrhynchos are widely established in New Zealand (NZ), but little is known regarding their reproductive ecology. We evaluated the nesting ecology of female Mallards at two study sites in NZ (Southland and Waikato) in 2014–15. We radiotagged 241 pre‐breeding females with abdominal‐implant transmitters and measured breeding incidence, nesting chronology and re‐nesting propensity. We monitored 271 nests to evaluate nest survival, clutch and egg size, egg hatchability and partial clutch depredation. Breeding incidence averaged (mean ± se) 0.91 ± 0.03, clutch size averaged 9.9 ± 0.1 eggs, 94 ± 2% of eggs hatched in successful nests, partial depredation affected 6 ± 1% of eggs in clutches that were not fully destroyed by predators, and re‐nesting propensity following failure of nests or broods was 0.50 ± 0.003. Nesting season (first nest initiated to last nest hatched) lasted 4.5 months and mean initiation date of first detected nest attempts was 28 August ± 3.3 days. Smaller females were less likely to nest, but older, larger or better condition females nested earlier, re‐nested more often and laid larger clutches than did younger, smaller or poorer condition females. Younger females in Southland had higher nest survival; cumulative nest survival ranged from 0.25 ± 0.007 for adult females in Waikato to 0.50 ± 0.007 for yearling females in Southland. Compared with Mallards in their native range, the nesting season in NZ was longer, clutches and eggs were larger, and nest survival was generally greater. Different predators and climate, introgression with native heterospecifics and/or the sedentary nature of Mallards in NZ may have contributed to these differences.     

The effect of egg size variability on thermoregulation of Mallard (Anas platyrhynchos) offspring and its implications for survival

Summary There is a range of egg size phenotypes in Mallards (Anas platyrhynchos) that has a large genetic component. It was hypothesized that egg size variation could play an important role in survival of newly hatched ducklings during their first few days out of the nest when they are most susceptible to thermal stress and starvation. Precocial young must be physiologically capable of maintaining homeothermy in order to spend adequate time foraging. Duckling size at hatching was highly correlated with egg mass, and those hatching from heavier eggs were able to maintain homeothermy at colder environmental temperatures than those from lighter eggs. Heavy ducklings had significantly lower mass-specific cooling rates, but lower critical temperature did not vary significantly among ducklings of different size. Although insulation and energy reserves were not proportionally greater in larger ducklings, those hatching from heavier eggs can survive starvation longer than those from lighter eggs. The relative cold tolerance of young from light and heavy eggs will affect the ratio of time spent foraging to time spent being brooded by the female parent. Although there is no direct evidence that selection is acting on egg size, variation in this trait within a population could be maintained by fluctuating environmental conditions at hatch.     

The influence of male dominance on social interactions in black ducks and mallards

The influence of male dominance on male-female social interactions was examined in black ducks, Anas rubripes, and mallards, A. platyrhynchos, that were raised from hatch in four different groups (male and female mallards; male and female black ducks; male mallards and female black ducks; male black ducks and female mallards). The mate preference of females, independent of the influence of male dominance, was determined by exposing females to four caged, isolated males, one from each different group. All females preferred the type of male they had been raised with since hatch. Females were then exposed to the same four males, now free-swimming, to determine how male dominance influenced the initial preference. Under these conditions, each female associated primarily with the most dominant of the four males, regardless of her initial mate preference. Mallards were the most dominant males in all tests. These results demonstrate that male dominance influences social interactions and is one factor contributing to hybridization between these two species.     

Simulated eutrophication and browning alters zooplankton nutritional quality and determines juvenile fish growth and survival

The first few months of life is the most vulnerable period for fish and their optimal hatching time with zooplankton prey is favored by natural selection. Traditionally, however, prey abundance (i.e., zooplankton density) has been considered important, whereas prey nutritional composition has been largely neglected in natural settings. High‐quality zooplankton, rich in both essential amino acids (EAAs) and fatty acids (FAs), are required as starting prey to initiate development and fast juvenile growth. Prey quality is dependent on environmental conditions, and, for example, eutrophication and browning are two major factors defining primary producer community structures that will directly determine the nutritional quality of the basal food sources (algae, bacteria, terrestrial matter) for zooplankton. We experimentally tested how eutrophication and browning affect the growth and survival of juvenile rainbow trout (Oncorhynchus mykiss) by changing the quality of basal resources. We fed the fish on herbivorous zooplankton (Daphnia) grown with foods of different nutritional quality (algae, bacteria, terrestrial matter), and used GC‐MS, stable isotope labeling as well as bulk and compound‐specific stable isotope analyses for detecting the effects of different diets on the nutritional status of fish. The content of EAAs and omega‐3 (ω‐3) polyunsaturated FAs (PUFAs) in basal foods and zooplankton decreased in both eutrophication and browning treatments. The decrease in ω‐3 PUFA and especially docosahexaenoic acid (DHA) was reflected to fish juveniles, but they were able to compensate for low availability of EAAs in their food. Therefore, the reduced growth and survival of the juvenile fish was linked to the low availability of DHA. Fish showed very low ability to convert alpha‐linolenic acid (ALA) to DHA. We conclude that eutrophication and browning decrease the availability of the originally phytoplankton‐derived DHA for zooplankton and juvenile fish, suggesting bottom‐up regulation of food web quality.     

Eutrophication in Australian Rivers, Reservoirs and Estuaries – A Southern Hemisphere Perspective on the Science and its Implications

Australian science has made rapid advances in the last decade in understanding eutrophication processes in inland waters and estuaries. The freshwater research on which these advances are based was triggered by well-publicised blooms of cyanobacteria during the 1980s and early 1990s, particularly a 1000 km long bloom on the Darling River. In estuaries the study which greatly enhanced our understanding but simultaneously served to stimulate further research into estuarine eutrophication, the Port Phillip Bay Study, was initially designed to address perceived problems of toxicants in the Bay but provided profound insights into drivers for, and ecosystem responses to, eutrophication. Subsequent estuarine research has largely been stimulated by management questions arising from Australia’s increasing coastal development for residential purposes. The research has shown that some of the beliefs extant at the time of the blooms were incorrect. For example, it is now clear that stratification and light penetration, not nutrient availability, are the triggers for blooms in the impounded rivers of southeastern Australia, although nutrient exhaustion limits the biomass of blooms. Again, nitrogen seems to play as important a role as phosphorus does in controlling the biomass of these freshwater blooms. The research has also shown that aspects of eutrophication, such as nutrient transport, are dominated by different processes in different parts of Australia. Many of the biophysical processes involved in eutrophication have now been quantified sufficiently for models to be developed of such processes as sediment-nutrient release, stratification, turbidity and algal growth in both freshwater and estuarine systems. In some cases the models are reliable enough for the knowledge gained in particular waterbodies to be applied elsewhere. Thus, there is now a firm scientific foundation for managers to rely upon when managing algal blooms. Whilst these findings have already been presented to managers and communities throughout Australia, there is still a considerable way to go before they are absorbed into their modus operandi.     

Development of a phytoplankton indicator system for the ecological assessment of brackish coastal waters (German Baltic Sea coast)

The implementation of the European Water Framework Directive (WFD) requires the development of ecologically-based classification systems for anthropogenically-induced eutrophication in all types of water bodies. Due to the inherent high temporal and spatial variability of hydrological and geochemical parameters of the coastal waters of the southern Baltic Sea, discrimination between anthropogenic impact and natural variability is necessary. The development of statistical methods for this discrimination was the main aim of this study. These methods were used to derive indicative phytoplankton parameters for different stages of eutrophication for the investigation area. For this purpose, a long-term phytoplankton data series was analysed, which covered a broad salinity and eutrophication gradient. In order to detect eutrophication effects, the analysis was restricted to phytoplankton spring bloom events and to the salinity range between 5 and 10 psu, i.e. superimposing seasonal and hydrodynamic effects were eliminated. An artificial abiotic degradation vector was developed based on four typical water quality parameters. A total of 11 potentially indicative phytoplankton parameters on different taxonomical levels arose from a correlation analysis with this degradation vector. These indicators were then tested for their ability to discriminate between three eutrophication levels. Finally, seven phytoplankton indices could be proposed: total phytoplankton biovolume, the percentage of diatoms and the biovolume of different size ranges of diatoms and one indicative species (Woronichinia compacta). Guest editors: A. Razinkovas, Z. R. Gasiūnaitė, J. M. Zaldivar & P. Viaroli European Lagoons and their Watersheds: Function and Biodiversity     

Factors influencing seaweed responses to eutrophication: some results from EU-project EUMAC

Seaweed responses to eutrophication and their role in coastal eutrophication processes were compared at 8 different sites along the European coasts from the Baltic to the Mediterranean as part of the EU-ENVIRONMENT Project Marine Eutrophication and benthic Macrophytes (EUMAC). Structural and functional changes of marine benthic vegetation typical of eutrophic waters, in particular mass development (blooms) of certain seaweeds, are not merely the result of increased nutrient loading, but must be attributed to complex interactions of primary and secondary effects during the eutrophication process. Due to species-specific physiological properties of the algae (nutrient kinetics, growth potential, light, temperature requirements), the combined effects of abiotic and biotic factors on juvenile or adult developmental stages control the development of algal blooms in different ways. In particular the role of light, temperature, water motion and oxygen depletion, as well as of grazers, on early and adult developmental stages of the algae are considered. The result are discussed in the context of coastal eutrophication control and management. This revised version was published online in June 2006 with corrections to the Cover Date.     

High influenza a virus infection rates in mallards bred for hunting in the camargue, South of france

During the last decade, the role of wildlife in emerging pathogen transmission to domestic animals has often been pointed out. Conversely, far less attention has been paid to pathogen transmission from domestic animals to wildlife. Here, we focus on the case of game restocking, which implies the release of millions of animals worldwide each year. We conducted a 2-year study in the Camargue (Southern France) to investigate the influence of hand-reared Mallard releases on avian influenza virus dynamics in surrounding wildlife. We sampled Mallards (cloacal swabs) from several game duck facilities in 2009 and 2010 before their release. A very high (99%) infection rate caused by an H10N7 strain was detected in the game bird facility we sampled in 2009. We did not detect this strain in shot ducks we sampled, neither during the 2008/2009 nor the 2009/2010 hunting seasons. In 2010 infection rates ranged from 0 to 24% in hand-reared ducks. The 2009 H10N7 strain was fully sequenced. It results from multiple reassortment events between Eurasian low pathogenic strains. Interestingly, H10N7 strains had previously caused human infections in Egypt and Australia. The H10 and N7 segments we sequenced were clearly distinct from the Australian ones but they belonged to the same large cluster as the Egyptian ones. We did not observe any mutation linked to increased virulence, transmission to mammals, or antiviral resistance in the H10N7 strain we identified. Our results indicate that the potential role of hand-reared Mallards in influenza virus epizootics must be taken into account given the likely risk of viral exchange between game bird facilities and wild habitats, owing to duck rearing conditions. Measures implemented to limit transmission from wildlife to domestic animals as well as measures to control transmission from domestic animals to wild ones need to be equally reinforced.     

Wasserlinsen als Nutzpflanzen

Duckweeds as crop plants Members of the plant family Lemnaceae (duckweeds) are not only interesting because they represent the smallest flowering plants; they possess also the fastest rates of producing biomass. As aquatic plants, duckweed production is not in competition with other agricultural crops that require fertile land while the cultivation of duckweeds does not contribute to further eutrophication of surface water. Instead, they can be cultivated on municipal or agricultural waste water and remove the nutrients during their propagation and growth. Duckweeds can thus be used for cleaning of waste water and the resulting biomass can be valuable starting material for animal feeds and the production of biofuels. Research focusing on these goals has begun to transfer from research laboratories to pilot plants in different parts of the world, e.g. in New Jersey and North Carolina, USA; Chengdu, P. R. China; and Armidale, Australia.     

MITOCHONDRIAL GENE TREES AND THE EVOLUTIONARY RELATIONSHIP OF MALLARD AND BLACK DUCKS

We assayed restriction site differences in mitochondrial DNA (mtDNA) within and among allopatric populations of the Mallard (Anas platyrhynchos) and the American Black Duck (A. rubripes). The observed mtDNA clones grouped into two phylogenetically distinct arrays that we estimate differ by about 0.8% in nucleotide sequence. Genotypes in one clonal array were present in both species, while genotypes in the other array were seen only in Mallards. In terms of the mtDNA “gene tree,” the assayed Mallards exhibit a paraphyletic relationship with respect to Black Ducks, meaning that genealogical separations among some extant haplotypes in the Mallard predate the species separation. Evidence is advanced that this pattern probably resulted from demographically based processes of lineage sorting, rather than recent, secondary introgressive hybridization. However, haplotype frequencies were most similar among conspecific populations, so the Mallard and Black Ducks cluster separately in terms of a population phenogram. The results provide a clear example of the distinction between a gene tree and a population tree, and of the distinction between data analyses that view individuals versus populations as operational taxonomic units (OTUs). Overall, the mtDNA data indicate an extremely close evolutionary relationship between Mallards and Black Ducks, and in conjunction with the geographic distributions suggest that the Black Duck is a recent evolutionary derivative of a more broadly distributed Mallard-Black ancestor.     

Evaluation of eutrophication control in Lake Chivero,Zimbabwe, by multivariate analysis of zooplankton

Zooplankton was a useful tool in evaluation of eutrophication control in Lake Chivero, a tropical man made impoundment. Principal component analysis revealed both spatial and temporal changes in zooplankton community structure which were related to nutrient inflows as well as changes in nutrient content of the lake. Studies on filtration rates of different algae by Bosmina longirostris and Daphnia lumholtzi demonstrated that the response of zooplankton to eutrophication was related to their inability to utilize colonial algal species that develop in a nutrient rich-environment.