The diet of breeding Dippers Cinclus cinclus and their nestlings in the catchment of the River Wye, mid-Wales: a preliminary study by faecal analysis

A preliminary study by faecal analysis into the diet of seven pairs of breeding Dippers and their nestlings was undertaken in the catchment of the River Wye in spring 1983. Macro-invertebrate items in faeces could be identified to family and quantified on the basis of mouthparts whilst the macro-invertebrate prey available to each pair of Dippers was assessed quantitatively by riffle-sampling.
In terms of numerical abundance, ephemeropteran nymphs, particularly Baetidae, and trichopteran larvae, particularly Hydropsychidae, were more important to adults and nestlings, respectively. When prey weights were taken into account, trichopteran larvae were the most important macro-invertebrate group to both, but especially to nestlings.
Some macro-invertebrate groups were selected in proportion to their abundance in the environment although indices of electivity indicated that prey selection for adults and nestlings, respectively, was negatively and positively correlated with prey weight. Significant changes in diet were also apparent with nestling age: smaller items, such as ephemeropteran nymphs, were eaten by younger nestlings whilst larger items, such as trichopteran larvae, became more frequent up to Day 15 after hatching.     

Variable energetic responses to clutch size manipulations in white-throated dippers Cinclus cinclus

We used the doubly-labelled water technique to measure daily energy expenditure (DEE) of a free-living uniparental incubator, the white-throated dipper Cinclus cinclus , in Scotland. DEE was 205±8 (s.e.m.) kJ d⁻¹ for 17 females incubating their natural clutch sizes, equivalent to 3.2±0.1×basal metabolic rate (BMR). To investigate the influence of clutch size on the energy budget, we measured the DEE of 14 females with clutches increased or reduced by a single egg. Birds with reduced clutch sizes had an energy expenditure with a mean and variance that did not differ from those of birds with unmanipulated clutches. Enlarging the clutch led to an increase in energy expenditure to over 4×BMR for some individuals but not for others, resulting in greater variance in energy expenditure for birds with enlarged clutches. Individual variation in energy expenditure could not be fully explained by environmental conditions, by patterns of behaviour or clutch size. Incubating females received a maximum of only 4 kJ d⁻¹ (2% of DEE) from provisioning by the male, and mobilised up to 6 kJ d⁻¹ (3% of DEE) from reserves. Females spent 2.9±0.2 h (n=20) away from the nest each day, so a foraging rate of 95 kJ h⁻¹ was required during incubation recesses to balance DEE. This 'required foraging rate' is double previous estimates of the maximum rates of energy acquisition for birds of this size. We suggest that the greater likelihood of a raised energy expenditure associated with larger clutches, combined with the difficulties in maintaining energy supplies, may constitute a constraint on avian clutch size.     

The distribution of breeding dippers (Cinclus cinclus (L.); Aves) in relation to stream acidity in upland Wales

SUMMARY. 1. The distribution of breeding dippers was assessed in 1984 over seventy-four sites on soft-water streams throughout mid and north Wales.
2. Sites without breeding dippers had significantly higher mean concentrations of filterable aluminium, lower mean pH, fewer trichopteran larvae and ephemeropteran nymphs, and were on rivers with more conifer afforestation on their catchments, than sites where dippers were present.
3. We speculate that influences which increase the acidity of soft-water streams could be detrimental to their populations of dippers, although further data are required on causative mechanisms.     

Foraging behavior predicts age at independence in juvenile Eurasian dippers (Cinclus cinclus)

Timing of foraging independence may be controlled by parents,offspiing, or both and may have consequences for dispersal,reproduction, and survival In a study of juvenile Eurasian dippers(Cinclus cinchus), I examined the relationship between individualdifferences in parental provisioning, the development of foraging,and the timing of independence. Young dippers forage very differentlyfrom adults, specializing on small, stationary pre such as simuliidlarvae (Diptera), and avoiding energetically costly and highlyskilled foraging techniques, such as diving, by which adultsobtain larger prey In my Welsh study area, age at independencevaried substantially within and between broods. Juveniles thatrelied more on simuliids and less on larger prey were independentsooner, as were juveniles with higher rates of intake whileforaging the effect was obvious 3–4 days after fledging.Provisioning rate and total caloric intake over the dependentperiod was unrelated to age at independence, although greaterreliance on parental feeding was associated with higher totalcaloric intake. Higher rates of intake from begging were correlatedwith reduced foraging time. The results demonstrate that theacquisition of the adult foraging strategy is not a prerequisitefor independence or, hence, dispersal. In fact, mastering theskills necessary to capture larger prey may delay independence.Individual juvenile dippers seem to find different solutionsto the trade-off between gaining sufficient food (either frombegging or from foraging for simuliids) and the eventual necessityof learning to forage like an adult.     

Timing of onset of breeding in three different Dipper Cinclus cinclus populations in France

Capsule Timing of breeding varies widely.     

Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

Capsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.

Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.

Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.

Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.

Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.     

The ecology of dippers Cinclus cinclus (L.) in relation to stream acidity in upland Wales: time-activity budgets and energy expenditure

Summary The time-activity budget and energy expenditure of a riverine bird, the dipper Cinclus cinclus, was studied from March 1988 to July 1989, across a range of streams of contrasting acidity in upland Wales. Differences in time-activity budgets of birds on acidic and circumneutral streams were consistent with documented differences in prey availability and diet. Birds spent a significantly greater proportion of their active day foraging, swimming and flying, and less time resting, on acidic streams. Activity measurements varied significantly through the year, through the day, and with river flow. Despite differences in time budgets, mean Daily Energy Expenditure (DEE) on acidic streams was only 4.5–7.0% greater than on circumneutral streams. However, rates of energy gain were greater for dippers on circumneutral streams for every month of the year, a pattern confirmed by differences in body condition. By spending more time feeding, dippers on acidic streams will have less time for other activities such as self-maintenance and predator surveillance; they may also be less able to meet the additional demands accompanying the initiation of breeding. These findings are discussed in relation to the feeding ecology and breeding performance of dippers on streams of contrasting water quality in upland Wales.     

The distribution of dippers, Cinclus cinclus (L.), in the acid‐sensitive region of Wales 1984–95

1. Dippers were surveyed along seventy-four acid-sensitive streams in upland Wales in 1984 and again in 1995. At forty-eight of the sites, and in an additional twenty-three to twenty-nine adjacent catchments, changes in acid-base status and macroinvertebrate communities were assessed. River habitat surveys (RHS) and GIS (Geographical Information System) provided information on habitat features. 2. pH means across all the sites were higher on average by 0.12 pH units in 1995 than 1984, accompanied by significantly increased total hardness and reduced sulphate. However, geometric mean aluminium concentrations increased significantly, while the abundances of important dipper prey either remained constant (Ephemeroptera) or declined (Trichoptera; Plecoptera). 3. As in 1984, dipper distribution in 1995 was related to acid-base status: aluminium concentrations were significantly higher, and pH significantly lower, at sites where dippers were absent. In both surveys, there were significantly more bankside broadleaves and fewer conifers where dippers were present. 4. Although dippers occurred in 1995 on a similar number of streams as in 1984, there were both gains and losses, and an overall significant reduction in the number of visits on which birds were recorded (= registrations) per survey reach. Gains and losses were not related to habitat structure or acid-base status, and might be stochastic. Mean aluminium concentrations increased more (P < 0.06) at sites where dipper registrations fell, than where they increased, but changes were large enough to explain altered occupancy at only five sites. Plecopteran abundances declined most at sites losing birds. 5. We conclude that recovery from acidification has not yet been large or sustained enough to allow widescale increase in the Welsh dipper population, and continued decline cannot be excluded. There is a need for better understanding of how recovery processes will permeate foodwebs to reach top predators in acidified streams, and of the geographical scale of recovery required to increase populations in dispersed organisms such as birds. Such uncertainties, together with the mismatch in trends revealed by stream chemistry and dippers, illustrate reasons why chemical data alone are inadequate for the assessment of changing river quality.     

Energetics of the annual cycle of Dippers Cinclus cinclus

Time-activity budgets and energy expenditure of Dippers Cinclus cinclus were studied in all months of the year and for every stage of the annual cycle. The commonest daytime activity was feeding (54%), then resting (43%) and flying (4%). On a 24-hr day basis the most marked changes in activity followed from changing daylengths. DEE (Daily Energy Expenditure), derived from time-activity budgets through the year and laboratory estimates of metabolism, averaged 201 kj d ^-1 in females and 228 kj d ^-1 in the larger males. Over a more restricted range of circumstances, direct estimates of DEE obtained from 77 Dippers using the doubly labelled water technique averaged 205 ± 43 kj d^-1 and 251 ± 55 kj d^-1 in females and males, respectively. Overall, the correspondence between these largely independent estimates of energy expenditure was reasonably close. DEE was highest during breeding (laying-females; rearing-males) and in late winter for both sexes. The lowest energy expenditures occurred during moult, amongst juveniles and in early winter. Incubating females and mate-guarding males also had low energy costs. Across all stages of the annual cycle body size, activity patterns, ambient temperature and river flow had significant effects on energy expenditure. The rate at which food was gathered to meet these changing energy demands varied widely. While some of this variation was imposed by a seasonal environment, it was also likely to reflect adaptive shifts in rates of food gathering, in some cases consequent upon the changing fitness benefits of various non-feeding activities.     

Factors influencing the abundance of breeding Dippers Cinclus cinclus in the catchment of the River Wye, mid-Wales

The number of breeding Dippers was assessed in relation to environmental features and benthic macro-invertebrates over 220 km of tributaries in or adjacent to the catchment of the Welsh River Wye in 1982. Bivariate regression and multivariate analysis revealed strong correlations between breeding abundance and combinations of variables describing stream-gradient and the relative abundances of larval Trichoptera and nymphal Plecoptera. The results are discussed in relation to aspects of the feeding ecology of breeding Dippers and their nestlings.     

Aspects of the breeding biology of Dippers Cinclus cinclus in the southern catchment of the River Wye,Wales

Laying dates, clutch and brood size, and the incidence of double brooding, in Dippers breeding in the southern catchment of the Welsh River Wye between 1978 and 1983 are described and compared with data from other regions of Britain and mainland Europe. The mean size of 222 clutches (4.78±0.08) was found to be significantly larger than values given by all other studies in Britain where data were sufficient for statistical comparison. This is discussed in relation to territory quality and other influences.     

Song function and territoriality in male and female White-throated Dippers Cinclus cinclus

Capsule: Male White-throated Dippers Cinclus cinclus are more likely and quicker to respond to the playback of song than females, but both sexes are more likely to respond before the onset of breeding than after.

Aims: Territoriality and the function of song in female birds have rarely been studied outside of the tropics or Australasia. We investigated territoriality and song function in males and females of a Northern temperate species, the White-throated Dipper.

Methods: We conducted playback trials on established pairs and compared the responses of males and females according to the sex of the simulated intruder and the timing of playback relative to the onset of breeding. A response was classified as movement towards the speaker, singing or both.

Results: Males were significantly more likely and quicker to respond to playback than females, but neither sex responded differently to the playback of male and female song. Both sexes were more likely to respond to playback before breeding had begun than after.

Conclusions: Our results suggest that both males and females are territorial but that males take the dominant role in defence. Female song appears to elicit a similar response to male song and may play a role in territoriality or mate defence.     

Molecular phylogenetics and the historical biogeography of dippers (Cinclus)

I used combined sequences of mitochondrial cytochrome  b and ND2 genes to determine the molecular phylogenetic relationships of all five extant species of dipper ( Cinclus ), as well as the relationships of Cinclidae to postulated nearest relatives. All methods of analysis resulted in a single best tree of dipper relationships, uniting the two South American taxa (as sisters) with the single North American exemplar, and the two Eurasian taxa forming a sister clade to the New World taxa. Further, each tree identified thrushes (Turdidae) as the closest relative to Cinclidae. Based on relationships within Cinclus , a Eurasian ancestral area is proposed, with subsequent movement into the New World. Dating of species divergences suggest that dippers arose approximately 4 mya, and achieved their present continental distributions soon after.     

Climate effects on population fluctuations of the white-throated dipper Cinclus cinclus

1. Climate change may have profound consequences for many organisms. We have studied fluctuations in a population of the white-throated dipper Cinclus cinclus during 31 years (1978-2008) in a river system in southern Norway in relation to both large-scale and local weather conditions occurring during the non-breeding season. 2. Multiple regression and partial least squares regression were used to model the growth rate of the population, accounting for population size in the previous year. 3. Population growth was influenced by North Atlantic Oscillation (NAO), mean winter temperature, precipitation and timing of ice formation on the main lake in the river system in autumn. These variables explained 84% of the variation in population growth over the 31-year study period. 4. Local winter conditions played a prominent role in explaining the population fluctuations, which is plausible because the dipper depends on open water for foraging. In the study area, winters can be harsh and rivers and lakes may freeze and severely affect the subsequent population size of the dipper in spring. 5. The breeding population of the dipper does not seem yet to have reached a level where all possible territories in the area have been occupied, even after mild winters, and the estimated carrying capacity is also decidedly lower (66 breeding pairs) than the number of available territories. If the trend of milder winters continues, the population might increase in the future. However, strong climate variation is expected to continue in the future, and hence periods of rapid growth of the dipper population will probably be followed by severe declines.     

Prevalence of haematozoan parasites in the White-throated Dipper Cinclus cinclus in southern Europe

Capsule White-throated Dippers from southern Europe were found to be infected by three haemosporidians.

Aims To examine the occurrence of blood parasites in dippers in the Iberian Peninsula and to investigate the relationship between prevalence levels, environmental factors and bird fitness.

Methods White-throated Dippers were trapped with mist-nets from five montane areas in the centre of the Iberian Peninsula. Parasites were detected from blood samples by polymerase chain reaction screening.

Results About half (51.3%) of 152 dippers showed some kind of infection. The genus Haemoproteus was recorded in 49.3% of the birds, and the genus Leucocytozoon in 19.1%, while Plasmodium was present only in 0.7%. Among the infected birds, 34.6% carried a double infection (Haemoproteus?+?Leucocytozoon). Prevalence did not differ between gender or age-classes, but it varied between study sites, being significantly higher in mountains with higher precipitation. There was a reduction in body mass owing to double infection in yearling males only.

Conclusion This is the first work assessing parasite prevalence in dippers from southern Europe. Infection of dippers by haemosporidians is likely to be related to a study site's climatic envelope.     

The adaptive significance of brood size and time of breeding in the dipper Cinclus cinclus (Aves: Passeriformes) as seen from post-fledging survival

We assessed the post-fledging survival of dippers Cinclus cinclus from 743 broods in relation to brood size, time of hatching and territory quality. We paid particular attention to assessing whether contrasting breeding performance along unproductive (i.e. acidic) and productive (i.e. circumneutral) rivers represented strategies which optimized the number of surviving young.
For all brood sizes, post-fledging survival varied significantly through the breeding season, with most survivors coming from attempts in the peak period of hatching. After correcting for these seasonal effects, the most common brood size overall, of four, was also the most productive as seen from post-fledging survival; differences in the frequency of occurrence and survival between broods of four and five were marginal. Moreover, a change in the modal brood size from five to four occurred as the season progressed. consistent with a shift in brood productivity.
Broods at acidic sites were significantly smaller than at circumneutral sites; while brood size four was the most productive at both types of site, brood size three was the second most productive at acidic sites, while brood size five was the second most productive at circumneutral sites. Dippers at acidic sites bred significantly later than at circumneutral sites, but post-fledging survival declined most rapidly through the season at the former.
These survival data provide evidence from both seasonal and spatial patterns that brood sizes in the dipper may be optimized in ways consistent with the enhancement of productivity. By contrast, delayed breeding at acidic sites contrasted with the patterns expected from optimization, instead reflecting resource scarcity.     

Extrapair paternity and genetic diversity: the white‐throated dipper Cinclus cinclus

Passerine bird species vary considerably in the frequency of extrapair paternity, but the factors causing this variation are not well understood. There is some comparative evidence that extrapair paternity is associated with the population level of genetic diversity, but there is no consensus of how genetic diversity should be measured and compared across species or populations. Here we report a low frequency of extrapair paternity (2% extrapair offspring) in a Norwegian population of the white‐throated dipper Cinclus cinclus, which shows strong signs of reduced genetic diversity. We encountered difficulties in constructing a robust parentage analysis system for the species, largely due to consistently low polymorphism levels in 100 heterologous microsatellite markers. Furthermore, single‐nucleotide polymorphisms (SNPs) were almost absent in intron sequences in 10 nuclear genes (>5 kb) that are much more polymorphic in other species. Hence, our results seem consistent with the genetic diversity hypothesis that predicts a low frequency of extrapair paternity in species with low genetic diversity. Heterologous microsatellite markers are generally unsuitable for interspecific comparisons of genetic diversity as they show strong phylogenetic dependency in polymorphism levels. We suggest that SNP rates at homologous nuclear introns, like those presented here, can provide a useful method for obtaining unbiased estimates of genome‐wide genetic diversity across populations and species.     

The potential influence of Atlantic salmon Salmo salar and brown trout Salmo trutta on density and breeding of the white‐throated dipper Cinclus cinclus

Interactions between birds and fish are often overlooked in aquatic ecosystems. We studied the influence of Atlantic salmon and brown trout on the breeding population size and reproductive output of the white‐throated dipper in a Norwegian river. Acidic precipitation led to the extinction of salmon, but salmon recolonized after liming was initiated in 1991. We compared the dipper population size and reproductive output before (1978–1992) and after (1993–2014) salmon recolonization. Despite a rapid and substantial increase in juvenile salmon, the breeding dipper population size and reproductive output were not influenced by juvenile salmon, trout, or total salmonid density. This might be due to different feeding strategies in salmonids and dippers, where salmonids are mainly feeding on drift, while the dipper is a benthic feeder. The correlation between the size of the dipper population upstream and downstream of a salmonid migratory barrier was similar before and after recolonization, indicating that the downstream territories were not less attractive after the recolonization of salmon. Upstream dipper breeding success rates declined before the recolonization event and increased after, indicating improved water quality due to liming, and increasing invertebrate prey abundances and biodiversity. Surprisingly, upstream the migratory barrier, juvenile trout had a weak positive effect on the dipper population size, indicating that dippers may prey upon small trout. It is possible that wider downstream reaches might have higher abundances of alternative food, rending juvenile trout unimportant as prey. Abiotic factors such as winter temperatures and acidic precipitation with subsequent liming, potentially mediated by prey abundance, seem to play the most important role in the life history of the dipper.     

Response of American dippers (Cinclus mexicanus) to variation in stream water quality

1. Semi‐aquatic birds may be sensitive to altered water quality. While avian species are not used in the bioassessment of streams, they may complement the more common use of benthic macroinvertebrates and fish. We estimated the extent to which water quality can predict attributes of the populations of one common semi‐aquatic bird, the American dipper (Cinclus mexicanus). 2. First, we estimated dipper presence/absence in relation to water quality as measured by a multimetric assessment index and individual bioassessment metrics. Second, we estimated dipper territory area and reproductive success in response to variation in water quality. We studied the diet, territory area and fecundity of dippers and sampled benthic macroinvertebrates, water chemistry and physical variables at 32 sites with and 17 sites without nesting dippers. 3. Dipper presence was only weakly related to chemical, physical and commonly recorded bioassessment metrics such as per cent Ephemeroptera, Plecoptera and Trichoptera (%EPT). Dippers were strongly related to the abundance of their common prey, Drunella and Heptageniidae, which are only a small component of the commonly recorded bioassessment metrics. The variances in territory area and reproductive success were weakly predicted by water quality variables. 4. Dipper presence reflected disturbance as measured by their common prey, showing that lower abundance of these stream invertebrates affected this semi‐aquatic bird. We suggest dipper presence/absence might be used in multimetric indices of biotic integrity for the bioassessment of streams.