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.     

Breeding density of Dippers Cinclus cinclus' Grey Wagtails Motacilla cinerea and Common Sandpipers Actitis hypoleucos in relation to the acidity of streams in south-west Scotland

The distribution of breeding pairs of Dippers Cinclus cinclus , Grey Wagtails Motacilla cinerea and Common Sandpipers Actitis hypoleucos was assessed along 18 streams in south-west Scotland during summer 1987, and the lengths of territories, occupied by 5 5 breeding pairs of Dippers, were measured. In each stream the water chemistry, physical structure (e.g. gradient, altitude, nature of bankside vegetation) and density of aquatic invertebrates was determined. The density of breeding pairs of dippers was significantly lower along streams of low pH (high acidity) and steep gradients, and territories were significantly longer at these sites compared with those of higher pH. Many of the invertebrate families that are important prey for Dippers were scarce at sites of low pH, particularly caddisfly larvae (Trichoptera) and mayfly nymphs (Ephemeroptera). The low numbers and large territories of breeding pairs of Dippers on acidic streams may be due to reduced density and diversity of macroinvertebrate prey. Stream pH was not an important correlate of the distribution or density of Grey Wagtails or Common Sandpipers. It is suggested that the lack of any similar effect of pH on these other two species compared with Dippers is attributable to differences in their feeding ecology.     

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.     

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.     

Organochlorines and mercury in the eggs of British Peregrines Falco peregrinus

The levels of various contaminants were measured in 550 addled and deserted Peregrine Falco peregrinus eggs obtained in Britain during 1963-86. In this period the population was recovering from a low level imposed by organochlorine pesticides. Over the whole period, HEOD levels declined in eggs from both inland and coastal regions, while DDE levels declined in eggs from inland regions. At the same time, shell-indices improved. PCB levels increased in eggs from inland regions. At any one time, levels of DDE and HEOD in eggs decreased and shell-indices increased from south to north within Britain. These gradients fitted with the extent of agricultural land (= pesticide use) and with the extent of Peregrine population decline (both greatest in the south). No south-north trend was apparent in levels of PCBs derived from industrial sources. In some regions eggs from coastal sites were more contaminated than eggs from inland sites, especially with PCBs and mercury. Significant relationships were found between brood sizes and DDE levels, between brood sizes and shell-indices, and between shell-indices and DDE levels. This implied that DDE influenced shell thickness and breeding success. Some evidence was obtained that mercury reduced brood sizes, but no evidence that HEOD and PCB (at the levels found) did so. Overall, DDE and mercury levels together accounted for 17% of the variance in brood sizes during 1963-86. On a regional basis, DDE had no obvious effect on mean productivity when the geometric mean DDE level in collected eggs was less than 3 p.p.m., and when mean shell-index was no more than 8% less than normal. At higher DDE levels, and lower shell-indices, productivity declined markedly. Recovery of regional populations was associated with geometric mean HEOD levels in eggs no greater than 0–7 p.p.m., DDE levels no greater than 15 p.p.m., shell indices no more than about 15–20% below normal, and a mean breeding success exceeding 0–6 young per territorial pair.     

Do American Dippers Obtain a Survival Benefit from Altitudinal Migration?

Studies of partial migrants provide an opportunity to assess the cost and benefits of migration. Previous work has demonstrated that sedentary American dippers (residents) have higher annual productivity than altitudinal migrants that move to higher elevations to breed. Here we use a ten-year (30 period) mark-recapture dataset to evaluate whether migrants offset their lower productivity with higher survival during the migration-breeding period when they occupy different habitat, or early and late-winter periods when they coexist with residents. Mark-recapture models provide no evidence that apparent monthly survival of migrants is higher than that of residents at any time of the year. The best-supported model suggests that monthly survival is higher in the migration-breeding period than winter periods. Another well-supported model suggested that residency conferred a survival benefit, and annual apparent survival (calculated from model weighted monthly apparent survival estimates using the Delta method) of residents (0.511 ± 0.038SE) was slightly higher than that of migrants (0.487 ± 0.032). Winter survival of American dippers was influenced by environmental conditions; monthly apparent survival increased as maximum daily flow rates increased and declined as winter temperatures became colder. However, we found no evidence that environmental conditions altered differences in winter survival of residents and migrants. Since migratory American dippers have lower productivity and slightly lower survival than residents our data suggests that partial migration is likely an outcome of competition for limited nest sites at low elevations, with less competitive individuals being forced to migrate to higher elevations in order to breed.     

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.     

The diet of breeding dippers Cinclus cinclus cinclus and their nestlings in southwestern Norway

The diet of dippers in south west Norway was investigated by faecal analysis during the 1985 breeding season. Overall, the diets of adults and nestlings were similar and consisted mostly of nymphs and larvae of Baetidae (mayflies). Leuctridae/Nemouridae (stoneflies). Hydropsychidae, Limnephilidae and Rhyaeophilidae (caddisflies). However, strong trends were apparent with nestling age: large items such as caddisflies declined in adult diet but increased in nestling diet as the chicks grew. The pattern probably reflected the growing energy demands of the brood in a central-place. Because some of the items of dietary importance are scarce in acidic streams, we stress the need for research on the influences of acidification on the ecology of Scandinavian dippers.     

Relationships between stream acidity and bacteria,macroinvertebrates, and fish: a comparison of north temperate and south temperate mountain streams,USA

A comparative study of relationships between stream acidity and bacteria, macroinvertebrates, and fish in the Adirondack Mountains of upper New York state and in the Southern Blue Ridge Mountains of eastern Tennessee, USA, was conducted. Although the study sites in both regions spanned a pH range from approximately 4.5 to 6.4, considerably greater seasonal variability in pH and higher monomeric Al concentrations characterized the Adirondack sites. Relationships between several biological characteristics and stream water acidity were similar in both regions, including lower production of epilithic bacteria and bacteria on decomposing leaves, lower leaf decomposition rates, lower density and generic richness of scraper/grazer macroinvertebrates, particularly Ephemeroptera, and lower fish abundance and survival in more acidic streams. Densities of total macroinvertebrates and densities of macroinvertebrates and bacteria inhabiting or closely associated with stream sediments were generally not related to stream water acidity.Regional differences occur in some of the relationships between biological characteristics and stream water acidity. Negative correlations between bacterial production on rocks and pH, between bacterial production on decomposing leaves and pH, and between densities of Ephemeroptera and scrapers and pH were stronger in the Adirondacks than in the Southern Blue Ridge. Higher Al concentrations in the Adirondacks may be responsible for the stronger relationships with pH there. The steeper slopes of the relationships between Ephemeroptera density and all forms of Al in the Adirondacks compared with the Southern Blue Ridge suggests that there may be some adaptation among a few acid/aluminum-tolerant species in the seasonally more constant acidic Southern Blue Ridge streams. Fish bioassays indicated longer survival times in acidic streams in the Adirondacks compared with the Southern Blue Ridge, but these results may be an artifact associated with the use in the Southern Blue Ridge of rainbow trout as the test species which is known to be more acid sensitive compared with brook trout, the test species used in the Adirondacks.     

DOES PREY BIOMASS OR MERCURY EXPOSURE AFFECT LOON CHICK SURVIVAL IN WISCONSIN?

Abstract: Past studies suggest that the productivity of common loons (Gavia immer) is lower on acidic lakes in northern Wisconsin, USA, than on neutral lakes. Two hypotheses have been proposed to explain low chick survival: (1) reduced food consumption related to changes in prey communities on lower pH lakes and (2) high mercury (Hg) exposure on lower pH lakes. To address these hypotheses, we quantified prey and Hg consumption by loon chicks on 51 lakes and survival on 55 lakes ranging in pH from 4.9 to 9.5 in northern Wisconsin in 1995 and 1996. The time adults spent providing prey to chicks was unrelated to lake pH but increased with number of chicks and chick age. The number of prey caught per provisioning time declined as lake pH declined because adults made fewer dives, not because success of prey capture declined. Chicks consumed more insect larvae on acidic lakes and more crayfish (Family Astacidae) on neutral lakes. Biomass consumed ranged from an average 1.99 ± 1.05 (SE) g/hr/chick during the first week of a chick's life to a peak of 7.93 ± 1.93 g/hr/chick during the eighth week. Biomass intake per chick body weight (g/W_g/hr) declined with lake acidity but was not related to chick survival (P = 0.25). Although the Hg concentration in the 3 major prey species was positively related to lake acidity and blood Hg level of chicks at a lake, total Hg consumption (μg/W_g/hr) was highest on moderately acidic lakes rather than on the most acidic lakes. We suggest that loon chick survival in northern Wisconsin lakes is more likely related to prey availability than to Hg exposure. When we removed from our analysis 1 lake where 2 11-day-old chicks were killed by predators, chick survival was negatively related to lake acidity but not to biomass or Hg consumption. We discuss mechanisms of Hg excretion that may allow young chicks to survive on acidic lakes in northern Wisconsin despite high Hg intake.     

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.     

Productivity in a social wasp: per capita output increases with swarm size

We measured the productivity of newly-founded colonies of Polybiaocddentalis, a Neotropical swarm-founding social wasp, overtheir first 25 days. By both of the measures we used, numberof nest cells built by the swarm and dry weight of brood produced,colony-level productivity was a significant positive quadraticfunction of the number of adults in the swarm, indicating thatper capita output increased with swarm size. Subdividing adultsinto queens and workers did not improve significantly on thesemodels, but the proportion of queens was a significant factorexplaining brood production in one of two sampling years. Earlierwork on P. ocddentalis suggests that the mechanism behind thepattern is that workers transferring materials to one anotherexperience increasing queuing delays as group size decreases.The largest colony in each of the two years produced unusuallylow outputs of brood. One interpretation is that the curve ofgroup-size related brood productivity peaks at intermediategroup size and that these colonies are on the downward partof the curve. That these same two colonies also had the lowestproportions of queens suggests a second interpretation: thesecolonies were constrained to low brood production by a low colony-leveloviposition rate. A third possibility is diat these were maturecolonies, and mature colonies may allocate a smaller fractionof resources to brood rearing than do younger colonies. Ourresult contradicts earlier findings for a variety of socialand subsocial Hymenoptera that per capita productivity declinesas group size increases. We suspect that Michener's result forswarm-founding wasps is an artifact of his having to lump coloniesof different species and different stages of development toobtain adequate sample sizes to plot. If our result for P. ocddentaliscan be generalized to other swarm-founders, then these waspshave evolved a mode of colony organization fundamentally differentfrom that of other wasps. Thus, our result places new significanceon the role of group dynamics as a factor affecting group sizein different taxa.     

Offspring growth, survival and reproductive success in the bank vole: a litter size manipulation experiment

To estimate the optimality of brood size, it is essential to study the effects of brood size manipulation on offspring survival and reproductive success. Moreover, testing the generality of the hypothesis of reproductive costs requires experimental data from a diversity of organisms. Here I present data on the growth, survival and reproductive success of bank vole Clethrionomys glareolus individuals from manipulated litters. Furthermore, the survival of mothers whose litter size was manipulated was studied. At weaning, the mean weight of pups from enlarged litters was lower and from reduced litters higher compared to control litters. After winter, at the start of the breeding season, individuals from enlarged litters, especially males, were still lighter than individuals from the other two treatments. Litter enlargements did not increase the number of reproducing female offspring per mother, nor did the litter sizes of female offspring differ between treatments. There were no differences between treatments in winter survival of offspring after weaning, but among female offspring, weaning weight explained the survival probabilities over winter. A higher weight of females at winter determined the probability of starting to reproduce in spring. The survival of mothers did not seem to be influenced by litter manipulation performed the previous year. According to the results, mothers nursing enlarged or reduced litters do not gain any fitness benefits in terms of number of offspring surviving to breeding. The results are consistent with the majority of experiments conducted in birds, which have found costs of enlarged brood appearing as offspring trade-offs rather than parent trade-offs. Received: 14 December 1997 / Accepted: 1 March 1998     

Urban flight: understanding individual and population-level responses of Nearctic-Neotropical migratory birds to urbanization

1. Despite the fact that studies of urban ecology have become commonplace in the literature, ecologists still lack empirical evidence of the underlying mechanisms responsible for relationships between urbanization and animal community structure. In an effort to understand the processes that govern an apparent avoidance of urban landscapes by many Nearctic-Neotropical migratory birds, we examined population- and individual-level responses of the Acadian flycatcher (Empidonax virescens) to urbanization within the landscapes surrounding 35 riparian forest stands in Ohio, USA. 2. From May to August 2001-06 we surveyed birds, banded 175 territorial flycatchers to estimate both condition and survival, tracked nest initiation dates, monitored success of 387 nests and estimated annual reproductive productivity of 163 breeding pairs. 3. Neither apparent annual survival of males (phi = 0.53 +/- 0.056 SE) nor females (phi = 0.23 +/- 0.064 SE) was related to the amount of urban development within the landscape. Similarly, daily survival rates of nests, which ranged from 0.92 to 0.98 across sites, was not associated significantly with urbanization. In contrast, reproductive productivity was related negatively to the amount of urbanization surrounding riparian forests, perhaps due in part to the greater incidence of brood parasitism and fewer numbers of nesting attempts made by pairs in urban compared to rural forests. 4. Forests within urban landscapes experienced higher levels of turnover in site occupancy, and birds settling in urban areas initiated nests later, had marginally smaller body sizes and exhibited lower return rates following nest predation than birds in more rural landscapes. In this way, behavioural processes governing habitat selection, settlement patterns and site fidelity probably contributed to the lower levels of reproductive productivity achieved by pairs nesting in urban landscapes. 5. This study provides evidence that the negative association between Acadian flycatchers and urbanization results from both population- and individual-level responses to urbanizing landscapes surrounding their riparian forest habitats.     

Factors affecting the probability of double brooding by Southern House Wrens

Seasonal fecundity of birds is influenced by clutch sizes and the number of successful breeding attempts during a breeding season. As such, understanding the factors that determine the decision to initiate multiple broods within a season and the consequences of this reproductive tactic is important. We examined the frequency of double brooding by Southern House Wrens (Troglodytes aedon musculus) in eastern Argentina. We analyzed inter‐ and intraseasonal variation in double brooding and evaluated the effect of weather conditions and laying date on the frequency and occurrence of this behavior. Finally, we assessed the effect of double brooding on the seasonal and lifetime productivity of female Southern House Wrens. During our 8‐year study, we found that ～43% (range = 17–83% each year) of breeding pairs attempted a second brood after successfully raising a first brood. The probability of females having a second brood was affected by the laying date of the first nesting attempt, but was independent of the number of young fledged. About 65% of females that started laying eggs before the first quarter of each breeding season produced a second brood, and this percentage decreased to ～40% after this period. In addition, variation in double‐brooding frequency among years was related to weather conditions, with the proportion of pairs double brooding increasing with increased precipitation early in the breeding season. More precipitation likely contributed to an increase in insect abundance. Although double brooding increased the seasonal and lifetime productivity of female Southern House Wrens, additional study of the survival and fate of fledglings from first and second broods is needed to assess the importance of multi‐brooding in the reproductive success of these wrens.     

Parental care trade‐offs in the inter‐brood phase in Barn Swallows Hirundo rustica

In seasonal environments with limited time and energy resources, double‐brooded birds face trade‐offs in the timing of their two reproductive attempts and in the effort allocated to the first and the second broods. In the Barn Swallow Hirundo rustica a long care period for the first brood enhances the survival of first‐brood chicks, but also delays the start of the second brood, which in turn reduces the survival prospects of second‐brood chicks. Probably as a response to this trade‐off, double‐brooded Barn Swallows reduce the period of post‐fledging care for first‐brood fledglings. By radiotracking whole families, we investigated the determinants of this behaviour and its consequences for the survival of the first‐brood fledglings. The end of the females’ investment in post‐fledging care of the first brood was related to the beginning of egg synthesis for the second clutch. With the start of egg synthesis, females significantly reduced provisioning rates to the first‐brood fledglings to less than one‐fifth of the previous rates, while the proportion of time they spent foraging remained high. Assuming that the females’ foraging success was constant, we conclude that their energy income was allocated to egg production rather than fledgling provision. Males did not compensate for the females’ reduced feeding rates. Thus the start of egg production for the second clutch had a marked effect on the quantity of food received by first‐brood fledglings. In parallel with the changes in parental behaviour and provisioning rates, we observed a marked drop in the daily survival rate of first‐brood chicks. These results support the hypothesis that females face a strong trade‐off in the allocation of energy to subsequent broods. Energy allocation to a second clutch involves a cost in terms of reduced provisioning, and as a result the survival of first‐brood chicks is compromised. This is probably outweighed by the improved success of an early second brood.     

Production and costs of nesting attempts in Snow Buntings Plectrophenax nivalis: why do they attempt second broods?

During 1988–1993, pairs of Snow Buntings Plectrophenax nivalis on our study sites in northeast Scotland reared an average of 1.2 broods. Clutch sizes of first and second broods were similar, but partial losses were greater in second broods, leading to a difference of at least 40% in overall nest productivity between successful first and second broods. Over and above this, total nest failure was four times higher in the second broods, and autumn sightings of ringed nestlings from second broods were only a third of those of first-brood nestlings. As a result, second broods produced a mere 10% of future recruits to the breeding population. However, there was also little evidence of costs associated with producing, or attempting to produce, second broods. Young from first broods were not less likely to reach independence if a second brood was attempted, between-year return rates of double-brooded adults were similar to those of single-brooded birds and double-brooded adults did not appear to be less fecund in the following summer. Therefore low costs to the adults of attempting second broods may allow the continuation of a strategy which appears to have only a marginal influence on overall reproductive success in the area studied.     

Field experiments on the effect of ticks on breeding success and chick health of cattle egrets

Abstract Two experiments compared broods that were naturally tick-infested with an equal number that were rendered tick-free by application of an acaricide to their nests and of barriers against further infestation. In the first experiment conducted in 1991–92 nestlings in tick-infested broods had up to 159 larval ticks at once and a mean infestation of 23.6 larval ticks per chick per day. The chick's mean tick load was inversely correlated with its longevity. Chicks that survived to at least 18 days post-hatching had significantly lower larval tick loads than those that died by 18 days, excluding the third-hatched chick in each brood whose survival rate was low irrespective of tick-infestation. At 7 days post-hatching, tick-infested chicks had a lower haematocrit and higher polychromasia than tick-free chicks. I infer that blood loss anaemia caused the deaths of tick-infested chicks in their first week. None died in their second week and the demise of those in their third week may have been due to the paralysis manifested prior to their death. I conclude that heavy tick infestation of their chicks reduced the breeding success of the parent egrets below that of the egrets whose chicks were kept free of ticks. In the second experiment in the 1992–93 season the mean level of tick infestation was much lower (5.1 per chick) and these chicks survived equally with tick-free chicks through fledging.     

Growth and population size of crayfish in headwater streams: individual- and higher-level consequences of acidification

1. Environmental stress may have indirect positive effects on population size through modification of food‐web interactions, despite having negative effects on individuals. Here we evaluate the individual‐ and population‐level effects of acidification on crayfish (Cambarus bartonii) in headwater streams of the Allegheny Plateau (PA, U.S.A.) with field experiments and survey data. Median baseflow pH of 24 study reaches in nine streams varied from 4.4 to 7.4, with substantial variation found both among and within streams. 2. Two bioassays were conducted to evaluate the relationship between stream pH and crayfish growth rates. Growth rates were always higher in circumneutral reaches than in acidic reaches. Crayfish originating in acidic water grew less when transplanted into neutral water than did crayfish originating in neutral water, providing some evidence for a cost of acclimation to acidity. 3. Stream surveys showed that fish were less abundant and crayfish more abundant in acidified streams than in circumneutral streams. Crayfish density was sixfold higher in reaches with the lowest pH relative to circumneutral reaches. Large crayfish made up a higher proportion of crayfish populations at sites with high fish biomass, consistent with the hypothesis that fish predation on small individuals may be limiting crayfish population size at these sites. 4. Although individual crayfish suffered lower growth in acidified streams, increased acidity appeared to cause an increase in crayfish population size and shifts in size structure, possibly by relieving predation pressure by fish.     

Fish species distribution in relation to water quality gradients in the North Branch of the Moose River Basin

The distribution of fish species in the North Branch of the Moose River (Lake Rondaxe to headwaters of Big Moose Lake) was determined by intensive netting and electrofishing surveys of lakes and streams in the watershed during 1982–83. A chronology of changes in fish species occurrence in the drainage system was reconstructed from earlier published surveys conducted in 1882 and 1931 and unpublished survey data obtained by the NYSDEC during the period 1948–1975. Native species present in 1882 were also collected in 1931. Smallmouth bass (Micropterus dolomieu) were introduced in the early 1900's and were present in collections made in 1931. Major changes in the fish community have taken place since 1931. The smallmouth bass and many of the native species found in the earlier surveys were either absent or restricted in occurrence to downstream sites (eg. L. Rondaxe and Moss L. sub-drainage) in 1982. Non-native species introduced after 1931 (yellow perch,Perca flavescens; central mudminnow,Umbra limi; banded killifish,Fundulus diaphanus) are currently widely distributed throughout the drainage system. In particular, the yellow perch is now a dominant species in the larger lakes of the basin. Comparisons of survival rates for caged fish transferred from high to low pH sites in the Big Moose drainage system demonstrated relatively greater acid tolerane of non-native species (yellow perch, mudminnow, killifish) than native cyprinids. Watershed acidity gradients (pH and aluminum concentrations) and relative physiological acid tolerance are major determinants of currently observed fish species distribution patterns in the North Branch of the Moose River. Differences in age and size structure of fish populations inhabiting acidic and non-acidic lakes of the drainage system were apparent, but difficult to interpret without additional information onpopulation size and potential density dependent parameters such as age specific growth and survival rates. Differential hatching success was observed for yellow perch eggs reciprocally transferred between acid (Big Moose) and neutral (Moss L.) lakes. Eggs transferred from Moss L. to Big Moose L. exhibited poor hatching success as a result of alterations in egg membrane structure that inhibited normal egg expansion and the hatching process. This effect was not evident in eggs from the same parents reared in Moss Lake nor in eggs from the Big Moose parents reared in both lakes. These experimental observations suggested possible genetic adaptation to acid stress by the yellow perch population inhabiting Big Moose Lake.