Breeding territory fidelity in a partial migrant, the American dipper Cinclus mexicanus

American dipper Cinclus mexicanus populations are frequently composed of resident individuals that occupy permanent territories year round and migratory individuals that overwinter with residents but migrate to breeding territories on higher elevation creeks each spring. Between 1999 and 2004 we examined how migratory strategy (resident/migratory) and sex differences influence breeding territory fidelity of American dippers occupying the Chilliwack River watershed, British Columbia, Canada. Counter to expectation we found that the migratory strategy of American dippers did not influence whether birds breeding in one year were found on their former breeding territory in the next. Migratory strategy also did not affect the probability that known surviving dippers occupied the same breeding territory in the following year. Males and females were equally likely to be found on their former territory in the following year (females 43%, males 41%) and known survivors had similar levels of breeding territory fidelity (females 74%, males 68%). However, breeding territory fidelity of males and females varied in response to different factors. Surviving female dippers were more likely to be found on their former breeding territory in the subsequent year following a successful breeding attempt than an unsuccessful breeding attempt. Prior reproductive performance did not influence whether surviving male dippers were found on their former breeding territory. Male dippers were more likely to be found on their former territory and, if they survived, have higher breeding territory fidelity when their mate also returned to that same territory. Mate retention also influenced whether females were found on their former territory in the following year but had no effect on the breeding territory fidelity of known survivors. We argue that sex‐specific dispersal decision rules in American dippers are driven by sex differences in the predictability of breeding performance between years and sex differences in how mate retention influences subsequent reproductive success.     

Macro-floral assemblages in upland Welsh streams in relation to acidity, and their importance to invertebrates

SUMMARY 1. Macro-flora (angiosperms. bryophytes and macroscopic algae) and macroinvertebrates were sampled in 1984 at eighty-eight sites on soft-water streams in upland Wales. Assemblage patterns were related to stream chemistry using TWINSPAN, DECORANA and multiple discriminant analysis.
2. Floral assemblages could be related most strongly to pH and aluminium concentration, with Scapania undulata, Nardia compressa and filamentous chlorophytes characterizing streams of mean pH5.2–5.8, whilst Fontinalis squamosa occurred mostly at pH 5.6–6.2 and Lemanea at pH 5.8–7.0. We propose an indicator system based on thesetaxa.
3. Assemblages of invertebrates and flora concorded highly significantly, sites with Scapania and Nardia holding impoverished faunas. Because some acid sensitive invertebrates (e.g. Ecdyonurus and Ancylus ) can feed from acid tolerant plants (e.g. Scapania ), we hypothesize that they are restricted physiologically from acid streams.     

Effects of flooding on avian top-predators and their invertebrate prey in a monsoonal Taiwan stream

1. The effects of flooding on top predators are poorly understood globally, but particularly in monsoonal streams. We therefore attempted to assess how inter-annual and intra-annual variations in flood magnitude affected an obligate riverine predator, the brown dipper ( Cinclus pallasii ), and its invertebrate prey, in the mountain Tachia River, Taiwan. Major flooding in one of the study years (2005) allowed an insight into the effects of abnormally large flows.
2. The abundance and biomass of insects, and the abundance of dippers, decreased steadily from 2003 to 2005 as flood magnitude grew, but then increased in 2006 when more typical discharge returned. Dipper abundance, insect abundance and insect biomass were all strongly positively inter-related, but negatively related to discharge. Insect biomass, rather than abundance, was more useful in predicting brown dipper abundance.
3. Aquatic insect composition fluctuated among sampling years, revealed by non-metric multidimensional scaling, and these fluctuations were also related to discharge. In turn, dipper abundance and the mean body size of aquatic insects declined with the shift in insect composition as flow increased.
4. These data illustrate how discharge fluctuations can have pronounced effects on top predators in streams, mediated in this case by fluctuating prey abundance. While contributions from bird movement, breeding performance and mortality were not clearly differentiated, our data reveal how dippers have strategies to accommodate varying discharge in river systems. We suggest that the effects of floods on dippers should be taken into account when using this group as indicators of river quality.     

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.     

Seasonal and spatial variation in zooplankton community structure and their relation to possible controlling variables in Lake Okeechobee

1. We examined the behavioural response of stream macroinvertebrates to real and simulated predatory activity by a river bird, the Eurasian dipper, Cinclus cinclus L.
2. In the field, we assessed whether invertebrate drift changed in response to live dippers in enclosures; we found no effects on the drift of any of the five families for which individual analyses were possible, either because it was infrequent, or involved distances too short to be detected (< 0.5 m).
3. In a laboratory stream, we observed prey during encounters with a model dipper which simulated flight, swimming, bill contact with the prey, and stone turning. Invertebrate families varied in their response. Simuliids and hydropsychid caddis lacked effective escape behaviour, consistent with heavy losses through predation by dippers in the wild. Other families either drifted (Baetidae, Gammaridae) or moved away (Heptageniidae, Ephemerellidae, Leuctridae, Perlidae) from the model dipper, but responded only to bill contact or simulated stone turning. Such delayed responses would not protect individuals directly targeted by foraging dippers and partly explain the lack of detectable effects by dippers on drift in the field.
4. We suggest why invertebrates do not show more marked escape responses to this important predator.     

Seasonal and spatial variation in zooplankton community structure and their relation to possible controlling variables in Lake Okeechobee

1. We examined the behavioural response of stream macroinvertebrates to real and simulated predatory activity by a river bird, the Eurasian dipper, Cinclus cinclus L.
2. In the field, we assessed whether invertebrate drift changed in response to live dippers in enclosures; we found no effects on the drift of any of the five families for which individual analyses were possible, either because it was infrequent, or involved distances too short to be detected (< 0.5 m).
3. In a laboratory stream, we observed prey during encounters with a model dipper which simulated flight, swimming, bill contact with the prey, and stone turning. Invertebrate families varied in their response. Simuliids and hydropsychid caddis lacked effective escape behaviour, consistent with heavy losses through predation by dippers in the wild. Other families either drifted (Baetidae, Gammaridae) or moved away (Heptageniidae, Ephemerellidae, Leuctridae, Perlidae) from the model dipper, but responded only to bill contact or simulated stone turning. Such delayed responses would not protect individuals directly targeted by foraging dippers and partly explain the lack of detectable effects by dippers on drift in the field.
4. We suggest why invertebrates do not show more marked escape responses to this important predator.     

The ordination and classification of macroinvertebrate assemblages in the catchment of the River Wye in relation to environmental factors

SUMMARY. 1. Macroinvertebrates were sampled from three habitats at forty-five sites in the catchment of the Welsh River Wye. Species assemblages were ordinated by DECORANA. classified by TWIN-SPAN and related to physico-chemical factors using correlation and multiple discriminant analysis respectively.
2. DECORANA axis 1 was correlated with pH or total hardness, whilst axis 2 correlated with slope or distance from source. TWINSPAN groupings were also related to hardness and, to a lesser extent, slope. Assemblages at soft-water sites (<15gm⁻³ CaCO₃) were composed mostly of Plecoptera but at hard-water sites, even at high slope (>10 m km⁻¹), the fauna was dominated numerically by Ephemerop-tera, net-spinning Trichoptera, Mollusca and Crustacea.
3. We suggest that our data do not support the River Continuum Concept unless there are modifications to allow for multiple gradients.     

Ecological methods for adult populations of Oryctes rhinoceros (Coleoptera, Scarabaeidae)

ABSTRACT. 1. For comparing assessment methods, O.rhinoceros (L.) populations were monitored in five 30–50 ha plots in southern Luzon, Philippines. No consistent correlations were found between number of beetles caught in traps, amount of palm damage and number of breeding sites. Apparently, plots were too small to account for fast dispersal of beetles.
2. Coconut cap traps baited with ethylchrysanthemumate attracted O.rhinoceros adults searching for breeding sites. Several factors were identified influencing catches but trapping was found to be unsuitable for accurately assessing pest populations.
3. A new method which gives estimates on the monthly number of O.rhinoceros attacks on coconut palms, was tested in the Maldives. Palms are climbed about once a year and the sequence of fronds, the number and the position of beetle cuts are recorded.
4. Reduction with age of the length of three teeth on the fore tibia of O. rhinoceros adults was studied in the Philippines. The data was insufficient to obtain a clear correlation between age and length of teeth.
5. The observations indicated that in the Philippines after leaving their site of pupation, O.rhinoceros adults spend about 5 weeks feeding on coconut palms. This is followed by a period of about 7 weeks in breeding sites and, on occasion, additional visits to palms. With these estimates it was possible to relate palm damage records with numbers of feeding adults.     

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 reproductive success of the dipper Cinclus cinclus in relation to the acidity of streams in south-west Scotland

1. The breeding success of dippers Cinclus cinclus was assessed in south-west Scotland over 3 years and related to the acidity of the streams along which they bred. 2. At sites of high acidity, clutch and brood sizes were significantly smaller than those at sites of lower acidity. 3. Egg weight and the incidence of second clutches increased significantly with pH, but there was no evidence to suggest an effect of acidity on hatching success. 4. The rate at which food was delivered to individual nestlings was significantly lower at acidic than non-acidic sites. Nestling weights and survival were lower at more acidic sites. 5. Reduced brood sizes, low nestling survival and the low incidence of second brooding attempts resulted in a significant reduction in total productivity (number of fledglings produced, per pair, per year) at acidic compared with non-acidic streams. 6. It is suggested that pH-related differences in the invertebrate fauna of streams result in low prey availability for dippers on acidic streams, leading to reduced productivity.     

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.     

Characteristics of Culicoides (Diptera, Ceratopogonidae) breeding places near Salisbury, Rhodesia

Abstract. 1. A survey of Culicoides breeding places was made around Salisbury. Out of forty species caught in light traps, only nineteen were reared from breeding places.
2. The seven most abundant species with their characteristic sites were C.pycnostictw in mud around water bodies, especially dams; C.nivosus very prevalent around puddles rich in organic matter: C. distinctipennis along drainage canals intermediate in organic matter; C.gulbenkiani in cow dung especially on damp soil; Caccraensis gp. only in rot holes and tree forks; Cschultzei gp. along streams and drainage canals in mud rather poor in organic matter; C.tropi-calis in drainage canals very low in organic matter; C.imicola possibly breeds widely dispersed mainly in a rich mixture of organic matter and damp soil, with cow pats and edges of water bodies being marginal.
3. The prevalent species are believed to breed all year round. Preponderance of females in the breeding places was found in C.pycnostictus, C.nivosus and C.accraensis gp. while the reverse was true in C.distinctipennis and C.gulbenkiani. The preponderance of males as indication of proximity to breeding sites is thus probably questionable.     

Haematocrit correlates with tail ornament size in three populations of the Barn Swallow (Hirundo rustica)

1. Handicap models of sexual selection propose that male ornaments are indicators of male quality and that honesty is enforced by the costs imposed by the exaggerated ornamental traits. In long-distance migratory birds that feed on the wing, the aerodynamic cost of exaggerated ornamental characters should be particularly high because the size of the ornaments deviates from the natural selection optimum. During migration, birds are expected to raise their oxygen consumption in relation to the energetic demands imposed by their morphology. An increase of haematocrit is an adaptive response to enhance oxygen uptake and efficiency of transfer to the muscular tissues during spells of intense muscular activity.
2. The change of haematocrit of Barn Swallows ( Hirundo rustica ) after their arrival to the breeding sites, and the relationships between haematocrit values recorded after migration and the size of ordinary and sexually selected morphological characters in three Barn Swallow populations were analysed.
3. Males had higher haematocrit values than females. Individual haematocrit values declined after arrival to the breeding sites. Haematocrit values of males were significantly and positively correlated with the size of their ornamental tail but not correlated with other characters, thus suggesting that well-ornamented males, in order to arrive early, have to raise their haematocrit above the level of short-tailed males.
4. Males and females of similar tail length did not differ in their haematocrit, thus suggesting that sexual dimorphism in haematocrit might be functionally related to dimorphism in tail length.
5. Our results are consistent with the handicap principle because long-tailed males experience lower mortality and larger seasonal reproductive success compared with short-tailed males.     

Significance of spring migration and flexibility in flight-muscle histolysis in waterstriders (Heteroptera, Gerridae)

Abstract. 1. Individuals of long-winged waterstrider (Gerridae) species were found in spring far from their breeding habitats, which indicates that they fly before reproduction.
2. Field samples and laboratory studies show that once they return to their breeding sites, many individuals of three waterstrider species ( Gerris odontogaster (Zett.), Gerris lacustris (L.) and Limnoporus rufoscutellatus (Lat.)) histolyse wing muscles and lose flight ability during their reproductive period.
3. The extent of flight-muscle histolysis varies with environmental factors. Food scarcity affects flight-muscle histolysis in G.odontogaster females. In G.Lacustris , flight-muscle histolysis was more common in the laboratory than in the field samples. Proportionately more females than males lost their flight ability by the end of the reproductive period.
4. Flight ability had direct costs in reproductive potential with (non-flyer) females, which histolysed their flight muscles, laying more eggs than (flyer) females, which maintained flight ability. This was also the case during food scarcity. Non-flyer males of G.odontogaster survived longer than flyer males.
5. Spring migration was distinguished from dispersal during the reproductive period, because these flights serve different functions. Flight-muscle histolysis of females during reproduction is a qualitative reproductive option, with a trade-off between dispersal ability and reproductive potential. Ability to change reproductive behaviour depending on environmental conditions increases an individual's ability to cope with a large variety of habitats.     

A comparison of Acroneuria lycorias (Plecoptera) production and growth in northern Michigan hard- and soft-water streams

1. Density, biomass, production and growth of a predaceous stonefly, Acroneuria lycorias, were compared between fourth-order hard- and soft-water streams in Michigan's upper peninsula, U.S.A. 2. Mean densities, estimated from Hess samples, were higher (100 ± 17 individuals m^?2) at the hard-water site than at the soft-water site (40 ± 9 ind. m^?2). Mean dry weight biomass was 4.9 times greater at the hard-water site. 3. Mean annual production, calculated using the size frequency method, was 5.0 times greater at the hard-water site (2.18 ± 0.44 g dry weight m^?2yr^?1) than at the soft-water site (0.43 ± 0.02g dry weight m^?2yr^?1). Annual production/mean biomass ratios were similar between sites. 4. Monthly growth rates of naturally occurring nymphs of paired cohorts were similar in both streams. Individual growth rates were similar for nymphs reared in artificial streams at high and low water hardnesses with unlimited food and space. 5. Stonefly production and growth rates were influenced more by indirect physical, biological, or habitat factors than by streamwater cation concentrations.     

The choice of foraging methods of the Brown Dipper,Cinclus pallasii (Aves: Cinclidae)

The Brown Dipper,Cinclus pallasii, forages for large prey by diving deeply among submerged rocks, and for small prey in shallow water by wading and pecking on the bottom. Foraging by diving entailed a higher energy cost but resulted in taking larger prey than foraging by wading-and-pecking. Foraging by diving was seldom observed from May to October, but increased from December to April, when the dippers came into breeding condition and the proportion of large prey in the benthic fauna increased. During the breeding season, adult dippers fed themselves mainly by foraging by wading-and-pecking, whereas those feeding nestlings and fledglings foraged by diving. The foraging methods employed was determined by the individuals being fed and the abundance of the large prey. Foraging by diving may be advantageous for individuals provisioning their young because of the high gross intake rate, but may be less advantageous for the foragers themselves because of the high energy cost.     

THE BREEDING BIRDS OF THE HULEH SWAMP AND LAKE (NORTHERN ISRAEL).

1. The Huleh swamp and lake, which are now under a draining scheme, are described in relation to their vegetation habitats and the birds breeding in them.
2. Data are presented on the breeding birds, concerning their distribution within Israel, laying season, nest site and clutch size.
3. The Common Heron, White-tailed Eagle, Marsh Harrier, Mallard, Baillon's Crake, Yellow Wagtail, Moustached Warbler and the Great Reed Warbler are here first recorded as breeding in Israel.
4. The relationship between Acrocephalus arundinaceus and A. stentoreus is discussed.
5. Some remarks are made on habitat selection of the warblers breeding in the area.     

Lethal and sub-lethal effects of acid, aluminium and lime on Gammarus pulex during repeated simulated episodes in a Welsh stream

SUMMARY. 1. A soft-water stream in upland mid-Wales was dosed with sulphuric acid, aluminium sulphate and limestone slurry to produce a zone of aluminium at low pH. and one of aluminium, low pH and added lime.
2. Three episodes of 24-h duration were induced, each separated by a 12-h inter-dosing period, and the responses of the crustacean Gammarus pulex monitored throughout.
3. G. pulex parasitized by the acanthocephalan Pomphorhynchus laevis were significantly more sensitive than unparasitized individuals.
4. Liming significantly reduced mortality during dosing but postexposure deaths were greatest for animals which had been exposed in the time zone.
5. The disruption of precopulalory behaviour in G. pulex was rapid and liming did not mitigate this response.     

Dynamics and Characterization of Aedes aegypti (L.) (Diptera: Culicidae) Key Breeding Sites

The present study aimed to analyze the dynamics of containers used as breeding sites by Aedes aegypti (L.) in the city of Aracaju, SE, one of the Northeast Brazilian states. A total of three entomological surveys were performed during different precipitation levels. Breeding sites were categorized according to their function into storage, disposable containers, and reusable containers. “Mean number of pupae” and “frequency of each type of breeding site” were the criteria considered to identify key breeding sites. House index and Breteau index were calculated in each survey. A total of 3,647 water reservoirs were found, of which 220 were breeding sites, where 22,880 immature forms were identified. There were no differences in the mean number of larvae of several types of breeding sites and in the number of larvae among surveys. Larval indices showed a reduction in the second visit, but with no effect on adult occurrence when the number of pupae was considered. Key breeding sites resulted from containers used for water storage. The area studied showed conditions favorable to a new epidemic of dengue fever.     

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.